Taxonomy of Rhodacaroidea mites (Acari: Mesostigmata) Raphael de Campos … · 2012. 3. 23. ·...

Universidade de São Paulo

Escola Superior de Agricultura “Luiz de Queiroz”

Taxonomy of Rhodacaroidea mites (Acari: Mesostigmata)

Raphael de Campos Castilho

Thesis submitted in partial fulfillment of the

requirements for the degree of Doctor in Science.

Area of concentration: Entomology

Piracicaba

2012

2

Raphael de Campos Castilho

Engenheiro Agrônomo


Adviser:

Prof. Dr. GILBERTO JOSÉ DE MORAES

Thesis submitted in partial fulfillment of the

requirements for the degree of Doctor in Science.

Area of concentration: Entomology

Piracicaba

2012

Dados Internacionais de Catalogação na Publicação DIVISÃO DE BIBLIOTECA - ESALQ/USP

Castilho, Raphael de Campos Taxonomy of Rhodacaroidea mites (Acari: Mesostigmata) / Raphael de Campos

Castilho. - - Piracicaba, 2012. 579 p. : il.

Tese (Doutorado) - - Escola Superior de Agricultura “Luiz de Queiroz”, 2012.

1. Ácaros predadores 2. Classificação 3. Ácaros de solo 4. Controle biológico I. Título

CDD 595.42 C352t

“Permitida a cópia total ou parcial deste documento, desde que citada a fonte – O autor”

3

To GOD

Source of perseverance and life,

To my mother

Sonia Regina de Campos

For her love, tenderness and comprehension.

To my partner

Karina Cezarete Semençato

for her love, patience and unfailing support to me

Offer

To Prof. Dr. Gilberto José de Moraes

For his valuable guidance, friendship and recognition of my

work

Special thanks

5

Ackanowledgements

To Escola Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de São

Paulo (USP), and especially to ―Departamento de Entomologia e Acarologia‖ for providing

all intellectual and material support necessary for the proper development of this work;

I am especially grateful to Carlos H. W. Flechtman (ESALQ/USP, Brazil), for his

valuable suggestions and for help with translation of German, Latim, French and English

papers;

To Italo Delalibera Jr. (ESALQ/USP, Brazil) for participating in the whole process of

the Doctorate training;

To Bruce Halliday (CSIRO, Australia) for his scientific and intelectual input;

To Maurice Sabelis, Izabela Lesna and Farid Faraji (Section Population Biology,

Institute of Biodiversity and Ecosystem Dynamics-IBED, University of Amsterdam, The

Netherlands) for their logistical support and valuable help in my ―Doctorate Sandwich‖;

To Bruce Halliday (CSIRO, Australia), João Paulo Z. Narita (ESALQ/USP, Brazil)

and Mahdi Jalaeian (Research Center of Khorasan Razavi, Iran) for coauthoring several

papers, resulting from this research;

To Hans Klompen (Ohio State University, Columbus, USA) and Debbie Creel (USDA

Systematic Entomology Laboratory, Beltsville, Maryland, USA) for valuable information

about the location of Hurlbutt‘s type specimens;

To Giuseppino Sabbatini and Roberto Nannelli (Istituto Sperimentale per la Zoologia

Agraria, Florence, Italy) for providing photos of species described by A. Berlese;

To María L. Moraza (Facultad de Ciencias, Universidad de Navarra, Spain), E.A.

Ueckermann (ARC-Plant Protection Research Institute, South Africa) and Evert E. Lindquist

(Agriculture & Agri-Food Canada, Canada) for important personal information.

To Jason Dunlop (Arachnologische Sammlung des Museums für Naturkunde, Berlin,

Germany), Axel Christian (Staatlichen Museum für Naturkunde Görlitz, Görlitz, Germany),

Anne Baker (Natural History Museum, London, England) and Pieter Theron (North-West

University - Potchefstroom Campus, Souh Africa) for providing access to types for this study.

To Acarologia (Serge Kreiter); Bydraes van die P.U. vir C.H.O., Reeks B:

Natuurwetenskappe (Frikkie van Niekerk, North West University); Deutsche Entomologische

Zeitschrift (Hannelore Hoch); Dopovidi Akademii Nauki Ukrainskoi RSR, Seriya B. (Galina

I. Shcherbak); Entomologist‘s Monthly Magazine (Ian Johnson, Pemberley Books); Hong

Kong University Press (Christy Leung); Systematic and Applied Acarology (Zhi-Qiang

Zhang); The Canadian Entomologist (Evert E. Lindquist); Zoologischer Anzeiger (Natalie

David, Elsevier) for permission to reproduce their figures.

To Frédéric Beaulieu (Agriculture & Agri-Food Canada, Canada), María L. Moraza

(Facultad de Ciencias, Universidad de Navarra, Spain), Shahrooz Kazemi (Tarbiat Modares

6

University, Iran) and Carlos H. W. Flechtmann (ESALQ/USP, Brazil) for their numerous

suggestions for the improvement of several papers, resulting from this research;

To Eliot W. Kitajima (ESALQ/USP), for his help in the SEM examination of the

mites;

To Abd-Allah Afifi, Ahmed Fouly, Bruce Halliday, Dariusz J. Gwiazdowicz, Heinrich

Schatz, Li-Ming Ma, Pavel Klimov, Peter Masan, Tetsuo Gotoh for providing copies of hard

papers;

To USP COMUT-Librarians for providing copies of many papers;

To Lásaro V. F. da Silva (ESALQ/USP, Brazil), for logístic supporte and laboratory

help;

To my friends of Acarology Laboratory of ESALQ/USP, Ana C. Cavalcanti, Alberto

D. G. Alvarado, Aníbal R. Oliveira, Camila Dainese, Daiane H. Nunes, Daniel C. Oliveira,

Diana M. Rueda, Edmilson S. Silva, Érika P. J. Britto, Fernanda de C. N. Esteca, Fernando R.

da Silva, Geraldo J. N. de Vasconcelos, Grazielle F. Moreira, Jandir C. Santos, João Paulo Z.

Narita, John J. S. Ausique, Leocádia S. Martinez, Letícia H. Azevedo, Marcos R. Bellini,

Marina F. C. Barbosa, Natasha Iwanicki, Olivia S. Camargo, Paula C. Lopes, Peterson R.

Demite, Ralf V. Araújo, Renan V. da Silva, Renata A. P. Freire, Renata A. Simões, Samuel

Roggia, Sheila Spongoski, Tatiane M. M. G. Castro, Thiago R. Castro, Vanessa S. Duarte and

Vitalis W. Wekesa for their help, support and friendship;

To colleagues of ―Departamento de Entomologia e Acarologia‖ of ESALQ/USP for

support and friendship;

To laboratory and administrative staff of ―Departamento de Entomologia e

Acarologia‖ of ESALQ/USP, Carolina D. Jorge, Claudete A. A. Marques, José L. F. Piedade,

Josenilton L. Mandro, Maria Marta Coletti, Marinalda S. Zambon, Regina C. B. de Moraes,

Rosâgela A. da Silva and Vera L. Durrer;

To Silvia M. Zinsly and Maria da Glória E. da Silva (Library of ESALQ/USP) for the

corrections of the references;

and for those who in one way or and other helped me in this work.

Special ackanowledgements

To my family, especially my father, Luis Carlos Castilho, and my brother, Vinicius de

Campos Castilho, and to the relatives of Karina C. Semençato, for their warm support and

love;

To dear researchers of Embrapa Algodão, Carlos Alberto Domingues da Silva and

José Ednilson Miranda, and Edmilson Santos Silva, of ―Universidade Federal de Alagoas‖,

for their continuous and strong encouragement in my academic activities.

This work was supported by CAPES and CNPq, Conselho Nacional de

Desenvolvimento Científico e Tecnológico, Brazil.

7

TABLE OF CONTENTS

ABSTRACT ............................................................................................................................. 11

RESUMO ................................................................................................................................. 13

1 INTRODUCTION .............................................................................................................. 15

References ............................................................................................................................. ...17

2 CATALOGUE OF THE MITE FAMILY RHODACARIDAE OUDEMANS, WITH

NOTES ON THE CLASSIFICATION OF THE RHODACAROIDEA (ACARI:

MESOSTIGMATA) ........................................................................................................... 19

Abstract ............................................................................................................................ ...19

Resumo ............................................................................................................................ ...20

2.1 Introduction ............................................................................................................... ...20

2.2 Materials and methods ............................................................................................... ...25

2.3 Results ....................................................................................................................... ...28

2.4 Discussion .................................................................................................................. ...99

References ..................................................................................................................... ...100

3 CATALOGUE OF THE MITE FAMILIES DIGAMASELLIDAE,

LAELAPTONYSSIDAE, OLOGAMASIDAE AND TERANYSSIDAE (ACARI:

RHODACAROIDEA: MESOSTIGMATA), AND A KEY TO THE WORLD GENERA

OF THESE FAMILIES .................................................................................................... 121

Abstract .......................................................................................................................... ...121

Resumo .......................................................................................................................... ...121

3.1 Introduction ............................................................................................................. ...122

3.2 Materials and methods ............................................................................................. ...126

3.3 Results ..................................................................................................................... ...128

References ..................................................................................................................... ...365

4 REVISION OF THE GENERA Interrhodeus, Pennarhodeus and Poropodalius (ACARI:

RHODACARIDAE) ......................................................................................................... 409

Abstract .......................................................................................................................... ...409

Resumo .......................................................................................................................... ...409

4.1 Introduction ............................................................................................................. ...409


4.3 Results ..................................................................................................................... ...410

8

4.4 Discussion ................................................................................................................ ...456

References ...................................................................................................................... ...457

5 REVISION OF THE GENUS Protogamasellopsis (ACARI: MESOSTIGMATA:

RHODACARIDAE) ......................................................................................................... 459

Abstract .......................................................................................................................... ...459

Resumo........................................................................................................................... ...459

5.1 Introduction .............................................................................................................. ...459


5.3 Results ...................................................................................................................... ...461

References ...................................................................................................................... ...482

6 RHODACARIDAE MITES (ACARI: MESOSTIGMATA: RHODACAROIDEA) FROM

THE STATE OF SÃO PAULO, BRAZIL, WITH DESCRIPTIONS OF A NEW GENUS

AND THREE NEW SPECIES ......................................................................................... 485

Abstract .......................................................................................................................... ...485

Resumo........................................................................................................................... ...485

6.1 Introduction .............................................................................................................. ...485


6.3 Results ...................................................................................................................... ...486

6.4 Discussion ................................................................................................................ ...503

References ...................................................................................................................... ...503

7 TWO NEW SPECIES OF RHODACARIDAE (MESOSTIGMATA:

RHODACAROIDEA) FROM IRAN ............................................................................... 507

Abstract .......................................................................................................................... ...507

Resumo........................................................................................................................... ...507

7.1 Introduction .............................................................................................................. ...507


7.3 Results ...................................................................................................................... ...508

7.4 Discussion ................................................................................................................ ...517

References ...................................................................................................................... ...517

8 A NEW SPECIES OF Gamasiphis (ACARI: OLOGAMASIDAE) FROM BRAZIL,

WITH A KEY TO SPECIES FROM THE NEOTROPICAL REGION .......................... 521

Abstract .......................................................................................................................... ...521

Resumo........................................................................................................................... ...521

8.1 Introduction .............................................................................................................. ...521

9


8.3 Results ..................................................................................................................... ...523

References ..................................................................................................................... ...537

9 THREE NEW SPECIES OF Gamasiphis (ACARI: MESOSTIGMATA:

OLOGAMASIDAE) FROM BRAZIL, WITH COMPLEMENTARY INFORMATION

ABOUT Gamasiphis plenosetosus KARG AND A KEY TO THE WORLD SPECIES OF

THE GENUS .................................................................................................................... 541

Abstract .......................................................................................................................... ...541

Resumo .......................................................................................................................... ...541

9.1 Introduction ............................................................................................................. ...542


9.3 Results ..................................................................................................................... ...543

9.4 Discussion ................................................................................................................ ...570

References ..................................................................................................................... ...572

10 FINAL CONSIDERATIONS ........................................................................................... 579

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ABSTRACT


The mite superfamily Rhodacaroidea is a member of the order Mesostigmata. The

families Digamasellidae Evans, Halolaelapidae Karg, Laelaptonyssidae Womersley,

Ologamasidae Ryke, Rhodacaridae Oudemans and Teranyssidae Halliday constitute this

superfamily. The main morphological characteristic considered in this study for inclusion of

mites in the Rhodacaroidea is the insertion of seta st4 on the sternal shield. These mites have

been found in soil, litter, rodent nests, mosses, lichens, termite nests, on Coleoptera [mainly

bark beetles (Curculionidae: Scolytinae)] and in galleries made by them in tree trunks.

Digamasellids, ologamsids and rhodacarids are commonly mentioned in the literature as

predators of nematodes, small insects, mites and springtails, and at least one species, appears

to have potential as a biological control agent against insect and mite pests in the soil. The

taxonomic concepts of the families belonging to Rhodacaroidea are confusing do to the

frequent taxonomic changes over time. As a consequence, it has been quite often difficult to

determine to which family a determined species of this group belongs. Several authors

contributed significantly to the taxonomic knowledge of Rhodacaroidea, but few are still

professionally active. The general objective of this thesis was to establish the bases to

facilitate the identification of Rhodacaroidea mites. Diagnoses of the genera of Rhodacaridae,

a key for their identification and an updated list of the species within each genus were

prepared. This family is presently composed of 148 species arranged in 15 genera. A

dichotomous key to the genera as well as updated and complemented lists of the species

within each genus of Digamasellidae, Laelaptonyssidae, Ologamasidae and Teranyssidae

were also prepared. Digamasellidae is composed of 277 species arranged in 11 genera,

Laelaptonyssidae, 8 species in one genus, Ologamasidae 450 species in 44 genera and

Teranyssidae, a single species. The species of the rhodacarid genera Interrhodeus Karg,

Pennarhodeus Karg, Poropodalius Karg and Protogamasellopsis Evans and Purvis were

examined and a characterization of the genera, diagnoses of each species, complementary

descriptions of some species and a key to help in the separation of the species of each genus

were provided. The re-examination of those species allowed the conclusion that they are

correctly placed in the Rhodacaridae. Also as result of this work, a new genus and five new

species of Rhodacaridae and four new species of Ologamasidae were described and new

records of mites of those groups were determined, as the result of analyses of mites from

southern Brazil and Iran. A taxonomic key to separate the 60 species of Gamasiphis

(Ologamasidae) described from different parts of the world was prepared. The study

conducted represents a significant contribution to the knowledge of the taxonomy of

Rhodacaroidea.

Keywords: Soil mites; Predators; Biological control; Digamasellidae; Laelaptonyssidae;

Ologamasidae; Rhodacaridae; Teranyssidae

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RESUMO

Taxonomia de ácaros Rhodacaroidea (Acari: Mesostigamata)

Os ácaros da superfamília Rhodacaroidea são membros da ordem Mesostigmata. As

famílias Digamasellidae Evans, Halolaelapidae Karg, Laelaptonyssidae Womersley,

Ologamasidae Ryke, Rhodacaridae Oudemans e Teranyssidae Halliday constituem esta

superfamília. A principal característica morfológica considerada neste estudo para a inclusão

de ácaros em Rhodacaroidea é a inserção de seta st4 no escudo esternal. Estes ácaros são

encontrados no solo, folhedo, ninhos de roedores, musgos, liquens, ninhos de cupins, em

Coleoptera [principalmente besouros de casca (Curculionidae: Scolytinae)] e em galerias

feitas por estes em troncos de árvores. Os Digamasellidae, Ologamsidae e Rhodacaridae são

comumente citados na literatura como predadores de nematóides, pequenos insetos, ácaros e

Collembola. Pelo menos uma espécie parece ter potencial como agente de controle biológico

contra insetos e ácaros pragas que ocorrem no solo. Os conceitos taxonômicos das famílias

pertencentes à Rhodacaroidea são confusos devido às mudanças taxonômicas freqüentes ao

longo do tempo. Como conseqüência, freqüentemente, é difícil de se determinar a que família

uma determinada espécie deste grupo pertence. Vários autores contribuíram

significativamente para o conhecimento taxonômico das espécies de Rhodacaroidea, mas

poucos desses ainda estão profissionalmente ativos. O objetivo geral desta tese foi estabelecer

as bases para facilitar a identificação de ácaros Rhodacaroidea.. A elaboração da diagnose dos

gêneros de Rhodacaridae, uma chave para a identificação destes e uma lista atualizada das

espécies dentro de cada gênero foram preparadas. Esta família é atualmente composta por 148

espécies distribuídas em 15 gêneros. Uma chave dicotômica para a separação dos gêneros,

bem como uma atualização e complementação das listas das espécies dentro de cada gênero

de Digamasellidae, Laelaptonyssidae, Ologamasidae e Teranyssidae também foram

preparadas. Digamasellidae é composta de 277 espécies de 11 gêneros, Laelaptonyssidae, 8

espécies de um gênero, Ologamasidae 450 espécies de 44 gêneros e Teranyssidae, uma única

espécie. As espécies de Interrhodeus Karg, Pennarhodeus Karg, Poropodalius Karg e

Protogamasellopsis Evans e Purvis, todos Rhodacaridae, foram examinadas, realizando-se a

caracterização dos gêneros, a diagnose de cada espécie, descrições complementares de

algumas espécies e uma chave para ajudar na separação das espécies de cada gênero. A re-

avaliação dessas espécies permitiu a conclusão de que estas estão corretamente colocadas em

Rhodacaridae. Também como resultado deste trabalho, um novo gênero e cinco novas

espécies de Rhodacaridae assim como quatro novas espécies de Ologamasidae foram descritas

e novos registros de ácaros desses grupos foram estabelecidos, como resultado do estudo de

ácaros do sudeste do Brasil e Iran. Uma chave taxonômica para separar as 60 espécies de

Gamasiphis (Ologamasidae) descritas de diferentes partes do mundo foi preparada. O estudo

realizado representa uma significativa contribuição para o conhecimento da taxonomia de

Rhodacaroidea.

Palavras-chave: Ácaros de solo; Predadores; Controle biológico; Digamasellidae;

Laelaptonyssidae; Ologamasidae; Rhodacaridae; Teranyssidae

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1 INTRODUCTION

The mite superfamily Rhodacaroidea is a member of the order Mesostigmata. In a

recent publication, Lindquist; Krantz and Walter (2009) considered Digamasellidae Evans,

Halolaelapidae Karg, Laelaptonyssidae Womersley, Ologamasidae Ryke, Rhodacaridae

Oudemans and Teranyssidae Halliday to constitute superfamily. The main morphological

characteristic considered in this study for inclusion of mites in the Rhodacaroidea is the

insertion of seta st4 on the sternal shield. Members of all of these families, except the

ologamasid Oriflammella Halliday and the members of Halolaelapidae, have this

characteristic. The latter family is in need of a detailed revision to determine whether or not it

should be included as Rhodacaroidea. Therefore, Halolaelapidae is not considered in this

study.

Digamasellidae, Ologamasidae and Rhodacaridae mites have been found in soil, litter,

rodent nests, mosses, lichens, and other types of organic matter in contact with the soil. In

addition, digamasellids have been found on Coleoptera, mainly bark beetles (Curculionidae:

Scolytinae), and in galleries made by them in tree trunks. Digamasellids, ologamsids and

rhodacarids are commonly mentioned in the literature as predators of nematodes, small

insects, mites and springtails (KARG, 1971; LEE, 1974; MOSER, 1975; ENDA; TAMURA,

1977; WALTER; HUNT; ELLIOTT, 1988; ABOU-EL-SOUD; SHOEIB, 2000; BEAULIEU;

WALTER, 2007). At least one species, Protogamasellopsis posnaniensis Wiśniewski and

Hirschmann (Rhodacaridae), appears to have potential as a biological control agent against

insect and mite pests in the soil (CASTILHO et al., 2009).

Laelaptonyssidae and Teranyssidae mites have been found from termite nests, but the

details of the relationship between those organisms are not known. Krantz (2000) suggested

laelaptonyssid to be phoretic on termites rather than parasitic, feeding on nematodes in the

termites' nest material.

However, the taxonomic concepts of the families belonging to Rhodacaroidea are

confusing, leading to a very complex taxonomic history because of the frequent changes over

time. As a consequence, it has been quite often difficult to determine to which family a

determined species of this group belongs. Most descriptions of species and other taxonomic

works on mites of this group referred and still refer to most species as Rhodacaridae.

The whole world is going through a lack of specialists in the taxonomy of many

different types of organisms, and the same has been observed in relation to mites of this

group. Several authors contributed significantly to the taxonomic knowledge of

16

Rhodacaroidea species, namely C. Willmann, W. Hirschmann, P.A.J. Ryke, G.C. Loots, E.E.

Linquist, W. Karg, D.C. Lee, G.I. Shcherbak, J. Wiśniewski, R.B. Halliday and others.

However, few of those are still active.

Rhodacaroids as a group have potential as biological control agents of soil pests

(CASTILHO et al., 2009) and are commonly found in tropical regions (MINEIRO;

MORAES, 2001; SILVA; MORAES; KRANTZ, 2004). However, the success of the search

process to the use of predators depends on the correct identification of the organisms

involved. A question then arises: how to change this situation in order to facilitate the

identification of species of this group?

The general objective of this thesis is to establish the bases to facilitate the

identification of Rhodacaroidea mites. Therefore, specific objectives were established:

Review the literature concerning the taxonomy of Rhodacaridae, to present a new

characterisation of the family and the included genera, to construct keys for identification of

genera, as well as to update and complement the a list of the species considered to belong to

the Rhodacaridae, with relevant taxonomic information about each;

Review the literature concerning the taxonomy of Digamasellidae, Laelaptonyssidae,

Ologamasidae and Teranyssidae, to construct a key for the identification of genera of these

groups, as well as to update and complement the lists of species considered to belong to the

these families, with relevant taxonomic information about them;

Provide redescriptions of Interrhodeus Karg, Pennarhodeus Karg; Poropodalius Karg and

Protogamasellopsis Evans and Purvis and their included species, and the preparation of keys

for identification of species of these genera;

Provide the description of one new genus and three new species, and report the rhodacarid

species found in surveys conducted in southeastern Brazil;

Provide the description of two new species of Rhodacaridae found in surveys conducted in

Iran;

Provide the description of four new species of Gamasiphis (Ologamasidae) from

Piracicaba, Brazil and a taxonomic key to separate the world species of this genus.

17

References

ABOU-EL-SOUD, A.B.; SHOEIB, A.A. Root-knot nematode Meloidogyne javanica and

acarid mite, Acarus siro L. predation by digamasellid mite, Dendrolaelaps rasmii Nasr and

Mersal. Al-Azhar Journal of Agricultural Research, Cairo, v. 31 p. 45-50, 2000.

BEAULIEU, F.; WALTER, D.E. Predation in suspended and forest floor soils observations

on Australian mesostigmatic mites. Acarologia, Paris, v. 47, p. 43-54, 2007.

CASTILHO, R.C.; MORAES, G.J. de; SILVA, E.S.; SILVA L.O. Predation potential and

biology of Protogamasellopsis posnaniensis Wisniewski & Hirschmann (Acari:

Rhodacaridae). Biological Control, Orlando, v. 48, p. 164-167, 2009.

ENDA, N.; TAMURA, H. Nematophagous mites found on adults of the Japanese pine

sawyer. Shinrin Boeki [Forest Pests], Tokyo, v. 26, p. 188-190, 1977. [in Japanese]

KARG, W. Acari (Acarina), Milben: Unterordnung Anactinochaeta (Parasitiformes):

Die freilebenden Gamasina (Gamasides), Raubmilben. Jena: Gustav Fischer Verlag, 1971.

475 p.

KRANTZ, G.W. Two new species of the genus Laelaptonyssus Womersley from North

America and Australia, with observations supporting the reinstatement to family level of the

subfamily Laelaptonyssinae sensu Lee, 1970 (Acari: Mesostigmata: Rhodacaroidea).

Acarologia, Paris, v. 41, p. 25-38, 2000.

LEE, D.C. Rhodacaridae (Acari: Mesostigmata) from near Adelaide, Australia. III. –

Behaviour and development. Acarologia, Paris, v. 16, p. 21-44, 1974.

LINDQUIST, E.E.; KRANTZ, G.W.; WALTER, D.E. Mesostigmata. In: KRANTZ, G.W.;

WALTER, D.E. (Eds.). A manual of acarology. 3rd

ed. Lubbock: Texas Tech University

Press, 2009. p. 124-232.

MINEIRO, J.L.C.; MORAES, G.J de. Gamasida (Arachnida: Acari) edáficos de Piracicaba,

Estado de São Paulo. Neotropical Entomology, Londrina, v. 30, n. 3, p. 379-385, 2001.

18

MOSER, J.C. Mite predators of the southern pine beetle. Annals of the Entomological

Society of America, College Park, v. 68, p. 1113-1116, 1975.

SILVA, E.S.; MORAES G.J. de; KRANTZ, G.W. Diversity of edaphic rhodacaroid mites

(Acari: Mesostigmata: Rhodacaroidea) in natural ecosystems in the State of São Paulo, Brazil.

Neotropical Entomology, Londrina, v. 33, n. 5, p. 547-555, 2004.

WALTER D.E.; HUNT, H.W.; ELLIOTT, E.T. Guilds or functional groups? An analysis of

predatory arthropods from a shortgrass steppe soil. Pedobiologia, Jena, v. 31, p. 247–260,

1988.

19

2 CATALOGUE OF THE MITE FAMILY RHODACARIDAE OUDEMANS, WITH

NOTES ON THE CLASSIFICATION OF THE RHODACAROIDEA (ACARI:

MESOSTIGMATA)

Abstract

The Rhodacaridae Oudemans are free living, cosmopolitan, edaphic mites, which are

considered in the literature to be predators. The taxonomic concept of the family has changed

considerably over time, making it very difficult for non-taxonomists to decide on the correct

placement of many species. Even taxonomists sometimes find it difficult to determine

whether a given species belongs to this family or not, because many of the old descriptions

are not sufficiently detailed. Also, the family placement of some genera has been very

unstable. A historic review of the literature on the classification of the Rhodacaridae is

presented. Diagnoses are given for the families of Rhodacaroidea is given, followed by a

checklist of genera and sub-genera for each family. The families included are Digamasellidae

Evans, Halolaelapidae Karg, Laelaptonyssidae Womersley, Ologamasidae Ryke,

Rhodacaridae and Teranyssidae Halliday. Diagnoses are given for the genera of

Rhodacaridae, and a key for their identification, derived from a standardized database of

character states. Finally, a list of species within each genus of this family was updated and

complemented, giving relevant taxonomic information about the respective types, and

providing references to nomenclatural changes, synonymy, and redescriptions of each species.

In total, 148 rhodacarid species and one subspecies are listed in this work, arranged in 15

genera. The most diverse genera are Afrodacarellus Hurlbutt and Rhodacarus Oudemans,

each with about 20% of the valid species. Five of the genera are monotypic. Taxonomic

confusion surrounds some groups of species, especially in the genus Rhodacarus. It appears

that Rhodacarus calcarulatus Berlese, R. coronatus Berlese, R. pallidus Hull, R. reconditus

Athias-Henriot and R. roseus Oudemans have often been misidentified, and these species are

in need of detailed revision.

Keywords: Catalogue; Dichotomous key; Rhodacaroidea; Soil mites; Taxonomy

20

Resumo

Rhodacaridae Oudemans são ácaros de vida livre, cosmopolitas, edáficos e

considerados na literatura como predadores. O conceito taxonômico da família mudou

consideravelmente ao longo do tempo, tornando-se muito difícil para não-taxonomistas

decidir a classificação correta de muitas espécies. Mesmo para taxonomistas, às vezes é difícil

determinar se uma determinada espécie pertence a esta família ou não, porque muitas das

descrições antigas não são suficientemente detalhadas. Além disso, a classificação de alguns

gêneros da família tem sido muito instável. Uma revisão da literatura sobre a classificação dos

Rhodacaridae é apresentada. Diagnósticos são dados para as famílias de Rhodacaroidea,

seguindo-se pela apresentação de uma lista de gêneros e sub-gêneros para cada família. As

famílias incluídas são Digamasellidae Evans, Halolaelapidae Karg, Laelaptonyssidae

Womersley, Ologamasidae Ryke, Rhodacaridae e Teranyssidae Halliday. Diagnósticos e uma

chave para a identificação são dados para os gêneros de Rhodacaridae. Estes foram

elaborados tendo como base um banco de dados padronizado da caracterização das espécies.

Por fim, uma lista de espécies de cada gênero foi atualizada e complementada, fornecendo

informações taxonômicas relevantes sobre os respectivos tipos, e fornecendo referências a

mudanças nomenclaturais, sinonímias e redescrições de cada espécie. No total, 148 espécies e

uma subespécie de Rhodacaridae arranjados em 15 gêneros estão listadas neste trabalho. Os

gêneros mais diversos são Afrodacarellus Hurlbutt e Rhodacarus Oudemans, cada um com

cerca de 20% das espécies válidas. Cinco dos gêneros são monotípicos. Incertezas ainda

existem em relação a alguns grupos de espécies, especialmente em Rhodacarus.

Aparentemente, Rhodacarus calcarulatus Berlese, R. coronatus Berlese, R. pallidus Hull, R.

reconditus Athias-Henriot e R. roseus Oudemans têm sido freqüentemente identificados

incorretamente. Essas espécies necessitam de uma revisão detalhada.

Palavras-chave: Catálogo; Chave dicotômica; Rhodacaroidea; Ácaros de solo; Taxonomia

2.1 Introduction

The mite family Rhodacaridae Oudemans is a member of the superfamily

Rhodacaroidea in the order Mesostigmata. The Rhodacaridae are free living, cosmopolitan

edaphic mites found mainly in the top few centimetres of the mineral soil layer. They are

often reported in ecological surveys of the arthropod fauna of agricultural soils (for example

21

EL TITI, 1984, 1986; KOEHLER, 1991), but they are also found in mosses, lichens, leaf

litter, and other types of organic matter in contact with the soil. Rhodacarids are commonly

mentioned in the literature as predators, but very few papers have dealt with their feeding

behaviour. Some species have been found to feed on Collembola, nematodes and mites

(KARG, 1971; LEE, 1974; WALTER; HUNT; ELLIOTT, 1988), and at least one species,

Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, appears to have potential as a

biological control agent for insect and mite pests in soil (CASTILHO et al., 2009).

The taxonomic concept of the Rhodacaridae has changed considerably over time,

making it difficult for non-taxonomists to understand which species belong to this group.

Quite often, taxonomists also have difficulty in determining whether or not a described

species belongs to this family, given that many of the old descriptions are not sufficiently

detailed.

Oudemans (1902) established Rhodacarinae as a subfamily of Parasitidae Oudemans,

to contain his new genus and species Rhodacarus roseus Oudemans, 1902. Halbert (1915)

raised the subfamily to family level. Willmann (1935) described two new genera of

Rhodacaridae, Rhodacaropsis Willmann, 1935 and Rhodacarellus Willmann, 1935.

Oudemans (1939a) included Rhodacarellus in his newly-created family Gamasolaelaptidae

Oudemans, but Vitzthum (1941) and Baker and Wharton (1952) included all three genera in

their concept of Rhodacaridae. Evans (1957) extended the concept of the Rhodacaridae,

including in this family all genera of Mesostigmata having a three-tined palp tarsal claw and a

divided dorsal shield, namely Leitneria Evans, 1957, Rhodacarus, Rhodacarellus,

Halolaelaps Berlese and Trouessart, 1889, Saprolaelaps Leitner, 1946, Euryparasitus

Oudemans, 1902 and Cyrtolaelaps Berlese, 1887. He made no reference to Rhodacaropsis.

Willmann (1959) described the new genus Rhodacaroides Willmann, 1959,

presumably in the Rhodacaridae, although he did not clearly state its family placement. Two

new genera of Rhodacaridae were described in the early 1960s: Allogamasellus Athias-

Henriot, 1961 and Evanssellus Ryke, 1961a. Ryke (1961b) described Gamaselliphis Ryke,

1961b as a subgenus of Cyrtolaelaps, and this subgenus was raised to generic status by Lee

(1970).

Ryke (1962a) further expanded the concept of Rhodacaridae, including in this family

all genera of Mesostigmata whose deutonymphs had separate podonotal and opisthonotal

shields, even when the palp tarsal claw was two-tined. He divided the family into two

subfamilies, Rhodacarinae, containing species whose adults also had separate podonotal and

opisthonotal dorsal shields, and Ologamasinae, whose adults had these shields fused. His

22

subfamily Rhodacarinae included Antennoseius Berlese, 1916, Asca von Heyden, 1826,

Cyrtolaelaps, Evanssellus, Halolaelaps, Leitneria, Longoseius Chant, 1961, Rhodacarus,

Saintdidieria Oudemans, 1939b and Saprolaelaps. In this concept, the subfamily

Ologamasinae included Antennolaelaps Womersley, 1956a, Epiphis Berlese, 1916,

Gamasiphis Berlese, 1904, Gamasiphoides Womersley, 1956b, Gamasitus Womersley,

1956b, Hydrogamasus Berlese, 1892, Laelaptiella Womersley, 1956b, Megaliphis Willmann,

1938, Micriphis Berlese, 1914, Neogamasiphis Trägårdh, 1952, Ologamasus Berlese, 1888,

Onchogamasus Womersley, 1956a, Pachyseius Berlese, 1910, Parasitiphis Womersley,

1956b, Periphis Berlese, 1914, Physallolaelaps Berlese, 1908, Queenslandolaelaps

Womersley, 1956a, Sessiluncus Canestrini, 1898 and Trachygamasus Berlese, 1906. Ryke

(1962a) classified Rhodacarellus and Rhodacaropsis as sub-genera of Rhodacarus, and

Gamasellus Berlese, 1892, Digamasellus Berlese, 1905, Euryparasitus and Gamaselliphis as

sub-genera of Cyrtolaelaps. This very inclusive concept of Rhodacaridae included many

genera that are now placed in other families.

Evans (1963) analysed the leg chaetotaxy of the free-living Gamasina, including the

Rhodacaridae. In the initial part of that publication, he included the following genera in this

family: Asca, Cyrtolaelaps, Digamasellus, Euryparasitus, Gamasellus, Gamasiphis,

Halolaelaps, Hydrogamasus, Ologamasus, Rhodacarellus, Rhodacarus and Sessiluncus.

However, after reviewing the leg chaetotaxy of these and other genera, he considered that

Asca, Halolaelaps and Digamasellus probably did not belong to the same group as the other

genera, which he called the Rhodacarus-group. The latter was characterised as having the

following leg chaetotaxy: coxa 2, 2, 2, 1; trochanter 6, 5, 5, 5; femur 13, 11, 6, 6; genu 13, 11,

9, 10; and tibia 14, 10, 8, 10. Evans (1963) followed Ryke (1962a) in considering

Rhodacaropsis a subgenus of Rhodacarus, but treated Rhodacarellus as a genus.

Karg (1965) used a more restricted concept of this family. He considered the

Rhodacaridae as mites having scleronoduli on the podonotal shield and males with a hook-

shaped spermatodactyl, a group that included Dendrolaelaps Halbert, 1915, Protogamasellus

Karg, 1962, Rhodacarus and Rhodacarellus. He described Dendroseius Karg, 1965 as

subgenus of Dendrolaelaps; this subgenus was raised to genus level by Hirschmann (1974).

Between 1965 and 1968, several new genera of Rhodacaridae were described:

Notogamasellus Loots and Ryke, 1965, with sub-genera Notogamasellus (Notogamasellus)

Loots and Ryke, 1965 and Notogamasellus (Podonotogamasellus) Loots and Ryke, 1965,

Gamasellopsis Loots and Ryke, 1966, Gamasellevans Loots and Ryke, 1967,

Neogamasellevans Loots and Ryke, 1967, Afrogamasellus Loots and Ryke, 1968 and

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Paragamasellevans Loots and Ryke, 1968. Emberson (1968) reviewed the history of the

family, using a definition that included some genera that are now placed in other families.

Lee (1970) characterised the Rhodacaridae largely based on the type of leg

chaetotaxy used by Evans (1963) to define his Rhodacarus-group. In addition, he

characterised the family as including males and females with separate podonotal and

opisthonotal shields and the palp tarsal claw usually three-tined, never with associated hyaline

flap, and if two-tined, then with four ventral setae on tibia I; female with posteriorly truncate

genital shield separated from a conspicuous ventrianal shield; seta st4 usually on the sternal

shield (cited as sterno-metasternal shield); and male with distally free spermatodactyl;

presternal genital orifice; and seta av of femur II larger than that of the female and usually

considerably modified as a conspicuous spur. Lee (1970) considered that in this sense the

Rhodacaridae corresponded approximately to the Gamasellini of Hirschmann (1962),

Rhodacaridae plus Cyrtolaelapidae Berlese of Johnston (1968), Rhodacaridae (in part,

excluding Digamasellidae Evans) and Gamasellinae of Karg (1965) and Rhodacaridae (in

part, excluding some Ascidae Voigts and Oudemans, the Digamasellidae and the

Halolaelapidae Karg) of Ryke (1962a). Lee (1970) divided the family into six subfamilies:

Gamasiphinae, containing Euepicrius Womersley, 1942, Gamaselliphis, Gamasiphis,

Gamasiphoides, Hydrogamasus, Laelaptiella and the new genus Caliphis Lee, 1970;

Laelaptonyssinae, containing Laelaptonyssus Womersley, 1956b; Ologamasinae, containing

Allogamasellus, Cyrtolaelaps, Euryparasitus, Evanssellus, Gamasellus, Heterogamasus

Trägårdh, 1907, Heydeniella Richters, 1907, Hydrogamasellus Hirschmann, 1966,

Laelogamasus Berlese, 1905, Neogamasellevans, Notogamasellus, Ologamasus, Parasitiphis,

Periseius Womersley, 1961, Rhodacaroides, and the new genera Acugamasus Lee, 1970,

Cymiphis Lee, 1970, Geogamasus Lee, 1970, Hiniphis Lee, 1970, Litogamasus Lee, 1970,

Pilellus Lee, 1970, Pyriphis Lee, 1970, and Rykellus Lee, 1970; Rhodacarinae, containing

Afrogamasellus, Rhodacarellus, Rhodacaropsis and Rhodacarus; Sessiluncinae, containing

Antennolaelaps, Gamasellevans, Gamasellopsis, Gamasitus, Onchogamasus,

Paragamasellevans, Queenslandolaelaps, Sessiluncus and Stylochirus Canestrini and

Canestrini, 1882; and Tangaroellinae, containing Tangaroellus Luxton, 1968.

In the 1970s, the following new genera of Rhodacaridae were described: Athiasella

Lee, 1973a, Solugamasus Lee, 1973a, Afrodacarellus Hurlbutt, 1974, Mediorhodacarus

Shcherbak, 1976 and Orientolaelaps Bregetova and Shcherbak, 1977.

Krantz (1978) characterised the Rhodacaridae as having a three-tined palp tarsal

claw; podonotal shield not fused to opisthonotal shield; adults usually with distinctive

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scleronoduli between setae j5 and j6; anterior portion of sternal shield weakly defined, usually

carrying seta st1; ten setae each on genu and tibia IV; adult male with spermatodactyl often

recurved basally and with seta st5 on sternogenital shield (cited as sterno-metasterno-genital

shield). This concept of Rhodacaridae, corresponded to the strict sense of Johnston (1968) and

to the Rhodacarinae of Lee (1970). Actually, Johnston (1968) did not discuss the

characteristics and the constitution of the group, but just provided a re-drawn illustration of

details of the female and the male of R. roseus, the type species of Rhodacarus. Krantz (1978)

also characterised the Rhodacaroidea for the first time, including in this superfamily the

families Rhodacaridae, Digamasellidae and Ologamasidae Ryke. The superfamily was

considered to consist of mites with undivided sternal shield that generally bears four pairs of

setae, and the genital shield usually rounded anteriorly and not fused with the ventrianal

shield. It was considered that in some Rhodacaroidea seta st4 could be inserted in the

unsclerotised integument and seta st1 could be inserted on a weakly defined anteromarginal

extension of the sternal shield.

Shcherbak (1980) characterised the Rhodacaridae as a family whose adults and

deutonymphs had the dorsal idiosoma covered by two shields of subequal sizes; podonotal

shield with three or four scleronoduli, which in deutonymphs were sometimes difficult to

discern; female with undivided sternal shield (referred to as fused sternal and metasternal

shields) bearing four pairs of setae; females and males with seven pairs of setae on the venter

of the opisthosoma in addition to the three circumanal setae; epistome usually with three

processes of equal or different lengths; males with spermatodactyl S-shaped, basally fused to

movable cheliceral digit and apically free and with femur II bearing a spine-like seta.

Apparently referring to both sexes, she mentioned that Rhodacaridae have the hypostomal

setae simple in all developmental stages, and a deutosternal groove with six to eight

transverse rows of denticles. She divided Rhodacaridae into three subfamilies: Rhodacarinae,

containing Mediorhodacarus, Rhodacaropsis and Rhodacarus; Rhodacarellinae, containing

Rhodacarellus and the new genus Minirhodacarellus Shcherbak, 1980; and Dendrolaelapinae,

containing Dendrolaelaps, Dendrolaelaspis Lindquist, 1975, Dendroseius Karg, 1965,

Longoseius, Multidendrolaelaps Hirschmann, 1974, Orientolaelaps, and two new genera,

Oligodentatus Shcherbak, 1980 and Insectolaelaps Shcherbak, 1980.

Shcherbak (1980) also described Multidentorhodacarus as a subgenus of

Rhodacarus, and this subgenus was raised to genus level by Karg (2000b). However,

Shcherbak (1980) did not designate a type species for Multidentorhodacarus, so the name is

not available from that date (International Code of Zoological Nomenclature, Article 13.3).

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Karg (2000b) was the first to both publish a diagnosis of this genus and specify its type

species, so the authorship of this name becomes Multidentorhodacarus Karg, 2000b.

Shcherbak (1983) also described the new genus Dendrolobatus Shcherbak, 1983 in

Rhodacaridae.

Evans and Purvis (1987) described the genus Protogamasellopsis Evans and Purvis,

1987 in the family Ascidae. This genus was transferred to Rhodacaridae by Karg (1994a).

Further valuable discussions about this genus were presented by Karg (1994b) and Halliday;

Walter and Lindquist (1998), and it is here considered to belong to the Rhodacaridae. Six

other new genera have been described since then: Pararhodacarus Jordaan, Loots and

Theron, 1988; Pachymasiphis Karg, 1996, Interrhodeus Karg, 2000, Pennarhodeus Karg,

2000, Poropodalius Karg, 2000 and Binodacarus Castilho and Moraes, 2010. The original

descriptions of the genera Interrhodeus, Pennarhodeus and Poropodalius were brief and

lacked some important details, but recent re-examination of these genera has confirmed that

they are correctly placed in the Rhodacaridae (CASTILHO; MORAES; HALLIDAY, 2012).

Lindquist; Krantz and Walter (2009) provided a revised classification of the

Mesostigmata, with an expanded concept of the Rhodacaroidea that included the

Digamasellidae, Halolaelapidae, Laelaptonyssidae Womersley, Ologamasidae, Rhodacaridae

and Teranyssidae Halliday. Dowling and O‘Connor (2010) proposed an alternative hypothesis

in which the Rhodacaroidea as understood here is paraphyletic. However, until formal

taxonomic changes are made, we have used the classification of Lindquist; Krantz and Walter

(2009).

The purpose of this work is to review the literature concerning the taxonomy of

Rhodacaridae, to present a new characterisation of the family and the included genera, to

construct keys for identification of genera, as well as to update and complement the a list of

the species considered to belong to the Rhodacaridae, with relevant taxonomic information

about each. We also attempt a summary of the genus-level classification of the

Rhodacaroidea, to establish the family placement for every genus in the superfamily.

2.2 Materials and Methods

This publication includes papers published up to June 2011. The search for

information was initiated by considering the literature available in the personal reprint

collections of each of the authors. New references were detected by a search of electronic

databases and by the evaluation of references listed in each paper available to the authors. Of

26

fundamental importance in the initial part of the work were the publication of Lee (1970) and

the Rhodacaridae database of Hallan (2005); the latter was consulted periodically between

March 2006 and June 2011.

After obtaining copies of all the literature cited in this publication, we constructed a

spreadsheet cross referencing each of the species considered to belong to Rhodacaridae, with

characteristics mentioned in the respective descriptions or redescriptions. We examined the

available types of Interrhodeus, Multidentorhodacarus, Pennarhodeus, Poropodalius and

Protogamasellopsis to complement the information available in the literature. The types of

Protogamasellopsis dioscorus (Manson) and Protogamasellopsis posnaniensis Wiśniewski

and Hirschmann were not examined given their sufficiently detailed descriptions for the

purpose of this work. Despite our efforts, we were unable to examine the types of

Multidentorhodacarus denticulatus (Berlese), Multidentorhodacarus ruwenzoriensis (Loots),

Multidentorhodacarus sogdianus (Shcherbak) and Multidentorhodacarus thysi (Jordaan,

Loots and Theron). Missing information was added as much as possible to build a standard set

of characters that was used to characterise the whole family and the included genera. This

analysis made it necessary to move some species from one genus to another. The final

spreadsheet was then used to prepare a dichotomous key to the genera.

During our study of the Rhodacaridae, we also collected information about the

supraspecific taxa of other families of Rhodacaroidea. Thus, in this document we first present

diagnostic characteristics of each famly of the superfamily and a checklist of the genera and

sub-genera of the respective families. We then present a thorough diagnosis of the

Rhodacaridae and of each genus within this family, and a key to families of Rhodacaroidea

and genera of Rhodacaridae. The catalogue proper then includes a list of species of

Rhodacaridae, presented in alphabetical order of genera and of species within each genus. We

have not used intermediate categories such as subfamily, subgenus, or species groups, because

some species are not described in enough detail to allow these decisions to be done. For each

genus, the following information is provided:

Name and author;

Original designation of the genus (even if described at subgeneric level), author, year

of the original description, page on which the description begins, family in which the

genus was initially placed, type species, and further references to descriptions or re-

descriptions of the genus;

Synonyms, each followed by its author, page on which the corresponding original

description started, family in which the genus was initially placed, type species,

27

reference to the paper in which each corresponding synonymy was established, type

species, and further references providing descriptive information about the junior

synonym.

For each species, the following information is provided:

Current generic combination of the species, with its author and date of description;

Name of the species in its original combination, with reference to the author, date, and

the page on which the description started;

References to subsequent literature on the species, including different combinations or

variations of the name, and including publications that provide information on the

morphology of the species, other than the original description;

Synonyms, each followed by its author, date, and page number, and the reference in

which the synonymy was established;

Type depository, the institution where the name-bearing type specimens are deposited.

The types of most species described by Hurlbutt (1974) were located in the US

National Museum, Beltsville, Maryland, but some have not been found;

Type locality (first mentioning the country – to be more geographically informative,

Saint Helena and Galapagos Islands were mentioned as such, without referring to the

country to which they belong – followed by the location within the country, from the

more specific to the more general geographic information); and habitat in which the

type specimens were collected. In some cases we have added complementary locality

information, in square brackets.

Occasionally a note is added after the details of a genus or species, to explain an

unusual or complicated taxonomic or nomenclatural problem.

The terminology used for anatomical structures is that of Evans and Till (1979). Lee

(1970), Shcherbak (1982) and Hirschmann and Wiśniewski (1983) used different systems of

notation for the dorsal idiosomal chaetotaxy of Rhodacaroidea, but we have used the widely

accepted system developed by Lindquist and Evans (1965) and Lindquist (1994).

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2.3 Results

2.3.1 World genera and sub-genera of Rhodacaroidea

The Rhodacaridae and related families have had a very complex taxonomic history. In

order to provide a satisfactory summary of the genera of Rhodacaridae, it was necessary to

account for the placement of genera in all related families. We therefore present here a list of

the genera and sub-genera of Rhodacaroidea. We have tried to account for every genus-group

name that has been applied to taxa in the superfamily. The classification used here was

compiled from information in Lee (1970), Shcherbak (1980), Antony (1986), Silva (2007),

Halliday (2008a, 2008b), Lindquist; Krantz and Walter (2009) and other sources as cited.

Karg (1977) divided Afrogamasellus into five sub-genera. These are included in the checklist,

but they are not used in the list of species, because it is not possible to unambiguously assign

every species in the genus to one of the sub-genera. Antony (1986) introduced the new genus

name Mediodacarellus, but that name is here considered to be unavailable for nomenclatural

purposes.

Many of the genera listed here in other families have been placed in the

Rhodacaridae by previous authors, but are here excluded for various reasons. The genera that

are here placed in the Ologamasidae have a three-tined palp tarsal claw, seta st4 on sternal

shield, and genu and tibia IV with 9-10 setae each, but they are distinguished from

Rhodacaridae by lacking scleronoduli (present in some Gamasellus and some Rhodacaroides)

and desclerotised punctate bands on the dorsal and ventral shields. As in the Rhodacaridae,

genera here placed in the Digamasellidae also have separate podonotal and opisthonotal

shields (notal shield entire in Lindquistoseius Genis, Loots and Ryke, 1969 and Panteniphis

Willmann, 1949), seta st4 on sternal shield, and scleronoduli usually present (absent in

Digamasellus, Lindquistoseius, Panteniphis and Pontiolaelaps Luxton, 1984), but they have

two-tined palp tarsal claw, lack desclerotised punctate bands on the dorsal and ventral shields,

and each of their genu and tibia IV has only 6-8 setae. The genera that are here placed in the

Halolaelapidae have a three-tined palp tarsal claw, separate podonotal and opisthonotal

shields, and genu IV with 9-10 setae, but they have seta st4 on metasternal shield or on the

soft integument posterior to sternal shield, lack scleronoduli and desclerotised punctate bands

on the dorsal and ventral shields, and their tibia IV has 8-10 setae. Laelaptonyssus (and its

senior synonym Starkovia Lombardini, 1947) is considered to belong to Laelaptonyssidae, as

mentioned by Womersley (1956b) when he described the genus. Despite having a three-tined

29

palp tarsal claw and seta st4 on sternal shield, species in this genus do not have completely

separated podonotal and opisthonotal shields, lack scleronoduli and desclerotised and punctate

bands on the dorsal and ventral shields, and each of their genu and tibia IV has 7-10 setae.

Teranyssus Halliday, 2006 is considered to belong to Teranyssidae, as mentioned by Halliday

(2006) when he described the genus. Despite having seta st4 on sternal shield, species in this

genus have a two-tined palp tarsal claw, entire dorsal shield, lack scleronoduli and

desclerotised punctate bands on the dorsal and ventral shields, and their genu and tibia IV

have 11 and 10 setae, respectively.

Antennoseius, Asca, Protogamasellus [including Protogamasellus

(Protogamasellodes) Evans and Purvis, 1987], Pachyseius and Trachygamasus are excluded

from the Rhodacaroidea, either because they have two-tined palp tarsal claw, lack

scleronoduli (present in some Protogamasellus) and desclerotised punctate bands on dorsal

and ventral shields, do not have completely separate podonotal and opisthonotal shields, and

mainly because have seta st4 on metasternal shield or on the soft integument posterior to

sternal shield.

Tangaroellus was provisionally placed in the Rhodacaridae by Luxton (1968), Lee

(1970) and Lindquist; Krantz and Walter (2009). However, it is here considered incertae

sedis, following Halliday; Walter and Lindquist (1998). Despite having separate podonotal

and opisthonotal shields, and genu and tibia IV each with ten setae, the single species in this

genus has a two-tined palp tarsal claw, lacks scleronoduli and desclerotised punctate bands on

the dorsal and ventral shields, has a fully sclerotised sternal shield with only three pairs of

setae, and seta st4 on soft integument posterior to sternal shield. It therefore does not fit the

concept of Rhodacaridae as understood here.

Instabilty in the concept of the family Rhodacaridae means that some species that

were originally placed in this family are now placed elsewhere. Examples are Rhodacaropsis

angustiventris Athias-Henriot 1961, Rhodacaropsis cognatus Athias-Henriot 1961,

Rhodacaropsis massula Athias-Henriot 1961 [transferred to Protogamasellus (Ascidae) by

Lindquist and Evans (1965)], and Rhodacarus costai Sheals 1962 [transferred to

Rhodacaroides (Ologamasidae) by Lee (1970)].

In the families Digamasellidae and Ologamasidae, our arrangement of taxa does not

imply any taxonomic statements about their validity or lack of it, and is not the result of

taxonomic research on our part. We list these names only as they appear in the literature, as a

point of reference for future taxonomic revisions.

30

Rhodacaroidea Oudemans, 1902

Digamasellidae Evans, 1957

Dendrolaelaps Halbert, 1915

Dendrolaelaps (Apophyseodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Cornodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Daeleidendrolaelaps) Wiśniewski and Hirschmann, 1990

Dendrolaelaps (Dendrolaelaps) Halbert, 1915

Dendrolaelaps (Disetodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Duplodendrolaelaps) Wiśniewski and Hirschmann, 1991b

Dendrolaelaps (Epistodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Foveodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Ipidodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Majestidendrolaelaps) Wiśniewski and Hirschmann, 1989a

Dendrolaelaps (Monodendrolaelaps) Wiśniewski and Hirschmann, 1989b

Dendrolaelaps (Presepodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Sellnickidendrolaelaps) Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Stanidendrolaelaps) Wiśniewski and Hirschmann, 1993a

Dendrolaelaps (Xylodendrolaelaps) Wiśniewski and Hirschmann, 1993b

Dendrolaelaspis Lindquist, 1975

Dendrolobatus Shcherbak, 1983

Dendroseius Karg, 1965

Digamasellus Berlese, 1905a

= Dendrolaelaps (Tridendrolaelaps) Hirschmann, 1974

Insectolaelaps Shcherbak, 1980

Lindquistoseius Genis, Loots and Ryke, 1969

Longoseius Chant, 1961

Longoseius (Longoseius) Chant, 1961

Longoseius (Longoseiulus) Lindquist, 1975

Multidendrolaelaps Hirschmann, 1974

Oligodentatus Shcherbak, 1980

Orientolaelaps Bregetova and Shcherbak, 1977a

31

Panteniphis Willmann, 1949

Pontiolaelaps Luxton, 1984

Halolaelapidae Karg, 1965

Halodarcia Karg, 1969

Halolaelaps Berlese and Trouessart, 1889

= Saintdidieria Oudemans, 1939b

= Saprolaelaps Leitner, 1946

= Saprogamasellus Willmann, 1957

= "Halogamasellus" Błaszak and Ehrnsberger, 1995 (unavailable name)

= Haloseius Błaszak and Ehrnsberger, 1998

Leitneria Evans, 1957

Leitneria (Leitneria) Evans, 1957

Leitneria (Saproseius) Karg, 1965

Saprosecans Karg, 1964

Laelaptonyssidae Womersley, 1956b

Starkovia Lombardini, 1947

= Laelaptonyssus Womersley, 1956b

= Puchihlungia Samšiňák, 1964

Ologamasidae Ryke, 1962

Acugamasus Lee, 1970

Acuphis Karg, 1998

Allogamasellus Athias-Henriot, 1961

Antennolaelaps Womersley, 1956a

= Stylogamasus Womersley, 1956a

Athiasella Lee, 1973a

Caliphis Lee, 1970

Cymiphis Lee, 1970

Cyrtolaelaps Berlese, 1887

32

= Protolaelaps Trägårdh, 1912

Desectophis Karg, 2003a

Euepicrius Womersley, 1942

Euryparasitus Oudemans, 1902

= Eurylaelaps Oudemans, 1902 (lapsus)

Evanssellus Ryke, 1961a

Gamasellevans Loots and Ryke, 1967a

Gamaselliphis Ryke, 1961b

Gamasellopsis Loots and Ryke, 1966

Gamasellus Berlese, 1892a

Gamasiphis Berlese, 1904

= Micriphis Berlse, 1914

= Heteroiphis Trägårdh, 1952

= Neogamasiphis Trägårdh, 1952

Gamasiphoides Womersley, 1956b

Gamasitus Womersley, 1956b

Geogamasus Lee, 1970

Heterogamasus Trägårdh, 1907

Heydeniella Richters, 1907

Hiniphis Lee, 1970

Hydrogamasellus Hirschmann, 1966

Hydrogamasus Berlese, 1892b

Laelaptiella Womersley, 1956b

Laelogamasus Berlese, 1905b

Litogamasus Lee, 1970

Neogamasellevans Loots and Ryke, 1967b

Notogamasellus Loots and Ryke, 1965

Ologamasus Berlese, 1888

= Hologamasus Berlese, 1892a (incorrect subsequent spelling)

= Ologamasellus Berlese, 1914

Onchogamasus Womersley, 1956a

Oriflammella Halliday, 2008a

Parasitiphis Womersley, 1956b

= Austrohydrogamasus Hirschmann, 1966

33

Periseius Womersley, 1961

Periseius (Periseius) Womersley, 1961

Periseius (Psammonsella) Haq, 1965

Pilellus Lee, 1970

Podonotogamasellus Loots and Ryke, 1965

Pyriphis Lee, 1970

Queenslandolaelaps Womersley, 1956a

Rhodacaroides Willmann, 1959

Rhodacaroides (Rhodacaroides) Willmann, 1959

Rhodacaroides (Nodacaroides) Karg, 1977

Rhodacaroides (Tenacaroides) Karg, 1977

Rykellus Lee, 1970

Sessiluncus G. Canestrini, 1898

Solugamasus Lee, 1973a

Stylochirus G. Canestrini and R. Canestrini, 1882 (justified emendation following Berlese,

1882)

= Stilochirus G. Canestrini and R. Canestrini, 1882

= Iphidosoma Berlese, 1892c

= Physallolaelaps Berlese, 1908

= Gamasiphis (Periphis) Berlese, 1914

= Gamasiphis (Epiphis) Berlese, 1916b

= Gamasiphis (Megaliphis) Willmann, 1938

Rhodacaridae Oudemans, 1902

Afrodacarellus Hurlbutt, 1974

Afrogamasellus (Foliogamasellus) Karg, 1977

Afrogamasellus (Latogamasellus) Karg, 1977

Afrogamasellus Loots and Ryke, 1968

Afrogamasellus (Afrogamasellus) Loots and Ryke, 1968

= Afrogamasellus (Jugulogamasellus) Karg, 1977

Afrogamasellus (Podalogamasellus) Karg, 1977

Binodacarus Castilho and Moraes, 2010

Interrhodeus Karg, 2000

34

Mediorhodacarus Shcherbak, 1976

Minirhodacarellus Shcherbak, 1980

Multidentorhodacarus Karg, 2000

Paragamasellevans Loots and Ryke, 1968

= Paragamasellus Loots and Ryke, 1968 (lapsus)

Pararhodacarus Jordaan, Loots and Theron, 1988

Pennarhodeus Karg, 2000

Poropodalius Karg, 2000

Protogamasellopsis Evans and Purvis, 1987

= Rhodacarella Moraza, 2004

Rhodacarellus Willmann, 1935

Rhodacaropsis Willmann, 1935

Rhodacarus Oudemans, 1902

Teranyssidae Halliday, 2006

Teranyssus Halliday, 2006

Unplaced

Tangaroellus Luxton, 1968

Nomina nuda

Pachymasiphis Karg, 1996

2.3.2 Definition of the family Rhodacaridae

Adult females

Movable digit of chelicera with 2-6 teeth, fixed digit with 3-15 teeth. Palp tarsal claw three-

tined. Epistome with anterior section either triangular; or with an anterocentral extension that

may be of about uniform width along its length, narrower or wider at the base, often flanked

by one or more pairs of anterolateral extensions, these extensions longer, as long as, or shorter

35

than anterocentral extension; margin serrate or smooth. Hypostome with seta h2 about in

transverse line with h3, or both arranged roughly in longitudinal line with seta h1 (no

reference is made to this characteristic in most descriptions of rhodacarid species, making it

impossible to refer to it in the characterisation of each genus).

Idiosoma elongate or oval. Podonotal shield not fused to opisthonotal shield, except

in Afrogamasellus luberoensis Loots, in which a groove is present at the line of fusion.

Podonotal shield smooth or ornamented; with or without punctate band along posterior

margin; with or without a transverse or V-shaped line between setae j4 and j5; with 14-23

pairs of setae; 0-4 scleronoduli present between setae j5 and j6 (in Poropodalius basisetae

Karg, between setae j6 and z6). Opisthonotal shield smooth or ornamented, with or without

punctate band along anterior margin and with 14-20 pairs of setae.

Dorsal idiosoma with 0-8 pairs of setae on soft integument along lateral margins of

podonotal shield, and 0-5 pairs of setae on soft integument along lateral margins of

opisthonotal shield (Protogamasellopsis has 6-10 pairs of setae on soft integument along

lateral margins of opisthonotal shield).

Ventral idiosoma with 0-4 pairs of presternal plates, some of which may be partially

subdivided. Sternal shield (also referred by other authors as sterno-metasternal shield) longer

than wide or as long as wide, with 3-4 pairs of setae; anterior margin distinct or not; when

indistinct, the region of the sternal shield anterior to the first pair of lyrifissures (iv1) is lightly

sclerotised and punctate; posterior margin straight, concave, convex or with spine-like central

projection; seta st1 on presternal plate or on either lightly sclerotised or well sclerotised

regions of sternal shield; seta st4 always inserted on sternal shield. Genital shield extending

posteriorly beyond hind margin of coxa IV; longer, as long as or shorter than length of its

posterior margin; the latter straight, concave or convex; with a pair of lateral setae.

Opisthogastric integument with or without lightly sclerotised plates between genital and

ventrianal shields. Ventral and anal shields fused to form a ventrianal shield. Ventrianal shield

longer than wide, as long as wide or wider than long, smooth or ornamented, with 1-8 pairs of

preanal setae in addition to paranal and postanal setae; anterior margin straight, concave or

convex. With 0-6 pairs of setae on soft integument around ventrianal shield. With 0-3 pairs of

rounded, elongate or triangular metapodal plates. Peritreme extending anteriorly to mid-level

of coxa II, except in Rhodacarellus liuzhiyingi Ma and Rhodacarellus yalujiangensis Ma, in

which the peritreme extends anteriorly to mid-level of coxa I (but these species are probably

not Rhodacaridae). Peritrematal shield distinct or not; anteriorly fused or not fused to

podonotal shield.

36

Leg I with or without pretarsus; legs II-IV with pretarsi similar to each other; seta pl4

of tarsus IV present or absent. Genu IV with ten setae (2-2/1,3/1-1) and tibia IV with ten setae

(2-1/1,3/1-2).

Adult males. Spermatodactyl often recurved basally. Seta st5 inserted on sternogenital shield.

2.3.3 Definitions of world genera of Rhodacaridae

Afrodacarellus Hurlbutt

Movable and fixed cheliceral digits with 2-5 and 4-6 teeth, respectively. Arthrodial process of

chelicera shaped like a long cylindrical brush. Epistome with an anterocentral extension that

may be of about uniform width along its length, narrower or wider at the base; usually flanked

by one or more pairs of anterolateral extensions longer, as long as or shorter than the

anterocentral extension [with anterior region triangular in A. euungulae (Karg)]. Idiosoma

elongate or oval. Podonotal shield smooth or ornamented, with or without punctate band

along posterior margin, with 21-23 pairs of setae, two pairs of which along anterior margin,

without transverse line between setae j4 and j5 and with four scleronoduli between setae j5

and j6. Soft integument along lateral margins of podonotal shield with 0-2 pairs of setae.

Opisthonotal shield smooth or ornamented, with or without punctate band along anterior

margin and with 18-20 pairs of setae. Soft integument along lateral margins of opisthonotal

shield with 0-1 pair of setae. Peritreme extending anteriorly to level between anterior margin

of coxa III and median region of coxa II. Peritrematal shield anteriorly fused or not fused to

podonotal shield. Presternal plates absent. Sternal shield longer than wide, with anterior

margin indistinct and posterior margin straight, concave or with central spine-like projection.

Genital shield longer than length of its posterior margin; the latter straight. Opisthogastric

integument without plates between genital and ventrianal shields. Ventrianal shield longer

than wide, smooth or ornamented, with 5-7 pairs of preanal setae; anterior margin straight,

slightly concave or slightly convex. Soft integument around ventrianal shield with 0-2 pairs of

setae. With 1-2 pairs of rounded or elongate metapodal plates; when with two pairs, these

separated or partially fused. Pretarsus I present. Seta pl4 of tarsus IV absent (present in A.

camaxiloensis).

Afrogamasellus Loots and Ryke

37


chelicera shaped like a long cylindrical brush. Epistome with anterior region triangular; or

with an anterocentral extension wider at the base, not flanked by anterolateral extensions

[with an anterocentral extension narrower at the base, flanked by a pair of anterolateral

extensions in A. congoensis (Ryke and Loots) and A. uviraensis (Ryke and Loots)]. Idiosoma

elongate or oval. Podonotal shield smooth or ornamented, with or without punctate band

along posterior margin, with 18-23 pairs of setae, two pairs of which along anterior margin,

without transverse line between setae j4 and j5 and with four scleronoduli between setae j5

and j6 (absent in A. luberoensis kalibuensis Loots). Soft integument along lateral margins of

podonotal shield with 0-1 pair of setae. Opisthonotal shield smooth or ornamented, with or

without punctate band along anterior margin and with 17-20 pairs of setae. Soft integument

along lateral margins of opisthonotal shield with 0-3 pairs of setae. Peritreme extending

anteriorly to level between anterior margin of coxa III and median region of coxa II.

Peritrematal shield anteriorly fused or not fused to podonotal shield. With 0-1 pair of

presternal plates. Sternal shield longer than wide, with anterior margin distinct and posterior

margin straight, concave or with central spine-like projection. Genital shield as long as or

shorter than length of its posterior margin; the latter straight. Opisthogastric integument

without plates between genital and ventrianal shields. Ventrianal shield longer than wide,

smooth or ornamented, with 5-7 pairs of preanal setae; anterior margin straight or concave.

Soft integument around ventrianal shield with 0-2 pairs of setae. With 1-2 pairs of rounded,

elongate or triangular metapodal plates; when with two pairs, these separated or partially

fused. Pretarsus I present. Seta pl4 of tarsus IV absent.

Binodacarus Castilho and Moraes

Movable and fixed cheliceral digits with three and four teeth, respectively. Arthrodial process

of chelicera in the form of a short coronet-like fringe. Epistome with anterior region

triangular. Idiosoma elongate. Podonotal shield smooth, with punctate band along posterior

margin, with 14 pairs of setae, two pairs of which along anterior margin, without transverse

line between setae j4 and j5 and with two scleronoduli between setae j5 and j6. Soft

integument along lateral margins of podonotal shield with eight pairs of setae. Opisthonotal

shield smooth, with punctate band along anterior margin and with 14 pairs of setae. Soft

integument along lateral margins of opisthonotal shield with five pairs of setae. Peritreme

greatly reduced, not reaching anterior margin of coxa IV. Peritrematal shield indistinct.

Presternal plates absent. Sternal shield longer than wide, with anterior margin indistinct and

38

posterior margin concave. Genital shield longer than length of its posterior margin; the latter

straight. Opisthogastric integument with two lightly sclerotised plates between genital and

ventrianal shields. Ventrianal shield longer than wide, mostly smooth, except for a punctate

anteromedian lobe and for few diagonal lateral striae; with five pairs of preanal setae. Soft

integument around ventrianal shield with two pairs of setae. With a pair of metapodal plates

subdivided into a small anterior fragment and a larger posterior fragment. Pretarsus I present.

Seta pl4 of tarsus IV absent.

Interrhodeus Karg

Movable and fixed cheliceral digits with four and five teeth, respectively. Arthrodial process

of chelicera in the form of a short coronet-like fringe. Epistome with a serrate anterocentral

extension about uniform width along its length, flanked by a pair of serrate anterolateral

extensions slightly longer than the anterocentral extension. Idiosoma elongate. Podonotal

shield ornamented, with punctate band along posterior margin, with 22 pairs of setae, two

pairs of which along anterior margin, without transverse line between setae j4 and j5 and

without distinct scleronoduli. Soft integument along lateral margins of podonotal shield

without setae. Opisthonotal shield ornamented, with punctate band along anterior margin and

with 20 pairs of setae. Soft integument along lateral margins of opisthonotal shield without

setae. Peritreme extending anteriorly to median level of coxa II. Peritrematal shield not fused

to podonotal shield. Presternal plates absent. Sternal shield longer than wide, with anterior

margin indistinct and posterior margin straight. Genital shield longer than length of its

posterior margin; the latter straight. Opisthogastric integument without plates between genital

and ventrianal shields. Ventrianal shield longer than wide, ornamented, with five pairs of

preanal setae; anterior margin straight. Soft integument around ventrianal shield with two

pairs of setae. With a pair of elongate metapodal plates. Pretarsus I present. Seta pl4 of tarsus

IV absent.

Mediorhodacarus Shcherbak

Movable and fixed cheliceral digits with four and nine teeth, respectively. Form of arthrodial

process of chelicera unknown. Epistome with an anterocentral extension wider at the base,

totally smooth or with distal half slightly serrate, and flanked by a pair of smooth or slightly

serrate anterolateral extensions shorter than the anterocentral extension. Idiosoma elongate.

Podonotal shield smooth, with punctate band along posterior margin, with 23 pairs of setae,

four pairs of which along anterior margin, with transverse line between setae j4 and j5 and

39

with four scleronoduli between setae j5 and j6. Soft integument along lateral margins of

podonotal shield without setae. Opisthonotal shield smooth, without punctate band along

anterior margin and with 16 pairs of setae. Soft integument along lateral margins of

opisthonotal shield with three pairs of setae. Peritreme extending anteriorly to level of anterior

margin of coxa III. Peritrematal shield indistinct. With two pairs of presternal plates. Sternal

shield longer than wide, with anterior margin indistinct and posterior margin convex. Genital

shield longer than length of its posterior margin; the latter convex. Opisthogastric integument

without plates between genital and ventrianal shields. Ventrianal shield longer than wide,

ornamented, with five pairs of preanal setae; anterior margin slightly convex. Soft integument

around ventrianal shield with two pairs of setae. With a pair of elongate metapodal plates.

Pretarsus I absent. Seta pl4 of tarsus IV absent.

Minirhodacarellus Shcherbak

Movable and fixed cheliceral digits with three and 4-5 teeth, respectively. Arthrodial process

of chelicera in the form of a short coronet-like fringe. Epistome with a smooth anterocentral

extension about uniform width along its length, flanked by a pair of serrate anterolateral

extensions shorter than the anterocentral extension. Idiosoma elongate. Podonotal shield

smooth, with punctate band along posterior margin, with 22 pairs of setae, three pairs of

which along anterior margin, with transverse line between setae j4 and j5 and with four

scleronoduli between setae j5 and j6. Soft integument along lateral margins of podonotal

shield with a pair of setae. Opisthonotal shield smooth, without punctate band along anterior

margin and with 15 pairs of setae. Soft integument along lateral margins of opisthonotal

shield with four pairs of setae. Peritreme extending anteriorly to level of posterior margin of

coxa II. Peritrematal shield indistinct. Presternal plates absent. Sternal shield longer than

wide, with anterior margin indistinct and posterior margin convex. Genital shield longer than

length of its posterior margin; the latter convex. Opisthogastric integument without plates

between genital and ventrianal shields. Ventrianal shield longer than wide, smooth, with five

pairs of preanal setae; anterior margin slightly concave. Soft integument around ventrianal

shield with two pairs of setae. Without metapodal plates. Pretarsus I present. Seta pl4 of tarsus

IV absent.

Multidentorhodacarus Karg

Movable and fixed cheliceral digits with 4-6 and 9-15 teeth, respectively. Arthrodial process

of chelicera in the form of a short coronet-like fringe. Epistome with an anterocentral

40

extension slightly wider at the base, smooth or with the distal half slightly serrate, and flanked

by at least a pair of smooth or serrate anterolateral extensions shorter than the anterocentral

extension. Idiosoma elongate or slightly oval. Podonotal shield smooth, with or without

punctate band along posterior margin, with 21-23 pairs of setae, four pairs of which along

anterior margin, with V-shaped line posterior of setae j4, z3 and s2 and with three


shield with 0-1 pair of setae. Opisthonotal shield smooth, with or without punctate band along

anterior margin and with 16-19 pairs of setae. Soft integument along lateral margins of

opisthonotal shield with 1-3 pairs of setae. Peritreme extending anteriorly to level between

median region of coxa III and posterior margin of coxa II. Peritrematal shield not fused to

podonotal shield. Presternal plates absent. Sternal shield longer than wide, with anterior

margin indistinct and posterior margin straight or with spine-like central projection. Genital

shield longer than length of its posterior margin; the latter straight or convex. Opisthogastric

integument without plates between genital and ventrianal shield. Ventrianal shield longer than

wide, smooth, with 4-5 pairs of preanal setae; anterior margin slightly convex, concave or

straight. Soft integument around ventrianal shield with 2-3 pairs of setae. With or without a

pair of elongate metapodal plates. Pretarsus I absent. Seta pl4 of tarsus IV absent.

Paragamasellevans Loots and Ryke

Movable and fixed cheliceral digits with three and five teeth. respectively. Arthrodial process


extension wider at the base; with one to several pairs of spines along its median region; not

flanked by anterolateral extensions. Idiosoma elongate. Podonotal shield ornamented, with

punctate band along posterior margin, with 21-22 pairs of setae, three pairs of which along

anterior margin, without transverse line between setae j4 and j5 and with four scleronoduli

between setae j5 and j6. Soft integument along lateral margins of podonotal shield without

setae. Opisthonotal shield ornamented, with punctate band along anterior margin and with 15-

20 pairs of setae. Soft integument along lateral margins of opisthonotal shield with 0-3 pairs

of setae. Peritreme extending anteriorly to level of posterior margin of coxa II. Peritrematal

shield anteriorly fused or not fused to podonotal shield. With a pair of presternal plates.

Sternal shield longer than wide, with anterior margin distinct and posterior margin concave.

Genital shield longer than length of its posterior margin; the latter straight. Opisthogastric

integument without plates between genital and ventrianal shields. Ventrianal shield longer

than wide, ornamented, with six pairs of preanal setae; anterior margin slightly concave. Soft

41

integument around ventrianal shield with a pair of setae. With or without a pair of elongate

metapodal plates. Pretarsus I present. Seta pl4 of tarsus IV present.

Pararhodacarus Jordaan, Loots and Theron

Movable and fixed cheliceral digits with three and five teeth, respectively. Arthrodial process

of chelicera shaped like a long cylindrical brush. Epistome with anterior region triangular.

Idiosoma elongate. Podonotal shield ornamented, without punctate band along posterior

margin, with 21 pairs of setae, three pairs of which along anterior margin, without transverse

line between setae j4 and j5 and with four scleronoduli between setae j5 and j6. Soft

integument along lateral margins of podonotal shield without setae. Opisthonotal shield

ornamented, without punctate band along anterior margin and with 15 pairs of setae. Soft

integument along lateral margins of opisthonotal shield with four pairs of setae. Peritreme

extending anteriorly to median level of coxa III. Peritrematal shield not fused to podonotal

shield. With a pair of presternal plates. Sternal shield longer than wide, with anterior margin

indistinct and posterior margin straight. Genital shield longer than length of its posterior

margin; the latter convex. Opisthogastric integument with three lightly sclerotised plates

between genital and ventrianal shields. Ventrianal shield longer than wide, ornamented, with

seven pairs of preanal setae; anterior margin straight. Soft integument around ventrianal shield

without setae. With two pairs of rounded and one pair of elongate metapodal plates. Pretarsus

I absent. Seta pl4 of tarsus IV present.

Pennarhodeus Karg


of chelicera in the form of a short coronet-like fringe or three-pointed. Epistome with a

smooth anterocentral extension that may be of about uniform width along its length or slightly

wider at the base, and flanked by a pair of smooth or serrate anterolateral extensions shorter

than the anterocentral extension. Idiosoma elongate. Podonotal shield ornamented, with

punctate band along posterior margin, with 22-23 pairs of setae, two pairs of which along

anterior margin, without transverse line between setae j4 and j5 and without distinct

scleronoduli. Soft integument along lateral margins of podonotal shield without setae.

Opisthonotal shield ornamented, with punctate band along anterior margin and with 15 pairs

of setae. Soft integument along lateral margins of opisthonotal shield with 4-5 pairs of setae.

Peritreme extending anteriorly to level between anterior margin of coxa III and median region

of coxa II. Peritrematal shield not fused to podonotal shield. Presternal plates absent. Sternal

42

shield longer than wide, with anterior margin indistinct and posterior margin straight or

slightly concave. Genital shield longer or shorter than length of its posterior margin; the latter

straight. Opisthogastric integument with or without lightly sclerotised plates between genital

and ventrianal shields. Ventrianal shield wider than long, ornamented, with five pairs of

preanal setae; anterior margin straight or slightly convex. Soft integument around ventrianal

shield with two pairs of setae. With a pair of rounded metapodal plates. Pretarsus I present.

Seta pl4 of tarsus IV absent.

Poropodalius Karg

Movable and fixed cheliceral digits with three and five teeth, respectively. Arthrodial process

of chelicera in the form of a short coronet-like fringe. Epistome with a smooth anterocentral

extension that may be of about uniform width along its length or wider at the base, and

flanked by a pair of smooth or serrate anterolateral extensions shorter than the anterocentral

extension. Idiosoma elongate. Podonotal shield ornamented, with punctate band along

posterior margin, with 21-22 pairs of setae, two pairs of which along anterior margin, without

transverse line between setae j4 and j5 and with four scleronoduli between setae j5 and j6

(between setae j6 and z6 in P. basisetae). Soft integument along lateral margins of podonotal

shield with 1-2 pairs of setae. Opisthonotal shield ornamented, with punctate band along

anterior margin and 15 pairs of setae. Soft integument along lateral margins of opisthonotal

shield with 4-5 pairs of setae. Peritreme extending anteriorly to level of median region of coxa

II. Peritrematal shield not fused to podonotal shield. Presternal plates absent. Sternal shield

longer than wide, with anterior margin indistinct and posterior margin straight or concave.

Genital shield longer or shorter than length of its posterior margin; the latter straight.

Opisthogastric integument with or without lightly sclerotised plates between genital and

ventrianal shields. Ventrianal shield wider than long, ornamented, with 5-6 pairs of preanal

setae; anterior margin concave or straight. Soft integument around ventrianal shield with two

pairs of setae. With a pair of variously shaped metapodal plates (without metapodal plates in

P. acutus Karg). Pretarsus I present. Seta pl4 of tarsus IV absent.

Protogamasellopsis Evans and Purvis

Movable and fixed cheliceral digits with two and 6-8 teeth, respectively. Arthrodial process of

chelicera in the form of a short coronet-like fringe. Epistome with anterior region acuminate.

Idiosoma elongate. Podonotal shield smooth, with or without punctate band along posterior

margin, with 16 pairs of setae, three pairs of which along anterior margin, without transverse

43

line between setae j4 and j5 and without distinct scleronoduli. Soft integument along lateral

margins of podonotal shield with 5-6 pairs of setae and an elongate plate running from the

level of z1 to the level of z4, bearing or not seta r3. Opisthonotal shield smooth, with or

without punctate band along anterior margin and with 15 pairs of setae. Soft integument along

lateral margins of opisthonotal shield with 6-10 pairs of setae. Peritreme extending anteriorly

to level of median region of coxa II. Peritrematal shield not fused to podonotal shield. With 0-

4 pairs of transverse series of presternal plates, each series consisting of 1-2 platelets. Sternal

shield longer than wide, with anterior margin indistinct and posterior margin concave. Genital

shield longer than length of its posterior margin; the latter straight. Opisthogastric integument

with two pairs of setae and with or without lightly sclerotised plates between genital and

ventrianal shields. Ventrianal shield longer than wide, smooth, with 1-2 pairs of preanal setae;

anterior margin convex. Soft integument around ventrianal shield with 3-4 pairs of setae. With

0-3 pairs of elongate or rounded metapodal plates. Pretarsus I present. Seta pl4 of tarsus IV

present.

Rhodacarellus Willmann


chelicera in the form of a short coronet-like fringe. Epistome with a smooth or serrate

anterocentral extension that may be of about uniform width along its length or wider at the

base, and usually flanked by at least a pair of smooth or serrate anterolateral extensions

longer, as long as or shorter than the anterocentral extension. Idiosoma elongate or oval.

Podonotal shield smooth or ornamented, with punctate band along posterior margin, with 16-

23 pairs of setae, 2-3 pairs of which along anterior margin, entire or with a lateral fissure at

level of setae z1, z2 and z3, without transverse line between setae j4 and j5 and with four


shield with 0-6 pairs of setae. Opisthonotal shield smooth or ornamented, with punctate band

along anterior margin and with 15-20 pairs of setae. Soft integument along lateral margins of

opisthonotal shield with 0-5 pairs of setae. Peritreme extending anteriorly to level between

anterior margin of coxa III and median region of coxa II [peritreme extending anteriorly to

median region of coxa I in R. liuzhiyingi and R. yalujiangensis (but these species are probably

not Rhodacaridae)]. Peritrematal shield anteriorly fused or not fused to podonotal shield.

Presternal plates absent. Sternal shield longer than wide, with anterior margin indistinct and

posterior margin concave. Genital shield longer than length of its posterior margin; the latter

straight or convex. Opisthogastric integument with or without lightly sclerotised plates

44

between genital and ventrianal shields. Ventrianal shield as long as wide, smooth or

ornamented, with 3-8 pairs of preanal setae; anterior margin straight or slightly concave. Soft

integument around ventrianal shield with 0-4 pairs of setae. With 1-2 pairs of elongate or

rounded metapodal plates; when with two pairs, these separated or partially fused. Pretarsus I

present. Seta pl4 of tarsus IV absent.

Rhodacaropsis Willmann


of chelicera in the form of a short coronet-like fringe. Epistome with a anterocentral extension

wider at the base, totally smooth or with distal half slightly serrate, and flanked by a pair of

smooth anterolateral extensions shorter than the anterocentral extension. Idiosoma elongate.

Podonotal shield smooth, with or without punctate band along posterior margin, with 21-22

pairs of setae, four pairs of which along anterior margin, with transverse line between setae j4

and j5 and with three scleronoduli between setae j5 and j6. Soft integument along lateral

margins of podonotal shield with 1-2 pairs of setae. Opisthonotal shield smooth, with or


along lateral margins of opisthonotal shield with 0-5 pairs of setae. Peritreme greatly reduced,

not reaching anterior margin of coxa IV. Peritrematal shield absent or restricted to a band

along peritreme. With two pairs of presternal plates. Sternal shield longer than wide, with

anterior margin distinct and posterior margin convex. Genital shield longer than length of its

posterior margin; the latter convex. Opisthogastric integument without plates between genital

and ventrianal shields. Ventrianal shield longer than wide, smooth, with 4-7 pairs of preanal

setae; anterior margin straight or convex. Soft integument around ventrianal shield with 0-3

pairs of setae. With a pair of rounded metapodal plates. Pretarsus I present. Seta pl4 of tarsus

IV absent.

Rhodacarus Oudemans



extension that may be of about uniform width along its length or wider at the base, totally

smooth or with distal half serrate, and flanked by at least a pair of smooth or serrate

anterolateral extensions shorter than the anterocentral extension (with anterior region

triangular in R. rhodacaropsis Ryke). Idiosoma elongate. Podonotal shield smooth, with or

without punctate band along posterior and lateral margins, with 18-23 pairs of setae, four

45

pairs of which along anterior margin, with or without V-shaped line posterior of setae j4, z3

and s2 and with three scleronoduli between setae j5 and j6. Soft integument along lateral

margins of podonotal shield with 0-4 pairs of setae. Opisthonotal shield smooth, with or


along lateral margins of opisthonotal shield with 0-6 pairs of setae. Peritreme extending

anteriorly to level between median region of coxa III and posterior margin of coxa II.

Peritrematal shield anteriorly fused or not fused to podonotal shield. Presternal plates absent

(with a pair of presternal plates in R. berrisfordi Loots and R. rhodacaropsis). Sternal shield

longer than wide, with anterior margin indistinct and posterior margin straight or with a

central spine-like projection. Genital shield longer than length of its posterior margin; the

latter straight or convex. Opisthogastric integument with or without lightly sclerotised plates

between genital and ventrianal shields. Ventrianal shield longer than wide, smooth, with 4-6

pairs of preanal setae; anterior margin straight, slightly or distinctly convex. Soft integument

around ventrianal shield with 1-3 pairs of setae. With 1-2 pairs of elongate or rounded

metapodal plates. Pretarsus I absent. Seta pl4 of tarsus IV absent.

2.3.4 Key to world families of Rhodacaroidea and genera of Rhodacaridae (adult

females)

1. Seta st4 on metasternal shield or on soft integument .......... Halolaelapidae [Fig. 2.1 A-B]

- Seta st4 on sternal shield [except for Oriflammella (Ologamasidae), which has st4 on

metasternal platelets, but has the dorsal shield setae strongly pilose and mounted on

long stalks] ........................................................................................................................ 2

2. With anal shield, without preanal setae.................................. Teranyssidae [Fig. 2.2 A-C]

- With ventrianal shield, with 1-9 pairs of preanal setae .................................................. ...3

3. With attenuate chelicera; palp with 4 segments (fused tibia and tarsus) ............................

......................................................................................... Laelaptonyssidae [Fig. 2.3 A-C]

- With normal chelicera, not attenuate; palp with 5 segments ......................................... ...4

4. Palp tarsal claw two-tined; genu and tibia IV with six or eight setae .................................

............................................................................................ Digamasellidae [Fig. 2.4 A-D]

- Palp tarsal claw three-tined; genu and tibia IV with nine or ten setae .............................. 5

46

st4

A B

Figure 2.1 - Halolaelaps posidonis Halliday [after Halliday, 2008] – A. Dorsal idiosoma; B. Ventral idiosoma

Figure 2.2 - Teranyssus howardensis Halliday – A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial)

view of chelicera

Figure 2.3 - Starkovia lacticolus (Halliday) – A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial)

view of chelicera

47

A B

C

D

Figure 2.4 - Digamasellus australis Lindquist [after Lindquist, 1975] – A. Dorsal idiosoma; B. Ventral idiosoma;

C. Lateral (antiaxial) view of chelicerae; D. Epistome

5. Usually without densely sclerotised strutuctures between setae j5 and j6 (scleronoduli);

podonotal and opisthonotal shields fused or not; without desclerotised bands of punctate

integument ............................................................................ Ologamasidae [Fig. 2.5 A-D]

- With scleronoduli (except in Interrhodeus, Protogamasellopsis and Pennarhodeus);

podonotal and opisthonotal shields not fused (except in Afrogamasellus luberoensis);

usually with desclerotised bands of punctate integument along posterior margin of

podonotal, anterior margin of opisthonotal, anterior margin of sternal, posterior margin

of genital and anterior margin of ventrianal shields ................................ Rhodacaridae...6

Figure 2.5 - Neogamasellevans ornata Karg [after Karg, 1975] – A. Dorsal idiosoma; B. Ventral idiosoma; C.

Lateral (antiaxial) view of chelicerae; D. Epistome

48

6. Peritreme greatly reduced, not reaching anterior margin of coxa IV ................................ 7

- Peritreme extending anteriorly at least to level of median region of coxa III ................... 8

7. Podonotal shield with 14 pairs of setae, without transverse line between setae j4 and j5

and with two scleronoduli..................... Binodacarus Castilho and Moraes [Fig. 2.6 A-D]

- Podonotal shield with 21-22 pairs of setae, with transverse line between setae j4 and j5

and with three scleronoduli................................ Rhodacaropsis Willmann [Fig. 2.7 A-D]

Figure 2.6 - Binodacarus brasiliensis Castilho and Moraes – A. Dorsal idiosoma; B. Ventral idiosoma; C.


Figure 2.7 - Rhodacaropsis cheungae Luxton [after Luxton, 1992] – A. Dorsal idiosoma; B. Ventral idiosoma;


49

8. Podonotal shield with 16 pairs of setae; ventrianal shield with one or two pairs of

preanal setae ................................... Protogamasellopsis Evans and Purvis [Fig. 2.8 A-D]

- Podonotal shield with 17 or more pairs of setae (Rhodacarellus arcanus with 16);

ventrianal shield with at least four pairs of preanal setae ................................................. 9

Figure 2.8 - Protogamasellopsis posnaniensis Wiśniewski and Hirschmann – A. Dorsal idiosoma; B. Ventral

idiosoma; C. Lateral (antiaxial) view of chelicerae; D. Epistome

9. Scleronoduli absent; arthrodial process of chelicera in the form of a short coronet-like

fringe or three-pointed..................................................................................................... 10

- Scleronoduli present or absent; arthrodial process of chelicera in the form of a short

coronet-like fringe or shaped like a long cylindrical brush) (if scleronoduli absent, in

Afrogamasellus luberoensis kalibuensis, arthrodial process shaped like a long

cylindrical brush) ............................................................................................................ 11

10. Podonotal and opisthonotal setae serrated .................. Pennarhodeus Karg [Fig. 2.9 A-D]

- Podonotal and opisthonotal setae smooth ................... Interrhodeus Karg [Fig. 2.10 A-D]

11. Podonotal shield with three scleronoduli; arthrodial process of chelicera in the form of a

short coronet-like fringe .................................................................................................. 12

- Podonotal shield with four scleronoduli; arthrodial process of chelicera in the form of a

short coronet-like fringe or shaped like a long cylindrical brush.................................... 13

50

Figure 2.9 - Interrhodeus brevicornus Karg – A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial)

view of chelicerae; D. Epistome

Figure 2.10 - Pennarhodeus decoris Karg – A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial) view

of chelicerae; D. Epistome

12. Fixed cheliceral digit with at least nine teeth……………………………………………..

. ......................................................................Multidentorhodacarus Karg [Fig. 2.11 A-D]

- Fixed cheliceral digit with at most six teeth ........ Rhodacarus Oudemans [Fig. 2.12 A-D]

51

Figure 2.11 - Multidentorhodacarus paulista Castilho and Moraes – A. Dorsal idiosoma; B. Ventral idiosoma; C.


Figure 2.12 - Rhodacarus matatlanticae Castilho and Moraes – A. Dorsal idiosoma; B. Ventral idiosoma; C.


13. With two pairs of presternal plates............. Mediorhodacarus Shcherbak [Fig. 2.13 A-D]

- With 0-1 pair of presternal plates .................................................................................... 14

14. Arthrodial process of chelicera shaped like a long cylindrical brush ............................. 15

- Arthrodial process of chelicera in the form of a short coronet-like fringe...................... 17

52

Figure 2.13 - Mediorhodacarus tetranodulosus Shcherbak [after Shcherbak, 1976] – A. Dorsal idiosoma; B.

Ventral idiosoma; C. Lateral (antiaxial) view of chelicerae; D. Epistome

15. With three pairs of metapodal plates; basitarsus IV with four setae (seta pl4 present) .......

............................................ Pararhodacarus Jordaan, Loots and Theron [Fig. 2.14 A-D]

- With 1-2 pairs of metapodal plates; basitarsus IV with three setae (seta pl4 absent,

except in Afrodacarellus camaxiloensis) ......................................................................... 16

Figure 2.14 - Pararhodacarus intermedius Jordaan, Loots and Theron [after Jordaan; Loots and Theron, 1988] –

A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial) view of chelicerae; D. Epistome

16. Epistome with anterior region triangular or with an anterocentral extension wider at the

base, not flanked by anterolateral extensions [in A. congoensis (Ryke and Loots) and A.

uviraensis (Ryke and Loots) anterior region of epistome with an anterocentral extension

53

narrower at the base, flanked by a pair of anterolateral extensions]; with 0-1 pair of

presternal plates; sternal shield with anterior margin distinct; genital shield as long as or

shorter than length of its posterior margin………………………………………………...

………………………………………..Afrogamasellus Loots and Ryke [Fig. 2.15 A-D]

- Epistome with an anterocentral extension that may be of about uniform width along its

length, narrower or wider at the base; usually flanked by one or more pairs of

anterolateral extensions [A. euungulae (Karg), with anterior region triangular]; without

presternal plates; sternal shield with anterior margin indistinct and with region anterior

to first pair of lyrifissures (iv1) lightly sclerotised and punctate; genital shield longer

than length of its posterior margin ..... …………Afrodacarellus Hurlbutt [Fig. 2.16 A-D]

Figure 2.15 - Afrogamasellus isthmus Hurlbutt [after Hurlbutt, 1974] – A. Dorsal idiosoma; B. Ventral idiosoma;


Figure 2.16 - Afrodacarellus femoratus Hurlbutt [after Hurlbutt, 1974] – A. Dorsal idiosoma; B. Ventral

idiosoma; C. Lateral (antiaxial) view of chelicerae; D. Epistome

54

17. Podonotal shield with a transverse line between setae j4 and j5 .........................................

................................................................... Minirhodacarellus Shcherbak [Fig. 2.17 A-D]

- Podonotal shield without transverse line between setae j4 and j5 ................................... 18

Figure 2.17 - Minirhodacarellus minimus (Karg) [after Karg, 1961] – A. Dorsal idiosoma; B. Ventral idiosoma;


18. Epistome with an anterocentral extension (with one to several pairs of spines along its

median region) wider at the base, not flanked by anterolateral extensions; basitarsus IV

with four setae (pl4 present) ............ Paragamasellevans Loots and Ryke [Fig. 2.18 A-D]

- Epistome with a smooth or serrate anterocentral extension that may be of about uniform

width along its length or wider at the base, usually flanked by at least a pair of smooth

or serrate anterolateral extensions; basitarsus IV with three setae (pl4 absent) .............. 19

Figure 2.18 - Paragamasellevans michaeli Loots and Ryke [after Loots and Ryke, 1968] – A. Dorsal idiosoma;

B. Ventral idiosoma; C. Lateral (antiaxial) view of chelicerae; D. Epistome

55

19. Podonotal shield ornamented; with a pair of roundish metapodal plates (indistinct or

absent em P. acutus Karg)…………………. ............ Poropodalius Karg [Fig. 2.19 A-D]

- Podonotal shield smooth (ornamented in R. unicus Karg); with a pair of elongate

metapodal plates (if second pair of metapodal plates are present, this are rounded and

small .................................................................. Rhodacarellus Willmann [Fig. 2.20 A-D]

Figure 2.19 - Poropodalius hexapennatus Karg – A. Dorsal idiosoma; B. Ventral idiosoma; C. Lateral (antiaxial)

view of chelicerae; D. Epistome

Figure 2.20 - Rhodacarellus epigynialis Sheals [after Sheals, 1956] – A. Dorsal idiosoma; B. Ventral idiosoma;


56

2.3.5 Catalogue of world species of Rhodacaridae

Genus Afrodacarellus Hurlbutt, 1974

Afrodacarellus Hurlbutt, 1974: 589 (described in Rhodacaridae Oudemans).

Type species: Afrodacarellus femoratus Hurlbutt, 1974, by original designation.

Afrogamasellus (Foliogamasellus) Karg, 1977: 345 (described in Rhodacaridae Oudemans)

Afrogamasellus (Foliogamasellus).— Karg, 2003b: 26; Karg and Schorlemmer, 2009: 65.

Type species: Afrogamasellus camaxiloensis Loots, 1969, by original designation.

Afrogamasellus (Latogamasellus) Karg, 1977: 345 (described in Rhodacaridae Oudemans).

Afrogamasellus (Latogamasellus).— Karg, 2003b: 26; Karg and Schorlemmer, 2009: 65.

Type species: Afrogamasellus squamosus Karg, 1977, by original designation.

01. Afrodacarellus bakeri (Hurlbutt, 1974)

Afrogamasellus bakeri Hurlbutt, 1974: 586.

Mediodacarellus bakeri.— Antony, 1986: 152.

TYPE DEPOSITORY: unknown.

TYPE LOCALITY AND HABITAT: Tanzania, track from Morningside to the top of Mount

Bondwa, Uluguru Mountains, [Morogoro], alt. 1450 m, 1 December 1967, in rotting

log at edge of rain forest.

NOTE: Mediodacarellus is not available for nomenclatural purposes, and is used here for

information only.

02. Afrodacarellus bipilosus (Karg, 1979)

Afrogamasellus bipilosus Karg, 1979: 207.

Afrogamasellus (Latogamasellus) bipilosus.— Karg, 1979: 207; Karg, 2003b: 27.

Latogamasellus bipilosus.— Antony, 1986: 148.

TYPE DEPOSITORY: Ungarische Naturwissenschaftliches Museum, Budapest, Hungary.

TYPE LOCALITY AND HABITAT: Argentina, Mount Piltriquitron [= Cerro Piltriquitron],

El Bolsón, Rio Negro, 19 October 1961, in leaf litter.

03. Afrodacarellus camaxiloensis (Loots, 1969)

57

Afrogamasellus camaxiloensis Loots, 1969a: 60.

Afrogamasellus camaxiloensis.— Lee, 1970: 34; Loots, 1971: 3.

Afrodacarellus camaxiloensis.— Hurlbutt, 1974: 591; Antony, 1986: 151.

Afrogamasellus (Foliogamasellus) camaxiloensis.— Karg, 1977: 345; Karg and

Schorlemmer, 2009: 66.

TYPE DEPOSITORY: Museu do Dundo, Dundo, Angola.

TYPE LOCALITY AND HABITAT: Angola, Tshihumbwe River, branch of the Lubalo

River, Camaxilo, Lunda, 6 July 1962, in forest soil.

04. Afrodacarellus citri (Loots, 1969)

Afrogamasellus citri Loots, 1969a: 75.

Afrogamasellus citri.— Hurlbutt, 1974: 588; Castilho and Moraes, 2010: 396.

Afrogamasellus (Foliogamasellus) citri.— Karg, 1977: 344.


TYPE LOCALITY AND HABITAT: South Africa, Nelspruit, [Mpumalanga], 1965, in soil

under Citrus sp. [Rutaceae].

05. Afrodacarellus concavus Hurlbutt, 1974

Afrodacarellus concavus Hurlbutt, 1974: 596.

TYPE DEPOSITORY: United States National Museum of Natural History Acari Collection,

Beltsville, Maryland, USA.

TYPE LOCALITY AND HABITAT: Tanzania, summit of Mount Bondwa, Uluguru

Mountains, [Morogoro], alt. 2120 m, 1 May 1968, in humus and lichens under

Philippia [Ericaceae].

06. Afrodacarellus euungulae (Karg, 2003)

Afrogamasellus euungulae Karg, 2003a: 245.

TYPE DEPOSITORY: Arachnida und Myriapoda Sammlung (cited as Arachnologische

Sammlung), Museum für Naturkunde, Berlin, Germany.

TYPE LOCALITY AND HABITAT: Ecuador, Cardy [= Carchi?], 1989, in prairie soil.

07. Afrodacarellus femoratus Hurlbutt, 1974

Afrodacarellus femoratus Hurlbutt, 1974: 593.

58



TYPE LOCALITY AND HABITAT: Tanzania, in the upper part of a valley between the

Morogoro Regional Forestry Office and Mount Lupanga, Morogoro, alt. 1100 m, 30

August 1967, on leaves and twigs under clump of trees.

08. Afrodacarellus filofissus (Karg and Schorlemmer, 2009)

Afrogamasellus (Foliogamasellus) filofissus Karg and Schorlemmer, 2009: 65.



TYPE LOCALITY AND HABITAT: Venezuela, on the road from Caracas to La Guaira,

1973, in litter and soil.

09. Afrodacarellus furculatus (Karg, 1979)

Afrogamasellus furculatus Karg, 1979: 208.

Afrogamasellus (Latogamasellus) furculatus.— Karg, 1979: 207; Karg, 2003b: 27.


TYPE LOCALITY AND HABITAT: Argentina, Mount Piltriquitron [= Cerro Piltriquitron],

El Bolsón, Rio Negro, 16 May 1961, in soil with grass under a tree trunk.

10. Afrodacarellus kivuensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) kivuensis Ryke and Loots, 1966: 141.

Afrogamasellus kivuensis.— Loots and Ryke, 1968: 2; Lee, 1970: 33.

Afrodacarellus kivuensis.— Hurlbutt, 1974: 591.

Afrogamasellus (Foliogamasellus) kivuensis.— Karg, 1977: 344.

TYPE DEPOSITORY: Musée Royal de l‘Afrique Centrale, Tervuren, Belgium.

TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Mont Kabobo,

Kyimbi, Katanga, October 1958, in soil.

11. Afrodacarellus leleupi (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) leleupi Ryke and Loots, 1966: 139.

Afrogamasellus leleupi.— Loots and Ryke, 1968: 2; Lee, 1970: 33.

Afrodacarellus leleupi.— Hurlbutt, 1974: 591.

Afrogamasellus (Foliogamasellus) leleupi.— Karg, 1977: 344.

59


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Kivu Province, 27

December 1950, in soil.

12. Afrodacarellus longipodus Hurlbutt, 1974

Afrodacarellus longipodus Hurlbutt, 1974: 594.


TYPE LOCALITY AND HABITAT: Tanzania, hill south of Morogoro Agricultural College,

[Morogoro], alt. 900 m, 26 April 1968, in soil under patch of trees.

13. Afrodacarellus lubalensis (Loots, 1969)

Afrogamasellus lubalensis Loots, 1969a: 79.

Afrogamasellus (Latogamasellus) lubalensis.— Karg, 1979: 206; Karg, 2003b: 27.




14. Afrodacarellus lunguensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) lunguensis Ryke and Loots, 1966: 149.

Afrogamasellus lunguensis.— Loots and Ryke, 1968: 1; Lee, 1970: 33.

Afrodacarellus lunguensis.— Hurlbutt, 1974: 590.

Afrogamasellus (Foliogamasellus) lunguensis.— Karg, 1977: 344.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Lac Lungue, Kivu

Province, May 1958, in soil in bamboo forest [Poaceae].

15. Afrodacarellus lupangaensis Hurlbutt, 1974

Afrodacarellus lupangaensis Hurlbutt, 1974: 602.


TYPE LOCALITY AND HABITAT: Tanzania, base of Mount Lupanga, Uluguru Mountains,

Morogoro, alt. 1350 m, 30 October 1966, in leaf litter and humus from regenerated

rain forest.

16. Afrodacarellus machadoi (Loots, 1969)

60

Afrogamasellus machadoi Loots, 1969a: 65.

Afrodacarellus machadoi.— Hurlbutt, 1974: 598.

Afrogamasellus (Foliogamasellus) machadoi.— Karg, 1977: 344; Karg and Schorlemmer,

2009: 66.


TYPE LOCALITY AND HABITAT: Angola, by Tshihumbwe River, branch of the Lubalo


17. Afrodacarellus minutus Hurlbutt, 1974

Afrodacarellus minutus Hurlbutt, 1974: 609.




Morogoro Regional Forestry Office and Mount Lupanga, near Morogoro, alt. 1150

m, 22 May 1968, on leaves and twigs under clump of trees.

18. Afrodacarellus mongii (Hurlbutt, 1974)

Afrogamasellus mongii Hurlbutt, 1974: 574.

Afrogamasellus mongii.— Antony, 1986: 148.


TYPE LOCALITY AND HABITAT: Tanzania, near Marangu track, Mount Kilimanjaro, alt.

2100 m, 10 June 1972, in forest litter.

19. Afrodacarellus mossi Hurlbutt, 1974

Afrodacarellus mossi Hurlbutt, 1974: 606.



Mountains, alt. 2120 m, 10 June 1968, in moss on branch of tree.

20. Afrodacarellus msituni Hurlbutt, 1974

Afrodacarellus msituni Hurlbutt, 1974: 604.


61


Morogoro Regional Forestry Office and Mount Lupanga, [Morogoro], alt. 1100 m,

23 January 1968, on dead grass stems and soil near clump of trees.

21. Afrodacarellus myersi (Loots, 1969)

Afrogamasellus myersi Loots, 1969a: 77.

Afrogamasellus myersi.— Hurlbutt, 1974: 570.

Afrogamasellus (Latogamasellus) meyersi [sic].— Karg, 1979: 207.

Afrogamasellus (Latogamasellus) myersi.— Karg, 2003b: 27.


TYPE LOCALITY AND HABITAT: South Africa, Nelspruit, [Mpumalanga], 1965, in soil

under Citrus sp. [Rutaceae].

22. Afrodacarellus ngorongoroensis Hurlbutt, 1974

Afrodacarellus ngorongoroensis Hurlbutt, 1974: 597.


TYPE LOCALITY AND HABITAT: Tanzania, rim of Ngorongoro Crater, [Arusha Region],

alt. 2300 m, 3 January 1968, among twigs and bits of wood under trees.

23. Afrodacarellus novembus Hurlbutt, 1974

Afrodacarellus novembus Hurlbutt, 1974: 602.



TYPE LOCALITY AND HABITAT: Tanzania, near path through rain forest, track from

Morningside to the top of Mount Bondwa, Uluguru Mountains, [Morogoro], alt.

1500 m, 4 November 1967, in soil and on fern roots [Pteridophyta].

24. Afrodacarellus pili Hurlbutt, 1974

Afrodacarellus pili Hurlbutt, 1974: 600.




Mountains, [Morogoro], alt. 2120 m, 10 June 1968, on leaves and humus from elfin

forest.

62

25. Afrodacarellus pocsi Hurlbutt, 1974

Afrodacarellus pocsi Hurlbutt, 1974: 607.



Mountains, [Morogoro], alt. 2125 m, 30 May 1972, in moss on branch of tree.

26. Afrodacarellus reticulatus (Loots, 1969)

Afrogamasellus reticulatus Loots, 1969a: 68.

Afrodacarellus reticulatus.— Hurlbutt, 1974: 591.

Afrogamasellus (Foliogamasellus) reticulatus.— Karg, 1977: 344; Karg and Schorlemmer,

2009: 66.




27. Afrodacarellus ruwenzoriensis (Loots, 1969)

Afrogamasellus ruwenzoriensis Loots, 1969a: 71.

Afrodacarellus ruwenzoriensis.— Hurlbutt, 1974: 594.

Afrogamasellus (Foliogamasellus) ruwenzoriensis.— Karg, 1977: 344; Karg and


TYPE DEPOSITORY: Natural History Museum, London, England.

TYPE LOCALITY AND HABITAT: Uganda, near Nyinabitaba, Ruwenzori, alt. 8650 feet,

1952, in soil.

28. Afrodacarellus squamosus (Karg, 1977)

Afrogamasellus (Latogamasellus) squamosus Karg, 1977: 345.

Afrogamasellus (Latogamasellus) squamosus.— Karg, 1979: 207; Karg, 2003b: 27.

Latogamasellus squamosus.— Antony, 1986: 154.


TYPE LOCALITY AND HABITAT: Argentina, near Norquinco and El Bolsón, Rio Negro,

23 February 1961, in leaf litter of Mulinum spinosum [Apiaceae].

29. Afrodacarellus succinctus (Berlese, 1916)

63

Gamasellus succinctus Berlese, 1916a: 160.

Cyrtolaelaps (Gamasellus) succinctus.— Ryke, 1962b: 48; Ryke and Loots, 1966: 124.

Afrogamasellus succinctus.— Loots and Ryke, 1968: 2; Lee, 1970: 33; Hurlbutt, 1974: 571.

Afrogamasellus (Foliogamasellus) succinctus.— Karg, 1977: 344.

Gamasellus succinctus.— Castagnoli and Pegazzano, 1985: 404.

TYPE DEPOSITORY: Istituto Sperimentale per la Zoologia Agraria, Florence, Italy.

TYPE LOCALITY AND HABITAT: East Africa, on unspecified substrate.

30. Afrodacarellus unospinae (Karg, 2003)

Afrogamasellus unospinae Karg, 2003b: 27.

Afrogamasellus (Latogamasellus) unospinae.— Karg, 2003b: 27.

TYPE DEPOSITORY: Staatliches Museum für Naturkunde Görlitz, Görlitz, Germany.

TYPE LOCALITY AND HABITAT: Ecuador, Calderon, 1990, in litter under bamboo forest

[Poaceae].

Genus Afrogamasellus Loots and Ryke, 1968

Afrogamasellus Loots and Ryke, 1968: 2 (described in Rhodacaridae Oudemans).

Afrogamasellus.— Loots, 1969a: 60; Loots, 1969b: 359; Loots, 1969c: 182; Lee, 1970: 30;

Hurlbutt, 1974: 568; Karg, 1977: 343; Loots, 1979: 2; Karg, 2003b: 26; Karg and


Afrogamasellus (Afrogamasellus).— Karg and Schorlemmer, 2009: 65.

Type species: Cyrtolaelaps (Gamasellus) franzi Loots and Ryke, 1968, by original

designation.

Afrogamasellus (Podalogamasellus) Karg, 1977: 345 (described in Rhodacaridae Oudemans).

Afrogamasellus (Podalogamasellus).— Karg, 2003b: 26; Karg and Schorlemmer, 2009: 65.

Type species: Gamasellus tetrastigma Berlese, 1916a, by original designation.

Afrogamasellus (Jugulogamasellus) Karg, 1977: 345 (described in Rhodacaridae Oudemans)

[objective junior synonym of Afrogamasellus (Afrogamasellus)].

Type species: Cyrtolaelaps (Gamasellus) franzi Loots and Ryke, 1968, by original

designation.

64

NOTE: Karg (1977) designated as type of Afrogramasellus (Jugulogamasellus) the species

designated by Loots and Ryke (1968) as type of the Afrogamasellus, which makes that

subgenus an objective junior synonym of Afrogramasellus (Afrogamasellus).

31. Afrogamasellus celisi Loots, 1969

Afrogamasellus celisi Loots, 1969b: 367.

Afrogamasellus celisi.— Lee, 1970: 34; Hurlbutt, 1974: 570.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], near Lubero,

Kivu Province, 3 July 1957, in soil.

32. Afrogamasellus congoensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) uviraensis congoensis Ryke and Loots, 1966: 146.

Afrogamasellus uviraensis congoensis.— Loots and Ryke, 1968: 2 (implicit); Lee, 1970: 33.

Afrodacarellus congoensis.— Hurlbutt, 1974: 590.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], near Lake

Tanganyika, Uvira, Kivu Province, January 1960, in soil.

33. Afrogamasellus evansi Loots, 1969

Afrogamasellus evansi Loots, 1969b: 377.

Afrogamasellus evansi.— Lee, 1970: 34; Hurlbutt, 1974: 570.



1952, in soil.

34. Afrogamasellus franzi (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) franzi Ryke and Loots, 1966: 124.

Afrogamasellus franzi.— Loots and Ryke, 1968: 2; Lee, 1970: 33; Loots, 1971: 3; Hurlbutt,

1974: 571.

Afrogamasellus (Jugulogamasellus) franzi.— Karg, 1977: 345.

TYPE DEPOSITORY: Institute for Zoological Research of the Potchefstroom University,

Potchefstroom, South Africa.

65

TYPE LOCALITY AND HABITAT: Kenya, on western side of Mount Meru, alt. 2600 m, 9

July 1962, in moss and litter in a Hagenia forest [Rosaceae].

35. Afrogamasellus franzoides Hurlbutt, 1974

Afrogamasellus franzoides Hurlbutt, 1974: 573.


TYPE LOCALITY AND HABITAT: Tanzania, upper part of a valley between the Morogoro

Regional Forestry Office and Mount Lupanga, Morogoro, alt. 3600 feet, 12 June

1966, in soil under tree.

36. Afrogamasellus isthmus Hurlbutt, 1974

Afrogamasellus isthmus Hurlbutt, 1974: 576.



TYPE LOCALITY AND HABITAT: Tanzania, adjacent to Morogoro River, Morogoro, alt.

550 m, 11 May 1966, in duff under bushes and trees.

37. Afrogamasellus kahusiensis Loots, 1969

Afrogamasellus kahusiensis Loots, 1969b: 372.

Afrogamasellus kahusiensis.— Lee, 1970: 34; Hurlbutt, 1974: 570.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Kaleke, northeast

of Kahusi, Kivu Province, 29 June 1951, in soil.

38. Afrogamasellus kilimanjaroensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) kilimanjaroensis Ryke and Loots, 1966: 133.

Afrogamasellus kilimanjaroensis.— Loots and Ryke, 1968: 2; Lee, 1970: 33; Hurlbutt, 1974:

573.



TYPE LOCALITY AND HABITAT: Tanzania, above Maskame, Mount Kilimanjaro, alt.

2400 m, in mosses on trees in a mountain forest.

39. Afrogamasellus latigynia Hurlbutt, 1974

66

Afrogamasellus latigynia Hurlbutt, 1974: 575.




Regional Forestry Office and Mount Lupanga, Morogoro, alt. 1100 m, 12 June 1966,

in soil under tree.

40. Afrogamasellus lokelei Van Daele, 1976

Afrogamasellus lokelei Van Daele, 1976: 339.

TYPE DEPOSITORY: Chair of Zoology, Faculty of Agricultural Sciences, State University

of Ghent, Belgium.

TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Yangambi, 1

November 1974, in litter at the base of a decaying banana plant [Musaceae].

41. Afrogamasellus lootsi Hurlbutt, 1974

Afrogamasellus lootsi Hurlbutt, 1974: 578.


TYPE LOCALITY AND HABITAT: Tanzania, adjacent to Morogoro River, Morogoro, alt.

550 m, 15 February 1968, on dead leaves, twigs and in humus under bushes and

trees.

42. Afrogamasellus luberoensis luberoensis Loots, 1968

Afrogamasellus luberoensis luberoensis Loots, 1968: 366.

Afrogamasellus luberoensis luberoensis.— Antony, 1986: 147.

Afrogamasellus luberoensis.— Lee, 1970: 34; Hurlbutt, 1974: 569 (part).


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], near Kakolwe

River, Lubero, Kivu Province, 21 December 1953, in soil.

42a. Afrogamasellus luberoensis kalibuensis Loots, 1968

Afrogamasellus luberoensis kalibuensis Loots, 1968: 370.

Afrogamasellus luberoensis kalibuensis.— Lee, 1970: 34; Antony, 1986: 147.

Afrogamasellus luberoensis.— Hurlbutt, 1974: 569 (part).


67

TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], near

Kaliba River, Ruwenzori, 23 January 1954, in soil.

43. Afrogamasellus lyamunguensis Hurlbutt, 1974

Afrogamasellus lyamunguensis Hurlbutt, 1974: 581.


TYPE LOCALITY AND HABITAT: Tanzania, Lyamungu Research and Training Centre,

Lyamungu, [Hai, Kilimanjaro Region], alt. 1300 m, 9 June 1972, fine fragments of

leaves and twigs in forest.

44. Afrogamasellus maskamensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) maskamensis Ryke and Loots, 1966: 135.

Afrogamasellus maskamensis.— Loots and Ryke, 1968: 2; Lee, 1970: 33; Hurlbutt, 1974:

583.



TYPE LOCALITY AND HABITAT: Tanzania, above Maskame, Mount Kilimanjaro, alt.

2300 m, July 1962, in forest soil.

45. Afrogamasellus mitigatus (Berlese, 1923)

Gamasellus mitigatus Berlese, 1923: 250.

Cyrtolaelaps (Gamasellus) mitigatus.— Ryke, 1962b: 46.

Afrogamasellus mitigatus.— Loots and Ryke, 1968: 2; Lee, 1970: 34; Hurlbutt, 1974: 571.

Gamasellus mitigatus.— Castagnoli and Pegazzano, 1985: 257.



46. Afrogamasellus muhiensis Loots, 1969

Afrogamasellus muhiensis Loots, 1969b: 369.

Afrogamasellus muhiensis.— Lee, 1970: 34; Hurlbutt 1974: 570.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], Mount Muhi,

Kivu Province, July 1955, in soil humus.

68

47. Afrogamasellus nyinabitabaensis Loots, 1969

Afrogamasellus nyinabitabaensis Loots, 1969b: 381.

Afrogamasellus nyinabitabaensis.— Lee, 1970: 34; Hurlbutt, 1974: 570.



1952, in soil.

48. Afrogamasellus paratruncatus Hurlbutt, 1974

Afrogamasellus paratruncatus Hurlbutt, 1974: 585.




Mountains, [Morogoro], alt. 6900 feet, 1 May 1968, in moss.

49. Afrogamasellus quadrisigillatus (Berlese, 1916)

Gamasellus quadrisigillatus Berlese, 1916a: 160.

Cyrtolaelaps (Gamasellus) quadrisigillatus.— Ryke, 1962b: 48; Ryke and Loots, 1966: 124.

Afrogamasellus quadrisigillatus.— Loots and Ryke, 1968: 2; Lee, 1970: 33; Hurlbutt, 1974:

571.

Gamasellus quadrisigillatus.— Castagnoli and Pegazzano, 1985: 348.



50. Afrogamasellus rugegensis Loots, 1969

Afrogamasellus rugegensis Loots, 1969b: 375.

Afrogamasellus rugegensis.— Lee, 1970: 34; Hurlbutt, 1974: 570.


TYPE LOCALITY AND HABITAT: Rwanda, Rugege Forest, May 1951, in soil.

51. Afrogamasellus tetrastigma (Berlese, 1916)

Gamasellus tetrastigma Berlese, 1916a: 161.

Cyrtolaelaps (Gamasellus) tetrastigma.— Ryke, 1962b: 46.

Afrogamasellus tetrastigma.— Loots and Ryke, 1968: 2; Loots, 1969b: 361; Lee, 1970: 34;

Loots, 1971: 3; Hurlbutt, 1974: 571.

69

Afrogamasellus (Podalogamasellus) tetrastigma.— Karg, 1977: 345.

Gamasellus tetrastigma.— Castagnoli and Pegazzano, 1985: 414.



52. Afrogamasellus truncatus Hurlbutt, 1974

Afrogamasellus truncatus Hurlbutt, 1974: 583.




Regional Forestry Office and Mount Lupanga, above Morogoro, alt. 1100 m, 18

January 1967, on dead leaves and twigs under clump of trees.

53. Afrogamasellus uluguruensis Hurlbutt, 1974

Afrogamasellus uluguruensis Hurlbutt, 1974: 578.




Mountains, [Morogoro], alt. 1500 m, in leaf litter from rain forest.

54. Afrogamasellus uviraensis (Ryke and Loots, 1966)

Cyrtolaelaps (Gamasellus) uviraensis uviraensis Ryke and Loots, 1966: 143.

Afrogamasellus uviraensis.— Loots and Ryke, 1968: 2; Lee, 1970: 33.

Afrodacarellus uviraensis.— Hurlbutt, 1974: 590.

Afrogamasellus (Foliogamasellus) uviraensis.— Karg, 1977: 344.


TYPE LOCALITY AND HABITAT: [Democratic Republic of the Congo], near Lake

Tanganyika, Uvira, Kivu Province, November 1959, in soil.

Genus Binodacarus Castilho and Moraes, 2010

Binodacarus Castilho and Moraes, 2010: 387 (described in Rhodacaridae Oudemans).

Type species: Binodacarus brasiliensis Castilho and Moraes, 2010, by original

designation.

70

55. Binodacarus brasiliensis Castilho and Moraes, 2010

Binodacarus brasiliensis Castilho and Moraes, 2010: 389.

TYPE DEPOSITORY: Departamento de Entomologia e Acarologia, Escola Superior de

Agricultura ―Luiz de Queiroz‖, Universidade de São Paulo, Piracicaba, State of São

Paulo, Brazil.

TYPE LOCALITY AND HABITAT: Brazil, Pirassununga, São Paulo, 3 May 2000, in soil

under Psidium guajava [Myrtaceae].

Genus Interrhodeus Karg, 2000

Interrhodeus Karg, 2000a: 258 (described in Rhodacaridae Oudemans).

Type species: Interrhodeus brevicornus Karg, 2000, by original designation.

56. Interrhodeus brevicornus Karg, 2000

Interrhodeus brevicornus Karg, 2000a: 259.



TYPE LOCALITY AND HABITAT: Costa Rica, La Selva, 11 June 1993, in soil.

Genus Mediorhodacarus Shcherbak, 1976

Mediorhodacarus Shcherbak, 1976: 949 (described in Rhodacaridae Oudemans).

Mediorhodacarus.— Shcherbak, 1980: 37.

Type species: Mediorhodacarus tetranodulosus Shcherbak, 1976, by original

designation.

57. Mediorhodacarus tetranodulosus Shcherbak, 1976

Mediorhodacarus tetranodulosus Shcherbak, 1976: 951.

Mediorhodacarus tetranodulosus.— Shcherbak, 1980: 37.

TYPE DEPOSITORY: Institute of Zoology and Biology of the All-Ukrainian Academy of

Sciences (AUAS), Kiev, Ukraine.

TYPE LOCALITY AND HABITAT: Ukraine, Lanzheron Beach, Odessa, 11 July 1974, in

sand (depth 30-40 cm), 3 metres from the water edge.

71

Genus Minirhodacarellus Shcherbak, 1980

Minirhodacarellus Shcherbak, 1980: 92 (described in Rhodacaridae Oudemans).

Minirhodacarellus.— Karg, 1993: 336.

Type species: Rhodacarellus minimus Karg, 1961, by original designation.

58. Minirhodacarellus minimus (Karg, 1961)

Rhodacarellus minimus Karg, 1961: 128.

Rhodacarus minimus.— Hirschmann, 1962: 49; Karg, 1971: 317.

Rhodacarellus minimus.— Emberson 1968: 156; Lee, 1970: 37; Bregetova and Shcherbak,

1977b: 275; Antony, 1986: 148.

Minirhodacarellus minimus.— Shcherbak, 1980: 92; Karg, 1993: 336.

TYPE DEPOSITORY: Institut für Pflanzenschutzforschung – Biologische Zentralanstalt der

Deutschen Akademie der Landwirtschaftswissenschaften zu Berlin, Kleinmachnow,

Germany.

TYPE LOCALITY AND HABITAT: Germany, Berlin, in cultivated soil and pasture.

Genus Multidentorhodacarus Karg, 2000

Multidentorhodacarus Karg, 2000b: 144 (described in Rhodacaridae Oudemans).

Rhodacarus (Multidentorhodacarus) Shcherbak, 1980: 72 (nomen nudum).

Rhodacarus (Multidentorhodacarus).— Karg, 1996: 170 (nomen nudum).

Rhodacarus (Multidentrhodacarus) [sic].— Karg, 1998: 186 (nomen nudum).

Multidentorhodacarus.— Karg, 2000a: 259 (nomen nudum).

Type species: Rhodacarus denticulatus Berlese, 1920 (by original designation, Karg,

2000b).

NOTE: Multidentorhodacarus was not made available by Shcherbak (1980) because a type

species for the subgenus was not fixed (International Code of Zoological Nomenclature,

Article 13.3). The name did not become available until Karg (2000b) published a description

of Multidentorhodacarus as a genus, and stated that its type species was M. denticulatus

(Berlese, 1920). The names of species described in the genus Multidentorhodacarus before

72

2000 are available, even though the genus name was not available at the time (International

Code of Zoological Nomenclature, Article 11.9.3.1).

59. Multidentorhodacarus ananasi (Ryke, 1962)

Rhodacarus ananasi Ryke, 1962c: 82.

Rhodacarus (Rhodacarus) ananasi.— Loots, 1969a: 50.

Rhodacarus ananasi.— Lee, 1970: 29.

Multidentorhodacarus ananasi.— Karg, 2000b: 145.



TYPE LOCALITY AND HABITAT: South Africa, Bathurst, June 1956, in soil of pineapple

fields [Bromeliaceae].

60. Multidentorhodacarus angustacuminis (Karg, 1998)

Rhodacarus (Multidentrhodacarus) [sic] angustacuminis Karg, 1998: 187.

Multidentorhodacarus angustacuminis.— Karg, 2000b: 148.



TYPE LOCALITY AND HABITAT: Ecuador, between Pifo and Papallacta, Pichincha

Province, alt. 4100 m, 14 April 1989, in moss and withered plant debris from under

bushes.

61. Multidentorhodacarus brevicuspidis Karg, 2000

Multidentorhodacarus brevicuspidis Karg, 2000c: 211.


TYPE LOCALITY AND HABITAT: Costa Rica, near La Selva, December 1993, in soil.

62. Multidentorhodacarus brevisetosus Karg, 2000

Multidentorhodacarus brevisetosus Karg, 2000c: 212.


TYPE LOCALITY AND HABITAT: Costa Rica, near La Selva, August 1993, in soil.

63. Multidentorhodacarus denticulatus (Berlese, 1920)

Rhodacarus denticulatus Berlese, 1920: 164.

73

Rhodacarus denticulatus.— Willmann, 1951: 119; Athias-Henriot, 1961: 499; Emberson,

1968: 156; Lee, 1970: 29; Karg, 1971: 318; Shcherbak and Furman, 1975: 47;

Bregetova and Shcherbak, 1977b: 266; Castagnoli and Pegazzano, 1985: 111; Karg,

1993: 331; Karg, 1996: 170; Karg, 1998: 187.

Rhodacarus (Multidentorhodacarus) denticulatus.— Shcherbak, 1980: 72; Farrier and

Hennessey, 1993: 133.

Rhodacarus (Multidentrhodacarus) [sic] denticulatus.— Karg, 1998: 187.

Multidentorhodacarus denticulatus.— Karg, 2000b: 145.

Rhodacarus guevarai Guevara-Benitez, 1974: 207 (synonymy by Shcherbak, 1980: 72).

TYPE DEPOSITORY of M. denticulatus: Istituto Sperimentale per la Zoologia Agraria,

Florence, Italy; of R. guevarai: Laboratorio de Acarología del Instituto ―Lopez-

Neyra‖ de Parasitologia de Granada, Granada, Spain.

TYPE LOCALITY AND HABITAT of M. denticulatus: Indonesia, Semarang, Java, and

USA, Lake City, Columbia County, Florida, on unspecified substrate; of R. guevarai:

Spain, Jardín Experimental del Instituto ―Lopez-Neyra‖ de Parasitologia de Granada,

[Armilla], Granada, June 1971, in soil under Piptatherum miliaceum [Poaceae].

NOTE: Shcherbak and Furman (1975) listed M. ruwenzoriensis Loots, 1969 as a synonym of

M. denticulatus, but Shcherbak (1980) considered this synonymy as questionable

(see below). Emberson (1968) also pointed out that Berlese's original type series was

probably heterogeneous.

64. Multidentorhodacarus differentis Karg and Schorlemmer, 2009

Multidentorhodacarus differentis Karg and Schorlemmer, 2009: 67.



TYPE LOCALITY AND HABITAT: Ecuador, between Puerto Napo and Ahuano Tenatol,

1990, in litter of a cacao plantation [Sterculiaceae].

65. Multidentorhodacarus minutocorpus (Karg, 1998)

Rhodacarus (Multidentrhodacarus) [sic] minutocorpus Karg, 1998: 186.

Multidentorhodacarus minutocorpus.— Karg, 2000b: 145.



74

TYPE LOCALITY AND HABITAT: Ecuador, Pia Santa Rosa, near San Francisco, Pichincha

Province, alt. 2990 m, 13 April 1989, in moss and soil from a roadside.

66. Multidentorhodacarus paulista Castilho and Moraes, 2010

Multidentorhodacarus paulista Castilho and Moraes, 2010: 392.



Paulo, Brazil.

TYPE LOCALITY AND HABITAT: Brazil, Piracicaba, São Paulo, 11 November 2000, in

soil under Syagrus oleracea [Arecaceae].

67. Multidentorhodacarus pennacornutus Karg, 2000

Multidentorhodacarus pennacornutus Karg, 2000a: 259.

Multidentorhodacarus pennacornutus.— Karg, 2000b: 145.



TYPE LOCALITY AND HABITAT: Cuba, Pinar del Rio, 1975, in litter.

68. Multidentorhodacarus ruwenzoriensis (Loots, 1969)

Rhodacarus (Rhodacarus) ruwenzoriensis Loots, 1969a: 54.

Multidentorhodacarus ruwenzoriensis.— Karg, 2000b: 148.


TYPE LOCALITY AND HABITAT: Uganda, near Nyabitaba, Ruwenzori, alt. 8650 feet,

1952, in soil.

NOTE: Shcherbak and Furman (1975) synonymised R. ruwenzoriensis with R. denticulatus.

However, Shcherbak (1980) stated that these two species were very similar, but the

synonymy could not be confirmed until they were examined more closely. We

therefore treat these two species as separate for the moment.

69. Multidentorhodacarus sogdianus (Shcherbak, 1980)

Rhodacarus (Multidentorhodacarus) sogdianus Shcherbak, 1980: 74.

Multidentorhodacarus sogdianus.— Karg, 2000b: 145.



75

TYPE LOCALITY AND HABITAT: Tajikistan, Ramidskoe Valley, Gissar Range, 23 April

1978, in soil under rock along a brook.

70. Multidentorhodacarus squamosus Karg, 2000

Multidentorhodacarus squamosus Karg, 2000b: 144.



TYPE LOCALITY AND HABITAT: Costa Rica, La Selva, March 1993, in primary forest.

71. Multidentorhodacarus sublapideus (Ryke, 1962)

Rhodacarus sublapideus Ryke, 1962c: 82.

Rhodacarus (Rhodacarus) sublapidius [sic].— Loots, 1969a: 50.

Rhodacarus sublapideus.— Lee, 1970: 29.

Multidentorhodacarus sublapideus.— Karg, 2000b: 145.



TYPE LOCALITY AND HABITAT: South Africa, University campus, Potchefstroom,

March 1960, in soil in termite‘s nest under stone.

72. Multidentorhodacarus tertius (Karg, 1996)

Rhodacarus (Multidentorhodacarus) tertius Karg, 1996: 171.

Multidentorhodacarus tertius.— Karg, 2000b: 148.



TYPE LOCALITY AND HABITAT: New Caledonia, near Lifou, 20 February 1977, in

primary forest.

73. Multidentorhodacarus thysi (Jordaan, Loots and Theron, 1988)

Rhodacarus thysi Jordaan, Loots and Theron, 1988: 279.



TYPE LOCALITY AND HABITAT: South Africa, 20 km south of Kuruman, Northern Cape,

30 October 1977, in soil under Tarchonanthus sp. [Asteraceae].

76

74. Multidentorhodacarus triramulus (Karg, 1998)

Rhodacarus (Multidentrhodacarus) [sic] triramulus Karg, 1998: 187.

Multidentorhodacarus triramulus.— Karg, 2000b: 145.



TYPE LOCALITY AND HABITAT: Ecuador, Rio Guajalito, Las Palmeras, Pichincha

Province, alt. 1850 m, 18 April 1989, in moss from the vertical river bank.

Genus Paragamasellevans Loots and Ryke, 1968

Paragamasellevans Loots and Ryke, 1968: 3 (described in Rhodacaridae Oudemans).

Paragamasellevans.— Loots, 1969c: 183; Lee, 1970: 193; Loots, 1979: 2.

Type species: Paragamasellevans michaeli Loots and Ryke, 1968, by original

designation.

75. Paragamasellevans michaeli Loots and Ryke, 1968

Paragamasellevans michaeli Loots and Ryke, 1968: 5.

Paragamasellevans michaeli.— Loots, 1971: 7; Silva, 2007: 84.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, Transvaal, 1963, in

forest soil.

76. Paragamasellevans vandenbergi Loots and Ryke, 1968

Paragamasellevans vandenbergi Loots and Ryke, 1968: 12.

Paragamasellevans vandenbergi.— Antony, 1986: 146; Silva, 2007: 84.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, Transvaal, 1963, in

forest soil.

Genus Pararhodacarus Jordaan, Loots and Theron, 1988

77

Pararhodacarus Jordaan, Loots and Theron, 1988: 275 (described in Rhodacaridae

Oudemans).

Type species: Pararhodacarus intermedius Jordaan, Loots and Theron, 1988, by

original designation.

77. Pararhodacarus intermedius Jordaan, Loots and Theron, 1988

Pararhodacarus intermedius Jordaan, Loots and Theron, 1988: 276.



TYPE LOCALITY AND HABITAT: South Africa, 28 km north of Kuruman, Northern Cape,

2 December 1977, in soil under Tarchonanthus sp. [Asteraceae].

Genus Pennarhodeus Karg, 2000

Pennarhodeus Karg, 2000a: 255 (described in Rhodacaridae Oudemans).

Type species: Pennarhodeus pennatus Karg, 2000, by original designation.

78. Pennarhodeus brevipennatus Karg, 2000

Pennarhodeus brevipennatus Karg, 2000c: 211.


TYPE LOCALITY AND HABITAT: Costa Rica, 3 August 1993, in soil and litter.

NOTE: Species described based only an adult male.

79. Pennarhodeus decoris Karg, 2000

Pennarhodeus decoris Karg, 2000a: 255.

Pennarhodeus decoris.— Karg, 2000c: 211.



TYPE LOCALITY AND HABITAT: Cuba, Pinar del Rio, 1976, in soil.

80. Pennarhodeus pennatus Karg, 2000

Pennarhodeus pennatus Karg, 2000a: 255.

Pennarhodeus pennatus.— Karg, 2000c: 211.

78




81. Pennarhodeus turris Karg, 2000

Pennarhodeus turris Karg, 2000a: 257.

Pennarhodeus turris.— Karg, 2000c: 211.




NOTE: Species described based only an adult male.

Genus Poropodalius Karg, 2000

Poropodalius Karg, 2000a: 252 (described in Rhodacaridae Oudemans).

Poropodalius.— Karg and Schorlemmer, 2009: 66.

Type species: Poropodalius hexapennatus Karg, 2000, by original designation.

82. Poropodalius acutus Karg, 2000

Poropodalius acutus Karg, 2000a: 253.

Poropodalius acutus.— Karg and Schorlemmer, 2009: 67.



TYPE LOCALITY AND HABITAT: Cuba, Sierra Esperon, 1977, in soil.

83. Poropodalius basisetae Karg, 2000

Poropodalius basisetae Karg, 2000a: 255.

Poropodalius basisetae.— Karg and Schorlemmer, 2009: 67.




84. Poropodalius crispus Karg, 2000

Poropodalius crispus Karg, 2000a: 253.

79

Poropodalius crispus.— Karg and Schorlemmer, 2009: 67.




85. Poropodalius hexapennatus Karg, 2000

Poropodalius hexapennatus Karg, 2000a: 252.

Poropodalius hexapennatus.— Karg and Schorlemmer, 2009: 67.




86. Poropodalius medioflagelli Karg and Schorlemmer, 2009

Poropodalius medioflagelli Karg and Schorlemmer, 2009: 66.



TYPE LOCALITY AND HABITAT: Venezuela, near the street from Caracas to La Guaira,

1973, in litter and soil.

Genus Protogamasellopsis Evans and Purvis, 1987

Protogamasellopsis Evans and Purvis, 1987: 855 (described in Ascidae Voigts and

Oudemans).

Protogamasellopsis.— Karg, 1994a: 123; Karg, 1994b: 207; Halliday et al., 1998: 2; Karg,

2007: 123.

Type species: Protogamasellopsis corticalis Evans and Purvis, 1987, by original

designation.

Rhodacarella Moraza, 2004: 2 (described in Rhodacaridae Oudemans) (synonymy, following

M.L. Moraza, personal communication, 2007).

Type species: Rhodacarella cavernicola Moraza, 2004, by monotypy.

87. Protogamasellopsis corticalis Evans and Purvis, 1987

Protogamasellopsis corticalis Evans and Purvis, 1987: 856.

80

Protogamasellopsis corticalis.— Karg, 1994a: 124; Karg, 1994b: 208; Shaw, 1999: 45; Karg,

2007: 124.


TYPE LOCALITY AND HABITAT: Saint Helena [South Atlantic island], Jamestown, public

gardens, under the dead bark of a Citrus sp. [Rutaceae] (possibly imported from

Cape Province, South Africa).

88. Protogamasellopsis dioscorus (Manson, 1972)

Protogamasellus dioscorus Manson, 1972: 437.

Protogamasellopsis dioscorus.— Evans and Purvis, 1987: 855; Karg, 1994a: 123; Karg,

1994b: 208; Karg, 2007: 124; Castilho and Moraes, 2010: 397.

TYPE DEPOSITORY: Collection of the Department of Agriculture, Levin, New Zealand.

TYPE LOCALITY AND HABITAT: New Zealand, Auckland International Airport, 19 June

1969, intercepted on yam [Dioscoreaceae] from Tonga.

89. Protogamasellopsis granulosus Karg, 1994

Protogamasellopsis granulosus Karg, 1994b: 208.

Protogamasellopsis granulosus.— Karg, 2007: 124.



TYPE LOCALITY AND HABITAT: Galapagos Islands, Cueva Bella Vista, Bella Vista,

Santa Cruz, 15-25 May 1985, in trap with manure.

90. Protogamasellopsis leptosomae Karg, 1994

Protogamasellopsis leptosomae Karg, 1994b: 210.

Protogamasellopsis leptosomae.— Karg, 2007: 123.



TYPE LOCALITY AND HABITAT: Galapagos Islands, Puntudo, Santa Cruz, 10 March

1985, in litter of fern [Pteridophyta] and pieces of wood.

91. Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, 1991

Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, 1991a: 189.

81

Protogamasellopsis posnaniensis.— Karg, 1993: 369; Karg, 1994b: 208; Karg, 2007: 123;

Castilho et al., 2009: 165.

Rhodacarella cavernicola Moraza, 2004: 4 (synonymy, following M.L. Moraza, personal

communication, 2007).

TYPE DEPOSITORY of P. posnaniensis: Department of Forest Protection, University of Life

Sciences, Poznań, Poland; of R. cavernicola: Florida Collection of Arthropods,

Division of Plant Industry, Gainesville, Florida, USA.

TYPE LOCALITY AND HABITAT of P. posnaniensis: Poland, Poznań, July to August

1989, in litter of Phoenix sp. [Aracaceae] in greenhouse; of R. cavernicola: USA,

Kartchner Caverns State Park, Cochise County, Arizona, 19 May 1990, on bat guano.

92. Protogamasellopsis praeendopodalis Karg, 1994

Protogamasellopsis praeendopodalis Karg, 1994a: 123.

Protogamasellopsis praeendopodalis.— Karg, 1994b: 208; Karg, 2007: 123.



TYPE LOCALITY AND HABITAT: Galapagos Islands, Fernandina, 18 March 1985, in litter

of grass and sand in coastal area.

93. Protogamasellopsis transversus Karg, 2000

Protogamasellopsis transversus Karg, 2000a: 252.

Protogamasellopsis transversus.— Karg, 2007: 123.



TYPE LOCALITY AND HABITAT: Ecuador, Cotopaxi Province, 1973, in paramo.

Genus Rhodacarellus Willmann, 1935

Rhodacarellus Willmann, 1935: 429 (described in Rhodacaridae Oudemans).

Rhodacarellus.— Evans, 1957: 221; Sheals, 1958: 302; Lee, 1970: 36; Karg, 1971: 323;

Bregetova and Shcherbak, 1977b: 272; Evans and Till, 1979: 206; Shcherbak, 1980:

76; Fouly, 1992: 436; Karg, 1993: 336.

Type species: Rhodacarellus subterraneus Willmann, 1935, by original designation.

82

94. Rhodacarellus apophyseus Karg, 1971

Rhodacarellus apophyseus Karg, 1971: 324.

Rhodacarellus apophyseus.— Bregetova and Shcherbak, 1977b: 279; Shcherbak, 1980: 86;

Karg, 1993: 339.



Germany.

TYPE LOCALITY AND HABITAT: Germany, near Halberstadt, Holtemme, 5 September

1963, flood plain.

95. Rhodacarellus arcanus (Athias-Henriot, 1961)

Rhodacaropsis arcanus Athias-Henriot, 1961: 497.

Rhodacarellus arcanus.— Loots, 1969a: 47; Lee, 1970: 37; Bregetova and Shcherbak, 1977b:

276; Shcherbak, 1980: 91; Shcherbak, 1982: 63.

TYPE DEPOSITORY: Laboratoire d‘Acarologie de l‘École Pratique des Hautes Études,

Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, l'Oued Bouzaréah, 3 January 1961, in soil

under Laurus nobilis [Lauraceae].

96. Rhodacarellus citri Fouly, 1992

Rhodacarellus citri Fouly, 1992: 436.

TYPE DEPOSITORY: Collection of Department of Plant Protection, Faculty of Agriculture,

Mansoura University, Mansoura, Egypt.

TYPE LOCALITY AND HABITAT: Egypt, farm of the Faculty of Agriculture, Mansoura

University, Mansoura, 1990-91, in 10 cm depth from soil surface under Citrus sp.

orchards [Rutaceae].

97. Rhodacarellus corniculatus Willmann, 1935

Rhodacarellus corniculatus Willmann, 1935: 432.

Rhodacarellus corniculatus.— Athias-Henriot, 1961: 486; Karg, 1961: 128; Lee, 1970: 37;

Bregetova and Shcherbak, 1977b: 280; Shcherbak, 1980: 79; Krantz, 1986: 634;

Karg, 1993: 340.

TYPE DEPOSITORY: Zoologische Staatssammlung München, Munich, Germany.

83

TYPE LOCALITY AND HABITAT: Germany, Leipzig, in forest soil on the edge of a

stream.

98. Rhodacarellus epigynialis Sheals, 1956

Rhodacarellus epigynialis Sheals, 1956: 102.

Rhodacarellus epigynialis.— Sheals, 1958: 302; Athias-Henriot, 1961: 486; Karg, 1961: 128;

Lee, 1970: 37; Karg, 1971: 325; Bregetova and Shcherbak, 1977b: 275; Shcherbak,

1980: 83; Karg, 1993: 336.


TYPE LOCALITY AND HABITAT: Scotland, Bellahouston Park, Glasgow, June 1951-

October 1952, in soil of old grassland.

99. Rhodacarellus francescae Athias-Henriot, 1961

Rhodacarellus francescae Athias-Henriot, 1961: 491.

Rhodacarellus francescae.— Hirschmann, 1962: 50; Lee, 1970: 37; Bregetova and

Shcherbak, 1977b: 276; Shcherbak, 1980: 89; Shcherbak, 1982: 63.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, l'Oued Bouzaréah, 3 January 1961, in soil

under Laurus nobilis [Lauraceae].

100. Rhodacarellus kreuzi Karg, 1965

Rhodacarellus kreuzi Karg, 1965: 296.

Rhodacarellus kreuzi.— Lee, 1970: 37; Karg, 1971: 326; Shcherbak and Furman, 1975: 50;

Bregetova and Shcherbak, 1977b: 280; Shcherbak, 1980: 85; Karg, 1993: 340.



Germany.

TYPE LOCALITY AND HABITAT: Germany, near Delitzsch, Lemsel, Leipzig, 1960, in

flooded pasture.

101. Rhodacarellus liuzhiyingi Ma, 1995

Rhodacarellus liuzhiyingi Ma, 1995: 50.

Rhodacarellus liuzhiyingi.— Ma, 2005: 17.

84

TYPE DEPOSITORY: National Base of Plague and Brucellosis Control, Baicheng, China.

TYPE LOCALITY AND HABITAT: China, Baicheng, Jilin Province, July-September 1993,

under the decomposed bark of poplar [Salicaceae].

NOTE: This species is only provisionally placed in this genus, given the absence of key

morphological information about the species in the original description. When the

specimens are examined in detail, we believe this species will be transferred to the

family Digamasellidae.

102. Rhodacarellus maxidactylus Karg, 2000

Rhodacarellus maxidactylus Karg, 2000c: 208.


TYPE LOCALITY AND HABITAT: Costa Rica, 1 November 1993, in soil.

103. Rhodacarellus moneli Solomon, 1978

Rhodacarellus moneli Solomon, 1978: 85.


TYPE LOCALITY AND HABITAT: Romania, Voinesti (Iaşi) forest, Iasi, 1976, in litter of

carpineto-fagetum association.

104. Rhodacarellus montanus Shcherbak, 1980

Rhodacarellus montanus Shcherbak, 1980: 92.



TYPE LOCALITY AND HABITAT: Turkmenistan, summit of Mountain Dushak, alt. 2000

m, 17 May 1978, in soil under Juniperus seravschanica [Cupressaceae].

105. Rhodacarellus perspicuus Halašková, 1959

Rhodacarellus epigynialis perspicuus Halašková, 1959: 98.

Rhodacarellus perspicuus.— Athias-Henriot, 1961: 486; Karg, 1971: 324; Bregetova and

Shcherbak, 1977b: 279; Shcherbak, 1980: 84; Karg, 1993: 336.

Rhodacarellus epigynialis perspicuus.— Lee, 1970: 37.

TYPE DEPOSITORY: Unknown.

85

TYPE LOCALITY AND HABITAT: Czech Republic, Strančice, Řičany, South Prague, 15

January 1956, in soil of a Picea excelsa vulgaris [Pinaceae] forest sparsely covered

with Oxalis acetosella [Oxalidaceae].

106. Rhodacarellus shandongensis Ma, 2008

Rhodacarellus shandongensis Ma, 2008: 127.

TYPE DEPOSITORY: Entomology Gallery, Institute of Microbiology and Epidemiology,

Academy of Military Medical Sciences, Beijing, China.

TYPE LOCALITY AND HABITAT: China, Tai‘an (36°15'N, 117°08'E), Shandong Province,

15 July 2000, under bark of tree.





107. Rhodacarellus silesiacus Willmann, 1936

Rhodacarellus silesiacus Willmann, 1936: 282.

Rhodacarellus silesiacus.— Sheals, 1958: 302; Athias-Henriot, 1961: 488; Karg, 1961: 128;

Hirschmann, 1962: 49; Emberson, 1968: 160; Lee, 1970: 37; Karg, 1971: 324; Lee,

1973a: 3; Shcherbak and Furman, 1975: 50; Bregetova and Shcherbak, 1977b: 279;

Shcherbak, 1980: 80; Krantz, 1986: 634; Farrier and Hennessey, 1993: 132; Karg,

1993: 339; Meng et al., 2007: 242.

TYPE DEPOSITORY: Zoologische Staatssammlung München, Germany.

TYPE LOCALITY AND HABITAT: Germany, near Hundsfeld, Mähwiese; Germany,

Waldenburg, Bergland; [Poland], Breslau [Wrocław]; in pasture and prairie.

108. Rhodacarellus subterraneus Willmann, 1935

Rhodacarellus subterraneus Willmann, 1935: 430.

Rhodacarellus subterranus [sic].— Schweizer, 1961: 89.

Rhodacarellus subterraneus.— Athias-Henriot, 1961: 488; Karg, 1961: 127; Lee, 1970: 37;

Karg, 1971: 324; Bregetova and Shcherbak, 1977b: 276; Shcherbak, 1980: 78;

Shcherbak, 1982: 63; Farrier and Hennessey, 1993: 132; Karg, 1993: 339.


86

TYPE LOCALITY AND HABITAT: Germany, Leipzig, Saxony, in forest soil of a riparian

zone.

109. Rhodacarellus tadchikistanicus Shcherbak, 1980

Rhodacarellus tadchikistanicus Shcherbak, 1980: 87.



TYPE LOCALITY AND HABITAT: Tajikistan, Ramidskoe Valley, Gissar Range, 23 April

1978, in soil under rock along a stream.

NOTE: Species described based on the stages of protonymph, deutonymph and adult male.

110. Rhodacarellus tebeenus Hafez and Nasr, 1979

Rhodacarellus tebeenus Hafez and Nasr, 1979: 78.

Rhodacarellus tebeenus.— Zaher, 1986: 17.


TYPE LOCALITY AND HABITAT: Egypt, El-Tebeen, Helwan, Cairo Governate, in soil

cultivated with banana [Musaceae].

111. Rhodacarellus unicus Karg, 2000

Rhodacarellus unicus Karg, 2000a: 251.




112. Rhodacarellus vervacti (Athias-Henriot, 1961)

Rhodacaropsis vervacti Athias-Henriot, 1961: 497.

Rhodacarellus vervacti.— Lee, 1970: 37.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, road to Gué-de-Constantine, Baraki, May-June

1959, in soil under Scolymus spp. [Asteraceae].

NOTE: This species was described only from the deutonymph. Lee (1970) placed it in

Rhodacarellus, apparently based only on the original description.

87

113. Rhodacarellus yalujiangensis Ma, 2003

Rhodacarellus yalujiangensis Ma, 2003: 85.

Rhodacarellus yalujiangensis.— Ma, 2005: 18.


TYPE LOCALITY AND HABITAT: China, Linjiang County, Jilin Province, 2 August 1999,

under decomposed bark.





Genus Rhodacaropsis Willmann, 1935

Rhodacaropsis Willmann, 1935: 426 (described in Rhodacaridae Oudemans).

Rhodacarus (Rhodacaropsis).— Loots, 1969a: 56; Loots, 1969c: 182; Loots, 1979: 2.

Rhodacaropsis.— Lee, 1970: 37; Bregetova and Shcherbak, 1977b: 270; Shcherbak, 1980:

32; Luxton, 1992: 237; Karg, 1993: 336.

Type species: Rhodacaropsis inexpectatus Willmann, 1935, by original designation.

114. Rhodacaropsis attenuatus (Loots, 1969)

Rhodacarus (Rhodacaropsis) attenuatus Loots, 1969a: 56.

Rhodacaropsis attenuatus.— Shcherbak, 1980: 33; Antony, 1986: 151; Luxton, 1992: 241.

Rhodacaropsis attenuata.— Krantz, 1986: 634.


TYPE LOCALITY AND HABITAT: South Africa, Blauwberg Beach, Cape Town, [Western

Cape], 1966, in soil in the intertidal zone.

115. Rhodacaropsis botosaneanui (Petrova and Beron, 1973)

Rhodacarus (Rhodacaropsis) botosaneanui Petrova and Beron, 1973: 317.

Rhodacaropsis botosaneanui.— Shcherbak, 1980: 33; Krantz, 1986: 634; Luxton, 1992: 241.

TYPE DEPOSITORY: Mite Collection of the Institute of Zoology of the Bulgarian Academy

of Sciences, Sofia, Bulgaria.

TYPE LOCALITY AND HABITAT: Cuba, Baracoa, Guantanamo Province, 4 April 1969, in

sand of track that separates the Rio Miel and Atlantic Ocean.

88

NOTE: Species described based only on adult males.

116. Rhodacaropsis cheungae Luxton, 1992

Rhodacaropsis cheungae Luxton, 1992: 238.


TYPE LOCALITY AND HABITAT: Hong Kong, Cape d‘Aguilar, in beach sand.

117. Rhodacaropsis cubanus (Petrova and Beron, 1973)

Rhodacarus (Rhodacaropsis) cubanus Petrova and Beron, 1973: 319.

Rhodacaropsis cubanus.— Shcherbak, 1980: 33; Luxton, 1992: 242; Farrier and Hennessey,

1993: 133.

Rhodacaropsis cubana.— Krantz, 1986: 634.

TYPE DEPOSITORY: Mite Collection of the Institute of Zoology of the Bulgarian Academy

of Sciences, Sofia, Bulgaria.

TYPE LOCALITY AND HABITAT: Cuba, Baracoa, Guantanamo Province, 4 April 1969, in

sand of track that separates the Rio Miel and Atlantic Ocean.

118. Rhodacaropsis inexpectatus Willmann, 1935

Rhodacaropsis inexpectatus Willmann, 1935: 427.

Rhodacaropsis inexpectatus.— Athias-Henriot, 1961: 491; Emberson, 1968: 157; Lee, 1970:

38; Bregetova and Shcherbak, 1977b: 271; Shcherbak, 1980: 33; Antony, 1986: 151;

Krantz and Ainscough, 1990: 618; Luxton, 1992: 242; Karg, 1993: 336; Farrier and

Hennessey, 1993: 133.

Rhodacaropsis inexpectata.— Krantz, 1986: 634.

TYPE DEPOSITORY: Zoologische Staatssammlung München, Germany.

TYPE LOCALITY AND HABITAT: Germany, Kiel Harbour, in coastal groundwater.

119. Rhodacaropsis ponticus Shcherbak, 1980

Rhodacaropsis ponticus Shcherbak, 1980: 34.

Rhodacaropsis ponticus.— Karg, 1993: 336.



TYPE LOCALITY AND HABITAT: Ukraine, Fox Bay, near Kara Dag Mountain, Black Sea

littoral, Crimea, in sand (depth 2-13 cm), 7 metres from the water edge.

89

Genus Rhodacarus Oudemans, 1902

Rhodacarus Oudemans, 1902: 5 (described in Parasitidae Oudemans).

Rhodacarus.— Evans, 1957: 221; Sheals, 1958: 298; Ryke, 1962c: 81; Loots, 1969a: 49:

Loots, 1969c: 182; Loots, 1979; 2; Lee, 1970: 26; Bregetova and Shcherbak, 1977b:

259; Shcherbak, 1980: 39; Karg, 1993: 331.

Rhodacarus (Rhodacarus).— Shcherbak, 1980: 42.

Type species: Rhodacarus roseus Oudemans, 1902, by monotypy.

120. Rhodacarus aequalis Karg, 1971

Rhodacarus aequalis Karg, 1971: 322.

Rhodacarus aequalis.— Bregetova and Shcherbak, 1977b: 267; Karg, 1993: 335; Kalúz,

1994: 675.

Rhodacarus (Rhodacarus) aequalis.— Shcherbak, 1980: 71.



Germany.

TYPE LOCALITY AND HABITAT: Germany, Krägenriss, near Wörlitz, 13 April 1967, in

humus.

121. Rhodacarus agrestis Karg, 1971

Rhodacarus agrestis Karg, 1971: 322.

Rhodacarus agrestis.— Bregetova and Shcherbak, 1977b: 269; Karg, 1993: 335; Kalúz, 1994:

675.

Rhodacarus (Rhodacarus) agrestis.— Shcherbak, 1980: 70.



Germany.

TYPE LOCALITY AND HABITAT: Germany, Harz, Halberstadt, 11 October 1967, in

cultivated soil.

122. Rhodacarus angustiformis Willmann, 1951

Rhodacarus angustiformis Willmann, 1951: 119.

90

Rhodacarus angustiformis.— Athias-Henriot, 1961: 498; Lee, 1970: 29; Karg, 1971: 318;

Karg, 1993: 331.

Rhodacarus angustiformes [sic].— Bregetova and Shcherbak, 1977b: 269; Shcherbak, 1980:

42.


TYPE LOCALITY AND HABITAT: Austria, Leitha, Purbach am Neusiedlersee,

Burgenland, in litter.

123. Rhodacarus berrisfordi Loots, 1969

Rhodacarus (Rhodacarus) berrisfordi Loots, 1969a: 50.

Rhodacarus berisfordi [sic].— Shcherbak, 1980: 202.

Rhodacarus berrisfordi.— Antony, 1986: 149.

Rhodacarus berresfordi [sic].— Krantz, 1986: 634.


TYPE LOCALITY AND HABITAT: South Africa, Durban Beach, Durban, [Kwazulu-Natal],

1966, in soil in the upper intertidal zone.

124. Rhodacarus calcarulatus Berlese, 1920

Rhodacarus calcarulatus Berlese, 1920: 164.

Rhodacarus pallidus f. calcarulatus.— Sheals, 1958: 299.

Rhodacarus calcarulatus.— Lee, 1970: 29; Bregetova and Shcherbak, 1977b: 269; Castagnoli

and Pegazzano, 1985: 57; Karg, 1993: 335; Kalúz, 1994: 675.

Rhodacarus (Rhodacarus) calcarulatus.— Shcherbak, 1980: 67.

Rhodacarus elbius Karg, 1971: 322 (synonymy by Bregetova and Shcherbak, 1977b: 269).

TYPE DEPOSITORY of R. calcarulatus: Istituto Sperimentale per la Zoologia Agraria,

Florence, Italy; of R. elbius: Institut für Pflanzenschutzforschung – Biologische

Zentralanstalt der Deutschen Akademie der Landwirtschaftswissenschaften zu

Berlin, Kleinmachnow, Germany.

TYPE LOCALITY AND HABITAT of R. calcarulatus: Italy, Portici, Napoli, in moss; of R.

elbius: Germany, margin of Elbe River, Vockerode, [Wittenberg], 1967, in humus.

NOTE1: Sheals (1958) considered R. calcarulatus to be a junior synonym of Rhodacarus

pallidus Hull, naming it Rhodacarus pallidus f. calcarulatus. In the same

publication, he mentioned that T. pallidus had a V-shaped groove posterior to setae

91

j4, z3 and s2, and that R. pallidus f. calcarulatus did not have it. Thus, we understand

that those species should not be considered synonyms.

NOTE2: Rhodacarus calcarulatus Berlese sensu Athias-Henriot (1961) and Karg (1971) were

misidentifications of R. reconditus Athias-Henriot, 1961 (see below). The two

species may be distinguished by the V-shaped groove in the podonotal shield - absent

in R. calcarulatus, present in R. reconditus.

125. Rhodacarus clavulatus Athias-Henriot, 1961

Rhodacarus clavulatus Athias-Henriot, 1961: 502.

Rhodacarus clavulatus.— Lee, 1970: 29; Guevara-Benitez, 1974: 210; Karg, 1993: 334.

Rhodacarus (Rhodacarus) clavulatus.— Shcherbak, 1980: 47.

Rhodacarus ancorae Karg, 1971: 320 (synonymy by Shcherbak, 1980: 47).

TYPE DEPOSITORY of R. clavulatus: Laboratoire d‘Acarologie de l‘École Pratique des

Hautes Études, Paris, France; of R. ancorae: Institut für Pflanzenschutzforschung –

Biologische Zentralanstalt der Deutschen Akademie der

Landwirtschaftswissenschaften zu Berlin, Kleinmachnow, Germany.

TYPE LOCALITY AND HABITAT of R. clavulatus: Algeria, Bouzareah valley, 3 January

1961, in soil and litter under Laurus nobilis [Lauraceae]; of R. ancorae: Germany,

Kemnitztal, near Oelsnitz, 26 September 1967, in moss.

126. Rhodacarus coronatus Berlese, 1920

Rhodacarus coronatus Berlese, 1920: 165.

Rhodacarus (?) coronatus [sic].— Athias-Henriot, 1961: 502.

Rhodacarus coronatus.— Lee, 1970: 29; Karg, 1971: 321; Bregetova and Shcherbak, 1977b:

261; Shcherbak, 1977: 76; Castagnoli and Pegazzano, 1985: 91; Karg, 1993: 333.

Rhodacarus (Rhodacarus) coronatus.— Shcherbak, 1980: 50.

Rhodacarus (?) coronatus forma simplex Athias-Henriot, 1961: 502 (unavailable

infrasubspecific name, International Code of Zoological Nomenclature, Articles 15.2

and 45.6.3).


TYPE LOCALITY AND HABITAT: Italy, Firenze, Boboli and Vallombrosa, in moss.

NOTE: Athias-Henriot (1961) provisionally identified specimens from Italy as Rhodacarus

(?) coronatus, and from Algeria as Rhodacarus (?) coronatus forma simplex. This

92

ambiguity suggests that the name Rhodacarus coronatus conceals more than one

species, and all identifications under this name must be considered as unconfirmed.

127. Rhodacarus cuneatus Athias-Henriot, 1961

Rhodacarus cuneatus Athias-Henriot, 1961: 499.

Rhodacarus cuneatus.— Lee, 1970: 29; Karg, 1971: 321; Bregetova and Shcherbak, 1977b:

264; Karg, 1993: 332.

Rhodacarus (Rhodacarus) cuneatus.— Shcherbak, 1980: 57.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, Bouzaréah, January-March 1958, in litter.

128. Rhodacarus fatrensis Kalúz, 1994

Rhodacarus fatrensis Kalúz, 1994: 675.

TYPE DEPOSITORY: Slovak National Museum, Bratislava, Slovakia.

TYPE LOCALITY AND HABITAT: Slovakia, State Nature Reserve Šrámková, National

Park Malá Fatra montains, 2 June 1982, on unspecified substrate.

129. Rhodacarus furmanae Shcherbak, 1975

Rhodacarus furmanae Shcherbak, in Shcherbak and Furman, 1975: 50.

Rhodacarus furmanae.— Bregetova and Shcherbak, 1977b: 264; Karg, 1993: 332.

Rhodacarus (Rhodacarus) furmanae.— Shcherbak, 1980: 61.



TYPE LOCALITY AND HABITAT: Ukraine, Velikomikhaylovskiy District, Odessa Region,

4 September 1965, in soil.

130. Rhodacarus gracilis Shcherbak, 1980

Rhodacarus (Rhodacarus) gracilis Shcherbak, 1980: 67.



TYPE LOCALITY AND HABITAT: Turkmenistan, summit of Mountain Dushak, alt. 2000

m, 17 May 1978, in soil with roots of Juniperus sp. [Cupressaceae].

93

131. Rhodacarus haarlovi Shcherbak, 1977

Rhodacarus haarlovi Shcherbak, 1977: 79.

Rhodacarus (Rhodacarus) haarlovi.— Shcherbak, 1980: 53.

Rhodacarus haarlovi.— Karg, 1993: 333.

Rhodacarus roseus.— Haarlov, 1957: 19 (misidentification, according to Shcherbak, 1977:

79).



TYPE LOCALITY AND HABITAT: Lithuania, Klaipeda County, 25 September 1964, soil in

sugar beet field [Chenopodiaceae].

132. Rhodacarus laureti Athias-Henriot, 1961

Rhodacarus laureti Athias-Henriot, 1961: 501.

Rhodacarus laureti.— Lee, 1970: 29; Karg, 1993: 335.

Rhodacarus (Rhodacarus) laureti.— Shcherbak, 1980: 46.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, l'Oued Bouzaréah valley, 3 January 1961, in

soil under Laurus nobilis [Lauraceae].

NOTE: Bregetova and Shcherbak (1977): 269 suggested that R. roseus sensu Sheals (1958)

could be a misidentification of R. laureti.

133. Rhodacarus longisetosus Shcherbak, 1980

Rhodacarus (Rhodacarus) furmanae longisetosus Shcherbak, 1980: 64.

Rhodacarus longisetosus.— Karg, 1993: 332.



TYPE LOCALITY AND HABITAT: Ukraine, Alushta, Crimea, 16 November 1960, in soil

under stone.

134. Rhodacarus mandibularis Berlese, 1920

Rhodacarus mandibularis Berlese, 1920: 165.

Rhodacarus mandibularis.— Lee, 1970: 29; Bregetova and Shcherbak, 1977b: 261;

Shcherbak, 1977: 74; Castagnoli and Pegazzano, 1985: 239; Karg, 1993: 333.

94

Rhodacarus (Rhodacarus) mandibularis.— Shcherbak, 1980: 48.


TYPE LOCALITY AND HABITAT: Italy, Florence and Udine, in moss and humus.

NOTE: Sheals (1958) considered this species to be a junior synonym of R. roseus. However,

in subsequent publications they have been considered as different species (e.g.

Shcherbak, 1980; Karg, 1993). We agree with this conclusion, on the basis of the

redescriptions of R. roseus by Lee (1970) and of R. mandibularis by Shcherbak

(1980).

135. Rhodacarus mandibularosimilis Shcherbak and Kadite, 1979

Rhodacarus mandibularosimilis Shcherbak and Kadite, 1979: 84.

Rhodacarus (Rhodacarus) mandibularosimilis.— Shcherbak, 1980: 55.

Rhodacarus mandibularosimilis.— Karg, 1993: 333.



TYPE LOCALITY AND HABITAT: Ukraine, Lyutezh, Kiev Region, in soil (0-5 cm) of

mixed forest.

136. Rhodacarus marksae Domrow, 1957

Rhodacarus marksae Domrow, 1957: 200.

Rhodacarus marksae.— Lee, 1970: 29.

TYPE DEPOSITORY: Queensland Museum, Brisbane, Australia.

TYPE LOCALITY AND HABITAT: Australia, Low Isles [Great Barrier Reef, Queensland],

Green Ant Island, 24 August 1954, on leaf mould.

137. Rhodacarus matatlanticae Castilho and Moraes, 2010

Rhodacarus matatlanticae Castilho and Moraes, 2010: 394.



Paulo, Brazil.

TYPE LOCALITY AND HABITAT: Brazil, Cananéia, São Paulo, 12 July 2000, in soil under

Bactris setosa [Arecaceae].

138. Rhodacarus olgae Shcherbak, 1975

95

Rhodacarus olgae Shcherbak, in Shcherbak and Furman, 1975: 47.

Rhodacarus olgae.— Bregetova and Shcherbak, 1977b: 264; Karg, 1993: 332.

Rhodacarus (Rhodacarus) olgae.— Shcherbak, 1980: 59; Shcherbak, 1982: 63.



TYPE LOCALITY AND HABITAT: Ukraine, Shelekhovo, Ananievskiy District, Odessa

Region, 19 September 1965, in forest soil (depth of 10-20 cm).

139. Rhodacarus pallidus Hull, 1918

Rhodacarus pallidus Hull, 1918: 57.

Rhodacarus roseus var. pallidus.— Halbert, 1920: 115.

Rhodacarus pallidus f. typica Sheals, 1958: 299.

Rhodacarus (?) pallidus.— Athias-Henriot, 1961: 501.

Rhodacarus pallidus.— Lee, 1970: 29; Bregetova and Shcherbak, 1977b: 267; Krantz, 1986:

634; Karg, 1993: 334; Farrier and Hennessey, 1993: 133.

Rhodacarus (Rhodacarus) pallidus.— Shcherbak, 1980: 44.


TYPE LOCALITY AND HABITAT: England, Allendale (cited as West Allendale),

Tynedale, [Northumberland], under deeply embedded stones with Pergamasus

hamatus [Acari: Parasitidae].

NOTE1: Sheals (1958) considered R. calcarulatus to be a junior synonym of Rhodacarus

pallidus Hull, naming it Rhodacarus pallidus f. calcarulatus. In the same

publication, he mentioned that T. pallidus had a V-shaped groove posterior to setae

j4, z3 and s2, and that R. pallidus f. calcarulatus did not have it. Thus, we understand

that those species should not be considered synonyms. Krantz (1986) drew attention

to possible misidentifications of this species, and ambiguity over its doubtful

synonymy with R. calcarulatus, as suggested by Sheals (1958).

NOTE2: The species reported by Willmann (1935): 422 as Rhodacarus pallidus Hull was

described as Rhodacarus willmanni Karg, 1993: 333.

140. Rhodacarus reconditus Athias-Henriot, 1961

Rhodacarus reconditus Athias-Henriot, 1961: 503.

Rhodacarus reconditus.— Lee, 1970: 29; Bregetova and Shcherbak, 1977b: 261; Karg, 1993:

333.

96

Rhodacarus (Rhodacarus) reconditus.— Shcherbak, 1980: 57.

Rhodacarus calcarulatus.— Athias-Henriot, 1961: 501; Karg, 1971: 321 (misidentification).


Paris, France.

TYPE LOCALITY AND HABITAT: Spain, Isla Cies Norte, 26 July 1955, under cut Ulex

europaeus [Fabaceae].

141. Rhodacarus rhodacaropsis Ryke, 1962

Rhodacarus rhodacaropsis Ryke, 1962c: 83.

Rhodacarus (Rhodacarus) rhodacaropsis.— Loots, 1969a: 50.

Rhodacarus rhodacaropsis.— Lee, 1970: 29.



TYPE LOCALITY AND HABITAT: South Africa, Potchefstroom, March 1960, in humus

under trees on river bank.

NOTE: This species is only provisionally placed in this genus.

142. Rhodacarus roseus Oudemans, 1902

Rhodacarus roseus Oudemans, 1902: 50.

Genus and species unknown, number 81.— Oudemans, 1896: 136.

Rhodacarus roseus f. typica Sheals, 1958: 299.

Rhodacarus roseus.— Athias-Henriot, 1961: 498; Hirschmann, 1962: 49; Lee, 1970: 29;

Karg, 1971: 322; Lee, 1973a: 2; Lee, 1973b: 141; Bregetova and Shcherbak, 1977b:

267; Evans and Till, 1979: 206; Zaher, 1986: 16; Krantz, 1986: 634; Fouly and

Nawar, 1990: 336; Krantz and Ainscough, 1990: 618; Farrier and Hennessey, 1993:

133; Karg, 1993: 334.

Rhodacarus (Rhodacarus) roseus.— Shcherbak, 1980: 43.

TYPE DEPOSITORY: National Museum of Natural History – Naturalis, Leiden, Netherlands.

TYPE LOCALITY AND HABITAT: Netherlands, Haarlem, on decaying leaves.

NOTES: Sheals (1958) recognised two forms of Rhodacarus roseus - R. roseus f. typica, with

a V-shaped groove on the podonotal shield, and R. roseus f. simplex, without a

groove. Sheals' R. roseus f. typica appears to match R. roseus, but his R. roseus f.

simplex remains unidentified. Schweizer (1961) illustrated R. roseus without a

podonotal groove, so the identity of those specimens must also be considered as

97

doubtful. The name Rhodacarus roseus has been used in a large number of

publications reporting faunistic and ecological studies of soil arthropods (e.g. Block,

1966; Costa, 1966; Lee, 1973b; Sardar and Murphy, 1987). In view of the difficulty

of identifying R. roseus and related species, it seems likely that this name has been

applied to a number of different species. Bregetova and Shcherbak (1977) suggested

that R. roseus sensu Sheals (1958) could be a misidentification of R. laureti. Sheals

(1958) also considered Rhodacarus roseus to be a senior synonym of R.

mandibularis. However, we follow other recent authors in considering these to be

two different species (see under R. mandibularis).

143. Rhodacarus salarius Karg, 1993

Rhodacarus salarius Karg, 1993: 334.



TYPE LOCALITY AND HABITAT: Germany, Simonsberg, near Husum, [Nordfriesland,

Schleswig-Holstein], 7 December 1987, in salt marshes on the North Sea coast, 4-8

cm deep.

144. Rhodacarus solimani Fouly and Nawar, 1990

Rhodacarus solimani Fouly and Nawar, 1990: 337.

TYPE DEPOSITORY: Agricultural Zoology Department, Faculty of Agriculture, Cairo

University, Giza, Egypt.

TYPE LOCALITY AND HABITAT: Egypt, farm of the Faculty of Agriculture, Cairo

University, Giza, in debris under pear trees [Rosaceae].

145. Rhodacarus strenzkei Willmann, 1957

Rhodacarus strenzkei Willmann, 1957: 166.

Rhodacarus strenzkei.— Athias-Henriot, 1961: 498; Karg, 1971: 322; Bregetova and

Shcherbak, 1977b: 265; Krantz, 1986: 634; Karg, 1993: 332.

Rhodacarus stenzkei [sic].— Lee, 1970: 29.

Rhodacarus (Rhodacarus) strenzkei.— Shcherbak, 1980: 56.


TYPE LOCALITY AND HABITAT: Germany, margin of Eider River, Kiel, in soil.

98

NOTE: Lombardini (1962) described a small variety of this species from Italy, as

"Rhodacarus strenzkei Willmann var. strictior n. var.". The name strictior is not

available from this paper, because it is infrasubspecific (International Code of

Zoological Nomenclature, Articles 15.2 and 45.6.3). It has not been used by later

authors, so it remains unavailable.

146. Rhodacarus tribaculatus Athias-Henriot, 1961

Rhodacarus tribaculatus Athias-Henriot, 1961: 501.

Rhodacarus tribaculatus.— Lee, 1970: 29; Bregetova and Shcherbak, 1977b: 269; Karg,

1993: 335; Kalúz, 1994: 675.

Rhodacarus (Rhodacarus) tribaculatus.— Shcherbak, 1980: 68.


Paris, France.

TYPE LOCALITY AND HABITAT: France, Port Provençal, Ajaccio, Corsica, April 1957, in

litter of Cistus sp. (Cistaceae).

147. Rhodacarus willmanni Karg, 1993

Rhodacarus willmanni Karg, 1993: 333.

Rhodacarus pallidus.— Willmann, 1935: 422; Karg, 1971: 320 (misidentification).

Rhodacarus roseus var. pallidus.— Willmann, 1935: 422 (misidentification).


TYPE LOCALITY AND HABITAT: Germany, Kiel Harbour, in coastal groundwater.

NOTE: This species was described on the basis of the specimens previously identified by

Willmann (1935): 422 as Rhodacarus pallidus Hull.

148. Rhodacarus zaheri Fouly and Nawar, 1990

Rhodacarus zaheri Fouly and Nawar, 1990: 336.

TYPE DEPOSITORY: Agricultural Zoology Department, Faculty of Agriculture, Cairo

University, Giza, Egypt.


University, Giza, in debris under pear trees [Rosaceae].

99

2.4 Discussion

We have attempted to compile a complete list of all species of the Rhodacaridae, and

to place them in a coherent generic classification. We may have missed some species that are

presently placed in other families and should be transferred to the Rhodacaridae, or were

published in papers that we have overlooked. However, we believe that the present catalogue

will provide a useful foundation for future taxonomic research, and will help in the

identification of the Rhodacaridae that are frequently collected in ecological studies of the soil

fauna.

In this compilation we draw attention to some areas where detailed research will be

necessary to resolve taxonomic problems. Taxonomic confusion surrounds some groups of

species, especially in the genus Rhodacarus. It appears that Rhodacarus calcarulatus, R.

coronatus, R. pallidus, R. reconditus and R. roseus have often been misidentified, and all of

these species are in need of detailed revision. It is very likely that some published records of

R. roseus actually refer to other species, given the several citations of this species on different

continents despite the insufficient information available about the morphology of this species

(LEE, 1973b). Bregetova and Shcherbak (1977) drew attention to the very subtle differences

that separate some species, including R. clavulatus, R. ancorae, R. pallidus, and R. laureti,

and their possible misidentification.

Most species of Rhodacarus have a transverse V-shaped groove across the podonotal

shield (e.g. R. roseus, R. pallidus) while others do not (e.g. R. calcarulatus, R. tribaculatus).

Bregetova and Shcherbak (1977) used this character in their key to species, and Kalúz (1994)

recognised the roseus-group, with a podonotal groove, and the calcarulatus-group, without a

groove. It is not clear whether this character has any real taxonomic value, and if so, at what

level.

Rhodacarellus liuzhiyingi and Rhodacarellus yalujiangensis are the only rhodacarid

species with the peritreme extending anteriorly to the median region of coxa I; in other

species, it reaches at most the median region of coxa II. The median notch at the anterior

margin of the opisthonotal shield, the longer Z5 and S5 in relation to other opisthonotal setae,

the paired lightly sclerotised and punctate anterior areas of the sternal shield and the elongate

and medially constricted ventrianal shield suggest these species to belong to Dendrolaelaps

(Digamasellidae). Conclusive placement of these species requires the determination of the

number of tines of the palp tarsal claw and the number of setae on genu and tibia IV, which

are not described in the original descriptions of the species. Rhodacarellus shandongensis

100

shares most of those characteristics, except that the peritreme is longer, reaching the median

region of coxa III; also for this species, characteristics of of the palp tarsal claw and setal

counts are not known. When these three species are examined more closely, we believe that

they will be transferred to the Digamasellidae.

There are many published records of rhodacarids identified only to the genus level,

including, among others, Rhodacarellus sp., Rhodacaroides sp., and Rhodacarus sp., in

records compiled by Farrier and Hennessey (1993), and Rhodacaroides new species from

Australia (HEATWOLE; DONE; CAMERON, 1981). A future taxonomic revision of the

family should include identification of these taxa. We have also commented on some cases of

possible synonymy that have not been resolved, and on some species for which the adult

female is not yet known. There are some questions about the familial placement of some

genera, especially Paragamasellevans and Tangaroellus, and these would benefit from further

study. A convincing phylogenetically-based classification of the Rhodacaroidea will only be

possible when all the speces listed here have been re-examined in more detail than they have

in the past.

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3 CATALOGUE OF THE MITE FAMILIES DIGAMASELLIDAE,

LAELAPTONYSSIDAE, OLOGAMASIDAE AND TERANYSSIDAE (ACARI:

RHODACAROIDEA: MESOSTIGMATA), AND A KEY TO THE WORLD GENERA

OF THESE FAMILIES

Abstract

The taxonomic concepts of Digamasellidae Evans, Laelaptonyssidae Womersley,

Ologamasidae Ryke and Teranyssidae Halliday, families belonging to Rhodacaroidea, are

confusing and have changed considerably over time, making it difficult to determine to which

family a determined species of this group belongs. A brief historic review of the literature on

the classification of these families is presented. A dichotomous key to the genera is provided,

derived from a standardised database of character states. A list of species within each genus

was updated and complemented, giving relevant taxonomic information about the respective

types, and providing references to nomenclatural changes, synonymy and redescriptions of

each species. In total, 277 digamasellid species and one subspecies arranged in 11 genera,

eight laelaptonyssid species arranged in one genus, 450 ologamasid species arranged in 44

genera and one teranyssid species are listed in this work.

Keywords: Catalogue; Soil mites; Taxonomy

Resumo

Os conceitos taxonômicos de Digamasellidae Evans, Laelaptonyssidae Womersley,

Ologamasidae Ryke e Teranyssidae Halliday, famílias pertencentes a Rhodacaroidea, são

confusos e têm mudado consideravelmente ao longo do tempo, tornando difícil determinar a

qual família pertence uma determinada espécie deste grupo. Uma breve revisão histórica da

literatura sobre a classificação dessas famílias é apresentada. Uma chave dicotômica para os

gêneros é fornecida, derivada de um banco de dados padronizado da caracterização das

espécies. Uma lista de espécies de cada gênero foi atualizada e complementada, fornecendo

informações taxonômicas relevantes sobre os respectivos tipos, e fornecendo referências a

mudanças nomenclaturais, sinonímias e redescrições de cada espécie. No total, 277 espécies

de Digamasellidae arranjadas em 11 gêneros, oito espécies de Laelaptonyssidae arranjadas em

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um gênero, 450 espécies de Ologamasidae arranjadas em 44 gêneros e uma espécie de

Teranyssidae estão listadas neste trabalho.

Palavras-chave: Catálogo; Ácaros de solo; Taxonomia

3.1 Introduction

The mite families Digamasellidae Evans, Halolaelapidae Karg, Laelaptonyssidae

Womersley, Ologamasidae Ryke, Rhodacaridae Oudemans and Teranyssidae Halliday are

considered today to be the member of the superfamily Rhodacaroidea (LINDQUIST;

KRANTZ; WALTER, 2009).

The taxonomic concept of the families belonging to Rhodacaroidea are confusing and

have changed considerably over time, making it difficult to determine which family belongs a

determined species of this group. Most descriptions of species and other works on this group

referred and still refers to most species of this group as Rhodacaridae.

Krantz (1978) named and characterised the Rhodacaroidea for the first time, including

in this superfamily the families Digamasellidae, Ologamasidae and Rhodacaridae. The

superfamily was considered to consist of mites with undivided sternal shield that generally

bearing four pairs of setae, genital shield usually rounded anteriorly and not fused with the

ventrianal shield. It was considered that in some Rhodacaroidea seta st4 could be inserted in

the unsclerotised integument and seta st1 could be inserted on a weakly defined

anteromarginal extension of the sternal shield.

Lindquist; Krantz and Walter (2009) provided an expanded concept of the

Rhodacaroidea including the Digamasellidae, Halolaelapidae, Laelaptonyssidae,

Ologamasidae, Rhodacaridae and Teranyssidae. Dowling and OConnor (2010) proposed an

alternative hypothesis in which the Rhodacaroidea as understood here would be paraphyletic.

However, until formal taxonomic changes are made, we have used the classification of

Lindquist; Krantz and Walter (2009).

Rhodacaridae was the first supra-generic taxon of Rhodacaroidea created, when

Oudemans (1902a) established Rhodacarinae as a subfamily of Parasitidae Oudemans, to

contain his new genus and species Rhodacarus roseus Oudemans, 1902. Halbert (1915) raised

the subfamily to family level. As understood today, Rhodacaridae contains 158 described

species distributed in 15 genera (CASTILHO; MORAES; HALLIDAY, 2012).

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Evans (1957) created and characterised the Digamasellidae as a family whose

chelicerae are dentate; palp tarsal claw two-tined; epistome produced into a long median

process; dorsal shield in both sexes completely divided into two shields of approximately

equal sizes and with more than 23 pairs of setae; female genital shield female truncated

posteriorly or very slightly convex, with no more than one pair of setae and lying close to a

ventrianal shield or remote from an anal shield; male with sternogenital shield and a separate

ventrianal or anal shield; leg II in the male spurred; setae added to the shield (or shields) in

the deutonymphal stage. He included in this family Asca von Heyden, 1826 and Digamasellus

Berlese, 1905b.

Digamasellidae presently contains 277 described species distributed in the following

11 genera: Dendrolaelaps Halbert, 1915, Dendrolaelaspis Lindquist, 1975, Dendroseius

Karg, 1965, Digamasellus, Insectolaelaps Shcherbak, 1980, Longoseius Chant, 1961,

Multidendrolaelaps Hirschmann, 1974, Oligodentatus Shcherbak, 1980, Orientolaelaps

Bregetova and Shcherbak, 1977b, Panteniphis Willmann, 1949, Pontiolaelaps Luxton, 1989.

Revisionary works referring to mites of this group were published by Leitner (1949),

Hirschmann (1960), Karg (1971), Linquist (1975), Shcherbak (1980) and Hirschmann and

Wiśniewski (1982).

Hirschmann and Wiśniewski (1982, 1989b, 1989c, 1989d, 1990, 1991b, 1993b,

1993c) divided Dendrolaelaps into 14 sub-genera. These are not considered further in this

work, because it is not possible to unambiguously assign every species in the genus to the

respective sub-genera.

Digamasellidae mites have been found in soil, litter, on Coleoptera or in galleries

made by them in tree trunks, rodent nests, and other types of organic matter in contact with

the soil. Digamasellids are commonly mentioned in the literature as predators, which some

species have been found to feed on first stages of Coleoptera, Collembola, nematodes and

mites (KARG, 1971; MOSER, 1975; ENDA; TAMURA, 1977; ABOU-EL-SOUD; SHOEIB,

2000).

Ryke (1962c) included in Rhodacaridae all genera of Mesostigmata whose

deutonymphs had separate podonotal and opisthonotal shields, and palp tarsal claw two- or

three-tined. He divided the family into two subfamilies, Rhodacarinae, containing species

whose adults also had separate podonotal and opisthonotal dorsal shields, and a new

subfamily, Ologamasinae, whose adults had these shields fused. In this concept, the subfamily

Ologamasinae included Antennolaelaps Womersley, 1956a, Epiphis Berlese, 1916b,

Gamasiphis Berlese, 1904, Gamasiphoides Womersley, 1956b, Gamasitus Womersley,

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1956b, Hydrogamasus Berlese, 1892c, Laelaptiella Womersley, 1956b, Megaliphis

Willmann, 1938, Micriphis Berlese, 1914, Neogamasiphis Trägårdh, 1952, Ologamasus

Berlese, 1888, Onchogamasus Womersley, 1956a, Pachyseius Berlese, 1910b, Parasitiphis

Womersley, 1956b, Periphis Berlese, 1914, Physallolaelaps Berlese, 1908,

Queenslandolaelaps Womersley, 1956a, Sessiluncus G. Canestrini, and Trachygamasus

Berlese, 1906.

Ologamasidae presently contains 450 described species distributed in the following 44

genera: Acugamasus Lee, 1970, Acuphis Karg, 1998, Allogamasellus Athias-Henriot, 1961a,

Antennolaelaps, Athiasella Lee, 1973, Caliphis Lee, 1970, Cymiphis Lee, 1970, Cyrtolaelaps

Berlese, 1887b, Desectophis Karg, 2003, Euepicrius Womersley, 1942, Euryparasitus

Oudemans, 1902a, Evanssellus Ryke, 1961a, Gamasellevans Loots and Ryke, 1967a,

Gamaselliphis Ryke, 1961b, Gamasellopsis Loots and Ryke, 1966, Gamasellus Berlese,

1892f, Gamasiphis, Gamasiphoides, Gamasitus, Geogamasus Lee, 1970, Heterogamasus

Trägårdh, 1907, Heydeniella Richters, 1907, Hiniphis Lee, 1970, Hydrogamasellus

Hirschmann, 1966b, Hydrogamasus, Laelaptiella, Laelogamasus Berlese, 1905a,

Litogamasus Lee, 1970, Neogamasellevans Loots and Ryke, 1967b, Notogamasellus Loots

and Ryke, 1965, Ologamasus, Onchogamasus, Oriflammella Halliday, 2008, Parasitiphis,

Periseius Womersley, 1961, Pilellus Lee, 1970, Podonotogamasellus Loots and Ryke, 1965,

Pyriphis Lee, 1970, Queenslandolaelaps, Rhodacaroides Willmann, 1959, Rykellus Lee,

1970, Sessiluncus, Solugamasus Lee, 1973 and Stylochirus G. Canestrini and R. Canestrini,

1882.

Lee (1970) conducted an extensive revision of the genera placed today in

Ologamasidae, but he considered them all as members of Rhodacaridae. The family was

divided into six subfamilies, three of which have genera considered to belong to

Ologamasidae as the family is here understood: Gamasiphinae, containing Caliphis,

Euepicrius, Gamaselliphis, Gamasiphis, Gamasiphoides, Hydrogamasus and Laelaptiella;

Ologamasinae, containing Acugamasus, Allogamasellus, Cymiphis, Cyrtolaelaps,

Euryparasitus, Evanssellus, Gamasellus, Geogamasus, Heterogamasus, Heydeniella,

Hiniphis, Hydrogamasellus, Laelogamasus, Litogamasus, Neogamasellevans,

Notogamasellus, Ologamasus, Parasitiphis, Periseius, Pilellus, Pyriphis, Rhodacaroides and

Rykellus; and Sessiluncinae, containing Antennolaelaps, Gamasellevans, Gamasellopsis,

Gamasitus, Onchogamasus, Paragamasellevans, Queenslandolaelaps, Sessiluncus and

Stylochirus. The other subfamilies were Rhodacarinae, containing Afrogamasellus,

Rhodacarellus, Rhodacaropsis and Rhodacarus (today genera of Rhodacaridae);

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Laelaptonyssinae, containing Laelaptonyssus Womersley, 1956b (today in Laelaptonyssidae);

and Tangaroellinae, containing Tangaroellus Luxton, 1968 (today genus incertae sedis).

A first list of species of Ologamasidae was presented by Silva (2007). He included in

this family 383 species arranged in 45 genera.

Ologamasidae mites are found in soil, litter, mosses, lichens, rodent nests, and other

types of organic matter in contact with the soil. These are some of the most common

Mesostigmata in the soils of São Paulo State, Brazil (MINEIRO; MORAES, 2001; SILVA;

MORAES; KRANTZ, 2004). Ologamasids are commonly mentioned in the literature as

predators, some species having been found to feed on Collembola, nematodes and mites

(KARG, 1971; LEE, 1974; WALTER; HUNT; ELLIOTT, 1988; BEAULIEU; WALTER,

2007).

Womersley (1956b) erected the new family Laelaptonyssidae, with the new genus

Laelaptonyssus. Lee (1970) and Halliday (1987) placed Laelaptonyssus in the Rhodacaridae,

in a subfamily Laelaptonyssinae. However, Halliday (1998) and Krantz (2000) reverted to

original usage of the family Laelaptonyssidae. Castilho; Moraes and Halliday (2012) verified

that Laelaptonyssus is junior synonym of Starkovia Lombardini, 1947. So actually this family

contains eight species in a single genus, Starkovia.

Mites of this family were described from termite nests. Starkovia mites are often

found riding on the heads or bodies of termites, dying soon after the death of their hosts

(WANG; POWER; O‘CONNOR, 2002), which suggests an intimate relationship, but the

details of that relationship is not known. Krantz (2000) suggested Starkovia to be phoretic on

termites rather than parasitic, feeding on nematodes in the termites' nest material.

Halliday (2006) erected Teranyssidae, to contain his new genus and species

Teranyssus howardensis Halliday, 2006. T. howardensis is presently the single species of the

family. It was also described from termite nests, but the nature of this relationship between the

mite and the termites is not known.

The last family to be included in Rhodacaroidea was Halolaelapidae, by Lindquist;

Krantz and Walter (2009). This family consists of 80 species in four genera. This group is in

need of a detailed revision to determine whether or not it should be included as

Rhodacaroidea, mainly because seta st4 of members of this family is inserted on the soft

integument rather than on the sternal shield, a characteristic shared by all other

Rhodacaroidea, except Oriflammella. Therefore, Halolaelapidae is not considered in this

study.

126

The purpose of this work is to review the literature concerning the taxonomy of

Digamasellidae, Laelaptonyssidae, Ologamasidae and Teranyssidae, to construct a key for the

identification of genera of these groups, as well as to update and complement the lists of the

species considered to belong to the these families, with relevant taxonomic information about

them.

3.2 Materials and Methods

This chapter includes papers published up to December 2011. The search for

information was initiated by considering the literature available in the reprint collection of

Prof. C.H.W. Flechtmann and Prof. G.J. de Moraes, of Departamento de Entomologia e

Acarologia, Escola Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de

São Paulo (USP). New references were detected by searches in electronic databases and by

the evaluation of references listed in each paper available. Many publications were obtained

with the help of the librarian of ‖Biblioteca Central, ESALQ-USP‖, as well as by direct

contacts with authors and collaborators from different countries. Of fundamental importance

in the initial part of the work were the publications of Lee (1970), Silva (2007) and the

Rhodacaroidea database of Hallan (2005); the latter was consulted periodically between

March 2006 and June 2011.

After obtaining copies of all the literature cited in this publication, we constructed a

spreadsheet in an attempt to standardize the morphological characteristics of each of the

species considered to belong to Digamasellidae, Laelaptonyssidae, Ologamasidae and

Teranyssidae, based on information from the respective descriptions and/or redescriptions.

Missing information was added as much as possible to build a standard set of characters. This

analysis made it necessary to move some species from one genus to another. The final

spreadsheet was then used to prepare a dichotomous key to the genera.

The list of species proper includes all members of Digamasellidae, Laelaptonyssidae,

Ologamasidae and Teranyssidae, presented in alphabetical order within each taxonomic

group. We have not used intermediate categories such as subfamily, subgenus, or species

groups, because some species are not described in sufficient detail to allow these decisions to

be taken. For each genus, the following information is provided:

Name and author;

Original designation of the genus (even if described at subgeneric level), author, year

of the original description, page on which the description begins, family in which the

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genus was initially placed, type species, and further references to descriptions or re-

descriptions of the genus;

Synonyms, each followed by its author, page on which the corresponding original

description started, family in which the genus was initially placed, type species,

reference to the paper in which each corresponding synonymy was established, type

species, and further references providing descriptive information about the junior

synonym.

For each species, the following information is provided:

Current generic combination of the species, with its author and date of description;

Name of the species in its original combination, with reference to the author, date, and

the page on which the description started;

References to subsequent literature on the species, including different combinations or

variations of the name, and including publications that provide information on the

morphology of the species, other than the original description;

Synonyms, each followed by its author, date, and page number, and the reference in

which the synonymy was established;

Type depository, the institution where the name-bearing type specimens are deposited;

Type locality (first mentioning the country – to be more geographically informative,

Saint Helena, Galapagos Islands, Puerto Rico and Hawaii were mentioned as such,

without referring to the country to which they belong – followed by the location

within the country, from the more specific to the more general geographic

information); and habitat in which the type specimens were collected. In some cases

we have added complementary locality information, in square brackets.

Occasionally a note is added after the details of a genus or species, to explain unusual

or complicated taxonomic or nomenclatural problems.

The terminology used for anatomical structures is that of Evans and Till (1979). Lee

(1970), Shcherbak (1982a) and Hirschmann and Wiśniewski (1983) used different systems of

notation for the dorsal idiosomal chaetotaxy of Rhodacaroidea, but we have used the widely

accepted system developed by Lindquist and Evans (1965) and Lindquist (1994).

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3. 3 Results

3.3.1 Key to world families of Rhodacaroidea and genera of Digamasellidae,

Laelaptonyssidae, Ologamasidae and Teranyssidae (adult females)

1. Seta st4 on metasternal shield or on soft integument .................................. Halolaelapidae

- Seta st4 on sternal shield [if on metasternal platelets (Oriflammella, Ologamasidae),

dorsal shield setae strongly pilose and set on long stalks] ................................................ 2

2. With anal shield, without preanal setae ..................... Teranyssidae – Teranyssus Halliday

- With ventrianal shield, with 1-9 pairs of preanal setae .................................................. ...3

3. With attenuate chelicera; palp with 4 segments (fused tibia and tarsus) .............................

.......................................................................... Laelaptonyssidae – Starkovia Lombardini

- With normal chelicera, not attenuate; palp with 5 segments .......................................... ...4

4. Palp tarsal claw two-tined; genu and tibia IV with six or eight setae .................................

........................................................................................................ Digamasellidae……..6

- Palp tarsal claw three-tined; genu and tibia IV with nine or ten setae .............................. 5

5. Usually without densely sclerotised strutuctures between setae j5 and j6 (scleronoduli);

podonotal and opisthonotal shields fused or not; without desclerotised bands of punctate

integument ...................................................................................... Ologamasidae……..16

- With scleronoduli (except in Interrhodeus, Protogamasellopsis and Pennarhodeus);

podonotal and opisthonotal shields not fused (except in Afrogamasellus luberoensis);

usually with desclerotised bands of punctate integument along posterior margin of

podonotal, anterior margin of opisthonotal, anterior margin of sternal, posterior margin

of genital and anterior margin of ventrianal shields .................................... Rhodacaridae

6. Podonotal and opisthonotal shields fused ...................................... Panteniphis Willmann

- Podonotal and opisthonotal shields totally separated ...................................................... 7

7. Seta Zv1 on ventrianal shield; anal opening conspicuously enlarged, longer than 1/5 the

length of the ventrianal shield ........................................................ Digamasellus Berlese

- Seta Zv1 on unsclerotised integument next to anterior margin of ventrianal shield; anal

opening not enlarged, shorter than 1/5 the length of the ventrianal shield ...................... 8

8. Posterior margin of idiosoma bilobed ..................................... Dendrolaelaspis Lindquist

- Posterior margin of idiosoma rounded ............................................................................ 9

9. Epistome with anterior region triangular; podonotal shield with 23 pairs of setae (seta r6

present) ............................................................ Orientolaelaps Bregetova and Shcherbak

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- Epistome with two or three anterior extensions; podonotal shield with less than 23 pairs

of setae (seta r6 absent) .................................................................................................. 10

10. Movable cheliceral digit with at least five teeth ............................................................ 11

- Movable cheliceral digit with 1-4 teeth ......................................................................... 13

11. Scleronoduli absent ......................................................................... Pontiolaelaps Luxton

- Scleronoduli present ....................................................................................................... 12

12. Spermatheca opening at base of coxa III ....................... Multidendrolaelaps Hirschmann

- Spermatheca opening at base of coxa IV .................................. Insectolaelaps Shcherbak

13. Movable cheliceral digit with three teeth; anterior margin of the opisthonotal shield

without median notch ..................................................................................................... 14

- Movable cheliceral digit with one, two or four teeth; anterior margin of the opisthonotal

shield usually with median notch ................................................................................... 15

14. With seven rows of deutosternal denticles; setae r4 and r5 on unsclerotised integument

along lateral margins of podonotal shield ............................................. Dendroseius Karg

- With six rows of deutosternal denticles; setae r4 and r5 on podonotal shield....................

. .................................................................................................. Oligodentatus Shcherbak

15. Trochanter III with five setae; genu and tibia III with eight or nine setae; basitarsi II and

III each with four setae, and basitarsi IV usually with three or four setae .........................

…………………………………………………………………….Dendrolaelaps Halbert

- Trochanter III with four setae; genu and tibia III with seven setae or less; basitarsi II and

III each with two or three setae, and basitarsi IV usually with one to three setae

............................................................................................................... Longoseius Chant

16. Podonotal and opisthonotal shields totally separated; epistome never with club-shaped

antero-medial extension; peritremal shield fused or not fused posteriorly to exopodal

shield [if fused, connection between them wider than width of peritreme] ................... 17

- Podonotal and opisthonotal shields fused [if not fused (some Geogamasus), epistome

with club-shaped antero-medial extension, and peritremal and exopodal shields fused by

a narrow connection posterior to stigma] ....................................................................... 37

17. Dorsal shield setae set on long stalks .............................................. Oriflamellla Halliday

- Dorsal shield setae not set on stalks ............................................................................... 18

18. All dorsal idiosomal setae densely pilose ...................................... Laelogamasus Berlese

- Dorsal idiosomal setae not densely pilose ..................................................................... 19

19. Opisthonotal and ventrianal shields fused ..................................................................... 20

- Opisthonotal and ventrianal shields not fused ............................................................... 24

130

20. Presternal plates absent and pretarsus I absent .............................. Euepicrius Womersley

- Presternal plates present and pretarsus I present ........................................................... 21

21. With one pair of presternal plates ...................................................... Gamaselliphis Ryke

- With two pairs of presternal plates ................................................................................ 22

22. Opithogastric region without latero-diagonal fissure; peritreme extending anteriorly to

anterior margin of coxa II .............................................................................. Hiniphis Lee

- Opithogastric region with a distinct latero-diagonal fissure; peritreme extending

anteriorly to region of coxa I ......................................................................................... 23

23. Peritremal shield fused only to exopodal shield near coxa IV .............. Desectophis Karg

- Peritremal shield fused to exopodal shield near coxa IV and ventrianal shield ................

………………………………………………………………………………Acuphis Karg

24. Dorsal podosomal region with at least 28 pairs of setae .............................................. 25

- Dorsal podosomal region with less than 25 pairs of setae ............................................. 27

25. Dorsal opisthosomal region with more than 30 pairs of setae ........................ Pilellus Lee

- Dorsal opisthosomal region with less than 21 pairs of setae ......................................... 26

26. Podonotal shield with 32 pairs and one odd seta (posterior and mediad to j3); trocanther

IV with six, genu III with ten and tibia III with nine setae……………………………….

....................................................................................... Notogamasellus Loots and Ryke

- Podonotal shield with 28 pairs of setae; trocanther IV with five, genu III with nine and

tibia III with eight setae ..........................................Podonotogamasellus Loots and Ryke

27. Peritreme extending anteriorly to median region of coxa III ............................................

……………………………………………………………Allogamasellus Athias-Henriot

- Peritreme extending anteriorly at least to median region of coxa II ............................. 28

28. Seta j1 distinctly longer than r3 and Z5 ..................................... Heterogamasus Trägårdh

- Seta j1 shorter than r3 and Z5 ........................................................................................ 29

29. Without presternal plates. ................................................................. Cyrtolaelaps Berlese

- With 1-5 pairs of presternal plates. ................................................................................ 30

30. With one pair of presternal plates [if with two pairs (Rhodacaroides aegypticus),

peritremal shield not fused with exopodal shield near coxa IV] ................................... 31

- With 2-5 pairs of presternal plates [if with two pairs, peritremal shield fused with

exopodal shield near coxa IV] ....................................................................................... 34

31. Seta j1 on prominent protuberance. ....................................................... Evanssellus Ryke

- Seta j1 not set on protuberance ...................................................................................... 32

32. Dorsal shields with some setae pilose and/ or spatulate ......................... Acugamasus Lee

131

- Dorsal shields without pilose or spatulate setae ............................................................. 33

33. Peritremal shield not fused with exopodal shield near coxa IV .........................................

…………………………………………………………………Rhodacaroides Willmann

- Peritremal shield fused with exopodal shield near coxa IV ...... Euryparasitus Oudemans

34. Sternal shield fused with endopodal shield near coxa IV ................ Periseius Womersley

- Sternal shield not fused with endopodal shield near coxa IV ........................................ 35

35. Dorsal opisthosomal region with 12 pairs of setae ................................ Solugamasus Lee

- Dorsal opisthosomal region with more than 12 pairs of setae ....................................... 36

36. With 2-3 pairs of presternal plates ..................................................... Gamasellus Berlese

- With five pairs of presternal plates ........................................................ Litogamasus Lee

37. With a distinct and complete line of fusion between podonotal and opisthonotal shields

........................................................................................................................................ 38

- Line of fusion between podonotal and opisthonotal shields generally indistinct [if

distinct (Onchogamasus communis), then interrupted between setae j6] ...................... 39

38. Line of fusion between podonotal and opisthonotal shields straight; peritremal shield

and exopodal shield near coxa IV not fused ................... Gamasellevans Loots and Ryke

- Line of fusion between podonotal and opisthonotal shields V-shaped; peritremal shield

and exopodal shield near coxa IV fused ....................................................... Rykellus Lee

39. Dorsal shield with more than 50 pairs of setae ............................................. Caliphis Lee

- Dorsal shield with less than 45 pairs of setae ................................................................ 40

40. Dorsal shield not fused with ventrianal shield [if fused (some Gamasiphoides), with two

pairs of presternal plates, sternal shield not fused with endopodal shield near coxa IV,

and exopodal shield near coxae II-III interrupted at level of median region of coxa III]

........................................................................................................................................ 41

- Dorsal shield fused with ventrianal shield; 1-2 pairs of presternal plates; sternal shield

fused or not fused with endopodal shield near coxa IV; exopodal shield near coxae II-III

interrupted or not at level of median region of coxa III ................................................. 53

41. Peritreme sinuous ................................................................... Antennolaelaps Womersley

- Peritreme straight ........................................................................................................... 42

42. Genu I with 12 setae (two ventral setae) .......................... Gamasellopsis Loots and Ryke

- Genu I with 13 setae (three ventral setae) ...................................................................... 43

43. Peritremal shield not fused with exopodal shield [if fused (Onchogamasus communis),

with a distinct line of fusion between podonotal and opisthonotal shields, except for

region between setae j6] ................................................................................................. 44

132

- Peritremal shield fused with exopodal shield; line of fusion between podonotal and

opisthonotal shields indistinct ....................................................................................... 47

44. Sternal shield not fused with endopodal shield near coxa IV; epistome with club-shaped

antero-medial extension. ............................................ Neogamasellevans Loots and Ryke

- Sternal shield fused with endopodal shield near coxa IV shield; epistome without club-

shaped antero-medial extension. .................................................................................... 45

45. Genu III with nine setae (two ventral setae) and genu IV with ten setae (five dorsal and

two ventral setae) ................................................................... Onchogamasus Womersley

- Genu III with eight setae (one ventral seta) and genu IV with eight or nine setae (four or

five dorsal and one ventral setae) .................................................................................. 46

46. Pretarsus I pedunculate. ................................................................. Gamasitus Womersley

- Pretarsus I sessile ....................................................................... Sessiluncus G. Canestrini

47. With two pairs of presternal plates ........................................ Gamasiphoides Womersley

- With one pair of presternal plates, partially fused or not fused to sternal shield .......... 48

48. Seta st3 mediad and slightly posterior to st2 .......................... Pachymasiphis Karg, 1996

- Seta st3 approximately in longitudinal line with st2 and st4 ......................................... 49

49. Connection between peritremal and exopodal shields behind stigma narrower than

peritreme ........................................................................................................................ 50

- Connection between peritremal and exopodal shields behind stigma wider than

peritreme ........................................................................................................................ 51

50. Peritreme extending anteriorly to anterior margin of coxa II ................. Geogamasus Lee

- Peritreme extending anteriorly to region of coxa I ..................... Parasitiphis Womersley

51. Sternal shield fused with endopodal shield near coxa IV; epistome with three

extensions, the antero-medial club-shaped .................... Queenslandolaelaps Womersley

- Sternal shield not fused with endopodal shield near coxa IV; epistome not as above .. 52

52. Dorsal podosomal region with less than 21 pairs of setae; peritreme extending anteriorly

to anterior margin of coxa II ....................................................................... Athiasella Lee

- Dorsal podosomal region with more than 20 pairs of setae; peritreme extending

anteriorly to region of coxa I or to median region of coxa III ...........................................

…………………………………………………………….Hydrogamasellus Hirschmann

53. Peritremal shield not fused with exopodal shield near coxa IV .................................... 54

- Peritremal shield fused with exopodal shield near coxa IV .......................................... 55

54. With two pairs of presternal plates; peritreme extending anteriorly to region of coxa I….

..................................................................................................... Laelaptiella Womersley

133

- With one pair of presternal plates; peritreme extending anteriorly to median region of

coxa II............................................................ Stylochirus G. Canestrini and R. Canestrini

55. Sternal shield fused with endopodal shield near coxa IV. ............................................. 56

- Sternal shield not fused with endopodal shield near coxa IV ........................................ 58

56. Exopodal shield near coxae II-III entire; with 0-3 pairs of presternal plates .....................

..............................................................................................................Gamasiphis Berlese

- Exopodal shield near coxae II-III interrupted on median region of coxa III; with one pair

of presternal plates ......................................................................................................... 57

57. Dorsal and ventrianal shields with some pilose setae .................................. Cymiphis Lee

- Dorsal and ventrianal shields with smooth setae ............................. Heydeniella Richters

58. With two pairs of presternal plates................................................ Hydrogamasus Berlese

- With one pair of presternal plates .................................................................................. 59

59. Setae j1 e r3 inserted on prominent protuberance. ........................................ Pyriphis Lee

- Setas j1 e r3 not inserted on protuberance. ....................................... Ologamasus Berlese

3.3.2 Catalogue of world species of Digamasellidae Evans

Genus Dendrolaelaps Halbert, 1915

Dendrolaelaps Halbert, 1915: 68 [described in Parasitidae Oudemans (also referred to as

Gamasidae Leach, name mentioned in the original description of the genus].

Dendrolaelaps.— Lindquist, 1975: 14; Bregetova and Shcherbak, 1977a: 282.

Dendrolaelaps (Dendrolaelaps).— Lindquist, 1975: 15; Hirschmann and Wiśniewski, 1982:

66.

Type species: Dendrolaelaps oudemansi Halbert, 1915, by original designation.

Dendrolaelaps (Punctodendrolaelaps) Hirschmann and Wiśniewski, 1982: 43.

Punctodendrolaelaps.— Karg, 1993c: 351.

Type species: Dendrolaelaps punctatulus Hirschmann, 1960, by original designation.

Dendrolaelaps (Sellnickidendrolaelaps) Hirschmann and Wiśniewski, 1982: 62.

Type species: Dendrolaelaps sellnicki Hirschmann, 1960, by original designation.

134

Dendrolaelaps (Cornodendrolaelaps) Hirschmann and Wiśniewski, 1982: 89.

Cornodendrolaelaps.— Karg, 1993c: 354.

Type species: Dendrolaelaps cornutulus Hirschmann, 1960, by original designation.

Dendrolaelaps (Apophyseodendrolaelaps) Hirschmann and Wiśniewski, 1982: 107.

Type species: Dendrolaelaps apophyseus Hirschmann, 1960, by original designation.

Dendrolaelaps (Disetodendrolaelaps) Hirschmann and Wiśniewski, 1982: 115.

Type species: Dendrolaelaps disetus Hirschmann, 1960, by original designation.

Dendrolaelaps (Foveodendrolaelaps) Hirschmann and Wiśniewski, 1982: 118.

Type species: Digamasellus foveolatus Leitner, 1949, by original designation.

Dendrolaelaps (Presepodendrolaelaps) Hirschmann and Wiśniewski, 1982: 135.

Type species: Gamasellus (Digamasellus) presepum Berlese, 1918, by original

designation.

Dendrolaelaps (Majestidendrolaelaps) Wiśniewski and Hirschmann, 1989b: 317.

Type species: Dendrolaelaps (Majestidendrolaelaps) majesticus Wiśniewski and

Hirschmann, 1989, by original designation.

Dendrolaelaps (Luxtondendrolaelaps) Wiśniewski and Hirschmann, 1989c: 325.

Type species: Dendrolaelaps (Luxtondendrolaelaps) luxtoni Wiśniewski and


Dendrolaelaps (Monodendrolaelaps) Wiśniewski and Hirschmann, 1989d: 117.

Type species: Dendrolaelaps (Monodendrolaelaps) monodentatus Wiśniewski and


Dendrolaelaps (Daeleidendrolaelaps) Wiśniewski and Hirschmann, 1990: 284.

Type species: Dendrolaelaps bidentatus Van Daele, 1977, by original designation.

Dendrolaelaps (Duplodendrolaelaps) Wiśniewski and Hirschmann, 1991b: 403.

135

Type species: Dendrolaelaps (Duplodendrolaelaps) duplodens Wiśniewski and


Dendrolaelaps (Stanidendrolaelaps) Wiśniewski and Hirschmann, 1993b: 290.

Type species: Dendrolaelaps (Stanidendrolaelaps) stanislavi Wiśniewski and


Dendrolaelaps (Xylodendrolaelaps) Wiśniewski and Hirschmann, 1993c: 322.

Type species: Dendrolaelaps (Xylodendrolaelaps) xylophilus Wiśniewski and


001. Dendrolaelaps aberratus Hirschmann and Wiśniewski, 1984

Dendrolaelaps (Cornodendrolaelaps) aberratus Hirschmann and Wiśniewski, 1984: 90.

TYPE DEPOSITORY: Ungarischess Naturwissenschaftliches Museum, Budapest, Hungary.

TYPE LOCALITY AND HABITAT: Ecuador, between Baeza and Quito, 1971, from

unspecified substrate.

002. Dendrolaelaps abietis Hirschmann, 1960

Dendrolaelaps abietis Hirschmann, 1960: 8.

Dendrolaelaps abietis.— Hirschmann, 1971d: 17; Karg, 1971: 328; Shcherbak, 1980: 118;

Karg, 1993c: 347.

Dendrolaelaps (Dendrolaelaps) abietis.— Hirschmann, 1974: 62; Hirschmann and

Wiśniewski, 1982: 78.

TYPE DEPOSITORY: Hirschmann‘s Milbensammlung, Nuremberg, Germany.

TYPE LOCALITY AND HABITAT: Germany, Zell, Schwarzwald, on stump of Abies sp.

[Pinaceae].

003. Dendrolaelaps acornutosimilis Hirschmann, 1960

Dendrolaelaps acornutosimilis Hirschmann, 1960: 7.

Dendrolaelaps acornutosimilis.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 12;

Hirschmann, 1971c: 16; Hirschmann, 1971d: 17; Hirschmann, 1971e: 22;

Hirschmann, 1971f: 27; Karg, 1993c: 348.

Dendrolaelaps (Dendrolaelaps) acornutosimilis.— Hirschmann, 1974: 62; Hirschmann and


136


TYPE LOCALITY AND HABITAT: Germany, Zell, Schwarzwald, on stumps of Picea sp.

[Pinaceae], Abies sp. [Pinaceae] and Fagus sp. [Fagaceae]; Germany, Bückeberg, on

stump of Quercus sp. [Fagaceae]; Switzerland, Pontresina, [Maloja], on stump of

Pinus sp. [Pinaceae].

NOTE1: Karg (1971): 340 considered D. acornutosimilis as junior synonymy of

Dendrolaelaps septentrionalis (Sellnick, 1958), but Hirschmann and Wiśniewski

(1982): 77 revoked that synonymy.

NOTE2: Shcherbak (1980): 118 considered D. acornutosimilis as junior synonymy of

Dendrolaelaps oudemansi Halbert, 1915, but Hirschmann and Wiśniewski (1982): 77

revoked that synonymy.

004. Dendrolaelaps acornutus Hirschmann, 1960

Dendrolaelaps acornutus Hirschmann, 1960: 4.

Dendrolaelaps acornutus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,

1971c: 16; Hirschmann, 1971d: 17; Hirschmann, 1971e: 22; Hirschmann, 1971f: 27;

Karg, 1971: 330; Shcherbak, 1980: 118; Karg, 1993c: 349.

Dendrolaelaps (Dendrolaelaps) acornutus.— Hirschmann, 1974: 62; Hirschmann and



TYPE LOCALITY AND HABITAT: Germany, Bückeberg, on stump of Picea sp.

[Pinaceae]; Germany, Lüneburg, on stump of Picea sp.; Germany, Nuremberg, on

stumps of Pinus sp. [Pinaceae] and Betula sp. [Betulaceae], on Ernoporus fagi

[Coleoptera: Curculionidae: Scolytinae] and in gallery of Dryocoetes autographus

[Coleoptera: Curculionidae: Scolytinae]; Germany, Oberstdorf, in gallery of D.

autographus; Germany, Süderlügum, Schleswig-Holstein, on Myelophilus piniperda

[Coleoptera: Curculionidae: Scolytinae] and Dendroctonus micans [Coleoptera:

Curculionidae: Scolytinae]; Germany, Bonn, on root of Robinia pseudacacia

[Fabaceae].

005. Dendrolaelaps adelaideae Womersley, 1954

Dendrolaelaps adelaideae Womersley, 1954: 113.

Dendrolaelaps adelaideae.— Hirschmann, 1960: 8.

Cyrtolaelaps (Digamasellus) adelaideae.— Ryke, 1962a: 110.

137

Dendrolaelaps (Punctodendrolaelaps ?) adelaideae [sic].— Hirschmann and Wiśniewski,

1982: 60.

Dendrolaelaps concinna Womersley, 1954: 115 [junior synonymy by Hirschmann and

Wiśniewski, 1982: 60].

Dendrolaelaps concinna.— Hirschmann, 1960: 8.

Cyrtolaelaps (Digamasellus) concinnus.— Ryke, 1962a: 110.

TYPE DEPOSITORY: D. adelaideae: South Australian Museum, Adelaide, Australia; D.

concinna: South Australian Museum, Adelaide, Australia.

TYPE LOCALITY AND HABITAT: D. adelaideae: Australia, Adelaide, May 1952, from

frass of bark-boring beetles Ips sp. [Coleoptera: Curculionidae: Scolytinae]; D.

concinna: Australia, Adelaide, May 1951, from frass of bark-boring beetles Ips sp..

006. Dendrolaelaps aegypticus Metwally and Mersal, 1985

Dendrolaelaps aegypticus Metwally and Mersal, 1985 apud Mostafa, 2011: 855.


TYPE LOCALITY AND HABITAT: Unknown.

007. Dendrolaelaps apophyseosimilis Hirschmann, 1960

Dendrolaelaps apophyseosimilis Hirschmann, 1960: 7.

Dendrolaelaps apophyseosimilis.— Hirschmann, 1971b: 13; Hirschmann, 1971c: 16;

Hirschmann, 1971e: 20; Karg, 1971: 330; Karg, 1993c: 348.

Dendrolaelaps (Dendrolaelaps) apophyseosimilis.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) apophyseosimilis.— Hirschmann and Wiśniewski,

1982: 111.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stumps of Picea sp.

[Pinaceae] and Alnus sp. [Betulaceae]; Switzerland, Ofenpass, on stump of Pinus

mugo [Pinaceae]; Germany, Lüneburg, on stump of Picea sp.; Germany, Fürth, in

gallery of Scolytus ratzeburgi [Coleoptera: Curculionidae: Scolytinae].

NOTE: Shcherbak (1980): 136 considered D. apophyseosimilis as junior synonymy of

Dendrolaelaps disetosimilis Hirschmann, 1960, but Hirschmann and Wiśniewski


008. Dendrolaelaps apophyseus Hirschmann, 1960

138

Dendrolaelaps apophyseus Hirschmann, 1960: 7.

Dendrolaelaps apophyseus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,

1971c: 16; Hirschmann, 1971d: 18; Hirschmann, 1971e: 21; Karg, 1971: 331;

Shcherbak, 1980: 138; Karg, 1993c: 349.

Dendrolaelaps (Dendrolaelaps) apophyseus.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) apophyseus.— Hirschmann and Wiśniewski, 1982:

110.


TYPE LOCALITY AND HABITAT: Germany, on stumps of Picea sp. [Pinaceae] and Pinus

sp. [Pinaceae], and in galleries of Ips typographus [Coleoptera: Curculionidae:

Scolytinae], Hylurgops palliates [Coleoptera: Curculionidae: Scolytinae] and

Dryocoetes autographus [Coleoptera: Curculionidae: Scolytinae]; Switzerland,

Pontresina, [Maloja], on stump of Larix sp. [Pinaceae].

009. Dendrolaelaps arenarioides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Foveodendrolaelaps) arenarioides Hirschmann and Wiśniewski, 1982: 129.



TYPE LOCALITY AND HABITAT: Central Europe, in sand of sea shore, on roots of Cakile

maritime [Brassicaceae], on algae, humus of Scirpus sp. [Cypercaceae] and in beech

Fagus sp. [Fagaceae] litter.

NOTE: Described on the basis of specimens reported by Shcherbak (1980): 144 as

Dendrolaelaps arenarius Karg, 1971.

010. Dendrolaelaps arenarius Karg, 1971

Dendrolaelaps arenarius Karg, 1971: 339.

Dendrolaelaps (Dendrolaelaps) arenarius.— Hirschmann, 1974: 63.

Dendrolaelaps (Foveodendrolaelaps) arenarius.— Hirschmann and Wiśniewski, 1982: 129.

Dendrolaelaps arenarius.— Karg, 1993c: 351.

TYPE DEPOSITORY: Institut für Pflanzenschutzforschung Kleinmachnow, Kleinmachnow,

Germany.

TYPE LOCALITY AND HABITAT: Germany, Darss, [Fischland-Darss-Zingst Peninsula], 5

September 1967, on roots of Ammophila sp. [Poaceae].

139

NOTE: Specimens reported by Shcherbak (1980): 144 as Dendrolaelaps arenarius Karg,

1971 were described as Dendrolaelaps (Foveodendrolaelaps) arenarioides

Hirschmann and Wiśniewski, 1982: 129.

011. Dendrolaelaps arvicolus (Leitner, 1949)

Digamasellus arvicolus Leitner, 1949: 61.

Digamasellus arvicolus.— Franz, 1954: 341.

Dendrolaelaps arvicolus.— Hirschmann, 1960: 7; Hirschmann, 1971c: 15; Hirschmann,

1971d: 19; Hirschmann, 1971e: 21; Hirschmann, 1971f: 26; Karg, 1971: 338;

Shcherbak, 1980: 101; Huhta, 1982: 226.

Cyrtolaelaps (Digamasellus) arvicolus.— Ryke, 1962a: 108.

Dendrolaelaps (Dendrolaelaps) arvicolus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) arvicolus.— Hirschmann and Wiśniewski, 1982: 51.

Punctodendrolaelaps arvicolus.— Karg, 1993c: 353.


TYPE LOCALITY AND HABITAT: Austria, Admont, [Styria], in arable soil (10-17 cm).

NOTE: Shcherbak (1980): 101 considered Dendrolaelaps insignis Hirschmann, 1960 as

junior synonymy of D. arvicolus, but Hirschmann and Wiśniewski (1982): 51 revoked

that synonymy.

012. Dendrolaelaps australicornutus Hirschmann, 1972

Dendrolaelaps australicornutus Hirschmann, 1972: 31.

Dendrolaelaps (Dendrolaelaps) australicornutus.— Hirschmann, 1974: 62; Hirschmann and



TYPE LOCALITY AND HABITAT: Brazil, Ibirama, [Santa Catarina State], 1964, in phloem

of Araucaria angustifolia [Araucariaceae] damaged by various insects.

013. Dendrolaelaps baixuelii Ma, 1997

Dendrolaelaps baixuelii Ma, 1997b: 137.

Dendrolaelaps baixuelii.— Ma, 2001b: 13; Ma, 2005a: 18.


TYPE LOCALITY AND HABITAT: China, Baicheng (45°37‘N, 122°49‘E), Jilin Province, 6

August 1994, on rotten dregs of Populus sp. [Salicaceae].

140

014. Dendrolaelaps balazyi Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Cornodendrolaelaps) balazyi Hirschmann and Wiśniewski, 1982: 101.

Cornodendrolaelaps balazyi.— Karg, 1993c: 355.

TYPE DEPOSITORY: Department of Forest Protection, University of Life Sciences, Poznań,

Poland.

TYPE LOCALITY AND HABITAT: Poland, Taborz, [Olsztyn], September 1961, under bark

of Picea sp. [Pinaceae] in gallery of an unidentified beetle [Coleoptera].

015. Dendrolaelaps bengalensis Pramanik and Raychaudhuri, 1978

Dendrolaelaps bengalensis Pramanik and Raychaudhuri, 1978: 33.

Dendrolaelaps (Punctodendrolaelaps) bengalensis.— Hirschmann, 1983b: 72.

TYPE DEPOSITORY: Entomology Laboratory, Department of Zoology, University of

Calcutta, Calcutta, India.

TYPE LOCALITY AND HABITAT: India, Barasat, North 24 Parganas, West Bengal, 7

August 1974, in soil litter.

016. Dendrolaelaps bhattacharyyai Hirschmann, 1974

Dendrolaelaps (Dendrolaelaps) bhattacharyyai Hirschmann, 1974: 52.

Dendrolaelaps (Apophyseodendrolaelaps) bhattacharyyai.— Hirschmann and Wiśniewski,

1982: 114.

TYPE DEPOSITORY: Zoological Survey of India, Calcutta, India.

TYPE LOCALITY AND HABITAT: India, Kameng Elephant Reserve, Arunachal Pradesh

(cited as North-East Frontier Agency), 23 December 1965, under bark of a dead tree.

NOTE1: Described on the basis of specimens reported by Bhattacharyya (1969): 69 as

allotype adult male of Digamasellus orientalis Bhattacharyya, 1969.

NOTE2: Described from the adult male.

017. Dendrolaelaps bidentatus Van Daele, 1977

Dendrolaelaps bidentatus Van Daele, 1977: 199.

Dendrolaelaps (Apophyseodendrolaelaps) bidentatus.— Hirschmann and Wiśniewski, 1982:

114.

Dendrolaelaps bidentatus.— Wiśniewski and Hirschmann, 1990: 271; Karg, 1993c: 350.

Dendrolaelaps (Daeleidendrolaelaps) bidentatus.— Wiśniewski and Hirschmann, 1990: 284.

141

TYPE DEPOSITORY: Chair of Zoology, Faculty of Agricultural Sciences, State University

of Ghent, Belgium.

TYPE LOCALITY AND HABITAT: Belgium, Ghent, [East Flanders], 12 February 1976, on

substrate of Dracaena fragrans var. massangeana [Ruscaceae] in glasshouses.

018. Dendrolaelaps bisetus (Berlese, 1891)

Zercon bisetus Berlese, 1891: LVIII, 7.

Cyrtolaelaps bisetus.— Oudemans, 1902a: 29.

Gamasellus bisetus.— Berlese, 1920a: 23; Bernini, Castagnoli and Nannelli, 1995: 20.

Cyrtolaelaps (Digamasellus) bisetus.— Ryke, 1962a: 91.

Dendrolaelaps (Punctodendrolaelaps) bisetus.— Hirschmann and Wiśniewski, 1982: 59.

Cyrtolaelaps captator Berlese, 1892c: LXVIII, 8 [junior synonymy by Berlese, 1920a: 6].

Cyrtolaelaps (Gamasellus) captator.— Berlese, 1892f: 61.

Gamasellus captator.— Schweizer, 1922: 34.

Dendrolaelaps captator.— Hirschmann, 1960: 8.

Dendrolaelaps (Dendrolaelaps) captator.— Hirschmann, 1974: 62.

TYPE DEPOSITORY: P. bisetus and C. captator: Istituto Sperimentale per la Zoologia

Agraria, Florence, Italy.

TYPE LOCALITY AND HABITAT: P. bisetus: Italy, Portici, Province of Naples, in

grooves; C. captator: Italy, Portici, Province of Naples, in forest litter.

019. Dendrolaelaps brasiliensis Wiśniewski and Hirschmann, 1984

Dendrolaelaps (Foveodendrolaelaps) brasiliensis Wiśniewski and Hirschmann, in



TYPE LOCALITY AND HABITAT: Brazil, Cotia, [São Paulo State], 16 January 1975, on

Bothynus deiphobus [Coleoptera: Dynastidae].

NOTE: Described from the deutonymph.

020. Dendrolaelaps brevipilis (Leitner, 1949)

Digamasellus brevipilis Leitner, 1949: 62.

Digamasellus brevipilis.— Franz, 1954: 341; Willmann, 1951: 143.

Dendrolaelaps brevipilis.— Hirschmann, 1960: 8; Hirschmann, 1971d: 18; Hirschmann,

1971e: 20; Karg, 1971: 332; Karg, 1993c: 350; Huhta and Karg, 2010: 347.

142

Cyrtolaelaps (Digamasellus) brevipilis.— Ryke, 1962a: 109.

Dendrolaelaps (Dendrolaelaps) brevipilis.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) brevipilis.— Hirschmann and Wiśniewski, 1982: 128.


TYPE LOCALITY AND HABITAT: Austria, Admont, [Styria], in nest of Talpa europaea

[Mammalia: Talpidae].

NOTE: Specimens reported by Shcherbak (1980): 145 as Dendrolaelaps brevipilis (Leitner,

1949) were described as Dendrolaelaps (Foveodendrolaelaps) brevipiloides


021. Dendrolaelaps brevipiloides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Foveodendrolaelaps) brevipiloides Hirschmann and Wiśniewski, 1982: 128.



TYPE LOCALITY AND HABITAT: Ukraine; and Kazakhstan, Karaganda, Karagandy

Province, in soil and litter of a mixed forest, in compost and in rodent nests

[Mammalia: Rodentia].


Dendrolaelaps brevipilis (Leitner, 1949).

022. Dendrolaelaps camponoti Wiśniewski and Hirschmann, 1983

Dendrolaelaps (Dendrolaelaps) camponoti Wiśniewski and Hirschmann, 1983a: 109.


TYPE LOCALITY AND HABITAT: Poland, Śnieżnik Mountain, Miedzylesie, 17 May 1979,

in a tree nest of Camponotus herculeanus [Hymenoptera: Formicidae].

NOTE: Described from the adult male.

023. Dendrolaelaps capensis (Berlese, 1920)

Gamasellus (Digamasellus) capensis Berlese, 1920b: 161.

Dendrolaelaps capensis.— Hirschmann, 1960: 8.

Dendrolaelaps (Tridendrolaelaps) capensis.— Hirschmann, 1974: 60.

? Dendrolaelaps capensis [sic].— Hirschmann and Wiśniewski, 1982: 148.


143

TYPE LOCALITY AND HABITAT: Cape of Good Hope, [Atlantic Coast of Africa], in

humus.

NOTE: Specimens reported by Ryke (1962a): 92 as Cyrtolaelaps (Digamasellus) capensis

(Berlese, 1921) were described as Dendrolaelaps (Tridendrolaelaps) rykei

Hirschmann, 1974: 60.

024. Dendrolaelaps carolinensis McGraw and Farrier, 1969

Dendrolaelaps carolinensis McGraw and Farrier, 1969: 105.

Dendrolaelaps (Dendrolaelaps) carolinensis.— Hirschmann, 1974: 63.

Dendrolaelaps (Cornodendrolaelaps) carolinensis.— Hirschmann and Wiśniewski, 1982:

102.


TYPE LOCALITY AND HABITAT: USA, Wake County, North Carolina, 31 March 1966,

on Dendroctonus terebrans [Coleoptera: Curculionidae: Scolytinae] attacking Pinus

taeda [Pinaceae].

025. Dendrolaelaps casualis Huhta and Karg, 2010

Dendrolaelaps casualis Huhta and Karg, 2010: 346.

TYPE DEPOSITORY: Zoological Museum, University of Helsinki, Finland.

TYPE LOCALITY AND HABITAT: Finland, Kittilä, Ylläs, Lapland, 5 August 2005, on a

dry toilet compost [= dry human feces?] on the roadside by a popular tourist route.

026. Dendrolaelaps coleopterophilus (Hirschmann, 1954)

Digamasellus coleopterophilus Hirschmann, 1954b: 247.

Dendrolaelaps coleopterophilus.— Hirschmann, 1960: 9; Hirschmann, 1971c: 14; Shcherbak,

1980: 122.

Dendrolaelaps (Dendrolaelaps) coleopterophilus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) coleopterophilus [sic].— Hirschmann and



TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stump of Picea sp.

[Pinaceae], and under the elytra of Elater sanguineus [Coleoptera: Elateridae] and

Pyrochroa sp. [Coleoptera: Pyrochroidae].


144

027. Dendrolaelaps comatus Hirschmann, 1960

Dendrolaelaps comatus Hirschmann, 1960: 7.

Dendrolaelaps comatus.— Hirschmann, 1971b: 12; Hirschmann, 1971c: 16; Hirschmann,


Shcherbak, 1980: 110.

Dendrolaelaps (Dendrolaelaps) comatus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) comatus.— Hirschmann and Wiśniewski, 1982: 55.

Punctodendrolaelaps comatus.— Karg, 1993c: 352.


TYPE LOCALITY AND HABITAT: Germany, on stumps of Picea sp. [Pinaceae] and Pinus

sp. [Pinaceae], and in galleries of Ips typographus [Coleoptera: Curculionidae:

Scolytinae], Hylurgops palliates [Coleoptera: Curculionidae: Scolytinae],

Dendroctonus micans [Coleoptera: Curculionidae: Scolytinae] and Pityokteines

curvidens [Coleoptera: Curculionidae: Scolytinae].

028. Dendrolaelaps cornutodaelei Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) cornutodaelei Hirschmann and Wiśniewski, 1982: 86.


TYPE LOCALITY AND HABITAT: Belgium, in Merelbeke and Ghent, [East Flanders], 23

July 1982, intercepted on cuttings of Dracaena fragrans [Ruscaceae] imported from

Brazil.

029. Dendrolaelaps cornutohirschmanni Wiśniewski, 1979

Dendrolaelaps cornutohirschmanni Wiśniewski, 1979a: 150.

Dendrolaelaps (Dendrolaelaps) cornutohirschmanni.— Hirschmann and Wiśniewski, 1982:

87.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Potasze, Poznań County, 20 July 1975, under

bark of Picea abies [Pinaceae] in galleries of unidentified Coleoptera.


030. Dendrolaelaps cornutulus Hirschmann, 1960

145

Dendrolaelaps cornutulus Hirschmann, 1960: 7.

Dendrolaelaps cornutulus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,


Karg, 1971: 338; Shcherbak, 1980: 123.

Dendrolaelaps (Dendrolaelaps) cornutulus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) cornutulus.— Hirschmann and Wiśniewski, 1982: 96.

Cornodendrolaelaps cornutulus.— Karg, 1993c: 363.


TYPE LOCALITY AND HABITAT: Germany, on stumps of Picea sp. [Pinaceae], Abies sp.

[Pinaceae], Fagus sp. [Fagaceae], Quercus sp. [Fagaceae], Betula sp. [Betulaceae],

Alnus sp. [Betulaceae] and Sorbus sp. [Rosaceae], and in galleries of Dendroctonus

micans [Coleoptera: Curculionidae: Scolytinae].

NOTE: Franz (1954): 341 cited Digamasellus cornutulus Hirschmann n. sp., but did not

characterize it morphologically; thus, that citation constitutes a nomen nudum.

031. Dendrolaelaps cornutus (Kramer, 1886)

Sejus cornutus Kramer, 1886: 257.

Cyrtolaelaps (Gamasellus) cornutus.— Berlese, 1892f: 61.

Dendrolaelaps cornutus.— Halbert, 1923: 366; Hirschmann, 1971a: 11; Hirschmann, 1971b:

13; Hirschmann, 1971c: 16; Hirschmann, 1971d: 17; Hirschmann, 1971e: 22;

Hirschmann, 1971f: 27; Karg, 1971: 330.

Digamasellus cornutus.— Schweizer, 1961: 137.

Cyrtolaelaps (Digamasellus) cornutus.— Ryke, 1962a: 103.

Dendrolaelaps (Dendrolaelaps) cornutus.— Hirschmann, 1974: 62; Hirschmann and


Dendrolaelaps bicornis Hull, 1918: 57 [junior synonymy by Halbert, 1923: 366].

Dendrolaelaps (Dendrolaelaps) bicornis.— Hirschmann and Wiśniewski, 1982: 88.

TYPE DEPOSITORY: D. cornutus: Zoologisches Museum, Hamburg, Germany; D. bicornis:

British Museum (Natural History), London, England.

TYPE LOCALITY AND HABITAT: D. cornutus: Germany, from unspecified substrate; D.

bicornis: England, Allendale [cited as West Allendale], Tynedale [cited as Tyne

Province], under bark of fallen trees.

146

NOTE1: Specimens reported by Vitzthum (1926b): 411 as Dendrolaelaps cornutus (Kramer,

1886) were described as Dendrolaelaps (Dendrolaelaps) vitzthumicornutus


NOTE2: Specimens reported by Hirschmann (1960): 6 as Dendrolaelaps cornutus (Kramer,

1886) were described as Dendrolaelaps (Dendrolaelaps) nostricornutus Hirschmann

and Wiśniewski, 1982: 80.

NOTE3: Specimens reported by Shcherbak (1980): 114 as Dendrolaelaps cornutus (Kramer,

1886) sensu Hirschmann, 1960 were described as Dendrolaelaps (Dendrolaelaps)

shcherbakaecornutus Hirschmann and Wiśniewski, 1982: 80.

NOTE4: Described from the larve and adult male.

032. Dendrolaelaps crassipes (Schweizer, 1961)

Digamasellus crassipes Schweizer, 1961: 141.

Dendrolaelaps (Dendrolaelaps) crassipes.— Hirschmann, 1974: 63; Hirschmann and


TYPE DEPOSITORY: Naturhistorisches Museum Basel, Basel, Switzerland.

TYPE LOCALITY AND HABITAT: Switzerland, Alp Stabelchod (alt. 1,963 m), Swiss

National Park, under lumber.


033. Dendrolaelaps crassitarsalis (Willmann, 1951)

Digamasellus crassitarsalis Willmann, 1951: 142.

Dendrolaelaps crassitarsalis.— Hirschmann, 1960: 8; Hirschmann, 1971d: 19; Hirschmann,

1971e: 19; Karg, 1971: 332; Shcherbak, 1980: 141; Karg, 1993c: 351.

Cyrtolaelaps (Digamasellus) crassitarsalis.— Ryke, 1962a: 109.

Dendrolaelaps (Dendrolaelaps) crassitarsalis.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) crassitarsalis.— Hirschmann and Wiśniewski, 1982:

133.


TYPE LOCALITY AND HABITAT: Germany, Wangerooge Island, [Friesland], 19 June

1949, in soil with grass in a horse pasture.

NOTE: Shcherbak (1980): 141 considered Dendrolaelaps rectus Karg, 1962 as junior

synonymy of D. crassitarsalis, but Hirschmann and Wiśniewski (1982): 134 revoked

that synonymy.

147

034. Dendrolaelaps cubae Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Disetodendrolaelaps) cubae Wiśniewski and Hirschmann, 1993a: 75.


TYPE LOCALITY AND HABITAT: Cuba, Jardín Botánico de Cienfuegos, Cienfuegos, 4

April 1987, under tree bark.

035. Dendrolaelaps cylindricus (Berlese, 1918)

Gamasellus (Digamasellus) cylindricus Berlese, 1918: 135.

Dendrolaelaps cylindricus.— Hirschmann, 1960: 8.

Cyrtolaelaps (Digamasellus) cylindricus.— Ryke, 1962a: 104.

Dendrolaelaps (Dendrolaelaps) cylindricus.— Hirschmann, 1974: 62: Hirschmann and



TYPE LOCALITY AND HABITAT: ―North America, Cl. Crosby, Columbia‖ [sic], on

humus and litter.

036. Dendrolaelaps debilipes (Berlese, 1920)

Gamasellus (Digamasellus) debilipes Berlese, 1920b: 160.

Dendrolaelaps debilipes.— Hirschmann, 1960: 8.

Cyrtolaelaps (Digamasellus) debilipes.— Ryke, 1962a: 104.

Dendrolaelaps (Dendrolaelaps) debilipes.— Hirschmann, 1974: 62.

Dendrolaelaps (Punctodendrolaelaps) debilipes.— Hirschmann and Wiśniewski, 1982 : 59.


TYPE LOCALITY AND HABITAT: USA, Columbia, in humus.

037. Dendrolaelaps disetosimilis Hirschmann, 1960

Dendrolaelaps disetosimilis Hirschmann, 1960: 7.

Dendrolaelaps disetosimilis.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,

1971c: 15; Hirschmann, 1971d: 18; Hirschmann, 1971e: 20; Karg, 1971: 331;

Shcherbak, 1980: 136; Karg, 1993c: 349.

Dendrolaelaps (Dendrolaelaps) disetosimilis.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) disetosimilis.— Hirschmann and Wiśniewski,

1982: 111.

148


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stump of Pinus sp. [Pinaceae]

and in galleries of Dryocoetes autographus [Coleoptera: Curculionidae: Scolytinae].

NOTE: Shcherbak (1980): 136 considered Dendrolaelaps apophyseosimilis Hirschmann,

1960 as junior synonymy of D. disetosimilis, but Hirschmann and Wiśniewski (1982):

111 revoked that synonymy.

038. Dendrolaelaps disetus (Hirschmann, 1954)

Digamasellus disetus Hirschmann, 1954a: 107.

Digamasellus disetus.— Hirschmann, 1954b: 247

Dendrolaelaps disetus.— Hirschmann, 1960: 7; Hirschmann, 1971b: 13; Hirschmann, 1971c:

16; Hirschmann, 1971d: 17; Hirschmann, 1971e: 21; Hirschmann, 1971f: 28; Karg,

1971: 331; Shcherbak, 1980: 138.

Dendrolaelaps (Dendrolaelaps) disetus.— Hirschmann, 1974: 62.

Dendrolaelaps (Disetodendrolaelaps) disetus.— Hirschmann and Wiśniewski, 1982: 116.

Cornodendrolaelaps disetus.— Karg, 1993c: 354.


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stump of Fagus sp.

[Fagaceae]; Germany, Belchen, Schwarzwald, on stump of Fagus sp..

039. Dendrolaelaps doljensis Wiśniewski and Hirschmann, 1991

Dendrolaelaps (Sellnickidendrolaelaps) doljensis Wiśniewski and Hirschmann, 1991a: 230.


Poland.

TYPE LOCALITY AND HABITAT: Romania, Jiu River, [Dolj], 22 July 1988, in detritus of

Quercus cerris [Fagaceae] in a hillside erosion by a cave.

040. Dendrolaelaps duplodens Wiśniewski and Hirschmann, 1991

Dendrolaelaps (Duplodendrolaelaps) duplodens Wiśniewski and Hirschmann, 1991b: 404.

Cornodendrolaelaps duplodens.— Karg, 1993c: 361.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Głuszyca, Wałbrzych, 18 March 1990, in litter

of Acer sp. [Sapindaceae].

149

041. Dendrolaelaps eichhorni Wiśniewski, 1980

Dendrolaelaps eichhorni Wiśniewski, 1980: 7.

Dendrolaelaps (Punctodendrolaelaps) eichhorni.— Hirschmann and Wiśniewski, 1982: 53.

Punctodendrolaelaps eichhorni.— Karg, 1993c: 352.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Śnieżnik Landscape Park (alt. 750 m),

Międzygórze, 16 August 1979, in tree nests of Camponotus herculeanus

[Hymenoptera: Formicidae].

042. Dendrolaelaps elaterophilus (Hirschmann, 1954)

Digamasellus elaterophilus Hirschmann, 1954b: 247.

Dendrolaelaps elaterophilus.— Hirschmann, 1960: 9; Hirschmann, 1971b: 14; Hirschmann,

1971c: 14; Shcherbak, 1980: 122.

Dendrolaelaps (Dendrolaelaps) elaterophilus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) elaterophilus.— Hirschmann and Wiśniewski, 1982:

99.


TYPE LOCALITY AND HABITAT: Germany, Bremen, on stump of Quercus sp.

[Fagaceae]; Germany, Nuremberg, on stump of Picea sp. [Pinaceae], under the elytra

of Spondylis buprestoides [Coleoptera: Cerambycidae], Tetropium luridum

[Coleoptera: Cerambycidae], Rhagium bifasciatum [Coleoptera: Cerambycidae],

Elater balteatus [Coleoptera: Elateridae] and Elater sanguineus [Coleoptera:

Elateridae].

NOTE: Described from the protonymph and deutonymph.

043. Dendrolaelaps fageticola (Schmölzer, 1995)

Cornodendrolaelaps fageticola Schmölzer, 1995a: 500.


TYPE LOCALITY AND HABITAT: Austria, circuit ―Kolsche auf der Petzen‖, north of the

peak Feistritzer (alt. 1,375 m), Karawanken, Carinthia, 2 August 1990, in litter of a

beech Fagus sp. [Fagaceae] wood at the edge of a mixed forest of beech Fagus sp. and

spruce Picea sp. [Pinaceae] in an area surrounding a cottage.

150

044. Dendrolaelaps fallacoides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps) fallacoides Hirschmann and Wiśniewski, 1982: 54.



TYPE LOCALITY AND HABITAT: Ukraine; Russia, Novosibirsk Oblast; Kazakhstan,

Karaganda, Karagandy Province; in dung, carrion cane, soil under the trees of an oak

forest, old stumps, litter and soil under mushroom.


Dendrolaelaps fallax (Leitner, 1949).

045. Dendrolaelaps fallax (Leitner, 1949)

Digamasellus fallax Leitner, 1949: 60.

Digamasellus fallax.— Hirschmann, 1954a: 106; Franz, 1954: 341.

Dendrolaelaps fallax.— Hirschmann, 1960: 6; Hirschmann, 1971a: 11; Hirschmann, 1971b:

13; Hirschmann, 1971c: 16; Hirschmann, 1971d: 18; Hirschmann, 1971e: 21;


Cyrtolaelaps (Digamasellus) fallax.— Ryke, 1962a: 108.

Dendrolaelaps (Dendrolaelaps) fallax.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) fallax.— Hirschmann and Wiśniewski, 1982: 54.

Punctodendrolaelaps fallax.— Karg, 1993c: 352.


TYPE LOCALITY AND HABITAT: Austria, Saalbach-Hinterglemm, Zell am See, in

compost with predominance of evergreen aciculate litter.

NOTE: Specimens reported by Shcherbak (1980): 107 as Dendrolaelaps fallax (Leitner,

1949) were described as Dendrolaelaps (Punctodendrolaelaps) fallacoides


046. Dendrolaelaps forcipiformis Hirschmann, 1960

Dendrolaelaps forcipiformis Hirschmann, 1960: 7.

Dendrolaelaps forcipiformis.— Hirschmann, 1971d: 18; Hirschmann, 1971e: 21; Karg, 1971:

337; Shcherbak, 1980: 130.

Dendrolaelaps (Dendrolaelaps) forcipiformis.— Hirschmann, 1974: 62.

151

Dendrolaelaps (Cornodendrolaelaps) forcipiformis.— Hirschmann and Wiśniewski, 1982:

102.

Cornodendrolaelaps forcipiformis.— Karg, 1993c: 360.



and in gallery of Ips typographus [Coleoptera: Curculionidae: Scolytinae].

047. Dendrolaelaps formicarius (Huhta and Karg, 2010)

Punctodendrolaelaps formicarius Huhta and Karg, 2010: 339.


TYPE LOCALITY AND HABITAT: Finland, Kisko, 3 May 2005, in a hill of the ant

Formica rufa [Hymenoptera: Formicidae].

048. Dendrolaelaps fossilis Hirschmann, 1971

Dendrolaelaps fossilis Hirschmann, 1971g: 69.

Dendrolaelaps (Multidendrolaelaps) fossilis.— Hirschmann, 1974: 61.

? Dendrolaelaps fossilis [sic].— Hirschmann and Wiśniewski, 1982: 149.

TYPE DEPOSITORY: Museum of Paleontology, University of California, Berkeley, USA.

TYPE LOCALITY AND HABITAT: Mexico, Rancho San Jose Buenavista, south slope of

Cerro Balumtun, Las Cruces landslide, Chiapas, in amber of late Oligocene to early

Miocene age.


049. Dendrolaelaps foveolatosimilis Hirschmann, 1960

Dendrolaelaps foveolatosimilis Hirschmann, 1960: 8.

Dendrolaelaps foveolatosimilis.— Hirschmann, 1971c: 16; Hirschmann, 1971d: 17;

Hirschmann, 1971e: 20; Karg, 1971: 328; Shcherbak, 1980: 148; Karg, 1993c: 347.

Dendrolaelaps (Dendrolaelaps) foveolatosimilis.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) foveolatosimilis.— Hirschmann and Wiśniewski, 1982:

125.


TYPE LOCALITY AND HABITAT: Sweden, Jämtland, in nest of Formica rufa


152

NOTE: Sellnick (1958): 22 cited Digamasellus foveolatosimilis Hirschmann 1950, but did not


050. Dendrolaelaps foveolatus (Leitner, 1949)

Digamasellus foveolatus Leitner, 1949: 55.

Digamasellus toveolatus [sic].— Leitner, 1949: 59.

Digamasellus foveolatus.— Franz, 1954: 341.

Dendrolaelaps foveolatus.— Hirschmann, 1960: 8; Hirschmann, 1971c: 16; Hirschmann,


Shcherbak, 1980: 147; Karg, 1993c: 350.

Cyrtolaelaps (Digamasellus) foveolatus.— Ryke, 1962a: 108.

Dendrolaelaps (Dendrolaelaps) foveolatus.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) foveolatus.— Hirschmann and Wiśniewski, 1982: 126.

Dendrolaelaps hirschmanni Karg, 1962: 40 [junior synonymy by Karg, 1971: 341].

TYPE DEPOSITORY: D. foveolatus: Unknown; D. hirschmanni: Institut für

Pflanzenschutzforschung Kleinmachnow, Kleinmachnow, Germany.

TYPE LOCALITY AND HABITAT: D. foveolatus: Austria, Admont, [Styria], in arable soil

(3 cm); D. hirschmanni: Germany, Experimental Field of Biologische Zentralanstalt of

Kleinmachnow, [Potsdam-Mittelmark, Brandenburg], in grassland soil.

051. Dendrolaelaps frenzeli (Willmann, 1936)

Digamasellus frenzeli Willmann, 1936: 277.

Digamasellus frenzeli.— Leitner, 1949: 59; Willmann, 1951: 142.

Dendrolaelaps frenzeli.— Hirschmann, 1960: 7; Hirschmann, 1971c: 15; Karg, 1993c: 351.

Cyrtolaelaps (Digamasellus) frenzeli.— Ryke, 1962a: 108.

Dendrolaelaps (Dendrolaelaps) frenzeli.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) frenzeli.— Hirschmann and Wiśniewski, 1982: 132.

Dendrolaelaps (Dendrolaelaps) helvetiae Hirschmann, 1974: 50 [junior synonymy by

Hirschmann and Wiśniewski, 1982: 132].

TYPE DEPOSITORY: D. frenzeli: Unknown; D. helvetiae: Naturhistorisches Museum Basel,

Basel, Switzerland.

TYPE LOCALITY AND HABITAT: D. frenzeli: Poland, Hundsfeld, Wroclaw, in dry

meadow; D. helvetiae: Switzerland, Swiss National Park, S-chanf, [Maloja,

Graubünden], in soil of abandoned grassland.

153

NOTE: Dendrolaelaps (Dendrolaelaps) helvetiae Hirschmann, 1974 was described on the

basis of specimens reported by Schweizer (1961): 138 as Digamasellus oudemansi

(Halbert, 1915).

052. Dendrolaelaps fukikoae Ishikawa, 1977

Dendrolaelaps fukikoae Ishikawa, 1977: 102.

Dendrolaelaps (Cornodendrolaelaps) fukikoae.— Hirschmann and Wiśniewski, 1982: 97.

TYPE DEPOSITORY: Biological Laboratory of Matsuyama Shinonome Junior College,

Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, Sugeta, Ōzu, Ehime Prefecture, 3 July 1976, on

Monochamus alternatus [Coleoptera: Cerambycidae].

053. Dendrolaelaps glareoli Wiśniewski and Hirschmann, 1984

Dendrolaelaps (Cornodendrolaelaps ?) glareoli [sic] Wiśniewski and Hirschmann, in

Hirschmann and Wiśniewski, 1984: 94


TYPE LOCALITY AND HABITAT: Poland, Wielkopolska National Park, [Wielkopolska],

17 July 1970, on hairs of Clethrionomys glareolus [Mammalia: Rodentia: Cricetidae].


054. Dendrolaelaps halaskovae Schmölzer, 1995

Dendrolaelaps (Punctodendrolaelaps) halaškovae Schmölzer, 1995b: 99.


TYPE LOCALITY AND HABITAT: Austria, Vellacher Kotschna Reserve, Carinthia, 2 July

1992, in soil under Alnetum incanae [Betulaceae] from a mixed pine forest, and in a

humid area with Brachypodium silvaticum [Poaceae], Aegopodium podagraria

[Apiaceae] and Taraxacum paludosum [Asteraceae].

055. Dendrolaelaps halophilus (Willmann, 1951)

Digamasellus halophilus Willmann, 1951: 143.

Digamasellus halophilus.— Hirschmann, 1954a: 108.

Dendrolaelaps halophilus.— Hirschmann, 1960: 7; Hirschmann, 1971b: 13; Hirschmann,


Karg, 1971: 330; Shcherbak, 1980: 120; Karg, 1993c: 348.

154

Cyrtolaelaps (Digamasellus) halophilus.— Ryke, 1962a: 110.

Dendrolaelaps (Dendrolaelaps) halophilus.— Hirschmann, 1974: 62; Hirschmann and



TYPE LOCALITY AND HABITAT: Germany, Westturm, Wangerooge Island, [Friesland], 6

October 1949, on Salicornia sp. [Amaranthaceae] in a salt marsh outside the bird

sanctuary.

056. Dendrolaelaps heterotrichus Hirschmann, 1960

Dendrolaelaps heterotrichus Hirschmann, 1960: 9.

Dendrolaelaps (Dendrolaelaps) heterotrichus.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) heterotrichus.— Hirschmann and Wiśniewski,

1982: 112.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stump of Picea sp. [Pinaceae]

and in galleries of Hylurgops palliates [Coleoptera: Curculionidae: Scolytinae].


057. Dendrolaelaps hunteri Wiśniewski, 1979

Dendrolaelaps hunteri Wiśniewski, 1979a: 158.

Dendrolaelaps (Dendrolaelaps) hunteri.— Hirschmann and Wiśniewski, 1982: 88.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Swietokrzyskie Nationalpark, Debno, 7

November 1974, under bark of Abies alba [Pinaceae] in galleries of Pityokteines

vorontzovi [Coleoptera: Curculionidae: Scolytinae].


058. Dendrolaelaps hurlbutti Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) hurlbutti Hirschmann and Wiśniewski, 1982: 87.


TYPE LOCALITY AND HABITAT: USA, Bowie, Maryland, 15 April 1960, from



155

059. Dendrolaelaps imitopraetarsalis Ma and Lin, 2005

Dendrolaelaps imitopraetarsalis Ma and Lin, 2005: 350.

TYPE DEPOSITORY: Institute of Plant Protection of Fujian Academy of Agricultural

Science, Fuzhou, China.

TYPE LOCALITY AND HABITAT: China, Songshan Mountain (34.55°N, 113.05°E),

Dengfeng County, Henan Province, 17 July 2002, in nest of magpie [Aves: Corvidae].

060. Dendrolaelaps insignis Hirschmann, 1960

Dendrolaelaps insignis Hirschmann, 1960: 8.

Dendrolaelaps insignis.— Hirschmann, 1971b: 13; Hirschmann, 1971d: 19; Karg, 1971: 338.

Dendrolaelaps (Dendrolaelaps) insignis.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) insignis.— Hirschmann and Wiśniewski, 1982: 51.

Punctodendrolaelaps insignis.— Karg, 1993c: 353.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stump of Picea sp. [Pinaceae].

NOTE: Shcherbak (1980): 101 considered D. insignis as junior synonymy of Dendrolaelaps

arvicolus (Leitner, 1949), but Hirschmann and Wiśniewski (1982): 51 revoked that

synonymy.

061. Dendrolaelaps isochetus Shcherbak and Bregetova, 1980

Dendrolaelaps isochetus Shcherbak and Bregetova, in Shcherbak, 1980: 164.

Dendrolaelaps (Foveodendrolaelaps ?) isochetus [sic].— Hirschmann and Wiśniewski, 1982:

126.

Dendrolaelaps isochetus.— Karg, 1993c: 348.



TYPE LOCALITY AND HABITAT: Uzbekistan, Dzharkurgan, 15 June 1966, in litter of

saksaul Haloxylon sp. [Amaranthaceae] in a desert.

062. Dendrolaelaps krantzi Wiśniewski, 1979

Dendrolaelaps krantzi Wiśniewski, 1979a: 152.

Dendrolaelaps (Dendrolaelaps) krantzi.— Hirschmann and Wiśniewski, 1982: 87.

156


Poland.

TYPE LOCALITY AND HABITAT: Poland, Potasze, Poznań County, 13 June 1974, under

bark of Pinus sylvestris [Pinaceae] in galleries of Myelophilus piniperda [Coleoptera:

Curculionidae: Scolytinae].


063. Dendrolaelaps laetus Shcherbak, 1980

Dendrolaelaps laetus Shcherbak, 1980: 120.

Dendrolaelaps (Dendrolaelaps) laetus.— Hirschmann and Wiśniewski, 1982: 85.



TYPE LOCALITY AND HABITAT: Ukraine, Yunakovskoe Forest, Sumy Oblast, 17 July

1971, on a rotten stump of Quercus sp. [Fagaceae].

064. Dendrolaelaps langi Hirschmann and Wiśniewski, 1984

Dendrolaelaps (Cornodendrolaelaps) langi Hirschmann and Wiśniewski, 1984: 91.


TYPE LOCALITY AND HABITAT: Vietnam, Lai Khe, 50 km north of Ho Chi Minh City, 2

June 1968, on legs and prothorax of longhorn beetle [Coleoptera: Cerambycidae] from

a rubber [Euphorbiaceae] plantation.

NOTE1: Reported as Digamasellus sp. by Lang (1978): 58.

NOTE2: Described from the deutonymph.

065. Dendrolaelaps lasiophilus Hirschmann, 1960

Dendrolaelaps lasiophilus Hirschmann, 1960: 8.

Dendrolaelaps lasiophilus.— Hirschmann, 1971d: 19; Karg, 1971: 338.

Dendrolaelaps (Dendrolaelaps) lasiophilus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) lasiophilus [sic].— Hirschmann and Wiśniewski,

1982: 106.

Cornodendrolaelaps lasiophilus.— Karg, 1993c: 362.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, in nest of Lasius fuliginosus


157

NOTE: Shcherbak (1980): 156 considered D. lasiophilus as junior synonymy of

Dendrolaelaps populi Hirschmann, 1960, but Hirschmann and Wiśniewski (1982):

106 revoked that synonymy.

066. Dendrolaelaps latior (Leitner, 1949)

Digamasellus latior Leitner, 1949: 59.

Digamasellus latior.— Franz, 1954: 342.

Dendrolaelaps latior.— Hirschmann, 1960: 8; Hirschmann, 1971d: 19; Karg, 1971: 338.

Cyrtolaelaps (Digamasellus) latior.— Ryke, 1962a: 107.

Dendrolaelaps (Dendrolaelaps) latior.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) latior.— Hirschmann and Wiśniewski, 1982: 57.

Punctodendrolaelaps latior.— Karg, 1993c: 353; Huhta and Karg, 2010: 341.


TYPE LOCALITY AND HABITAT: Austria, Admont, [Styria], in old compost.

NOTE: Specimens reported by Shcherbak (1980): 103 as Dendrolaelaps latior (Leitner,

1949) were described as Dendrolaelaps (Punctodendrolaelaps) latioroides


067. Dendrolaelaps latioroides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps) latioroides Hirschmann and Wiśniewski, 1982: 57.



TYPE LOCALITY AND HABITAT: Russia, Novosibirsk and Chita Oblasts, on decaying

plant debris, hay, straw, leaves, fruits, soil and in ant nest [Hymenoptera].


Dendrolaelaps latior (Leitner, 1949).

068. Dendrolaelaps latoides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Cornodendrolaelaps) latoides Hirschmann and Wiśniewski, 1982: 97.



TYPE LOCALITY AND HABITAT: Russia, Voronezh Oblast; Russia, Buryatia; Russia,

Ussuriysk, Primorsky Krai, under bark of Picea sp. [Pinaceae] and Quercus sp.

[Fagaceae].

158


Dendrolaelaps latus Hirschmann, 1960.

069. Dendrolaelaps latus Hirschmann, 1960

Dendrolaelaps latus Hirschmann, 1960: 8.

Dendrolaelaps latus.— Hirschmann, 1971b: 13; Hirschmann, 1971d: 19; Karg, 1971: 338.

Dendrolaelaps (Dendrolaelaps) latus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) latus.— Hirschmann and Wiśniewski, 1982: 97.

Cornodendrolaelaps latus.— Karg, 1993c: 362.


TYPE LOCALITY AND HABITAT: Germany, Zell, Schwarzwald, on stumps of Abies sp.

[Pinaceae] and Fagus sp. [Fagaceae]; Germany, Bielefeld, on stump of Betula sp.

[Betulaceae]; Sweden, Jämtland, on bark of Salix caprea [Salicaceae].

NOTE: Specimens reported by Shcherbak (1980): 125 as Dendrolaelaps latus Hirschmann,

1960 were described as Dendrolaelaps (Cornodendrolaelaps) latoides Hirschmann


070. Dendrolaelaps lemani (Schweizer, 1961)

Digamasellus lemani Schweizer, 1961: 139.

Dendrolaelaps (Dendrolaelaps) lemani.— Hirschmann, 1974: 63.

Dendrolaelaps (Cornodendrolaelaps) lemani.— Hirschmann and Wiśniewski, 1982: 101.


TYPE LOCALITY AND HABITAT: Switzerland, Rhone River Delta, Villeneuve, 20 May

1918, under tree bark.

071. Dendrolaelaps lindquisti Wiśniewski, 1979

Dendrolaelaps lindquisti Wiśniewski, 1979a: 155.

Dendrolaelaps (Dendrolaelaps) cornutolindquisti Hirschmann and Wiśniewski, 1982: 87

[objective synonymy — unjustified replacement name].


Poland.

TYPE LOCALITY AND HABITAT: Poland, Potasze, Poznań County, 20 July, 1975, under

bark of Picea abies [Pinaceae] in galleries of Dryocoetes autographus [Coleoptera:


159


072. Dendrolaelaps liujingyuani Ma, 2008

Dendrolaelaps liujingyuani Ma, 2008: 129.



TYPE LOCALITY AND HABITAT: China, Lushan Mountain (29°32‘N, 115°55‖E), Jiangxi

Province, 29 August 1983, under tree bark.

073. Dendrolaelaps longiductus Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Cornodendrolaelaps) longiductus Wiśniewski and Hirschmann, 1993c: 331.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Port of Szczecin, 14 December 1987,

intercepted on wood of Entandrophragma cylindricum [Meliaceae] imported from

Cameroon.

074. Dendrolaelaps longifallax Hirschmann, 1960

Dendrolaelaps longifallax Hirschmann, 1960: 8.

Dendrolaelaps (Dendrolaelaps) longifallax.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) longifallax.— Hirschmann and Wiśniewski, 1982: 54.

Punctodendrolaelaps longifallax.— Karg, 1993c: 352.


TYPE LOCALITY AND HABITAT: Switzerland, Pontresina, [Maloja], on stump of Larix

sp. [Pinaceae].

075. Dendrolaelaps longiusculus (Leitner, 1949)

Digamasellus longiusculus Leitner, 1949: 61.

Digamasellus longiusculus.— Franz, 1954: 342.

Dendrolaelaps longiusculus.— Hirschmann, 1960: 7; Hirschmann, 1971c: 16; Hirschmann,

1971d: 17; Hirschmann, 1971e: 20; Karg, 1971: 328; Shcherbak, 1980: 131.

Cyrtolaelaps (Digamasellus) longiusculus.— Ryke, 1962a: 108.

Dendrolaelaps (Dendrolaelaps) longiusculus.— Hirschmann, 1974: 62.

160

Dendrolaelaps (Cornodendrolaelaps) longiusculus.— Hirschmann and Wiśniewski, 1982:

103.

Cornodendrolaelaps longiusculus.— Karg, 1993c: 354.


TYPE LOCALITY AND HABITAT: Austria, Admont, [Styria], in compost.

NOTE: Reported by Shcherbak (1980): 131 and Karg (1971): 328 as Dendrolaelaps

longiusculus Hirschmann, 1960, and by Karg (1993c): 354 as Cornodendrolaelaps

longiusculus (Hirschmann, 1960).

076. Dendrolaelaps louisianae Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps ?) louisianae [sic] Hirschmann and Wiśniewski, 1982:

62.


TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, 26 April 1966, phoretic

under the wing of Hylobius pales [Coleoptera: Curculionidae: Molytinae].


077. Dendrolaelaps lusikisikiae Hirschmann and Wiśniewski, 1984

Dendrolaelaps (Cornodendrolaelaps) lusikisikiae Hirschmann and Wiśniewski, 1984: 91.


TYPE LOCALITY AND HABITAT: South Africa, Lusikisiki, Transkei, 1979, from a coastal

evergreen forest.


078. Dendrolaelaps luxtoni Wiśniewski and Hirschmann, 1989

Dendrolaelaps (Luxtondendrolaelaps) luxtoni Wiśniewski and Hirschmann, 1989c: 326.


TYPE LOCALITY AND HABITAT: Australia, Valva, New South Wales, 22 June 1983, in

wet kelp [Phaeophyceae].

079. Dendrolaelaps macfarlanei (Ryke, 1962)

Cyrtolaelaps (Digamasellus) macfarlanei Ryke, 1962a: 100.

Dendrolaelaps (Dendrolaelaps) macfarlanei.— Hirschmann, 1974: 63; Hirschmann and


161



TYPE LOCALITY AND HABITAT: England, Roudsea Wood and Mosses National Nature

Reserve, Cumbria, 1957, on oak Quercus sp. [Fagaceae].

080. Dendrolaelaps magnus Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Stanidendrolaelaps) magnus Wiśniewski and Hirschmann, 1993b: 305.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Port of Szczecin, 19 April 1985, intercepted

under bark of Entandrophragma cylindricum [Meliaceae] imported from Cameroon.

081. Dendrolaelaps majesticus Wiśniewski and Hirschmann, 1989

Dendrolaelaps (Majestidendrolaelaps) majesticus Wiśniewski and Hirschmann, 1989b: 317.


Poland.

TYPE LOCALITY AND HABITAT: Cuba, Botanical Garden, Cienfuegos, 4 April 1987,

under tree bark.

082. Dendrolaelaps markewitschi Shcherbak, 1980

Dendrolaelaps markewitschi Shcherbak, 1980: 151.

Dendrolaelaps (Foveodendrolaelaps) markewitschi.— Hirschmann and Wiśniewski, 1982:

129.



TYPE LOCALITY AND HABITAT: Ukraine, Dnieper River, Cherkasy Oblast, 29 July 1976,

in nest of Riparia riparia [Aves: Hirundinidae] on a sandy soil by a river.

083. Dendrolaelaps marylandae (Hurlbutt, 1967)

Digamasellus marylandae Hurlbutt, 1967: 509.

Dendrolaelaps (Dendrolaelaps) marylandae.— Hirschmann, 1974: 63.

Dendrolaelaps (Sellnickidendrolaelaps) marylandae.— Hirschmann and Wiśniewski, 1982:

65.

162

TYPE DEPOSITORY: Smithsonian National Museum of Natural History, Washington,

District of Columbia, USA.

TYPE LOCALITY AND HABITAT: USA, Patuxent Wildlife Research Refuge, Maryland, 4

September 1959, in humus from a deciduoud forest.

084. Dendrolaelaps medius Shcherbak, 1980

Dendrolaelaps medius Shcherbak, 1980: 161.

Dendrolaelaps (Cornodendrolaelaps) medius.— Hirschmann and Wiśniewski, 1982: 106.



TYPE LOCALITY AND HABITAT: Russia, Ussurisky State Nature Reserve, Primorsky

Krai, 15 July 1977, on rotten wood of Ulmus sp. [Ulmaceae].

085. Dendrolaelaps metwallyi El-Halawany and Abdel-Samad, 1991

Dendrolaelaps metwallyi El-Halawany and Abdel-Samad, 1991: 180.


TYPE LOCALITY AND HABITAT: Egypt, Sidi Krier District, Alexandria, in debris under

fig trees.

086. Dendrolaelaps modestus Barilo, 1989

Dendrolaelaps (Dendrolaelaps) modestus Barilo, 1989: 138.



TYPE LOCALITY AND HABITAT: Uzbekistan, Samarkand, Samarqand Province, 10

October 1985, in litter.

087. Dendrolaelaps monodentatus Wiśniewski and Hirschmann, 1989

Dendrolaelaps (Monodendrolaelaps) monodentatus Wiśniewski and Hirschmann, 1989d:

117.

Dendrolaelaps monodentatus.— Karg, 1993c: 350; Huhta and Karg, 2010: 346.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Bielinek Nature Reserve, [Gmina Cedynia], 10

April 1988, in litter of Quercus sp. [Fagaceae].

163

088. Dendrolaelaps moseri (Hurlbutt, 1967)

Digamasellus moseri Hurlbutt, 1967: 514.

Dendrolaelaps (Dendrolaelaps) moseri.— Hirschmann, 1974: 63; Hirschmann and

Wiśniewski, 1982: 77; Shcherbak, 1984: 40.



TYPE LOCALITY AND HABITAT: USA, Delaware, Ohio, 1 February 1965, in galleries of

Scolytus multistriatus [Coleoptera: Curculionidae: Scolytinae].

089. Dendrolaelaps moserisimilis Shcherbak, 1984

Dendrolaelaps moserisimilis Shcherbak, 1984: 35.



TYPE LOCALITY AND HABITAT: Ukraine, Poltava, 4 April 1976, under bark of a poplar

Populus sp. [Salicaceae].

090. Dendrolaelaps myiaphilus (Karg, 2002)

Punctodendrolaelaps myiaphilus Karg, 2002: 238.


TYPE LOCALITY AND HABITAT: Sweden, Tyresta National Park, Stockholm, 14 April-5

June 2000, on the gall midge Polyardis silvalis [Diptera: Cecidomyiidae] from a

devastated forest area.


091. Dendrolaelaps myrmecophilus Hirschmann, 1960

Dendrolaelaps myrmecophilus Hirschmann, 1960: 17.

Dendrolaelaps myrmecophilus.— Hirschmann, 1971b: 12; Hirschmann, 1971c: 15;

Hirschmann, 1971d: 18; Hirschmann, 1971f: 26; Karg, 1971: 338; Shcherbak, 1980:

133.

Dendrolaelaps (Dendrolaelaps) myrmecophilus.— Hirschmann, 1974: 61.

Dendrolaelaps (Cornodendrolaelaps ?) myrmecophilus [sic].— Hirschmann and Wiśniewski,

1982: 104.

Cornodendrolaelaps myrmecophilus.— Karg, 1993c: 362.

164




092. Dendrolaelaps natans (Pintchuk, 1972)

Leioseius natans Pinchuk, 1972a: 60.

TYPE DEPOSITORY: Institute of Zoology and Parasitology SSR Moldova, Kishinev,

Moldova.

TYPE LOCALITY AND HABITAT: Moldova, Pashkany, 3 December 1968, in nest of Sorex

araneus [Mammalia: Soricidae].

093. Dendrolaelaps neocornutus (Hurlbutt, 1967)

Digamasellus neocornutus Hurlbutt, 1967: 510.

Dendrolaelaps neocornutus.— McGraw and Farrier, 1969: 111.

Dendrolaelaps (Dendrolaelaps) neocornutus.— Hirschmann, 1974: 63; Hirschmann and




TYPE LOCALITY AND HABITAT: USA, Knoxville, Mississipi, 27 November 1964, in

galleries of Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] in Pinus

taeda [Pinaceae].

094. Dendrolaelaps neodisetosimilis McGraw and Farrier, 1969

Dendrolaelaps neodisetosimilis McGraw and Farrier, 1969: 115.

Dendrolaelaps (Dendrolaelaps) neodisetosimilis.— Hirschmann, 1974: 63.

Dendrolaelaps (Apophyseodendrolaelaps) neodisetosimilis.— Hirschmann and Wiśniewski,

1982: 112.


TYPE LOCALITY AND HABITAT: USA, Yadkin County, North Carolina, 30 May 1966,

on Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] attacking Pinus

echinata [Pinaceae].

095. Dendrolaelaps neodisetus (Hurlbutt, 1967)

Digamasellus neodisetus Hurlbutt, 1967: 518.

165

Dendrolaelaps neodisetus.— McGraw and Farrier, 1969: 117.

Dendrolaelaps (Dendrolaelaps) neodisetus.— Hirschmann, 1974: 63.

Dendrolaelaps (Disetodendrolaelaps) neodisetus.— Hirschmann and Wiśniewski, 1982: 117.



TYPE LOCALITY AND HABITAT: USA, Bude, Mississipi, 29 September 1964, in galleries

of Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] in Pinus taeda

[Pinaceae].

096. Dendrolaelaps neozwoelferi Hirschmann, 1983

Dendrolaelaps (Apophyseodendrolaelaps) neozwoelferi Hirschmann, 1983c: 128.


TYPE LOCALITY AND HABITAT: Unspecified type locality and substrate.

NOTE: Described on the basis of specimens reported by Baker et al. (1958): 58 as

Digamasellidae.

097. Dendrolaelaps nikolai Shcherbak, 1978

Dendrolaelaps nikolai Shcherbak, 1978b: 665.

Dendrolaelaps nikolai.— Shcherbak, 1980: 149.

Dendrolaelaps (Foveodendrolaelaps) nikolai.— Hirschmann and Wiśniewski, 1982: 128.



TYPE LOCALITY AND HABITAT: Russia, Selenge River, Kabansky District, Buryatia, 26

July 1977, from an old aspen [Populus sp. (Salicaceae)] in a river floodplain.

098. Dendrolaelaps ningxiaensis Ma and Bai, 2009

Dendrolaelaps ningxiaensis Ma and Bai, 2009: 75.



TYPE LOCALITY AND HABITAT: China, Yinchuan (38°28‘N, 106°32‘E), Ningxia Hui

Autonomous Region, 9 April 1990, in a hayrick.

099. Dendrolaelaps nostricornutus Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) nostricornutus Hirschmann and Wiśniewski, 1982: 80.

166

Dendrolaelaps nostricornutus.— Karg, 1993c: 349.


TYPE LOCALITY AND HABITAT: Germany, under bark of Acer sp. [Sapindaceae],

Robinia sp. [Fabaceae], Tilia sp. [Malvaceae] and Ulmus sp. [Ulmaceae]; on stumps of

Carpinus sp. [Betulaceae], Fagus sp. [Fabaceae], Picea sp. [Pinaceae], Pinus sp.

[Pinaceae], Quercus sp. [Fabaceae] and Sorbus sp. [Rosaceae]; in galleries of

Ernopocerus fagi, Ips sexdentatus, Leperesinus fraxini, Pissodes piceae, Scolytus

scolytus and Tomicus piniperda [all Coleoptera: Curculionidae: Scolytinae].

NOTE: Described on the basis of specimens reported by Hirschmann (1960): 6 as

Dendrolaelaps cornutus (Kramer, 1886).

100. Dendrolaelaps oblitus Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) oblitus Hirschmann and Wiśniewski, 1982: 84.


TYPE LOCALITY AND HABITAT: Germany, on unspecified substrate.

101. Dendrolaelaps oligochetus Shcherbak, 1980

Dendrolaelaps oligochetus Shcherbak, 1980: 132.

Dendrolaelaps (Cornodendrolaelaps) oligochetus.— Hirschmann and Wiśniewski, 1982:

103.



TYPE LOCALITY AND HABITAT: Ukranie, Lyutezh, Kiev Oblast, 26 July 1976, in soil (0-

5 cm) from a mixed forest.

102. Dendrolaelaps ophidiotrichus Luxton, 1982

Dendrolaelaps ophidiotrichus Luxton, 1982: 331.

TYPE DEPOSITORY: Entomology Division, DSIR, Auckland, New Zealand.

TYPE LOCALITY AND HABITAT: New Zealand, Waikato, North Island, July 1967, from

peat pasture soils.

103. Dendrolaelaps opticus Barilo, 1989

Dendrolaelaps (Dendrolaelaps) opticus Barilo, 1989: 140.

167




November 1984, in cow manure.

104. Dendrolaelaps oudemansi Halbert, 1915

Dendrolaelaps oudemansi Halbert, 1915: 68.

Dendrolaelaps (Dendrolaelaps) oudemansi.— Vitzthum, 1926b: 424.

Dendrolaelaps oudemansi.— Hirschmann, 1960: 7.

Cyrtolaelaps (Digamasellus) oudemansi.— Ryke, 1962a: 103.

Dendrolaelaps (Dendrolaelaps) oudemansi.— Hirschmann, 1974: 61; Hirschmann and


TYPE DEPOSITORY: The National Museum, Dublin, Ireland.

TYPE LOCALITY AND HABITAT: Ireland, Westport, [Mayo County], under bark of

decayed trees; Ireland, Achill Island, on fallen pinecones; Ireland, Friarstown, under

bark of fir Abies sp. [Pinaceae] trees.

NOTE1: Specimens reported by Schweizer (1961): 138 as Digamasellus oudemansi (Halbert,

1915) were described as Dendrolaelaps (Dendrolaelaps) helvetiae Hirschmann, 1974:

50.

NOTE2: Specimens reported by Shcherbak (1980): 113 as Dendrolaelaps oudemansi Halbert,

1915 were described as Dendrolaelaps (Dendrolaelaps) oudemansiformis Hirschmann


NOTE3: Shcherbak (1980): 118 considered Digamasellus septentrionalis Sellnick, 1958 as

junior synonymy of D. oudemansi, but Hirschmann and Wiśniewski (1982): 77


NOTE4: Shcherbak (1980): 118 considered Dendrolaelaps acornutosimilis Hirschmann, 1960

as junior synonymy of D. oudemansi, but Hirschmann and Wiśniewski (1982): 77


105. Dendrolaelaps oudemansiformis Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) oudemansiformis Hirschmann and Wiśniewski, 1982: 76.



168

TYPE LOCALITY AND HABITAT: Russia, Chukotka Autonomous Okrug, in litter of

coniferous forests, under bark of old stumps of different tree species, in humus and

grass debris.


Dendrolaelaps oudemansi Halbert, 1915.

106. Dendrolaelaps papuae Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps ?) papuae [sic] Hirschmann and Wiśniewski, 1982: 61.


TYPE LOCALITY AND HABITAT: New Guinea, on Batocera sp. [Coleoptera:

Cerambycidae].


107. Dendrolaelaps paradoxa Shcherbak, 1982

Dendrolaelaps paradoxa Shcherbak, 1982b: 74.

Dendrolaelaps (Foveodendrolaelaps ?) paradoxa [sic].— Hirschmann and Wiśniewski, 1982:

131.



TYPE LOCALITY AND HABITAT: Ukraine, Kaniv, Cherkasy Oblast, 9 June 1977, in litter

and nest of Riparia riparia [Aves: Hirundinidae].

108. Dendrolaelaps passalorum (Pearse and Wharton, 1936)

Zercon passalorum Pearse and Wharton, in Pearse et al., 1936: 477.

Dendrolaelaps passalorum.— Delfinado and Baker, 1975: 50.

Dendrolaelaps (Cornodendrolaelaps) passalorum.— Hirschmann and Wiśniewski, 1982:

100.



TYPE LOCALITY AND HABITAT: USA, Duke Forest, Durham County, North Carolina, 19

August 1934, under wings of Odontotaenius disjunctus (cited as Passalus cornutus)

[Coleoptera: Passalidae].

109. Dendrolaelaps peruensis Wiśniewski and Hirschmann, 1989

169

Dendrolaelaps (Cornodendrolaelaps) peruensis Wiśniewski and Hirschmann, 1989a: 327.


TYPE LOCALITY AND HABITAT: Peru, Qiroz Junin, 5 November 1933, on unidentified

Curculionidae [Coleoptera] deposited in the beetle collection of ―Drozda Kollection‖

of Muzeum Górnoślaskie, Bytom, Poland.


110. Dendrolaelaps piriformis Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Punctodendrolaelaps) piriformis Hirschmann and Wiśniewski, 1982: 52.



TYPE LOCALITY AND HABITAT: Russia, Buryatia, on old stumps of Cedrus sp.

[Pinaceae], Fagus sp. [Fagaceae], Larix sp. [Pinaceae] and Prunus cerasifera

[Rosaceae].


Dendrolaelaps trapezoides Hirschmann, 1960.

111. Dendrolaelaps populi Hirschmann, 1960

Dendrolaelaps populi Hirschmann, 1960: 7.

Dendrolaelaps populi.— Hirschmann, 1971c: 15; Hirschmann, 1971d: 18; Hirschmann,

1971e: 21; Karg, 1971: 336.

Dendrolaelaps (Dendrolaelaps) populi.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) populi [sic].— Hirschmann and Wiśniewski, 1982:

105.

Cornodendrolaelaps populi.— Karg, 1993c: 359.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, on stump of Populus sp.

[Salicaceae].

NOTE1: Specimens reported by Shcherbak (1980): 156 as Dendrolaelaps populi Hirschmann,

1960 were described as Dendrolaelaps (Cornodendrolaelaps ?) populoides [sic]


NOTE2: Shcherbak (1980): 156 considered Dendrolaelaps lasiophilus Hirschmann, 1960 as

junior synonymy of D. populi, but Hirschmann and Wiśniewski (1982): 106 revoked

that synonymy.

170

112. Dendrolaelaps populoides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Cornodendrolaelaps ?) populoides [sic] Hirschmann and Wiśniewski, 1982:

105.



TYPE LOCALITY AND HABITAT: Ukraine, on stump of Quercus sp. [Fagaceae] and in

nest of Lasius fuliginosus [Hymenoptera: Formicidae].


Dendrolaelaps populi Hirschmann, 1960.

113. Dendrolaelaps posnaniensis Wiśniewski and Hirschmann, 1984

Dendrolaelaps (Cornodendrolaelaps ?) posnaniensis [sic] Wiśniewski and Hirschmann, in

Hirschmann and Wiśniewski, 1984: 93


TYPE LOCALITY AND HABITAT: Poland, Poznań, 15 January 1984, in litter of Populus

sp. [Salicaceae].

114. Dendrolaelaps praetarsalis Wiśniewski and Hirschmann, 1985

Dendrolaelaps praetarsalis Wiśniewski and Hirschmann, 1985: 71.

TYPE DEPOSITORY: Zoologischen Museum, Universität Hamburg, Hamburg, Germany.

TYPE LOCALITY AND HABITAT: Germany, Hafen, Hamburg, 12 June 1962, intercepted

on Brazil nuts [Lecythidaceae] imported from Manaus, Brazil.

115. Dendrolaelaps presepum (Berlese, 1918)

Gamasellus (Digamasellus) presepum Berlese, 1918: 136.

Digamasellus presepum.— Leitner, 1949: 57; Hirschmann, 1954a: 107; Franz, 1954: 342.

Dendrolaelaps presepum.— Hirschmann, 1960: 7; Hirschmann, 1971d: 18; Hirschmann,

1971e: 20; Karg, 1971: 336; Shcherbak, 1980: 160.

Cyrtolaelaps (Digamasellus) presepum.— Ryke, 1962a: 91.

Dendrolaelaps (Dendrolaelaps) presepum.— Hirschmann, 1974: 62.

Dendrolaelaps (Presepodendrolaelaps) presepum.— Hirschmann and Wiśniewski, 1982:

136; Wiśniewski and Hirschmann, 1983b: 124.

171

Cornodendrolaelaps presepum.— Karg, 1993c: 359; Bernini, Castagnoli and Nannelli, 1995:

19.

Leioseius codrensis Pinchuk, 1972a: 63 [junior synonymy by Shcherbak, 1980: 160].

TYPE DEPOSITORY: D. presepum: Istituto Sperimentale per la Zoologia Agraria, Florence,

Italy; L. codrensis: Institute of Zoology and Parasitology SSR Moldova, Kishinev,

Moldova.

TYPE LOCALITY AND HABITAT: D. presepum: Italy, Florence, Tuscany, in litter of grass

from a stable; L. codrensis: Ukraine, Lozovaya Forest, Kharkiv, 11 July 1969, in nest

of Muscardinus avellanarius [Mammalia: Rodentia: Gliridae].

116. Dendrolaelaps procornutoides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) procornutoides Hirschmann and Wiśniewski, 1982: 79.



TYPE LOCALITY AND HABITAT: Russia, in Leningrad Oblast and in Buryatia, under bark

of Betula sp. [Betulaceae].


Dendrolaelaps procornutus Hirschmann, 1960.

117. Dendrolaelaps procornutus Hirschmann, 1960

Dendrolaelaps procornutus Hirschmann, 1960: 8.

Dendrolaelaps procornutus.— Hirschmann, 1971d: 17; Karg, 1971: 330.

Dendrolaelaps (Dendrolaelaps) procornutus.— Hirschmann, 1974: 62; Hirschmann and

Wiśniewski, 1982: 79; Karg, 1993c: 348.


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stump of Pinus sp.

[Pinaceae].

NOTE: Specimens reported by Shcherbak (1980): 117 as Dendrolaelaps procornutus

Hirschmann, 1960 were described as Dendrolaelaps (Dendrolaelaps) procornutoides


118. Dendrolaelaps proprius Shcherbak, 1985

Dendrolaelaps proprius Shcherbak, 1985b: 68.

172



TYPE LOCALITY AND HABITAT: Russia, Maly Khamardaban montain ridge (southern

boundary), Buryatia, 17 July 1977, on birch [Betula sp. (Betulaceae)] in a mountain of

taiga.

119. Dendrolaelaps proteae (Ryke, 1962)

Cyrtolaelaps (Digamasellus) proteae Ryke, 1962a: 94.

Dendrolaelaps (Dendrolaelaps) proteae.— Hirschmann, 1974: 63.

Dendrolaelaps (Apophyseodendrolaelaps) proteae.— Hirschmann and Wiśniewski, 1982:

114.



TYPE LOCALITY AND HABITAT: South Africa, Grabouw, [Western Cape], January 1955,

from a closed dry flower of Protea mellifera [Proteaceae].

120. Dendrolaelaps punctatosimilis Hirschmann, 1960

Dendrolaelaps punctatosimilis Hirschmann, 1960: 7.

Dendrolaelaps punctatosimilis.— Hirschmann, 1971b: 13; Hirschmann, 1971c: 15;

Hirschmann, 1971d: 18; Hirschmann, 1971e: 21; Karg, 1971: 337; Shcherbak, 1980:

128.

Dendrolaelaps (Dendrolaelaps) punctatosimilis.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) punctatosimilis.— Hirschmann and Wiśniewski, 1982:

99.

Cornodendrolaelaps punctatosimilis.— Karg, 1993c: 360.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, in gallery of Ips typographus

[Coleoptera: Curculionidae: Scolytinae].

121. Dendrolaelaps punctatulus Hirschmann, 1960

Dendrolaelaps punctatulus Hirschmann, 1960: 7.

Dendrolaelaps punctatulus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 12; Hirschmann,


Karg, 1971: 338; Shcherbak, 1980: 102.

173

Dendrolaelaps (Dendrolaelaps) punctatulus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) punctatulus.— Hirschmann and Wiśniewski, 1982: 49.

Punctodendrolaelaps punctatulus.— Karg, 1993c: 353.


TYPE LOCALITY AND HABITAT: Germany, on stumps of Betula sp. [Betulaceae], Fagus

sp. [Fagaceae], Picea sp. [Pinaceae], Pinus sp. [Pinaceae] and Quercus sp. [Fagaceae];

Germany, Munich, in gallery of Hylesinus fraxini [Coleoptera: Curculionidae:

Scolytinae]; Sweden, Jämtland, in nest of Formica rufa [Hymenoptera: Formicidae].

NOTE: Sellnick (1958): 22 cited Digamasellus punctatulus Hirschmann, 1950, but did not


122. Dendrolaelaps punctatus (Hirschmann, 1954)

Digamasellus punctatus Hirschmann, 1954b: 247.

Dendrolaelaps punctatus.— Hirschmann, 1960: 7; Hirschmann, 1971b: 14; Hirschmann,


Karg, 1971: 337; Shcherbak, 1980: 127.

Dendrolaelaps (Dendrolaelaps) punctatus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) punctatus.— Hirschmann and Wiśniewski, 1982: 98.

Cornodendrolaelaps punctatus.— Karg, 1993c: 359.


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stumps of Picea sp.

[Pinaceae] and Pinus sp. [Pinaceae], in gallery of Dryocoetes autographus

[Coleoptera: Curculionidae: Scolytinae] and under the elytra of Tetropium luridum

[Coleoptera: Cerambycidae]; Germany, Flossenbürg, on stump of Picea sp.

[Pinaceae]; Austria, Admont, [Styria], on stump of Picea sp..

NOTE: Franz (1954): 342 cited Digamasellus punctatus Hirschmann n. sp., but did not


123. Dendrolaelaps puntperivi (Schweizer, 1961)

Digamasellus puntperivi Schweizer, 1961: 140.

Dendrolaelaps (Dendrolaelaps) puntperivi.— Hirschmann, 1974: 63.

Dendrolaelaps (Sellnickidendrolaelaps) puntperivi.— Hirschmann and Wiśniewski, 1982: 66.

Punctodendrolaelaps puntperivi.— Karg, 1993c: 352.


174

TYPE LOCALITY AND HABITAT: Switzerland, the waterfront of Spöl River, Punt Periv,

Swiss National Park, in drenched moss.

124. Dendrolaelaps quadricrinus (Berlese, 1920)

Gamasellus (Digamasellus) quadricrinus Berlese, 1920b: 162.

Dendrolaelaps quadricinus [sic].— Hirschmann, 1960: 8.

Cyrtolaelaps (Digamasellus) quadricrinus.— Ryke, 1962a: 105.

Dendrolaelaps (Dendrolaelaps) quadricinus [sic].— Hirschmann, 1974: 62.

Dendrolaelaps (Punctodendrolaelaps) quadricrinus.— Hirschmann and Wiśniewski, 1982 :

59.


TYPE LOCALITY AND HABITAT: USA, Lake City, Columbia County, Florida, in humus.

125. Dendrolaelaps quadripilus (Berlese, 1920)

Gamasellus quadripilus Berlese, 1920b: 159.

Dendrolaelaps quadripilus.— Hirschmann, 1960: 8.

Cyrtolaelaps (Gamasellus) quadripilus.— Ryke, 1962b: 53.

Dendrolaelaps (Multidendrolaelaps) quadripilus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) quadripilus.— Hirschmann and Wiśniewski, 1982: 59.

Gamasellus quadripilus.— Bernini, Castagnoli and Nannelli, 1995: 20.


TYPE LOCALITY AND HABITAT: Italy, San Vincenzo and Pisa, Tuscany, in humus.

126. Dendrolaelaps quadritorus (Robillard, 1971)

Digamasellus quadritorus Robillard, 1971: 1763.

Dendrolaelaps quadritorus.— Lindquist, 1975: 32.

Dendrolaelaps (Cornodendrolaelaps) quadritorus.— Hirschmann and Wiśniewski, 1982:

100.



TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, 28 June 1967, from inner

bark of Pinus elliotii [Pinaceae] with Ips grandicollis [Coleoptera: Curculionidae:

Scolytinae].

175

127. Dendrolaelaps rackae Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Sellnickidendrolaelaps) rackae Hirschmann and Wiśniewski, 1982: 65.

TYPE DEPOSITORY: Zoologischen Museum, Universität Hamburg, Hamburg, Germany.

TYPE LOCALITY AND HABITAT: Germany, Obergünzburg, [Ostallgäu], 7-11 September

1973, on Cheilotrichia cinerascens [Diptera: Limoniidae]; Germany, Eichholz (near

Kempten), [Allgäu], 15 September 1973, on C. cinerascens; Germany, Kempten,

[Allgäu], 30 June 1962, on C. cinerascens; Austria, Lunz am See, 27 June 1970, on C.

cinerascens; Germany, Birgsau, Allgäu Alps, 15-19 September 1974, on Cheilotrichia

staryi [Diptera: Limoniidae].


128. Dendrolaelaps rasmii Nasr and Mersal, 1986

Dendrolaelaps rasmii Nasr and Mersal, 1986 apud Abou-El-Soud and Shoeib, 2000: 45.



129. Dendrolaelaps rectus Karg, 1962

Dendrolaelaps rectus Karg, 1962: 36.

Dendrolaelaps rectus.— Karg, 1971: 332; Karg, 1993c: 351.

Dendrolaelaps (Dendrolaelaps) rectus.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) rectus.— Hirschmann and Wiśniewski, 1982: 134.


Germany.

TYPE LOCALITY AND HABITAT: Germany, Experimental Field of Biologische

Zentralanstalt of Teltow, [Potsdam-Mittelmark, Brandenburg], in grassland soil.

NOTE: Shcherbak (1980): 141 considered D. rectus as junior synonymy of Dendrolaelaps

crassitarsalis (Willmann, 1951), but Hirschmann and Wiśniewski (1982): 134


130. Dendrolaelaps remotus Karg, 1977

Dendrolaelaps remotus Karg, 1977: 347.

? Dendrolaelaps remotus [sic].— Hirschmann and Wiśniewski, 1982: 145.

TYPE DEPOSITORY: Ungarischess Naturwissenschaftliches Museum, Budapest, Hungary.

176

TYPE LOCALITY AND HABITAT: Argentina, valley of Rio Azul, El Bolson, Rio Negro

Province, 8 August 1961, in litter from a Nothofagus dombeyi [Nothofagaceae] forest.

131. Dendrolaelaps reticulatus (Berlese, 1920)

Gamasellus (Digamasellus) reticulatus Berlese, 1920b: 161.

Cyrtolaelaps (Digamasellus) reticulatus.— Ryke, 1962a: 103.

Dendrolaelaps (Dendrolaelaps) reticulatus.— Hirschmann, 1974: 62.

Digamasellus reticulatus.— Bernini, Castagnoli and Nannelli, 1995: 20.


TYPE LOCALITY AND HABITAT: Italy, Florence, in litter with coniferous bark.


132. Dendrolaelaps reticulosus Hirschmann, 1960

Dendrolaelaps reticulosus Hirschmann, 1960: 7.

Dendrolaelaps reticulosus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 14; Hirschmann,


Karg, 1971: 336; Shcherbak, 1980: 159.

Dendrolaelaps (Dendrolaelaps) reticulosus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) reticulosus [sic].— Hirschmann and Wiśniewski,

1982: 104.

Cornodendrolaelaps reticulosus.— Karg, 1993c: 355.


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stumps of Picea sp.

[Pinaceae] and Pinus sp. [Pinaceae]; Germany, Erlangen, in compost.

133. Dendrolaelaps rotoni (Hurlbutt, 1967)

Digamasellus rotoni Hurlbutt, 1967: 508.

Dendrolaelaps rotoni.— McGraw and Farrier, 1969: 128.

Dendrolaelaps (Dendrolaelaps) rotoni.— Hirschmann, 1974: 63.

Dendrolaelaps (Punctodendrolaelaps) rotoni.— Hirschmann and Wiśniewski, 1982: 55.



177

TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, in galleries of

Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] in Pinus taeda

[Pinaceae]; USA, Hardin, Texas, 2 September 1963, on inner bark of Pinus taeda.

134. Dendrolaelaps rotundus Hirschmann, 1960

Dendrolaelaps rotundus Hirschmann, 1960: 8.

Dendrolaelaps rotundus.— Hirschmann, 1971d: 19; Karg, 1971: 338; Shcherbak, 1980: 106.

Dendrolaelaps (Dendrolaelaps) rotundus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) rotundus.— Hirschmann and Wiśniewski, 1982: 58.

Punctodendrolaelaps rotundus.— Karg, 1993c: 354; Huhta and Karg, 2010: 341.




135. Dendrolaelaps ruhmi Hirschmann, 1972

Dendrolaelaps rühmi Hirschmann, 1972: 31.

Dendrolaelaps (Dendrolaelaps) rühmi.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) rühmi [sic].— Hirschmann and Wiśniewski, 1982:

106.


TYPE LOCALITY AND HABITAT: Chile, Lonquimay, [Malleco Province], 1961-1963, in

rotten phloem of Araucaria araucana [Araucariaceae], under the elytra of Araucarius

medius [Coleoptera: Curculionidae: Cossoninae] and Araucarius minor [Coleoptera:

Curculionidae: Cossoninae] in A. araucana, and in galleries of A. minor in A.

araucana; Chile, Nahuelbuta National Park, [Araucanía Region], 1961-1963, under

the elytra of A. minor in A. araucana, and in galleries of A. minor in A. araucana.

136. Dendrolaelaps rykei Hirschmann, 1974

Dendrolaelaps (Tridendrolaelaps) rykei Hirschmann, 1974: 60.

? Dendrolaelaps rykei [sic].— Hirschmann and Wiśniewski, 1982: 148.



TYPE LOCALITY AND HABITAT: South Africa, Potchefstroom, [North West], July and

August 1952, in a decaying straw heap, in a sheep‘s fold and in humus.

178

NOTE: Described on the basis of specimens reported by Ryke (1962a): 92 as Cyrtolaelaps

(Digamasellus) capensis (Berlese, 1921).

137. Dendrolaelaps samsinaki Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Foveodendrolaelaps) samsinaki Hirschmann and Wiśniewski, 1982: 127.

Dendrolaelaps samsinaki.— Karg, 1993c: 350.


TYPE LOCALITY AND HABITAT: Czech Republic, Kdanice, Sobotka, 24 October 1961, in

nest of Lasius fuliginosus [Hymenoptera: Formicidae].


138. Dendrolaelaps saprophilus Huhta, 1982

Dendrolaelaps saprophilus Huhta, 1982: 225.

Dendrolaelaps (Punctodendrolaelaps) saprophilus.— Hirschmann and Wiśniewski, 1982: 52.

Dendrolaelaps saprophilus.— Ma and Bai, 2009: 75.

TYPE DEPOSITORY: Zoological Museum, University of Helsinki, Helsinki, Finland.

TYPE LOCALITY AND HABITAT: Finland, Tikkurila (ca. 20 km north of Helsinki), 1977,

in a mixture of fresh activated sewage sludge and crushed bark.

139. Dendrolaelaps schauenburgi (Schweizer, 1961)

Digamasellus schauenburgi Schweizer, 1961: 139.

Dendrolaelaps (Dendrolaelaps) schauenburgi.— Hirschmann, 1974: 63.

Dendrolaelaps (Punctodendrolaelaps) schauenburgi.— Hirschmann and Wiśniewski, 1982:

50.


TYPE LOCALITY AND HABITAT: Switzerland, Schauenburger Fluh, Jura Mountains, on

rotten roots in a forest.

NOTE: Described from the nymph and adult male.

140. Dendrolaelaps schweizeri Hirschmann, 1960

Dendrolaelaps schweizeri Hirschmann, 1960: 8.

Dendrolaelaps schweizeri.— Hirschmann, 1971c: 14; Hirschmann, 1971e: 20; Karg, 1971:

339; Shcherbak, 1980: 155.

Dendrolaelaps (Dendrolaelaps) schweizeri.— Hirschmann, 1974: 62.

179

Dendrolaelaps (Sellnickidendrolaelaps) schweizeri.— Hirschmann and Wiśniewski, 1982:

66.


TYPE LOCALITY AND HABITAT: Germany, Schöhsee, Schleswig-Holstein, in soil of a

shore meadow.

NOTE: Described from the deutonymph and adult male.

141. Dendrolaelaps sellnicki Hirschmann, 1960

Dendrolaelaps sellnicki Hirschmann, 1960: 8.

Dendrolaelaps sellnicki.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,


Karg, 1971: 337; Shcherbak, 1980: 152.

Dendrolaelaps (Dendrolaelaps) sellnicki.— Hirschmann, 1974: 62.

Dendrolaelaps (Sellnickidendrolaelaps) sellnicki.— Hirschmann and Wiśniewski, 1982: 64.

Punctodendrolaelaps sellnicki.— Karg, 1993c: 353.



[Hymenoptera: Formicidae]; Germany, Schöngeising, Oberbayern, in bee nest

[Hymenoptera].

NOTE: Specimens reported by Shcherbak (1980): 152 as adult female of Dendrolaelaps

sellnicki Hirschmann, 1960 were described as Dendrolaelaps (Sellnickidendrolaelaps)

sellnickiformis Hirschmann and Wiśniewski, 1982: 64.

142. Dendrolaelaps sellnickiformis Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Sellnickidendrolaelaps) sellnickiformis Hirschmann and Wiśniewski, 1982:

64.



TYPE LOCALITY AND HABITAT: Kazakhstan, Karaganda, Karagandy Province, in

hollows of rotten wood, in overrotten compost and in nests of red wood ant and wild

bees [Hymenoptera].

NOTE: Described on the basis of specimens reported by Shcherbak (1980): 152 as adult

female of Dendrolaelaps sellnicki Hirschmann, 1960.

180

143. Dendrolaelaps septentrionalis (Sellnick, 1958)

Digamasellus septentrionalis Sellnick, 1958: 19.

Dendrolaelaps septentrionalis.— Hirschmann, 1960: 4; Karg, 1971: 330; Karg, 1993c: 348.

Cyrtolaelaps (Digamasellus) septentrionalis.— Ryke, 1962a: 110.

Dendrolaelaps (Dendrolaelaps) septentrionalis.— Hirschmann and Wiśniewski, 1982: 76.


TYPE LOCALITY AND HABITAT: Sweden, Järvsö, [Ljusdal], 22 May 1953, on rye leaves

[Poaceae]; Sweden, 13 June – 28 July 1953, on leaves and ears of grasses and cereals.

NOTE1: Karg (1971): 340 considered Dendrolaelaps acornutosimilis Hirschmann, 1960 as

junior synonymy of D. septentrionalis, but Hirschmann and Wiśniewski (1982): 77


NOTE2: Shcherbak (1980): 118 considered Digamasellus septentrionalis Sellnick, 1958 as

junior synonymy of Dendrolaelaps oudemansi Halbert, 1915, but Hirschmann and

Wiśniewski (1982): 77 revoked that synonymy.

144. Dendrolaelaps serratus Hirschmann, 1960

Dendrolaelaps serratus Hirschmann, 1960: 27.

Dendrolaelaps serratus.— Hirschmann, 1971c: 16.

Dendrolaelaps (Dendrolaelaps) serratus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps ?) serratus [sic].— Hirschmann and Wiśniewski, 1982:

106.



[Pinaceae].


145. Dendrolaelaps shcherbakaecornutus Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) shcherbakaecornutus Hirschmann and Wiśniewski, 1982: 80.



TYPE LOCALITY AND HABITAT: Russia, Arkhangelsk Oblast, under bark of coniferous

trees, in galleries of various bark beetles [Coleoptera].


Dendrolaelaps cornutus (Kramer, 1886) sensu Hirschmann, 1960.

181

146. Dendrolaelaps shennongjiaensis Ma and Liu, 2003

Dendrolaelaps shennongjiaensis Ma and Liu, in Ma, Liu and Ye, 2003: 252.

TYPE DEPOSITORY: Institute of Parasitic Diseases, Hubey Academy of Medical Science,

Wuhan, China.

TYPE LOCALITY AND HABITAT: China, Shenongjia (31°15‘N, 109°56‘E, alt. 400 m),

Hubei Province, 16 August 1999, from bat nesting area [Mammalia: Chiroptera].

147. Dendrolaelaps sibiriae Wiśniewski and Michalski, 1983

Dendrolaelaps (Cornodendrolaelaps ?) sibiriae [sic] Wiśniewski and Michalski, 1983: 101.


TYPE LOCALITY AND HABITAT: Russia, Ust-Kut, [Irkutsk Oblast], 5 July 1969, under

the elytra of Monochamus sutor [Coleoptera: Cerambycidae]; Russia, Kirensk,

[Irkutsk Oblast], 20-21 July 1969, under the elytra of M. sutor; Russia, Tunkinskij,

Buryatia, 28 July 1969, under the elytra of Trichius fasciatus [Coleoptera:

Scarabaeidae].


148. Dendrolaelaps simplicis Wiśniewski and Hirschmann, 1991

Dendrolaelaps (Monodendrolaelaps) simplicis Wiśniewski and Hirschmann, 1991a: 223.


Poland.

TYPE LOCALITY AND HABITAT: Poland, near Oder River, Bielinek, 20 August 1988, in

litter of Acer sp. [Sapindaceae], Fagus sp. [Fagaceae] and Quercus sp. [Fagaceae].

149. Dendrolaelaps sinodendronis Wiśniewski and Hirschmann, 1989

Dendrolaelaps sinodendronis Wiśniewski and Hirschmann, 1989a: 331.


Poland.

TYPE LOCALITY AND HABITAT: Bulgaria, near Kamchiya River, 15 July 1983, on

Sinodendron cylindricum [Coleoptera: Lucanidae] deposited in the beetle collection of

Zoologisches Institut der Jagiellonischen Universität, Kraków, Poland.


182

150. Dendrolaelaps sitalaensis (Bhattacharyya, 1978)

Digamasellus sitalaensis Bhattacharyya, 1978: 82.

? Dendrolaelaps (Cornodendrolaelaps) sitalaensis [sic].— Hirschmann, 1983b: 71.


TYPE LOCALITY AND HABITAT: India, Sitala, Rajpur Sonarpur, South 24 Parganas

District, West Bengal, 17 November 1963, in soil under grass.

151. Dendrolaelaps songshanensis Ma and Lin, 2005

Dendrolaelaps songshanensis Ma and Lin, 2005: 350.

TYPE DEPOSITORY: Institute of Plant Protection of Fujian Academy of Agricultural

Science, Fuzhou, China.

TYPE LOCALITY AND HABITAT: China, Songshan Mountain (34°55‘N, 113°05‘E),

Dengfeng County, Henan Province, 15 July 2002, in tree hole.

152. Dendrolaelaps stammeri Hirschmann, 1960

Dendrolaelaps stammeri Hirschmann, 1960: 7.

Dendrolaelaps stammeri.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,


Karg, 1971: 332; Karg, 1993c: 350.

Dendrolaelaps (Dendrolaelaps) stammeri.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) stammeri.— Hirschmann and Wiśniewski, 1982: 130.


TYPE LOCALITY AND HABITAT: Germany, Lippe, on the inner side of a dike.

NOTE: Specimens reported by Shcherbak (1980): 156 as Dendrolaelaps stammeri

Hirschmann, 1960 were described as Dendrolaelaps (Foveodendrolaelaps)

stammeriformis Hirschmann and Wiśniewski, 1982: 130.

153. Dendrolaelaps stammeriformis Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Foveodendrolaelaps) stammeriformis Hirschmann and Wiśniewski, 1982:

130.



TYPE LOCALITY AND HABITAT: Ukraine, in decaying plant debris on the bank of a

canal.

183


Dendrolaelaps stammeri Hirschmann, 1960.

154. Dendrolaelaps stanislavi Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Stanidendrolaelaps) stanislavi Wiśniewski and Hirschmann, 1993b: 291.


Poland.


intercepted under bark of Entandrophragma cylindricum [Meliaceae] imported from

Cameroon.

155. Dendrolaelaps strenzkei (Hirschmann, 1958)

Digamasellus strenzkei Hirschmann, in Sellnick, 1958: 21.

Dendrolaelaps (Punctodendrolaelaps) strenzkeiformis Hirschmann and Wiśniewski, 1982: 56

[objective junior synonym— unjustified replacement name].

Punctodendrolaelaps strenzkeiformis.— Karg, 1993c: 353; Huhta and Karg, 2010: 341.


TYPE LOCALITY AND HABITAT: Sweden, Mora and Orsa, [Dalama], 30 May 1953, on

roots, stems and leaf sheaths of cereal crops.

NOTE: Dendrolaelaps strenzkei Hirschmann, 1960: 7 is a junior homonym of Digamasellus

strenzkei Hirschmann, in Sellnick, 1958: 21, as Hirschmann and Wiśniewski, 1982: 56

considered them distinct species of the same genus.

156. Dendrolaelaps strenzkei Hirschmann, 1960

Dendrolaelaps strenzkei Hirschmann, 1960: 7.

Dendrolaelaps strenzkei.— Athias-Henriot, 1961a: 468; Hirschmann, 1971c: 14; Hirschmann,


Shcherbak, 1980: 105.

Dendrolaelaps (Dendrolaelaps) strenzkei.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) strenzkei.— Hirschmann and Wiśniewski, 1982: 56.

Punctodendrolaelaps strenzkei.— Karg, 1993c: 354; Huhta and Karg, 2010: 341.


184


[Hymenoptera: Formicidae] and on grass; Germany, Süseler Moor, Ostholstein, in

cow manure; Germany, Berlin, from a potato field.

NOTE: Dendrolaelaps strenzkei Hirschmann, 1960: 7 is a junior homonym of Digamasellus

strenzkei Hirschmann, in Sellnick, 1958: 21, as Hirschmann and Wiśniewski, 1982: 56

considered them distinct species of the same genus. Conversely to the interpretation of

Hirschmann and Wiśniewski (1982): 56, the species in need of a new name is

Dendrolaelaps strenzkei Hirschmann, 1960.

157. Dendrolaelaps tauricus Shcherbak, 1983

Dendrolaelaps tauricus Shcherbak, 1983: 81.



TYPE LOCALITY AND HABITAT: Ukraine, Karadag Nature Reserve, Crimea, 7 August

1980, in soil with rotten wood under an ash tree Fraxinus sp. [Oleaceae].

158. Dendrolaelaps tenuipiloides Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) tenuipiloides Hirschmann and Wiśniewski, 1982: 83.



TYPE LOCALITY AND HABITAT: Russia, Arkhangelsk Oblast and Kamchatka, on stumps

of Alnus sp. [Betulaceae] and Pinus sp. [Pinaceae], and under the bark of Betula sp.

[Betulaceae].


Dendrolaelaps tenuipilus Hirschmann, 1960.

159. Dendrolaelaps tenuipilus Hirschmann, 1960

Dendrolaelaps tenuipilus Hirschmann, 1960: 7.

Dendrolaelaps tenuipilus.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 13; Hirschmann,


Karg, 1971: 330; Karg, 1993c: 349.

Dendrolaelaps (Dendrolaelaps) tenuipilus.— Hirschmann, 1974: 62; Hirschmann and



185

TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stumps of Alnus sp.

[Betulaceae] and Picea sp. [Pinaceae]; Germany, Blauen, Schwarzwald, on stump of

Picea sp..

NOTE: Specimens reported by Shcherbak (1980): 116 as Dendrolaelaps tenuipilus

Hirschmann, 1960 were described as Dendrolaelaps (Dendrolaelaps) tenuipiloides


160. Dendrolaelaps transkeiensis Hirschmann and Wiśniewski, 1984

Dendrolaelaps (Cornodendrolaelaps) transkeiensis Hirschmann and Wiśniewski, 1984: 92.


TYPE LOCALITY AND HABITAT: South Africa, Lusikisiki, Transkei, 1979, from coastal

evergreen forest.


161. Dendrolaelaps transportabilis Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Punctodendrolaelaps) transportabilis Wiśniewski and Hirschmann, 1993c:

317.


TYPE LOCALITY AND HABITAT: Poland, Port of Szczecin, 19 April 1985, intercepted

under bark of logs imported from Cameroon.


162. Dendrolaelaps transvaalensis (Ryke, 1962)

Cyrtolaelaps (Digamasellus) transvaalensis Ryke, 1962a: 100.

Dendrolaelaps (Tridendrolaelaps) transvaalensis.— Hirschmann, 1974: 60.

Dendrolaelaps (Foveodendrolaelaps) transvaalensis.— Hirschmann and Wiśniewski, 1982:

126.



TYPE LOCALITY AND HABITAT: South Africa, Boskop, [North West], June 1958, from

damp soil.

163. Dendrolaelaps trapezoides Hirschmann, 1960

Dendrolaelaps trapezoides Hirschmann, 1960: 8.

186

Dendrolaelaps trapezoides.— Hirschmann, 1971d: 19; Karg, 1971: 338.

Dendrolaelaps (Dendrolaelaps) trapezoides.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) trapezoides.— Hirschmann and Wiśniewski, 1982: 52.

Punctodendrolaelaps trapezoides.— Karg, 1993c: 353.


TYPE LOCALITY AND HABITAT: Germany, Bielefeld, on stump of Fagus sp. [Fagaceae];

Germany, Oberstdorf, on stump of Alnus sp. [Betulaceae]; Switzerland, Ofenpass,

[Graubünden], on stump of Pinus mugo [Pinaceae].

NOTE: Specimens reported by Shcherbak (1980): 107 as Dendrolaelaps trapezoides

Hirschmann, 1960 were described as Dendrolaelaps (Punctodendrolaelaps) piriformis


164. Dendrolaelaps tritrichus Hirschmann, 1960

Dendrolaelaps tritrichus Hirschmann, 1960: 22.

Dendrolaelaps (Dendrolaelaps) tritrichus.— Hirschmann, 1974: 61.

Dendrolaelaps (Punctodendrolaelaps) tritrichus.— Hirschmann and Wiśniewski, 1982: 57.


TYPE LOCALITY AND HABITAT: Sweden, from unspecified substrate.


165. Dendrolaelaps tuberosus Hirschmann, 1960

Dendrolaelaps tuberosus Hirschmann, 1960: 7.

Dendrolaelaps tuberosus.— Hirschmann, 1971e: 20; Shcherbak, 1980: 113.

Dendrolaelaps (Dendrolaelaps) tuberosus.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) tuberosus.— Hirschmann and Wiśniewski, 1982:

138.


TYPE LOCALITY AND HABITAT: Germany, Bremen, under bark.


166. Dendrolaelaps tumulus Luxton, 1982

Dendrolaelaps tumulus Luxton, 1982: 328.


187

TYPE LOCALITY AND HABITAT: New Zealand, Waikato, North Island, July 1967, from

peat pasture soils.

167. Dendrolaelaps uncinatus Hirschmann, 1960

Dendrolaelaps uncinatus Hirschmann, 1960: 7.

Dendrolaelaps uncinatus.— Hirschmann, 1971d: 18; Hirschmann, 1971e: 20; Karg, 1971:

337; Shcherbak, 1980: 129.

Dendrolaelaps (Dendrolaelaps) uncinatus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) uncinatus.— Hirschmann and Wiśniewski, 1982: 102.

Cornodendrolaelaps uncinatus.— Karg, 1993c: 360.


TYPE LOCALITY AND HABITAT: Germany, Nuremberg, on stump of Pinus sp.

[Pinaceae].

168. Dendrolaelaps undulatus Hirschmann, 1960

Dendrolaelaps undulatus Hirschmann, 1960: 8.

Dendrolaelaps undulatus.— Hirschmann, 1971b: 14; Hirschmann, 1971d: 18; Karg, 1971:

336; Shcherbak, 1980: 125.

Dendrolaelaps (Dendrolaelaps) undulatus.— Hirschmann, 1974: 62.

Dendrolaelaps (Cornodendrolaelaps) undulatus.— Hirschmann and Wiśniewski, 1982: 101.

Cornodendrolaelaps undulatus.— Karg, 1993c: 355.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, on stump of Populus sp.

[Salicaceae] and in nest of Lasius fuliginosus [Hymenoptera: Formicidae].

169. Dendrolaelaps validulus (Berlese, 1920)

Gamasellus (Digamasellus) validulus Berlese, 1920b: 163.

? Gamasellus (di) validulus [sic].— Lombardini, 1941: 183.

Dendrolaelaps validulus.— Hirschmann, 1960: 5.

Cyrtolaelaps (Digamasellus) validulus.— Ryke, 1962a: 90.

Dendrolaelaps (Dendrolaelaps) validulus.— Hirschmann, 1974 : 62.

Dendrolaelaps (Punctodendrolaelaps) validulus.— Hirschmann and Wiśniewski, 1982 : 59.


TYPE LOCALITY AND HABITAT: Somalia, from a forest.

188

170. Dendrolaelaps varipunctatus (Hurlbutt, 1967)

Digamasellus varipunctatus Hurlbutt, 1967: 521.

Dendrolaelaps varipunctatus.— McGraw and Farrier, 1969: 131.

Dendrolaelaps (Dendrolaelaps) varipunctus [sic].— Hirschmann, 1974: 63.

Dendrolaelaps (Cornodendrolaelaps) varipunctatus.— Hirschmann and Wiśniewski, 1982:

98.



TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, in galleries of

Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] in Pinus taeda

[Pinaceae].

171. Dendrolaelaps vermicularis Ma, 2001

Dendrolaelaps vermicularis Ma, 2001a: 231.


TYPE LOCALITY AND HABITAT: China, Dunhua Conty (43°21‘N, 128°13‘E), Jilin

Province, August 1990, in forest soil.

172. Dendrolaelaps viator (Vitzthum, 1921)

Gamasellus viator Vitzthum, 1921: 7.

Gamasellus (Digamasellus) viator.— Vitzthum, 1923: 117.

Cyrtolaelaps (Digamasellus) viator.— Ryke, 1962a: 106.

Dendrolaelaps (Dendrolaelaps) viator.— Hirschmann and Wiśniewski, 1982: 82.


TYPE LOCALITY AND HABITAT: Germany, lower Elbe River, downstream of Hamburg,

July 1914, on Bombus sp. [Hymenoptera: Apidae].


173. Dendrolaelaps vitzthumicornutus Hirschmann and Wiśniewski, 1982

Dendrolaelaps (Dendrolaelaps) vitzthumicornutus Hirschmann and Wiśniewski, 1982: 80.

Dendrolaelaps vitzthumicornutus.— Karg, 1993c: 349.


189

TYPE LOCALITY AND HABITAT: Austria, Waidhofen an der Thaya, May 1921, in gallery

of Scolytus laevis [Coleoptera: Curculionidae: Scolytinae] in Ulmus montanus

[Ulmaceae].

NOTE: Described on the basis of specimens reported by Vitzthum (1926b): 411 as

Dendrolaelaps cornutus (Kramer, 1886).

174. Dendrolaelaps wangfengzheni Ma, 1995

Dendrolaelaps wangfengzheni Ma, 1995: 52.


TYPE LOCALITY AND HABITAT: China, Baicheng, Jilin Province, July to September

1993, under the decomposed bark of poplar Populus sp. [Salicaceae].

175. Dendrolaelaps wengrisae Wiśniewski, 1979

Dendrolaelaps wengrisae Wiśniewski, 1979b: 53.

Dendrolaelaps (Punctodendrolaelaps) wengrisae.— Hirschmann and Wiśniewski, 1982: 50.


Poland.

TYPE LOCALITY AND HABITAT: Poland, near Poznań, 1961, in nest of Formica

polyetena [Hymenoptera: Formicidae].

NOTE: Hirschmann and Wiśniewski (1982): 50 suspected Dendrolaelaps punctatulus

Hirschmann, 1960 sensu Shcherbak (1980): 102 to correspond to Dendrolaelaps

(Punctodendrolaelaps) wengrisae Wiśniewski, 1979.

176. Dendrolaelaps willmanni Hirschmann, 1960

Dendrolaelaps willmanni Hirschmann, 1960: 8.

Dendrolaelaps willmanni.— Hirschmann, 1971c: 15; Hirschmann, 1971d: 19; Hirschmann,

1971e: 20; Karg, 1971: 332; Shcherbak, 1980: 149; Karg, 1993c: 350.

Dendrolaelaps (Dendrolaelaps) willmanni.— Hirschmann, 1974: 62.

Dendrolaelaps (Foveodendrolaelaps) willmanni.— Hirschmann and Wiśniewski, 1982: 132.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, in cow manure.

177. Dendrolaelaps xylophilus Wiśniewski and Hirschmann, 1993

Dendrolaelaps (Xylodendrolaelaps) xylophilus Wiśniewski and Hirschmann, 1993c: 322.

190


Poland.


intercepted on wood of Entandrophragma cylindricum [Meliaceae] imported from

Cameroon.

178. Dendrolaelaps zaheri Metwally and Mersal, 1985

Dendrolaelaps zaheri Metwally and Mersal, 1985.



NOTE: Description not seen.

179. Dendrolaelaps zwoelferi Hirschmann, 1960

Dendrolaelaps zwoelferi Hirschmann, 1960: 7.

Dendrolaelaps zwoelferi.— Hirschmann, 1971d: 17; Karg, 1971: 330; Shcherbak, 1980: 143;

Karg, 1993c: 349.

Dendrolaelaps (Dendrolaelaps) zwoelferi.— Hirschmann, 1974: 62.

Dendrolaelaps (Apophyseodendrolaelaps) zwoelferi.— Hirschmann and Wiśniewski, 1982:

113.


TYPE LOCALITY AND HABITAT: Germany, Schöngeising, in bee [Hymenoptera] nest;

Germany, Berlin, on grass seeds.

Genus Dendrolaelaspis Lindquist, 1975

Dendrolaelaps (Dendrolaelaspis) Lindquist, 1975: 16 (described in Digamasellidae Evans).

Dendrolaelaspis.— Shcherbak, 1980: 175.

Dendrolaelaps (Dendrolaelaspis).— Hirschmann and Wiśniewski, 1982: 137.

Type species: Digamasellus angulosus Willmann, 1936, by original designation.

Dendrolobatus Shcherbak, 1983: 74 (described in Rhodacaridae Oudemans) [junior synoym].

Type species: Dendrolaelaps (Dendrolaelaspis) longisetosus Shcherbak, 1977, by


191

180. Dendrolaelaspis angulosus (Willmann, 1936)

Digamasellus (?) angulosus [sic] Willmann, 1936: 280.

Digamasellus angulosus.— Leitner, 1949: 59; Willmann, 1951: 142; Franz, 1954: 341;

Hurlbutt, 1967: 498.

Dendrolaelaps angulosus.— Hirschmann, 1960: 8; Athias-Henriot, 1961a: 467; Hirschmann,

1971a: 11; Hirschmann, 1971b: 12; Hirschmann, 1971c: 15; Hirschmann, 1971d: 19;

Karg, 1971: 338.

Cyrtolaelaps (Digamasellus) angulosus.— Ryke, 1962a: 109.

Dendrolaelaps (Dendrolaelaps) angulosus.— Hirschmann, 1974: 62.

Dendrolaelaps (Dendrolaelaspis) angulosus.— Lindquist, 1975: 16; Shcherbak, 1978a: 1437;

Karg, 1979: 205; Hirschmann and Wiśniewski, 1982: 143.

Dendrolaelaspis angulosus.— Shcherbak, 1980: 178; Karg, 1993c: 345; Karg, 1998: 189;

Karg and Schorlemmer, 2009: 69.


TYPE LOCALITY AND HABITAT: Germany, in Hundsfeld and Waldenburger, 1933 and

1934, in meadows.

181. Dendrolaelaspis baculus Karg, 2003

Dendrolaelaspis baculus Karg, 2003: 244.



TYPE LOCALITY AND HABITAT: Ecuador, Pichincha Province, 1990, in soil.

182. Dendrolaelaspis baloghi (Hirschmann, 1974)

Dendrolaelaps (Dendrolaelaps) baloghi Hirschmann, 1974: 54.

Dendrolaelaps (Dendrolaelaspis) baloghi.— Lindquist, 1975: 43; Shcherbak, 1978a: 1437;


Dendrolaelaspis baloghi.— Shcherbak, 1980: 178; Karg, 1998: 189; Karg and Schorlemmer,

2009: 69.


TYPE LOCALITY AND HABITAT: Cuba, Sierra del Rosario, Pinar del Río Province, 24

January 1970, in litter from evergreen forest.

192

183. Dendrolaelaspis bistilus (Karg, 1979)

Dendrolaelaps (Dendrolaelaspis) bistilus Karg, 1979: 201.

Dendrolaelaps (Dendrolaelaspis) bistilus.— Hirschmann and Wiśniewski, 1982: 142.

Dendrolaelaspis bistilus.— Karg, 1998: 188; Karg and Schorlemmer, 2009: 68.

TYPE DEPOSITORY: Ungarisches Naturwissenschaftliches Museum, Budapest, Hungary.

TYPE LOCALITY AND HABITAT: Argentina, Arroyo Negro, Rio Negro Province, 6

March 1961, in litter.

184. Dendrolaelaspis bregetovae (Shcherbak, 1978)

Dendrolaelaps (Dendrolaelaspis) bregetovae Shcherbak, 1978a: 1434.

Dendrolaelaspis bregetovae.— Shcherbak, 1980: 179; Karg, 1993c: 344; Karg, 1998: 189;


Dendrolaelaps (Dendrolaelaspis) bregetovae.— Hirschmann and Wiśniewski, 1982: 144.

TYPE DEPOSITORY: Zoological Institute of the Russian Academy of Sciences, Saint-

Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Luzsky District, Kirov Oblast, 12 May 1960, on

a tussock of moss on a meadow on the shore of a river.

185. Dendrolaelaspis brevisetosus (Shcherbak, 1978)

Dendrolaelaps (Dendrolaelaspis) brevisetosus Shcherbak, 1978a: 1436.

Dendrolaelaspis brevisetosus.— Shcherbak, 1980: 178; Karg, 1998: 189; Karg and


Dendrolaelaps (Dendrolaelaspis) brevisetosus.— Hirschmann and Wiśniewski, 1982: 144.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Panama, Lerida Farm, Boquete District, Chiriquí

Province, 14 March 1959, in litter.

186. Dendrolaelaspis cienfuegi (Wiśniewski and Hirschmann, 1989)

Dendrolaelaps (Dendrolaelaspis) cienfuegi Wiśniewski and Hirschmann, 1989b: 310.


Poland.


under tree bark.

193

187. Dendrolaelaspis crassilaciniae (Wiśniewski and Hirschmann, 1983)

Dendrolaelaps (Dendrolaelaspis) crassilaciniae Wiśniewski and Hirschmann, 1983a: 103.


TYPE LOCALITY AND HABITAT: Poland, Biedrusko Botanic Garden, Oborniki, 11 April

1983, under bark of Robinia sp. [Fagaceae]; Poland, Czolowo Botanic Garden, Babki,

8 May 1983 and 7 July 1983, under bark of Pinus sp. [Pinaceae] and Populus sp.

[Salicaceae].

188. Dendrolaelaspis eucrinis (Karg, 1979)

Dendrolaelaps (Dendrolaelaspis) eucrinis Karg, 1979: 203.

Dendrolaelaps (Dendrolaelaspis) eucrinis.— Hirschmann and Wiśniewski, 1982: 142.

Dendrolaelaspis eucrinis.— Karg and Schorlemmer, 2009: 68.


TYPE LOCALITY AND HABITAT: Argentina, Mount Piltriquitron, El Bolson, Rio Negro

Province, 19 September 1961, in litter of Libocedrus [Cupressaceae] and Lomatia

[Proteaceae] forest.

189. Dendrolaelaspis geminus Karg, 1998

Dendrolaelaspis geminus Karg, 1998: 188.

Dendrolaelaspis geminus.— Karg and Schorlemmer, 2009: 69.



TYPE LOCALITY AND HABITAT: Ecuador, between Pifo and Papallacta, Pichincha

Province, 14 April 1989, in moss and withered plant-debris under bushes.


190. Dendrolaelaspis hungaricus (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Dendrolaelaspis) hungaricus Hirschmann and Wiśniewski, 1982: 141.

Dendrolaelaspis hungaricus.— Karg, 1993c: 344.


TYPE LOCALITY AND HABITAT: Hungary, from unspecified substrate.

191. Dendrolaelaspis lindquisti (Shcherbak, 1978)

194

Dendrolaelaps (Dendrolaelaspis) lindquisti Shcherbak, 1978a: 1435.

Dendrolaelaspis lindquisti.— Shcherbak, 1980: 178; Karg, 1998: 189; Karg and


Dendrolaelaspis (Dendrolaelaspis) lindquisti.— Hirschmann and Wiśniewski, 1982: 144.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: México, 3 miles north of San Cristóbal de las Casas,

Chiapas, on mushrooms on the rotting bark of a log.

192. Dendrolaelaspis lobatus (Shcherbak and Chelebiev, 1977)

Dendrolaelaps (Dendrolaelaspis) lobatus Shcherbak and Chelebiev, 1977: 471.

Dendrolaelaspis lobatus.— Shcherbak, 1980: 180; Karg, 1998: 189; Karg and Schorlemmer,

2009: 69.

Dendrolaelaps (Dendrolaelaspis) lobatus.— Hirschmann and Wiśniewski, 1982: 144.



TYPE LOCALITY AND HABITAT: Kazakhstan, Botanical Garden, Karaganda, Karagandy

Province, 14 May 1976, in compost.

193. Dendrolaelaspis longisetosus (Shcherbak, 1977)

Dendrolaelaps (Dendrolaelaspis) longisetosus Shcherbak, in Shcherbak and Gomelauri,

1977: 210.

Dendrolaelaspis longisetosus.— Shcherbak, 1980: 181; Karg and Schorlemmer, 2009: 68.

Dendrolaelaps (Dendrolaelaspis) longisetosus.— Hirschmann and Wiśniewski, 1982: 142.

Dendrolobatus longisetosus.— Shcherbak, 1983: 74.



TYPE LOCALITY AND HABITAT: Georgia, Shiraki, 24 May 1960, in nest of Microtus

socialis [Mammalia: Rodentia: Cricetidae].

194. Dendrolaelaspis miniangulosus (Shcherbak, 1978)

Dendrolaelaps (Dendrolaelaspis) miniangulosus Shcherbak, 1978a: 1437.

Dendrolaelaspis miniangulosus.— Shcherbak, 1980: 178; Karg, 1998: 189; Karg and


195

Dendrolaelaps (Dendrolaelaspis) miniangulosus.— Hirschmann and Wiśniewski, 1982: 144.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: México, 30 miles east of San Cristóbal de las Casas,

Chiapas, 4 May 1969, on Polyporaceae fungus on an oak deck.

195. Dendrolaelaspis orientalis (Bhattacharyya, 1969)

Digamasellus orientalis Bhattacharyya, 1969: 69.

Dendrolaelaps (Dendrolaelaspis) orientalis.— Lindquist, 1975: 17; Shcherbak, 1978a: 1437;


Dendrolaelaspis orientalis.— Shcherbak, 1980: 177; Karg and Schorlemmer, 2009: 68.


TYPE LOCALITY AND HABITAT: India, Kameng Elephant Reserve, Arunachal Pradesh

(cited as North-East Frontier Agency), 23 December 1965, under bark of a dead tree.

NOTE: Specimens reported as allotype adult male of Digamasellus orientalis Bhattacharyya,

1969 were described as Dendrolaelaps (Dendrolaelaps) bhattacharyyai Hirschmann,

1974: 52.

196. Dendrolaelaspis piscis (Karg, 1979)

Dendrolaelaps (Dendrolaelaspis) piscis Karg, 1979: 203.

Dendrolaelaps (Dendrolaelaspis) piscis.— Hirschmann and Wiśniewski, 1982: 142.

Dendrolaelaspis piscis.— Karg and Schorlemmer, 2009: 68.


TYPE LOCALITY AND HABITAT: Argentina, Mount Piltriquitron, El Bolson, Rio Negro

Province, 10 June 1961, in litter.

197. Dendrolaelaspis poltavae Shcherbak and Sklar, 1983

Dendrolaelaspis poltavae Shcherbak and Sklar, 1983: 77.



TYPE LOCALITY AND HABITAT: Ukraine, Poltava, 12 April 1983, under bark of a

Populus sp. [Salicaceae].

198. Dendrolaelaspis sexsetosus Karg and Schorlemmer, 2009

196

Dendrolaelaspis sexsetosus Karg and Schorlemmer, 2009: 68.



TYPE LOCALITY AND HABITAT: Ecuador, between Puerto Napo and Ahuano, [Napo

Province], 1990, in litter of cacao [Sterculiaceae] plantation.

Genus Dendroseius Karg, 1965

Dendrolaelaps (Dendroseius) Karg, 1965: 297 (described in Rhodacaridae Oudemans).

Dendrolaelaps (Dendroseius).— Karg, 1971: 317.

Dendroseius.— Lindquist, 1975: 12; Shcherbak, 1980: 172.

Type species: Dendrolaelaps scotarius Sheals, 1958, by original designation

(objective junior synonym of Digamasellus reticulatus Sheals, 1956).

199. Dendroseius amoliensis Faraji, Sakenin-Chelav and Karg, 2006

Dendroseius amoliensis Faraji, Sakenin-Chelav and Karg, 2006: 64.

TYPE DEPOSITORY: National Museum of Natural History, Leiden, Netherlands.

TYPE LOCALITY AND HABITAT: Iran, Amol, [Māzandarān Province], 25 August 2004, in

soil and debris.

200. Dendroseius badenhorsti (Ryke, 1962)

Cyrtolaelaps (Digamasellus) badenhorsti Ryke, 1962a: 96.

Dendrolaelaps (Tridendrolaelaps) badenhorsti.— Hirschmann, 1974: 60.

Dendroseius badenhorsti.— Lindquist, 1975: 42; Faraji, Sakenin-Chelav and Karg, 2006: 67.



TYPE LOCALITY AND HABITAT: South Africa, Viljoenskroon, March 1958, in manure.

201. Dendroseius congoensis Wiśniewski and Hirschmann, 1992

Dendroseius congoensis Wiśniewski and Hirschmann, 1992: 63.

Dendroseius congoensis.— Faraji, Sakenin-Chelav and Karg, 2006: 67.


Poland.

197

TYPE LOCALITY AND HABITAT: Poland, Port of Szczecin, 14 March 1989, intercepted

under bark of Entandrophragma cylindricum [Meliaceae] imported from the

Democratic Republic of the Congo.


202. Dendroseius gujarati Wiśniewski and Hirschmann, 1989

Dendroseius gujarati Wiśniewski and Hirschmann, 1989a: 319.

Dendroseius gujarati.— Faraji, Sakenin-Chelav and Karg, 2006: 67.


Poland.

TYPE LOCALITY AND HABITAT: India, Dhrangadhra, Gujarat, 8 August 1986, on

unidentified Scarabaeidae [Coleoptera].


203. Dendroseius reticulatus (Sheals, 1956)

Digamasellus reticulatus Sheals, 1956: 101.

Dendrolaelaps (Dendroseius) reticulatus.— Karg, 1971: 327.

Dendroseius reticulatus.— Shcherbak, 1980: 173; Karg, 1993c: 342; Faraji, Sakenin-Chelav

and Karg, 2006: 66.

Digamasellus scotarius Sheals, 1958: 304 [objective junior synonym by Shcherbak, 1980:

172 — unjustified replacement name].

Dendrolaelaps scotarius.— Athias-Henriot, 1961a: 468.

Cyrtolaelaps (Digamasellus) scotarius.— Ryke, 1962a: 111.

Dendrolaelaps (Dendroseius) scotarius.— Karg, 1965: 297.

Dendroseius scotarius.— Lindquist, 1975: 12.

TYPE DEPOSITORY: British Museum (Natural History), London, England.

TYPE LOCALITY AND HABITAT: Scotland, Bellahouston Park, Glasgow, June 1951-

October 1952, in soil of old grassland.

Genus Digamasellus Berlese, 1905

Gamasellus (Digamasellus) Berlese, 1905b: 234 [presumed to be described in Parasitidae

Oudemans (also referred to as Gamasidae Leach)].

198

Digamasellus.— Leitner, 1949: 51; Lindquist, 1975: 13; Bregetova and Shcherbak, 1977a:

286.

Type species: Gamasellus (Digamasellus) perpusillus Berlese, 1905, by original

designation [junior synonym of Digamasellus punctum Berlese, 1904].

Dendrolaelaps (Tridendrolaelaps) Hirschmann, 1974: 60 [objective junior synoym by

Lindquist, 1975: 40].

Type species: Cyrtolaelaps (Gamasellus) punctum Berlese, 1904, by original

designation.

204. Digamasellus australis Lindquist, 1975

Digamasellus australis Lindquist, 1975: 21.

TYPE DEPOSITORY: South Australian Museum, Adelaide, Australia.

TYPE LOCALITY AND HABITAT: Australia, One Tree Island, Capricorn Islands Group,

off east coast of Queensland, 3 August 1969, from relatively dry humus of Sesuvium

portulacastrum [Aizoaceae], to which guano was added by seabirds, near margin of a

brackish pond.

205. Digamasellus punctum (Berlese, 1904)

Cyrtolaelaps (Gamasellus) punctum Berlese, 1904: 262.

Dendrolaelaps punctum.— Hirschmann, 1960: 6; Hirschmann, 1971c: 15; Hirschmann,

1971d: 19; Hirschmann, 1971e: 20; Karg, 1971: 339.

Dendrolaelaps (Tridendrolaelaps) punctum.— Hirschmann, 1974: 60.

Digamasellus punctum.— Lindquist, 1975: 13; Karg, 1993c: 346; Bernini, Castagnoli and

Nannelli, 1995: 20.

Gamasellus (Digamasellus) perpusillus Berlese, 1905b: 234 [junior synonymy by Lindquist,

1975: 13].

Gamasellus (Digamasellus) perpusillus.— Berlese, 1910a: 199; Schweizer, 1922: 34.

Digamasellus perpusillus.— Leitner, 1949: 57; Franz, 1954: 342.

Cyrtolaelaps (Digamasellus) perpusillus.— Ryke, 1962a: 89.

TYPE DEPOSITORY: D. punctum and G. perpusillus: Istituto Sperimentale per la Zoologia

Agraria, Florence, Italy.

TYPE LOCALITY AND HABITAT: D. punctum: Italy, Florence, in piled manure; G.

perpusillus: Italy, Tiarno, Province of Trento, on rotten wood.

199

206. Digamasellus variabilis Wiśniewski and Hirschmann, 1989

Digamasellus variabilis Wiśniewski and Hirschmann, 1989b: 305.


Poland.


under tree bark.

Genus Insectolaelaps Shcherbak, 1980

Insectolaelaps Shcherbak, 1980: 192 (described in Rhodacaridae Oudemans).

Dendrolaelaps (Insectolaelaps).— Hirschmann and Wiśniewski, 1982: 31.

Insectolaelaps.— Karg, 1993c: 368.

Type species: Dendrolaelaps armatus Hirschmann, 1960, by original designation.

Dendrolaelaps (Ipidodendrolaelaps) Hirschmann and Wiśniewski, 1982: 38 [junior

synonymy by Karg, 1993c: 368].

Type species: Gamasellus (Digamasellus) quadrisetus Berlese, 1920, by original

designation.

207. Insectolaelaps armatus (Hirschmann, 1954)

Digamasellus armatus Hirschmann, 1954b: 247.

Dendrolaelaps armatus.— Hirschmann, 1960: 7; Hirschmann, 1971a: 11; Hirschmann,

1971b: 12; Hirschmann, 1971c: 15; Hirschmann, 1971d: 18; Hirschmann, 1971e: 21;


Dendrolaelaps (Multidendrolaelaps) armatus.— Hirschmann, 1974: 61.

Insectolaelaps armatus.— Shcherbak, 1980: 197; Karg, 1993c: 368.

Dendrolaelaps (Insectolaelaps) armatus.— Hirschmann and Wiśniewski, 1982: 34.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stump of Picea sp.

[Pinaceae]; Germany, Sieber, on stump of Picea sp. and in gallery of Ips typographus

[Coleoptera: Curculionidae: Scolytinae]; Germany, Bückeberg, on stump of Picea sp.;

Austria, Admont, [Styria], on stump of Picea sp..

200

NOTE1: Specimens reported by Hurlbutt (1967): 507 as Digamasellus armatus (Hirschmann)

were described as Dendrolaelaps (Insectolaelaps) neoarmatus Hirschmann and


NOTE2: Specimens reported by McGraw and Farrier (1969): 100 as Dendrolaelaps armatus

Hirschmann were described as Dendrolaelaps (Insectolaelaps) latoarmatus


NOTE3: Specimens reported by Ishikawa (1980): 31 as Dendrolaelaps armatus Hirschmann

were described as Dendrolaelaps (Insectolaelaps) japanoarmatus Hirschmann and


NOTE4: Franz (1954): 341 cited Digamasellus armatus Hirschmann n. sp., but did not

characterize it morphologically; thus, that constitutes a nomen nudum.

208. Insectolaelaps bialowiezae (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Insectolaelaps) bialowiezae Hirschmann and Wiśniewski, 1982: 35.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Bialowieza Nationalpark, [Podlaskie

Voivodeship], 1 June 1980, on Cerambycidae [Coleoptera].


209. Insectolaelaps euarmatus (Hirschmann, 1960)

Dendrolaelaps euarmatus Hirschmann, 1960: 7.

Dendrolaelaps euarmatus.— Hirschmann, 1971c: 16; Hirschmann, 1971d: 18; Hirschmann,

1971e: 20; Karg, 1971: 335.

Dendrolaelaps (Multidendrolaelaps) euarmatus.— Hirschmann, 1974: 61.

Insectolaelaps euarmatus.— Shcherbak, 1980: 199; Karg, 1993c: 368.

Dendrolaelaps (Insectolaelaps) euarmatus.— Hirschmann and Wiśniewski, 1982: 36.


TYPE LOCALITY AND HABITAT: Germany, Bielefeld, on stump of Picea sp. [Pinaceae];

Germany, Bückeberg, in gallery of Dryocoetes autographus [Coleoptera:


210. Insectolaelaps eustructurus (Hirschmann, 1960)

Dendrolaelaps eustructurus Hirschmann, 1960: 21.

201

Dendrolaelaps eustructurus.— Hirschmann, 1971c: 16

Dendrolaelaps (Multidendrolaelaps) eustructurus.— Hirschmann, 1974: 61.

Dendrolaelaps (Insectolaelaps) eustructurus.— Hirschmann and Wiśniewski, 1982: 37.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, under the elytra of Hylurgus

ligniperda [Coleoptera: Curculionidae: Scolytinae].


211. Insectolaelaps hirsutus (Hirschmann, 1960)

Dendrolaelaps hirsutus Hirschmann, 1960: 21.

Dendrolaelaps hirsutus.— Hirschmann, 1971c: 16.

Dendrolaelaps (Multidendrolaelaps) hirsutus.— Hirschmann, 1974: 61.

Insectolaelaps hirsutus.— Shcherbak, 1980: 192.

Dendrolaelaps (Insectolaelaps) hirsutus.— Hirschmann and Wiśniewski, 1982: 37.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, under the elytra of Hylurgus

ligniperda [Coleoptera: Curculionidae: Scolytinae].


212. Insectolaelaps ipidoquadrisetus (Wiśniewski and Hirschmann, 1983)

Dendrolaelaps (Ipidodendrolaelaps) ipidoquadrisetus Wiśniewski and Hirschmann, 1983b:

124.



TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, August 1965, on inner bark

of Pinus taeda [Plantae: Pinaceae].

NOTE: Described on the basis of specimens reported by Hurlbutt (1967): 502 as

Digamasellus quadrisetosimilis (Hirschmann and Rühm, 1955).

213. Insectolaelaps japanoarmatus (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Insectolaelaps) japanoarmatus Hirschmann and Wiśniewski, 1982: 35.


TYPE LOCALITY AND HABITAT: Japan, in phoretic association with beetles [Coleoptera].

202

NOTE: Described on the basis of specimens reported by Ishikawa (1980): 31 as

Dendrolaelaps armatus Hirschmann, 1960.

214. Insectolaelaps javae (Wiśniewski and Hirschmann, 1989)

Dendrolaelaps (Insectolaelaps ?) javae [sic] Wiśniewski and Hirschmann, 1989a: 322.

‗TYPE DEPOSITORY: Department of Forest Protection, University of Life Sciences,

Poznań, Poland.

TYPE LOCALITY AND HABITAT: Indonesia, Java, on Thyrsia sp. [Coleoptera:

Cerambycidae] deposited in the beetle collection of Bereich Biologie der Sektion

Forstwirtschaft, Tharandt, Germany.


215. Insectolaelaps kielczewskii (Skorupski and Gwiazdowicz, 1992)

Dendrolaelaps (Insectolaelaps) kielczewskii Skorupski and Gwiazdowicz, 1992: 225.


Poland.

TYPE LOCALITY AND HABITAT: Poland, Pieniny National Park, 4 September 1991, on

fir [Abies sp. (Pinaceae)] bark; Poland, Białowieża Forest, 17 June 1990, in galleries

of Coleoptera under bark of spruce [Picea sp. (Pinaceae)]; Poland, Forestry Kalina,

Kaczory Forest District, 20 July 1991, in galleries of Coleoptera under bark of spruce

[Picea sp.]; Poland, Oborniki Wlkp. Forest District, 5 May 1992, on stump of Pinus

sp. [Pinaceae]; Poland, Wielkopolska National Park, Forestry Osowa Góra, 5 May

1992, under bark of Pinus sp..

216. Insectolaelaps latoarmatus (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Insectolaelaps) latoarmatus Hirschmann and Wiśniewski, 1982: 35.


TYPE LOCALITY AND HABITAT: USA, Beaufort County, North Carolina, on

Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] attacking Pinus

echinata [Pinaceae]; USA, Yadkin County, North Carolina, on Ips avulses

[Coleoptera: Curculionidae: Scolytinae] attacking Pinus taeda [Pinaceae]; USA,

Amelia County, Virginia, on Ips grandicollis [Coleoptera: Curculionidae: Scolytinae]

attacking Pinus virginiana [Pinaceae].

203

NOTE: Described on the basis of specimens reported by McGraw and Farrier (1969): 100 as

Dendrolaelaps armatus Hirschmann, 1960.

217. Insectolaelaps latopini (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Insectolaelaps) latopini Hirschmann and Wiśniewski, 1982: 36.

Dendrolaelaps (Insectolaelaps) latopini.— Wiśniewski, 1984: 120.



TYPE LOCALITY AND HABITAT: Azerbaijan, Caucasus, along the Black Sea, in litter,

under bark of Picea sp. [Pinaceae] and under wings of Hylastes sp. [Coleoptera:



Insectolaelaps pini (Hirschmann, 1960).

218. Insectolaelaps neoarmatus (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Insectolaelaps) neoarmatus Hirschmann and Wiśniewski, 1982: 34.



TYPE LOCALITY AND HABITAT: USA, Moscow, Latah, Idaho, in galleries of

Dendroctonus pseudotsugae [Coleoptera: Curculionidae: Scolytinae].

NOTE: Described on the basis of specimens reported by Hurlbutt (1967): 507 as

Digamasellus armatus (Hirschmann).

219. Insectolaelaps nidiphilus (Wiśniewski and Hirschmann, 1983)

Dendrolaelaps (Insectolaelaps) nidiphilus Wiśniewski and Hirschmann, 1983a: 109.


TYPE LOCALITY AND HABITAT: Poland, Forestry Czolowo, Babki Forest District, 8 May

1983, in bird nest on a broken Pinus sp. [Pinaceae].

220. Insectolaelaps pini (Hirschmann, 1954)

Digamasellus pini Hirschmann, 1954b: 247.

Dendrolaelaps pini.— Hirschmann, 1960: 7; Hirschmann, 1971a: 11; Hirschmann, 1971b: 12;

Hirschmann, 1971c: 16; Hirschmann, 1971d: 18; Hirschmann, 1971e: 21;


204

Dendrolaelaps (Multidendrolaelaps) pini.— Hirschmann, 1974: 61.

Insectolaelaps pini.— Karg, 1993c: 368.

Dendrolaelaps (Insectolaelaps) pini.— Wiśniewski and Hirschmann, 1982: 36.



and under the elytra of Hylurgus ligniperda [Coleoptera: Curculionidae: Scolytinae]

and Hylastes sp. [Coleoptera: Curculionidae: Scolytinae].

NOTE: Specimens reported by Shcherbak (1980): 196 as Insectolaelaps pini (Hirschmann,

1960) were described as Dendrolaelaps (Insectolaelaps) latopini Hirschmann and


221. Insectolaelaps pinisimilis (Hirschmann, 1960)

Dendrolaelaps pinisimilis Hirschmann, 1960: 8.

Dendrolaelaps pinisimilis.— Hirschmann, 1971c: 15.

Dendrolaelaps (Multidendrolaelaps) pinisimilis.— Hirschmann, 1974: 61.

Dendrolaelaps (Insectolaelaps) pinisimilis.— Hirschmann and Wiśniewski, 1982: 37.


TYPE LOCALITY AND HABITAT: Germany, Sieber, in gallery of Ips laricis [Coleoptera:

Curculionidae: Scolytinae]; Germany, Süderlügum, Schleswig-Holstein, in galleries of

Hylastes ater [Coleoptera: Curculionidae: Scolytinae] and Dendroctonus micans

[Coleoptera: Curculionidae: Scolytinae].

222. Insectolaelaps quadrisetoides (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Ipidodendrolaelaps) quadrisetoides Hirschmann and Wiśniewski, 1982: 42.



TYPE LOCALITY AND HABITAT: Russia, Arkhangelsk Oblast and Primorsky Krai, under

bark and in galleries of Ips sp. [Coleoptera: Curculionidae: Scolytinae].


Insectolaelaps quadrisetus (Berlese, 1920).

223. Insectolaelaps quadrisetosimilis (Hirschmann and Rühm, 1955)

Digamasellus quadrisetosimilis Hirschmann and Rühm, 1955: 235.

Dendrolaelaps quadrisetosimilis.— Hirschmann, 1960: 15; Hirschmann, 1971c: 15.

205

Dendrolaelaps (Multidendrolaelaps) quadrisetosimilis.— Hirschmann, 1974: 61.

Insectolaelaps quadrisetosimilis.— Shcherbak, 1980: 192; Karg, 1993c: 368.

Dendrolaelaps (Ipidodendrolaelaps) quadrisetosimilis.— Hirschmann and Wiśniewski, 1982:

41; Wiśniewski and Hirschmann, 1983b: 123.


TYPE LOCALITY AND HABITAT: Germany, Sieber, on stump of Picea sp. [Pinaceae];

Germany, Bückeberg, on stump of Picea sp..

NOTE1: Described from the deutonymph.

NOTE2: Specimens reported by Hurlbutt (1967): 502 as Digamasellus quadrisetosimilis

(Hirschmann and Rühm, 1955) were described as Dendrolaelaps (Ipidodendrolaelaps)

ipidoquadrisetus Wiśniewski and Hirschmann, 1983b: 124.

NOTE3: The specimens reported as Digamasellus quadrisetosimilis (Hirschmann, 1960) by

Hurlbutt (1967): 502 were re-identified as Dendrolaelaps quadrisetus (Berlese, 1920)

by McGraw and Farrier (1969): 121, with what Hirschmann and Wiśniewski (1982):

42 did not agree.

224. Insectolaelaps quadrisetus (Berlese, 1920)

Gamasellus (Digamasellus) quadrisetus Berlese, 1920b: 159.

Gamasellus (Digamasellus) quadrisetus.— Vitzthum, 1923: 115.

Dendrolaelaps quadrisetus.— Vitzthum, 1926b: 425; Oudemans, 1936: 194; Hirschmann,

1960: 5; McGraw and Farrier, 1969: 121; Hirschmann, 1971a: 11; Hirschmann,

1971b: 12; Hirschmann, 1971c: 15; Hirschmann, 1971d: 17; Hirschmann, 1971e: 21;


Digamasellus quadrisetus.— Schweizer, 1949: 33; Franz, 1954: 342; Hirschmann and Rühm,

1955: 235; Hurlbutt, 1967: 499.

Cyrtolaelaps (Digamasellus) quadrisetus.— Ryke, 1962a: 104.

Dendrolaelaps (Multidendrolaelaps) quadrisetus.— Hirschmann, 1974: 61.

Insectolaelaps quadrisetus.— Karg, 1993c: 368.

Dendrolaelaps (Ipidodendrolaelaps) quadrisetus.— Hirschmann and Wiśniewski, 1982: 40.


TYPE LOCALITY AND HABITAT: Italy, Florence, Tuscany, in litter of conifers.

NOTE: Specimens reported by Shcherbak (1980): 193 as Insectolaelaps quadrisetus (Berlese,

1920) were described as Dendrolaelaps (Ipidodendrolaelaps) quadrisetoides


206

225. Insectolaelaps volnyi (Wiśniewski and Hirschmann, 1991)

Dendrolaelaps (Insectolaelaps) volnyi Wiśniewski and Hirschmann, 1991a: 227.


Poland.

TYPE LOCALITY AND HABITAT: China, Pedang Mountains, on Xylotrupes sp.

[Coleoptera: Scarabaeidae] deposited in the Coleoptera collection of Entomologické

Oddělení, The Moravian Museum, Brno, Czech Republic.


226. Insectolaelaps zvoleniensis (Wiśniewski and Hirschmann, 1984)

Dendrolaelaps (Insectolaelaps) zvoleniensis Wiśniewski and Hirschmann, in Hirschmann and



TYPE LOCALITY AND HABITAT: Slovakia, Kováčová, Zvolen District, September 1983,

under the elytron of Cucujus cinnabarinus sp. [Coleoptera: Cucujidae], in galleries of

Coleoptera under bark of Pinus sp. [Pinaceae], under bark of Quercus sp. [Fagaceae]

and in nest of Formica polyctena [Hymenoptera: Formicidae].

NOTE: Described from the deutonymph and adult male.

Genus Longoseius Chant, 1961

Longoseius Chant, 1961: 11 (described in Digamasellidae Evans).

Longoseius.— Lindquist, 1975: 18; Shcherbak, 1980: 170; Hirschmann and Wiśniewski,

1982: 149.

Longoseius (Longoseius).— Lindquist, 1975: 19; Hirschmann and Wiśniewski, 1982: 155.

Type species: Longoseius cuniculus Chant, 1961, by original designation.

Longoseius (Longoseiulus) Lindquist, 1975: 21 (described in Digamasellidae Evans).

Dendrolaelaps (Longoseiulus).— Shcherbak, 1980: 164.

Longoseius (Longoseiulus).— Hirschmann and Wiśniewski, 1982: 156.

Type species: Dendrolaelaps longulus Hirschmann, 1960, by original designation.

207

227. Longoseius aberrans (Hirschmann, 1960)

Dendrolaelaps aberrans Hirschmann, 1960: 7.

Dendrolaelaps aberrans.— Hirschmann, 1971e: 20.

Dendrolaelaps (Dendrolaelaps) aberrans.— Hirschmann, 1974: 62.

Longoseius (Longoseiulus) aberrans.— Lindquist, 1975: 21; Hirschmann and Wiśniewski,

1982: 157.

Dendrolaelaps (Longoseiulus) aberrans.— Shcherbak, 1980: 169.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, on stump of Pinus sp. [Pinaceae].


228. Longoseius brachypoda (Hurlbutt, 1967)

Digamasellus brachypoda Hurlbutt, 1967: 525.

Dendrolaelaps brachypoda.— McGraw and Farrier, 1969: 103.

Dendrolaelaps (Dendrolaelaps) brachypoda.— Hirschmann, 1974: 63.

Longoseius (Longoseiulus) brachypoda.— Lindquist, 1975: 21; Hirschmann and Wiśniewski,

1982: 157.

Dendrolaelaps brachipoda [sic].— Shcherbak, 1980: 169.



TYPE LOCALITY AND HABITAT: USA, Elizabeth, Louisiana, in vacated galleries of

Dendroctonus frontalis [Coleoptera: Curculionidae: Scolytinae] in the inner bark of

Pinus taeda [Pinaceae].

229. Longoseius cuniculus Chant, 1961

Longoseius cuniculus Chant, 1961: 11.

Dendrolaelaps (Dendrolaelaps) cuniculus.— Hirschmann, 1974: 66.

Longoseius (Longoseius) cuniculus.— Lindquist, 1975: 20; Hirschmann and Wiśniewski,

1982: 155.

Longoseius cuniculus.— Hurlbutt, 1967: 527; Lindquist, 1975: 24; Shcherbak, 1980: 170.



TYPE LOCALITY AND HABITAT: USA, Orono, Maine, 20 July 1959, in galleries of

Monochamus notatus [Coleoptera: Cerambycidae] in pine.

208

230. Longoseius longuloides Hirschmann and Wiśniewski, 1982

Longoseius (Longoseiulus) longuloides Hirschmann and Wiśniewski, 1982: 156.



TYPE LOCALITY AND HABITAT: Ukraine, on stumps of Quercus sp. [Fagaceae] and of

other conifers and hardwoods, in litter of a Quercus forest, on stumps, and in galleries

of Cerambyx sp. [Coleoptera: Cerambycidae] and Elater sp. [Coleoptera: Elateridae].


Dendrolaelaps (Longoseiulus) longulus Hirschmann, 1960.

231. Longoseius longulus (Hirschmann, 1960)

Dendrolaelaps longulus Hirschmann, 1960: 7.

Dendrolaelaps longulus.— Hirschmann, 1971b: 12; Hirschmann, 1971c: 15; Hirschmann,

1971d: 18; Hirschmann, 1971e: 19; Karg, 1971: 342.

Dendrolaelaps (Dendrolaelaps) longulus.— Hirschmann, 1974 : 62.

Longoseius (Longoseiulus) longulus.— Lindquist, 1975: 21; Hirschmann and Wiśniewski,

1982: 156.

Longoseius longulus.— Karg, 1993c: 366.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, on stumps of Betula sp.

[Betulaceae], Fagus sp. [Fagaceae] and Pinus sp. [Pinaceae]; Germany, Nuremberg,

under the elytra of Cerambyx sp. [Coleoptera: Cerambycidae] and Elater sp.

[Coleoptera: Elateridae].

NOTE: Specimens reported by Shcherbak (1980): 165 as Dendrolaelaps (Longoseiulus)

longulus Hirschmann, 1960 were described as Longoseius (Longoseiulus) longuloides


232. Longoseius longus (Hirschmann, 1954)

Digamasellus longus Hirschmann, 1954b: 247.

Dendrolaelaps longus.— Hirschmann, 1960: 15; Hirschmann, 1971c: 15.

Dendrolaelaps (Dendrolaelaps) longus.— Hirschmann, 1974 : 62.

Longoseius (Longoseius) longus.— Lindquist, 1975: 20; Hirschmann and Wiśniewski, 1982:

156.

209


TYPE LOCALITY AND HABITAT: Germany, Sieber, Herzberg am Harz, on stump of Picea

sp. [Pinaceae]; Germany, Erlangen, under the elytra of Elater sanguineus [Coleoptera:

Elateridae].


233. Longoseius nobilis (Barilo, 1989)

Dendrolaelaps (Longoseiulus) nobilis Barilo, 1989: 140



TYPE LOCALITY AND HABITAT: Uzbekistan, gardens east of Samarkand, Samarqand

Province, 22 October 1984, in the dust of a hollow plant of Populus sp. [Salicaceae].

234. Longoseius ornatosimilis (Shcherbak, 1980)

Dendrolaelaps (Longoseiulus) ornatosimilis Shcherbak, 1980: 167.

Longoseius (Longoseiulus) ornatosimilis.— Hirschmann and Wiśniewski, 1982: 157.



TYPE LOCALITY AND HABITAT: Russia, Selenge River, Kabansky District, Buryatia, 26

July 1977, on rotting bark of birch wood [Betulaceae] in the mouth of the river.

235. Longoseius ornatus (Hirschmann, 1960)

Dendrolaelaps ornatus Hirschmann, 1960: 8.

Dendrolaelaps ornatus.— Hirschmann, 1971c: 14; Hirschmann, 1971d: 17; Karg, 1971: 341.

Dendrolaelaps (Dendrolaelaps) ornatus.— Hirschmann, 1974: 62.

Longoseius (Longoseiulus) ornatus.— Lindquist, 1975: 21; Hirschmann and Wiśniewski,

1982: 157.

Dendrolaelaps (Longoseiulus) ornatus.— Shcherbak, 1980: 167.

Longoseius ornatus.— Karg, 1993c: 363.


TYPE LOCALITY AND HABITAT: Germany, Bückeberg, on stump of Quercus sp.

[Fagaceae]; Germany, Erlangen, under the elytra of Pyrochroa coccinea [Coleoptera:

Pirochroidae].

210

Genus Multidendrolaelaps Hirschmann, 1974

Dendrolaelaps (Multidendrolaelaps) Hirschmann, 1974: 61 (described in Parasitiformes

Reuter).

Dendrolaelaps (Multidendrolaelaps).— Lindquist, 1975: 41; Hirschmann and Wiśniewski,

1982: 19.

Multidendrolaelaps.— Shcherbak, 1980: 182; Karg, 1993c: 366.

Type species: Dendrolaelaps ulmi Hirschmann, 1960, by original designation.

Dendrolaelaps (Epistodendrolaelaps) Hirschmann and Wiśniewski, 1982: 24 [junior

synonymy by Karg, 1993c: 366].

Type species: Dendrolaelaps euepistomus Hirschmann, 1960, by original designation.

236. Multidendrolaelaps acriluteus (Athias-Henriot, 1961)

Dendrolaelaps acriluteus Athias-Henriot, 1961a: 468.

Dendrolaelaps (Multidendrolaelaps) acriluteus.— Hirschmann, 1974: 61.

Dendrolaelaps (Epistodendrolaelaps ?) acriluteus [sic].— Hirschmann and Wiśniewski,

1982: 28.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, l‘École Nationale d‘Agriculture, Argel, on

vegetable and ornamental crops.


237. Multidendrolaelaps baddeley (Hirschmann and Wiśniewski, 1984)

Dendrolaelaps (Epistodendrolaelaps) baddeley Hirschmann and Wiśniewski, 1984: 97.


TYPE LOCALITY AND HABITAT: South Africa, Lusikisiki, Transkei, 1979, from coastal

evergreen forest.


238. Multidendrolaelaps bakeri (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Epistodendrolaelaps) bakeri Hirschmann and Wiśniewski 1982: 30.

211

Multidendrolaelaps bakeri.— Shcherbak, 1985a: 25.


TYPE LOCALITY AND HABITAT: USA, Washington, District of Columbia, 25 July 1958,

under bark of black locust Robinia pseudoacacia [Fabaceae].

239. Multidendrolaelaps bispinosus (Karg, 1971)

Dendrolaelaps bispinosus Karg, 1971: 331.

Dendrolaelaps (Multidendrolaelaps) bispinosus.— Hirschmann, 1974: 61; Hirschmann and


Multidendrolaelaps bispinosus.— Shcherbak, 1980: 188; Karg, 1993c: 367; Huhta and Karg,

2010: 345.


Germany.

TYPE LOCALITY AND HABITAT: Germany, Wiesenburg, Potsdam-Mittelmark, on rotten

tree stump in a forest of Fagus [Fagaceae] with Rhododendron [Ericaceae].

240. Multidendrolaelaps camerunis (Wiśniewski and Hirschmann, 1984)

Dendrolaelaps (Epistodendrolaelaps) camerunis Wiśniewski and Hirschmann, 1984: 149.


Poland.

TYPE LOCALITY AND HABITAT: Cameroon, Barombi Lake, [Meme], on Chloridolum sp.

[Coleoptera: Cerambycidae] and on unidentified Cerambycidae [Coleoptera] deposited

in the beetle collection of ―Bewnnigsen‖ of ―Zoologisches Institut der Polnischen

Akademie der Wissenschaften‖, Warsaw, Poland.


241. Multidendrolaelaps carinthiacus Schmölzer, 1995

Multidendrolaelaps carinthiacus Schmölzer, 1995a: 500.


TYPE LOCALITY AND HABITAT: Austria, Heiliger-Geist-Gatter (alt. 1,400 m), Saint

Leonhard, Karawanken, Carinthia, 28 July 1979, from the upper soil layer on the edge

of a meadow with a predominance of Arnica montana [Asteraceae].

242. Multidendrolaelaps daelei (Hirschmann and Wiśniewski, 1982)

212

Dendrolaelaps (Epistodendrolaelaps) daelei Hirschmann and Wiśniewski, 1982: 29.


TYPE LOCALITY AND HABITAT: Belgium, Ghent, 19 May 1982, intercepted in boxes

with seeds of Kentia palm Howea sp. [Arecaceae] imported from Lord Howe Island

[Pacific Ocean].

243. Multidendrolaelaps epistospinosus Shcherbak, 1985

Multidendrolaelaps epistospinosus Shcherbak, 1985a: 33.

TYPE DEPOSITORY: The Canadian National Collection of Insects, Arachnids and

Nematodes, Ottawa, Canada.

TYPE LOCALITY AND HABITAT: Canada, Pasadena, Newfoundland, 2 August 1976, in

old Polyporus [Fungi: Polyporaceae] with moss and lichens on trunk of birch Betula

sp. [Betulaceae].

244. Multidendrolaelaps euepistomoides (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Epistodendrolaelaps) euepistomoides Hirschmann and Wiśniewski, 1982: 27.



TYPE LOCALITY AND HABITAT: Ukraine; and Russia, Kamchatka, Russian Far East,

among decaying birch bark.


Multidendrolaelaps eupistomus [sic] (Hirschmann, 1960).

245. Multidendrolaelaps euepistomosimilis (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Epistodendrolaelaps) euepistomosimilis Hirschmann and Wiśniewski, 1982:

28.

Multidendrolaelaps euepistomosimilis.— Shcherbak, 1985a: 35.


TYPE LOCALITY AND HABITAT: Poland, Lopunchowko Forest, Brzezna, 8 and 29 July

1980, in nests of Formica fusca [Hymenoptera: Formicidae].


246. Multidendrolaelaps euepistomus (Hirschmann, 1960)

Dendrolaelaps euepistomus Hirschmann, 1960: 7.

213

Dendrolaelaps euepistomus.— Hirschmann, 1971b: 12; Hirschmann, 1971c: 16; Hirschmann,

1971d: 18; Hirschmann, 1971e: 21; Hirschmann, 1971f: 25; Karg, 1971: 337.

Dendrolaelaps (Multidendrolaelaps) euepistomus.— Hirschmann, 1974: 61.

Dendrolaelaps (Epistodendrolaelaps) euepistomus.— Hirschmann and Wiśniewski, 1982: 27.

Multidendrolaelaps euepistomus.— Shcherbak, 1985a: 34; Karg, 1993c: 367; Huhta and

Karg, 2010: 346.



[Pinaceae]; Germany, Bückeberg, on stumps of Fagus sp. [Fagaceae], Pinus sp.

[Pinaceae] and Quercus sp. [Fagaceae]; Germany, Oberstdorf, on stump of Alnus sp.

[Betulaceae].

NOTE: Specimens reported by Shcherbak (1980): 183 as Multidendrolaelaps eupistomus [sic]

(Hirschmann, 1960) were described as Dendrolaelaps (Epistodendrolaelaps)

euepistomoides Hirschmann and Wiśniewski, 1982: 27.

247. Multidendrolaelaps hurlbutti Shcherbak, 1985

Multidendrolaelaps harlbutti Shcherbak, 1985a: 31.

Multidendrolaelaps hurlbutti.— Shcherbak, 1985a: 32.



TYPE LOCALITY AND HABITAT: Canada, Pasadena, Newfoundland, 30 July 1976, in

small polyporous fungi on the bark of a log.

248. Multidendrolaelaps hexaspinosus (Hirschmann, 1954)

Dendrolaelaps hexaspinosus Hirschmann, 1954a: 107.

Dendrolaelaps hexaspinosus.— Hirschmann, 1960: 6; Hirschmann, 1971c: 15; Hirschmann,

1971d: 17; Hirschmann, 1971e: 20; Hirschmann, 1971f: 24; Karg, 1971: 330.

Dendrolaelaps (Multidendrolaelaps) hexaspinosus.— Hirschmann, 1974: 61; Hirschmann


Multidendrolaelaps hexaspinosus.— Shcherbak, 1980: 190; Karg, 1993c: 367; Huhta and

Karg, 2010: 345.


214

TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stumps of Fagus sp.

[Fagaceae] and Picea sp. [Pinaceae], and in galleries of Hylurgops palliates

[Coleoptera: Curculionidae: Scolytinae]; Austria, Admont, [Styria], on Dryocoetes

autographus [Coleoptera: Curculionidae: Scolytinae].

NOTE: Franz (1954): 342 cited Digamasellus hexaspinosus Hirschmann n. sp., but did not


249. Multidendrolaelaps inconstans Shcherbak, 1985

Multidendrolaelaps inconstans Shcherbak, 1985a: 28.



TYPE LOCALITY AND HABITAT: Canada, Information Centre Area, Kouchibouguac

National Park, New Brunswick, 7 August 1977, under bark of Pinus strobus

[Pinaceae] infested with Ips pini [Coleoptera: Curculionidae: Scolytinae] and

Pityogenes hopkinsi [Coleoptera: Scolytidae].

250. Multidendrolaelaps isodentatus (Hurlbutt, 1967)

Digamasellus isodentatus Hurlbutt, 1967: 504.

Dendrolaelaps isodentatus.— McGraw and Farrier, 1969: 108.

Dendrolaelaps (Multidendrolaelaps) isodentatus.— Hirschmann, 1974: 61.

Dendrolaelaps (Epistodendrolaelaps) isodentatus.— Hirschmann and Wiśniewski, 1982: 29.

Multidendrolaelaps isodentatus.— Shcherbak, 1985a: 25.



TYPE LOCALITY AND HABITAT: USA, Hardin, Texas, 2 September 1963, on inner bark

of Pinus taeda [Pinaceae].

251. Multidendrolaelaps kargi (Hirschmann, 1966)

Dendrolaelaps kargi Hirschmann, 1966c: 39.

Dendrolaelaps (Multidendrolaelaps) kargi.— Hirschmann, 1974: 61.

? Dendrolaelaps kargi [sic].— Hirschmann and Wiśniewski, 1982: 147.


TYPE LOCALITY AND HABITAT: Macquarie Island [between Australia and Antarctica],

Garden Cove, in nest of Eudyptes chrysocome (cited as Euclyptes chrysocome) [Aves:

215

Spheniscidae]; Macquarie Island, Isthmus Sand, on drift wood; Macquarie Island,

North Head, on Agrostis magellanca [Poaceae].

252. Multidendrolaelaps kribii (Wiśniewski and Hirschmann, 1984)

Dendrolaelaps (Epistodendrolaelaps) kribii Wiśniewski and Hirschmann, 1984: 151.


Poland.

TYPE LOCALITY AND HABITAT: Cameroon, Kribi, on Pseudhammus oculifrons (cited as

Chloridolum oculifrons) [Coleoptera: Cerambycidae] deposited in the beetle collection

of ―Bennigsen‖ of ―Zoologisches Institut der Polnischen Akademie der

Wissenschaften‖, Warsaw, Poland.


253. Multidendrolaelaps manualkrantzi (Hirschmann and Wiśniewski, 1982)

Dendrolaelaps (Epistodendrolaelaps) manualkrantzi Hirschmann and Wiśniewski, 1982: 163.

Dendrolaelaps (Epistodendrolaelaps) manualkrantzi.— Hirschmann, 1983c: 128.


TYPE LOCALITY AND HABITAT: USA, Applegate, Oregon, 6 January 1961, on

Physiphora demandata (cited as Chrysomyza demandata) [Diptera: Otitidae].

NOTE: Described on the basis of specimens reported by Krantz (1970): 108 as Digamasellus

sp., and by Krantz (1978): 175 as Dendrolaelaps sp..

254. Multidendrolaelaps multidentatus (Leitner, 1949)

Digamasellus multidentatus Leitner, 1949: 62.

Digamasellus multidentatus.— Franz, 1954: 342.

Dendrolaelaps multidentatus.— Hirschmann, 1960: 7; Athias-Henriot, 1961a: 467;

Hirschmann, 1971d: 18; Karg, 1971: 336; Hirschmann, 1971e: 21.

Cyrtolaelaps (Digamasellus) multidentatus.— Ryke, 1962a: 109.

Dendrolaelaps (Multidendrolaelaps) multidentatus.— Hirschmann, 1974: 61.

Multidendrolaelaps multidentatus.— Shcherbak, 1980: 185; Karg, 1993c: 366; Huhta and

Karg, 2010: 346.

Dendrolaelaps (Epistodendrolaelaps) multidentatus.— Hirschmann and Wiśniewski, 1982:

27.


216

TYPE LOCALITY AND HABITAT: Austria, Säusenstein, Melk District, in compost.

255. Multidendrolaelaps putte Huhta and Karg, 2010

Multidendrolaelaps putte Huhta and Karg, 2010: 341.


TYPE LOCALITY AND HABITAT: Finland, Lammi, 7 September 2007, on decaying trunk

of aspen Populus sp. [Salicaceae].

256. Multidendrolaelaps querci (Hirschmann, 1960)

Dendrolaelaps querci Hirschmann, 1960: 7.

Dendrolaelaps querci.— Hirschmann, 1971b: 12; Hirschmann, 1971c: 15; Hirschmann,

1971e: 20; Hirschmann, 1971f: 24.

Dendrolaelaps (Multidendrolaelaps) querci.— Hirschmann, 1974: 61; Hirschmann and

Wiśniewski, 1982 : 22.

Multidendrolaelaps querci.— Shcherbak, 1980: 186; Karg, 1993c: 367; Huhta and Karg,

2010: 345.


TYPE LOCALITY AND HABITAT: Germany, Hasbruch, [Oldenburg], on stump of Quercus

sp. [Fagaceae].

NOTE: Described from the protonymph, deutonymph and adult male.

257. Multidendrolaelaps schusteri (Hirschmann, 1966)

Dendrolaelaps schusteri Hirschmann, 1966c: 39.

Dendrolaelaps (Multidendrolaelaps) schusteri.— Hirschmann, 1974: 61.

? Dendrolaelaps schusteri [sic].— Hirschmann and Wiśniewski, 1982: 147.



Camp Hill, in grassland soil and in litter of Poa halmitoni [Poaceae]; Macquarie

Island, Aerial Cove, on Colobanthus muscoides [Caryophyllaceae]; Macquarie Island,

Nuggets Point, in litter of Stilbocarpa sp. [Areliaceae], Cotula sp. [Asteraceae] and

Colobanthus sp. [Caryophyllaceae].

258. Multidendrolaelaps spinosus (Hirschmann, 1960)

Dendrolaelaps spinosus Hirschmann, 1960: 6.

217

Dendrolaelaps spinosus.— Hirschmann, 1971a: 10; Hirschmann, 1971b: 12; Hirschmann,

1971c: 16; Hirschmann, 1971d: 17; Hirschmann, 1971f: 24; Karg, 1971: 328.

Dendrolaelaps (Multidendrolaelaps) spinosus.— Hirschmann, 1974: 61; Hirschmann and


Multidendrolaelaps spinosus.— Shcherbak, 1980: 185; Shcherbak, 1985a: 30; Karg, 1993c:

367; Huhta and Karg, 2010: 345.


TYPE LOCALITY AND HABITAT: Germany, Oberstdorf, on stump of Alnus sp.

[Betulaceae]; Germany, Erlangen, in a nest of Myrmica ruginodis [Hymenoptera:

Formicidae] and under the elytron of Lamia textor [Coleoptera: Cerambycidae].

258a. Multidendrolaelaps spinosus elongatus Shcherbak, 1985

Multidendrolaelaps spinosus elongatus Shcherbak, 1985a: 30.



TYPE LOCALITY AND HABITAT: Canada, Waterton Lakes National Park, Alberta,

2-5 September 1980, in strips of small bracket fungi on spruce [Picea sp.

(Pinaceae)].

259. Multidendrolaelaps subcorticalis Huhta and Karg, 2010

Multidendrolaelaps subcorticalis Huhta and Karg, 2010: 344.

TYPE DEPOSITORY: Zoological Museum, University of Turku, Finland.

TYPE LOCALITY AND HABITAT: Finland, Parainen, 28 March 1983, in old scolytid

[Coleoptera: Curculionidae: Scolytinae] galleries under bark of pine [Pinaceae].

260. Multidendrolaelaps templei (Hunter, 1970)

Digamasellus templei Hunter, 1970: 59.

? Dendrolaelaps templei [sic].— Hirschmann and Wiśniewski, 1982: 147.

TYPE DEPOSITORY: CSIRO, Canberra, Australia.

TYPE LOCALITY AND HABITAT: Heard Island [Southern Ocean], Polly Gully, 6 February

1965, from burrow of Pachyptila desolata [Aves: Procellariidae].

261. Multidendrolaelaps tetraspinosus (Hirschmann, 1954)

Dendrolaelaps tetraspinosus Hirschmann, 1954a: 107.

218

Dendrolaelaps tetraspinosus.— Hirschmann, 1960: 6; McGraw and Farrier, 1969: 128;

Hirschmann, 1971a: 10; Hirschmann, 1971b: 12; Hirschmann, 1971c: 16;

Hirschmann, 1971d: 17; Hirschmann, 1971e: 20; Hirschmann, 1971f: 24; Karg, 1971:

330.

Dendrolaelaps (Multidendrolaelaps) tetraspinosus.— Hirschmann, 1974: 61; Hirschmann


Multidendrolaelaps tetraspinosus.— Shcherbak, 1980: 191; Shcherbak, 1985a: 25; Karg,

1993c: 367; Huhta and Karg, 2010: 345.


TYPE LOCALITY AND HABITAT: Germany, Erlangen, on stump of Picea sp. [Pinaceae];

Germany, Nuremberg, on stump of Pinus sp. [Pinaceae]; Austria, Admont, [Styria], on

stump of Abies sp. [Pinaceae]; Germany, Schönau im Schwarzwald, in gallery of

Pissodes piceae [Coleoptera: Curculionidae: Molytinae].

NOTE: Franz (1954): 342 cited Digamasellus tetraspinosus Hirschmann n. sp., but did not


262. Multidendrolaelaps trispinosus Shcherbak, 1985

Multidendrolaelaps trispinosus Shcherbak, 1985a: 25.



TYPE LOCALITY AND HABITAT: Canada, Central Experimental Farm, Ottawa, Ontario,

16 June 1971, on stump of Acer sp. [Sapindaceae] or [sic] Ulmus sp. [Ulmaceae].

263. Multidendrolaelaps ulmi (Hirschmann, 1960)

Dendrolaelaps ulmi Hirschmann, 1960: 7.

Dendrolaelaps ulmi.— Hirschmann, 1971a: 11; Hirschmann, 1971b: 12; Hirschmann, 1971c:

16; Hirschmann, 1971d: 17; Hirschmann, 1971e: 20; Hirschmann, 1971f: 24; Karg,

1971: 330.

Dendrolaelaps (Multidendrolaelaps) ulmi.— Hirschmann, 1974: 61; Hirschmann and


Multidendrolaelaps ulmi.— Shcherbak, 1980: 188; Shcherbak, 1985a: 28; Karg, 1993c: 367;

Huhta and Karg, 2010: 345.


219

TYPE LOCALITY AND HABITAT: Germany, Erlangen and Munich, in gallery of Scolytus

scolytus [Coleoptera: Curculionidae: Scolytinae] in Ulmus sp. [Ulmaceae].

264. Multidendrolaelaps unispinatus (Ishikawa, 1977)

Dendrolaelaps unispinatus Ishikawa, 1977: 99.

Dendrolaelaps (Epistodendrolaelaps) unispinatus.— Hirschmann and Wiśniewski, 1982: 30.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, Ikata, Nishiuwa District, Ehime Prefecture, 9

July 1976, on Monochamus alternatus [Coleoptera: Cerambycidae].

265. Multidendrolaelaps watsoni (Hirschmann, 1966)

Dendrolaelaps watsoni Hirschmann, 1966c: 38.

Dendrolaelaps (Multidendrolaelaps) watsoni.— Hirschmann, 1974: 61.

? Dendrolaelaps watsoni [sic].— Hirschmann and Wiśniewski, 1982: 146.



Aerial Cove, on Colobanthus sp. [Caryophyllaceae]; Macquarie Island, Gadget Gully,

in moss; Macquarie Island, Wireless Hill, in moss; Macquarie Island, Garden Cove, in

litter of Pleurophyllum sp. [Asteraceae]; Macquarie Island, North Head, on algae on

rocks.

Genus Oligodentatus Shcherbak, 1980

Oligodentatus Shcherbak, 1980: 173 (described in Rhodacaridae Oudemans).

Dendrolaelaps (Oligodentatus).— Hirschmann and Wiśniewski, 1982: 18.

Type species: Oligodentatus tridentatus Shcherbak and Bregetova, 1980, by original

designation.

266. Oligodentatus certus Barilo, 1989

Oligodentatus certus Barilo, 1989: 142.



220


October 1986, in cow manure.

267. Oligodentatus fimetarius (Karg, 1965)

Dendrolaelaps fimetarius Karg, 1965: 297.

Dendrolaelaps (Tridendrolaelaps) fimetarius.— Karg, 1971: 335.

Dendrolaelaps (Tridendrolaelaps) fimetarius.— Hirschmann, 1974: 60.

Oligodentatus fimetarius.— Shcherbak, 1980: 175.

Dendrolaelaps (Oligodentatus ?) fimetarius [sic].— Hirschmann and Wiśniewski, 1982: 18.

Punctodendrolaelaps fimetarius.— Karg, 1993c: 351.

TYPE DEPOSITORY: Institut für Pflanzenschutzforschung.— Biologische Zentralanstalt der

deutschen Akademie der Landwirtschaftswissenschaften zu Berlin, Kleinmachnow,

Germany.

TYPE LOCALITY AND HABITAT: Germany, Manschnow, [Märkisch-Oderland,

Brandenburg], 1962, in one- year-old manure.

268. Oligodentatus shcherbakae Barilo, 1989

Oligodentatus shcherbakae Barilo, 1989: 141.




January 1985, in the dust of a hollow plant of Populus sp. [Salicaceae].

269. Oligodentatus tridentatus Shcherbak and Bregetova, 1980

Oligodentatus tridentatus Shcherbak and Bregetova, in Shcherbak, 1980: 174.

Dendrolaelaps (Oligodentatus) tridentatus.— Hirschmann and Wiśniewski, 1982: 18.



TYPE LOCALITY AND HABITAT: Russia, Sakhalin Islands, 1 May 1959, in nest of

Pteromys volans [Mammalia: Rodentia: Sciuridae].

Genus Orientolaelaps Bregetova and Shcherbak, 1977

221

Orientolaelaps Bregetova and Shcherbak, 1977b: 175 (described in Rhodacaridae

Oudemans).

Orientolaelaps.— Shcherbak, 1980: 200.

Type species: Orientolaelaps eutamiasi Bregetova and Shcherbak, 1977, by original

designation.

270. Orientolaelaps eutamiasi Bregetova and Shcherbak, 1977

Orientolaelaps eutamiasi Bregetova and Shcherbak, 1977b: 175.

Orientolaelaps eutamiasi.— Shcherbak, 1980: 201.



TYPE LOCALITY AND HABITAT: Russia, summit of Sikhote-Alin, Primorsky Kray, 6

June 1961, from nest of Eutamias sibiricus [Mammalia: Rodentia: Sciuridae].

Genus Panteniphis Willmann, 1949

Panteniphis Willmann, 1949: 342 (described in Parasitiformes Reuter).

Panteniphis.— Athias-Henriot, 1969: 70; Bregetova, 1977: 183; Hirschmann, 1983a: 18;

Karg, 1993c: 369.

Panteniphis (Panteniphis).— Hirschmann, 1983a: 18.

Type species: Panteniphis mirandus Willmann, 1949, by original designation.

Lindquistoseius Genis, Loots and Ryke, 1969: 109 (described in Ascidae Oudemans) [junior

synonymy by Hurlbutt, 1975: 36].

Panteniphis (Lindquistoseius).— Hurlbutt, 1975: 36; Hirschmann, 1983a: 19.

Type species: Lindquistoseius africanus Genis, Loots and Ryke, 1969, by original

designation.

271. Panteniphis africanus Genis, Loots and Ryke, 1969

Lindquistoseius africanus Genis, Loots and Ryke, 1969: 109.

TYPE DEPOSITORY: Museu do Dundo, Lunda-Norte, Angola.

222

TYPE LOCALITY AND HABITAT: Angola, Tshitengo River (7°40‘S, 21°45‘E), affluent of

the Luango River and subafluente of the Kasai River, Lunda, 6 February 1963, in

forest soil.

NOTE: Specimens reported by Hurlbutt (1975): 36 as Panteniphis (Lindquistoseius) africanus

(Genis, Loots and Ryke, 1969) were described as Panteniphis (Lindquistoseius)

tanzaniae Hirschmann, 1983a: 19.

272. Panteniphis athiasae Hirschmann, 1983

Panteniphis (Panteniphis) athiasae Hirschmann, 1983a: 18.


Paris, France.

TYPE LOCALITY AND HABITAT: France, Station F/150, few km east of Lormes, Morvan,

30 May 1966, in a marshy meadow.

NOTE1: Described on the basis of specimen reported by Athias-Henriot (1969): 72 as adult

male of Panteniphis mirandus Willmann, 1949.

NOTE2: Described from the adult male.

273. Panteniphis mirandus Willmann, 1949

Panteniphis mirandus Willmann, 1949: 344.

Panteniphis mirandus.— Hirschmann, 1983a: 18; Karg, 1993c: 369; Gwiazdowicz, 2000:

465.


TYPE LOCALITY AND HABITAT: Germany, Panten, Herzogtum Lauenburg, Schleswig-

Holstein, July-August 1943, from unspecified substrate.

NOTE: Specimen reported by Athias-Henriot (1969): 72 as adult male of Panteniphis

mirandus Willmann, 1949 was described as Panteniphis (Panteniphis) athiasae

Hirschmann, 1983a: 18.

274. Panteniphis tanzaniae Hirschmann, 1983

Panteniphis (Lindquistoseius) tanzaniae Hirschmann, 1983a: 19.


Paris, France.

TYPE LOCALITY AND HABITAT: Tanzania, near Lyamungu Research and Training

Centre (alt. 1,300 m), Mount Kilimanjaro, 9 June 1972, on dead leaves and sticks in a

223

forest patch; Tanzania, near Morogoro (alt. 1,000 m), 18 January 1967, on dead grass

near a stream; Tanzania, near Morogoro (alt. 1,100 m), 30 August 1967, in litter the

edge of a clearing; Tanzania, near Morogoro (alt. 550 m), 26 March 1968, in litter

under bushes.

NOTE: Described on the basis of specimens reported by Hurlbutt (1975): 36 as Panteniphis

(Lindquistoseius) africanus (Genis, Loots and Ryke, 1969).

Genus Pontiolaelaps Luxton, 1989

Pontiolaelaps Luxton, 1989: 413 (described in Digamasellidae Evans).

Type species: Dendrolaelaps (Pontiolaelaps) crenatus Luxton, 1984, by original

designation.

Dendrolaelaps (Pontiolaelaps) Luxton, 1984: 83 (nomen nudum).

NOTE: Pontiolaelaps was described as a subgenus of Dendrolaelaps by Luxton (1984): 83,

but the name was not made available because the type species of the subgenus was not

fixed (International Code of Zoological Nomenclature, Article 13.3). The name only

became available when Luxton (1989): 413 raised it to genus level and defined

Dendrolaelaps (Pontiolaelaps) crenatus Luxton, 1984 as type species.

275. Pontiolaelaps crenatus (Luxton, 1984)

Dendrolaelaps (Pontiolaelaps) crenatus Luxton, 1984: 84.

Pontiolaelaps crenatus.— Luxton, 1989: 415.

TYPE DEPOSITORY: Auckland Museum, Auckland, New Zealand.

TYPE LOCALITY AND HABITAT: New Zealand, Omaha Cove, Leigh, North Auckland, 5

December 1968, in the intertidal zone of a sheltered rock platform.

276. Pontiolaelaps salinus Luxton, 1989

Pontiolaelaps salinus Luxton, 1989: 413.


224

TYPE LOCALITY AND HABITAT: New Zealand, below [sic] Marine Biology Laboratory,

Portobello, Otago Peninsula, February 1968, on the alga Bostrychia sp.

[Rhodomelaceae].

277. Pontiolaelaps terebratus (Luxton, 1984)

Dendrolaelaps (Pontiolaelaps) terebratus Luxton, 1984: 92.

Pontiolaelaps terebratus.— Luxton, 1989: 415.


TYPE LOCALITY AND HABITAT: New Zealand, opposite Goat Island, Leigh, North

Auckland, 5 December 1968, from Elminius sp. [Crustacea: Balanidae] on rock of an

exposed rock platform.

Species incertae sedis

001. Asca muricata Fox, 1947

.Asca muricata Fox, 1947: 600

Cyrtolaelaps (Digamasellus) muricatus.— Ryke, 1961a: 109.

TYPE DEPOSITORY: Entomological Collection of the School of Tropical Medicine, San

Juan, Puerto Rico.

TYPE LOCALITY AND HABITAT: Puerto Rico, San Juan, 16 May 1947, from Rattus

norvegicus [Mammalia: Rodentia: Muridae].

002. Cyrtolaelaps (Gamasellus) armatus Berlese, 1904

Cyrtolaelaps (Gamasellus) armatus Berlese, 1904: 279.

Digamasellus armatus.— Hirschmann, 1954b: 246; Franz, 1954: 341.

Cyrtolaelaps (Digamasellus) armatus.— Ryke, 1962a: 107.

Gamasellus spalacis Oudemans, 1912: 261 [objective synonym — unjustified replacement

name].

Cyrtolaelaps (Gamasellus) spalacis.— Ryke, 1962b: 52.

Gamasellus spalacis.— Lee, 1970: 150.


TYPE LOCALITY AND HABITAT: Germany, Bremen, in a mole nest [Mammalia:

Talpidae].

225

NOTE: Described on the basis of specimens reported by Oudemans (1904): 78 as adult male

of Parasitus spinipes (Koch).

003. Digamasellus arcuatus Willmann, 1939

Digamasellus arcuatus Willmann, 1939: 456.

Cyrtolaelaps (Digamasellus) arcuatus.— Ryke, 1962a: 107.


TYPE LOCALITY AND HABITAT: Glatzer Schneeberg, between Czech Republic and

Poland, in soil.


004. Digamasellus gradatus Willmann, 1938

Digamasellus gradatus Willmann, 1938: 159.

Cyrtolaelaps (Digamasellus) gradatus.— Ryke, 1962a: 106.


TYPE LOCALITY AND HABITAT: Slovakia, Kremnica (cited as Körmöcbánya), 25 May

1933, in soil.

005. Gamasellus inermis Halbert, 1920

Gamasellus inermis Halbert, 1920: 117.

Cyrtolaelaps (Digamasellus) inermis.— Ryke, 1962a: 103.


TYPE LOCALITY AND HABITAT: Ireland, Malahide, February and September 1919, in

fissures and between flakes [sic] on the seashore in an intertidal zone.

006. Gamasellus (di) claviger Lombardini, 1941

Gamasellus (di) claviger Lombardini, 1941: 183.

Cyrtolaelaps (Digamasellus) claviger.— Ryke, 1962a: 107.


TYPE LOCALITY AND HABITAT: Somalia, Caschei, 20 July 1939, on Onitis alexis

[Coleoptera: Scarabaeidae] near a river.

007. Gamasellus (Di) cultriger Lombardini, 1940

Gamasellus (Di) cultriger Lombardini, 1940: 13.

226

Cyrtolaelaps (Digamasellus) cultriger.— Ryke, 1962a: 107.


TYPE LOCALITY AND HABITAT: Brazil, under the elytron of a passalid beetle

[Coleoptera].

NOTE: Described from the nymph.

008. Gamasellus (Digamasellus) gracilis Berlese, 1920

Gamasellus (Digamasellus) gracilis Berlese, 1920b: 159.

Cyrtolaelaps (Digamasellus) gracilis.— Ryke, 1962a: 104.

Digamasellus gracilis.— Bernini, Castagnoli and Nannelli, 1995: 19.


TYPE LOCALITY AND HABITAT: Italy, Florence, Tuscany, in litter of conifers.

009. Gamasellus (Digamasellus) innumerus Berlese, 1920

Gamasellus (Digamasellus) innumerus Berlese, 1920b: 161.

Dendrolaelaps innumerus.— Hirschmann, 1960: 9.

Cyrtolaelaps (Digamasellus) innumerus.— Ryke, 1962a: 105.


TYPE LOCALITY AND HABITAT: Indonesia, Java Island, under wings of Copridis molussi

[sic] [= Catharsius molossus ?] [Coleoptera: Scarabaeidae]; South Africa, Cape of

Good Hope, on C. hamadryadis [sic] [= Heliocopris hamadryas ?] [Coleoptera:

Scarabaeidae].

010. Gamasellus (Digamasellus) rhodacaroides Berlese, 1920

Gamasellus (Digamasellus) rhodacaroides Berlese, 1920b: 162.

Cyrtolaelaps (Digamasellus) rhodacaroides.— Ryke, 1962a: 105.

Digamasellus rhodacaroides.— Bernini, Castagnoli and Nannelli, 1995: 20.


TYPE LOCALITY AND HABITAT: Italy, Florence, Tuscany, in litter and humus in pots in a

greenhouse.

011. Gamasellus (Digamasellus) simplex Berlese, 1920

Gamasellus (Digamasellus) simplex Berlese, 1920b: 163.

Cyrtolaelaps (Digamasellus) simplex.— Ryke, 1962a: 106.

227

Digamasellus simplex.— Bernini, Castagnoli and Nannelli, 1995: 20.


TYPE LOCALITY AND HABITAT: Italy, Portici, Province of Naples, in moss.

012. Panteniphis rhombus Ma and Lin, 2007

Panteniphis rhombus Ma and Lin, 2007: 7.

TYPE DEPOSITORY: Medical Entomology Gallery, Institute of Microbiology and

Epidemiology, Academy of Military Medical Sciences, Beijing, China.

TYPE LOCALITY AND HABITAT: China, Hua‘an (25°00‘N, 117°34‘E), Fujian Province,

23 October 1996, in tree holes.

Nomina nuda

001. Digamasellus nidophilus Hirschmann, 1958

Digamasellus nidophilus Hirschmann, in Sellnick, 1958: 22.

NOTE: The author did not provide any morphological characteristics about the species,

constituting a nomen nudum.

Species previously considered in genera of Digamasellidae and now placed in other

families

Cyrtolaelaps (Digamasellus) seminudus Ryke, 1962a: 98 [Gamasellodes seminudus (Ascidae)

— Hurlbutt, 1967: 498].

Digamasellus circuliformis Leitner, 1949: 59 [junior synonymy of Gamasellodes bicolor

(Berlese, 1918) (Ascidae) — Bernhard, 1963: 105].

? Digamasellus concina [sic] Womersley, 1942: 159 [Gamasellus concinus (Ologamasidae)

— Lee, 1970: 131].

? Digamasellus punctatus [sic] Womersley, 1942: 160 [Acugamasus punctatus

(Ologamasidae) — Lee, 1970: 142].

228

? Digamasellus semipunctatus [sic] Womersley, 1942: 163 [Acugamasus semipunctatus

(Ologamasidae) — Lee, 1970: 142].

Digamasellus shealsi Costa, 1962: 486 [junior synonymy of Gamasellodes bicolor (Berlese,

1918) (Ascidae) — Hurlbutt, 1970: 475].

Digamasellus tasmanicus Womersley, 1956b: 538 [Gamasellus tasmanicus (Ologamasidae)

— Lee, 1970: 137].

? Digamasellus tragardhi [sic] Womersley, 1942: 161 [Gamasellus tragardhi (Ologamasidae)

— Lee, 1970: 135].

Gamasellus americanus Garman, 1948: 9 [junior synonym of Gamasellodes bicolor (Berlese,

1918) (Ascidae) — Hurlbutt, in Baker, Delfinado and Abbatiello, 1976: 63].

Gamasellus (Digamasellus) bicolor Berlese, 1918: 135 [Gamasellodes bicolor (Ascidae) —

Hurlbutt, 1970: 475].

Gamasellus (Digamasellus) magnituberculatus Vitzthum, 1925: 5 [Asca magnituberculatus

[sic] (Ascidae) — Wharton, 1946: 180].

Gamasus racovitzai Trouessart, 1903: 8 [cited as Gamasellus (Digamasellus) racovitzai.—

Berlese, 1917: 5] [Hydrogamasellus racovitzai (Ologamasidae) — Lee, 1970].

3.3.3 Catalogue of world species of Laelaptonyssidae Womersley

Genus Starkovia Lombardini, 1947

Starkovia Lombardini, 1947: 175 (described in Parasitidae Womersley).

Type species: Starkovia termitophila Lombardini, 1947, by monotypy.

229

Laelaptonyssus Womersley, 1956b: 543 (described in Laelaptonyssidae Womersley) [junior

synonymy]

Laelaptonyssus.— Lee, 1970: 72; Halliday, 1987: 85; Krantz, 2000: 26; Halliday, 2006: 29.

Type species: Laelaptonyssus mitis Womersley, 1956, by original designation.

Puchihlungia Samšiňák, 1964: 39 (described in Rhodacaridae Oudemans) [junior synonymy

of Laelaptonyssus by Lee, 1970: 72].

Type species: Puchihlungia chinensis Samšiňák, 1964, by original designation.

001. Starkovia chinensis (Samšiňák, 1964)

Puchihlungia chinensis Samšiňák, 1964: 39.

Laelaptonyssus chinensis.— Lee, 1970: 74; Halliday, 1987: 86; Krantz, 2000: 29.

TYPE DEPOSITORY: Institute of Zoology, Chinese Academy of Sciences, Beijing, China.

TYPE LOCALITY AND HABITAT: China, near Kao-Ho, Guangdong [cited as Kanton

Province], 1963, on Coptotermes formosanus [Isoptera: Rhinotermitidae].

002. Starkovia darwiniensis (Halliday, 1987)

Laelaptonyssus darwiniensis Halliday, 1987: 89.

Laelaptonyssus darwiniensis.— Krantz, 2000: 29.

TYPE DEPOSITORY: Australian National Insect Collection, CSIRO, Canberra, Australia.

TYPE LOCALITY AND HABITAT: Australia, Darwin, Northern Territory, July 1984, in

nest of Coptotermes acinaciformis [Isoptera: Rhinotermitidae].

003. Starkovia hallidayi (Krantz, 2000)

Laelaptonyssus hallidayi Krantz, 2000: 31.

Laelaptonyssus hallidayi.— Halliday, 2006: 29.


TYPE LOCALITY AND HABITAT: Australia, Brindabella Range, Australian Capital

Territory, 31 May 1978, in nest of Porotermes adamsoni [Isoptera: Termopsidae].

004. Starkovia lacticolus (Halliday, 2006)

Laelaptonyssus lacticolus Halliday, 2006: 29.


230

TYPE LOCALITY AND HABITAT: Australia, Tallaganda State Forest, near Captain's Flat

(35°38'S 149°33'E), New South Wales, 10 August 1989, in nest of Coptotermes

lacteus (Isoptera: Rhinotermitidae).

005. Starkovia lenzi (Halliday, 1987)

Laelaptonyssus lenzi Halliday, 1987: 86.

Laelaptonyssus lenzi.— Krantz, 2000: 29; Halliday, 2006: 29.


TYPE LOCALITY AND HABITAT: Australia, Termeil, near Batemans Bay, New South

Wales, 16 November 1983, in nest of Coptotermes lacteus [Isoptera: Rhinotermitidae].

006. Starkovia mitis (Womersley, 1956)

Laelaptonyssus mitis Womersley, 1956b: 543.

Laelaptonyssus mitis.— Lee, 1970: 74; Krantz, 2000: 29; Halliday, 1987: 85.


TYPE LOCALITY AND HABITAT: Australia, Department of Zoology, University of

Western Australia, [Perth], October 1950, from a housefly colony.

007. Starkovia setosus (Krantz, 2000)

Laelaptonyssus setosus Krantz, 2000: 29.



TYPE LOCALITY AND HABITAT: USA, Stoneville, Washington County, Mississippi, 2

July 1999, associated with Reticulitermes flavipes [Isoptera: Rhinotermitidae].

008. Starkovia termitophila Lombardini, 1947

Starkovia termitophila Lombardini, 1947: 177.


TYPE LOCALITY AND HABITAT: Italy, Lake Albano, [Roma Province], 1947, in nest of

Reticulitermes lucifugus [Isoptera: Rhinotermitidae] under a stone on the banks of the

lake.

231

Species previously considered in a genus of Laelaptonyssidae and now placed in another

family

Laelaptonyssus phytoseioides Baker and Johnston, 1959: 275 [Treatia phytoseioides

(Otopheidomenidae) — Krantz and Khot, 1962: 535].

3.3.4 Catalogue of world species of Ologamasidae Ryke

Genus Acugamasus Lee, 1970

Acugamasus Lee, 1970: 139 (described in Rhodacaridae Oudemans).

Acugamasus.— Karg, 1977: 342; Karg, 1993a: 52; Karg, 1997b: 38.

Type species: Digamasellus punctatus Womersley, 1942, by original designation.

001. Acugamasus cursor Lee, 1970

Acugamasus cursor Lee, 1970: 142.

Acugamasus cursor.— Karg, 1993a: 53; Karg, 1997b: 38.


TYPE LOCALITY AND HABITAT: Australia, Beauchamp Falls, near Beech Forest, Otway

Ranges, Victoria, 9 December 1965, on moss on rocks and rotting tree stumps.

002. Acugamasus drakensbergensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) drakensbergensis Ryke, 1962b: 24.

Acugamasus drakensbergensis.— Lee, 1970: 144.

TYPE DEPOSITORY: Natal Museum, Pietermaritzburg, South Africa.

TYPE LOCALITY AND HABITAT: South Africa, Champagne Castle, [Drakensberg],

KwaZulu-Natal, January 1944, on forest floor.

003. Acugamasus elachyaspis Lee, 1973

Acugamasus elachyaspis Lee, 1973: 21.

Acugamasus elachyaspis.— Karg, 1993a: 54; Karg, 1997b: 38.


232

TYPE LOCALITY AND HABITAT: Australia, Grange Golf Club (approximately 1.5 km

from the sea), Adelaide, South Australia, 23 May 1965, on moss.

004. Acugamasus grahami (Ryke, 1962)

Cyrtolaelaps (Gamasellus) grahami Ryke, 1962b: 38.

Acugamasus grahami.— Lee, 1970: 144; Halliday, 2005: 41.

TYPE DEPOSITORY: Institute for Zoological Research, Potchefstroom University,


TYPE LOCALITY AND HABITAT: South Africa, Bathurst, [Eastern Cape], January 1956,

in bush soil cores.

005. Acugamasus hluhluwensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) hluhluwensis Ryke, 1962b: 23.

Acugamasus hluhluwensis.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, Hluhluwe Game Reserve, KwaZulu-

Natal, 1941, on forest floor.

006. Acugamasus knysnaensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) knysnaensis Ryke, 1962b: 32.

Acugamasus knysnaensis.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, Knysna Forest, December 1943, on forest

floor.

007. Acugamasus losovensis (Pinchuk, 1972)

Gamasellus losovensis Pinchuk, 1972b: 69.

TYPE DEPOSITORY: Institute of Zoology and Parasitology SSR Moldova, Chişinău,

Moldova.

TYPE LOCALITY AND HABITAT: Moldova, Chişinău, 20 April 1969, in nest of Apodemus

agrarius [Mammalia: Rodentia: Muridae].

008. Acugamasus macrosetosus (Ryke, 1962)

Cyrtolaelaps (Gamasellus) macrosetosus Ryke, 1962b: 30.

233

Acugamasus macrosetosus.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, Knysna Forest, December 1943, on forest

floor.

009. Acugamasus montanus (Willmann, 1936)

Cyrtolaelaps montanus Willmann, 1936: 273.

Gamasellus montanus.— Franz, 1954: 340; Hirschmann, 1962: 49; Lee, 1970: 131; Karg,

1971: 350; Bregetova and Shcherbak, 1977a: 302; Davydova, 1982: 9; Karg, 1993c:

371.

Cyrtolaelaps (Gamasellus) montanus.— Ryke, 1962b: 54.


TYPE LOCALITY AND HABITAT: Glatzer Schneeberg (alt. 1,200 m), between Czech

Republic and Poland, 6 May 1934 and 25 August 1934, in soil of Avena sp. [Poaceae]

meadows.

NOTE1: Hirschmann (1962): 52 considered Gamasellus silvestris Halašková, 1958 as junior

synonymy of Acugamasus montanus (Willmann, 1936), but Davydova (1982): 9


NOTE2: Bregetova and Shcherbak (1977a): 302 considered Gamasellus alifanovi Davydova,

1970: 82 as junior synonymy of Acugamasus montanus (Willmann, 1936), but

Davydova (1982): 9 revoked that synonymy.

NOTE3: The specimens identified by Ishikawa (1983): 113 as Gamasellus montanus

(Willmann) belong to a different species of Gamasellus (not of Acugamasus).

010. Acugamasus natalensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) natalensis Ryke, 1962b: 27.

Acugamasus natalensis.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, High Flats, South Coast, KwaZulu-Natal,

December 1943, South Africa, on forest floor.

011. Acugamasus nepotulus (Berlese, 1908)

Gamasellus nepotulus Berlese, 1908: 13.

Cyrtolaelaps (Gamasellus) nepotulus.— Ryke, 1962b: 44.

234

Gamasellus nepotulus.— Lee, 1970: 131.


TYPE LOCALITY AND HABITAT: Italy, Rosignano, from unspecified substrate.

012. Acugamasus neotasmanicus (Ryke, 1962)

Cyrtolaelaps (Gamasellus) neotasmanicus Ryke, 1962b: 36.

Acugamasus neotasmanicus.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, Richards Bay, KwaZulu-Natal, December

1943, from unspecified substrate.

013. Acugamasus paranatalensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) paranatalensis Ryke, 1962b: 30.

Acugamasus paranatalensis.— Lee, 1970: 144.


TYPE LOCALITY AND HABITAT: South Africa, Champagne Castle, [Drakensberg],

KwaZulu-Natal, January 1944, on forest floor.

014. Acugamasus parvipectus Karg, 1977

Acugamasus parvipectus Karg, 1977: 342.

Acugamasus parvipectus.— Karg, 1993a: 54; Karg, 1997b: 38.


TYPE LOCALITY AND HABITAT: Chile, Cuesta El Melón, [Valparaíso Province], 3

November 1965, in humus under bushes.

015. Acugamasus plumitergus Karg, 1997

Acugamasus plumitergus Karg, 1997b: 38.



TYPE LOCALITY AND HABITAT: New Caledonia, Mont Panié (alt. 400 m), 1977, on

moss and fern roots [Pteridophyta].

016. Acugamasus punctatus (Womersley, 1942)

? Digamasellus punctatus [sic] Womersley, 1942: 160.

235

Cyrtolaelaps punctatus.— Womersley, 1961: 194.

Cyrtolaelaps (Gamasellus) punctatus.— Ryke, 1962b: 54.

Acugamasus punctatus.— Lee, 1970: 142; Lee, 1973: 20; Karg, 1993a: 54; Karg, 1997b: 38.


TYPE LOCALITY AND HABITAT: Australia, Adelaide, South Australia, June 1935, from

unspecified substrate; Australia, Long Gully, Belair National Park, South Australia,

August 1938, from unspecified substrate.

017. Acugamasus semipunctatus (Womersley, 1942)

? Digamasellus semipunctatus [sic] Womersley, 1942: 163.

Digamasellus semipunctatus.— Womersley, 1956b: 538.

Cyrtolaelaps semipunctatus.— Womersley, 1961: 194.

Cyrtolaelaps (Gamasellus) semipunctatus.— Ryke, 1962b: 55.

Acugamasus semipunctatus.— Lee, 1970: 142; Lee, 1973: 23; Karg, 1993a: 54; Karg, 1997b:

38.


TYPE LOCALITY AND HABITAT: Australia, Bridgewater, South Australia, August 1942,

on moss.

018. Acugamasus tuberculatus Karg, 1993

Acugamasus tuberculatus Karg, 1993a: 52.

Acugamasus tuberculatus.— Karg, 1997b: 38.



TYPE LOCALITY AND HABITAT: New Caledonia, Koumac, 15 February 1977, in litter in

a cave.

019. Acugamasus watsoni (Hirschmann, 1966)

Gamasellus (Gamasellus) watsoni Hirschmann, 1966c: 29.

Acugamasus watsoni.— Lee, 1970: 142; Lee and Hunter, 1974: 318; Karg, 1993a: 53; Karg,

1997b: 38.


TYPE LOCALITY AND HABITAT: Macquarie Island, [between Australia and Antarctica],

in grassland soil; in litter of Stilbocarpa sp. [Areliaceae], Pleurophyllum sp.

236

[Asteraceae], Colobanthus sp. [Caryophyllaceae], Stellaria media [Caryophyllaceae],

Poa foliosa [Poaceae] and Festuca erecta [Poaceae]; in soil with Cotula plumosa

[Asteraceae].

Genus Acuphis Karg, 1998

Acuphis Karg, 1998: 192 (described in Ologamasidae Ryke).

Acuphis.— Karg, 2006: 163; Karg and Schorlemmer, 2011b: 218.

Type species: Acuphis euarcus Karg, 1998, by original designation.

020. Acuphis euarcus Karg, 1998

Acuphis euarcus Karg, 1998: 193.

Acuphis euarcus.— Karg, 2006: 163; Karg and Schorlemmer, 2011b: 218.



TYPE LOCALITY AND HABITAT: Ecuador, 53 km from Otavalo (alt. 2,850 m), Província

de Imbabura, 20 April 1989, on moss hanging from trees.

021. Acuphis octornatus Karg, 1998

Acuphis octornatus Karg, 1998: 193.

Acuphis octornatus.— Karg, 2006: 163; Karg and Schorlemmer, 2011b: 218.



TYPE LOCALITY AND HABITAT: Ecuador, Rio Guajalito (alt. 1,850 m), Las Palmeras,

Pichincha Province, 18 April 1989, in litter and upper soil of primary forest.

022. Acuphis tetrapennatus Karg, 2006

Acuphis tetrapennatus Karg, 2006: 162.

Acuphis tetrapennatus.— Karg and Schorlemmer, 2011b: 218.



TYPE LOCALITY AND HABITAT: Ecuador, Flavio Alfaro, [Manabi Province], 20 April

1990, in litter in a cacao [Sterculiaceae] plantation.

237

Genus Allogamasellus Athias-Henriot, 1961

Allogamasellus Athias-Henriot, 1961a: 473 (described in Rhodacaridae Oudemans).

Allogamasellus.— Lee, 1970: 144.

Type species: Allogamasellus aquafortensis Athias-Henriot, 1961, by original

designation.

023. Allogamasellus aquafortensis Athias-Henriot, 1961

Allogamasellus aquafortensis Athias-Henriot, 1961a: 475.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, near the bridge of l‘Oued Hamiz, Bordj el

Kiffan [cited as Fort de l‘Eau], [Algiers], 17 February 1958, in a disturbed fragment of

Oleo-Lentiscetum [communities] on sandstone rock.

024. Allogamasellus squalidus Athias-Henriot, 1961

Allogamasellus squalidus Athias-Henriot, 1961a: 475.


Paris, France.

TYPE LOCALITY AND HABITAT: Algeria, Mechta Baiou, Batna, 24 June 1959, on roots

of Phillyrea media [Oleaceae].

Genus Antennolaelaps Womersley, 1956

Antennolaelaps Womersley, 1956a: 112 (described in Neoparasitidae Oudemans).

Antennolaelaps.— Ryke, 1962c: 159; Lee, 1970: 178; Karg, 1993a: 45.

Type species: Antennolaelaps affinis Womersley, 1956, by original designation.

Stylogamasus Womersley, 1956a: 113 [junior synonymy by Lee, 1970: 178].

Type species: Stylogamasus convexa Womersley, 1956, by original designation.

238

025. Antennolaelaps affinis Womersley, 1956

Antennolaelaps affinis Womersley, 1956a: 112.

Antennolaelaps affinis.— Lee, 1970: 183; Karg, 1993a: 48.


TYPE LOCALITY AND HABITAT: Australia, Brookfield, Queensland, 31 May to 10 June

1949, in litter.


026. Antennolaelaps alveolaris Karg, 1993

Antennolaelaps alveolaris Karg, 1993a: 46.



TYPE LOCALITY AND HABITAT: New Caledonia, Île des Pius [= Île des Pins?], 24

February 1977, from unspecified substrate.

027. Antennolaelaps aremenae Lee, 1973

Antennolaelaps aremenae Lee, 1973: 30.

Antennolaelaps aremenae.— Karg, 1993a: 48.


TYPE LOCALITY AND HABITAT: Australia, near the First Waterfall, Waterfall Gully,

Adelaide, South Australia, 24 May 1968, on moss.

028. Antennolaelaps brevisetae Karg, 1996

Antennolaelaps brevisetae Karg, 1996: 172.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, 1977, in humus between

roots.

029. Antennolaelaps celox Lee, 1973

Antennolaelaps celox Lee, 1973: 32.

Antennolaelaps celox.— Karg, 1993a: 48.


239

TYPE LOCALITY AND HABITAT: Australia, near the summit of Mount Lofty (alt. 715 m),

Adelaide, South Australia, 9 May to 4 July 1968, on moss and plant litter.

030. Antennolaelaps convexus (Womersley, 1956)

Stylogamasus convexa Womersley, 1956a: 113.

Antennolaelaps convexus.— Lee, 1970: 183; Karg, 1993a: 47.



1949, in soil debris.

031. Antennolaelaps heterosetae Karg, 1993

Antennolaelaps heterosetae Karg, 1993a: 45.



TYPE LOCALITY AND HABITAT: New Caledonia, E. d‘Amien [=Napué Amien?], near a

house.

032. Antennolaelaps testudo Lee, 1970

Antennolaelaps testudo Lee, 1970: 184.

Antennolaelaps testudo.— Karg, 1993a: 48; Halliday, 2001: 303.


TYPE LOCALITY AND HABITAT: Australia, Brookfield, near Brisbane, Queensland, 7

September 1966, on moss.

NOTE: According to Lee (1970): 184, the specimens reported by Domrow (1957): 202 as

Sessiluncus heterotarsus (Canestrini, 1897) are Antennolaelaps testudo.

Genus Athiasella Lee, 1973

Athiasella Lee, 1973: 10 (described in Rhodacaridae Oudemans).

Athiasella.— Lee and Hunter, 1974: 306; Karg, 1993a: 56.

Type species: Hydrogamasus dentatus Womersley, 1942, by original designation.

Heydeniella - dentata-complex.— Lee, 1970: 101.

240

033. Athiasella australica (Womersley, 1942)

Hydrogamasus australicus Womersley, 1942: 153.

Hydrogamasus australicus.— Womersley, 1956b: 529.

Gamasellus (Hydrogamasellus) australicus.— Hirschmann, 1966b: 7.

Heydeniella australica.— Lee, 1970: 105; Karg, 1976c: 199.

Athiasella australica.— Lee, 1973: 10; Karg, 1993a: 62.


TYPE LOCALITY AND HABITAT: Australia, Brisbane, Queensland, October 1934, on

moss.

034. Athiasella biconi Karg, 1993

Athiasella biconi Karg, 1993a: 57.



TYPE LOCALITY AND HABITAT: New Caledonia, E. d‘Amien [= Napué Amien?], 1977,

in a house.

035. Athiasella caverna Halliday, 2001

Athiasella caverna Halliday, 2001: 303.

TYPE DEPOSITORY: Australian National Insect Collection, Canberra, Australia.

TYPE LOCALITY AND HABITAT: Australia, Caves Reserve, between 3 Dolines Valley

and Wiburds Bluff, Jenolan Caves, New South Wales, 20 April 1993, in decomposing

leaf litter on the ground surface from sites adjacent to cave entrance.

036. Athiasella coniuncta Karg, 1993

Athiasella coniuncta Karg, 1993a: 58.



TYPE LOCALITY AND HABITAT: New Caledonia, Koumac, 15 February 1977, in litter of

a cave.

037. Athiasella dentata (Womersley, 1942)

Hydrogamasus dentatus Womersley, 1942: 149.

Gamasellus (Hydrogamasellus) dentatus.— Hirschmann, 1966b: 7.

241

Heydeniella dentata.— Lee, 1970: 105; Karg, 1976c: 199.

Athiasella dentata.— Lee, 1973: 10; Karg, 1993a: 61.


TYPE LOCALITY AND HABITAT: Australia, Mount Lofty Ranges and Long Gully, South

Australia, August and September 1938, on moss.

038. Athiasella goei (Lee, 1970)

Heydeniella goei Lee, 1970: 108.

Heydeniella goei.— Karg, 1976c: 199.

Athiasella goei.— Lee, 1973: 10; Karg, 1993a: 62.


TYPE LOCALITY AND HABITAT: England, Chislehurst Caves, Kent, 20 February 1955,

under mats of fur-like mould on decaying wooden pit props in chalk-mine.

039. Athiasella hami Karg, 1993

Athiasella hami Karg, 1993a: 60.



TYPE LOCALITY AND HABITAT: New Caledonia, Monts Koghi, 23 February 1977, from


040. Athiasella longiseta Lee and Hunter, 1974

Athiasella longiseta Lee and Hunter, 1974: 308.


TYPE LOCALITY AND HABITAT: New Zealand, Courrejolles Peninsula, Campbell Island,

13 February 1963, on moss and lichen on rocks.

041. Athiasella markmitchelli (Lee, 1970)

Heydeniella markmitchelli Lee, 1970: 109.

Heydeniella markmitchelli.— Karg, 1976c: 199.

Athiasella markmitchelli.— Lee, 1973: 10; Karg, 1993a: 62.


TYPE LOCALITY AND HABITAT: Australia, Flinders Ranges, near Wilmington, South

Australia, 25 September 1958, on moss.

242

042. Athiasella pecten Lee and Hunter, 1974

Athiasella pecten Lee and Hunter, 1974: 310.


TYPE LOCALITY AND HABITAT: New Zealand, Tucker Cove, Campbell Island, 1

February 1963, on Dracophyllum sp. [Ericaceae] leaf mould.

043. Athiasella relata (Womersley, 1942)

Hydrogamasus relatus Womersley, 1942: 151.

Gamasellus (Hydrogamasellus) relatus.— Hirschmann, 1966b: 7.

Heydeniella relata.— Lee, 1970: 105; Karg, 1976c: 199.

Athiasella relata.— Lee, 1973: 11; Karg, 1993a: 62.


TYPE LOCALITY AND HABITAT: Australia, Glen Osmond, South Australia, June and July

1934, on moss.

NOTE: According to Lee (1973): 11, the specimens reported by Womersley (1956b): 530 as

Hydrogamasus relatus Womersley, 1942 were misidentified.

044. Athiasella relicta (Womersley, 1942)

Hydrogamasus relictus Womersley, 1942: 152.

Gamasellus (Hydrogamasellus) relictus.— Hirschmann, 1966b: 7.

Heydeniella relicta.— Lee, 1970: 105; Karg, 1976c: 199.

Athiasella relicta.— Lee, 1973: 10; Karg, 1993a: 61.


TYPE LOCALITY AND HABITAT: Australia, Brisbane, Queensland, October 1934, on

moss; Australia, Adelaide, South Australia, 1935, on moss; Australia, Glen Osmond,

South Australia, 1935, on pine needles.

044a. Athiasella relicta major (Womersley, 1942)

Hydrogamasus relictus var. major Womersley, 1942: 153.

Heydeniella relicta var. major.— Lee, 1970: 105.

Athiasella relicta var. major.— Lee, 1973: 10.


243

TYPE LOCALITY AND HABITAT: Australia, Sassafras, Victoria, December 1931,

on moss; New Zealand, Bourke‘s Bush, Waimamaku, Auckland, October


045. Athiasella scaphosternum Lee and Hunter, 1974

Athiasella scaphosternum Lee and Hunter, 1974: 310.


TYPE LOCALITY AND HABITAT: New Zealand, Saint Col Peak (alt. 250 m), Campbell

Island, 23 February 1963, on tussock base.

046. Athiasella sellaris Karg, 1996

Athiasella sellaris Karg, 1996: 186.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, January 1977, in litter

accumulated in a rock fissure.

047. Athiasella stefani Halliday, 2001

Athiasella stefani Halliday, 2001: 303.


TYPE LOCALITY AND HABITAT: Australia, Caves Reserve, between 3 Dolines Valley

and Wiburds Bluff, Jenolan Caves, New South Wales, 20 April 1993, in decomposing

leaf litter on the ground surface from sites adjacent to cave entrance.

048. Athiasella tridentata (Karg, 1976)

Heydeniella tridentata Karg, 1976c: 193.

Athiasella tridentata.— Karg, 1993a: 61.


TYPE LOCALITY AND HABITAT: Chile, Collipulli, Malleco Province, 27 October 1965,

under stones.

049. Athiasella tuberculata Karg, 1993

Athiasella tuberculata Karg, 1993a: 60.

244




a cave.

050. Athiasella viripileus Lee and Hunter, 1974

Athiasella viripileus Lee and Hunter, 1974: 308.


TYPE LOCALITY AND HABITAT: New Zealand, summit of Mount Dumas (alt. 500 m),

Campbell Islands, 6 February 1963, under stones.

Genus Caliphis Lee, 1970

Caliphis Lee, 1970: 52 (described in Rhodacaridae Oudemans).

Type species: Caliphis calvus Lee, 1970, by original designation.

051. Caliphis calvus Lee, 1970

Caliphis calvus Lee, 1970: 53.


TYPE LOCALITY AND HABITAT: Australia, Lena Valley Track, Mount Wellington, near

Hobart, Tasmania, 14 December 1966, on moss.

052. Caliphis eugenitalis Karg, 1993

Caliphis eugenitalis Karg, 1993a: 56.



TYPE LOCALITY AND HABITAT: New Caledonia, Poindimié, 30 January 1977, in litter,

on moss and between roots on rocks.

053. Caliphis hickmani (Womersley, 1956)

Gamasiphis (Neogamasiphis) hickmani Womersley, 1956b: 519.

Caliphis hickmani.— Lee, 1970: 53.


245

TYPE LOCALITY AND HABITAT: Australia, Mount Wellington (alt. 3,000 feet),

Tasmania, 27 July 1943, from unspecified substrate.

054. Caliphis minisetae (Karg, 1993)

Gamasiphis minisetae Karg, 1993b: 173

Gamasiphis minisetae.— Karg, 1995: 16.





055. Caliphis novaezelandiae (Womersley, 1956)

Gamasiphis (Neogamasiphis) novae-zelandiae Womersley, 1956b: 524.

Caliphis novaezelandiae.— Lee, 1970: 53.


TYPE LOCALITY AND HABITAT: New Zealand, Beachlands, North Island, March 1949,

from unspecified substrate.

056. Caliphis queenslandicus (Womersley, 1956)

Gamasiphis (Neogamasiphis) queenslandicus Womersley, 1956b: 522.

Caliphis queenslandicus.— Lee, 1970: 53.


TYPE LOCALITY AND HABITAT: Australia, Taringa, Brisbane, Queensland, 29 to 31

January 1949, from unspecified substrate.

057. Caliphis schusteri (Hirschmann, 1966)

Gamasellus (Hydrogamasellus) schusteri Hirschmann, 1966c: 42.

Caliphis schusteri.— Lee, 1970: 53; Lee and Hunter, 1974: 299.


TYPE LOCALITY AND HABITAT: Macquarie Island, [between Australia and Antarctica],

in litter of Stilbocarpa sp. [Areliaceae], Cotula sp. [Asteraceae], Pleurophyllum sp.

[Asteraceae], Colobanthus sp. [Caryophyllaceae], Poa foliosa [Poaceae] and Festuca

erecta [Poaceae]; on moss; in grassland soil; on rocks.

246

058. Caliphis tamborinensis (Womersley, 1956)

Gamasiphis (Neogamasiphis) hickmani var. tamborinensis Womersley, 1956b: 521.

Caliphis tamborinensis.— Lee, 1970: 53.


TYPE LOCALITY AND HABITAT: Australia, Mount Tamborine, Queensland, 14 May

1952, on card placed on the ground in the scrub.

Cymiphis Lee, 1970

Cymiphis Lee, 1970: 90 (described in Rhodacaridae Oudemans).

Type species: Ologamasus cymosus Lee, 1966, by original designation.

Ologamasus - cymosus-group.— Lee, 1966: 211.

059. Cymiphis cymosus (Lee, 1966)

Ologamasus cymosus Lee, 1966: 212.

Cymiphis cymosus.— Lee, 1970: 91.


TYPE LOCALITY AND HABITAT: New Zealand, Botanical Gardens, Wellington, North

Island, December 1960, on leaf mould.

060. Cymiphis dumosus (Lee, 1966)

Ologamasus dumosus Lee, 1966: 219.

Cymiphis dumosus.— Lee, 1970: 91.




061. Cymiphis leptosceles (Lee, 1966)

Ologamasus leptosceles Lee, 1966: 217.

Cymiphis leptosceles.— Lee, 1970: 91.


TYPE LOCALITY AND HABITAT: New Zealand, Khandallah, Wellington, North Island,

19 December 1961, on leaf mould.

247

062. Cymiphis mansoni (Lee, 1966)

Ologamasus mansoni Lee, 1966: 213.

Cymiphis mansoni.— Lee, 1970: 92.




063. Cymiphis nucilis (Lee, 1966)

Ologamasus nucilis Lee, 1966: 220.

Cymiphis nucilis.— Lee, 1970: 92.




064. Cymiphis validus (Lee, 1966)

Ologamasus validus Lee, 1966: 222.

Cymiphis validus.— Lee, 1970: 92.


TYPE LOCALITY AND HABITAT: New Zealand, Waimamaku, North Island, 21 October



065. Cymiphis watsoni (Hirschmann, 1966)

Gamasiphis watsoni Hirschmann, 1966c: 41.

Cymiphis watsoni.— Lee, 1970: 92; Lee and Hunter, 1974: 312.



in Garden Cove, First Gully, Gadget Gully and Nugget Point, in litter of Stilbocarpa

sp. [Areliaceae].

248

Cyrtolaelaps Berlese, 1887

Cyrtolaelaps Berlese, 1887b: 5 [presumed to be described in Parasitidae Oudemans (also

referred to as Gamasidae Leach)].

Cyrtolaelaps.— Berlese, 1892b: 3; Berlese, 1892f: 60; Berlese, 1913b: 86; Vitzthum, 1931a:

142; Vitzthum, 1943: 758; Baker and Wharton, 1952: 64; Lee, 1970: 146; Karg, 1971:

349; Bregetova and Shcherbak, 1977a: 302; Karg, 1993c: 370.

Cyrtolaelaps (Cyrtalaelaps).— Ryke, 1962b: 4; Ryke, 1962c: 156.

Type species: Gamasus mucronatus G. Canestrini and R. Canestrini, 1881, by

subsequent monotypy.

Protolaelaps Trägårdh, 1912: 563 (described in Parasitidae Oudemans) [junior synonymy by

Vitzthum, 1931a: 142].

Type species: Gamasellus (?) brevispinosus Trägårdh, 1910, by original designation.

066. Cyrtolaelaps aster (Berlese, 1918)

Gamasellus (Protolaelaps) aster Berlese, 1918: 137.

Gamasellus (Protolaelaps) aster.— Schweizer, 1922: 35.

Cyrtolaelaps (Cyrtolaelaps) aster.— Ryke, 1962b: 9.

Cyrtolaelaps aster.— Lee, 1970: 150; Karg, 1971: 350; Bregetova and Shcherbak, 1977a:

305; Karg, 1993c: 370.


TYPE LOCALITY AND HABITAT: Italy, Asuni, Sardinia, in nest of Microtus agrestis

(cited as Murium agrestium) [Mammalia: Rodentia: Cricetidae].

NOTE: Specimens reported by Schweizer (1961): 91 as Cyrtolaelaps aster (Berlese, 1918)

were described as Cyrtolaelaps chiropterae Karg, 1971: 349.

067. Cyrtolaelaps berlesei Chelebiev, 1984

Cyrtolaelaps berlessei [sic] Chelebiev, 1984: 140.

Cyrtolaelaps berlesei.— Chelebiev, 1984: 141.


Petersburg, Russia.

249

TYPE LOCALITY AND HABITAT: Kazakhstan, Karaganda, Karagandy Province, 4 July

1984, in the nest of Spermophilus erythrogenys (cited as Citellus erythrogenys)

[Mammalia: Rodentia: Sciuridae].


068. Cyrtolaelaps chiropterae Karg, 1971

Cyrtolaelaps chiropterae Karg, 1971: 349.

Cyrtolaelaps chiropterae.— Bregetova and Shcherbak, 1977a: 304; Chelebiev, 1984: 140;

Karg, 1993c: 370.


TYPE LOCALITY AND HABITAT: Switzerland, Grotte de chemin de fer, Gorges de

l´Areuse, Canton of Neuchâtel, in bat detritus [Animalia: Chiroptera].

NOTE: Described on the basis of specimens reported by Schweizer (1961): 91 as

Cyrtolaelaps aster (Berlese, 1918).

069. Cyrtolaelaps kasakstanicus (Chelebiev, 1978)

Euryparasitus kasakstanicus Chelebiev, 1978: 1273.

Euparasitus [sic] kasakstanicus.— Chelebiev, 1978: 1276.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Kazakhstan, Saran, 14 April 1973, in nest of Lagurus

lagurus [Mammalia: Rodentia: Cricetidae].

070. Cyrtolaelaps minor Willmann, 1952

Cyrtolaelaps minor Willmann, 1952: 422.

Cyrtolaelaps (Cyrtolaelaps) minor.— Ryke, 1962b: 10.

Cyrtolaelaps minor.— Lee, 1970: 150; Karg, 1971: 350; Bregetova and Shcherbak, 1977a:

305; Chelebiev, 1984: 141; Karg, 1993c: 370.


TYPE LOCALITY AND HABITAT: Germany, Hamburg, 1 January 1937, in nest of

Apodemus flavicollis [Mammalia: Rodentia: Muridae].

071. Cyrtolaelaps mucronatus (G. Canestrini and R. Canestrini, 1881)

Gamasus mucronatus G. Canestrini and R. Canestrini, 1881: 1081.

250

Gamasus mucronatus.— G. Canestrini and R. Canestrini, 1882: 54; G. Canestrini, 1885: 78.

Cyrtolaelaps mucronatus.— Berlese, 1887b: 5; Berlese, 1892f: 60; Trägårdh, 1912: 566;

Willmann, 1935: 7; Willmann, 1941: 20; Willmann, 1952: 422; Schweizer, 1961: 90;

Lee, 1970: 150; Karg, 1971: 349; Bregetova and Shcherbak, 1977a: 305; Chelebiev,

1984: 141; Karg, 1993c: 370.

? Cyrtolaelaps mucronatus [sic].— Berlese, 1892f: 61.

Gamasellus (Protolaelaps) mucronatus.— Berlese, 1918: 188; Halbert, 1923: 365; Vitzthum,

1926a: 7.

Gamasellus (Protolaelaps) murcronatus [sic].— Schweizer, 1922: 35.

Cyrtolaelaps (Protolaelaps) mucronatus.— Sellnick, 1940: 26.

Cyrtolaelaps (Cyrtolaelaps) mucronatus.— Ryke, 1962b: 5; Ryke, 1962c: 157.

Asca affinis Oudemans, 1902a: 45 [junior synonymy by Wilmann, 1935: 8].

Asca affinis.— Oudemans, 1904: 92; Sellnick, 1940: 26.

Gamasellus (?) brevispinosus [sic] Trägårdh, 1910: 422 [suspected junior synonymy by

Trägårdh, 1912: 566; junior synonymy by Berlese, 1918: 188].

Protolaelaps brevispinosus.— Trägårdh, 1912: 566.

Gamasellus alienus Hull, 1918: 78 [junior synonymy apud Ryke, 1962b: 6].

Gamasellus rubicundus Hull, 1918: 78 [junior synonymy apud Ryke, 1962b: 5].

Gamasellus major Pinchuk, 1972b: 67 [junior synonymy by Bregetova and Shcherbak,

1977a: 305].

TYPE DEPOSITORY: C. mucronatus: Istituto Sperimentale per la Zoologia Agraria,

Florence, Italy; A. affinis: National Museum of Natural History – Naturalis, Leiden,

Netherlands; G. brevispinosus: Naturhistorika Riksmuseum, Stockholm, Sweden; G.

alienus: British Museum (Natural History) London, England; G. rubicundus: British

Museum (Natural History) London, England; G. major: Institute of Zoology and

Parasitology SSR Moldova, Chişinău, Moldova.

TYPE LOCALITY AND HABITAT: C. mucronatus: Italy, Padua, 23 March 1881, on

equine´s feces; A. affinis: Netherlands, Utrecht, in humus and on Mustela vulgaris and

Mustela putorius [Mammalia: Carnivora: Mustelidae]; G. brevispinosus: Sweden,

Säkok, 12 July 1907, in nest of lemmings [Mammalia: Rodentia: Cricetidae]; G.

alienus: England, Allendale [cited as West Allendale], Tynedale [cited as Tyne

Province], in nests of moles [Mammalia: Talpidae] and on moss; G. rubicundus:

England, Ninebanks, [Northumberland], under a dead fowl; G. major: Moldova,

251

Chişinău, 20 April 1969, in nest of Apodemus agrarius [Mammalia: Rodentia:

Muridae].

072. Cyrtolaelaps qinghaiensis Ma, 1988

Cyrtolaelaps qinghaiensis Ma, 1988: 149.

TYPE DEPOSITORY: First Institute of Endemic Diseases Research, Jilin Province, China.

TYPE LOCALITY AND HABITAT: China, Heka (35°54‘N, 100°00‘E), Xinghai, Qinghai

Province, August 1961, in nest of Marmota himalayana [Mammalia: Rodentia:

Sciuridae].

073. Cyrtolaelaps rectus (Berlese, 1920)

Gamasellus (Protolaelaps) rectus Berlese, 1920b: 163.

Cyrtolaelaps (Cyrtolaelaps) rectus.— Ryke, 1962b: 9.

Cyrtolaelaps rectus.— Lee, 1970: 151.


TYPE LOCALITY AND HABITAT: USA, Columbia, Missouri, 1904-1906, in humus.

074. Cyrtolaelaps spurius (Holzmann, 1969)

Euryparasitus spurius Holzmann, 1969: 51.

Cyrtolaelaps spurius.— Lee, 1970: 150.


TYPE LOCALITY AND HABITAT: Central Europe, in nest of moles [Mammalia: Talpidae].


075. Cyrtolaelaps subnudus (Berlese, 1918)

Gamasellus (Protolaelaps) subnudus Berlese, 1918: 138.

Gamasellus (Protolaelaps) subnudus? [sic].— Halbert, 1923: 365.

Cyrtolaelaps (Cyrtolaelaps) subnudus.— Ryke, 1962b: 11.


TYPE LOCALITY AND HABITAT: Italy, Florence, in poultry [Animalia: Aves] manure.

252

Genus Desectophis Karg, 2003

Desectophis Karg, 2003: 238 (described in Gamasiphidae Lee).

Desectophis.— Karg and Schorlemmer, 2011b: 217.

Type species: Desectophis magnosimilis Karg, 2003, by original designation.

076. Desectophis eulateris (Karg, 1998)

Acuphis eulateris Karg, 1998: 193.

Acuphis eulateris.— Karg, 2006: 163.

Desectophis eulateris.— Karg and Schorlemmer, 2011b: 218.



TYPE LOCALITY AND HABITAT: Ecuador, between Pifo and Papallacta (alt. 4,100 m),

Pichincha Province, 14 April 1989, on moss from the soil beside water-course.

077. Desectophis flagellatus Karg and Schorlemmer, 2011

Desectophis flagellatus Karg and Schorlemmer, 2011b: 217.



TYPE LOCALITY AND HABITAT: Ecuador, Volcán Guagua Pichincha [cited as Aguagua

volcano], Pichincha Province, 1987, in soil and plant detritus from a depression

temporarily filled with water.

078. Desectophis magnosimilis Karg, 2003

Desectophis magnosimilis Karg, 2003: 239.

Desectophis magnosimilis.— Karg and Schorlemmer, 2011b: 218.



TYPE LOCALITY AND HABITAT: Ecuador, Imbabura Province, 1989, in litter and moss.

079. Desectophis pulcher Karg, 2003

Desectophis pulcher Karg, 2003: 241.

Desectophis pulcher.— Karg and Schorlemmer, 2011b: 218.

253



TYPE LOCALITY AND HABITAT: Ecuador, Tenatol (= Tena?), 1990, in litter.

Genus Euepicrius Womersley, 1942

Euepicrius Womersley, 1942: 170 (described in Macrochelidae Vitzthum).

Euepicrius.— Womersley, 1956a: 104; Lee, 1966: 206; Lee, 1970: 55; Karg, 1993a: 42; Karg,

1997a: 80.

Type species: Euepicrius filamentosus Womersley, 1942, by original designation.

080. Euepicrius bipeltatus Karg, 1997

Euepicrius bipeltatus Karg, 1997a: 82.




moss.

081. Euepicrius brevicruris Karg, 1993

Euepicrius brevicruris Karg, 1993a: 42.

Euepicrius brevicruris.— Karg, 1997a: 82.



TYPE LOCALITY AND HABITAT: New Caledonia, Mont Mandjelia (alt. 650 m), February

1977, in soil of a deciduous forest.

082. Euepicrius caesariatus Lee and Hunter, 1974

Euepicrius caesariatus Lee and Hunter, 1974: 299.


TYPE LOCALITY AND HABITAT: New Zealand, Ranui Island, Auckland Islands, 3

January 1963, on Metrosideros [Myrtaceae] leaf mould.

083. Euepicrius femuralis Karg, 1993

254

Euepicrius femuralis Karg, 1993a: 44.

Euepicrius femuralis.— Karg, 1997a: 82.



TYPE LOCALITY AND HABITAT: New Caledonia, E. d‘Amien [= Napué Amien?], near a

house.

084. Euepicrius filamentosus Womersley, 1942

Euepicrius filamentosus Womersley, 1942: 170.

Euepicrius filamentosus.— Lee, 1973: 8; Karg, 1993a: 44; Karg, 1997a: 81.


TYPE LOCALITY AND HABITAT: Australia, Glen Osmond, South Australia, June 1933

and July 1935, on moss; Australia, Long Gully, South Australia, August 1938, on

moss.

NOTE: According to Lee (1973): 8, an adult female of an undescribed species from

Waimamaku, New Zealand, was incorrectly identified as E. filamentosus in the

original description

085. Euepicrius lootsi Lee, 1970

Euepicrius lootsi Lee, 1970: 57.

Euepicrius lootsi.— Karg, 1993a: 45; Karg, 1997a: 81.


TYPE LOCALITY AND HABITAT: Australia, beside Wannon River, near Yarram Gap,

Grampians, Victoria, 14 May 1966, on moss and grass.

086. Euepicrius multipori Karg, 1993

Euepicrius multipori Karg, 1993a: 42.

Euepicrius multipori.— Karg, 1997a: 82.



TYPE LOCALITY AND HABITAT: New Caledonia, E. d‘Amien [= Napué Amien?], in a

house.

087. Euepicrius queenslandicus Womersley, 1956

255

Euepicrius queenslandicus Womersley, 1956a: 105.

Euepicrius queenslandicus.— Karg, 1993a: 44; Karg, 1997a: 81.


TYPE LOCALITY AND HABITAT: Australia, Brookfield, Brisbane, Queensland, between

May and July 1949, in litter.

Genus Euryparasitus Oudemans, 1902

Euryparasitus Oudemans, 1902a: 30 (described in Parasitidae Oudemans).

Eurylaelaps.— Oudemans, 1902a: 8 [lapsus for Euryparasitus Oudemans, 1902].

Euryparasitus.— Vitzthum, 1931a: 142; Vitzthum, 1943: 757; Baker and Wharton, 1952: 71;

Lee, 1970: 151; Bregetova and Shcherbak, 1977a: 305; Hagele et al., 2005: 3.

Cyrtolaelaps (Euryparasitus).— Ryke, 1962c: 157; Ryke, 1962a: 112.

Type species: Gamasus terribilis Michael, 1886, by subsequent monotypy (junior

synonymy of Gamasus emarginatus Koch, 1839).

088. Euryparasitus calcarator (Banks, 1910)

Gamasus calcarator Banks, 1910: 4.

Parasitus calcarator.— Banks, 1915: 83.

Euryparasitus calcarator.— Lee, 1970: 154; Hennessey and Farrier, 1988: 16; Hagele et al.,

2005: 14.

TYPE DEPOSITORY: Museum of Comparative Zoology, Harvard University, Cambridge,

USA

TYPE LOCALITY AND HABITAT: USA, Falls Church, Virginia, November 1909, in nest

of field mouse [Mammalia: Rodentia].

089. Euryparasitus changanensis Gu and Huang, 1992

Euryparasitus changanensis Gu and Huang, 1992: 202.

TYPE DEPOSITORY: Sanitation and Anti-Epidemic Station, Linyi, Shaanxi Province,

China.

TYPE LOCALITY AND HABITAT: China, Changan (34º16‘N, 108º89‘E), Shaanxi

Province, 10 October 1982, in nest of Cricetus triton [Mammalia: Rodentia:

Cricetidae].

256

090. Euryparasitus citelli Bai, Chen and Gu, 1988

Euryparasitus citelli Bai, Chen and Gu, 1988: 369.

TYPE DEPOSITORY: Institute of Endemic Disease Control, Yinchuan, Ningxia Hui

Autonomous Region, China.

TYPE LOCALITY AND HABITAT: China, Haiyuan (36.5°N, 105.6°E), Ningxia Hui

Autonomous Region, April 1985, in nest of Spermophilus alaschanicus (cited as

Citellus alaschanicus) [Mammalia: Rodentia: Sciuridae].

091. Euryparasitus davydovae Bondarchuk and Buyakova, 1978

Euryparasitus davydovae Bondarchuk and Buyakova, 1978: 1576.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Kalarsky District, Chita Region, Siberia, 9 July

1962, in nest of Clethrionomys rutilus [Mammalia: Rodentia: Muridae].

092. Euryparasitus emarginatus (Koch, 1839)

Gamasus emarginatus Koch, 1839a: 17.

Poecilochirus emarginatus.— Berlese, 1892d: 4.

Parasitus emarginatus.— Oudemans, 1904: 75.

Gamasus (?) emarginatus [sic].— Berlese, 1906: 266.

Gamasus (Gamasus?) emarginatus [sic].— Berlese, 1906: 279.

Euryparasitus emarginatus.— Oudemans, 1913a: 165; Oudemans, 1936: 195; Willmann,

1941: 20; Lee, 1970: 154; Davydova, 1970: 88; Zuevsky, 1971: 1407; Karg, 1971:

351; Bregetova and Shcherbak, 1977a: 307; Karg, 1993c: 371.

Cyrtolaelaps (Euryparasitus) emarginatus.— Ryke, 1962c: 157; Ryke, 1962a: 113.

Gamasus setiger Koch, 1839b: 2 [junior synonymy apud Oudemans 1936: 196].

Gamasus terribilis Michael, 1886: 265 [junior synonymy by Oudemans, 1913a: 165].

Gamasus (Eugamasus) horribilis [sic].— Berlese, 1892f: 62.

Eurylaelaps [sic] terribilis.— Oudemans, 1902a: 8.

Euryparasitus terribilis.— Oudemans, 1902a: 30; Oudemans, 1904: 83; Oudemans, 1913b: 4.

TYPE DEPOSITORY: E. emarginatus: Unknown.; Gamasus setiger: Unknown.; G.

terribilis: British Museum (Natural History), London, England.

257

TYPE LOCALITY AND HABITAT: E. emarginatus: Germany, Frauenholz, near

Regensburg, on moss on forest soil; Gamasus setiger: Germany, Frauenholz, near

Regensburg, in wet forest soil; G. terribilis: England, December 1885, in mole nest

[Mammalia].

NOTE1: Specimens reported by Trägårdh (1912): 549 as Euryparasitus terribilis (Michael,

1886) were described as Euryparasitus tragardhi Bregetova, in Bregetova and

Shcherbak (1977a): 307.

NOTE2: Species reported by Koch (1879): 119 as Gamasus emarginatus (Koch, 1839) was

renamed by Trägård, (1901): 61 as Cyrtolaelaps kochi, presently placed in Veigaia

(Veigaiidae).

093. Euryparasitus goncharovi Bondarchuk and Buyakova, 1976

Euryparasitus goncharovi Bondarchuk and Buyakova, 1976: 927.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Kalarsky District, Chita Region, Siberia, 8

September 1964, in a rodent nest [Mammalia: Rodentia].

094. Euryparasitus laxiventralis Gu and Guo, 1995

Euryparasitus laxiventralis Gu and Guo, 1995: 179.

TYPE DEPOSITORY: Department of Parasitology, Medical School, Nanjing University,

Nanjing, China.

TYPE LOCALITY AND HABITAT: China, Sinan, Guizhou Province, November 1988, off

Rattus norvegicus [Mammalia: Rodentia: Muridae].

095. Euryparasitus longicheta Bondarchuk and Buyakova, 1978

Euryparasitus longicheta Bondarchuk and Buyakova, 1978: 1578.

Euryparasitus longicheta.— Hagele et al., 2005: 10.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Kalarsky District, Chita Region, Siberia, 26

October 1960, associated with Eutamias sibericus [Mammalia: Rodentia: Sciuridae].


258

096. Euryparasitus maseri Whitaker and Klompen, 2005

Euryparasitus maseri Whitaker and Klompen, in Hagele et al., 2005: 12.

TYPE DEPOSITORY: Acarology Laboratory at Ohio State University, Columbus, Ohio,

USA.

TYPE LOCALITY AND HABITAT: USA, 10 km southeast of Whitehorse Ranch (43°15'N

117°40'W), Malheur, Oregon, 19 August 1976, on Onychomys leucogaster

[Mammalia: Rodentia: Cricetidae].


097. Euryparasitus medius Zuevsky, 1971

Euryparasitus medius Zuevsky, 1971: 1406.

Euryparasitus medius.— Bregetova and Shcherbak, 1977a: 307.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Nizhnevartovsk, Khanty-Mansi Autonomous

Okrug, 16 September 1965, on Clethrionomys rutilus [Mammalia: Rodentia:

Muridae].


098. Euryparasitus occidentalis Hagele, Kaufman, Whitaker and Klompen, 2005

Euryparasitus occidentalis Hagele, Kaufman, Whitaker and Klompen, 2005: 5.

TYPE DEPOSITORY: Acarology Laboratory at Ohio State University, Columbus, Ohio,

USA.

TYPE LOCALITY AND HABITAT: Canada, 11.5 miles southwest of Hope (49°23'N,

121°26'W), British Columbia, 8 July 1973, on Neurotrichus gibbsii [Mammalia:

Talpidae].


099. Euryparasitus pagumae Ishikawa, 1988

Euryparasitus pagumae Ishikawa, 1988: 14.

TYPE DEPOSITORY: Department of Zoology, National Science Museum (Natural History),

Tokyo, Japan.

259

TYPE LOCALITY AND HABITAT: Japan, Saredani, Nakayama, Lyo District, Ehime

Prefecture, Island of Shikoku, 9 December 1981, from mine adits of a nest of Paguma

larvata [Mammalia: Carnívora: Viverridae].

100. Euryparasitus taojiangensis Ma, 1982

Euryparasitus taojiangensis Ma, 1982: 115.

TYPE DEPOSITORY: First Institute of Endemic Diseases Research, Jilin Province, China.

TYPE LOCALITY AND HABITAT: China, Gansu Province, 5 June 1960, on Microtus sp.

[Mammalia: Rodentia: Cricetidae].

101. Euryparasitus tori Davydova, 1970

Euryparasitus tori Davydova, 1970: 87.

Euryparasitus tori.— Zuevsky, 1971: 1407; Bregetova and Shcherbak, 1977a: 307.

TYPE DEPOSITORY: Siberian Zoological Museum, Novosibirsk, Russia.

TYPE LOCALITY AND HABITAT: Russia, Lake Teletskoye, vicinity of Mount Kolushtu,

[Altai Mountains], July 1963, on red vole [Mammalia: Rodentia: Cricetidae].


102. Euryparasitus tragardhi Bregetova, 1977

Euryparasitus tragardhi Bregetova, in Bregetova and Shcherbak, 1977a: 307.


TYPE LOCALITY AND HABITAT: France, Pietralbello Cave, Ponte-Leccia, Morosaglia,

Corsica, 9 January 1907, from unspecified substrate.

NOTE1: Described from the deutonymph and adult male.

NOTE2: Described on the basis of specimens reported by Trägårdh (1912): 549 as

Euryparasitus terribilis (Michael, 1886).

Genus Evanssellus Ryke, 1961

Evanssellus Ryke, 1961a: 245 (described in Rhodacaridae Oudemans).

Heterogamasus (Evanssellus).— Lee, 1967: 505.

Evanssellus.— Ryke, 1962c: 157; Lee, 1970: 154.

Type species: Evanssellus foliatus Ryke, 1961, by original designation.

260

103. Evanssellus foliatus Ryke, 1961

Evanssellus foliatus Ryke, 1961a: 245.

Heterogamasus (Evanssellus) foliatus.— Lee, 1967: 510.

Evanssellus foliatus.— Lee, 1970: 155; Lee and Hunter, 1974: 320.

TYPE DEPOSITORY: Arachnida Section of the British Museum (Natural History), London,

England.

TYPE LOCALITY AND HABITAT: New Zealand, Queenstown, July 1954, in beech litter.

104. Evanssellus medusa (Lee, 1967)

Heterogamasus (Evanssellus) medusa Lee, 1967: 510.

Evanssellus medusa.— Lee, 1970: 155.


TYPE LOCALITY AND HABITAT: Australia, Hordern Vale, Cape Otway, Victoria, 28

August 1965, on moss and litter among tree ferns [Pteridophyta] and Eucalyptus

[Myrtaceae].

Genus Gamasellevans Loots and Ryke, 1967

Gamasellevans Loots and Ryke, 1967a: 212 (described in Rhodacaridae Oudemans).

Gamasellevans.— Lee, 1970: 184.

Type species: Gamasellevans epigynialis Loots and Ryke, 1967, by original

designation.

105. Gamasellevans bispermadactylus Loots and Ryke, 1967

Gamasellevans bispermadactylus Loots and Ryke, 1967a: 229.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, 1963, from forest floor

of the evergreen indigenous montane forest.

106. Gamasellevans epigynialis Loots and Ryke, 1967

Gamasellevans epigynialis Loots and Ryke, 1967a: 213.

261



TYPE LOCALITY AND HABITAT: South Africa, Agricultural Research Institute,

Potchefstroom, between September 1962 and September 1963, in pasture soil.

107. Gamasellevans evansi Loots and Ryke, 1967

Gamasellevans evansi Loots and Ryke, 1967a: 223.





108. Gamasellevans magoebaensis Loots and Ryke, 1967

Gamasellevans magoebaensis Loots and Ryke, 1967a: 234.





109. Gamasellevans reticulatus Loots and Ryke, 1967

Gamasellevans reticulatus Loots and Ryke, 1967a: 237.





110. Gamasellevans spermadactylus Loots and Ryke, 1967

Gamasellevans spermadactylus Loots and Ryke, 1967a: 219.



TYPE LOCALITY AND HABITAT: South Africa, Agricultural Research Institute,

Potchefstroom, between September 1962 and September 1963, in pasture soil.

111. Gamasellevans vandenbergi Loots and Ryke, 1967

262

Gamasellevans vandenbergi Loots and Ryke, 1967a: 232.





Genus Gamaselliphis Ryke, 1961

Cyrtolaelaps (Gamaselliphis) Ryke, 1961b: 99 (described in Rhodacaridae Oudemans).

Cyrtolaelaps (Gamaselliphis).— Ryke, 1962c: 157.

Gamaselliphis.— Lee, 1970: 57.

Type species: Cyrtolaelaps (Gamaselliphis) potchefstroomensis Ryke, 1961, by


112. Gamaselliphis cathkini (Ryke, 1961)

Cyrtolaelaps (Gamaselliphis) cathkini Ryke, 1961b: 107.

Gamaselliphis cathkini.— Lee, 1970: 59.


TYPE LOCALITY AND HABITAT: South Africa, Cathkin Peak, [Drakensberg], September

1943, on forest floor.

113. Gamaselliphis grahamstowni (Ryke, 1961)

Cyrtolaelaps (Gamaselliphis) grahamstowni Ryke, 1961b: 105.

Gamaselliphis grahamstowni.— Lee, 1970: 59.


TYPE LOCALITY AND HABITAT: South Africa, Grahamstown, October 1943, in humus.

114. Gamaselliphis lawrencei (Ryke, 1961)

Cyrtolaelaps (Gamaselliphis) lawrencei Ryke, 1961b: 107.

Gamaselliphis lawrencei.— Lee, 1970: 59.


TYPE LOCALITY AND HABITAT: South Africa, High Flats, South Coast, KwaZulu-Natal,

September 1943, from unspecified substrate.

263

115. Gamaselliphis montanellus (Ryke, 1961)

Cyrtolaelaps (Gamaselliphis) montanellus Ryke, 1961b: 103.

Gamaselliphis montanellus.— Lee, 1970: 59.


TYPE LOCALITY AND HABITAT: South Africa, Cathkin Peak, [Drakensberg], September


116. Gamaselliphis potchefstroomensis (Ryke, 1961)

Cyrtolaelaps (Gamaselliphis) potchefstroomensis Ryke, 1961b: 101.

Cyrtolaelaps (Gamaselliphis) potchefstroomensis.— Ryke, 1962c: 157.

Gamaselliphis potchefstroomensis.— Lee, 1970: 59; Halliday, 2005: 41.



TYPE LOCALITY AND HABITAT: South Africa, Potchefstroom, August 1952 and January

1959, in humus.

Genus Gamasellopsis Loots and Ryke, 1966

Gamasellopsis Loots and Ryke, 1966: 551 (described in Rhodacaridae Oudemans).

Gamasellopsis.— Lee, 1970: 186.

Type species: Gamasellopsis curtipilus Loots and Ryke, 1966, by original designation.

117. Gamasellopsis curtipilus Loots and Ryke, 1966

Gamasellopsis curtipilus Loots and Ryke, 1966: 552.

Gamasellopsis curtipilus.— Hirschmann, 1968: 22.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, 1963, in soil of the

indigenous evergreen forest.

118. Gamasellopsis longipilus Loots and Ryke, 1966

Gamasellopsis longipilus Loots and Ryke, 1966: 558.

264



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, 1963, in soil of the

indigenous evergreen forest.

119. Gamasellopsis magoebaensis Loots and Ryke, 1966

Gamasellopsis magoebaensis Loots and Ryke, 1966: 559.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, 1963, in the forest floor.

120. Gamasellopsis vandenbergi Loots and Ryke, 1966

Gamasellopsis vandenbergi Loots and Ryke, 1966: 562.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, 1963, in the forest floor.

Genus Gamasellus Berlese, 1892

Cyrtolaelaps (Gamasellus) Berlese, 1892f: 61 [presumed to be described in Parasitidae


Gamasellus.— Berlese, 1906: 101; Berlese, 1917: 5; Vitzthum, 1931a: 142; Vitzthum, 1943:

758; Baker and Wharton, 1952: 64; Lee, 1970: 129; Karg, 1971: 350; Bregetova and

Shcherbak, 1977a: 299; Davydova, 1982: 5; Karg, 1993c: 370.

Cyrtolaelaps (Gamasellus).— Ryke, 1962b: 14; Ryke, 1962c: 157.

Type species: Gamasus falciger G. Canestrini and R. Canestrini, 1881, by subsequent

designation by Berlese, 1906: 101.

Gamasellus (Eurysellus) Davydova, 1982: 32.

Type species: Gamasellus silvestris Halašková, 1958, by original designation.

Gamasellus (Brevisellus) Davydova, 1982: 60.

Type species: Gamasellus vibrissatus Emberson, 1967, by original designation.

265

121. Gamasellus acutus Karg, 1997

Gamasellus acutus Karg, 1997a: 79.



TYPE LOCALITY AND HABITAT: Chile, Azapa, 2 December 1965, in soil trap in a dry

shrubbery.

122. Gamasellus alexandrovae Davydova, 1982

Gamasellus (Brevisellus) alexandrovae Davydova, 1982: 78.


TYPE LOCALITY AND HABITAT: Russia, Molokovka, Chita Oblast, Siberia, 28 May

1975, in litter of a larch-birch forest.

123. Gamasellus alpinus Schweizer, 1949

Gamasellus falciger var. alpinus Schweizer, 1949: 34.

Gamasellus alpinus.— Schweizer, 1961: 93; Karg, 1971: 350; Bregetova and Shcherbak,

1977a: 301; Karg, 1993c: 371.

Cyrtolaelaps (Gamasellus) alpinus.— Ryke, 1962b: 56.


TYPE LOCALITY AND HABITAT: Switzerland, Alp Tablasot (alt. 2,761 m), on moss on a

rock.


124. Gamasellus bellavistae Emberson, 1967

Gamasellus bellavistae Emberson, 1967: 298.

TYPE DEPOSITORY: Lyman Entomological Museum, Sainte-Anne-de-Bellevue, Canada.

TYPE LOCALITY AND HABITAT: Canada, Morgan Arboretum, [McGill University

Macdonald Campus, Sainte-Anne-de-Bellevue], Quebec, 17 March 1965, on moss.

125. Gamasellus borealis (Koch, 1879)

Gamasus borealis Koch, 1879: 120.

Cyrtolaelaps borealis.— Trägårdh, 1901: 61.

Gamasellus borealis.— Lee, 1970: 131.

266


TYPE LOCALITY AND HABITAT: Russia, Yenisei River, Cape Schaitanskoj, Siberia,


126. Gamasellus caucasicus Bregetova and Troitsky, 1981

Gamasellus caucasicus Bregetova and Troitsky, 1981: 76.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, near Village Tagardon, Kurtatinskoe Valley,

Caucasus Mountains, 24 September 1974, in litter of Carpinus sp. [Betulaceae] in a

forest.

127. Gamasellus changbaiensis Bei and Yin, 1995

Gamasellus changbaiensis Bei and Yin, 1995: 63.

TYPE DEPOSITORY: Department of Plant Protection, Shenyang Agricultural University,

Shenyang, China.

TYPE LOCALITY AND HABITAT: China, Changbai Mountain, Jilin Province, 25


128. Gamasellus concinus (Womersley, 1942)

? Digamasellus concina [sic] Womersley, 1942: 159.

Digamasellus concinnus.— Womersley, 1956b: 537.

Cyrtolaelaps concinnus.— Womersley, 1961: 194.

Cyrtolaelaps (Gamasellus) concinnus.— Ryke, 1962b: 54.

Gamasellus concinnus.— Lee, 1970: 131; Lee, 1973: 15.

Gamasellus concinus.— Halliday, 1998: 180.


TYPE LOCALITY AND HABITAT: Australia, Long Gully, South Australia, August 1938,

on moss.

NOTE: In the original description, the species name was spelled as concina, but according to

Womersley (1956b): 537 this was a lapsus calumi, then he changed it to concinnus.

Halliday (1998): 180 restored the original spelling because he did not consider that a

lapsus calumi had occurred.

267

129. Gamasellus cooperi (Womersley, 1961)

Cyrtolaelaps cooperi Womersley, 1961: 194.

Gamasellus cooperi.— Lee, 1970: 131.


TYPE LOCALITY AND HABITAT: Australia, Ravine des Casoars Wilderness Protection

Area, Kangaroo Island, South Australia, October 1951, on moss of the littoral zone.

NOTE: Described based on specimens reported by Womersley (1956b): 537 as Digamasellus

trägårdhi Womersley, 1942.

130. Gamasellus cophinus Lee, 1973

Gamasellus cophinus Lee, 1973: 17.

Gamasellus cophinus.— Karg, 1997a: 79.



Adelaide, South Australia, 7 June 1968 – 12 September 1968, on moss.

131. Gamasellus davydovae Vinnik, 1993

Gamasellus (Brevisellus) davydovae Vinnik, 1993: 27.



TYPE LOCALITY AND HABITAT: Georgia, beyond the pass of Hino, Kintrishi Reserve

(alt. 1,900 m), Adjara, 15 August 1988, in litter of a beech [Fagaceae] forest.

132. Gamasellus deepdalensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) deepdalensis Ryke, 1962b: 18.

Gamasellus deepdalensis.— Lee, 1970: 131; Hurlbutt, 1979: 174.


TYPE LOCALITY AND HABITAT: South Africa, Deepdale, KwaZulu-Natal, August 1940,

on forest floor.

133. Gamasellus discutatus (Lee, 1966)

Ologamasus discutatus Lee, 1966: 226.

Gamasellus discutatus.— Lee, 1970: 137.


268

TYPE LOCALITY AND HABITAT: Australia, 2-6 miles north of Sardine Creek (just south

of the Australian Alps), Victoria, 23 November 1959, in leaf litter.

134. Gamasellus dunhuaensis Ma, 2003

Gamasellus dunhuaensis Ma, 2003: 313.


TYPE LOCALITY AND HABITAT: China, Dunhua (43°21‘N, 128°13‘E), Jilin Province,

August 1990, in forest soil.

135. Gamasellus exiquns Davydova, 1982

Gamasellus (Brevisellus) exiquns Davydova, 1982: 81.


TYPE LOCALITY AND HABITAT: Russia, ridge of Kumrach, Kamchatka, 5 June 1978, in

litter, moss and lichen of tundra.

136. Gamasellus ezoensis Ishikawa, 1983

Gamasellus ezoensis Ishikawa, 1983: 114.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, Mount Asahidake (alt. 1,600 m), Hokkaido, 6

July 1970, in litter of Pinus pumila [Pinaceae].

137. Gamasellus falciger (G. Canestrini and R. Canestrini, 1881)

Gamasus falciger G. Canestrini and R. Canestrini, 1881: 1080.

Gamasus falciger.— G. Canestrini and R. Canestrini, 1882: 53; G. Canestrini, 1885: 77.

Cyrtolaelaps falciger.— Berlese, 1892a: 4.

Cyrtolaelaps (Gamasellus) falciger.— Berlese, 1892f: 61; Ryke, 1962b: 16; Ryke, 1962c:

157.

Gamasellus falciger.— Berlese, 1906: 278; Schweizer, 1922: 34; Vitzthum, 1931a: 142;

Baker and Wharton, 1952: 64; Ryke, 1958: 121; Lee, 1970: 131.

Gamasus hungaricus Supino, 1894: 195 [junior synonymy by Berlese, 1906: 282].

Gamasus hungaricus.— Tietze, 1899: 195

Cyrtolaelaps sertatus Willmann, 1941: 21 [junior synonymy by Ryke, 1962b: 16].

Gamasellus sertatus.— Lee, 1970: 131.

269

TYPE DEPOSITORY: G. falciger: Unknown.; G. hungaricus: Unknown.; G. sertatus:

Zoologische Staatssammlung München, Munich, Germany.

TYPE LOCALITY AND HABITAT: G. falciger: Italy, Oliero, near Bassano del Grappa, 16

May 1881, on moss; G. hungaricus: Hungary; C. sertatus: Bosnia and Herzegovina,

Dubrava, 30 March 1913, from unspecified substrate.

138. Gamasellus falculatus Athias-Henriot, 1961

Gamasellus falculatus Athias-Henriot, 1961a: 504.


Paris, France.

TYPE LOCALITY AND HABITAT: Corse Island, Porto Pollo Road, near Stillico,

Vizzavona, 1957, in litter under Fagus sp. [Fagaceae].

139. Gamasellus grishinae Davydova, 1982

Gamasellus (Eurysellus) grishinae Davydova, 1982: 55.


TYPE LOCALITY AND HABITAT: Russia, Uspenka, Tuva Republic, 16 June 1976, from a

foodplain forest.

140. Gamasellus grossi Lee, 1973

Gamasellus grossi Lee, 1973: 19.


TYPE LOCALITY AND HABITAT: Australia, Grange Golf Course, Adelaide, South

Australia, 10 June 1965, on moss.

141. Gamasellus heteropilus (Karg, 1977)

Allogamasellus heteropilus Karg, 1977: 339.


TYPE LOCALITY AND HABITAT: Chile, Nevados de Payachatas, [Parque Nacional Lauca,

Parinacota Province], 29 November 1965, in muddy soil under bushes.

142. Gamasellus humosus Ishikawa, 1969

Gamasellus humosus Ishikawa, 1969: 48.

Gamasellus humosus.— Ishikawa, 1983: 116.

270


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, ―Otanomosu-no-taira‘ (alt. 1,750 m), western

side of Mount Shiga, Shiga Kogen, Nagano Prefecture, 19 October 1967, in soil and

litter of a coniferous forest.

143. Gamasellus kurilensis Bregetova and Troitsky, 1981

Gamasellus kurilensis Bregetova and Troitsky, 1981: 78.

Gamasellus (Gamasellus) kurilensis.— Klimov, 1998: 14.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Paramushir, Kuril Islands, 14 September 1968,

in litter.

144. Gamasellus lanceolatus Liang and Ishikawa, 1989

Gamasellus lanceotatus [sic] Liang and Ishikawa, 1989: 143.

Gamasellus lanceolatus.— Liang and Ishikawa, 1989: 143.

TYPE DEPOSITORY: Department of Environmental and Resources Biology, Fudan

University, Shangai, China.

TYPE LOCALITY AND HABITAT: China, Qi-li-ting (alt. 960 m), West Tian-mu Mountain,

Zhejiang Province, 2 September 1989, in litter of evergreen broadleaved and

coniferous trees such as Litsea auriculata [Lauraceae] and Cryptomeria fortunei

[Cupressaceae].

145. Gamasellus lativentralis Ishikawa, 1983

Gamasellus lativentralis Ishikawa, 1983: 118.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, Daisen Mountain, Tottori Prefecture, 29

September 1969, in litter.

146. Gamasellus leggetti (Ryke, 1962)

Cyrtolaelaps (Gamasellus) leggetti Ryke, 1962b: 19.

Gamasellus leggetti.— Lee, 1970: 131.

271

TYPE DEPOSITORY: Institute for Zoological Research, Potchefstroom University,


TYPE LOCALITY AND HABITAT: USA, Texas, 9 February 1956, in litter.

147. Gamasellus litoprothrix (Lee, 1966)

Ologamasus litoprothrix Lee, 1966: 227.

Gamasellus litoprothrix.— Lee, 1970: 137.


TYPE LOCALITY AND HABITAT: Australia, Turtons Track, Otway Ranges, Victoria, 18

January 1962, in leaf litter and moss.

148. Gamasellus morogoroensis Hurlbutt, 1979

Gamasellus morogoroensis Hurlbutt, 1979: 179.

TYPE DEPOSITORY: United States National Museum of Natural History, Beltsville,

Maryland, USA.

TYPE LOCALITY AND HABITAT: Tanzania, near Morningside (alt. 1,330 m), south of

Morogoro, 24 May 1972, in litter and twigs from patch of trees.

149. Gamasellus muscosus Hurlbutt, 1979

Gamasellus muscosus Hurlbutt, 1979: 177.


Maryland, USA.

TYPE LOCALITY AND HABITAT: Tanzania, summit of Bondwa Peak (alt. 2,120 m),

Uluguru Mountains, Morogoro, 30 May 1972, on moss cushion of elfin forest.

150. Gamasellus nivalis Schweizer, 1949

Gamasellus nivalis Schweizer, 1949: 34.

Gamasellus nivalis.— Schweizer, 1961: 94; Karg, 1971: 350; Bregetova and Shcherbak,

1977a: 302; Karg, 1993c: 371.

Cyrtolaelaps (Gamasellus) nivalis.— Ryke, 1962b: 55.


TYPE LOCALITY AND HABITAT: Switzerland, Alp Tablasot (alt. 2,850 m), Schadler (alt.

2,950 m) and Piz Lischana (alt. 3,109 m), on moss and other low-growing plants.

272

151. Gamasellus orientalis Davydova, 1982

Gamasellus (Eurysellus) orientalis Davydova, 1982: 50.


TYPE LOCALITY AND HABITAT: Russia, Shkotovsky District, Primorsky Territory, 18

August 1978, from a mixed forest.

152. Gamasellus peninsularis Ishikawa, 1976

Gamasellus peninsularis Ishikawa, 1976: 246.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Malaysia, Pasoh Forest Reserve, 31 January 1971, in

soil.

153. Gamasellus plumatilis Karg, 1993

Gamasellus plumatilis Karg, 1993a: 41.

Gamasellus plumatilis.— Karg, 1997a: 77.



TYPE LOCALITY AND HABITAT: New Caledonia, Île des Pius [= Île des Pins?], 24

February 1977, in litter.

154. Gamasellus plumosus Ishikawa, 1983

Gamasellus plumosus Ishikawa, 1983: 112.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, Mount Naragara (alt. 900 m), Ehime Prefecture,

23 November 1968, in litter of Fagus crenata [Fagaceae].

155. Gamasellus puberulus Davydova, 1982

Gamasellus (Eurysellus) puberulus Davydova, 1982: 47.


TYPE LOCALITY AND HABITAT: Russia, Shkotovsky District, Primorsky Territory, 5

August 1978, in litter of a mixed forest.

273

156. Gamasellus pulcherimus Davydova, 1982

Gamasellus (Brevisellus) pulcherimus Davydova, 1982: 70.


TYPE LOCALITY AND HABITAT: Russia, Ussuriysky Reserve, Primorsky Territory, 31

July 1978, in a rot tree stump in a mixed forest.

157. Gamasellus pyriformis Berlese, 1916

Gamasellus pyriformis Berlese, 1916a: 161.

Cyrtolaelaps (Gamasellus) pyriformis.— Ryke, 1962b: 44.

Gamasellus pyriformis.— Lee, 1970: 138; Hurlbutt, 1979: 175; Karg, 1997a: 77.


TYPE LOCALITY AND HABITAT: East Africa, in soil and litter.

158. Gamasellus quartornatus Karg, 1997

Gamasellus quartornatus Karg, 1997a: 78.



TYPE LOCALITY AND HABITAT: New Caledonia, Col des Rousettes, 1977, in litter.

159. Gamasellus quintornatus Karg, 1996

Gamasellus quintornatus Karg, 1996: 186.

Gamasellus quintornatus.— Karg, 1997a: 77.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, in litter.

160. Gamasellus radicolus (Karg, 1977)

Allogamasellus? radicolus [sic] Karg, 1977: 341.


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 23 November 1965, from

moist leaf litter.


161. Gamasellus robustipes Berlese, 1908

274

Gamasellus robustipes Berlese, 1908: 13.

Cyrtolaelaps (Gamasellus) robustipes.— Ryke, 1962b: 52.


TYPE LOCALITY AND HABITAT: Italy, Florence, from unspecified substrate.

162. Gamasellus sexornatus Karg, 1997

Gamasellus sexornatus Karg, 1997a: 78.





163. Gamasellus shcherbakae Davydova, 1982

Gamasellus (Brevisellus) shcherbakae Davydova, 1982: 67.


TYPE LOCALITY AND HABITAT: Russia, Tandinsky District, Tyva Republic, 12 July

1976, in a Caragana [Fabaceae] brushwood.

164. Gamasellus shongweniensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) shongweniensis Ryke, 1962b: 20.

Gamasellus shongwiensis [sic].— Lee, 1970: 131.

Gamasellus shongweniensis.— Hurlbutt, 1979: 174.


TYPE LOCALITY AND HABITAT: South Africa, Shongweni Dam, KwaZulu-Natal, July


165. Gamasellus silvaticus Davydova, 1982

Gamasellus (Eurysellus) silvaticus Davydova, 1982: 42.


TYPE LOCALITY AND HABITAT: Russia, Algaysky Reserve, [Saratov Oblast], 15

September 1965, in litter and soil in a cedar woodland.

166. Gamasellus silvestris Halašková, 1958

Gamasellus silvestris Halašková, 1958: 347.

275

Gamasellus (Eurysellus) silvestris.— Davydova, 1982: 36.

Gamasellus alifanovi Davydova, 1970: 82 [junior synonymy by Davydova, 1982: 9].

TYPE DEPOSITORY: G. silvestris: Author´s private collection; G. alifanovi: Siberian

Zoological Museum, Novosibirsk, Russia.

TYPE LOCALITY AND HABITAT: G. silvestris: Czech Republic, Herlíkovice, Vrchlabí, 27

February 1955, in soil of spruce forest; G. alifanovi: Russia, Yamalo-Nenets

Autonomous Okrug, 14 August 1965, in lichens.

NOTE1: Hirschmann (1962): 52 considered Gamasellus silvestris Halašková, 1958 as junior

synonymy of Acugamasus montanus (Willmann, 1936), but Davydova (1982): 9


NOTE2: Bregetova and Shcherbak (1977a): 302 considered Gamasellus alifanovi Davydova,

1970 as junior synonymy of Acugamasus montanus (Willmann, 1936), but Davydova


166a. Gamasellus silvestris italicus Lombardini, 1962

Gamasellus silvestris v. italica Lombardini, 1962: 193.


TYPE LOCALITY AND HABITA: Italy, Malga Coltrondo, Comelico Superiore,

Belluno, 27 August 1959, in soil.

167. Gamasellus simpliciseta Liang and Ishikawa, 1989

Gamasellus simpliciseta Liang and Ishikawa, 1989: 149.






[Cupressaceae].

168. Gamasellus southcotti (Lee, 1966)

Ologamasus southcotti Lee, 1966: 232.

Gamasellus southcotti.— Lee, 1970: 137.


276

TYPE LOCALITY AND HABITAT: Australia, Gogerley‘s Point (about 20 miles south of

Sydney), New South Wales, 7 August 1965, from Eucalyptus [Myrtaceae] leaf litter

and grass.

169. Gamasellus spiricornis (G. Canestrini and R. Canestrini, 1882)

Gamasus spiricornis G. Canestrini and R. Canestrini, 1882: 52.

Gamasus spiricornis.— G. Canestrini, 1885: 77.

Cyrtolaelaps spiricornis.— Berlese, 1892c: 9.

Cyrtolaelaps (Gamasellus) spiricornis.— Berlese, 1892f: 61; Ryke, 1962b: 50.

Gamasellus spiricornis.— Berlese, 1906: 288; Schweizer, 1922: 34; Cooreman, 1943: 18;

Schweizer, 1961: 92; Karg, 1971: 350; Bregetova and Shcherbak, 1977a: 301; Karg,

1993c: 370.

Gamasellus (?) spiricornis [sic].— Trägårdh, 1910: 425.


TYPE LOCALITY AND HABITAT: Italy, Trento, August, in litter.

170. Gamasellus sternopunctatus Vinnik, 1993

Gamasellus (Brevisellus) sternopunctatus Vinnik, 1993: 29.



TYPE LOCALITY AND HABITAT: Georgia, Kintrishi Reserve (alt. 800 m), Adjara, 22

August 1988, on the roots of plants in a fissure of an old chestnut.

171. Gamasellus taeniatus Davydova, 1982

Gamasellus (Eurysellus) taeniatus Davydova, 1982: 45.


TYPE LOCALITY AND HABITAT: Russia, Stolby Reserve, Krasnoyarsk Territory, 25 June

1974, in rocky soil from an elevation in a taiga region.

172. Gamasellus tasmanicus (Womersley, 1956)

Digamasellus tasmanicus Womersley, 1956b: 538.

Cyrtolaelaps tasmanicus.— Womersley, 1961: 194.

Cyrtolaelaps (Gamasellus) tasmanicus.— Ryke, 1962b: 55.

Gamasellus tasmanicus.— Lee, 1970: 137.

277


TYPE LOCALITY AND HABITAT: Australia, Mount Wellington, Tasmania, 27 July 1943,

on moss.

173. Gamasellus tengkuofani (Bai, Yan and Wei, 2010)

Euryparasitus tengkuofani Bai, Yan and Wei, in Bai et al., 2010: 174.

TYPE DEPOSITORY: Institute of Microbiology and Epidemiology, Academy of Military

Medical Sciences, Beijing, China.

TYPE LOCALITY AND HABITAT: China, Liupan Mountain Natural Reserve (106°13‘N,

35°30‘E, alt. 2,140 m), Ningxia Hui Autonomous Region, 4 June 2006, on Apodemus

peninsulae [Mammalia: Rodentia: Muridae].

174. Gamasellus tianmuensis Liang and Ishikawa, 1989

Gamasellus tianmuensis Liang and Ishikawa, 1989: 147.






[Cupressaceae].

175. Gamasellus tindalei (Lee, 1966)

Ologamasus tindalei Lee, 1966: 230.

Gamasellus tindalei.— Lee, 1970: 137.


TYPE LOCALITY AND HABITAT: Australia, Mount Ginini (alt. 5,700 feet, northern edge

of Australian Alps), Australian Capital Territory, 17 October 1965, on moss and litter

among ―snow grass‖ near ―snow gums‖ [Eucalyptus niphophila (Myrtaceae)].

176. Gamasellus tragardhi (Womersley, 1942)

? Digamasellus tragardhi [sic] Womersley, 1942: 161.

Cyrtolaelaps trägårdhi.— Womersley, 1961: 194.

Cyrtolaelaps (Gamasellus) tragardhi.— Rike, 1962b: 55.

Gamasellus tragardhi.— Lee, 1970: 135; Lee, 1973: 20.

278


TYPE LOCALITY AND HABITAT: Australia, Adelaide, June 1935, on moss.

NOTE: Specimens reported by Womersley (1956b): 537 as Digamasellus trägårdhi

Womersley, 1942 were described as Cyrtolaelaps cooperi Womersley, 1961: 194.

177. Gamasellus tschucotensis Davydova, 1982

Gamasellus (Eurysellus) tschucotensis Davydova, 1982: 39.


TYPE LOCALITY AND HABITAT: Russia, Chukotka Autonomous Okrug, 25 September

1972, in soil.

178. Gamasellus tundriensis Davydova, 1982

Gamasellus (Brevisellus) tundriensis Davydova, 1982: 75.


TYPE LOCALITY AND HABITAT: Russia, Taymyr Peninsula, 1 August 1977, on moss in

tundra.

179. Gamasellus tuvinycus Davydova, 1982

Gamasellus (Eurysellus) tuvinycus Davydova, 1982: 58.


TYPE LOCALITY AND HABITAT: Russia, Tandinsky District, Tyva Republic, 2 July

1976, in litter under a Caragana [Fabaceae] thicket bush.

180. Gamasellus uluguruensis Hurlbutt, 1979

Gamasellus uluguruensis Hurlbutt, 1979: 181.


Maryland, USA

TYPE LOCALITY AND HABITAT: Tanzania, Uluguru Mountains (alt. 1,600 m), 16

September 1966, on dead leaves from forest floor.

181. Gamasellus venustus Ishikawa, 1983

Gamasellus venustus Ishikawa, 1983: 117.


Matsuyama, Japan.

279

TYPE LOCALITY AND HABITAT: Japan, Machigasawa, Mount Tanigawa, Gunma

Prefecture, 17 November 1976, in litter of Fagus crenata [Fagaceae].

182. Gamasellus vibrissatus Emberson, 1967

Gamasellus vibrissatus Emberson, 1967: 294.

Gamasellus vibrissatus.— Bregetova and Shcherbak, 1977a: 301; Ma, Kuang and Lin, 2007:

246.

Gamasellus (Brevisellus) vibrissatus.— Davydova, 1982: 64.

TYPE DEPOSITORY: Lyman Entomological Museum, Sainte-Anne-de-Bellevue, Canada.

TYPE LOCALITY AND HABITAT: Canada, Canadian International Paper Company Tree

Farm, Harrington, Quebec, 31 October 1964, in litter of Abies sp. [Pinaceae] and

Picea mariana [Pinaceae].

183. Gamasellus villosus Davydova, 1982

Gamasellus (Eurysellus) villosus Davydova, 1982: 53.


TYPE LOCALITY AND HABITAT: Russia, Sharypovsky District, Krasnoyarsk Territory,

14 July 1978, in litter of a mixed forest.

184. Gamasellus virgosus (Lee, 1966)

Ologamasus virgosus Lee, 1966: 233.

Gamasellus virgosus.— Lee, 1970: 137.


TYPE LOCALITY AND HABITAT: Australia, Gogerley‘s Point (about 20 miles south of

Sydney), New South Wales, 7 August 1965, from Eucalyptus [Myrtaceae] leaf litter

and grass.

185. Gamasellus virguncula (Lee, 1973)

Onchogamasus virguncula Lee, 1973: 34



Adelaide, South Australia, 5 August 1968, in plant litter.

186. Gamasellus volkovi Davydova, 1982

280

Gamasellus (Brevisellus) volkovi Davydova, 1982: 73.


TYPE LOCALITY AND HABITAT: Russia, Hehtsirsky reserve, Khabarovsk Territory, 25

October 1979, in litter of a Pinus [Pinaceae] forest.

187. Gamasellus xini Liang and Ishikawa, 1989

Gamasellus xini Liang and Ishikawa, 1989: 144.






[Cupressaceae].

188. Gamasellus yastrebtsovi Vinnik, 1993

Gamasellus (Eurysellus) yastrebtsovi Vinnik, 1993: 31.



TYPE LOCALITY AND HABITAT: Georgia, Hino, Kintrishi Reserve (alt. 2,050 m), Adjara,

16 August 1988, in remains of a fallen tree.

189. Gamasellus yosiianus Ishikawa, 1999

Gamasellus yosiianus Ishikawa, 1999: 20.

TYPE DEPOSITORY: Department of Zoology, National Science Museum (Natural History),

Tokyo, Japan.

TYPE LOCALITY AND HABITAT: Japan, Mawaribuchi-do Cave (alt. 810 m), Enchi,

Odamiyama, Oda-chô, Kamiukena, Ehime Prefecture, Shikoku, 5 May 1990, in a

limestone cave.

Genus Gamasiphis Berlese, 1904

Gamasiphis Berlese, 1904: 261 [presumed to be described in Parasitidae Oudemans (also


281

Gamasiphis.— Berlese, 1906: 101; Berlese, 1914: 137; Vitzthum, 1931a: 142; Womersley,

1942: 155; Vitzthum, 1943: 756; Baker and Wharton, 1952: 73; Trägårdh, 1952: 54;

Ryke, 1962c: 159; Bregetova, 1977a: 308; Lee, 1970: 42; Zaher, 1986: 32; Karg,

1987: 302; Karg, 1990: 321; Karg, 1993b: 169; Karg, 1993c: 373; Karg, 1995: 15;

Karg, 1996: 173; Karg, 1997a: 73; Karg, 1998: 196; Karg, 2007: 125; Karg and

Schorlemmer, 2009: 83; Castilho, Moraes and Narita, 2010: 32.

Gamasiphis (Gamasiphis).— Trägårdh, 1952: 55; Womersley, 1956b: 518.

Type species: Gamasus pulchellus Berlese, 1887, by monotypy.

Ologamasellus (Micriphis) Berlese, 1914: 140 [junior synonymy by Lee, 1970: 42].

Ologamasus (Micriphis).— Vitzthum, 1943: 756; Baker and Wharton, 1952: 73.

Micriphis.— Ryke, 1962c: 160.

Type species: Gamasiphis gamasellus Berlese, 1913, by monotypy.

Gamasiphis (Heteroiphis) Trägårdh, 1952: 55 [junior synonymy by Ryke, 1962c: 160].

Gamasiphis (Heteroiphis).— Womersley, 1956b: 526.

Heteroiphis.— Ryke, 1962c: 160; Bhattacharyya, 1968: 530.

Type species: Gamasiphis (Heteroiphis) arcuatus Trägårdh, 1952, by original

designation.

Neogamasiphis Trägårdh, 1952: 57 [junior synonymy by Lee, 1970: 42].

Gamasiphis (Neogamasiphis).— Womersley, 1956b: 517.

Neogamasiphis.— Ryke, 1962c: 160.

Type species: Neogamasiphis hamifer Trägårdh, 1952, by original designation.

190. Gamasiphis adanalis Karg, 1990

Gamasiphis adanalis Karg, 1990: 327.

Gamasiphis adanalis.— Karg, 1993b: 187; Karg, 1997a: 77; Karg, 2007: 125; Castilho,

Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Lesser Antilles [Caribbean Region (cited as Central

America)], Saint Lucia, 1980, in soil.

282

191. Gamasiphis aduncus Ma, 2004

Gamasiphis aduncus Ma, 2004: 23.


TYPE LOCALITY AND HABITAT: China, Changchun (43°54‘N, 125°18‘E), Jilin Province,

15 June 1990, in a forest humus.

192. Gamasiphis anguis Karg, 1993

Gamasiphis anguis Karg, 1993b: 177.



TYPE LOCALITY AND HABITAT: New Caledonia, Poindimié, 30 January 1977, in litter

between roots on a rock.

193. Gamasiphis appendicularis Karg, 1993

Gamasiphis appendicularis Karg, 1993b: 177.




a cave.

194. Gamasiphis arcuatus Trägårdh, 1952

Gamasiphis (Heteroiphis) arcuatus Trägårdh, 1952: 55.

Heteroiphis arcuatus.— Bhattacharyya, 1968: 530.

Gamasiphis arcuatus.— Lee, 1970: 49; Karg, 1987: 305; Karg, 1990: 333; Karg, 1993b: 172;

Karg, 1996: 174.

TYPE DEPOSITORY: Bishop Museum, Honolulu, Hawaii.

TYPE LOCALITY AND HABITAT: French Polynesia, Arihiri, Pare, Tahiti, 16 March 1934,



195. Gamasiphis ardor Karg, 1993

Gamasiphis ardor Karg, 1993b: 176.

Gamasiphis ardor.— Karg, 1997a: 74.

283




and moss between roots on a rock.

196. Gamasiphis australicus Womersley, 1956

Gamasiphis (Heteroiphis) australicus Womersley, 1956b: 527.

Gamasiphis australicus.— Lee, 1970: 50; Lee, 1973: 3; Karg, 1987: 305; Karg, 1990: 333;

Karg, 1993b: 172; Karg, 1996: 175.


TYPE LOCALITY AND HABITAT: Australia, Mylor, South Australia, 28 June 1948, on

moss.

NOTE: According to Lee (1970): 50, specimens reported by Domrow (1957): 202 as

Gamasiphis (Heteroiphis) australicus are Gamasiphis setosus Womersley, 1956.

197. Gamasiphis bengalensis Bhattacharyya, 1966

Gamasiphis (Neogamasiphis) bengalensis Bhattacharyya, 1966: 151.

Gamasiphis bengalensis.— Lee, 1970: 49; Bhattacharyya, 1978: 83; Karg, 1987: 306; Karg,

1990: 334; Karg, 1993b: 182; Karg, 1996: 179.

TYPE DEPOSITORY: Author´s private collection.

TYPE LOCALITY AND HABITAT: India, Pond Sitala, Sonarpur, 24 Parganas District, West

Bengal, 2 December 1963, in litter under decaying Pistia stratiotes [Araceae].

198. Gamasiphis benoiti Loots, 1980

Gamasiphis benoiti Loots, 1980: 748.


TYPE LOCALITY AND HABITAT: Seychelles [Indian Ocean], Morne Blanc, Mahé Centre,

24-25 August 1972, from unspecified substrate.


199. Gamasiphis breviflagelli Karg, 1996

Gamasiphis breviflagelli Karg, 1996: 177.



284

TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, 29 January 1977, in litter

beetwen rocks.

200. Gamasiphis brevigenitalis Karg, 1993

Gamasiphis brevigenitalis Karg, 1993b: 187.

Gamasiphis brevigenitalis.— Karg, 1997a: 75.





201. Gamasiphis caper Karg, 1995

Gamasiphis caper Karg, 1995: 18.



TYPE LOCALITY AND HABITAT: New Caledonia, Mont Mandjelia (alt. 600 m), 1977, in

litter of a deciduous forest.

202. Gamasiphis conciliator Berlese, 1916

Gamasiphis (Periphis) conciliator Berlese, 1916a: 159.

Gamasiphis (Periphis) conciliator.— Berlese, 1923b: 123.

Gamasiphis conciliator.— Lee, 1970: 50; Karg, 1987: 306; Karg, 1990: 333; Karg, 1993b:

182; Karg, 1996: 179.


TYPE LOCALITY AND HABITAT: New Caledonia, Mont Panié, from unspecified

substrate.

203. Gamasiphis coniunctus Karg, 1995

Gamasiphis coniunctus Karg, 1995: 17.





285

204. Gamasiphis decoris Karg, 1990

Gamasiphis decoris Karg, 1990: 327.

Gamasiphis decoris.— Karg, 1993b: 183; Karg, 1998: 197; Karg, 2007: 126; Castilho,






205. Gamasiphis denticus Hafez and Nasr, 1979

Gamasiphis denticus Hafez and Nasr, 1979: 80.

Gamasiphis denticus.— Zaher, 1986: 34.



University, Giza, in debris under orange [Rutaceae] trees.

206. Gamasiphis elegantellus Berlese, 1910

Gamasiphis elegantellus Berlese, 1910b: 253.

Gamasiphis elegantellus.— Berlese, 1913a: 81; Berlese, 1914: 143; Lee, 1970: 49; Karg,

1987: 306; Karg, 1990: 334; Karg, 1993b: 182; Karg, 1996: 179; Karg, 2007: 125.


TYPE LOCALITY AND HABITAT: Indonesia, Bogor [cited as Buitenzorg], Java, in humus.

207. Gamasiphis ellipticus Karg, 1996

Gamasiphis ellipticus Karg, 1996: 176.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, in humus beetwen roots.

208. Gamasiphis elongatellus Berlese, 1910

Gamasiphis elongatellus Berlese, 1910c: 372.

Gamasiphis elongatellus.— Berlese, 1913a: 81; Berlese, 1914: 142; Lee, 1970: 49; Karg,

1987: 306; Karg, 1990: 334; Karg, 1993b: 183.


286

TYPE LOCALITY AND HABITAT: Indonesia, Semarang, Java, from unspecified substrate.

209. Gamasiphis erinaceus Karg, 1993

Gamasiphis erinaceus Karg, 1993b: 185.






210. Gamasiphis euincisus Karg, 1996

Gamasiphis euincisus Karg, 1996: 175.




roots in a deciduous forest.

211. Gamasiphis eumagnus Karg, 1996

Gamasiphis eumagnus Karg, 1996: 176.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, 29 January 1977, in litter

beetwen rocks.

212. Gamasiphis flagelli Karg, 1993

Gamasiphis flagelli Karg, 1993b: 177.




between roots on a rock.

213. Gamasiphis foliatus Karg, 1993

Gamasiphis foliatus Karg, 1993b: 173.

Gamasiphis foliatus.— Karg, 1995: 16.

287





214. Gamasiphis fornicatus Lee, 1970

Gamasiphis fornicatus Lee, 1970: 51.

Gamasiphis fornicatus.— Lee, 1973: 4; Karg, 1987: 307; Karg, 1990: 334; Karg, 1993b: 186.


TYPE LOCALITY AND HABITAT: Australia, Mount Remarkable, South Australia, 9

August 1966, on moss amongst Pteridium sp. [Pteridophyta] and Eucalyptus sp.

[Myrtaceae] trees in a gully.

215. Gamasiphis furcatus Karg, 1990

Gamasiphis furcatus Karg, 1990: 322.

Gamasiphis furcatus.— Karg, 1993b: 187; Karg, 1997a: 75; Karg, 2007: 125; Castilho,






216. Gamasiphis gamasellus Berlese, 1913

Gamasiphis gamasellus Berlese, 1913a: 80.

Ologamasellus (Micriphis) gamasellus.— Berlese, 1914: 140.

Ologamasus (Micriphis) gamasellus.— Baker and Wharton, 1952: 73.

Micriphis gamasellus.— Ryke, 1962c: 160.

Gamasiphis gamasellus.— Lee, 1970: 51; Karg, 1987: 307.


TYPE LOCALITY AND HABITAT: Indonesia, Semarang, Java, in humus.

217. Gamasiphis gandensius Van Daele, 1975

Gamasiphis gandensius Van Daele, 1975: 267.

288

Gamasiphis gandensius.— Karg, 1987: 305; Karg, 1990: 333; Karg, 1993b: 172; Karg,

1993c: 373; Karg, 1996: 175.

TYPE DEPOSITORY: Zoology Museum, Faculty of Sciences, Ghent University, Ghent,

Belgium.

TYPE LOCALITY AND HABITAT: Belgium, Heusden, Ghent, 14-15 September 1974, in

litter of azalea [Ericaceae] culture in an outdoor nursery.

218. Gamasiphis hamatellus Karg, 1998

Gamasiphis hamatellus Karg, 1998: 196.

Gamasiphis hamatellus.— Castilho, Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Ecuador, Rio Guajalito (alt. 1,850 m), Las Palmeras,

Pichincha Province, 18 April 1989, on moss from a rocky roadside.

219. Gamasiphis hamifer (Trägårdh, 1952)

Neogamasiphis hamifer Trägårdh, 1952: 57.

Gamasiphis (Neogamasiphis) hamifer.— Womersley, 1956b: 526.

Neogamasiphis hamifer.— Ryke, 1962c: 160.

Gamasiphis hamifer.— Lee, 1970: 49; Karg, 1987: 306; Karg, 1990: 333; Karg, 1993b: 179;

Karg, 2007: 125.

TYPE DEPOSITORY: Bishop Museum, Honolulu, Hawaii.

TYPE LOCALITY AND HABITAT: Flint Island [Pacific Ocean], 16 October 1934, under

coconut trash; Rapa Island [Pacific Ocean], 16 July 1935, from ground with land-

shells.

220. Gamasiphis hemicapillus Karg, 1990

Gamasiphis hemicapillus Karg, 1990: 327.

Gamasiphis hemicapillus.— Karg, 1993b: 183; Karg, 1998: 197; Karg, 2007: 126; Castilho,






289

221. Gamasiphis holocapillus Karg, 1990

Gamasiphis holocapillus Karg, 1990: 322.

Gamasiphis holocapillus.— Karg, 1993b: 172; Karg, 1996: 173; Castilho, Moraes and Narita,

2010: 41.





222. Gamasiphis hyalinus Karg, 2003

Gamasiphis hyalinus Karg, 2003: 242.

Gamasiphis hyalinus.— Castilho, Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Costa Rica, La Selva, Heredia Province, April 1993, in

soil.

223. Gamasiphis illotus Fox, 1949

Gamasiphis illotus Fox, 1949: 37.

Gamasiphis illotus.— Lee, 1970: 49; Karg, 1987: 307; Castilho, Moraes and Narita, 2010: 41.


Juan, Puerto Rico.

TYPE LOCALITY AND HABITAT: Puerto Rico, Santurce, San Juan, 1947, from Rattus


224. Gamasiphis incisus Karg, 1993

Gamasiphis incisus Karg, 1993b: 170.

Gamasiphis incisus.— Karg, 1996: 175.



TYPE LOCALITY AND HABITAT: New Caledonia, Poindimié, 30 January 1977, in litter,

moss and between roots on a rock.

290

225. Gamasiphis incudis Karg, 1993

Gamasiphis incudis Karg, 1993b: 172.

Gamasiphis incudis.— Karg, 1995: 16.




a cave.

226. Gamasiphis indicus Bhattacharyya, 1978

Gamasiphis indicus Bhattacharyya, 1978: 82.

Gamasiphis indicus.— Karg, 2007: 125.


TYPE LOCALITY AND HABITAT: India, Sitala, Sonarpur, 24 Parganas District, West

Bengal, 13 August 1971, from unspecified substrate.

227. Gamasiphis krieli Van Driel, Loots and Marais, 1977

Gamasiphis krieli Van Driel, Loots and Marais, 1977: 318.

Gamasiphis krieli.— Karg, 1987: 307; Karg, 1990: 334; Karg, 1993b: 183; Karg, 2007: 126.


TYPE LOCALITY AND HABITAT: Saint Helena Island [Atlantic Ocean], High Peak, April

1967, in soil.

228. Gamasiphis lanceolatus Karg, 1987

Gamasiphis lanceolatus Karg, 1987: 301.

Gamasiphis lanceolatus.— Karg, 1990: 334; Karg, 1993b: 183; Karg, 1993c: 373; Karg,

1998: 197; Karg, 2007: 126.


Germany.

TYPE LOCALITY AND HABITAT: Germany, Rostock, 22 July 1982, in soil of a cucumber

[Curcubitaceae] crop in a greenhouse.

229. Gamasiphis lenifornicatus Lee, 1973

Gamasiphis lenifornicatus Lee, 1973: 4.


291


Adelaide, South Australia, 24 April 1969, in plant litter.

230. Gamasiphis longiorsetosus Karg, 1997

Gamasiphis longiorsetosus Karg, 1997a: 73.



TYPE LOCALITY AND HABITAT: New Caledonia, Monts Koghi, 23 January 1977, from


231. Gamasiphis longirimae Karg, 1997

Gamasiphis longirimae Karg, 1997a: 77.



TYPE LOCALITY AND HABITAT: New Caledonia, Mont Panié (alt. 400 m), 1977,

between moss and ferns [Pteridophyta].

232. Gamasiphis macrorbis Karg, 1993

Gamasiphis macrorbis Karg, 1993b: 180.

Gamasiphis macrorbis.— Karg, 1996: 179; Karg, 2007: 126.




a cave.


233. Gamasiphis maheensis Loots, 1980

Gamasiphis maheensis Loots, 1980: 745.


TYPE LOCALITY AND HABITAT: Seychelles [Indian Ocean], Morne Blanc (alt. 667 m),

Mahé Centre, 24-25 August 1972, from unspecified substrate.

234. Gamasiphis mediosetosus Karg, 2003

Gamasiphis mediosetosus Karg, 2003: 242.

292

Gamasiphis mediosetosus.— Castilho, Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Ecuador, 1990, in soil.

235. Gamasiphis minoris Karg, 1996

Gamasiphis minoris Karg, 1996: 179.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, 1977, on moss with

coniferous needles on a rock.


236. Gamasiphis novipulchellus Ma and Yin, 1998

Gamasiphis novipulchellus Ma and Yin, 1998: 319.

Gamasiphis novipuljchellus [sic].— Ma, 2004: 26.


TYPE LOCALITY AND HABITAT: China, Liangshui Natural Reserve (47°10‘N,

128°53‘E), Heilongjiang Province, August 1995, in forest soil.

237. Gamasiphis ovoides Karg, 1993

Gamasiphis ovoides Karg, 1993b: 185.




on a rock.

238. Gamasiphis parpulchellus Nasr and Mersal, 1986

Gamasiphis parpulchellus Nasr and Mersal, 1986 apud Zaher, 1986: 35.

Gamasiphis parpulchellus.— Zaher, 1986: 35.


TYPE LOCALITY AND HABITAT: Egypt, Giza and El Qualyobia, in litter.

239. Gamasiphis paulista Castilho, Moraes and Narita, 2010

293

Gamasiphis paulista Castilho, Moraes and Narita, 2010: 32.


Agricultura ―Luiz de Queiroz‖, Universidade de São Paulo, Piracicaba, Brazil.

TYPE LOCALITY AND HABITAT: Brazil, ―Salvador de Toledo Piza Junior‖ building,

campus of Escola Superior de Agricultura ―Luiz de Queiroz‖ (22°42‘30‖S,

47°38‖00‖W), Universidade de São Paulo, Piracicaba, São Paulo State, July 2008, in

litter and soil of a inner garden.

240. Gamasiphis pilosellus Berlese, 1913

Gamasiphis pilosellus Berlese, 1913a: 81.

Gamasiphis pilosellus.— Vitzthum, 1931b: 65; Lee, 1970: 49; Karg, 1987: 306; Karg, 1990:

334; Karg, 1993b: 182; Karg, 1996: 179.


TYPE LOCALITY AND HABITAT: Indonesia, Semarang, Java, in humus.

241. Gamasiphis pinguis Karg, 1990

Gamasiphis pinguis Karg, 1990: 330.

Gamasiphis pinguis.— Karg, 1993b: 183; Karg, 2007: 125; Castilho, Moraes and Narita,

2010: 41.





242. Gamasiphis pinnatus Karg, 1998

Gamasiphis pinnatus Karg, 1998: 196.

Gamasiphis pinnatus.— Castilho, Moraes and Narita, 2010: 41.




Pichincha Province, 14 April 1989, on moss and withered plant-debris under bushes.

294

243. Gamasiphis plenosetosus Karg, 1994

Gamasiphis plenosetosus Karg, 1994b: 210.

Gamasiphis plenosetosus.— Castilho, Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Galapagos Islands, near Media Luna (alt. 590 m), Santa

Cruz Island, 6 February 1985, in litter, roots and wood pieces in a Miconia sp.

[Melastomataceae] area.

244. Gamasiphis productellus Berlese, 1923

Gamasiphis productellus Berlese, 1923a: 250.

Gamasiphis productellus.— Lee, 1970: 49; Karg, 1987: 307.


TYPE LOCALITY AND HABITAT: China, from unspecified substrate.

245. Gamasiphis pulchellus (Berlese, 1887)

Gamasus pulchellus Berlese, 1887a: 4.

Gamasus (Hologamasus) pulchellus.— Berlese, 1892f: 62.

Gamasiphis pulchellus.— Berlese, 1904: 261; Berlese, 1906: 287; Lee, 1970: 49; Bregetova,

1977a: 308; Hafez and Nasr, 1979: 78; Zaher, 1986: 33; Karg, 1987: 306; Hennessey

and Farrier, 1988: 17; Karg, 1990: 334; Karg, 1993b: 182; Karg, 1993c: 373; Karg,

1996: 179; Ma and Yin, 1998: 319; Halliday, 2005: 42.

Laelaps bermudaensis Ewing, 1920: 287 [junior synonymy by Hennessey and Farrier, 1988:

17].

TYPE DEPOSITORY: G. pulchellus: Istituto Sperimentale per la Zoologia Agraria, Florence,

Italy; L. bermudaensis: Smithsonian National Museum of Natural History,

Washington, District of Columbia, USA.

TYPE LOCALITY AND HABITAT: G. pulchellus: unspecified type locality, in humus; L.

bermudaensis: Bermuda Islands [Atlantic Ocean], Pembroke Parish, from Musa sp.

[Musaceae] and Cedrus sp. [Pinaceae] plantations.

246. Gamasiphis quadruplicis Karg, 1990

Gamasiphis quadruplicis Karg, 1990: 332.

295

Gamasiphis quadruplicis.— Karg, 1993b: 180; Karg, 1996: 179; Castilho, Moraes and Narita,

2010: 41.





247. Gamasiphis saccus Lee, 1973

Gamasiphis saccus Lee, 1973: 7.


TYPE LOCALITY AND HABITAT: Australia, near First Waterfall, Waterfall Gully,

Adelaide, South Australia, 21 June 1968, on moss.

248. Gamasiphis setosus Womersley, 1956

Gamasiphis (Neogamasiphis) setosus Womersley, 1956b: 521.

Gamasiphis setosus.— Lee, 1970: 49; Karg, 1987: 306; Karg, 1990: 333; Karg, 1993b: 176;

Karg, 1995: 16.


TYPE LOCALITY AND HABITAT: Australia, Carney‘s Creek, Queensland, 19 February


NOTE: According to Lee (1970): 50, specimens reported by Domrow (1957): 202 as

Gamasiphis (Heteroiphis) australicus Womersley, 1956 are Gamasiphis setosus.

249. Gamasiphis sextus Vitzthum, 1921

Gamasiphis (Gamasiphis) sextus Vitzthum, 1921: 10.

Gamasiphis sextus.— Karg, 1971: 352; Karg, 1987: 306; Karg, 1990: 334; Karg, 1993b: 182;

Karg, 1993c: 373; Karg, 1996: 179; Karg, 2007: 126.


TYPE LOCALITY AND HABITAT: Germany, Weimar, in a greenhouse where orchids

[Orchidaceae] were grown.

250. Gamasiphis silvestris Karg, 2007

Gamasiphis silvestris Karg, 2007: 124.

Gamasiphis silvestris.— Castilho, Moraes and Narita, 2010: 41.

296


TYPE LOCALITY AND HABITAT: Ecuador, between Calderón and Quevedo, 1990, in

litter in a rainforest.

251. Gamasiphis spinulosus Karg, 1995

Gamasiphis spinulosus Karg, 1995: 19.





252. Gamasiphis superardor Karg, 1993

Gamasiphis superardor Karg, 1993b: 180.

Gamasiphis superardor.— Karg, 1996: 179; Karg, 2007: 125.



TYPE LOCALITY AND HABITAT: New Caledonia, Koumac, 30 January 1977, in litter of a

cave.


253. Gamasiphis trituberosus Karg, 1990

Gamasiphis trituberosus Karg, 1990: 325.

Gamasiphis trituberosus.— Karg, 1993b: 187; Karg, 1997a: 77; Karg, 2007: 125.






254. Gamasiphis turgicalcareus Ma, 2009

Gamasiphis turgicalcareus Ma, 2009: 78.


TYPE LOCALITY AND HABITAT: China, Zunyi (27°43‘N, 106°51‘E), Guizhou Province,

10 June 2004, in leaf litter.

297

255. Gamasiphis uncifer Trägårdh, 1931

Gamasiphis uncifer Trägårdh, 1931a: 604.

Gamasiphis (Neogamasiphis) uncifer.— Womersley, 1956b: 526.

Gamasiphis uncifer.— Lee, 1970: 49; Karg, 1987: 307.


TYPE LOCALITY AND HABITAT: Juan Fernández Islands [South Pacific Ocean],

Robinson Crusoe Island [cited as Masatierra], among dry leaves.

256. Gamasiphis undulatus Karg and Schorlemmer, 2009

Gamasiphis undulatus Karg and Schorlemmer, 2009: 83.

Gamasiphis undulatus.— Castilho, Moraes and Narita, 2010: 41.



TYPE LOCALITY AND HABITAT: Ecuador, near Loreto, 1990, in litter of a coffee

[Rubiaceae] plantation.

257. Gamasiphis vinculi Karg, 1994

Gamasiphis vinculi Karg, 1994a: 122.

Gamasiphis vinculi.— Castilho, Moraes and Narita, 2010: 42.



TYPE LOCALITY AND HABITAT: Galapagos Islands, near Los Gemelos, Santa Cruz

Island, 16 April 1985, in moist litter of a Scalesia sp. [Asteraceae] area.

Genus Gamasiphoides Womersley, 1956

Gamasiphis (Gamasiphoides) Womersley, 1956b: 528 (described in Neoparasitidae

Oudemans).

Gamasiphoides.— Ryke, 1962c: 160; Lee, 1970: 59; Karg, 1976d: 3; Karg, 1993a: 48; Karg,

1996: 180; Halliday, 2005: 42; Karg and Schorlemmer, 2011a: 19.

Type species: Gamasiphis (Gamasiphoides) propinqua Womersley, 1956, by original

designation.

298

258. Gamasiphoides acanthioides Karg and Schorlemmer, 2011

Gamasiphoides acanthioides Karg and Schorlemmer, 2011a: 19.



TYPE LOCALITY AND HABITAT: Brazil, Pirapora do Bom Jesus [cited as Bom Jesus de

Pirapora], [São Paulo State], 1970, in humus.

259. Gamasiphoides aitkeni Lee, 1970

Gamasiphoides aitkeni Lee, 1970: 63.

Gamasiphoides aitkeni.— Karg, 1976d: 4; Karg, 1993a: 51; Karg, 1996: 183; Karg and

Schorlemmer, 2011a: 25.


TYPE LOCALITY AND HABITAT: Australia, near Glenelg River, near Nelson, Victoria, 28

January 1965, on moss and litter under shrubs in a cliff (top of 100 feet).

260. Gamasiphoides baloghi Karg, 1976

Gamasiphoides baloghi Karg, 1976d: 16.

Gamasiphoides baloghi.— Karg, 1993a: 51; Karg, 1996: 183; Karg and Schorlemmer, 2011a:

26.


TYPE LOCALITY AND HABITAT: Chile, road to Valdivia, La Unión, 26 October 1965, in

litter.

261. Gamasiphoides brevisetis Karg, 1976

Gamasiphoides brevisetis Karg, 1976d: 10.

Gamasiphoides brevisetae [sic].— Karg, 1976d: 10.

Gamasiphoides brevisetis.— Karg, 1993a: 51; Karg, 1996: 182; Karg and Schorlemmer,

2011a: 25.


TYPE LOCALITY AND HABITAT: Chile, near Lauca River, Tarapacá Region, 1 December

1965, in soil trap.

262. Gamasiphoides caudatae Karg, 1996

299

Gamasiphoides caudatae Karg, 1996: 183.



TYPE LOCALITY AND HABITAT: New Caledonia, Lifou, [Loyalty Islands], 20 February

1977, in jungle.


263. Gamasiphoides coniunctus Karg, 1976

Gamasiphoides coniunctus Karg, 1976d: 15.

Gamasiphoides coniunctus.— Karg, 1993a: 51; Karg, 1996: 183; Karg and Schorlemmer,

2011a: 26.



November 1965, under stones.

264. Gamasiphoides costai Lee and Hunter, 1974

Gamasiphoides costai Lee and Hunter, 1974: 302.

Gamasiphoides costai.— Karg, 1993a: 51; Karg and Schorlemmer, 2011a: 26.


TYPE LOCALITY AND HABITAT: New Zealand, Ocean Island, Auckland Islands, 29

December 1962, on Stilbocarpa [Araliaceae] leaf mould.

265. Gamasiphoides femoralis (Banks, 1916)

Cyrtolaelaps femoralis Banks, 1916: 228.

Gamasiphis femoralis.— Womersley, 1942: 156; Lee, 1970: 49; Karg, 1987: 302; Karg,

1990: 333; Karg, 1993b: 173; Karg, 1995: 16.

Gamasiphis (Neogamasiphis) femoralis.— Womersley, 1956b: 518.

Gamasiphoides femoralis.— Lee and Hunter, 1974: 302


TYPE LOCALITY AND HABITAT: Australia, Evandale, Tasmania, in nest of

Rhytidoponera metallica (cited as Ectatomma metallicum) [Hymenoptera:

Formicidae].

300

266. Gamasiphoides gamasiphioides (Sheals, 1962)

Hydrogamasus gamasiphioides Sheals, 1962: 85.

Gamasiphoides gamasiphoides [sic].— Balogh, 1963: 487; Karg, 1976d: 4; Karg and

Schorlemmer, 2011a: 25.

Gamasellus (Hydrogamasellus) gamasiphioides.— Hirschmann, 1966b: 7.

Gamasiphoides gamasiphioides.— Lee, 1970: 63; Karg, 1993a: 51.


TYPE LOCALITY AND HABITAT: Argentina, Villa Mascardi, Parque Nacional Nahuel

Huapi, 4 March 1959, on Nothofagus dombeyi [Nothofagaceae].

267. Gamasiphoides leptogenitalis Karg, 1993

Gamasiphoides leptogenitalis Karg, 1993a: 48.

Gamasiphoides leptogenitalis.— Karg and Schorlemmer, 2011a: 26.





268. Gamasiphoides linealis Karg, 1976

Gamasiphoides linealis Karg, 1976d: 13.

Gamasiphoides linealis.— Karg, 1993a: 51; Karg, 1996: 182; Karg and Schorlemmer, 2011a:

25.


TYPE LOCALITY AND HABITAT: Chile, Caquena, Tarapacá, 28 February 1965, in a moist

soil with manure.

269. Gamasiphoides longocuspis Karg, 1976

Gamasiphoides longocuspis Karg, 1976d: 6.

Gamasiphoides longocuspis.— Karg, 1993a: 50; Karg and Schorlemmer, 2011a: 25.


TYPE LOCALITY AND HABITAT: Chile, Sapahuira [= Zapahuira?] (alt. 3,100 m),

Tarapacá, 1 December 1965, in soil trap.

270. Gamasiphoides longosetis Karg, 1976

301

Gamasiphoides longosetis Karg, 1976d: 5.

Gamasiphoides longisetis [sic].— Karg, 1976d: 5.

Gamasiphoides longosetis.— Karg, 1993a: 50; Karg, 1996: 180; Karg and Schorlemmer,

2011a: 25.


TYPE LOCALITY AND HABITAT: Chile, base of Volcán Guallatiri, 29 November 1965,

under stones.

271. Gamasiphoides longoventris Karg, 1976

Gamasiphoides longoventris Karg, 1976d : 12.

Gamasiphoides longoventris.— Karg, 1993a: 51; Karg and Schorlemmer, 2011a: 25.


TYPE LOCALITY AND HABITAT: Chile, foothill of Volcán Guallatiri, 29 December 1965,

under stones.

272. Gamasiphoides lootsi Halliday, 2005

Gamasiphoides lootsi Halliday, 2005: 42.

TYPE DEPOSITORY: National Collection of Mites, ARC Plant Protection Research

Institute, Pretoria, South Africa.

TYPE LOCALITY AND HABITAT: South Africa, Hermanus, Western Cape, 28 August

1994, on clover Trifulium sp. [Fabaceae] and weeds.

273. Gamasiphoides macquariensis (Hirschmann, 1966)

Gamasellus (Hydrogamasellus) macquariensis Hirschmann, 1966c: 25.

Gamasiphoides macquariensis.— Lee, 1970: 63; Lee and Hunter, 1974: 304; Karg, 1976d: 4;

Karg, 1993a: 50; Karg, 1996: 182; Karg and Schorlemmer, 2011a: 25.



in litter of Pleurophyllum sp. [Asteraceae] in a subplateau.

274. Gamasiphoides octosetae Karg, 1976

Gamasiphoides octosetae Karg, 1976d: 8.

Gamasiphoides octosetae.— Karg, 1993a: 50; Karg, 1996: 182; Karg and Schorlemmer,

2011a: 25.

302


TYPE LOCALITY AND HABITAT: Chile, 5 km from Cóncon, direction from Quintero,

Valparaiso Province, 10 October 1965, on humid rotten reed by the sea.

275. Gamasiphoides postanalis Karg, 1993

Gamasiphoides postanalis Karg, 1993a: 50.

Gamasiphoides postanalis.— Karg and Schorlemmer, 2011a: 26.




a cave.

276. Gamasiphoides procerus Karg and Schorlemmer, 2011

Gamasiphoides procerus Karg and Schorlemmer, 2011a: 24.



TYPE LOCALITY AND HABITAT: Paraguay, 50 m above Acaray waterfall, Ciudad del

Este [cited as Puerto Presidente Stroessner], 1965, in litter and soil from a liana

shrubbery.

277. Gamasiphoides propinquus (Womersley, 1956)

Gamasiphis (Gamasiphoides) propinqua Womersley, 1956b: 528.

Gamasiphoides propinqua.— Ryke, 1962c: 160; Lee, 1970: 63; Karg, 1976d: 4; Karg, 1993a:

52; Karg, 1996: 183; Karg and Schorlemmer, 2011a: 26.

Gamasiphoides propinquus.— Lee, 1973: 8.


TYPE LOCALITY AND HABITAT: Australia, Belair National Park, Belair, South Australia,


278. Gamasiphoides rykei Halliday, 2005

Gamasiphoides rykei Halliday, 2005: 45.

TYPE DEPOSITORY: National Collection of Mites, ARC Plant Protection Research

Institute, Pretoria, South Africa.

303

TYPE LOCALITY AND HABITAT: South Africa, Plettenberg Bay, Western Cape, 18

August 1994, on clover Trifulium sp. [Fabaceae] and weeds in a roadside picnic area.

279. Gamasiphoides setosus Karg, 1976

Gamasiphoides setosus Karg, 1976d: 18.

Gamasiphoides setosus.— Karg, 1993a: 51; Karg and Schorlemmer, 2011a: 26.


TYPE LOCALITY AND HABITAT: Chile, north slope of Cerro San Cristobal [cited as

Mount San Cristobal], Santiago, 26 September 1965, under stones.

Genus Gamasitus Womersley, 1956

Gamasitus Womersley, 1956b: 531 (described in Neoparasitidae Oudemans).

Gamasitus.— Ryke, 1962c: 160; Lee, 1970: 187.

Type species: Gamasitus obscurus Womersley, 1956, by original designation.

280. Gamasitus obscurus Womersley, 1956

Gamasitus obscurus Womersley, 1956b: 531.

Gamasitus obscura.— Womersley, 1956b: 532.

Gamasitus obscurus.— Lee, 1970: 188.


TYPE LOCALITY AND HABITAT: Australia, Mount Wellington, Tasmania, 2 December

1934, on moss.

Genus Geogamasus Lee, 1970

Geogamasus Lee, 1970: 92 (described in Rhodacaridae Oudemans).

Geogamasus.— Karg, 1976a: 23; Karg, 1997a: 68; Karg, 1998: 189; Karg and Schorlemmer,

2011b: 214.

Type species: Geogamasus skoshi Lee, 1970, by original designation.

281. Geogamasus apophyseus Karg, 1976

304

Geogamasus apophyseus Karg, 1976a: 30.

Geogamasus apophyseus.— Karg and Schorlemmer, 2011b: 214.


TYPE LOCALITY AND HABITAT: Chile, Cerillos, Santiago Province, 3 October 1965, in

soil.

282. Geogamasus arcus Karg, 1976

Geogamasus arcus Karg, 1976a: 33.

Geogamasus arcus.— Karg and Schorlemmer, 2011b: 215.


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 23 November 1965, in moist

litter.

283. Geogamasus ardoris Karg, 1976

Geogamasus ardoris Karg, 1976a: 42.

Geogamasus ardoris.— Karg, 1997a: 69; Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, El Manzano, Santiago Province, 30 October

1965, under equine excrements and under stones.

284. Geogamasus bicirrus Karg, 1976

Geogamasus bicirrus Karg, 1976a: 25.

Geogamasus bicirrus.— Karg, 1998: 191; Karg and Schorlemmer, 2011b: 214.


TYPE LOCALITY AND HABITAT: Chile, Arica, Arica Province, 20 November 1965, on

moss.

285. Geogamasus bisetosus Karg, 1976

Geogamasus bisetosus Karg, 1976a: 39.

Geogamasus bisetosus.— Karg and Schorlemmer, 2011b: 214.


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 2 December 1965, in an

irrigation canal.

305

286. Geogamasus brevisetosus Karg, 1997

Geogamasus brevisetosus Karg, 1997a: 70.

Geogamasus brevisetosus.— Karg and Schorlemmer, 2011b: 216.



TYPE LOCALITY AND HABITAT: Chile, Quebrada La Plata, Maipú, Santiago Province,

28 September 1965, from unspecified substrate.

287. Geogamasus brevitondentis Karg, 1998

Geogamasus brevitondentis Karg, 1998: 189.

Geogamasus brevitondentis.— Karg and Schorlemmer, 2011b: 216.



TYPE LOCALITY AND HABITAT: Ecuador, Antisana (alt. 3,300 m), Pichincha Province,

15 April 1989, on dry moss and soil from rocks about 150 m above the level of the

brook, in an earthworm experimental station.

288. Geogamasus cochlearis Karg, 1976

Geogamasus cochlearis Karg, 1976a: 29.

Geogamasus cochlearis.— Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, Sapahuira [= Zapahuira?], Tarapacá, 1 December

1965, in soil trap.

289. Geogamasus coxalis (Sheals, 1962)

Gamasellus coxalis Sheals, 1962: 95.

Gamasellus (Hydrogamasellus) coxalis.— Hirschmann, 1966b: 7.

Geogamasus coxalis.— Lee, 1970: 95.


TYPE LOCALITY AND HABITAT: Argentina, Puerto Blest, Parque Nacional Nahuel

Huapi, 7 March 1959, from unspecified substrate.

290. Geogamasus cuneatus Karg, 1998

Geogamasus cuneatus Karg, 1998: 191.

306

Geogamasus cuneatus.— Karg and Schorlemmer, 2011b: 216.




15 April 1989, on scale-moss from a vertical wall in an earhworm experiment station.


291. Geogamasus delamarei (Sheals, 1962)

Hydrogamasus delamarei Sheals, 1962: 89.

Hydrogamasus delamarea [sic].— Balogh, 1963: 489.

Gamasellus (Hydrogamasellus) delamarei.— Hirschmann, 1966b: 7.

Geogamasus delamarei.— Lee, 1970: 95; Karg, 1976a: 47; Karg and Schorlemmer, 2011b:

215.


TYPE LOCALITY AND HABITAT: Argentina, Parque Nacional Nahuel Huapi, from


292. Geogamasus diffindentis Karg, 1997

Geogamasus diffindentis Karg, 1997a: 68.

Geogamasus diffindentis.— Karg and Schorlemmer, 2011b: 215.





293. Geogamasus fibularis Karg, 1976

Geogamasus fibularis Karg, 1976a: 40.

Geogamasus fibularis.— Karg, 1997a: 70; Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Panama, Cristóbal, [Colon], 12 September 1965, under

stones and branches in the coast.

294. Geogamasus filicuspidis Karg, 1976

Geogamasus filicuspidis Karg, 1976a: 24.

307


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 23 November 1965, in moist

litter.


295. Geogamasus flagellatus Karg, 1976

Geogamasus flagellatus Karg, 1976a: 37.

Geogamasus flagellatus.— Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 2 December 1965, in soil trap

in a dry river bed.

296. Geogamasus foliaceus Karg, 1976

Geogamasus foliaceus Karg, 1976a: 44.

Geogamasus foliaceus.— Karg, 1997a: 69; Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, La Unión, [Ranco Province], 26 October 1965,

in moist soil in a deep valley.


297. Geogamasus forcipis Karg, 1976

Geogamasus forcipis Karg, 1976a: 33.

Geogamasus forcipis.— Karg and Schorlemmer, 2011b: 215.


TYPE LOCALITY AND HABITAT: Chile, Campamento Experimental Misituni, Tarapacá, 1

December 1965, in soil trap in a wet place.

298. Geogamasus fornix Halliday, 2001

Geogamasus fornix Halliday, 2001: 305.

Geogamasus fornix.— Karg and Schorlemmer, 2011b: 215.


TYPE LOCALITY AND HABITAT: Australia, Bow Cave, Jenolan Caves, New South

Wales, 14 May 1988, in decomposing leaf litter along the sides of subterranean

streams inside the cave.

308

299. Geogamasus furcatius Karg, 1976

Geogamasus furcatius Karg, 1976a: 34.

Geogamasus furcatius.— Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, Concón, Valparaiso Province, 10 October 1965,

on herbs.


300. Geogamasus howardi Lee, 1970

Geogamasus howardi Lee, 1970: 96.

Geogamasus howardi.— Lee, 1973: 12; Karg, 1976a: 46; Karg and Schorlemmer, 2011b:

215.


TYPE LOCALITY AND HABITAT: Australia, Mount Burr, South Australia, 30 May 1966,

in plant litter and soil in a Pinus radiata [Pinaceae] forest.

301. Geogamasus incisus Karg, 1976

Geogamasus incisus Karg, 1976a: 24.

Geogamasus incisus.— Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, Pelchuquin, Valdivia Province, 25 October 1965,

in moist soil.

302. Geogamasus levispiritus Karg, 1998

Geogamasus levispiritus Karg, 1998: 191.

Geogamasus levispiritus.— Karg and Schorlemmer, 2011b: 216.




Pichincha Province, 14 April 1989, in felt-like debris from under dicotyledon plant

with big leaves and lilac flowers.

303. Geogamasus longifolii Karg and Schorlemmer, 2011

309

Neogamasellevans longifolii Karg and Schorlemmer, 2011b: 214.



TYPE LOCALITY AND HABITAT: Venezuela, near Maracay, 1973, in detritus and soil in a

forest next to the park of the University.

304. Geogamasus longisetosus Karg, 1976

Geogamasus longisetosus Karg, 1976a: 35.

Geogamasus longisetosus.— Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Chile, Campamento Experimental Misituni, Tarapacá,

25 November 1965, in peaty soil with gravel.

305. Geogamasus minimus Lee, 1973

Geogamasus minimus Lee, 1973: 12.

Geogamasus minimus.— Karg, 1976a: 46; Karg, 1997a: 68; Karg and Schorlemmer, 2011b:

215.



Adelaide, South Australia, 1968-1969, on moss or plant litter.

306. Geogamasus monocuspidis Karg, 1976

Geogamasus monocuspidis Karg, 1976a: 32.

Geogamasus monocuspidis.— Karg and Schorlemmer, 2011b: 215.


TYPE LOCALITY AND HABITAT: Chile, Farellones, Santiago Province, 6 October 1965,

on wet grass.

307. Geogamasus pentaspinosus Karg, 1979

Geogamasus pentaspinosus Karg, 1979: 212.


TYPE LOCALITY AND HABITAT: Argentina, Cerro Piltriquitron, El Bolson, Rio Negro

Province, 24 April 1961, from a pasture at the edge of a Libocedrus [Cupressaceae]

forest.

310

308. Geogamasus pisciformis Karg, 1997

Geogamasus pisciformis Karg, 1997a: 70.

Geogamasus pisciformis.— Karg and Schorlemmer, 2011b: 216.




on moist moss.

309. Geogamasus pugionis Karg, 1976

Geogamasus pugionis Karg, 1976a: 39.

Geogamasus pugionis.— Karg, 1997a: 69; Karg and Schorlemmer, 2011b: 215.



in leaf litter under bushes on rocks.

310. Geogamasus reticulatus Karg, 1976

Geogamasus reticulatus Karg, 1976a: 36.

Geogamasus reticulatus.— Karg and Schorlemmer, 2011b: 216.



in litter under bushes.

311. Geogamasus skoshi Lee, 1970

Geogamasus skoshi Lee, 1970: 95.

Geogamasus skoshi.— Karg, 1976a: 47; Karg and Schorlemmer, 2011b: 216.


TYPE LOCALITY AND HABITAT: Argentina, Universidad Nacional de Tucumán,

Tucumán, November 1957, in soil.

312. Geogamasus trispinosus Karg, 1976

Geogamasus trispinosus Karg, 1976a: 27.

Geogamasus trispinosus.— Karg, 1998: 191; Karg and Schorlemmer, 2011b: 214.


311

TYPE LOCALITY AND HABITAT: Chile, Sapahuira [= Zapahuira?], Tarapacá, 24

November 1965, in peaty soil.

313. Geogamasus tuberosus Karg, 1976

Geogamasus tuberosus Karg, 1976a: 29.

Geogamasus tuberosus.— Karg, 1998: 191; Karg and Schorlemmer, 2011b: 214.



under stones.

Genus Heterogamasus Trägårdh, 1907

Heterogamasus Trägårdh, 1907: 2 (family not specified in the description).

Heterogamasus.— Vitzthum, 1931a: 142; Lee, 1970: 156; Karg, 1977: 334.

Heterogamasus (Heterogamasus).— Lee, 1967: 500.

Type species: Heterogamasus claviger Trägårdh, 1907, by subsequent monotypy.

314. Heterogamasus calcarellus Lee, 1967

Heterogamasus (Heterogamasus) calcarellus Lee, 1967: 505.

Heterogamasus calcarellus.— Lee, 1970: 157; Lee and Hunter, 1974: 321.


TYPE LOCALITY AND HABITAT: Chile, Cordillera Darwin, Isla Grande de Tierra del

Fuego, 27 February and 1 March 1962, in litter of a Nothofagus sp. [Nothofagaceae]

forest.

315. Heterogamasus claviger Trägårdh, 1907

Heterogamasus claviger Trägårdh, 1907: 2.

Heterogamasus (Heterogamasus) claviger.— Lee, 1967: 501.

Heterogamasus claviger.— Lee, 1970: 157; Lee and Hunter, 1974: 321.

TYPE DEPOSITORY: Naturhistorika Riksmuseum, Stockholm, Sweden.

TYPE LOCALITY AND HABITAT: Falkland Islands [South Atlantic Ocean], east of Port

Stanley, 25 February 1902, under stones.


312

316. Heterogamasus euarmatus Karg, 1977

Heterogamasus euarmatus Karg, 1977: 337.


TYPE LOCALITY AND HABITAT: Chile, km 636 of the Panamerican Road, near

Collipulli, Malleco Province, 27 October 1965, under manure.


317. Heterogamasus inermus Karg, 1977

Heterogamasus inermus Karg, 1977: 335.



November 1965, in moist soil.

318. Heterogamasus spinosissimus (Balogh, 1963)

Gamasellus (?) spinosissimus [sic] Balogh, 1963: 489.

Heterogamasus (Heterogamasus) spinosissimus.— Lee, 1967: 501.

Heterogamasus spinosissimus.— Lee, 1970: 157.


TYPE LOCALITY AND HABITAT: Argentina, Cerro Piltriquitron (alt. 1,150 m), El Bolson,

Rio Negro Province, 16 November 1961, in litter from Maytenus boaria

[Celastreaceae] forest with Berberis darwinii [Berberidaceae] and Nothofagus

antarctica [Nothofagaceae], near a spring.

Genus Heydeniella Richters, 1907

Heydeniella Richters, 1907: 281 (described in Parasitidae Oudemans).

Heydeniella.— Lee, 1970: 96.

Heydeniella crozentensis-complex.— Lee, 1970: 97.

Heydeniella (in part).— Karg, 1976c: 193.

Type species: Heydeniella crozetensis Richters, 1907, by subsequent monotypy.

319. Heydeniella crozetensis Richters, 1907

313

Heydeniella crozetensis Richters, 1907: 281.

Gamasiphis crozetensis.— Trägårdh, 1907: 10.

Heydeniella crozetensis.— Lee, 1970: 100; Lee and Hunter, 1974: 312; Karg, 1976c: 199.

TYPE DEPOSITORY: According to Lee (1970): 100, specimen probably destroyed.

TYPE LOCALITY AND HABITAT: Crozet Islands [southern Indian Ocean], Île de la

Possession, from unspecified substrate.

320. Heydeniella leei Karg, 1976

Heydeniella leei Karg, 1976c: 194.



under manure.

321. Heydeniella loricata (Trägårdh, 1907)

Gamasiphis loricatus Trägårdh, 1907: 10.

Heydeniella loricata.— Lee, 1970: 101; Lee and Hunter, 1974: 312; Karg, 1976c: 199.


TYPE LOCALITY AND HABITAT: Falkland Islands [South Atlantic Ocean], east of Port

Stanley, 25 February 1902, under stones.


322. Heydeniella sherrae Lee and Hunter, 1974

Heydeniella sherrae Lee and Hunter, 1974: 312.


TYPE LOCALITY AND HABITAT: New Zealand, Rose Island, Auckland Islands, 8 January

1963, in a rabbit burrow.

323. Heydeniella womersleyi Lee and Hunter, 1974

Heydeniella womersleyi Lee and Hunter, 1974: 314.


TYPE LOCALITY AND HABITAT: New Zealand, Ocean Island, Auckland Islands, 29

December 1962, on Stilbocarpa [Araliaceae] leaf mould.

314

Genus Hiniphis Lee, 1970

Hiniphis Lee, 1970: 157 (described in Rhodacaridae Oudemans).

Type species: Hiniphis hinnus Lee, 1970, by original designation.

324. Hiniphis bipala Lee, 1973

Hiniphis bipala Lee, 1973: 24.



Adelaide, South Australia, 5 or 12 August 1968, in plant litter.

325. Hiniphis hinnus Lee, 1970

Hiniphis hinnus Lee, 1970: 159.


TYPE LOCALITY AND HABITAT: Australia, near Beech Forest, Otway Ranges, Victoria,

9 December 1965, on moss in a Pinus radiata [Pinaceae] plantation.

Genus Hydrogamasellus Hirschmann, 1966

Gamasellus (Hydrogamasellus) Hirschmann, 1966b: 6 (described in Eugamasidae

Hirschmann).

Hydrogamasellus.— Lee, 1970: 110; Karg, 1976b: 37; Karg, 1997a: 64; Karg and


Type species: Hydrogamasus antarcticus Trägårdh, 1907, by original designation.

326. Hydrogamasellus antarcticus (Trägårdh, 1907)

Hydrogamasus antarcticus Trägårdh, 1907: 12.

Gamasellus (Hydrogamasellus) antarcticus.— Hirschmann, 1966b: 7; Hunter, 1970: 61.

Hydrogamasus antarcticus.— Hunter, 1967a: 35; Hunter, 1967b: 31.

Hydrogamasellus antarcticus.— Lee, 1970: 115; Lee and Hunter, 1974: 316; Karg, 1976b:

53; Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.


315

TYPE LOCALITY AND HABITAT: Paulet Island [Antarctic Peninsula], 15 January 1902, in

wet moss.

NOTE: Specimens reported by Womersley (1937): 17 as Hydrogamasus antarcticus were

described as Parasitiphis aurora Lee, 1970: 123.

327. Hydrogamasellus antennatus Karg, 1976

Hydrogamasellus antennatus Karg, 1976b: 41.

Hydrogamasellus antennatus.— Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.



on moss.

328. Hydrogamasellus armatissimus Karg, 1976

Hydrogamasellus armatissimus Karg, 1976b: 37.

Hydrogamasellus armatissimus.— Karg, 1997a: 64; Karg and Schorlemmer, 2009: 82.



under manure.

329. Hydrogamasellus avium Karg, 1976

Hydrogamasellus avium Karg, 1976b: 49.

Hydrogamasellus avium.— Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.



under stones.

330. Hydrogamasellus brevipilus Karg, 1976

Hydrogamasellus brevipilus Karg, 1976b: 52.


TYPE LOCALITY AND HABITAT: Chile, road to Valdivia, La Unión, [Ranco Province],

26 October 1965, between litter and pieces of wood.


331. Hydrogamasellus brevispiritus Karg, 1998

316

Hydrogamasellus brevispiritus Karg, 1998: 191.

Hydrogamasellus brevispiritus.— Karg and Schorlemmer, 2009: 82.




Pichincha Province, 14 April 1989, on moss from the soil.

332. Hydrogamasellus bullarmatus Karg and Schorlemmer, 2009

Hydrogamasellus bullarmatus Karg and Schorlemmer, 2009: 81.



TYPE LOCALITY AND HABITAT: Ecuador, near Paramo, 1973, in detritus, litter and

moss.


333. Hydrogamasellus calculus Karg, 1997

Hydrogamasellus calculus Karg, 1997a: 67.

Hydrogamasellus calculus.— Karg and Schorlemmer, 2009: 83.




under stones in a stream.

334. Hydrogamasellus cicatricosus Karg, 1976

Hydrogamasellus cicatricosus Karg, 1976b: 44.

Hydrogamasellus cicatricosus.— Karg, 1997a: 67; Karg and Schorlemmer, 2009: 82.


TYPE LOCALITY AND HABITAT: Chile, foothill of Volcán Guallatiri, 29 November 1965,

under stones.

335. Hydrogamasellus coleoptratus (Berlese, 1888)

Hypoaspis coleoptratus Berlese, 1888: 198.

Ologamasellus coleoptratus.— Berlese, 1916a: 162.

317

Hydrogamasellus coleoptratus.— Lee, 1970: 116; Karg, 1976b: 53; Karg, 1997a: 67; Karg

and Schorlemmer, 2009: 83.


TYPE LOCALITY AND HABITAT: Argentina, Buenos Aires, under tree bark.

336. Hydrogamasellus conchatus Karg and Schorlemmer, 2009

Hydrogamasellus conchatus Karg and Schorlemmer, 2009: 79.



TYPE LOCALITY AND HABITAT: Venezuela, University, Maracay, 1973, in forest litter

and humus.

337. Hydrogamasellus contingentis Karg, 1997

Hydrogamasellus contingentis Karg, 1997a: 67.

Hydrogamasellus contingentis.— Karg and Schorlemmer, 2009: 82.



TYPE LOCALITY AND HABITAT: Chile, road to Quintero, Concón, Valparaiso Province,

10 October 1965, under tree bark lying on the shore in a meadow near a creek.

338. Hydrogamasellus crozetensis (Richters, 1907)

Gamasellus crozetensis Richters, 1907: 279.

Cyrtolaelaps (Gamasellus) crozetensis – Ryke, 1962b: 51.

Hydrogamasellus richtersi Lee, 1970: 115 [objective synonym — unjustified replacement

name for Gamasellus crozetensis Richters, 1907].

Hydrogamasellus crozetensis.— Lee and Hunter, 1974: 316.

Hydrogamasellus richtersi.— Karg, 1976b: 52; Karg, 1997a: 65; Karg, 1998: 192; Karg and


TYPE DEPOSITORY: According to Lee (1970): 100, specimen probably destroyed.


Possession, on moss.

339. Hydrogamasellus gaussi Lee, 1970

Hydrogamasellus gaussi Lee, 1970: 115.

318

Hydrogamasellus gaussi.— Lee and Hunter, 1974: 316; Karg, 1976b: 53; Karg, 1997a: 65;


TYPE DEPOSITORY: According to Lee (1970): 100, specimen probably destroyed..


Possession, from unspecified substrate.

NOTE: Hydrogamasellus gaussi is a replacement name for Neoparasitus crozetensis Richters,

1907: 279 which became a junior homonym of Hydrogamasellus crozetensis

(Richters, 1907): 279, originally described (as Gamasellus crozetensis) on the same

page, but immediately before N. crozetensis.

340. Hydrogamasellus iaculi Karg and Schorlemmer, 2009

Hydrogamasellus iaculi Karg and Schorlemmer, 2009: 80.



TYPE LOCALITY AND HABITAT: Brazil, Aracariguana [= Araçariguama, São Paulo

State?], 1970, in humus.

341. Hydrogamasellus longopilus Karg, 1976

Hydrogamasellus longopilus Karg, 1976b: 50.

Hydrogamasellus longopilus.— Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.


TYPE LOCALITY AND HABITAT: Chile, road to Valdivia, La Unión, [Ranco Province],

26 October 1965, between litter and pieces of wood.

342. Hydrogamasellus multospinosus Karg, 1976

Hydrogamasellus multospinosus Karg, 1976b: 47.

Hydrogamasellus multospinosus.— Karg, 1997a: 67; Karg and Schorlemmer, 2009: 83.


TYPE LOCALITY AND HABITAT: Chile, Cerro El Roble, Cordillera de la Costa, 29

September 1965, in litter.

343. Hydrogamasellus nasutus Karg, 1976

Hydrogamasellus nasutus Karg, 1976b: 45.

Hydrogamasellus nasutus.— Karg, 1997a: 67; Karg and Schorlemmer, 2009: 83.

319



in litter.

344. Hydrogamasellus racovitzai (Trouessart, 1903)

Gamasus racovitzai Trouessart, 1903: 8.

Gamasellus racovitzai.— Trägårdh, 1907: 7; Hunter, 1970: 61.

Gamasellus (Digamasellus) racovitzai.— Berlese, 1917: 5.

Cyrtolaelaps (Gamasellus) racovitzai.— Ryke, 1962b: 50; Hunter, 1967a: 35; Hunter, 1967b:

33.

Hydrogamasellus racovitzai.— Lee, 1970: 116; Lee and Hunter, 1974: 316; Karg, 1976b: 52;

Karg, 1997a: 65; Karg, 1998: 192; Karg and Schorlemmer, 2009: 82.

Zercon tuberculatus Trägårdh, 1907: 17 [junior synonymy by Hunter, 1967a: 35].

TYPE DEPOSITORY: H. racovitzai: Muséum National d‘Histoire Naturelle, Paris, France; Z.

tuberculatus: Naturhistorika Riksmuseum, Stockholm, Sweden.

TYPE LOCALITY AND HABITAT: H. racovitzai: Gerlache Strait [Antarctic Peninsula], on

moss; Z. tuberculatus: Graham Land [Antarctic Peninsula], Mount Bransfield, 7

December 1902, on moss.

344a. Hydrogamasellus racovitzai neorcadensis Trouessart, 1912

Gamasellus racovitzai neo-orcadensis Trouessart, 1912: 86.

Cyrtolaelaps (Gamasellus) racovitzai neo-orcadensis.— Ryke, 1962b: 50.


TYPE LOCALITY AND HABITAT: Laurie Island [Antarctic Circle], 18 December

1903, on moss on a cliff.

NOTE: Lee (1970): 117 suggested that this subspecies is the same as the nominal

subspecies

345. Hydrogamasellus stipulae Karg, 1998

Hydrogamasellus stipulae Karg, 1998: 191.

Hydrogamasellus stipulae.— Karg and Schorlemmer, 2009: 82.



320


15 April 1989, on dry moss and soil from rocks, ca. 150 m above the level of a brook

in an earthworm experiment station.

346. Hydrogamasellus striatus (Sheals, 1962)

Hydrogamasus striatus Sheals, 1962: 87.

Hydrogamasus striatus.— Balogh, 1963: 489.

Gamasellus (Hydrogamasellus) striatus.— Hirschmann, 1966b: 7.

Hydrogamasellus striatus.— Lee, 1970: 115; Karg, 1976b: 52; Karg, 1997a: 64; Karg and





347. Hydrogamasellus topali (Balogh, 1963)

Hydrogamasus topali Balogh, 1963: 489.

Gamasellus (Hydrogamasellus) topali.— Hirschmann, 1966b: 7.

Hydrogamasellus topali.— Lee, 1970: 115; Karg, 1976b: 53; Karg, 1997a: 65; Karg and



TYPE LOCALITY AND HABITAT: Argentina, Cerro Piltriquitron (alt. 360 m), El Bolson,

Rio Negro Province, 30 March 1961, in litter from a Libocedrus [Cupressaceae],

Lomatia [Proteaceae] and Aristotelia [Elaeocarpaceae] stand.

348. Hydrogamasellus ubatubaensis (Hirschmann, 1966)

Gamasellus (Hydrogamasellus) ubatubaensis Hirschmann, 1966c: 31.

Hydrogamasellus ubatubaensis.— Lee, 1970: 115.


TYPE LOCALITY AND HABITAT: Brazil, Ubatuba, [São Paulo State], on rocky coast.

321

Genus Hydrogamasus Berlese, 1892

Hydrogamasus Berlese, 1892c: 5 [presumed to be described in Parasitidae Oudemans (also


Hydrogamasus.— Vitzthum, 1931a: 142; Vitzthum, 1943: 756; Baker and Wharton, 1952:

73; Ryke, 1962c: 160; Hirschmann, 1966b: 7; Lee, 1970: 64.

Hydrogamasus (Hydrogamasus).— Hirschmann, 1966b: 10.

Type species: Gamasus littoralis G. Canestrini and R. Canestrini, 1881, by subsequent

designation (70.3 of International Code of Zoological Nomenclature) by Lee

(1970): 64, in replacement of Gamasus salinus Laboulbène, 1851, which had

been transferred to Pergamasus (Parasitidae) by Oudemans (1936): 163, but

which had been designated as the type species of Hydrogamasus by Vitzthum

(1943): 756; see explanation under Hydrogamasus littoralis.

349. Hydrogamasus giardi (Berlese and Trouessart, 1889)

Sejus giardi Berlese and Trouessart, 1889: 4.

Gamasus giardi.— Moniez, 1889: 193.

Hydrogamasus giardi.— Berlese, 1892c: 5; Berlese, 1892f: 72; Halbert, 1915: 65; Halbert,

1920: 121; Lee, 1970: 68; Karg, 1971: 352.

Hydrogamasus (Hydrogamasus) giardi.— Hirschmann, 1966c: 27.

TYPE DEPOSITORY: Muséum National d‘Histoire Naturelle, Paris, France.

TYPE LOCALITY AND HABITAT: France, Wimereux, [Pas-de-Calais], on Balanus

balanoides [Crustacea: Archaeobalanidae].

350. Hydrogamasus kensleri Luxton, 1967

Hydrogamasus kensleri Luxton, 1967: 76.

Hydrogamasus kensleri.— Lee, 1970: 69; Lee and Hunter, 1974: 306.

TYPE DEPOSITORY: Dominion Museum, Wellington, New Zealand.

TYPE LOCALITY AND HABITAT: New Zealand, Waimaru, Takatu Peninsula, North

Auckland, December 1964, in rock fissures on the upper shore.

351. Hydrogamasus littoralis (G. Canestrini and R. Canestrini, 1881)

Gamasus litoralis G. Canestrini and R. Canestrini, 1881: 1078.

322

Gamasus littoralis.— G. Canestrini and R. Canestrini, 1882: 48; G. Canestrini, 1885: 72;

Moniez, 1889: 186.

Hydrogamasus littoralis.— Berlese, 1892c: 6; Berlese, 1892f: 72; Halbert, 1920: 120; Lee,

1970: 68; Karg, 1993c: 374.


TYPE LOCALITY AND HABITAT: Italy, Lido de Venezia, under stones washed by salt

water in a lagoon.

NOTE1: In the original description, the species name was spelled as Gamasus litoralis. G.

Canestrini and R. Canestrini (1882): 537 unjustifiably (Article 33.3 of the fourth

edition of the International Code of Zoological Nomenclature) changed it to Gamasus

littoralis. All subsequent publications on the species used G. Canestrini and R.

Canestrini (1882)´s proposed spelling. Thus, because of the prevailing usage, G.

Canestrini and R. Canestrini (1882) spelling is maintained in this catalog.

NOTE2: Gamasus salinus Laboulbène, 1851: 297 had been regarded as senior synonym of

Gamasus littoralis G. Canestrini and R. Canestrini, 1881 by Oudemans (1902b): 286,

but Oudemans (1936): 163 revoked that synonymy and transferred the former species

to Pergamasus (Parasitidae).

352. Hydrogamasus vitzthumi Hirschmann, 1966

Hydrogamasus (Hydrogamasus) vitzthumi Hirschmann, 1966c: 25.

Hydrogamasus vitzthumi.— Lee, 1970: 68.


TYPE LOCALITY AND HABITAT: Italy, Porto di Duino, between algae on the jetty.

Genus Laelaptiella Womersley, 1956

Laelaptiella Womersley, 1956b: 512 (described in Ascaidae Oudemans).

Gamasiphis (Laelaptiella).— Domrow, 1957: 202.

Laelaptiella.— Ryke, 1962c: 160; Lee, 1970: 70; Karg, 1976d: 21; Karg, 1993a: 54.

Type species: Laelaptiella anomala Womersley, 1956, by original designation.

353. Laelaptiella anomala Womersley, 1956

Laelaptiella anomala Womersley, 1956b: 512.

323

Gamasiphis (Laelaptiella) anomala.— Domrow, 1957: 203.

Laelaptiella anomala.— Ryke, 1962c: 160; Lee, 1970: 72; Karg, 1976d: 21; Karg, 1993a: 56.


TYPE LOCALITY AND HABITAT: Australia, Adelaide, South Australia, June 1935, from


354. Laelaptiella cultrata Karg, 1993

Laelaptiella cultrata Karg, 1993a: 54.




a cave.

355. Laelaptiella eupodalia Karg, 1996

Laelaptiella eupodalia Karg, 1996: 180.



TYPE LOCALITY AND HABITAT: New Caledonia, Hienghène, 1977, in litter.

356. Laelaptiella mackerrasae (Domrow, 1957)

Gamasiphis (Laelaptiella) mackerrasae Domrow, 1957: 202.

Laelaptiella mackerrasae.— Lee, 1970: 72; Karg, 1976d: 22; Karg, 1993a: 54.

TYPE DEPOSITORY: Queensland Museum, Brisbane, Australia.

TYPE LOCALITY AND HABITAT: Australia, Low Isles, Great Barrier Reef, Queensland,

19 August 1954, on leaf mould on cay.

357. Laelaptiella media Karg, 1976

Laelaptiella media Karg, 1976d: 22.

Laelaptiella media.— Karg, 1993a: 56.



November 1965, in litter.

324

Genus Laelogamasus Berlese, 1905

Gamasus (Laelogamasus) Berlese, 1905a: 167 [presumed to be described in Parasitidae


Gamasus (Laelogamasus).— Berlese, 1906: 113.

Laelogamasus.— Vitzthum, 1931a: 142; Oudemans, 1939: 21; Vitzthum, 1943: 758; Baker

and Wharton, 1952: 64; Lee, 1970: 159.

Type species: Gamasus (Laelogamasus) simplex Berlese, 1905, by monotypy.

358. Laelogamasus simplex (Berlese, 1905)

Gamasus (Laelogamasus) simplex Berlese, 1905a: 167.

Gamasus (Laelogamasus) simplex.— Berlese, 1906: 115.

Laelogamasus simplex.— Vitzthum, 1931a: 142; Baker and Wharton, 1952: 64; Lee, 1970:

160.

Cyrtolaelaps (Gamasellus) simplex.— Ryke, 1962b: 51.


TYPE LOCALITY AND HABITAT: Indonesia, Bogor [cited as Buitenzorg], Java, from


Genus Litogamasus Lee, 1970

Litogamasus Lee, 1970: 160 (described in Rhodacaridae Oudemans).

Type species: Cyrtolaelaps setosus Kramer, 1898, by original designation.

359. Litogamasus falcipes Lee and Hunter, 1974

Litogamasus falcipes Lee and Hunter, 1974: 322.


TYPE LOCALITY AND HABITAT: New Zealand, Campbell Island, 1967, on rocky shore.

360. Litogamasus setosus (Kramer, 1898)

Cyrtolaelaps setosus Kramer, 1898a: 22.

Cyrtolaelaps (Cyrtolaelaps) setosus.— Ryke, 1962b: 10.

Litogamasus setosus.— Lee, 1970: 163; Lee and Hunter, 1974: 322.

325

TYPE DEPOSITORY: Zoologisches Museum, Hamburg, Germany.

TYPE LOCALITY AND HABITAT: Argentina, Ushuaia, Isla Grande de Tierra del Fuego,

27 and 30 October 1892 and 9 December 1892, under stones at the mouth of the creek

above the flood line; in seashore; lowest low tide beach.

Genus Neogamasellevans Loots and Ryke, 1967

Neogamasellevans Loots and Ryke, 1967b: 13 (described in Rhodacaridae Oudemans).

Neogamasellevans.— Lee, 1970: 117; Karg, 1975: 117; Karg and Schorlemmer, 2009: 70;

Karg and Schorlemmer, 2011b: 211.

Type species: Neogamasellevans preendopodalis Loots and Ryke, 1967, by original

designation.

361. Neogamasellevans ammonis Karg and Schorlemmer, 2009

Neogamasellevans ammonis Karg and Schorlemmer, 2009: 71.



TYPE LOCALITY AND HABITAT: Paraguay, Botanical Garden, Asunción, 1966, in litter

and soil from a jungle.

362. Neogamasellevans armata Karg and Schorlemmer, 2009

Neogamasellevans armata Karg and Schorlemmer, 2009: 71.



TYPE LOCALITY AND HABITAT: Paraguay, Ciudad del Este [cited as Puerto Presidente

Stroessner], 1965, in litter of a thick jungle.

363. Neogamasellevans brevisetosa Karg, 1997

Neogamasellevans brevisetosa Karg, 1997a: 71.

Neogamasellevans brevisetosa.— Karg and Schorlemmer, 2009: 73.



326

TYPE LOCALITY AND HABITAT: Chile, Región de Tarapacá, 25 November 1965, in soil

trap.

364. Neogamasellevans brevitremata Karg, 1975

Neogamasellevans brevitremata Karg, 1975: 125.

Neogamasellevans brevitremata.— Karg and Schorlemmer, 2009: 73.


Germany.


29 November 1965, on slimy algae sample from a small puddle with nematodes.

365. Neogamasellevans dentata Karg, 1975

Neogamasellevans dentata Karg, 1975: 120.

Neogamasellevans dentata.— Karg and Schorlemmer, 2009: 74.


Germany.


December 1965, in soil trap under bushes in the northern highlands.

366. Neogamasellevans furcatus Karg and Schorlemmer, 2011

Neogamasellevans furcatus Karg and Schorlemmer, 2011b: 211.



TYPE LOCALITY AND HABITAT: Venezuela, near Maracay, 30 March 1973, in detritus

and soil in a forest next to the park of the University.

367. Neogamasellevans gracilis Karg and Schorlemmer, 2011

Neogamasellevans gracilis Karg and Schorlemmer, 2011b: 213.



TYPE LOCALITY AND HABITAT: Venezuela, near Maracay, 30 March 1973, in detritus

and soil in a forest next to the park of the University.

368. Neogamasellevans longipes Karg and Schorlemmer, 2009

327

Neogamasellevans longipes Karg and Schorlemmer, 2009: 70.



TYPE LOCALITY AND HABITAT: Brazil, near Barueri, [São Paulo State], 1970, in humus.

369. Neogamasellevans longocalcaris Karg, 1975

Neogamasellevans longocalcaris Karg, 1975: 118.

Neogamasellevans longocalcaris.— Karg and Schorlemmer, 2009: 74.


Germany.


December 1965, in soil trap in a valley leading to the banks of the Lauca River in the

northern highlands.

370. Neogamasellevans macrochela Karg, 1975

Neogamasellevans macrochela Karg, 1975: 127.

Neogamasellevans macrochela.— Karg and Schorlemmer, 2009: 73.


Germany.

TYPE LOCALITY AND HABITAT: Chile, Sapahuira [= Zapahuira?] (alt. 3,100 m),

Tarapacá, 1 December 1965, in soil trap.

371. Neogamasellevans ornata Karg, 1975

Neogamasellevans ornata Karg, 1975: 130.

Neogamasellevans ornata.— Karg, 1997a: 72; Karg and Schorlemmer, 2009: 73.


Germany.



372. Neogamasellevans preendopodalis Loots and Ryke 1967

Neogamasellevans preendopodalis Loots and Ryke 1967b: 14.

Gamasellus (Hydrogamasellus) preendopodalis.— Hirschmann, 1968: 23.

Neogamasellevans praeendopodalis [sic].— Karg, 1975: 134.

328

Neogamasellevans preendopodalis.— Karg and Schorlemmer, 2009: 73.


TYPE LOCALITY AND HABITAT: Argentina, in soil litter.

373. Neogamasellevans serrata Karg, 1975

Neogamasellevans serrata Karg, 1975: 131.

Neogamasellevans serrata.— Karg and Schorlemmer, 2009: 73.


Germany.


29 November 1965, in nematode samples from llama dung, near a water puddle.

374. Neogamasellevans xylebori Van Daele, 1976

Neogamasellevans xylebori Van Daele, 1976: 330.

TYPE DEPOSITORY: Zoology Museum, Faculty of Sciences, Ghent University, Ghent,

Belgium.

TYPE LOCALITY AND HABITAT: Belgium, Merebelke, near Ghent, 18 February 1976, on

stems of Dracaena fragrans var. massangeana [Ruscaceae] cultivated under glass.

Genus Notogamasellus Loots and Ryke, 1965

Notogamasellus Loots and Ryke, 1965: 465 (described in Rhodacaridae Oudemans).

Notogamasellus (Notogamasellus).— Loots and Ryke, 1965: 465; Lee, 1970: 164.

Type species: Notogamasellus (Notogamasellus) vandenbergi Loots and Ryke, 1965,

by original designation.

375. Notogamasellus vandenbergi Loots and Ryke, 1965

Notogamasellus (Notogamasellus) vandenbergi Loots and Ryke, 1965: 467.

Notogamasellus (Notogamasellus) vandenbergi.— Lee, 1970: 164.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, Limpopo [cited as

Transvaal], 1962 and 1963, in soil of the indigenous forest.

329

Genus Ologamasus Berlese, 1888

Gamasus (Ologamasus) Berlese, 1888: 194 [presumed to be described in Parasitidae


Gamasus (Hologamasus).— Berlese, 1892f: 62.

Gamasus (Ologamasus).— Berlese, 1906: 242.

Ologamasus.— Berlese, 1913b: 202; Berlese, 1914: 137; Vitzthum, 1931a: 142; Baker and

Wharton, 1952: 73; Ryke, 1962c: 160; Lee, 1970: 84; Karg, 1976c: 185; Silva, Moraes

and Krantz, 2007: 61; Karg and Schorlemmer, 2009: 74.

Ologamasus (Ologamasus).— Vitzthum, 1943: 756; Baker and Wharton, 1952: 73.

Type species: Gamasus aberrans Berlese, 1888, by subsequent designation by

Oudemans (1936): 165 in replacement of Gamasus calcaratus Koch, the type

species of Holoparasitus Oudemans, 1936: 164 (mentioned in Parasitidae

Oudemans).

Ologamasellus Berlese, 1914: 139 [junior synonymy by Oudemans, 1939: 22].

Ologamasellus.— Berlese, 1916a: 162.

Ologamaselus [sic].— Vitzthum, 1931a: 142.

Type species: Gamasus aberrans Berlese, 1888, by original designation.

376. Ologamasus aberrans (Berlese, 1888)

Gamasus aberrans Berlese, 1888: 194.

Gamasus (Hologamasus) aberrans.— Berlese, 1892f: 62.

Gamasellus aberrans.— Berlese, 1906: 274.

Ologamasellus aberrans.— Berlese, 1914: 139.

Ologamasus aberrans.— Oudemans, 1936: 165; Vitzthum, 1943: 756; Ryke, 1962c: 160;

Lee, 1970: 89; Karg, 1976c: 186; Silva, Moraes and Krantz, 2007: 67; Karg and



TYPE LOCALITY AND HABITAT: Brazil, Matto-grosso [= Mato Grosso? or Mato Grosso

do Sul?], under tree bark.

377. Ologamasus brevidigitus Karg and Schorlemmer, 2009

330

Ologamasus brevidigitus Karg and Schorlemmer, 2009: 74.




378. Ologamasus brevisetosus Karg and Schorlemmer, 2009

Ologamasus brevisetosus Karg and Schorlemmer, 2009: 78.



TYPE LOCALITY AND HABITAT: Ecuador, Cotopaxi, 1973, in litter, moss and detritus of

a paramo [= páramo, Spanish term for a Neotropical habitat of high altitude].

379. Ologamasus cananeiae Silva, Moraes and Krantz, 2007

Ologamasus cananeiae Silva, Moraes and Krantz, 2007: 61.


Agricultura ―Luiz de Queiroz‖, Universidade de São Paulo, Piracicaba, Brazil.

TYPE LOCALITY AND HABITAT: Brazil, Cananéia, São Paulo State, 11 October 2000, in

forest litter.

380. Ologamasus cavei Sheals, 1962

Ologamasus cavei Sheals, 1962: 92.

Hydrogamasellus cavei.— Lee, 1970: 116; Karg, 1976b: 52; Karg, 1997a: 65; Karg and





381. Ologamasus distorta (Karg, 1976)

Heydeniella distorta Karg, 1976c: 197.


TYPE LOCALITY AND HABITAT: Chile, on the road to Valdivia, La Unión, [Ranco

Province], 26 October 1965, from unspecified substrate.

382. Ologamasus foliatus Karg, 1976

331

Ologamasus foliatus Karg, 1976c: 186.

Ologamasus foliatus.— Silva, Moraes and Krantz, 2007: 67; Karg and Schorlemmer, 2009:

79.


TYPE LOCALITY AND HABITAT: Chile, Bofedal de Bacuyo [=Bofedal de Parinacota?],

Tarapacá, 26 November 1965, between algae.

383. Ologamasus lanceolatus (Karg, 1976)

Hydrogamasellus lanceolatus Karg, 1976b: 41.

Hydrogamasellus lanceolatus.— Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.



in litter.

384. Ologamasus latoventer Karg and Schorlemmer, 2009

Ologamasus latoventer Karg and Schorlemmer, 2009: 77.



TYPE LOCALITY AND HABITAT: Paraguay, Botanical Garden, Asunción, 1966, in wet

soil-moss from shrubbery along a brooklet.

385. Ologamasus mahunkai (Karg, 1979)

Heydeniella mahunkai Karg, 1979: 208.



Province, 13 November 1961, in litter at the lower edge of a Nothofagus pumilio

[Nothofagaceae] forest.

386. Ologamasus membranosus Karg, 1976

Ologamasus membranosus Karg, 1976c: 188.

Ologamasus membranosus.— Silva, Moraes and Krantz, 2007: 67; Karg and Schorlemmer,

2009: 79.


332

TYPE LOCALITY AND HABITAT: Chile, Caquena, Tarapacá, 28 November 1965, in moist

soil under manure.

387. Ologamasus microcrinis (Karg, 1979)

Hydrogamasellus microcrinis Karg, 1979: 210.



Province, 16 November 1961, in litter in a Maytenus boaria [Celastreaceae] forest

near a source.

388. Ologamasus postpilus Karg and Schorlemmer, 2009

Ologamasus postpilus Karg and Schorlemmer, 2009: 77.




389. Ologamasus simplicior (Berlese, 1914)

Ologamasellus simplicior Berlese, 1914: 140.

Ologamasus simplicior.— Lee, 1970: 89, Karg, 1976c: 192; Silva, Moraes and Krantz, 2007:

67; Karg and Schorlemmer, 2009: 79.


TYPE LOCALITY AND HABITAT: Argentina, La Plata [Buenos Aires Province], from


390. Ologamasus simplicitus Karg and Schorlemmer, 2009

Ologamasus simplicitus Karg and Schorlemmer, 2009: 74.



TYPE LOCALITY AND HABITAT: Brazil, near Barueri, [São Paulo State], 1971, in nest of

Camponotus rufipes [Hymenoptera: Formicidae].

391. Ologamasus striolatosimilis Karg, 1976

Ologamasus striolatosimilis Karg, 1976c: 190.

333

Ologamasus striolatosimilis.— Silva, Moraes and Krantz, 2007: 67; Karg and Schorlemmer,

2009: 79.



under stones.

392. Ologamasus striolatus (Berlese, 1916)

Ologamasellus striolatus Berlese, 1916a: 163.

Ologamasus striolatus.— Lee, 1970: 90; Karg, 1976c: 192; Silva, Moraes and Krantz, 2007:

67; Karg and Schorlemmer, 2009: 79.


TYPE LOCALITY AND HABITAT: Argentina, La Plata and Olavarría, Buenos Aires

Province, under stones.

393. Ologamasus testudinis (Karg, 1976)

Hydrogamasellus testudinis Karg, 1976b: 39.

Hydrogamasellus testudinis.— Karg, 1997a: 65; Karg and Schorlemmer, 2009: 82.



November 1965, on herbs.

394. Ologamasus trituberculatus Karg and Schorlemmer, 2009

Ologamasus trituberculatus Karg and Schorlemmer, 2009: 76.



TYPE LOCALITY AND HABITAT: Brazil, Aracari-guana [=Araçariguama?, São Paulo

State], 1970, in humus.

Genus Onchogamasus Womersley, 1956

Onchogamasus Womersley, 1956a: 108 (described in Pseudoparasitidae Vitzthum).

Onchogamasus.— Ryke, 1962c: 160; Lee, 1970: 188.

Type species: Onchogamasus communis Womersley, 1956, by original designation.

334

395. Onchogamasus communis Womersley, 1956

Onchogamasus communis Womersley, 1956a: 108.

Onchogamasus communis.— Ryke, 1962c: 160; Lee, 1970: 190.




396. Onchogamasus heterosetae Karg, 1996

Onchogamasus heterosetae Karg, 1996: 172.



TYPE LOCALITY AND HABITAT: New Caledonia, E. d‘Amien [= Napué Amien?], 24

February 1977, in a house.

397. Onchogamasus pumilio Lee, 1970

Onchogamasus pumilio Lee, 1970: 191.


TYPE LOCALITY AND HABITAT: Australia, Formby Bay, Yorke Peninsula, South

Australia, 3 November 1965, on moss beneath white mallee, Eucalyptus sp.

[Myrtaceae], scrub.

398. Onchogamasus quasicurtipilus Lee, 1970

Onchogamasus quasicurtipilus Lee, 1970: 193.


TYPE LOCALITY AND HABITAT: Australia, Otway Ranges, Victoria, 28 August 1965, on

moss and litter beneath tree ferns [Pteridophyta] and Eucalyptus sp. [Myrtaceae].

Genus Oriflammella Halliday, 2008

Oriflammella Halliday, 2008: 43 (described in Ologamasidae Ryke).

Type species: Oriflammella lutulenta Halliday, 2008, by original designation.

335

399. Oriflammella lutulenta Halliday, 2008

Oriflammella lutulenta Halliday, 2008: 44.


TYPE LOCALITY AND HABITAT: Australia, Wishing Tree Track (28°14‘S, 153°08‘E),

O‘Reilly‘s, Lamington National Park, Queensland, 25 October 2006, in leaf litter in a

subtropical rainforest.

Genus Parasitiphis Womersley, 1956

Parasitiphis Womersley, 1956b: 535 (described in Pseudoparasitidae Vitzthum).

Parasitiphis.— Ryke, 1962c: 160; Lee, 1970: 118.

Type species: Parasitiphis littoralis Womersley, 1956, by original designation.

Hydrogamasus (Austrohydrogamasus) Hirschmann, 1966b: 10 [junior synonymy by Lee,

1970: 118].

Type species: Hydrogamasus (Austrohydrogamasus) watsoni Hirschmann, 1966, by


400. Parasitiphis aurora Lee, 1970

Parasitiphis aurora Lee, 1970: 123.

Parasitiphis aurora.— Lee and Hunter, 1974: 316.



28 October 1913, between tidemarks.

NOTE: Described based on specimens reported by Womersley (1937): 17 as Hydrogamasus

antarcticus Trägårdh, 1907.

401. Parasitiphis brunneus (Kramer, 1898)

Laelaps brunneus Kramer, 1898a: 24.

Parasitiphis brunneus.— Lee, 1970: 121; Lee and Hunter, 1974: 318.


TYPE LOCALITY AND HABITAT: Argentina, Ushuaia, [Isla Grande de Tierra del Fuego],

27 October and 13 November 1892, in roots of mangrove in a sea beach.

336

402. Parasitiphis jeanneli (André, 1947)

Gamasellus jeanneli André, 1947: 70.

Cyrtolaelaps (Gamasellus) jeanneli.— Ryke, 1962b: 55.

Parasitiphis jeanneli.— Lee, 1970: 124; Lee and Hunter, 1974: 318.

Hydrogamasus (Austrohydrogamasus) watsoni Hirschmann, 1966c: 25 [junior synonymy by

Lee, 1970: 124].

Hydrogamasus (Austrohydrogamasus) watsoni.— Hunter, 1970: 62.

TYPE DEPOSITORY: P. jeanneli: Muséum National d‘Histoire Naturelle, Paris, France; H.

(A.) watsoni: Hirschmann‘s Milbensammlung, Nuremberg, Germany.

TYPE LOCALITY AND HABITAT: P. jeanneli: Kerguelen Islands [Indian Ocean], Port

Hopeful, Port Louison and Île du Port, 10, 13 and 18 February 1939, on stones at low

tide; Macquarie Island [between Australia and Antarctica], on algae on coastal rocks;

under rocks on the beach; in mud; in nest of Pachyptila desolata [Aves:

Procellariidae]; in litter of Colobanthus muscoides [Caryophyllaceae], Poa annina and

Poa foliosa [Poaceae].

403. Parasitiphis littoralis Womersley, 1956

Parasitiphis littoralis Womersley, 1956b: 536.

Parasitiphis littoralis.— Ryke, 1962c: 160; Lee, 1970: 123.


TYPE LOCALITY AND HABITAT: Australia, mouth of American River, Kangaroo Island,

South Australia, January 1946, on littoral zone.

Genus Periseius Womersley, 1961

Periseius Womersley, 1961: 198 (described in Neoparasitidae Oudemans).

Periseius.— Hirschmann, 1966a: 2; Lee, 1970: 166; Karg, 1994a: 125.

Periseius (Periseius).— Hirschmann, 1966a: 5; Lee, 1970: 166.

Type species: Periseius littorale Womersley, 1961, by original designation [junior

synonym of Periseius hammeni (Womersley, 1961)].

337

Psammonsela Haq, 1965: 413 (described in Rhodacaridae Oudemans) [junior synonymy by

Hirschmann, 1966a: 5].

Periseius (Psammonsela).— Hirschmann, 1966a: 5; Lee, 1970: 168.

Type species: Psammonsella nobskae Haq, 1965, by original designation.

404. Periseius brasiliensis Hirschmann, 1966

Periseius (Periseius) brasiliensis Hirschmann, 1966c: 37.

Periseius (Periseius) braziliensis [sic].— Lee, 1970: 167.

Periseius brasiliensis.— Karg, 1994a: 125.


TYPE LOCALITY AND HABITAT: Brazil, Recife, [Pernambuco State], in coastal soil.

405. Periseius hammeni (Womersley, 1961)

Cyrtolaelaps hammeni Womersley, 1961: 190.

Periseius hammeni.— Lee, 1970: 167; Karg, 1994a: 125.

Periseius (Periseius) hammeni.— Lee, 1970: 167.

Periseius littorale Womersley, 1961: 198 [junior synonymy by Lee, 1970: 167].

TYPE DEPOSITORY: P. hammeni and P. littorale: Leiden Museum, Leiden, Netherlands.

TYPE LOCALITY AND HABITAT: P. hammeni and P. littorale: New Guinea, near the

Royal Netherlands Naval Base, Biak Island, 1953, on Cladophora socialis

[Cladophoraceae] and red algae [Eukaryote: Rhodophyta] on the stones in the

intertidal zone.

406. Periseius nobskae (Haq, 1965)

Psammonsella nobskae Haq, 1965: 413.

Periseius nobskae – Hirschmann, 1966a: 2; Karg, 1994a: 125.

Periseius (Psammonsella) nobskae.— Hirschmann, 1966a: 5; Lee, 1970: 169.



TYPE LOCALITY AND HABITAT: USA, Nobska Beach, Woods Hole, Massachusetts, in

beach sand.

407. Periseius plumosus Karg, 1994

Periseius plumosus Karg, 1994a: 124.

338



TYPE LOCALITY AND HABITAT: Galapagos Islands, Fernandina Island, 18 March 1985,

on grass and dry sand in a riparian zone.

408. Periseius schusteri Hirschmann, 1966

Periseius (Psammonsella) schusteri Hirschmann, 1966c: 37.

Periseius (Psammonsella) schusteri.— Lee, 1970: 169.

Periseius schusteri.— Karg, 1994a: 125.


TYPE LOCALITY AND HABITAT: Croatia, Saint Andrew's Island, Rovinj, in sand and

gravel; Greece, Epidaurus, Argolis, Peloponnese; Greece, Nea Krini, Thessaloniki Urban

Area; France, Carro and Marseille.

Genus Pilellus Lee, 1970

Pilellus Lee, 1970: 169 (described in Rhodacaridae Oudemans).

Type species: Cyrtolaelaps (Gamasellus) rykei Hunter, 1967, by original designation.

409. Pilellus rugipellis Lee and Hunter, 1974

Pilellus rugipellis Lee and Hunter, 1974: 324.


TYPE LOCALITY AND HABITAT: New Zealand, Hooker Hills, Auckland Island, 11

February 1963, on moss Dicronoloma billardieri [Dicranaceae].

410. Pilellus rykei (Hunter, 1967)

Cyrtolaelaps (Gamasellus) rykei Hunter, 1967b: 34.

Gamasellus rykei.— Hunter, 1970: 61.

Pilellus rykei.— Lee, 1970: 170; Lee and Hunter, 1974: 324.


TYPE LOCALITY AND HABITAT: South Sandwich Islands [Southern Atlantic Ocean],

Candlemas Island, 15 March 1964, on Polytrichum alpinum [Polytrichaceae] mat.

339

Genus Podonotogamasellus Loots and Ryke, 1965

Notogamasellus (Podonotogamasellus) Loots and Ryke, 1965: 471 (described in

Rhodacaridae Oudemans).

Notogamasellus (Podonotogamasellus).— Lee, 1970: 165.

Type species: Notogamasellus (Podonotogamasellus) magoebaensis Loots and Ryke,

1965, by original designation.

411. Podonotogamasellus magoebaensis (Loots and Ryke, 1965)

Notogamasellus (Podonotogamasellus) magoebaensis Loots and Ryke, 1965: 471.

Notogamasellus (Podonotogamasellus) magoebaensis.— Lee, 1970: 165.



TYPE LOCALITY AND HABITAT: South Africa, Magoebaskloof, Limpopo [cited as

Transvaal], 1962 and 1963, in soil in the indigenous forest.

Genus Pyriphis Lee, 1970

Pyriphis Lee, 1970: 125 (described in Rhodacaridae Oudemans).

Type species: Ologamasus pyrenoides Lee, 1966, by original designation.

412. Pyriphis pyrenoides (Lee, 1966)

Ologamasus pyrenoides Lee, 1966: 222.

Pyriphis pyrenoides.— Lee, 1970: 126.


TYPE LOCALITY AND HABITAT: Australia, Beachamps Falls, Otway Ranges, Victoria,

19 January 1962, from ―antarctic beech‖ (Nothofagus cunninghamii) [Nothofagaceae]

litter.

340

Queenslandolaelaps Womersley, 1956

Queenslandolaelaps Womersley, 1956a: 109 (described in Neoparasitidae Oudemans).

Queenslandolaelaps.— Ryke, 1962c: 160; Lee, 1970: 195.

Type species: Queenslandolaelaps vitzthumi Womersley, 1956, by original

designation.

413. Queenslandolaelaps berlesei Womersley, 1956

Queenslandolaelaps berlesei Womersley, 1956a: 111.

Neogamasellevans berlesei.— Lee, 1970: 203.




414. Queenslandolaelaps vitzthumi Womersley, 1956

Queenslandolaelaps vitzthumi Womersley, 1956a: 109.

Queenslandolaelaps vitzthumi.— Ryke, 1962c: 160; Lee, 1970: 198.




Genus Rhodacaroides Willmann

Rhodacaroides Willmann, 1959: 97 (presumed to be described in Rhodacaridae Oudemans).

Rhodacarus (Rhodacaroides).— Ryke, 1962c: 156.

Rhodacaroides.— Lee, 1970: 170; Karg, 1977: 328; Karg, 1979: 197.

Rhodacaroides (Rhodacaroides).— Karg, 1977: 329; Karg, 1979: 198.

Type species: Rhodacaroides aegyptiacus Willmann, 1959, by original designation.

Rhodacaroides (Tenacaroides) Karg, 1977: 329.

Rhodacaroides (Tenacaroides).— Karg, 1979: 198.

Type species: Rhodacaroides (Tenacaroides) calidus Karg, 1977, by original

designation [cited as Rhodacarus calidus].

341

Rhodacaroides (Nodacaroides) Karg, 1977: 330.

Rhodacaroides (Nodacaroides).— Karg, 1979: 198.

Type species: Rhodacaroides (Nodacaroides) coniunctus Karg, 1977, by original

designation.

415. Rhodacaroides aegyptiacus Willmann, 1959

Rhodacaroides aegyptiacus Willmann, 1959: 98.

Rhodacarus (Rhodacaroides) aegyptiacus.— Ryke, 1962c: 156.

Rhodacaroides aegyptiacus.— Lee, 1970: 172.

Rhodacaroides (Rhodacaroides) aegyptiacus.— Karg, 1979: 198.

TYPE DEPOSITORY: Zoologisches Institut der Universität zu Kiel, Kiel, Germany.

TYPE LOCALITY AND HABITAT: Egypt, Hurghada, coast of the Red Sea, 28 March 1956,

in coastal groundwater.

416. Rhodacaroides brevispiritus Karg, 1977

Rhodacaroides (Tenacaroides) brevispiritus Karg, 1977: 331.

Rhodacaroides brevispiritus.— Karg, 1977: 331.

Rhodacaroides (Tenacaroides) brevispiritus.— Karg, 1979: 198.


TYPE LOCALITY AND HABITAT: Chile, Azapa, Tarapacá, 2 December 1965, on the

bushes at the edge of a marsh.

417. Rhodacaroides calidus Karg, 1977

Rhodacaroides (Tenacaroides) calidus Karg, 1977: 330.

Rhodacarus calidus.— Karg, 1977: 329.

Rhodacaroides calidus.— Karg, 1977: 329.

Rhodacaroides (Tenacaroides) calidus.— Karg, 1979: 198.


TYPE LOCALITY AND HABITAT: Chile, La Rinconada, Santiago Province, 28

September1965, on moss and soil on rocks.

418. Rhodacaroides coniunctus Karg, 1977

Rhodacaroides (Nodacaroides) coniunctus Karg, 1977: 333.

342

Rhodacaroides coniunctus.— Karg, 1977: 332.

Rhodacaroides (Nodacaroides) coniunctus.— Karg, 1979: 198.


TYPE LOCALITY AND HABITAT: Chile, Azapa (alt. 1,000 m), Tarapacá, 2 December

1965, in soil trap.

419. Rhodacaroides costai (Sheals, 1962)

Rhodacarus costai Sheals, 1962: 85.

Rhodacaroides costai.— Lee, 1970: 203.

Rhodacaroides (Nodacaroides) costai.— Karg, 1979: 198.


TYPE LOCALITY AND HABITAT: Argentina, Parque Nacional Los Arrayanes, 2 May

1959, in soil.

420. Rhodacaroides crinitus Karg, 1979

Rhodacaroides (Nodacaroides) crinitus Karg, 1979: 199.



Province, 24 April 1961, in soil of a pasture at the edge of a Libocedrus

[Cupressaceae] forest.

421. Rhodacaroides leptinochaetus (Ma, 2005)

Gamasellus leptinochaetus Ma, 2005b: 538.

TYPE DEPOSITORY: National Base of Plague and Brucellosis Control, Baicheng City,

China.

TYPE LOCALITY AND HABITAT: China, Tai‘an (36°15‘N, 117°08‘E), Shandong

Province, July 2000, under fallen leaves.

422. Rhodacaroides levis Karg, 1977

Rhodacaroides (Nodacaroides) levis Karg, 1977: 333.

Rhodacaroides levis.— Karg, 1977: 334.

Rhodacaroides (Nodacaroides) levis.— Karg, 1979: 198.


343

TYPE LOCALITY AND HABITAT: Chile, Azapa (alt. 2,000 m), Tarapacá, 2 December

1965, in soil trap.

423. Rhodacaroides minyaspis Lee, 1973

Rhodacaroides minyaspis Lee, 1973: 26.



Adelaide, South Australia, 12 August 1968, in plant litter.

424. Rhodacaroides unguellus Karg, 1979

Rhodacaroides (Tenacaroides) unguellus Karg, 1979: 200.


TYPE LOCALITY AND HABITAT: Argentina, foothill of Cerro Pirque, Chubut Province, 3

April 1961, under stones in a meadow.

Genus Rykellus Lee, 1970

Rykellus Lee, 1970: 126 (described in Rhodacaridae Oudemans).

Rykellus.— Karg and Schorlemmer, 2009: 69.

Type species: Cyrtolaelaps (Gamasellus) darglensis Ryke, 1962, by original

designation.

425. Rykellus brevipellitus Karg and Schorlemmer, 2009

Rykellus brevipellitus Karg and Schorlemmer, 2009: 69.



TYPE LOCALITY AND HABITAT: Brazil, near Barueri, [São Paulo State], 1971, in nest of

Camponotus rufipes [Hymenoptera: Formicidae].

426. Rykellus darglensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) darglensis Ryke, 1962b: 39.

Gamasellus (Hydrogamasellus) darglensis.— Hirschmann, 1966b: 7.

Rykellus darglensis.— Lee, 1970: 127; Karg and Schorlemmer, 2009: 70.

344


TYPE LOCALITY AND HABITAT: South Africa, Dargle, KwaZulu-Natal, November 1942,

in litter.

427. Rykellus nkandhlaensis (Ryke, 1962)

Cyrtolaelaps (Gamasellus) nkandhlaensis Ryke, 1962b: 42.

Gamasellus (Hydrogamasellus) nkandhlaensis.— Hirschmann, 1966b: 7.

Rykellus nkandhlaensis.— Lee, 1970: 127; Karg and Schorlemmer, 2009: 70.


TYPE LOCALITY AND HABITAT: South Africa, Nkandhla Forest, KwaZulu-Natal,

November 1940, in litter.

Genus Sessiluncus G. Canestrini, 1898

Sessiluncus G. Canestrini, 1898: 486 [presumed to be described in Parasitidae Oudemans

(also referred to as Gamasidae Leach)].

Gamasellus (Sessiluncus).— Berlese, 1905a: 168.

Sessiluncus.— Vitzthum, 1931a: 142; Vitzthum, 1943: 758; Ryke, 1962c: 160; Lee, 1970:

175; Bregetova, 1977a: 311; Nasr and Afifi, 1984: 17; Zaher, 1986: 30.

Type species: Gamasus heterotarsus G. Canestrini, 1897, by subsequent monotypy.

428. Sessiluncus abalaae Datta and Bhattacharjee, 1991

Sessiluncus abalaae Datta and Bhattacharjee, 1991: 721.


TYPE LOCALITY AND HABITAT: India, Jorhat, Assam, 4 March 1984, in leaf litter.

429. Sessiluncus aegypticus Nasr and Afifi, 1984

Sessiluncus aegypticus Nasr and Afifi, 1984: 18.

Sessiluncus aegypticus.— Zaher, 1986: 31.

TYPE DEPOSITORY: National Research Centre, Cairo, Egypt.

TYPE LOCALITY AND HABITAT: Egypt, Mallawi, Minya Governorate, in soil cultivated

with sugar cane Saccharum officinarum L. [Poaceae].

345

430. Sessiluncus bengalensis Bhattacharyya, 1977

Sessiluncus bengalensis Bhattacharyya, 1977a: 47.


TYPE LOCALITY AND HABITAT: India, Uluberia, Howrah District, West Bengal, 12

March 1965, in soil litter of straw and cowdung.


431. Sessiluncus calcuttaensis Bhattacharyya, 1977

Sessiluncus calcuttaensis Bhattacharyya, 1977a: 44.


TYPE LOCALITY AND HABITAT: India, Esplanade, Calcutta, West Bengal, 11 June 1965,

in soil mixed with grass roots.

432. Sessiluncus cavensis Willmann, 1940

Sessiluncus cavensis Willmann, 1940: 213.

Sessiluncus cavensis.— Willmann, 1941: 23; Lee, 1970: 178; Bregetova, 1977a: 314; Nasr

and Afifi, 1984: 18.


TYPE LOCALITY AND HABITAT: Bosnia and Herzegovina, Petrinja, 25 March 1913, in a

cave.

433. Sessiluncus colchicus Bregetova, 1977

Sessiluncus colchicus Bregetova, 1977a: 314.

Sessiluncus colchicus.— Nasr and Afifi, 1984: 18.


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Russia, Akhun Montain, Krasnodar, 19 March 1962,

near the trunk of a hornbeam, Carpinus sp. [Betulaceae].

434. Sessiluncus femoralis Bhattacharyya, 1977

Sessiluncus femoralis Bhattacharyya, 1977a: 46.


TYPE LOCALITY AND HABITAT: India, Bandel, Hooghly District, West Bengal, 16

February 1966, in straw litter.

346

435. Sessiluncus heterotarsus (G. Canestrini, 1897)

Gamasus heterotarsus G. Canestrini, 1897: 473.

Sessiluncus heterotarsus.— G. Canestrini, 1898: 486; Ryke, 1962c: 160; Lee, 1970: 178;

Nasr and Afifi, 1984: 18.

Gamasellus (Sessiluncus) heterotarsus.— Berlese, 1905a: 168; Vitzthum, 1926a: 5.


TYPE LOCALITY AND HABITAT: Papua New Guinea, Madang [cited as Friedrich-

Wilhelmshafen], from unspecified substrate.

NOTE: According to Lee (1970): 184, the specimens reported by Domrow (1957): 202 as

Sessiluncus heterotarsus are Antennolaelaps testudo Lee, 1970.

436. Sessiluncus holostaspoides Canestrini, 1884

Gamasus holostaspoides Canestrini, 1884 apud Bregetova, 1977a: 314.

Sessiluncus holostaspoides.— Bregetova, 1977a: 314.


TYPE LOCALITY AND HABITAT: Italy apud Bregetova, 1977a: 314.

437. Sessiluncus hungaricus Karg, 1964

Sessiluncus hungaricus Karg, 1964: 73.

Sessiluncus hungaricus.— Lee, 1970: 178; Bregetova, 1977a: 313; Karg, 1993c: 378.

Sessiluncus hangaricus [sic].— Nasr and Afifi, 1984: 17.



TYPE LOCALITY AND HABITAT: Hungary, Tihany, 24 July 1962, in litter under oak,

Quercus sp. [Fagaceae] bushes.

438. Sessiluncus indicus Bhattacharyya, 1977

Sessiluncus indicus Bhattacharyya, 1977a: 43.


TYPE LOCALITY AND HABITAT: India, Botanical Garden, Shibpur, Howrah District,

West Bengal, 24 December 1964, in soil beneath Musa sp. [Musaceae].

439. Sessiluncus leei Datta and Bhattacharjee, 1991

347

Sessiluncus leei Datta and Bhattacharjee, 1991: 724.


TYPE LOCALITY AND HABITAT: India, Lumding, Assam, 8 March 1984, in leaf litter.

440. Sessiluncus oculatus Vitzthum, 1935

Sessiluncus oculatus Vitzthum, 1935: 150.

Sessiluncus oculatus.— Bhattacharyya, 1965: 151; Lee, 1970: 178; Bhattacharyya, 1977a: 47.

TYPE DEPOSITORY: Bernice P. Bishop Museum, Honolulu, Hawai.

TYPE LOCALITY AND HABITAT: French Polynesia, Papara-Tal, four miles from the

coast, Tahiti, 21 December 1928, on Zingiber sp. [Zingiberaceae].


441. Sessiluncus reticulatus Loots, 1980

Sessiluncus reticulatus Loots, 1980: 750.


TYPE LOCALITY AND HABITAT: Seychelles [Indian Ocean], Morne Blanc, Mahé Centre

(alt. 667 m), 24-25 August 1972, from unspecified substrate.

Genus Solugamasus Lee, 1973

Solugamasus Lee, 1973: 28 (described in Rhodacaridae Oudemans).

Type species: Solugamasus mustela Lee, 1973, by original designation.

442. Solugamasus mustela Lee, 1973

Solugamasus mustela Lee, 1973: 28.


TYPE LOCALITY AND HABITAT: Australia, near First Waterfall, Waterfall Gully,

Adelaide, South Australia, 9 May 1968 and 30 January 1969, on moss and plant litter.

348

Genus Stylochirus G. Canestrini and R. Canestrini, 1882

Stilochirus G. Canestrini and R. Canestrini, 1882: 56 [described in Parasitidae Oudemans

(also referred to as Gamasidae Leach), mentioned in the original description of the

genus as family Gamasini Canetrini and Fanzago].

Stilochirus.— G. Canestrini, 1885: 57.

Stylochirus.— Berlese, 1892e: 13; Berlese, 1892f: 45; Lee, 1970: 198; Karg, 1971: 351;

Bregetova, 1977a: 310; Mašán and Kaluz, 2001: 488.

Type species: Stilochirus rovennensis G. Canestrini and R. Canestrini, 1882, by

subsequent monotypy.

Iphidosoma Berlese, 1892d: 5 [presumed to be described in Parasitidae Oudemans (also

referred to as Gamasidae Leach)] [junior synonymy by Mašán and Kaluz, 2001: 484].

Iphidosoma.— Karg, 1971: 119; Bregetova, 1977b: 563; Karg, 1993c: 94; Mašán and

Halliday, 2010: 98.

Type species: Holostapis fimetaria Müller, 1860, by subsequent designation.

Physallolaelaps Berlese, 1908: 13 [described in Parasitidae Oudemans (also referred to as

Gamasidae Leach), mentioned in the original description of the genus as Gamasidi]

[junior synonymy by Lee, 1970: 198].

Physallolaelaps.— Athias-Henriot, 1961b: 256; Ryke, 1962c: 160.

Type species: Physallolaelaps ampulliger Berlese, 1908, by monotypy.

Gamasiphis (Periphis) Berlese, 1914: 142 (described in Cyrtolaelaptini Berlese) [junior

synonymy by Lee, 1970: 198].

Periphis.— Vitzthum, 1931a: 142; Oudemans, 1939: 22; Vitzthum, 1943: 756; Baker and

Wharton, 1952: 73; Ryke, 1962c: 160.

Gamasiphis (Periphis).— Trägårdh, 1952: 54; Womersley, 1956b: 517.

Type species: Iphis haemisphaericus Koch, 1839, by monotypy.

349

Gamasiphis (Epiphis) Berlese, 1916b: 302 [presumed to be described in Parasitidae

Oudemans (also referred to as Gamasidae Leach)] [junior synonymy by Lee, 1970:

198].

Epiphis.— Vitzthum, 1931a: 142; Oudemans, 1939: 22; Vitzthum, 1943: 756; Baker and

Wharton, 1952: 73; Ryke, 1962c: 159; Kethley, 1983: 2600.

Gamasiphis (Epiphis).— Womersley, 1956b: 517.

Type species: Gamasiphis (Epiphis) rarior Berlese, 1916, by monotypy.

Gamasiphis (Megaliphis) Willman, 1938: 161 (described in Parasitidae Oudemans) [junior

synonymy by Lee, 1970: 198].

Megaliphis.— Oudemans, 1939: 21; Vitzthum, 1943: 756; Baker and Wharton, 1952: 73;

Ryke, 1962c: 160.

Gamasiphis (Megaliphis).— Trägårdh, 1952: 55; Womersley, 1956b: 517.

Type species: Gamasiphis (Megaliphis) giganteus Willmann, 1938, by monotypy.

NOTE: In the original description, the genus name was spelled as Stilochirus. Berlese (1882):

2 unjustifiably (Article 33.3 of the fourth edition of the International Code of

Zoological Nomenclature) changed it to Stylochirus. All subsequent publications on

the species of this genus used Berlese‘s proposed spelling. Thus, because of the

prevailing usage, Berlese‘s spelling is maintained in this catalog.

443. Stylochirus ampulliger (Berlese, 1908)

Physallolaelaps ampulliger Berlese, 1908: 13.

Physallolaelaps ampulliger.— Berlese, 1913a: 84; Athias-Henriot, 1961b: 257; Ryke, 1962c:

160.

Stylochirus ampulliger.— Lee, 1970: 200.


TYPE LOCALITY AND HABITAT: Italy, Vallombrosa, [Reggelo, Florense Province], from



444. Stylochirus caucasicus Bregetova, 1977

Stylochirus caucasicus Bregetova, 1977a: 311.

350


Petersburg, Russia.

TYPE LOCALITY AND HABITAT: Georgia, Lagodekhi Nature Reserve, 6 October 1960,


445. Stylochirus fimetarius (Müller, 1859)

Holostapis fimetaria Müller, 1860: 182.

Poecilochirus fimetarius.— G. Canestrini and R. Canestrini, 1882: 57; G. Canestrini, 1885:

98.

Iphidosoma fimetarium.— Berlese, 1892d: 5; Willmann, 1941: 31; Karg, 1971: 121;

Bregetova, 1977b: 564; Karg, 1993c: 95.

Gamasoides fimetarium.— Schweizer, 1922: 34.

Iphidiosoma [sic] fimetarium.— Schweizer, 1961: 144.

Stylochirus fimetarius.— Mašán and Kaluz, 2001: 484; Mašán and Halliday, 2010: 98.

Copriphis janetscheki Schmölzer, 1956: 544 [junior synonymy by Kethley, 1983: 2599].

Copriphis janetski [sic].— Kethley, 1983: 2599.

TYPE DEPOSITORY: S. fimetarius: Author´s private collection; C. janetscheki: Unknown.

TYPE LOCALITY AND HABITAT: S. fimetarius: Czech Republic, Brno [cited as Brünn],

Moravia [cited as Mähren]; Czech Republic, Sudetes [cited as Sudeten], on Aphodius

fimetarius [Coleoptera: Scarabaeidae], Molytes germanus [Coleoptera: Curculionidae:

Molytinae] and Carabus glabratus [Coleoptera: Carabidae]; C. janetscheki: France,

the steep ridge to Le Flambeau, Refuge Temple-Ecrins, [Écrins National Park],

Dauphiné Alps, 31 July 1951, in patches of grass under a vertical wall in a Curvuletum

area.

446. Stylochirus giganteus (Willmann, 1938)

Gamasiphis (Megaliphis) giganteus Willmann, 1938: 161.

Megaliphis giganteus.— Oudemans, 1939: 21; Vitzthum, 1943: 756; Baker and Wharton,

1952: 73; Ryke, 1962c: 160; Willmann, 1953: 456.

Stylochirus giganteus.— Lee, 1970: 200; Karg, 1971: 351; Karg, 1993c: 376; Mašán and

Kaluz, 2001: 488.


TYPE LOCALITY AND HABITAT: Slovakia, Skalka Berg (alt. 1,188 m), Kremnica, from


351

447. Stylochirus haemisphaericus (Koch, 1839)

Iphis haemisphaericus Koch, 1839d: 16.

Eumaeus hemisphaericus [sic].— Koch, 1842: 95.

Gamasiphis (Periphis) haemisphaericus.— Berlese, 1914: 142.

Periphis haemisphaericus.— Vitzthum, 1931a: 142; Willman, 1938: 149.

Periphis hemisphaericus [sic].— Baker and Wharton, 1952: 73; Ryke, 1962c: 160.

Stylochirus haemisphaericus.— Lee, 1970: 201.


TYPE LOCALITY AND HABITAT: Germany, in marshy meadows near ditches or water

containers.

448. Stylochirus minor (Willmann, 1953)

Megaliphis minor Willmann, 1953: 456.

Stylochirus minor.— Lee, 1970: 200; Karg, 1971: 351; Karg, 1993c: 378; Mašán and Kaluz,

2001: 488.


TYPE LOCALITY AND HABITAT: Austria, Grieswiesalm, 15 May 1940, in litter of

Calluna [Ericaceae], Vaccinium [Ericaceae] and Rosa [Rosaceae].

449. Stylochirus rarior (Berlese, 1916)

Gamasiphis (Epiphis) rarior Berlese, 1916b: 303.

Epiphis rarior.— Vitzthum, 1931a: 142; Oudemans, 1939: 22; Vitzthum, 1943: 756; Baker

and Wharton, 1952: 73; Ryke, 1962c: 159; Kethley, 1983: 2600.

Stylochirus rarior.— Lee, 1970: 201.


TYPE LOCALITY AND HABITAT: USA, Columbia, Missouri, 1904-1906, on leaf mould.

450. Stylochirus rovennensis G. Canestrini and R. Canestrini, 1882

Stilochirus rovennensis G. Canestrini and R. Canestrini, 1882: 56.

Stilochirus rovennensis.— G. Canestrini, 1885: 100.

Stylochirus rovennensis.— Berlese, 1892e: 13; Lee, 1970: 200; Bregetova, 1977a: 310;

Mašán and Kaluz, 2001: 488.


352

TYPE LOCALITY AND HABITAT: Italy, Monte Rovenna, Val di Non, Province of Trento,

on moss.

Species incertae sedis

001. Asca muricata Fox, 1947

.Asca muricata Fox, 1947: 600

Cyrtolaelaps (Digamasellus) muricatus.— Ryke, 1961a: 109.


Juan, Puerto Rico.

TYPE LOCALITY AND HABITAT: Puerto Rico, San Juan, 16 May 1947, from Rattus


002. Cyrtolaelaps dama Berlese, 1905

Cyrtolaelaps dama Berlese, 1905a: 168.

Cyrtolaelaps dama.— Ryke, 1962b: 13.


TYPE LOCALITY AND HABITAT: Indonesia, Tjibodas, from unspecified substrate.

003. Cyrtolaelaps paraster Costa, 1961

Cyrtolaelaps paraster Costa, 1961: 275.

Cyrtolaelaps paraster.— Lee, 1970: 150; Bregetova and Shcherbak, 1977a: 304; Chelebiev,

1984: 141.

TYPE DEPOSITORY: Department of Parasitology, Hebrew University, Jerusalem, Israel.

TYPE LOCALITY AND HABITAT: Israel, Mishmar HaEmek, 3 January 1956, in nest of

Microtus guentheri [Mammalia: Rodentia: Cricetidae].

004. Cyrtolaelaps (Gamasellus) armatus Berlese, 1904

Cyrtolaelaps (Gamasellus) armatus Berlese, 1904: 279.

Digamasellus armatus.— Hirschmann, 1954b: 246; Franz, 1954: 341.

Cyrtolaelaps (Digamasellus) armatus.— Ryke, 1962a: 107.

Gamasellus spalacis Oudemans, 1912: 261 [objective synonym — unjustified replacement

name].

353

Cyrtolaelaps (Gamasellus) spalacis.— Ryke, 1962b: 52.

Gamasellus spalacis.— Lee, 1970: 150.


TYPE LOCALITY AND HABITAT: Germany, Bremen, in a mole nest [Mammalia:

Talpidae].

NOTE: Described on the basis of specimens reported by Oudemans (1904): 78 as adult male

of Parasitus spinipes (Koch).

005. Cyrtolaelaps (Gamasellus) grabouwensis Ryke, 1964

Cyrtolaelaps (Gamasellus) grabouwensis Ryke, 1964: 351.

Gamasellus grabouwensis.— Lee, 1970: 135.

TYPE DEPOSITORY: Department of Parasitology, Hebrew University, Jerusalem, Israel.

TYPE LOCALITY AND HABITAT: South Africa, Grabouw, Western Cape [cited as Cape

Province], January 1955, in a dry flower of Protea mellifera [Proteaceae].


006. Cyrtolaelaps (Gamasellus?) iphidiformis Berlese, 1904.

Cyrtolaelaps (Gamasellus?) iphidiformis [sic] Berlese, 1904: 261.


TYPE LOCALITY AND HABITAT: Italy, Boboli Gardens, Florence, in litter.

007. Gamasellus curvicrinus Berlese, 1911

Gamasellus curvicrinus Berlese, 1911: 433.

Cyrtolaelaps (Gamasellus) curvicrinus.— Ryke, 1962b: 52.


TYPE LOCALITY AND HABITAT: Indonesia, Java, on a unidentified Chrysomelidae

[Coleoptera].

008. Gamasellus gressitti Hunter, 1970

Gamasellus gressitti Hunter, 1970: 63.

Litogamasus gressitti.— Lee and Hunter, 1974: 324.

TYPE DEPOSITORY: Bernice P. Bishop Museum, Honolulu, Hawai.

TYPE LOCALITY AND HABITAT: South Georgia [southern Atlantic Ocean], Fresh Water

Bay, Bird Island, 7 April 1963, from Pachyptila desolata [Aves: Procellariidae].

354

009. Gamasellus inermis Halbert, 1920

Gamasellus inermis Halbert, 1920: 117.

Cyrtolaelaps (Digamasellus) inermis.— Ryke, 1962a: 103.


TYPE LOCALITY AND HABITAT: Ireland, Malahide, February and September 1919, in

fissures and between flakes [sic] on the seashore in an intertidal zone.

010. Gamasellus mycophagus Cooreman, 1942

Gamasellus mycophagus Cooreman, 1942: 11.

Cyrtolaelaps (Gamasellus) mycophagus.— Ryke, 1962b: 53.


TYPE LOCALITY AND HABITAT: Belgium, Gembloux, September 1942, in Inonotus

hispidus [Fungi: Hymenochaetaceae] in decomposition.

011. Gamasellus vulgaris Vitzthum, 1918

Gamasellus vulgaris Vitzthum, 1918: 9.

Cyrtolaelaps (Gamasellus) vulgaris.— Ryke, 1962b: 53.


TYPE LOCALITY AND HABITAT: Germany, Weimar, 1913, on Stomoxys calcitrans

[Diptera: Muscidae] and in litter; Sofia, Bulgaria, 1913, on Musca domestica [Diptera:

Muscidae].


012. Gamasellus (Di) cultriger Lombardini, 1940

Gamasellus (Di) cultriger Lombardini, 1940: 13.

Cyrtolaelaps (Digamasellus) cultriger.— Ryke, 1962a: 107.


TYPE LOCALITY AND HABITAT: Brazil, under the elytron of a passalid beetle

[Coleoptera].

NOTE: Described from the nymph.

013. Gamasus rotundatus Dugès, 1834

Gamasus rotundatus Dugès, 1834: 29.

355

Gamasus rotundatus.— Lucas, 1840: 481.

Sessiluncus ? rotundatus [sic].— Oudemans, 1936: 203.


TYPE LOCALITY AND HABITAT: Unspecified type locality and substrate.

014. Hologamasus quinquedentatus Ewing, 1920

Hologamasus quinquedentatus Ewing, 1920: 290.



TYPE LOCALITY AND HABITAT: USA, Ithaca, New York, under a stone.

015. Hydrogamasus japonicus Ishikawa, 1969

Hydrogamasus japonicus Ishikawa, 1969: 54.


Matsuyama, Japan.

TYPE LOCALITY AND HABITAT: Japan, ―Otanomosu-no-taira‘, Mount Shiga, Shiga

Kogen, Nagano Prefecture, 16 August 1967, in soil and litter of a coniferous forest.

016. Iphidosoma communis Nikolsky, 1981

Iphidosoma communis Nikolsky, 1981: 17.


TYPE LOCALITY AND HABITAT: Russia, Ussuriysky Reserve, Primorsky Territory, in

litter in the coniferous-deciduous forest.


017. Iphidosoma heterochaetum Nikolsky, 1981

Iphidosoma heterochaetum Nikolsky, 1981: 14.


TYPE LOCALITY AND HABITAT: Russia, Ussuriysky Reserve, Primorsky Territory, in

litter of the coniferous-deciduous forest.


018. Iphidosoma multiclavatum Willmann, 1953

Iphidosoma multiclavatum Willmann, 1953: 458.

356

Iphidosoma multiclavatum.— Karg, 1971: 119; Bregetova, 1977b: 564; Karg, 1993c: 94.


TYPE LOCALITY AND HABITAT: Austria, Kapruner Tal, 14 July 1939, in humus, litter

and rotten branches on a slope above a waterfall.


019. Iphidosoma ovatum Berlese, 1892

Iphidosoma ovatum Berlese, 1892d: 5.

Iphidosoma ovatum.— Karg, 1971: 121; Bregetova, 1977b: 563; Karg, 1993c: 94.


TYPE LOCALITY AND HABITAT: Italy, Vallombrosa, [Reggelo, Florence Province], on

moss.


020. Iphidosoma physogastris Karg, 1971

Iphidosoma physogastris Karg, 1971: 121.

Iphidosoma physogastris.— Bregetova, 1977b: 563; Karg, 1993c: 95.


Germany.

TYPE LOCALITY AND HABITAT: Central Europe, on moss, humus and litter in a

coniferous [Pinophyta] forest.


021. Iphidosoma razumovae Bregetova, 1977

Iphidosoma razumovae Bregetova, 1977b: 564.

Iphidosoma razumovae.— Karg, 1993c: 95; Peverieri, Gwiazdowicz and Sammarone, 2007:

174.


TYPE LOCALITY AND HABITAT: Georgia, from unspecified substrate.


022. Iphidosoma verrucosa Nikolsky, 1990

Iphidosoma verrucosa Nikolsky, 1990: 41.

Iphidosoma verrucosa.— Nikolsky, 1992: 111.

357


TYPE LOCALITY AND HABITAT: Russia, Irbit River, Ubsunur Hollow, Tuva Republic,

12 July 1976, in forest litter under willow Salix sp. [Salicaceae] and Caragana

[Fabaceae] of a floodplain.

023. Leioseius crassipes (Berlese, 1910)

Ameroseius crassipes Berlese, 1910c: 370.

Lasioseius (Leioseius) crassipes.— Berlese, 1916a: 33.

Leioseius crassipes.— Halliday, 1998: 118.


TYPE LOCALITY AND HABITAT: Australia, on Brontispa froggatti [junior synonym of

Brontispa longissima] [Coleoptera: Chrysomelidae].

NOTE: Bernhard (1963): 164 did not considered it to belong to Leioseius, but probably to

Ameroseius; Halliday (1998): 182 questioned its generic placement; Halliday (1997):

179 and Halliday, Walter and Lindquist (1998): 25 transferred it to Ologamasidae.

024. Lobocephalus acuminatus Kramer, 1898

Lobocephalus acuminatus Kramer, 1898b: 418.


TYPE LOCALITY AND HABITAT: German East Africa [corresponding today to Burundi,

Rwanda and Tanganyika], January 1888, from unspecified substrate.

Nomina nuda

001. Cyrtolaelaps leitnerae Franz, 1954

Cyrtolaelaps leitnerae Franz, 1954: 338.

NOTE: A nomen nudum was created when this name was used by Franz, 1954 without giving

a description or figure for the species.

002. Gamasiphis tylophagous El-Borolosy and El-Banhawy, 1999

Gamasiphis tylophagous El-Borolosy and El-Banhawy, in El-Banhawy et al., 1999: 25.

Genus Pachymasiphis Karg, 1996

358

Pachymasiphis Karg, 1996: 183.

Type species: not fixed.

NOTE: Pachymasiphis was described by Karg, 1996: 183, but the name was not made

available because the type species of the genus was not fixed (International Code of

Zoological Nomenclature, Article 13.3).

003. Pachymasiphis maior Karg, 1996

Pachymasiphis maior Karg, 1996: 185.




litter of a deciduous forest

004. Pachymasiphis porulatus Karg, 1996

Pachymasiphis porulatus Karg, 1996: 184.



TYPE LOCALITY AND HABITAT: New Caledonia, Lifou, [Loyalty Islands], 1977, in leaf

litter of a primary forest.

Species previously considered in genera of Ologamasidae and now placed in other

families

Cyrtolaelaps agilis Berlese, 1916b: 299 [Veigaia agilis (Veigaiidae) — Evans, 1955: 584].

Cyrtolaelaps (?) aurantiacus [sic] Berlese, 1903b: 241 [Gamasolaelaps aurantiacus

(Veigaiidae) — Berlese, 1903b: 241].

Cyrtolaelaps bouvieri Berlese, 1916a: 158 [Veigaia bouvieri (Veigaiidae) — Evans, 1955:

577].

359

Cyrtolaelaps capreolus Berlese, 1905a: 168 [Veigaia capreolus (Veigaiidae) — Vitzthum,

1925: 5].

Cyrtolaelaps captator Berlese, 1892c: 8 [junior synonym of Dendrolaelaps

(Punctodendrolaelaps) bisetus (Berlese, 1891) (Digamasellidae) — Hirschmann and

Wiśniewski, 1982: 59].

Cyrtolaelaps exiguus Berlese, 1916b: 300 [Veigaia exiguus (Veigaiidae) — Evans, 1955:

584].

Cyrtolaelaps goliathus Berlese, 1910c: 372 [junior synonym of Eugamasus immanis (Berlese,

1904) (Parasitidae) — Ryke, 1962b: 13].

Cyrtolaelaps gracilipes Banks, 1916: 228 [Pseudoparasitus gracilipes (Laelapidae) —

Halliday, 1998: 129].

Cyrtolaelaps grandipes Berlese, 1916b: 298 [Veigaia grandipes (Veigaiidae) — Ryke, 1962b:

12].

Cyrtolaelaps herculeanus Berlese, 1903b: 240 [Veigaia herculeana (Veigaiidae) —

Oudemans, 1905: 6].

Cyrtolaelaps humilis Hull, 1918: 75 [Veigaia humilis (Veigaiidae) — Evans, 1955: 583].

Cyrtolaelaps ibex Berlese, 1905b: 233 [Veigaia ibex (Veigaiidae) — Evans, 1955: 583].

Cyrtolaelaps kochi Tragardh, 1901: 61 [Veigaia kochi (Veigaiidae) — Willmann, 1932: 160]

NOTE: Name proposed to replace the species described by Koch (1879): 119 as Gamasus

emarginatus.

Cyrtolaelaps mitis Berlese, 1916b: 300 [Veigaia mitis (Veigaiidae) — Ryke, 1962b: 12].

Cyrtolaelaps nemorensis var. planicola Berlese, 1892a: 6 [Veigaia planicola (Veigaiidae) —

Evans, 1955: 583].

360

Cyrtolaelaps pusillus Berlese, 1916b: 301 [Veigaia pusilla (Veigaiidae) — Ryke, 1962b: 12].

Cyrtolaelaps transisalae Oudemans, 1902a: 28 [Veigaia transisalae (Veigaiidae) —

Trägårdh, 1931b: 13].

Cyrtolaelaps (Gamasellus) punctum Berlese, 1904: 262 [Digamasellus punctum

(Digamasellidae) — Lindquist, 1975: 13].

Gamasellus aeronauta Vitzthum, 1918: 12 [Halolaelaps aeronautus (Halolaelapidae) —

Karg, 1971: 296].

Gamasellus americanus Garman, 1948: 9 [junior synonym of Gamasellodes bicolor (Berlese,

1918) (Ascidae) — Hurlbutt, in Baker, Delfinado and Abbatiello, 1976: 63].

Gamasellus mitigatus Berlese, 1923a: 250 [Afrogamasellus mitigatus (Rhodacaridae) —

Loots and Ryke, 1968: 2].

Gamasellus octoclavatus Vitzthum, 1918: 14 [Halolaelaps octoclavatus (Halolaelapidae) —

Karg, 1971: 296].

Gamasellus quadripilus Berlese, 1920b: 159 [Dendrolaelaps quadripilus (Digamasellidae) —

Hirschmann, 1974: 61].

Gamasellus quadrisigillatus Berlese, 1916a: 160 [Afrogamasellus quadrisigillatus

(Rhodacaridae) — Loots and Ryke, 1968: 2].

Gamasellus succinctus Berlese, 1916a: 160 [Afrogamasellus succinctus (Rhodacaridae) —


Gamasellus tetrastigma Berlese, 1916a: 161 [Afrogamasellus tetrastigma (Rhodacaridae) —


361

Gamasellus viator Vitzthum, 1921: 7 [Dendrolaelaps (Dendrolaelaps) viator

(Digamasellidae) — Hirschmann and Wiśniewski, 1982: 82].

Gamasellus (Digamasellus) bicolor Berlese, 1918: 135 [Gamasellodes bicolor (Ascidae) —

Hurlbutt, 1970: 475].

Gamasellus (Digamasellus) capensis Berlese, 1920b: 161 [Dendrolaelaps capensis

(Digamasellidae) — Hirschmann, 1960: 1].

Gamasellus (Digamasellus) cylindricus Berlese, 1918: 135 [Dendrolaelaps cylindricus


Gamasellus (Digamasellus) debilipes Berlese, 1920b: 160 [Dendrolaelaps debilipes


Gamasellus (Digamasellus) gracilis Berlese, 1920b: 159 [Digamasellus gracilis

(Digamasellidae) — Bernini, Castagnoli and Nannelli, 1995: 18].

Gamasellus (Digamasellus) innumerus Berlese, 1920b: 161 [Dendrolaelaps innumerus


Gamasellus (Digamasellus) magnituberculatus Vitzthum, 1925: 5 [Asca magnituberculata

(Ascidae) — Wharton, 1946: 180].

Gamasellus (Digamasellus) perpusillus Berlese, 1905: 234 [junior synonym of Digamasellus

punctum (Berlese, 1904) (Digamasellidae) — Lindquist, 1975: 13].

Gamasellus (Digamasellus) presepum Berlese, 1918: 136 [Dendrolaelaps presepum


Gamasellus (Digamasellus) quadricrinus Berlese, 1920b: 162 [Dendrolaelaps quadricrinus


362

Gamasellus (Digamasellus) quadrisetus Berlese, 1920b: 159 [Insectolaelaps quadrisetus

(Digamasellidae) — Shcherbak, 1980: 193].

Gamasellus (Digamasellus) reticulatus Berlese, 1920b: 161 [Dendrolaelaps reticulatus


Gamasellus (Digamasellus) rhodacaroides Berlese, 1920b: 162 [Digamasellus rhodacaroides


Gamasellus (Digamasellus) simplex Berlese, 1920b: 163 [Digamasellus simplex


Gamasellus (Digamasellus) validulus Berlese, 1920b: 163 [Dendrolaelaps validulus


Gamasellus (Sessiluncus) eremita Berlese, 1918: 137 [Arctoseius eremitus (Ascidae) —

Evans, 1958: 223].

Gamasellus (Sessiluncus) latus Berlese, 1905a: 168 — Pachylaelapidae.

Gamasellus (Sessiluncus) solitarius Berlese, 1905a: 169 — Pachylaelapidae.

Gamasus calcaratus Koch, 1839c: 6 (cited as Ologamasus calcaratus.— Berlese, 1920a: 8)

— [Holoparasitus calcaratus (Parasitidae) — Oudemans, 1936: 165].

Gamasus cervus Kramer, 1876: 83 (cited as Cyrtolaelaps cervus.— Berlese, 1892c: 10) —

[Veigaia cervus (Veigaiidae) — Oudemans, 1905: 6].

Gamasus nemorensis Koch, 1839a: 18 (cited as Cyrtolaelaps nemorensis.— Berlese, 1892a:

5) — [Veigaia nemorensis (Veigaiidae) — Oudemans, 1905: 6].

Gamasus salinus Laboulbène, 1851: 297 (cited as Hydrogamasus salinus.— Oudemans,

1902b: 286; Hirschmann, 1966c: 28; Karg, 1971: 352) — [Pergamasus salinus

(Parasitidae) — Oudemans, 1936: 163].

363

Gamasus (Hologamasus) pollicipatus Berlese, 1903b: 238 — Parasitidae.

Gamasus (Ologamasus) calcaratus var. siculus Berlese, 1906: 248 — Parasitidae.

Gamasus (Ologamasus) calcaratus (?) var. excisus Berlese, 1906: 249 — Parasitidae.

Gamasus (Ologamasus) inornatus Berlese, 1906: 257 (cited as Ologamasus inornatus –

Berlese, 1916a: 156) — Parasitidae.

Gamasus (Ologamasus) pollicipatus var. apenninorum Berlese, 1906: 253 — Parasitidae.

Gamasus (Ologamasus) pollicipatus var. cultriger Berlese, 1906: 256 — Parasitidae.

Gamasus (Ologamasus) pollicipatus var. excipuliger Berlese, 1906: 252 — Parasitidae.

Gamasus (Ologamasus) pollicipatus var. peraltus Berlese, 1906: 256 — Parasitidae.

Gamasus (Ologamasus) pollicipatus var. pseudoperforatus Berlese, 1906: 254 — Parasitidae.

Holoparasitus coronarius Karg, 1971: 355 (cited as Ologamasus coronarius.— Schmölzer,

1991: 350) — [Holoparasitus coronarius (Parasitidae) — Karg, 1993c: 388].

Hydrogamasus silvestrii Berlese, 1903a: 27 [junior synonym of Pergamasus hamatus (Koch)

(Parasitidae) — Berlese, 1906: 289.

Iphidosoma belovae Davydova, 1975: 113 [Poecilochirus belovae (Parasitidae) — Mašán,

1999: 523].

Iphidosoma bennwili Schweizer, 1961: 145 [junior synonym of Uroiphis scabratus Berlese

(Eviphididae) — Mašán and Halliday, 2010: 91].

Iphidosoma insolentis Ma, 1997a: 39 [Alloseius insolentis (Eviphididae) — Mašán and

Halliday, 2009: 50].

364

Iphidosoma pratensis Karg, 1965: 263 [Alloseius pratensis (Eviphididae) — Mašán and

Halliday, 2009: 50].

Ologamasus absoloni Willmann, 1940: 212 [junior synonym of Holoparasitus

hemisphaericus (Vitzthum) (Parasitidae) — Witaliñski, 2006: 17].

Ologamasus hemisphaericus Vitzthum, 1923: 101 [Holoparasitus hemisphaericus

(Parasitidae) — Witaliñski, 2006: 17].

Ologamasus karawankianus Schmölzer, 1991: 349 [Holoparasitus karawankianus

(Parasitidae) — Karg, 1993c: 389].

Ologamasus longisetosus Schmölzer, 1995b: 102 [Holoparasitus longisetosus (Parasitidae)

— Schmölzer, 1998: 119].

Ologamasus rotulifer Willmann, 1940: 212 — Parasitidae.

Ologamasus (Ologamasiphis) minimus Holzmann, 1969: 49 [Holoparasitus minimus

(Parasitidae) — Karg, 1971: 360].

Ologamasus (Ologamasus) intermedius Holzmann, 1969: 50 — Parasitidae.

Parasitus sexclavatus Oudemans, 1903: 74 (cited as Gamasellus sexclavatus.— Oudemans,

1905: 6) — [Halolaelaps sexclavatus (Halolaelapidae) — Karg, 1971: 285].

Seius excisus Koch, 1879: 122 (cited as Cyrtolaelaps excisus.— Trägårdh, 1901: 61) —

[Gamasolaelaps excisus (Veigaiidae) — Halbert, 1920: 114].

Zercon bisetus Berlese, 1891: 7 (cited as Cyrtolaelaps bisetus.— Oudemans, 1902a: 29) —

[Dendrolaelaps bisetus (Digamasellidae) — Hirschmann and Wiśniewski, 1982: 59].

365

3.3.5 Catalogue of world species of Teranyssidae Halliday

Genus Teranyssus Halliday, 2006

Teranyssus Halliday, 2006: 33 (described in Teranyssidae Halliday).

Type species: Teranyssus howardensis Halliday, 2006, by original designation.

001. Teranyssus howardensis Halliday, 2006

Teranyssus howardensis Halliday, 2006: 33.


TYPE LOCALITY AND HABITAT: Australia, Howard Springs, 21 km SE of Darwin,

Northern Territory, 7 July 1985, in nest of Mastotermes darwiniensis [Isoptera:

Mastotermitidae].

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409

4 REVISION OF THE GENERA Interrhodeus, Pennarhodeus and Poropodalius

(ACARI: RHODACARIDAE)

Abstract

The genera Interrhodeus Karg, Pennarhodeus Karg and Poropodalius Karg are

redescribed on the basis of one species of Interrhodeus, four species of Pennarhodeus and

five species of Poropodalius, including the type species of each genus. All species are

redescribed on the basis of the type specimens, and keys for the separation of the species of

the latter two genera are provided. Re-examination of these species shows that these three

genera are correctly placed in the family Rhodacaridae.

Keywords: Rhodacaroidea; Soil mites; Taxonomy

Resumo

Os gêneros Interrhodeus Karg, Pennarhodeus Karg e Poropodalius Karg são

redescritos com base em uma espécie de Interrhodeus, quatro espécies de Pennarhodeus e

cinco espécies de Poropodalius, incluindo as espécies tipo de cada gênero. Todas as espécies

são redescritas com base em espécimes-tipo. As chaves para a separação das espécies dos dois

últimos gêneros são fornecidas. A reavaliação dessas espécies mostra que esses três gêneros

estão corretamente colocados na família Rhodacaridae.

Palavras-chave: Rhodacaroidea; Ácaros de solo; Taxonomia

4.1 Introduction

The Rhodacaridae Oudemans is a family of free living edaphic mites found mainly in

the top few centimetres of the soil surface. They have been commonly mentioned in the

literature as predators (LEE, 1970; LINDQUIST; KRANTZ; WALTER, 2009). The limited

information available in the literature shows that they can feed on insect larvae, springtails,

nematodes and mites (KARG, 1971; LEE, 1974; WALTER; HUNT; ELLIOTT, 1988;

CASTILHO et al., 2009).

410

The genera of Rhodacaridae have often been confused with each other and with

genera in several related families. That instability has been partly caused by incomplete

descriptions that do not define important diagnostic features of the taxa. The genera

Interrhodeus Karg, Pennarhodeus Karg and Poropodalius Karg consist of one, four and five

described species respectively. All of these species have been reported only from Central or

South America (KARG, 2000a, 2000b; KARG; SCHORLEMMER, 2009). The original

descriptions of these species and their respective genera are not sufficiently detailed, making

it difficult to decide whether these genera really belong to the Rhodacaridae. Thus, a re-

examination of the types of those species was considered necessary to allow the completion of

a catalogue of the Rhodacaridae (CASTILHO; MORAES; HALLIDAY, 2012), and to allow

the correct identification of mites of those groups in the American continent. The objective of

this paper is to provide redescriptions of these genera and their included species, and the

preparation of keys for identification of species of Pennarhodeus and Poropodalius.

4.2 Material and methods

The type specimens of species of these genera were borrowed from the

Arachnologische Sammlung des Museums für Naturkunde der Humboldt-Universität, Berlin

(MNHB) and the Staatliches Museum für Naturkunde Görlitz, Görlitz (SMNG), both in

Germany. They were examined under a phase contrast microscope provided with a camera

lucida, in the University of Amsterdam, The Netherlands. The specimens were illustrated and

measurements were taken of structures considered taxonomically important. In the following

redescriptions, setal nomenclature is based on Lindquist and Evans (1965). Measurements of

each structure are given in micrometres (µm) and as a range (or a single value when

measurement did not vary) representing the variation among all individuals examined. Leg

length was measured from the proximal edge of the coxa to the tip of the pretarsus.

4.3 Results

Genus Interrhodeus Karg

Interrhodeus Karg, 2000a: 258.

Type species: Interrhodeus brevicornus Karg, 2000, by original designation.

411

Diagnosis (adult female): Arthrodial process of chelicera with a short coronet-like fringe;

epistome with a serrate antero-medial projection of about uniform width, flanked on each side

by a slightly longer and serrate projection; seta h3 directly posterior to h1 and distinctly

postero-mediad to h2; podonotal shield with 22 pairs of setae (setae s1 and r1 absent) and

without scleronoduli; opisthonotal shield with 20 pairs of setae; all dorsal idiosomal setae

smooth; laciniae of tritosternum separated from each other only in the distal half; presternal

shields absent; ventrianal shield with five pairs of setae in addition to the circum-anal setae;

unsclerotised integument around ventrianal shield with two pairs of setae; with a pair of

elongate metapodal plates; peritreme extending anteriorly to mid-level of coxa II; pretarsus I

present; seta pl4 of tarsus IV absent. Numbers of setae on legs I-IV: coxa: 2, 2, 2, 1;

trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6; genu: 13, 11, 9, 10; tibia: 14, 10, 8, 10; tarsus II-IV:

18, 18, 17.

Interrhodeus brevicornus Karg, 2000

Interrhodeus brevicornus Karg, 2000a: 259.

Specimens examined: Holotype female: 6953 T, IB1418905, La Selva, Heredia, Costa Rica

(10°26‘0‖N 84°1‘0‖W), elev. 50-150 m, Coll. Alas, 6-XI-1993, ZMB Kat. Nr. 46010, [Costa

Rica INBIO CRI001418905]. Paratypes: one female, 6954, IB1418902, ZMB Kat. Nr. 46011,

[Costa Rica INBIO CRI001418902]; one female, 6955, IB1418922, ZMB Kat. Nr. 46012,

[Costa Rica INBIO CRI001418922]; one female, 6956, IB1419397, Ex. Hernandia

didymantha Hernandiaceae, 12-X-1994, ZMB Kat. Nr. 46013, [Costa Rica INBIO

CRI001419397]; one female, 6957, IB1422099, Suampo experimental, 20-I-1995, ZMB Kat.

Nr. 46014, [Costa Rica INBIO CRI001422099]; one deutonymph, D-N 6958, IB1419233,

SSO 350m, 3-IX-1993, ZMB Kat. Nr. 46015, [Costa Rica INBIO CRI001419233]; one

deutonymph, D-N 6959, IB1418871, ZMB Kat. Nr. 46016, [Costa Rica INBIO

CRI001418871] (for all paratypes, other data as for holotype). (in MNHB).

Diagnosis of adult: As for the genus.

Adult female (Fig. 4.1 A-F) (holotype and four paratypes): Dorsal and ventral idiosomal,

hypostomal, subcapitular and leg setae smooth.

412

Gnathosoma: Fixed cheliceral digit 34 long, with five teeth in addition to apical tooth and a

setiform pilus dentilis (Fig. 4.1A); movable cheliceral digit 35 long, with four teeth in

addition to apical tooth. Arthrodial process of chelicera with a short coronet-like fringe.

Epistome with a serrate antero-medial projection of about uniform width and provided with

two apical denticles, flanked on each side by a slightly longer, triangular and serrate

projection (Fig. 4.1B); margin of epistome smooth between antero-medial and antero-lateral

projections, and serrate laterad to antero-lateral projections. Deutosternal denticles in eight

rows, with seven to 13 denticles each; anterior row anteriorly convex, other rows straight and

roughly transverse, second last row narrow (Fig. 4.1C). Measurements of setae: h1 14-17, h2

13-15, h3 10-12, sc 11-14.

Dorsal idiosoma (Fig. 4.1D): Podonotal shield ornamented with roundish markings, except

for a punctate band along posterior margin; 158-163 long and 171-185 wide at widest point;

with 22 pairs of setae (setae s1 and r1 absent); without scleronoduli and with five pairs of

distinguishable lyrifissures. Opisthonotal shield imbricate, with roundish markings behind

setae Z4 and a punctate band along anterior margin; 131-146 long and 135-142 wide at widest

point; with 20 pairs of setae and four pairs of distinguishable lyrifissures. Measurements of

setae: j1 17-19, j2 18-19, j3 15-16, j4 13-15, j5 14-15, j6 13-14, z1 18-19, z2 16-18, z3 14-15,

z4 14-15, z5 12-14, z6 12-13, s2 15-16, s3 14-15, s4 14-15, s5 13-15, s6 12-13, r2 14-15, r3

18-20, r4 13-14, r5 13-14, r6 14-15, J1 12-13, J2 12-13, J3 12-14, J4 13-15, J5 15-16, Z1 13-

14, Z2 13-14, Z3 13-15, Z4 15-16, Z5 19-21, S1 12-14, S2 12-13, S3 14-15, S4 16-17, S5 19-

20, R1 12-13, R2 12-14, R3 14-15, R4 16-17, R5 19-20.

Ventral idiosoma (Fig. 4.1E): Base of tritosternum 14-21 long and 12-13 wide proximally

(Fig. 4.1F); laciniae 43-51 long, separated for about half of their total length, pilose. Sternal

shield lightly reticulate and with indistinct anterior margin; region anterior to the first pair of

lyrifissures (iv1) lightly sclerotised and punctate; posterior margin straight; approximately 88-

93 long, including the lightly sclerotised and punctate region, and 73-77 wide at widest point;

with four pairs of setae and three pairs of lyrifissures. Triangular remnants of exopodal shield

distinguishable between coxae II-III and III-IV. Elongate remnant of endopodal shield

distinguishable adjacent to coxa IV and slightly anterior to it. Genital shield with light striae

parallel to lateral margins, and a punctate band along most of its slightly convex posterior

margin; longer than wide; extending posteriorly behind coxa IV; distance between st5-st5 45-

48. Unsclerotised integument posterolaterad to st5 with a pair of lyrifissures. Ventrianal shield

413

Figure 4.1 - Interrhodeus brevicornus Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.

Hypostome; D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures enlarged for

improved visibility

414

reticulate, except for a punctate band along anterior margin; 91-102 long and 90-102 wide at

widest point, not fused to dorsal shield; with five pairs of setae (Jv1-Jv3, Zv1 and Zv2) in

addition to circum-anal setae; post-anal seta as long as para-anal setae; lyrifissures not

distinguishable. Unsclerotised integument around ventrianal shield with two pairs of setae

(Jv5 and Zv3) and a pair of elongate metapodal plates. Peritreme extending anteriorly to mid-

level of coxa II. Peritrematal shield fused with a section of exopodal shield next to coxa IV.

Measurements of setae: st1 18-20, st2 17-18, st3 16-18, st4 14-16, st5 14-15, Jv1 14-15, Jv2

15-16, Jv3 16-17, Jv5 14-16, Zv1 13-14, Zv2 15-16, Zv3 14-15, para-anal 15-16, post-anal 16-

18.

Legs: Lengths: I: 303-312; II: 223-234; III: 203-220; IV: 281-295. Pretarsi I-IV similar in

shape and length, an elongate ambulacral stalk, a pair of strong claws and three rounded

pulvillar lobes.

Deutonymph (Fig. 4.2 A-B) (two paratypes): Shape of setae as in adult female.

Gnathosoma: Fixed cheliceral digit 28-29 long, with five teeth in addition to apical tooth,

and a setiform pilus dentilis; movable cheliceral digit 28-30 long, with four teeth in addition

to apical tooth. Arthrodial process of chelicera with a short coronet-like fringe. Epistome,

deutosternum and position of hypostomal setae as in female. Measurements of setae: h1 16,

h2 12-13, h3 10, sc 14-15.

Dorsal idiosoma (Fig. 4.2A): Podonotal shield imbricate laterad to j setal series and with

roundish markings between j series; 132-133 long and 143-147 wide at widest point; with 21

pairs of setae (setae s1 and r1 absent); without scleronoduli and with five pairs of

distinguishable lyrifissures. Unsclerotised integument laterad to podonotal shield with a pair

of setae (r4). Opisthonotal shield mostly imbricate, except for a narrow band of roundish

markings along anterior margin; 110-122 long and 104 wide at widest point; with 15 pairs of

setae and four pairs of distinguishable lyrifissures. Unsclerotised integument laterad to

opisthonotal shield with five pairs of setae (R1-R5). Measurements of setae: j1 16-17, j2 18, j3

15, j4 13, j5 13-14, j6 11-13, z1 15-16, z2 15-16, z3 12-13, z4 12-13, z5 11-12, z6 11-12, s2

13, s3 12-13, s4 13, s5 12-13, s6 12, r2 13-14, r3 16-17, r4 11, r5 11-12, r6 11, J1 11-12, J2

11, J3 10-11, J4 11, J5 12, Z1 11-12, Z2 10-11, Z3 11-12, Z4 13-14, Z5 19-21, S1 11-12, S2

10-11, S3 10-11, S4 13, S5 15, R1 9-10, R2 8, R3 8-9, R4 10, R5 12-14.

415

Ventral idiosoma (Fig. 4.2B): Base of tritosternum 15-17 long and 11-13 wide proximally;

laciniae 40-41 long, otherwise as in adult female. Sternal shield smooth, narrowing behind

coxae II, with slightly rounded posterior margin; approximately 119-123 long and 58-59 wide

at widest point; with four pairs of setae and three pairs of lyrifissures. Setae st5 inserted on

unsclerotised integument at level of posterior end of sternal shield. Ventrianal shield

imbricate; 45-56 long along midline and 57-58 wide at widest point; with two pairs of setae

(Jv2 and Jv3) in addition to circum-anal setae; post-anal seta about as long as para-anal setae.

Unsclerotised integument around ventrianal shield with five pairs of setae (Jv1, Jv4 and Zv1-

Zv3). Peritreme extending anteriorly to mid-level of coxa II. Peritrematal shield indistinct.

Measurements of setae: st1 17-18, st2 16-17, st3 13-14, st4 11-13, st5 9-11, Jv1 11-13, Jv2 12,

Jv3 13, Jv4 10-11, Zv1 10-11, Zv2 10-11, Zv3 8-9, para-anal 13-15, post-anal 15-16.

Legs: Lengths: I: 267-271; II: 198-201; III: 180-181; IV: 236-241. Leg chaetotaxy and

pretarsi as in adult female.

Figure 4.2 - Interrhodeus brevicornus Karg. Deutonymph. A. Dorsal idiosoma; E. Ventral idiosoma. Lyrifissures

enlarged for improved visibility

416

Remarks

This species was described on the basis of the adult female holotype, 52 adult female

paratypes and seven deutonymph paratypes. The following characteristics appear in the

original description and illustrations of the species, but do not agree with our observations of

female type specimens: movable cheliceral digit with three teeth in addition to apical tooth;

most anterior row of deutosternal denticles roughly transverse; podonotal shield imbricate,

with 21 pairs of setae, and with structures suggesting the presence of scleronoduli [Karg

(2000a) mentioned that scleronoduli were not clear]; unsclerotised integument laterad to

podonotal shield with two pairs of setae; sternal shield smooth; exopodal shield consisting of

a single structure between coxae II-IV; no endopodal shields; genital shield smooth;

lyrifissures present only on podonotal (two pairs) and sternal shields (three pairs). No

information was provided about the tritosternum. The only measurements provided in the

original description referred to the length of the idiosoma (270-290), width of the idiosoma

(140-180), length of seta J5 (cited as I5) (18), range of lengths of other dorsal idiosomal setae

(15-16) and length of all st setae (18). Leg setal counts were given only for tibia IV (10 setae)

and genu IV (10 setae).

The deutonymph was illustrated in the original description of the species, but not

described (Figure 23 in Karg, 2000a). The following characteristics are present in the original

illustrations of the deutonymph, but do not agree with our observations of the paratype

specimens: podonotal shield smooth and with 23 pairs of setae; opisthonotal shield reticulate;

ventrianal shield smooth; with a longitudinal line extending from the anterior end of the

peritreme to the level of coxa I; lyrifissures present only on the sternal shield (two pairs).

Genus Pennarhodeus Karg

Pennarhodeus Karg, 2000a: 255.

Pennarhodeus.— Karg, 2000b: 211.

Type species: Pennarhodeus pennatus Karg, 2000, by original designation.

Diagnosis (adult female and male): Arthrodial process of chelicera with a short coronet-like

fringe (except P. decoris, arthrodial process elongate, with three points); epistome with a

smooth antero-medial projection either of about uniform width or slightly wider proximally,

flanked on each side by a shorter projection whose lateral margins are smooth or lightly

serrate; seta h3 directly posterior to h1 and mediad to h2; setae dorsal, circumanal and Jv4

usually pilose and other setae smooth; podonotal shield with 22-23 pairs of setae and without

417

scleronoduli (except male of P. brevipennatus, with two scleronoduli); unsclerotised

integument laterad to podonotal shield with zero or one pair of setae; opisthonotal shield with

15-17 pairs of setae; with four or five pairs of R setae on unsclerotised integument laterad to

opisthonotal shield in female and two to five pairs of R setae on unsclerotised integument

laterad to opisthonotal shield or on ventrianal shield in male; most dorsal idiosomal setae

pilose; presternal shields absent; ventrianal shield with five pairs of setae in female and seven

to nine pairs of setae in male; unsclerotised integument along margins of ventrianal shield

with two pairs of setae in female and without setae in male; with a pair of rounded metapodal

plates in female and without metapodal plates in male; peritreme extending anteriorly to level

between anterior margin of coxa III and median region of coxa II; pretarsus I, a single, sessile,

curved claw; seta pl4 of tarsus IV absent. Numbers of setae on legs I-IV: coxa: 2, 2, 2, 1;

trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6; genu: 13, 11, 9, 10 (9 in P. turris Karg); tibia: 14,

10, 8, 10 (9 in P. brevipennatus Karg); tarsus II-IV: 18, 18, 17. Adult male with

spermatodactyl abruptly bent dorsally or ventrally and with one or two spur-like ventral setae

on each of femur, genu and tibia II.

Pennarhodeus brevipennatus Karg

Pennarhodeus brevipennatus Karg, 2000b: 211.

Specimens examined: Holotype male: La Selva, Costa Rica (10°26‘N 84°01‘W), elevation

50-150 m, Primario (indicated as soil sample in the original description), 3-VIII-1993, Coll.

B., SMNG 93/38525 (in SMNG).

Diagnosis (adult male): Arthrodial process of chelicera with a short coronet-like fringe;

epistome with antero-medial projection about 1.2 times as long as antero-lateral projections,

and with margin serrate laterad to antero-lateral projections; corniculi rather straight; dorsal

idiosomal setae pilose, except r3, Z5 and S5, feather-like; podonotal and opisthonotal shields

imbricate; with two crescent-shaped scleronoduli between setae j5 and j6; posterior margin of

opisthonotal shield slightly concave between setae Z5; seta R5 absent; sternogenital shield

lightly reticulate; unsclerotised integument between sternogenital and ventrianal shields with

a subtriangular accessory platelet; peritreme extending anteriorly to level of posterior margin

of coxa II; with two spur-like ventral setae on each of femur, genu and tibia II.

418

Adult male (Fig. 4.3 A-G) (holotype): Dorsal idiosomal setae (except r3, Z5 and S5), para-

anal and post-anal setae pilose; setae r3, Z5, S5 and Jv5 feather-like; hypostomal,

subcapitular, ventral idiosomal (except Jv5, para-anal and post-anal) and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 36 long; movable digit 37 long; teeth and pilus dentilis

not visible because of position of chelicera. Spermatodactyl 77 long, abruptly bent ventrally at

an acute angle, distal portion narrower than proximal portion (Fig. 4.3A). Arthrodial process

of chelicera with a short coronet-like fringe. Epistome with a smooth antero-medial projection

slightly wider at the base and provided with three apical denticles, flanked on each side by

shorter, smooth projection, each provided with four or five apical denticles (Fig. 4.3B);

margin of epistome with two or three large teeth between antero-medial and antero-lateral

projections and serrate laterad to antero-lateral projections. Deutosternal denticles in eight

rows, each with seven to 13 denticles; anterior row ―V‖ shaped, other rows straight and

transverse (Fig. 4.3C). Corniculi rather straight. Measurements of setae: h1 14, h2 10, h3 11,

sc 12.

Dorsal idiosoma (Fig. 4.3D): Podonotal shield imbricate, except for a punctate band along

posterior margin; 160 long and 173 wide at widest point; with 22 pairs of setae (seta r1

absent); with two crescent-shaped scleronoduli between setae j5 and j6 and four pairs of


of setae (r4). Opisthonotal shield imbricate, except for a punctate band along anterior margin;

posterior margin slightly concave between setae Z5; 125 long and 141 wide at widest point;

with 18 pairs of setae and three pairs of distinguishable lyrifissures. Seta R1 on ventrianal

shield; seta R5 absent. Measurements of setae: j1 9, j2 8, j3 8, j4 10, j5 10, j6 11, z1 8, z2 not

seen (broken), z3 10, z4 10, z5 not seen (broken), z6 not seen (broken), s1 11, s2 9, s3 9, s4

11, s5 10, s6 10, r2 11, r3 15, r4 11, r5 not seen (broken), r6 14, J1 not seen (broken), J2 not

seen (broken), J3 not seen (broken), J4 11, J5 12, Z1 12, Z2 13, Z3 13, Z4 13, Z5 14, S1 12,

S2 11, S3 12, S4 12, S5 13, R1 8, R2 9, R3 10, R4 11.

Ventral idiosoma (Fig. 4.3E): Base of tritosternum 15 long and 12 wide proximally (Fig.

3F); laciniae 46 long, separated for about 90% of their total length, pilose. Sternogenital

shield lightly reticulate, with indistinct anterior margin and punctate band along straight

posterior margin; region anterior to the first pair of lyrifissures (iv1) lightly sclerotised and

419

Figure 4.3 - Pennarhodeus brevipennatus Karg. Male. A. Lateral (antiaxial) view of spermatodactyl; B.

Epistome; C. Hypostome; D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum; G.

Anterolateral view of femur, genu and tibia of leg II. Lyrifissures enlarged for improved

visibility

420

punctate; approximately 132 long, including the lightly sclerotised and punctate region, and

65 wide at widest point; with five pairs of setae and three pairs of lyrifissures; distance

between st5-st5 28; genital opening on anterior margin of shield. Elongate remnant of

endopodal shield adjacent to coxa IV. Unsclerotised integument between sternogenital and

ventrianal shields with a subtriangular accessory platelet and punctate near anterior margin of

ventrianal shield. Ventrianal shield imbricate; 95 long and 170 wide at widest point and

extending to dorso-lateral region of idiosoma, but not fused to dorsal shield; with nine pairs of

setae (R1, Jv1-Jv5 and Zv1-Zv3) in addition to circum-anal setae, a distinguishable pore and

two distinguishable lyrifissures; post-anal seta about as long as para-anal setae. Peritreme

extending anteriorly to level of posterior margin of coxa II. Peritrematal shield extending

posteriorly beyond mid-level of coxa IV. Measurements of setae: st1 13, st2 11, st3 13, st4 11,

st5 11, Jv1 11, Jv2 12, Jv3 12, Jv4 11, Jv5 12, Zv1 11, Zv2 11, Zv3 9, para-anal 13, post-anal

13.

Legs: Lengths: I: 240; II: 204; III: 177; IV: 230. Genu IV with ten setae and tibia IV with nine

setae. With two spur-like ventral setae on each of femur, genu and tibia II (Fig. 3G). Pretarsus

I, a single, sessile, curved claw; pretarsi II-IV, an elongate ambulacral stalk, a pair of strong

claws and three rounded pulvillar lobes.

Remarks

This species is known only from the adult male holotype. The following

characteristics appear in the original description and illustrations of the species, but do not

agree with our observations of the holotype: margin of epistome smooth between antero-

medial and antero-lateral projections; podonotal shield with 22 pairs of setae and four

crescent-shaped scleronoduli between setae j5 and j6; unsclerotised integument laterad to

podonotal shield without setae; opisthonotal shield with 17 pairs of setae; unsclerotised

integument laterad to opisthonotal shield with a pair of setae. Karg (2000b) provided no

information about the hypostome, tritosternum, ventral idiosoma, lyrifissures and leg setal

counts. The only measurements provided in the original description referred to the length of

the idiosoma (290), width of idiosoma (180), length of seta Z5 (13), range of other dorsal

idiosomal setae (8-10), length of legs I (220), II (190), III (170) and IV (220).

Pennarhodeus decoris Karg

Pennarhodeus decoris Karg, 2000a: 255.

421

Pennarhodeus decoris.— Karg, 2000b: 211.

Specimens examined: Holotype female: 6972 T, Kuba (= Cuba), 1976, Bodenpr. Nr. (soil

sample number) 48, ZMB Kat. Nr. 45977. (in MNHB).

Diagnosis (adult female): Arthrodial process of chelicera elongate, three-pointed; epistome

with three, well separated projections, the antero-medial about 1.2 times as long as the antero-

lateral ones; epistome margin irregularly serrate; corniculi rather straight; dorsal idiosomal

setae pilose; podonotal and opisthonotal shields imbricate; without scleronoduli; seta R5

present; genital shield lightly reticulate; unsclerotised integument between genital and

ventrianal shields with a transverse elongate accessory platelet; peritreme extending anteriorly

to level of anterior margin of coxa III.

Adult female (Fig. 4.4 A-F) (holotype): Dorsal idiosomal setae, Jv5, para-anal and post-anal

setae pilose; hypostomal, subcapitular, ventral idiosomal setae (except Jv5, para-anal and

post-anal) and leg setae smooth.

Gnathosoma: Chelicerae narrow and elongate (Fig. 4.4A); fixed cheliceral digit 88 long, with

three teeth in addition to apical tooth and a setiform pilus dentilis; movable cheliceral digit 90

long, with three teeth in addition to apical tooth. Arthrodial process of chelicerae elongate,

three-pointed. Epistome with three well separated projections; antero-medial projection

smooth of about uniform width and provided with two apical denticles; antero-lateral

projections slightly shorter, each provided with two or three lateral and three apical denticles

(Fig. 4.4B); epistome margin irregularly serrate. Deutosternal denticles in eight roughly

transverse rows, each with 11-16 denticles (Fig. 4.4C). With two pairs of rows of denticles

outside of the deutosternal groove, one between deutosternal denticle rows fifth and sixth and

the other in level with basal most row. Corniculi rather straight. Measurements of setae: h1

14, h2 10, h3 11, sc 12.

Dorsal idiosoma (Fig. 4.4D): Podonotal shield imbricate, except for a punctate band along

posterior margin; 217 long and 207 wide at widest point; with 22 pairs of setae (setae r1 and

r4 absent); without scleronoduli and with six pairs of distinguishable lyrifissures.

422

Figure 4.4 - Pennarhodeus decoris Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.


improved visibility

423

Opisthonotal shield imbricate, except for a punctate band along anterior margin; 150 long and

208 wide at widest point; with 15 pairs of setae and four pairs of distinguishable lyrifissures.

Unsclerotised integument laterad to opisthonotal shield with four pairs of setae (R2-R5); seta

R1 absent. Measurements of setae: j1 18, j2 25, j3 24, j4 26, j5 25, j6 24, z1 21, z2 27, z3 26,

z4 25, z5 25, z6 26, s1 17, s2 21, s3 25, s4 31, s5 31, s6 28, r2 29, r3 31, r5 27, r6 30, J1 26,

J2 27, J3 27, J4 32, J5 16, Z1 28, Z2 30, Z3 32, Z4 46, Z5 59, S1 32, S2 31, S3 32, S4 39, S5

48, R2 10, R3 7, R4 9, R5 8.


4.4F); laciniae 45, separated for about 90% of their total length, pilose. Sternal shield smooth,

except for a few light longitudinal lines along margin of posterior half, anterior margin

indistinct; region anterior to the first pair of lyrifissures (iv1) lightly sclerotised and punctate;

posterior margin slightly concave; approximately 115 long, including the lightly sclerotised

punctate region, 73 wide at widest point; with four pairs of setae and three pairs of

lyrifissures. Subtriangular remnants of exopodal shield present between coxae II-III and III-

IV. Genital shield lightly reticulate; longer than wide; extending posteriorly behind coxa IV;

distance between st5-st5 47; posterior margin straight. Unsclerotised integument between

genital and ventrianal shields with a transverse elongate accessory platelet, with setae Jv1 and

Zv1 and a pair of rounded metapodal plates. Ventrianal shield reticulate; 108 long and 158

wide at widest point, not fused to dorsal shield; with five pairs of setae (Jv2, Jv3, Jv5, Zv2 and

Zv3) in addition to circum-anal setae and three pairs of distinguishable lyrifissures; post-anal

seta longer than para-anal setae. Peritreme extending anteriorly to level of anterior margin of

coxa III. Peritrematal shield narrow, not extending beyond peritreme. Measurements of setae:

st1 12, st2 15, st3 15, st4 11, st5 14, Jv1 12, Jv2 13, Jv3 17, Jv5 44, Zv1 12, Zv2 17, Zv3 23,

para-anal 24, post-anal 35.

Legs: Lengths: I: 280; II: 220; III: 201; IV: 279. Genu and tibia IV each with ten setae.

Pretarsus I, a single, sessile, curved claw; pretarsi II-IV, an elongate ambulacral stalk, a pair

of strong claws and three rounded pulvillar lobes.

Remarks

This species is known only from the adult female holotype. The following


agree with our observations of the holotype: fixed cheliceral digit with two teeth in addition to

424

apical tooth; epistome with antero-medial projection about as long as antero-lateral

projections; podonotal shield without punctate band along posterior margin and with 20 pairs

of setae; opisthonotal shield without punctate band along anterior margin and with 15 pairs of

setae; unsclerotised integument laterad to opisthonotal shield with three pairs of setae; genital

shield smooth; peritrematal shield fused to podonotal shield; lyrifissures present only on

sternal shield (one pair). No information was provided about the tritosternum and leg setal

counts. The measurements provided in the original description referred to the length of the

movable cheliceral digit (90), length of the idiosoma (340), width of idiosoma (200), length of

setae j1 (cited as i1) (20), j5 (cited as i4) (25), r3 (cited as r5) (25), J1 (cited as I1) (25), Z4

(45), Z5 (53), S5 (50), range of all st (12-15), Jv5 (cited as V8) (45), post-anal (cited as Ps)

(35), range of other ventrianal setae (13-20), length of legs I (280), II (220), III (200) and IV

(280).

Pennarhodeus pennatus Karg

Pennarhodeus pennatus Karg, 2000a: 255.

Pennarhodeus pennatus.— Karg, 2000b: 211.

Specimens examined: Holotype female: 6973 T, Kuba (= Cuba), 1975, Bodenpr. (soil

sample) 8-9, ZMB Kat. Nr. 45980. Paratype male: 6974, Kuba (= Cuba), 1975, Bodenpr. (soil

sample) 11, ZMB Kat. Nr. 45981. (in MNHB).

Diagnosis (adult female and male): Adult female and male with arthrodial process of

chelicera with a short coronet-like fringe; epistome with antero-medial projection about 1.2

times as long as antero-lateral projections, and with margin smooth laterad to anterolateral

extensions; podonotal shield mostly with roundish markings except for reticulate antero-

lateral region, without scleronoduli; dorsal idiosomal setae pilose, except seta z1, smooth;

opisthonotal shield reticulate; seta R5 present, inserted on unsclerotised integument outside

lateral margins of opisthonotal shield; no accessory platelet between genital/ sternogenital and

ventrianal shields; peritreme extending anteriorly to mid-level of coxa II. Adult female with

genital shield smooth. Adult male with sternogenital shield lightly reticulate; with a spur-like

ventral seta on each of femur, genu and tibia II.

425

Adult female (Fig. 4.5 A-E) (holotype): Dorsal idiosomal setae (except z1), Jv5, para-anal

and post-anal setae pilose; setae z1, hypostomal, subcapitular, ventral idiosomal (except Jv5,

para-anal and post-anal) and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 35 long; movable cheliceral digit 36 long; teeth and pilus

dentilis not visible because of position of chelicera. Arthrodial process of chelicera with a

short coronet-like fringe. Epistome with a smooth antero-medial projection of about uniform

width and provided with five apical denticles, flanked on each side by a shorter, smooth

projection, each provided with three apical denticles (Fig. 4.5A); margin of epistome with one

or two denticles between antero-medial and antero-lateral projections, and smooth laterad to

antero-lateral projections. Deutosternal denticles in eight transverse rows, each with eight to

12 denticles (Fig. 4.5B). With two pairs of rows of denticles outside of the deutosternal

groove, slightly anterior to the sixth and to the seventh deutosternal denticle rows,

respectively. Measurements of setae: h1 11, h2 8, h3 9, sc 9.

Dorsal idiosoma (Fig. 4.5C): Podonotal shield mostly with roundish markings, except for

reticulate antero-lateral region and a punctate band along posterior margin; 156 long and 161

wide at widest point; with 22 pairs of setae (setae r1 and r4 absent); without scleronoduli and

with five pairs of distinguishable lyrifissures. Opisthonotal shield reticulate, except for a

punctate band along anterior margin; 146 long and 145 wide at widest point; with 15 pairs of

setae and four pairs of distinguishable lyrifissures. Unsclerotised integument laterad to

opisthonotal shield with five pairs of setae (R1-R5). Measurements of setae: j1 13, j2 14, j3

12, j4 11, j5 12, j6 13, z1 13, z2 12, z3 12, z4 13, z5 12, z6 14, s1 10, s2 13, s3 13, s4 14, s5

13, s6 12, r2 14, r3 20, r5 14, r6 15, J1 17, J2 17, J3 not seen (broken), J4 17, J5 9, Z1 18, Z2

18, Z3 18, Z4 17, Z5 16, S1 16, S2 15, S3 16, S4 17, S5 15, R1 6, R2 7, R3 7, R4 8, R5 9.

Ventral idiosoma (Fig. 4.5D): Base of tritosternum 15 long and 10 wide proximally (Fig.

4.5E); laciniae 31 long, separated for about 90% of their total length, pilose. Sternal shield

smooth and with indistinct anterior margin; region anterior to the first pair of lyrifissures (iv1)

lightly sclerotised and punctate; posterior margin straight; approximately 87 long, including

the lightly sclerotised punctate region, 70 wide at widest point; with four pairs of setae and

three pairs of lyrifissures. Genital shield smooth; slightly shorter than wide; extending

posteriorly behind coxae IV; distance between st5-st5 37; posterior margin straight.

Unsclerotised integument between genital and ventrianal shields with setae Jv1 and Zv1 and a

426

Figure 4.5 - Pennarhodeus pennatus Karg. Female. A. Epistome; B. Hypostome; C. Dorsal idiosoma; D. Ventral

idiosoma; E. Tritosternum. Lyrifissures enlarged for improved visibility

427

pair of rounded metapodal plates. Ventrianal shield reticulate; 88 long and 121 wide at widest

point, not fused to dorsal shield; with five pairs of setae (Jv2, Jv3, Jv5, Zv2 and Zv3) in

addition to circum-anal setae and three pairs of distinguishable lyrifissures; post-anal seta

longer than para-anal setae. Peritreme extending anteriorly to mid-level of coxa II.

Peritrematal shield narrow, not extending beyond peritreme. Measurements of setae: st1 13,

st2 14, st3 12, st4 13, st5 12, Jv1 11, Jv2 12, Jv3 not seen (broken), Jv5 11, Zv1 11, Zv2 12,

Zv3 12, para-anal 15, post-anal 25.

Legs: Lengths: I: 242; II: 191; III: 161; IV: 232. Genu and tibia IV each with ten setae.

Pretarsus I, a single, sessile, curved claw; pretarsi II-IV, an elongate ambulacral stalk, a pair

of strong claws and three rounded pulvillar lobes.

Adult male (Fig. 4.6 A-C) (paratype): Shape of setae as in adult female.


dentilis not visible because of position of chelicera. Spermatodactyl 48 long, abruptly bent

ventrally at an acute angle, tapering distally (Fig. 4.6A). Arthrodial process of chelicera with a

short coronet-like fringe. Epistome, deutosternum and position of hypostomal setae as in

female. Measurements of setae: h1 15, h2 8, h3 11, sc 9.

Dorsal idiosoma: Podonotal shield mostly with roundish markings, except for reticulate

antero-lateral region and a punctate band along posterior margin; 150 long and 162 wide at

widest point; with 22 pairs of setae (setae r1 and r4 absent); without scleronoduli and with

five pairs of distinguishable lyrifissures. Opisthonotal shield reticulate, except for a punctate

band along anterior margin; 139 long and 146 wide at widest point; with 15 pairs of setae and

four pairs of distinguishable lyrifissures. Unsclerotised integument laterad to opisthonotal

shield with five pairs of setae (R1-R5). Measurements of setae: j1 12, j2 14, j3 11, j4 10, j5 12,

j6 13, z1 12, z2 11, z3 11, z4 12, z5 13, z6 14, s1 9, s2 11, s3 12, s4 12, s5 13, s6 13, r2 12, r3

19, r5 12, r6 14, J1 16, J2 14, J3 15, J4 16, J5 10, Z1 16, Z2 17, Z3 17, Z4 16, Z5 14, S1 15,

S2 14, S3 15, S4 15, S5 13, R1 7, R2 7, R3 7, R4 8, R5 9.

Ventral idiosoma (Fig. 4.6B). Base of tritosternum 13 long and 10 wide proximally; laciniae

30 long, separated for about 90% of their total length, pilose. Sternogenital shield lightly

reticulate, with indistinct anterior margin, a small punctate region posterolaterad to seta st4

428

and a punctate band along straight posterior margin; region anterior to the first pair of

lyrifissures (iv1) lightly sclerotised and punctate; approximately 129 long, including the

lightly sclerotised and punctate region, and 69 wide at widest point; with five pairs of setae

and three pairs of lyrifissures; distance between st5-st5 22; genital opening on anterior margin

of shield. Ventrianal shield reticulate, except for a punctate band along anterior margin;

approximately 110 long and 126 wide at widest point, not fused to dorsal shield; with seven

pairs of setae (Jv1, Jv3, Jv5 and Zv1-Zv3) in addition to circum-anal setae and two pairs of

distinguishable lyrifissures; post-anal seta longer than para-anal setae. Peritreme extending

anteriorly to mid-level of coxa II. Peritrematal shield narrow, not extending beyond peritreme.

Measurements of setae: st1 11, st2 12, st3 11, st4 10, st5 10, Jv1 10, Jv2 not seen (broken),

Jv3 12, Jv5 11, Zv1 9, Zv2 10, Zv3 10, para-anal 15, post-anal 21.

Legs: Lengths: I: 247; II: 171; III: 168; IV: 225. Leg chaetotaxy as in adult female. With a

spur-like ventral seta on each of femur, genu and tibia II (Fig. 4.6C). Pretarsi as in adult

female.

Figure 4.6 - Pennarhodeus pennatus Karg. Male. A. Lateral (antiaxial) view of spermatodactyl; B. Ventral

idiosoma; C. Anterolateral view of femur, genu and tibia of leg II. Lyrifissures enlarged for

improved visibility

429

Remarks

This species was described on the basis of the adult female holotype, two adult female

paratypes and two adult male paratypes. The following characteristics appear in the original

description and illustrations of the species, but do not agree with our observations of the

female holotype: podonotal shield ornamented only with roundish markings, with 24 pairs of

setae and structures resembling scleronoduli (illustrated by Karg, 2000a, but not mentioned in

the text); seta z1 pilose; opisthonotal shield imbricate; unsclerotised integument laterad to

opisthonotal shield with three pairs of setae; genital shield longer than wide; peritrematal

shield fused with a section of exopodal shield next to coxa IV; lyrifissures present only on

sternal shield (three pairs). No information was provided about the tritosternum and leg setal


idiosoma (280-300), width of idiosoma (130-140), length of setae j1 (cited as i1) (12), j5

(cited as i4) (10), r3 (cited as r5) (18), J1 (cited as I1) (17), J3 (cited as I3) (16), J5 (cited as

I5) (14), Z5 (13), S5 (16), range of hypostomal setae (12-16), length of legs I (250), II (200),

III (160) and IV (250).

For the adult male paratype, the only characteristics provided in the original

description referred to details of the spermatodactyl and ventral setae of femur, genu and tibia

II, as well as measurements for length (250-280) and width (140) of the idiosoma.

Pennarhodeus turris Karg

Pennarhodeus turris Karg, 2000a: 257.

Pennarhodeus turris.— Karg, 2000b: 211.

Specimens examined: Holotype male: 6963 T, Kleine Antillen (indicated as Pinar del Rio,

Cuba in the original description), 1980 (indicated as 1976 in the original description), Boden

– Streu Pr. (litter sample) 19, leg. Mahunka, ZMB Kat. Nr. 45976. (in MNHB).

Diagnosis (adult male): Arthrodial process of chelicera with a short coronet-like fringe;

epistome with smooth margin, and with antero-medial projection about twice as long as

antero-lateral projections; podonotal shield mostly with roundish markings except for

reticulate antero-lateral region, without scleronoduli; with a narrow band projecting

posteriorly from region lateral to seta r3, bearing seta r4; dorsal idiosomal setae pilose, except

setae z1, z2, s1 and R1-R4, smooth; opisthonotal shield with polygons outlined by aligned

granules; seta R5 absent; sternogenital shield smooth; unsclerotised integument between

430

sternogenital and ventrianal shields without accessory platelet; peritreme extending anteriorly

to mid-level of coxa II; with a spur-like ventral seta on each of femur, genu and tibia II.

Adult male (Fig. 4.7 A-G) (holotype): Dorsal idiosomal setae (except z1, z2, s1 and R1-R4),

Jv5, para-anal and post-anal setae pilose; setae z1, z2, s1, R1-R4, hypostomal, subcapitular,

ventral idiosomal (except Jv5, para-anal and post-anal) and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 52 long, with three teeth in addition to apical tooth and a

setiform pilus dentilis (Fig. 4.7A); movable cheliceral digit 51 long, with one tooth in addition

to apical tooth. Spermatodactyl 110 long, abruptly bent dorsally, distal portion approximately

parallel to proximal portion, and almost twice as long. Arthrodial process of chelicera with a

short coronet-like fringe. Epistome with a smooth antero-medial projection, narrowed in

anterior one-third, with four apical denticles; flanked on each side by shorter, smooth

projection, each with five apical denticles (Fig. 4.7B); margin of epistome smooth between

antero-medial and antero-lateral projections and laterad to antero-lateral projections.

Deutosternal denticles in eight rows, each with 10-15 denticles; most anterior row shallow

―U‖ shaped, seventh row convex, other rows straight and transverse (Fig. 4.7C).

Measurements of setae: h1 18, h2 11, h3 15, sc 15.

Dorsal idiosoma (Fig. 4.7D): Podonotal shield mostly with roundish markings, except for

reticulate antero-lateral region, and a punctate band along its posterior margin, between setae

s6; 188 long and 222 wide at widest point; with 23 pairs of setae (seta r1 absent); without

scleronoduli and with four pairs of distinguishable lyrifissures; with a narrow band projecting

posteriorly from region lateral to seta r3, bearing seta r4. Opisthonotal shield with polygons

defined by aligned granules, except for a punctate band along anterior margin; 177 long and

154 wide at widest point; with 15 pairs of setae and four pairs of distinguishable lyrifissures.



z4 16, z5 20, z6 16, s1 12, s2 17, s3 17, s4 16, s5 18, s6 19, r2 18, r3 28, r4 12, r5 20, r6 20,

J1 21, J2 20, J3 18, J4 20, J5 13, Z1 21, Z2 21, Z3 20, Z4 20, Z5 25, S1 22, S2 21, S3 22, S4

22, S5 21, R1 10, R2 10, R3 11, R4 10.

431

Figure 4.7 - Pennarhodeus turris Karg. Male. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.

Hypostome; D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum; G. Anterolateral view of

femur, genu and tibia of leg II. Lyrifissures enlarged for improved visibility

432


4.7F); laciniae 63 long, separated for about 90% of their total length, pilose. Sternogenital

shield smooth and with indistinct anterior margin; region anterior to the first pair of

lyrifissures (iv1) lightly sclerotised and punctate; posterior margin straight; approximately 148

long, including the lightly sclerotised and punctate region, 74 wide at widest point; with five

pairs of setae and three pairs of lyrifissures; distance between st5-st5 34; genital opening on

anterior margin of shield. Ventrianal shield reticulate, except for a punctate band along

anterior margin, its anterior margin with a rounded protrusion, shield 125 long and 157 wide

at widest point, not fused to dorsal shield; with seven pairs of setae (Jv1, Jv3, Jv5 and Zv1-

Zv3) in addition to circum-anal setae and four pairs of distinguishable lyrifissures; post-anal

seta longer than para-anal setae. Peritreme extending anteriorly to mid-level of coxa II.


st2 15, st3 15, st4 13, st5 12, Jv1 16, Jv2 15, Jv3 16, Jv5 23, Zv1 14, Zv2 15, Zv3 15, para-anal

16, post-anal 22.

Legs: Lengths: I: 331; II: 267; III: 225; IV: 320. Genu IV with nine setae and tibia IV with ten

setae. With a spur-like ventral seta on each of femur, genu and tibia II (Fig. 7G). Pretarsus I, a

single, sessile, curved claw; pretarsi II-IV, an elongate ambulacral stalk, a pair of strong claws

and three rounded pulvillar lobes.

Remarks

This species is known only from the adult male holotype. The following


agree with our observations of the holotype: podonotal shield ornamented only with roundish

markings, with 24 pairs of setae and structures resembling scleronoduli (illustrated by Karg,

2000a but not mentioned in the text). No information was provided about the hypostome,

tritosternum, ventral idiosoma, lyrifissures and leg setal counts. The measurements provided

in the original description referred to the length of the idiosoma (350), width of idiosoma

(200), length of setae j1 (cited as i1) (18), j5 (cited as i4) (15), z6 (cited as i5) (16), J1 (cited

as I1) (20), J2 (cited as I2) (20), J4 (cited as I4) (20), Z5 (25), S5 (20), length of legs I (320),

II (250), III (220) and IV (300).

433

Key to Pennarhodeus species (known adult females)

1. Arthrodial process elongate, three-pointed; podonotal and opisthonotal shields

imbricate; genital shield lightly reticulate; unsclerotised integument between genital and

ventrianal shields with a transverse elongate accessory platelet; peritreme extending

anteriorly to level of anterior margin of coxa III ....... Pennarhodeus decoris Karg, 2000a

- Arthrodial process of chelicera with a short coronet-like fringe; podonotal shield mostly

with roundish markings, except for reticulate antero-lateral region, and opisthonotal

shield reticulate; genital shield smooth; unsclerotised integument between genital and

ventrianal shields without accessory platelet; peritreme extending anteriorly to mid-

level of coxa II ......................................................... Pennarhodeus pennatus Karg, 2000a

Key to Pennarhodeus species (known adult males)

1. Epistome with margin serrate laterad to antero-lateral projections; podonotal shield

imbricate; with two crescent-shaped scleronoduli between setae j5 and j6; unsclerotised

integument between sternogenital and ventrianal shields with a subtriangular accessory

platelet; with two spur-like ventral setae on each of femur, genu and tibia II ....................

……………………………………………….Pennarhodeus brevipennatus Karg, 2000b

- Epistome with margin smooth laterad to antero-lateral projections; podonotal shield

mostly with roundish markings, except for reticulate antero-lateral region; without

scleronoduli; unsclerotised integument between sternogenital and ventrianal shields

without accessory platelet; with one spur-like ventral seta on each of femur, genu and

tibia II ................................................................................................................................ 2

2. Epistome with antero-medial projection about 1.2 times as long as antero-lateral

projections; opisthonotal shield reticulate; seta R5 present, inserted on unsclerotised

integument; sternogenital shield lightly reticulate ... Pennarhodeus pennatus Karg, 2000a

- Epistome with antero-medial projection about twice as long as antero-lateral

projections; opisthonotal shield with reticules of aligned granules; seta R5 absent;

sternogenital shield smooth ........................................... Pennarhodeus turris Karg, 2000a

Genus Poropodalius Karg, 2000

Poropodalius Karg, 2000a: 252.

Poropodalius.— Karg and Schorlemmer, 2009: 66.

434

Type species: Poropodalius hexapennatus Karg, 2000, by original designation.

Diagnosis (adult female and male): Arthrodial process of chelicera with a short coronet-like

fringe; epistome with a smooth antero-medial projection, about uniform width or wider at the

base, flanked on each side by shorter projection, these either uniform width and smooth or

triangular and serrate; seta h3 directly posterior to h1 and mediad or slightly postero-mediad

to h2; dorsal idiosomal setae smooth or with distal half pilose; podonotal shield with 21-22

pairs of setae and four scleronoduli between setae j5 and j6 (except P. basisetae, where

scleronoduli are between setae j6 and z6); unsclerotised integument laterad to podonotal

shield with one or two pairs of setae; opisthonotal shield with 15 pairs of setae; with four or

five pairs of R setae on unsclerotised integument laterad to opisthonotal shield in female and

on ventrianal shield in male; presternal shields absent; ventrianal shield with five or six pairs

of setae in female and 12 pairs of setae in male; unsclerotised integument along anterior

margin of ventrianal shield with two pairs of setae in female and without setae in male; with

zero or one pair of metapodal plates in female and without metapodal plates in male;

peritreme extending anteriorly to level of median region of coxa II; pretarsus I either a single,

sessile, curved claw or an elongate ambulacral stalk, a pair of strong claws and three rounded

pulvillar lobes, with the same length or shorter to pretarsi of legs II-IV; seta pl4 of tarsus IV

absent. Numbers of setae on legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6,

6; genu: 13, 11, 9, 10; tibia: 14, 10, 8, 10; tarsus II-IV: 18, 18, 17. Adult male with

spermatodactyl abruptly bent dorsally and with a spur-like ventral seta on each of femur, genu

and tibia II.

Poropodalius acutus Karg, 2000

Poropodalius acutus Karg, 2000a: 253.

Poropodalius acutus.— Karg and Schorlemmer, 2009: 67.

Specimens examined: Holotype female: 6970 T, Kuba (= Cuba), 1977, Bodenprobe Nr. (soil

sample number) 1, ZMB Kat. Nr. 45925. Paratype male: 6971, Kuba (= Cuba), 1977,

Streudecke (litter sample), ZMB Kat. Nr. 45926. (in MNHB).

Diagnosis (adult female): Epistome with smooth antero-lateral projections of about uniform

width; dorsal idiosomal setae smooth; podonotal shield lightly imbricate, with four crescent-

shaped scleronoduli between setae j5 and j6; opisthonotal and sternal shields lightly

435

imbricate; unsclerotised integument between sternal and ventrianal shields without accessory

platelet; ventrianal shield with five pairs of setae (Jv2, Jv3, Jv5, Zv2 and Zv3) in addition to

circum-anal setae; peritrematal shield narrow, not extending beyond peritreme; pretarsi I-IV

similar in shape and length, an elongate ambulacral stalk, a pair of strong claws and three

rounded pulvillar lobes.

Adult female (Fig. 4.8 A-D) (holotype): Dorsal and ventral idiosomal, hypostomal,

subcapitular and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 30 long; movable digit 31 long; teeth and pilus dentilis

not visible because of position of chelicera. Arthrodial process of chelicera with a short

coronet-like fringe. Epistome with a smooth antero-medial projection of about uniform width,

with four apical denticles, flanked on each side by shorter, smooth projection, each with two

to four apical denticles (Fig. 4.8A); margin of epistome smooth between antero-medial and

antero-lateral projections and serrate laterad to lateral projections. Deutosternal denticles in

seven rows, each with eight to 12 denticles; fifth row anteriorly convex, other rows straight

and transverse (Fig. 4.8B). Seta h3 posterior to h1 and slightly postero-mediad to h2.

Measurements of setae: h1 13, h2 not seen (broken), h3 not seen (broken), sc 10.

Dorsal idiosoma (Fig. 4.8C): Podonotal shield lightly imbricate, except for a punctate band

along posterior margin; 137 long and 128 wide at widest point; with 21 pairs of setae (seta r1

absent); with four crescent-shaped scleronoduli between setae j5 and j6 and five pairs of

distinguishable lyrifissures. Unsclerotised integument laterad to podonotal shield with two

pairs of setae (r3 and r4). Opisthonotal shield lightly imbricate, except for a punctate band

along anterior margin; 132 long and 131 wide at widest point; with 15 pairs of setae and five

pairs of distinguishable lyrifissures. Unsclerotised integument laterad to opisthonotal shield

with five pairs of setae (R1-R5). Measurements of setae: j1 11, j2 10, j3 9, j4 8, j5 8, j6 not

seen (broken), z1 9, z2 7, z3 9, z4 8, z5 not seen (broken), z6 8, s1 6, s2 5, s3 8, s4 8, s5 9, s6

8, r2 7, r3 13, r4 7, r5 8, r6 10, J1 8, J2 8, J3 8, J4 10, J5 12, Z1 9, Z2 9, Z3 10, Z4 13, Z5 19,

S1 9, S2 8, S3 10, S4 14, S5 18, R1 6, R2 not seen (broken), R3 5, R4 5, R5 5.

Ventral idiosoma (Fig. 4.8D): Tritosternum not seen (broken). Sternal shield lightly

imbricate and with indistinct anterior margin; region anterior to the first pair of lyrifissures

436

Figure 4.8 - Poropodalius acutus Karg. Female. A. Epistome; B. Dorsal idiosoma; C. Hypostome; D. Ventral

idiosoma. Lyrifissures enlarged for improved visibility

437

(iv1) lightly sclerotised and punctate; posterior margin straight; approximately 85 long,

including the lightly sclerotised and punctate region, 79 wide at widest point; with four pairs

of setae and three pairs of lyrifissures. Genital shield lightly reticulate, except for a punctate

band along straight posterior margin; longer than wide; extending posteriorly behind coxae

IV; distance between st5-st5 42. Unsclerotised integument between genital and ventrianal

shields with setae Jv1 and Zv1. Metapodal plates not distinguishable. Ventrianal shield

reticulate; 88 long and 101 wide at widest point, not fused to dorsal shield; with five pairs of

setae (Jv2, Jv3, Jv5, Zv2 and Zv3) in addition to circum-anal setae and four pairs of

distinguishable lyrifissures. Peritreme extending anteriorly to mid-level of coxa II.


st2 not seen (broken), st3 not seen (broken), st4 not seen (broken), st5 not seen (broken), Jv1

10, Jv2 not seen (broken), Jv3 not seen (broken), Jv5 17, Zv1 8, Zv2 not seen (broken), Zv3

11, para-anal 14, post-anal not seen (broken).

Legs: Lengths: I: 248; II: 177; III: 153; IV not suitable for measurement. Pretarsi I-IV similar

in shape and length, an elongate ambulacral stalk, a pair of strong claws and three rounded

pulvillar lobes.

Remarks

This species was described on the basis of the adult female holotype and an adult male

paratype. The following characteristics appear in the original description and illustrations of

the species, but do not agree with our observations of the female holotype (the paratype male

was not re-examined in this study): unsclerotised integument laterad to opisthonotal shield

with two pairs of setae; genital shield smooth; a pair of rounded metapodal plates, containing

a circular structure; lyrifissures present only on podonotal shield (a pair) and sternal shield

(two pairs). No information was provided about the chelicera, hypostome, tritosternum and

leg setal counts. The only measurements provided for female in the original description

referred to the length of the idiosoma (260), width of idiosoma (140), range of dorsal

idiosomal setae (10-16), length of ventral idiosomal setae (10), length of legs I (250), II (170),

III (150) and IV (220).

Poropodalius basisetae Karg, 2000

Poropodalius basisetae Karg, 2000a: 255.

Poropodalius basisetae.— Karg and Schorlemmer, 2009: 67.

438

Specimens examined: Holotype female: 6966 T, Kleine Antillen (indicated as Cuba in the

original description), 1980 (indicated as 1976 in the original description), Streuschicht Pr.

(litter sample) 19, lg. Mahunka, ZMB Kat. Nr. 45927. (in MNHB).

Diagnosis of adult: Epistome with serrate, triangular antero-lateral projections; dorsal

idiosomal setae smooth; podonotal shield lightly imbricate, with four crescent-shaped

scleronoduli between setae j6 and z6; opisthonotal shield with anterior half lightly imbricate

and posterior half with indistinct roundish markings; sternal shield smooth; unsclerotised

integument between sternal and ventrianal shields with a transverse elongate accessory


circum-anal setae; peritrematal shield wider than peritreme, anterior and posterior ends

pointed; pretarsi I-IV similar in shape, an elongate ambulacral stalk, a pair of strong claws,

and three rounded pulvillar lobes; pretarsus I about half as long as other pretarsi.

Adult female (Fig. 4.9 A-F) (holotype): Dorsal and ventral idiosomal, hypostomal,


Gnathosoma: Fixed cheliceral digit 31 long, with five teeth in addition to apical tooth (pilus

dentilis indistinguishable) (Fig. 4.9A); movable cheliceral digit 32 long, with three teeth in


Epistome with a smooth antero-medial projection of about uniform width, with two apical

denticles, flanked on each side by shorter, triangular and serrate projection (Fig. 4.9B);

margin of epistome smooth between antero-medial and antero-lateral projections and serrate

laterad to antero-lateral projections. Deutosternal denticles in seven straight transverse rows,

each with eight to 12 denticles (Fig. 4.9C). Seta h3 directly posterior to h1 and mediad to h2.


Dorsal idiosoma (Fig. 4.9D): Podonotal shield lightly imbricate, except for a punctate band

along its posterior margin, between setae s6; 132 long and 129 wide at widest point; with 22

pairs of setae (seta r1 absent); with four crescent-shaped scleronoduli between setae j6 and z6

and five pairs of distinguishable lyrifissures. Unsclerotised integument laterad to podonotal

shield with a pair of setae (r4). Opisthonotal shield with anterior half lightly imbricate, and a

punctate band along anterior margin, and posterior half with indistinct roundish markings; 133

439

Figure 4.9 - Poropodalius basisetae Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.


improved visibility

440

long and 126 wide at widest point; with 15 pairs of setae and four pairs of distinguishable

lyrifissures. Unsclerotised integument laterad to opisthonotal shield with four pairs of setae

(R1-R4); seta R5 absent. Measurements of setae: j1 14, j2 18, j3 15, j4 16, j5 14, j6 14, z1 15,

z2 15, z3 16, z4 17, z5 15, z6 15, s1 12, s2 13, s3 15, s4 16, s5 15, s6 16, r2 9, r3 21, r4 9, r5

17, r6 21, J1 16, J2 18, J3 20, J4 24, J5 21, Z1 20, Z2 20, Z3 25, Z4 30, Z5 32, S1 20, S2 21,

S3 22, S4 27, S5 29, R1 8, R2 8, R3 10, R4 10.


4.9F); laciniae 28 long, separated for about 90% of their total length, pilose. Sternal shield


lightly sclerotised and punctate; posterior margin straight; approximately 92 long, including

the lightly sclerotised and punctate region, 58 wide at widest point; with four pairs of setae

and three pairs of lyrifissures. Genital shield lightly reticulate; shorter than wide; extending

posteriorly behind coxae IV; distance between st5-st5 35; posterior margin straight.

Unsclerotised integument between genital and ventrianal shields with a transverse elongate

accessory platelet and with setae Jv1 and Zv1; with a pair of elongate, curved metapodal

plates, each with a small median lobe on the convex paraxial margin. Ventrianal shield

reticulate; 84 long and 102 wide at widest point, not fused to dorsal shield; with five pairs of

setae (Jv2, Jv3, Jv5, Zv2 and Zv3) in addition to circum-anal setae and four pairs of

distinguishable lyrifissures; post-anal seta as long as para-anal setae. Peritreme extending

anteriorly to mid-level of coxa II. Peritrematal shield wider than peritreme, with pointed

anterior and posterior ends. Measurements of setae: st1 15, st2 16, st3 15, st4 13, st5 12, Jv1

11, Jv2 not seen (broken), Jv3 17, Jv5 26, Zv1 12, Zv2 16, Zv3 21, para-anal 17, post-anal 17.

Legs: Lengths: I: 231; II: 158; III: 156; IV: 210. Pretarsi I-IV similar in shape, an elongate

ambulacral stalk, a pair of strong claws, and three rounded pulvillar lobes; pretarsus I about

half as long as other pretarsi.

Remarks



agree with our observations of the holotype: podonotal shield reticulate, with 21 pairs of

setae, scleronoduli anterior to setae j6 (i5 in Karg's notation); unsclerotised integument laterad

to podonotal shield with a pair of setae; opisthonotal shield reticulate; genital shield smooth;

441

metapodal plates very large and with a median constriction on the paraxial margin;

lyrifissures present only on sternal shield (two pairs). No information was provided about the

chelicera, hypostome, tritosternum and leg setal counts. The measurements provided in the

original description referred to the length of the idiosoma (220-250), width of idiosoma (100-

110), length of setae j1 (cited as i1) (13), j3 (cited as i2) (16), j5 (cited as i4) (15), r3 (cited as

r5) (16), J1 (cited as I1) (16), J2 (cited as I2) (18), J3 (cited as I3) (18), J5 (cited as I5) (20),

Z3 (22), Z4 (25), Z5 (27), S1-S5 (20), Jv5 (cited as V8) (25), post-anal (15), length of other

ventral idiosomal setae (20) and length of metapodal plate (17).

Poropodalius crispus Karg, 2000

Poropodalius crispus Karg, 2000a: 253.

Poropodalius crispus.— Karg and Schorlemmer, 2009: 67.

Specimens examined: Holotype female: 6981 T, Kuba (= Cuba), 1975 (indicated as 1976 in

the original description), Bodenpr. (soil sample) 25-28 (indicated as litter sample number 48

in the original description), ZMB Kat. Nr. 45928. Paratypes: one male, 6982, Kuba (= Cuba),

1975, Bodenpr. (soil sample) 25-28, ZMB Kat. Nr. 45929; one male, 6980, Kuba (= Cuba),

1977, Bodenpr. (soil sample) 1, ZMB Kat. Nr. 45930. (in MNHB).

Diagnosis (adult female and male): Epistome with smooth antero-lateral projections, widest

at the base; setae j1, z2-z6, s5, r3, r6, J2-J5, Z1-Z5, S2-S5 pilose in distal half; other dorsal

idiosomal setae smooth; podonotal shield with four small circular scleronoduli between setae

j5 and j6; peritrematal shield broad and long, extending anteriorly onto the dorsum but not

fused to podonotal shield; pretarsus I, a single, sessile, curved claw. Adult female with

podonotal shield mostly with series of short, wavy lines mostly aligned transversely or

diagonally; opisthonotal shield imbricate with many imbrications containing series of short

longitudinal lines; sternal shield lightly reticulate; unsclerotised integument between genital

and ventrianal shields with three transverse elongate accessory platelets; ventrianal shield

with six pairs of setae (Jv2-Jv5, Zv2 and Zv3) in addition to circum-anal setae. Adult male

with podonotal shield imbricate; opisthonotal shield imbricate with imbrications without

series of short, longitudinal lines; sternogenital shield lightly reticulate; unsclerotised

integument between sternogenital and ventrianal shields with an ellipsoidal accessory platelet;

ventrianal shield with 12 pairs of setae (R1-R4, Jv1-Jv5 and Zv1-Zv3) in addition to circum-

anal setae; seta r4 on peritrematal shield.

442

Adult female (Fig. 4.10 A-E) (holotype): Setae j2-j6, z1, s1-s4, s6, r2, r4, r5, J1, S1, R1-R4,

hypostomal, subcapitular, ventral idiosomal (except Jv5) and leg setae smooth; setae j1, z2-z6,

s5, r3, r6, J2-J5, Z1-Z5, S2-S5 and Jv5 pilose in distal half.



short coronet-like fringe. Epistome with a smooth antero-medial projection, widest at the

base, with two apical denticles, flanked on each side by shorter, smooth projection, each with

three apical denticles (Fig. 4.10A); margin of epistome with two denticles between antero-

medial and antero-lateral projections and smooth laterad to antero-lateral projections.

Deutosternal denticles in eight rows, each with eight to 12 denticles; anterior row convex,

posterior row ―V‖ shaped, other rows straight and transverse (Fig. 4.10B). Seta h3 directly

posterior to h1 and slightly postero-mediad to h2. Measurements of setae: h1 24, h2 13, h3 not

seen (broken), sc 22.

Dorsal idiosoma (Fig. 4.10C): Podonotal shield with series of very short, wavy lines mostly

transversely or diagonally aligned, except for a punctate band along posterior margin; 224

long and 260 wide at widest point; with 22 pairs of setae (seta r1 absent); with four small

circular scleronoduli between setae j5 and j6 and four pairs of distinguishable lyrifissures.

Unsclerotised integument laterad to podonotal shield with a pair of short setae (r4).

Opisthonotal shield imbricate, except for a punctate band along its anterior margin, between

setae Z1; many imbrications containing series of short, longitudinal lines; shield 195 long and

258 wide at widest point; with 15 pairs of setae and five pairs of distinguishable lyrifissures.



z4 18, z5 20, z6 22, s1 14, s2 14, s3 15, s4 18, s5 20, s6 23, r2 20, r3 35, r4 10, r5 18, r6 27,

J1 19, J2 23, J3 22, J4 24, J5 17, Z1 23, Z2 23, Z3 30, Z4 32, Z5 35, S1 23, S2 23, S3 26, S4

30, S5 32, R1 10, R2 10, R3 13, R4 16.



lightly reticulate, with indistinct anterior margin; region anterior to the first pair of lyrifissures


443

Figure 4.10 - Poropodalius crispus Karg. Female. A. Epistome; B. Hypostome; C. Dorsal idiosoma; D. Ventral

idiosoma; E. Tritosternum. Lyrifissures enlarged for improved visibility

444

including the lightly sclerotised and punctate region, and 103 wide at widest point; with four

pairs of setae and three pairs of lyrifissures. Genital shield lightly reticulate; longer than wide;

extending posteriorly behind coxae IV; distance between st5-st5 62; posterior margin straight.

Unsclerotised integument between genital and ventrianal shields with three transverse

elongate accessory platelets, and with setae Jv1 and Zv1; with a pair of roundish metapodal

plates, each with a median constriction on the paraxial margin, and containing a circular

marking. Ventrianal shield reticulate; 124 long and 243 wide at widest point, not fused to

dorsal shield; with six pairs of setae (Jv2-Jv5, Zv2 and Zv3) in addition to circum-anal setae

and four pairs of distinguishable lyrifissures; post-anal seta as long as para-anal setae.

Peritreme extending anteriorly to mid-level of coxa II. Peritrematal shield smooth, broad and

long, extending anteriorly onto the dorsum but not fused to podonotal shield. Measurements

of setae: st1 16, st2 22, st3 not seen (broken), st4 16, st5 15, Jv1 not seen (broken), Jv2 19,

Jv3 25, Jv4 25, Jv5 30, Zv1 16, Zv2 18, Zv3 16, para-anal 23, post-anal 24.

Legs: Lengths: I: 354; II: 298; III: 279; IV: 357. Pretarsus I, a sessile, curved claw; pretarsi II-

IV, an elongate ambulacral stalk, a pair of strong claws and three rounded pulvillar lobes.

Adult male (Fig. 4.11 A-B) (two paratypes): Shape of setae as in adult female.

Gnathosoma: All structures damaged and unavailable for description.

Dorsal idiosoma: Podonotal shield imbricate, except for a punctate band along posterior

margin; 188 long and 195 wide at widest point; with 22 pairs of setae (seta r1 absent); with

four circular scleronoduli between setae j5 and j6 and four pairs of distinguishable

lyrifissures. Seta r4 on distal end of peritrematal shield. Opisthonotal shield imbricate, except

for a punctate band along its anterior margin, between setae Z1; imbrications without series of

short, longitudinal lines; 146-161 long and 171-189 wide at widest point; with 15 pairs of

setae and five pairs of distinguishable lyrifissures. Setae R1-R4 inserted on ventrianal shield;

seta R5 absent. Measurements of setae: j1 16, j2 not seen (broken), j3 16, j4 15-16, j5 15, j6

16, z1 13-14, z2 16, z3 not seen (broken), z4 16-17, z5 not seen (broken), z6 16-17, s1 not seen

(broken), s2 14, s3 16, s4 16, s5 18, s6 17, r2 14, r3 not seen (broken), r4 8-9, r5 15, r6 18, J1

18-19, J2 20, J3 19-20, J4 21, J5 14-15, Z1 21, Z2 20, Z3 23-24, Z4 26, Z5 28, S1 20, S2 20,

S3 22-23, S4 22, S5 24, R1 9-10, R2 11, R3 15, R4 14.

445

Ventral idiosoma (Fig. 4.11A): Tritosternum not seen (broken). Sternogenital shield lightly

reticulate and with indistinct anterior margin; region anterior to the first pair of lyrifissures


including the lightly sclerotised and punctate region, and 81 wide at widest point; with five

pairs of setae and three pairs of lyrifissures; distance between st5-st5 33-36; genital opening

on anterior margin of shield. Unsclerotised integument between sternogenital and ventrianal

shields with an ellipsoidal accessory platelet, punctate near anterior margin of ventrianal

shield. Ventrianal shield imbricate; 104-110 long and 192-201 wide at widest point, not fused

to dorsal shield; with 12 pairs of setae (R1-R4, Jv1-Jv5 and Zv1-Zv3) in addition to circum-

anal setae and four pairs of distinguishable lyrifissures; post-anal seta as long as para-anal

setae. Peritreme extending anteriorly to mid-level of coxa II. Peritrematal shield broad and

long, extending anteriorly onto the dorsum but not fused to podonotal shield. Measurements

of setae: st1 not seen (broken), st2 not seen (broken), st3 not seen (broken), st4 13, st5 12, Jv1

14, Jv2 15, Jv3 16-18, Jv4 16, Jv5 22, Zv1 not seen (broken), Zv2 15, Zv3 15-16, para-anal 16-

18, post-anal 20.

Figure 4.11 - Poropodalius crispus Karg. Male. A. Ventral idiosoma; B. Anterolateral view of femur, genu and

tibia of leg II. Lyrifissures enlarged for improved visibility

446

Legs: Lengths: I: 311; II: 220; III: 203-207; IV: 270. With a spur-like ventral seta on each of

femur, genu and tibia II (Fig. 4.11B). Pretarsi as in adult female.

Remarks

This species was described on the basis of the adult female holotype, one adult female

paratype and two adult male paratypes. The following characteristics appear in the original


female holotype: margin of epistome smooth between antero-medial and antero-lateral

projections; setae j1, j6, z4-z6, s4-s5, r3, r5-r6, J1-J5, Z2-Z5, S1-S5 and Jv5 totally pilose,

other setae smooth; genital shield smooth; peritrematal shield ornamented; lyrifissures present

only on podonotal shield (one pair) and sternal shield (three pairs). No information was

provided about the chelicera, hypostome, tritosternum and leg setal counts. The measurements

provided in the original description referred to the length of the idiosoma (320-400), width of

idiosoma (200-250), length of setae j1 (cited as i1) (20), j4 (cited as i3) (20), j5 (cited as i4)

(20), r3 (cited as r5) (35), J1 (cited as I1) (17), J3 (cited as I3) (25), J4 (cited as I4) (25), Z3

(25), Z4 (30), Z5 (30), S4 (27), S5 (32), Jv5 (cited as V8) (30) and length of other ventral

idiosomal setae (25).

For the adult male paratypes, the only characteristics provided referred to details of

the spermatodactyl and ventral setae of femur, genu and tibia II, as well as measurements for

length (380) and width (230) of the idiosoma.

Poropodalius hexapennatus Karg, 2000

Poropodalius hexapennatus Karg, 2000a: 252.

Poropodalius hexapennatus.— Karg and Schorlemmer, 2009: 67.

Specimens examined: Holotype female: 6976 T, Kuba (= Cuba), 1976, Bodenpr. (soil

sample) 48, ZMB Kat. Nr. 45932. Paratypes: one male, 6977, Kuba (= Cuba), 1976, Bodenpr.

(soils ample) 48, ZMB Kat. Nr. 45933; one male, 6978, Kuba (= Cuba), 1976, Bodenpr. (soil

sample) 20-24, ZMB Kat. Nr. 45934; one male, 6979, Kuba (= Cuba), 1977, Bodenpr. (soil

sample) 2, ZMB Kat. Nr. 45935. (in MHNB).

Diagnosis (adult female and male): Epistome with smooth antero-lateral projections of

about uniform width; setae r3, J4, J5, Z4, Z5, S5 pilose in distal half; other dorsal idiosomal

setae smooth; podonotal shield with indistinct roundish markings and four small circular

447

scleronoduli between setae j5 and j6; opisthonotal shield lightly reticulate; pretarsus I, a

single, sessile, curved claw. Adult female with sternal shield lightly imbricate; unsclerotised

integument between sternal and ventrianal shields with a transverse elongate accessory


circum-anal setae; peritrematal shield broad and long, extending anteriorly onto the dorsum

but not fused to podonotal shield. Adult male with sternogenital shield with indistinct

roundish markings; no accessory platelet between sternogenital and ventrianal shields;

ventrianal shield with 12 pairs of setae (R1-R5, Jv1-Jv3, Jv5 and Zv1-Zv3) in addition to

circum-anal setae; peritrematal shield narrow, not extending beyond peritreme.

Adult female (Fig. 4.12 A-E) (holotype): Dorsal idiosomal setae (except r3, J4, J5, Z4, Z5,

S5), hypostomal, subcapitular, ventral idiosomal (except post-anal) and leg setae smooth;

setae r3, J4-J5, Z4-Z5, S5 and post-anal setae pilose in distal half.

Gnathosoma: Fixed cheliceral digit 67 long, with five teeth in addition to apical tooth, and a

setiform pilus dentilis (Fig. 4.12A); movable cheliceral digit 66 long, with three teeth in


Epistome with a smooth antero-medial projection slightly wider at the base and provided with

two apical denticles, flanked on each side by a shorter, smooth projection, each provided with

three apical denticles (Fig. 4.12B); margin of epistome with two denticles between antero-

medial and antero-lateral projections and smooth laterad to antero-lateral projections.

Deutosternum, hypostomal and subcapitular setae not visible.

Dorsal idiosoma (Fig. 4.12C): Podonotal shield with indistinct roundish markings, except for

a punctate band along posterior margin; 201 long and 210 wide at widest point; with 22 pairs

of setae (seta r1 absent); with four small circular scleronoduli between setae j5 and j6 and five

pairs of distinguishable lyrifissures. Unsclerotised integument laterad to podonotal shield with

a pair of setae (r4). Opisthonotal shield lightly reticulate, except for a punctate band along

anterior margin; 167 long and 180 wide at widest point; with 15 pairs of setae and four pairs

of distinguishable lyrifissures. Unsclerotised integument laterad to opisthonotal shield with

five pairs of setae (R1-R5). Measurements of setae: j1 20, j2 20, j3 18, j4 17, j5 16, j6 19, z1

16, z2 18, z3 18, z4 20, z5 19, z6 20, s1 16, s2 18, s3 18, s4 19, s5 20, s6 21, r2 18, r3 25, r4

12, r5 20, r6 22, J1 20, J2 21, J3 22, J4 23, J5 21, Z1 21, Z2 22, Z3 25, Z4 30, Z5 33, S1 20,

S2 20, S3 22, S4 26, S5 29, R1 11, R2 11, R3 12, R4 13, R5 12.

448

Figure 4.12 - Poropodalius hexapennatus Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.

Dorsal idiosoma; D. Ventral idiosoma; E. Tritosternum. Lyrifissures enlarged for improved

visibility

449



lightly imbricate and with indistinct anterior margin; region anterior to the first pair of

lyrifissures (iv1) lightly sclerotised and punctate; posterior margin concave; approximately

103 long, including the lightly sclerotised and punctate region, and 96 wide at widest point;

with four pairs of setae and three pairs of lyrifissures. Genital shield lightly reticulate, except

for a punctate band along straight posterior margin; longer than wide; extending posteriorly

behind coxae IV; distance between st5-st5 50. Unsclerotised integument between genital and

ventrianal shields with a transverse elongate accessory platelet and with setae Jv1 and Zv1.

Metapodal plates irregular in shape, with a median constriction on the paraxial margin,

containing a circular marking. Ventrianal shield reticulate; 125 long and 152 wide at widest

point, not fused to dorsal shield; with five pairs of setae (Jv2, Jv3, Jv5, Zv2 and Zv3) in

addition to circum-anal setae and three pairs of distinguishable lyrifissures; post-anal seta as

long as para-anal setae. Peritreme extending anteriorly to mid-level of coxa II. Peritrematal

shield broad and long, extending anteriorly onto the dorsum but not fused to podonotal shield.

Measurements of setae: st1 17, st2 not seen (broken), st3 not seen (broken), st4 not seen

(broken), st5 17, Jv1 15, Jv2 not seen (broken), Jv3 20, Jv5 27, Zv1 14, Zv2 16, Zv3 15, para-

anal 20, post-anal 22.

Legs: Lengths: I: 332; II: 275; III: 226; IV: 310. Pretarsus I, a single, sessile, curved claw;

pretarsi II-IV, an elongate ambulacral stalk, a pair of strong claws and three rounded pulvillar

lobes.

Adult male (Fig. 4.13 A-C) (three paratypes): Shape of setae as in adult female.

Gnathosoma: Fixed cheliceral digit 42 long, with two teeth in addition to apical tooth and a

setiform pilus dentilis (Fig. 4.13A); movable cheliceral digit 41 long, with three teeth in

addition to apical tooth. Spermatodactyl 83 long, abruptly bent dorsally, distal portion

approximately parallel to proximal portion and double its length. Arthrodial process of

chelicera with a short coronet-like fringe. Epistome as in female. Deutosternum, hypostomal

and subcapitular setae not visible.

Dorsal idiosoma: Podonotal shield with indistinct roundish markings, except for a punctate

band along posterior margin; 176-192 long and 203-212 wide at widest point; with 22 pairs of

450

setae (seta r1 absent); with four circular scleronoduli between setae j5 and j6 and five pairs of


of setae (r4). Opisthonotal shield lightly reticulate, except for a punctate band along anterior

margin; 168-179 long and 179-187 wide at widest point; with 15 pairs of setae and four pairs

of distinguishable lyrifissures. Setae R1-R5 inserted on ventrianal shield. Measurements of

setae: j1 17, j2 17, j3 16-18, j4 15-17, j5 13-15, j6 not seen (broken), z1 14, z2 14-16, z3 15-

18, z4 16-18, z5 14-18, z6 16, s1 14, s2 16-18, s3 17, s4 17-20, s5 16-20, s6 20, r2 16, r3 21-

27, r4 11-13, r5 16-18, r6 20-22, J1 15-20, J2 18-20, J3 14-18, J4 16-21, J5 15-18, Z1 15-21,

Z2 15-21, Z3 18-22, Z4 20-26, Z5 23-28, S1 15-20, S2 17-20, S3 18-22, S4 20-24, S5 21-25,

R1 8-12, R2 9-12, R3 12-14, R4 10-14, R5 11-15.

Figure 4.13 - Poropodalius hexapennatus Karg. Male. A. Lateral (antiaxial) view of chelicera; B. Ventral

idiosoma; C. Anterolateral view of femur, genu and tibia of leg II. Lyrifissures enlarged for

improved visibility

Ventral idiosoma (Fig. 4.13B): Base of tritosternum 17 long and 11 wide proximally;

laciniae 51 long, separated for about 90% of their total length, pilose. Sternogenital shield

with indistinct roundish markings, indistinct anterior margin and a punctate band along

straight posterior margin; region anterior to the first pair of lyrifissures (iv1) lightly sclerotised

451

and punctate; approximately 136-154 long, including the lightly sclerotised and punctate

region, and 73-84 wide at widest point; with five pairs of setae and three pairs of lyrifissures;

distance between st5-st5 29-37; genital opening on anterior margin of shield. Unsclerotised

integument between sternogenital and ventrianal shields without accessory platelet.

Ventrianal shield reticulate, except for a punctate band along anterior margin; 135-155 long

and 201-230 wide at widest point, not fused to dorsal shield; with 12 pairs of setae (R1-R5,

Jv1-Jv3, Jv5 and Zv1-Zv3) in addition to circum-anal setae and three pairs of distinguishable

lyrifissures; post-anal seta as long as para-anal setae. Peritreme extending anteriorly to mid-

level of coxa II. Peritrematal shield narrow, not extending beyond peritreme. Measurements

of setae: st1 15-16, st2 14-16, st3 16-17, st4 13-15, st5 13-15, Jv1 15, Jv2 12-15, Jv3 13-16,

Jv5 20-23, Zv1 15, Zv2 11-13, Zv3 11-14, para-anal 15-20, post-anal 16-21.

Legs: Lengths: I: 311; II: 270; III: 210-235; IV: 265-305. With a spur-like ventral seta on

each of femur, genu and tibia II (Fig. 4.13C). Pretarsi as in adult female.

Remarks


paratypes and 15 adult male paratypes. The following characteristics appear in the original


holotype: epistome with antero-medial projection as long as antero-lateral projections;

podonotal shield ornamented with few irregular lines; unsclerotised integument laterad to

opisthonotal shield with only two pairs of setae; setae S5 and post-anal smooth and setae j1,

r3, J4, J5, Z4, Z5 and Jv5 totally pilose; unsclerotised integument between genital and

ventrianal shields with five accessory platelets; unsclerotised integument laterad to ventrianal

shield with four pairs of setae; peritrematal shield narrow, not extending beyond peritreme;

lyrifissures only on podonotal shield (one pair) and sternal shield (three pairs). No

information was provided about the hypostome, tritosternum and leg setal counts. The

measurements provided in the original description referred to the length of the idiosoma (310-

360), width of idiosoma (170-190), length of setae r3 (cited as r5) (28), J4 (cited as I4) (25),

Z4 (28), Z5 (30), range of other dorsal idiosomal setae (18-20), length of all st 17, Jv5 (cited

as V8) (25), range of other ventrianal idiosomal setae (16-20), length of legs I (290), II (250),

III (230) and IV (310).

452

For the male paratypes, the only characteristics provided referred to details of the

spermatodactyl and ventral setae of femur, genu and tibia II, as well as measurements for the

length (270-300) and width (150-190) of the idiosoma.

Poropodalius medioflagelli Karg and Schorlemmer

Poropodalius medioflagelli Karg and Schorlemmer, 2009: 66.

Specimens examined: Holotype female: 70161 T, Caracas, Venezuela, 1973, litter, soil,

ZMB Kat. Nr. 46165. (in MNHB).

Diagnosis (adult female): Epistome with smooth antero-lateral projections of about uniform

width; dorsal idiosomal setae smooth; podonotal shield with indistinct roundish markings and

four crescent-shaped scleronoduli between setae j5 and j6; opisthonotal shield lightly

imbricate; sternal shield with light reticulation along antero-lateral margin; no accessory

platelet between sternal and ventrianal shields; ventrianal shield with six pairs of setae (Jv2,

Jv3, Jv5, Zv2 and Zv3) in addition to circum-anal setae; peritrematal shield narrow, not

extending beyond peritreme; pretarsi I-IV similar in shape, an elongate ambulacral stalk, a

pair of strong claws and three rounded pulvillar lobes; pretarsus I about half as long as other

pretarsi.

Adult female (Fig. 4.14 A-C) (holotype): Dorsal and ventral idiosomal, hypostomal,




short coronet-like fringe. Epistome with a smooth antero-medial projection of about uniform

width, with two apical denticles, flanked on each side by shorter, smooth projection, each

with three apical denticles (Fig. 4.14A); margin of epistome smooth between antero-medial

and antero-lateral projections and serrate laterad to antero-lateral projections. Deutosternum,

hypostomal and subcapitular setae not visible.

Dorsal idiosoma (Fig. 4.14B): Podonotal shield with indistinct roundish markings, except for

a punctate band along posterior margin; 130 long and 119 wide at widest point; with 22 pairs

of setae (seta r1 absent); with four crescent-shaped scleronoduli between setae j5 and j6 and

453

five pairs of distinguishable lyrifissures. Unsclerotised integument laterad to podonotal shield

with a pair of setae (r4). Opisthonotal shield lightly imbricate, except for a punctate band

along anterior margin; 132 long and 125 wide at widest point; with 15 pairs of setae and four

pairs of distinguishable lyrifissures. Unsclerotised integument laterad to opisthonotal shield

with four pairs of setae (R1-R4); seta R5 absent. Measurements of setae: j1 17, j2 not seen

(broken), j3 20, j4 not seen (broken), j5 20, j6 not seen (broken), z1 16, z2 18, z3 21, z4 not

seen (broken), z5 18, z6 23, s1 12, s2 not seen (broken), s3 22, s4 22, s5 21, s6 23, r2 21, r3

29, r4 12, r5 21, r6 24, J1 25, J2 not seen (broken), J3 not seen (broken), J4 25, J5 22, Z1 25,

Z2 not seen (broken), Z3 24, Z4 25, Z5 27, S1 25, S2 24, S3 25, S4 25, S5 25, R1 10, R2 10,

R3 14, R4 16.

Figure 4.14 - Poropodalius medioflagelli Karg. Female. A. Epistome; B. Dorsal idiosoma; C. Ventral idiosoma.

Lyrifissures enlarged for improved visibility

454

Ventral idiosoma (Fig. 4.14C): Tritosternum not seen (broken). Sternal shield with light

reticulation along antero-lateral margin and with indistinct anterior margin; region anterior to

the first pair of lyrifissures (iv1) lightly sclerotised and punctate; posterior margin slightly

concave; approximately 77 long, including the lightly sclerotised and punctate region, and 61

wide at widest point; with four pairs of setae and three pairs of lyrifissures. Genital shield

with scanty reticulation and a punctate band along straight posterior margin; longer than wide;

extending posteriorly behind coxae IV; distance between st5-st5 38. Unsclerotised integument

between genital and ventrianal shields with setae Jv1 and Zv1. Metapodal plates kidney-

shaped, with narrow posterior end, containing a circular marking. Ventrianal shield reticulate;

87 long and 101 wide at widest point, not fused to dorsal shield; with five pairs of setae (Jv2,

Jv3, Jv5, Zv2 and Zv3) in addition to circum-anal setae and three pairs of distinguishable

lyrifissures. Peritreme extending anteriorly to mid-level of coxa II. Peritrematal shield narrow,

not extending beyond peritreme. Measurements of setae: st1 not seen (broken), st2 not seen

(broken), st3 not seen (broken), st4 not seen (broken), st5 not seen (broken), Jv1 not seen

(broken), Jv2 not seen (broken), Jv3 not seen (broken), Jv5 24, Zv1 not seen (broken), Zv2 not

seen (broken), Zv3 not seen (broken), para-anal not seen (broken), post-anal not seen

(broken).

Legs: Lengths: I: 197; II: 152; III: 140; IV: 181. Pretarsi I-IV similar in shape, an elongate

ambulacral stalk, a pair of strong claws and three rounded pulvillar lobes; pretarsus I about

half as long as other pretarsi.

Remarks



agree with our observations of the holotype: epistome with serrate anterior margin, a long

flagellum-like antero-medial projection and without antero-lateral projections; podonotal

shield reticulate; unsclerotised integument laterad to podonotal shield with two pairs of setae;

opisthonotal shield ornamented with transverse lines; genital shield smooth; ventrianal shield

with only a few weak transverse lines; lyrifissures present only on sternal shield (two pairs).

No information was provided about the chelicera, hypostome, tritosternum and leg setal


idiosoma (249), width of idiosoma (140), length of setae j1 (cited as i1) (17), j2 (cited as z1)

(20), j3 (cited as i2) (19), j5 (cited as i4) (20), z1 (cited as s1) (15), z2 (17), z5 (cited as z4)

455

(17), s2 (cited as r2) (10), r2 (cited as r3) (20), r6 (cited as r5) (25), J1 (cited as I1) (22), J4

(cited as I4) (25), J5 (cited as I5) (22), Z1-Z5 (24), S1 (23), S5 (25), length of all st (17), para-

anal (10), Jv5 (cited as lateral caudal setae) (22), length of other ventrianal idiosomal setae

(17), length of the ventrianal shield (81), width of ventrianal shield (102), length of legs I

(203), II (182), III (133) and IV (193).

Key to Poropodalius species (known adult females)

1. Dorsal idiosoma with at least one seta (r3) with distal half pilose; podonotal shield with

four circular scleronoduli; peritrematal shield broad and long, extending anteriorly onto

the dorsum but not fused to podonotal shield; pretarsus I, a single, sessile, curved claw ..

…………………………………………………………………………………………...2

- Dorsal idiosoma with all setae smooth; podonotal shield with four crescent-shaped

scleronoduli; peritrematal shield narrow, not extending beyond peritreme; pretarsus I, an

elongate ambulacral stalk, a pair of strong claws and three rounded pulvillar lobes ........ 3

2. Podonotal shield with series of short, wavy lines mostly transversely or diagonally

aligned; setae j1, z2-z5, s5, s6, r3 and r6 with distal half pilose, other podonotal setae

smooth; opisthonotal shield imbricate; sternal shield lightly reticulate; unsclerotised

integument between sternal and ventrianal shields with three transverse elongate

accessory platelets; ventrianal shield with six pairs of setae (Jv2-Jv5, Zv2 and Zv3) in

addition to circum-anal setae ....................................... Poropodalius crispus Karg, 2000a

- Podonotal shield with indistinct roundish markings; only seta r3 with distal half pilose,

other podonotal setae smooth; opisthonotal shield lightly reticulate; sternal shield lightly

imbricate; unsclerotised integument between sternal and ventrianal shields with one

transverse elongate accessory platelet; ventrianal shield with five pairs of setae (Jv2,

Jv3, Jv5, Zv2 and Zv3) in addition to circum-anal setae…………………………………..

............................................................................Poropodalius hexapennatus Karg, 2000a

3. Epistome with serrate antero-lateral projections triangular; scleronoduli between setae

j6 and z6; sternal shield smooth; unsclerotised integument between genital and

ventrianal shields with one transverse elongate accessory platelet……………………….

................................................................................... Poropodalius basisetae Karg, 2000a

- Epistome with smooth antero-lateral projections of about uniform width; scleronoduli

between setae j5 and j6; sternal shield ornamented; unsclerotised integument between

genital and ventrianal shields without accessory platelet ................................................. 4

456

4. Podonotal shield lightly imbricate; setae j3 and J1 not reaching the base of setae j4 and

J2, respectively; pretarsus I with the same size of other pretarsus………………………..

....................................................................................... Poropodalius acutus Karg, 2000a

- Podonotal shield with indistinct roundish markings; setae j3 and J1 reaching the base of

setae j4 and J2, respectively; pretarsus I with half the size of other pretarsi………………

…………………………………Poropodalius medioflagelli Karg and Schorlemmer, 2009

Key to Poropodalius species (known adult males)

Information for P. acutus is based on Karg (2000a), as the type specimens were not

available for study.

1. Spermatodactyl more than five times longer than movable cheliceral digit .......................

....................................................................................... Poropodalius acutus Karg, 2000a

- Spermatodactyl 2-3 times longer than movable cheliceral digit ....................................... 2

2. Podonotal shield imbricate; setae j1, z2-z5, s5, s6, r3 and r6 with distal half pilose, other

podonotal setae smooth; opisthonotal shield imbricate; sternogenital shield lightly

reticulate; unsclerotised integument between sternogenital and ventrianal shields with

an ellipsoidal accessory platelet; ventrianal shield imbricate; with four pairs of setae R

on ventrianal shield; peritrematal shield broad and long, extending anteriorly onto the

dorsum but not fused to podonotal shield .................... Poropodalius crispus Karg, 2000a

- Podonotal shield with indistinct roundish markings; only seta r3 with distal half pilose,

other podonotal setae smooth; opisthonotal shield lightly reticulate; sternogenital shield

with light roundish markings; unsclerotised integument between sternogenital and

ventrianal shields without accessory platelet; ventrianal shield reticulate; with five pairs

of setae R on ventrianal shield; peritrematal shield narrow, not extending beyond

peritreme ............................................................ Poropodalius hexapennatus Karg, 2000a

4.4 Discussion

Re-examination of these species has shown that the genera Interrhodeus,

Pennarhodeus and Poropodalius are correctly placed in the family Rhodacaridae, on the basis

of the following character states: sternal shield with four pairs of setae and with a punctate

band along is anterior margin (in some Rhodacaridae st1 inserted on pre-sternal shields or on

lightly sclerotised anterior region of sternal shield); podonotal shield with a punctate band

457

along its posterior margin, with 14-23 pairs of setae; opisthonotal shield with a punctate band

along its anterior margin, with 14-20 pairs of setae; ventral and anal shields of female fused to

form a ventrianal shield with 1-8 pairs of pre-anal setae in addition to circumanal setae; palp

tarsal claw three-tined; genu IV with 10 (rarely 9) setae, tibia IV with 10 (rarely 9) setae;

tarsus IV without pl4; and male with seta st5 inserted on sternogenital shield. Some of the

species in Interrhodeus and Pennarhodeus are unusual for Rhodacaridae in lacking

scleronoduli in the posterior part of the podonotal shield (the absence of scleronoduli in

rhodacarids may be more common than what has been reported). However, their other

characteristics are typical of the rhodacarids, and they will thus be included in the world-wide

catalogue of the family (CASTILHO; MORAES; HALLIDAY, 2012). Interrhodeus is most

similar to Afrodacarellus Hurlbutt, but the latter has species with the cheliceral arthrodial

process shaped like a long cylindrical brush, seta s1 present, podonotal shield with four

scleronoduli and tritosternum with laciniae separated for about 90% of their total length.

Pennrhodeus is also most similar to Afrodacarellus Hurlbutt, but the latter has species with

smooth dorsal setae [pilose in Afrodacarellus bipilosus (Karg)] and the cheliceral arthrodial

process as well as scleronoduli as previously mentioned. Poropodalius is most similar to

Rhodacarellus, but the latter has species with a smooth podonotal shield (ornamented in

Rhodacarellus unicus Karg), smooth idiosomal dorsal setae and a single pair of elongate

metapodal plates (rarely two, with the outer one small and rounded).

References









475 p.

458




243–262, 2000a.



72, p. 207–213, 2000b.






219, 1970.

LEE, D.C. Rhodacaridae (Acari: Mesostigmata) from near Adelaide, Australia. III. Behaviour

and development. Acarologia, Paris, v. 16, p. 21-44, 1974.







Press, 2009. p. 124-232.

WALTER D.E.; HUNT, H.W.; ELLIOTT E.T. Guilds or functional groups? An analysis of

predatory arthropods from a shortgrass steppe soil. Pedobiologia, Jena, v. 31, p. 247-260,

1988.

459

5 REVISION OF THE GENUS Protogamasellopsis (ACARI: MESOSTIGMATA:

RHODACARIDAE)

Abstract

Protogamasellopsis Evans and Purvis is revised based on the examination of the types

or specimens collected in Brazil of the seven species of this genus. A characterization of the

genus, diagnoses of the species examined, complementary descriptions of some species and a

key to help in the separation of the species of this genus are provided.

Keywords: Rhodacaroidea; Soil mites; Taxonomy

Resumo

Protogamasellopsis Evans and Purvis é revisado com base no exame dos tipos ou

espécimes coletados no Brasil das sete espécies deste gênero. Uma caracterização do gênero,

diagnoses das espécies examinadas, descrições complementares de algumas espécies e uma

chave para ajudar a separação das espécies deste gênero são apresentadas.

Palavras-chave: Rhodacaroidea; Ácaros de solo; Taxonomia

5.1 Introduction

Protogamasellopsis Evans and Purvis are free living edaphic mites found mainly in

the top soil layers. They are predators and at least one species appears to have potential as

biological control agent for soil insect and mite pests (CASTILHO et al., 2009).

Evans and Purvis (1987) described this genus in the family Ascidae Oudemans, where

it was initially included as a member of the Protogamasellus-group together with

Protogamasellus (Protogamasellus) Karg and Protogamasellus (Protogamasellodes) Evans

and Purvis, then described. This classification was based on the form and location of the

opening of the spermatheca (base of coxae III), number of setae of tibia I (13 setae), the

unarmed nature of male legs II and the spermatodactyl with recurved tip. They considered that

the resemblances between species of the Protogamasellopsis and the Rhodacaridae (the

460

colour, body form and sclerotization) was probably due to their common adaptations to life in

confined spaces and not necessarily because of close taxonomic relationship.

Karg (1994a) transferred Protogamasellopsis to Rhodacaridae, without presenting

arguments to support that decision, what was only done by Karg (1994b). According to the

latter publication, it should be included in that family because of the following characteristics:

setae j1, j2 and z1 inserted along anterior margin of podonotal shield; podonotal shield with a

punctate band along posterior margin; metasternal shield fused with sternal shield; post-anal

seta long; cheliceral digits long and epistome with long antero-medial extension.

Halliday; Walter and Lindquist (1998) presented further details about this genus. They

claimed that some character states used by Evans and Purvis (1987) to put

Protogamasellopsis in Ascidae were weak (some Rhodacaroidea have 13 setae on tibia I and

opening of the spermatheca located at the base of leg III; presence or absence of spine-like

setal armature on male leg II varies greatly in the Ascidae) and cited other character states to

support the inclusion of Protogamasellopsis in Rhodacaridae (palp tarsal claw three-tined;

coxa I with a dorsal spine; podonotal shield with desclerotised band of punctated cuticle

posterior to j6; seta st1 inserted on lightly sclerotised and punctate cuticle; female with st4

inserted on sternal shield; female with genital shield with a desclerotised band of punctate

cuticle along posterior margin; male with remnant of a separate platelet between the

sternogenital and ventrianal shields; male with Jv1 inserted on soft cuticle rather than on

ventrianal shield. In addition, Halliday; Walter and Lindquist (1998) cited that

Protogamasellopsis resembles the Rhodacaroidea in the shape of its epistome, the general

form of its gnathosoma, the position and relative lengths of its circum-anal setae, the

relatively slight extension of the genital shield‘s hyaline anterior margin anteriorly, and its

large, heavily sclerotised chelicerae. Although they agreed with Karg (1994a) and Karg

(1994b) in maintaining Protogamasellopsis in the Rhodacaridae, they considered

Protogamasellus to be an Ascidae.

Karg (2007) revised the Protogamasellus-group of Evans and Purvis (1987) and

included the genera Protogamasellopsis, Protogamasellus, Protogamasellodes and a new

genus Protofurcatus Karg in the group. But, differently from Evans and Purvis (1987), this

group was placed in Rhodacaridae.

Protogamasellopsis consists of seven described species (CASTILHO; MORAES;

HALLIDAY, 2012), wich have been reported mainly from South America (EVANS;

PURVIS, 1987; KARG, 1994a; KARG, 1994b; KARG, 2000; CASTILHO et al., 2009;

CASTILHO; MORAES, 2010). The descriptions of some of these species are not sufficiently

461

detailed, making it difficult to decide to which group they really belong. Thus, a re-

examination of the types of those species was considered necessary to allow the conclusion of

a project to elaborate a catalog of the rhodacarid species (CASTILHO; MORAES;

HALLIDAY, 2012), and to allow the correct identification of mites of those groups in the

South American continent. The objectives of this paper were to provide a characterization of

the genus, diagnoses of the species examined, complementary descriptions of some species

and a key to help in the separation of the species of this genus.


The type specimens of species examined were borrowed from the British Museum

(Natural History), London, England and the Arachnologische Sammlung des Museums für

Naturkunde der Humboldt-Universität, Berlin, Germany. They were examined under a phase

contrast microscope provided with a camera lucida, in the University of Amsterdam, The

Netherlands. The specimens were illustrated and measurements were taken of structures

considered taxonomically important. In the following redescriptions, setal nomenclature is

based on Lindquist and Evans (1965). Measurements of each structure are given in

micrometres (µm) and as a range (or a single value when measurement did not vary)

representing the variation among the specimens examined. Leg length was measured from the

proximal edge of the coxa to the tip of the pretarsus.

Despite our efforts, we were unable to examine the types of Protogamasellopsis

dioscorus (Manson) and Protogamasellopsis posnaniensis Wiśniewski and Hirschmann.

Thus, measurements given for those species were taken from specimens collected in Brazil.

For those species as well as for Protogamasellopsis corticalis Evans and Purvis, 1987, new

morphological information refers only to measurements, because the corresponding original

descriptions were considered sufficiently detailed to provide other informations.

5.3 Results

Genus Protogamasellopsis Evans and Purvis, 1987

Protogamasellopsis Evans and Purvis, 1987: 855 (described in Ascidae Voigts and

Oudemans).

Protogamasellopsis.— Karg, 1994a: 123; Karg, 1994b: 207; Halliday; Walter and Lindquist,

1998: 2; Karg, 2007: 123.

462

Type species: Protogamasellopsis corticalis Evans and Purvis, 1987, by original

designation.

Rhodacarella Moraza, 2004: 2 (described in Rhodacaridae Oudemans) [synonym, according

to M.L. Moraza, personal communication, 2007].

Type species: Rhodacarella cavernicola Moraza, 2004, by original designation.

Diagnosis (adult female): Movable and fixed cheliceral digits with two and 6-8 teeth,

respectively. Arthrodial process of chelicera in the form of a short coronet-like fringe.

Epistome with anterior region acuminate. Seta h3 directly posterior to h1 and slightly anterior

and mediad to h2. Idiosoma elongate. Podonotal shield smooth, with or without punctate band

along posterior margin, with 16 pairs of setae, three pairs of which (j1, j2 and z1) along

anterior margin and without distinct scleronoduli. Unsclerotised integument laterad to

podonotal shield with 5-6 pairs of setae and an elongate plate running from the level of z1 to

the level of z4, bearing or not seta r3. Opisthonotal shield smooth, with or without punctate

band along anterior margin and with 15 pairs of setae. Unsclerotised integument laterad to

opisthonotal shield with 6-10 pairs of setae. Peritreme extending anteriorly to mid-level of

coxa II. With 0-4 transverse groups of presternal plates, each group consisting of 1-2 platelets.

Sternal shield with anterior margin indistinct and posterior margin concave. Sclerotized

region of genital shield longer than length of its posterior margin; the latter straight.

Unsclerotised integument between genital and ventrianal shields with two pairs of setae and

with or without lightly sclerotised plates. Ventrianal shield longer than wide, smooth, with 1-2

pairs of preanal setae; anterior margin convex. Unsclerotised integument laterad to ventrianal

shield with 3-4 pairs of setae. Opisthogastric region with 0-7 pairs of elongate, round or

subquadrate plates. Numbers of setae on segments of legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6,

5, 5, 5; femur: 12, 11, 6, 6; genu: 13, 11, 8, 9; tibia: 13, 10, 8, 9; tarsus: not counted, 18, 18,

17 (pl4 present). Pretarsi I-IV similar in shape and length, with elongate ambulacral stalk, a

pair of strongly sclerotized claws and three round pulvillar lobes.

Protogamasellopsis corticalis Evans and Purvis, 1987

Protogamasellopsis corticalis Evans and Purvis, 1987: 856.

Protogamasellopsis corticalis.— Karg, 1994a: 124; Karg, 1994b: 208; Shaw, 1999: 45; Karg,

2007: 124.

463

Specimens examined: Adult female holotype, two adult female paratypes and one male

paratype collected in public gardens, Jamestown, Saint Helena [South Atlantic island], under

dead bark of a Citrus sp. plant [Rutaceae] (possibly imported from Cape Province, South

Africa). Types deposited at British Museum (Natural History), London, England.

Diagnosis (adult female): Margin of epistome finely denticulate except for a spine-like

anterocentral extension, smooth; setae j3 and j4 not reaching the base of j4 and j5,

respectively; unsclerotised integument laterad to opisthonotal shield with 10 pairs of setae

(R1-R5 and UR1-UR5); without presternal plates; unsclerotized integument between genital

and ventrianal shields with three small accessory plates; opisthogastric region with one pair of

elongate and three pairs of round plates laterad to ventrianal shield.

Adult female (Adult female holotype and two adult female paratypes).

Gnathosoma: Fixed cheliceral digit 80 (77-83) long; movable digit 77 (74-81) long.

Measurements of setae: h1 35 (35-36), h2 17 (16-17), h3 25 (24-25), sc 22 (21-23).

Dorsal idiosoma: Podonotal shield 225 (220-231) long and 136 (127-153) wide at widest

level. Opisthonotal shield 236-262 long and 86-92 wide at widest level. Measurements of

setae: j1 39 (38-41), j2 25 (23-26), j3 26 (25-27), j4 27 (27-28), j5 26 (25-27), j6 23 (22-24),

z1 22 (21-22), z2 25 (25-26), z3 26 (26-27), z4 26 (25-26), z5 26 (25-26), z6 29 (28-30), s2 24

(23-25), s3 25 (24-26), s4 27 (26-28), s5 27 (26-27), s6 27 (26-27), r2 29 (29-30), r3 35 (35-

36), r4 23 (22-23), r5 25 (24-26), r6 31 (31-32), J1 24 (23-25), J2 23 (22-25), J3 22 (21-23),

J4 25 (22-27), J5 12 (11-13), Z1 26 (26-27), Z2 24 (23-25), Z3 27 (25-28), Z4 32 (30-33), Z5

51 (50-52), S1 29 (28-30), S2 26 (25-28), S3 26 (25-26), S4 27 (25-30), S5 25 (25-26), R1 22

(21-24), R2 22 (21-23), R3 22 (21-23), R4 22 (21-23), R5 42 (41-43), UR1 21 (20-22), UR2

20 (19-21), UR3 24 (23-25), UR4 19 (19-20), UR5 24 (23-25).

Ventral idiosoma: Base of tritosternum 39 (37-42) long and 16 (15-16) wide proximally;

laciniae 106 (98-115) long, separated for about 70% of their total length. Sternal shield

approximately 137 (132-142) long, including the lightly sclerotized and punctate region, and

128 (127-129) wide at widest level. Distance between st5-st5 56 (55-57). Ventrianal shield

169 (167-171) long and 87 (86-88) wide at widest level. Measurements of setae: st1 31 (30-

32), st2 32 (32-33), st3 33 (33-34), st4 30 (29-31), st5 20 (20-21), Jv1 21 (20-22), Jv2 24 (24-

464

25), Jv3 26 (26-27), Jv4 32 (31-33), Zv1 23 (23-24), Zv2 16 (15-17), Zv3 23 (23-24), para-

anal 35 (33-36), post-anal 58 (56-61).

Legs: Lengths: I: 392 (379-412); II: 297 (281-313); III: 251 (244-257); IV: 381 (372-396).

Male (one paratype).

Gnathosoma: Fixed cheliceral digit 51 long; movable digit 49 long. Spermatodactyl 53 long.


Dorsal idiosoma: Podonotal shield 176 long and 111 wide at widest level. Opisthonotal

shield 176 long and 78 wide at widest level. Measurements of setae: j1 32, j2 23, j3 22, j4 24,

j5 23, j6 20, z1 17, z2 23, z3 22, z4 21, z5 21, z6 23, s2 20, s3 20, s4 23, s5 22, s6 23, r2 24, r3

30, r4 16, r5 17, r6 28, J1 20, J2 19, J3 20, J4 21, J5 10, Z1 22, Z2 21, Z3 24, Z4 27, Z5 41,

S1 25, S2 22, S3 23, S4 25, S5 23, R1 15, R2 16, R3 16, R4 17, R5 32.

Ventral idiosoma: Base of tritosternum 31 long and 15 wide proximally; laciniae 82 long,

otherwise as in adult female. Sternogenital shield approximately 183 long and 107 wide at

widest level. Distance between st5-st5 38. Ventrianal shield 122 long along midline and 133

wide at widest level. Measurements of setae: st1 26, st2 26, st3 30, st4 25, st5 18, Jv1 18, Jv2

24, Jv3 28, Jv4 32, Zv1 20, Zv2 16, Zv3 22, para-anal 28, post-anal 47.

Legs: Lengths: I: 319; II: 238; III: 197; IV: 303.

Remarks

This species was described on the basis of the adult female holotype, nine adult female

paratypes and three male paratypes. The original description of this species is very detailed,

but the only measurements provided for the adult females were: length of the idiosoma 545-

625, length of the podonotal shield 250-290, width of the podonotal shield at the level of seta

s4 168-203, length of the opisthonotal shield 285-325, width of the opisthonotal shield 105-

115, length of seta J5 9.5-12, length of seta Z5 57-71, length of the ventrianal shield 190-215

and width of the ventrianal shield 99-110. The only measurement provided for the adult males

was the length of the idiosoma 430-450.

465

Protogamasellopsis dioscorus (Manson, 1972)

Protogamasellus dioscorus Manson, 1972: 437.

Protogamasellopsis dioscorus.— Evans and Purvis, 1987: 855; Karg, 1994a: 123; Karg,

1994b: 208; Karg, 2007: 124; Castilho and Moraes, 2010: 397.

Specimens examined: Three females collected in Pirassununga, São Paulo State, Brazil, 1

November 2000, from soil under Psidium guajava [Myrtaceae]. Specimens deposited at

Departamento de Entomologia e Acarologia, Escola Superior de Agricultura ―Luiz de

Queiroz‖ (ESALQ), Universidade de São Paulo (USP), Piracicaba, São Paulo State, Brazil.

Diagnosis (adult female): Margin of epistome finely denticulate except for a spine-like



(R1-R5, UR1-UR5); without presternal plates; unsclerotized integument between genital and

ventrianal shields without accessory plates; opisthogastric region with one pair of elongate

and three pairs of round plates laterad to ventrianal shield plates.

Adult female

Gnathosoma: Fixed cheliceral digit 43 (37-48) long; movable digit 42 (36-47) long.



level. Opisthonotal shield 226 (224-228) long and 81 (80-82) wide at widest level.

Measurements of setae: j1 37 (36-37), j2 26 (25-27), j3 27 (26-27), j4 26 (25-26), j5 22 (21-

22), j6 21 (20-21), z1 20 (20-19), z2 35 (34-35), z3 26 (25-26), z4 20, z5 26, z6 34 (33-35), s2

17 (16-17), s3 22 (22-23), s4 25 (24-26), s5 39 (38-40), s6 39 (38-39), r2 34 (33-34), r3 43

(42-43), r4 26, r5 33 (32-33), r6 38 (37-39), J1 24, J2 24 (23-24), J3 22 (21-22), J4 26 (25-

26), J5 13 (12-14), Z1 22 (21-23), Z2 25 (24-25), Z3 29 (29-30), Z4 36 (35-36), Z5 49 (48-

49), S1 38 (36-38), S2 32, S3 32 (31-33), S4 37 (36-37), S5 32 (31-33), R1 33 (32-33), R2 29

(28-29), R3 30, R4 33 (32-34), R5 37 (36-37), UR1 30 (28-31), UR2 30 (29-30), UR3 29 (27-

30), UR4 39, UR5 43 (42-43).

466




98 (94-101) wide at widest level. Distance between st5-st5 49 (48-49). Ventrianal shield 155

(152-157) long and 61 (59-62) wide at widest level. Measurements of setae: st1 23 (22-24),

st2 31 (30-31), st3 28 (27-29), st4 24 (23-24), st5 19 (18-19), Jv1 19 (18-19), Jv2 24 (23-24),

Jv3 30, Jv4 30 (30-31), Zv1 23 (22-23), Zv2 21, Zv3 35 (34-36), para-anal 29 (27-31), post-

anal 53 (51-56).


Remarks.


paratypes, 10 male paratypes, seven deutonymph paratypes, four protonymph paratypes and

two larva paratypes collected from New Zealand (Auckland International Airport), 19 June

1969, intercepted on yam [Dioscoreaceae] imported from Tonga. All types are deposited at

Collection of the Department of Agriculture, Levin, New Zealand.

The original description of this species is very detailed, but the only measurementes

provided were: adult females – length of the idiosoma 372-485, width of the idiosoma 158-

224; adult males – length of the idiosoma 348-393, width of the idiosoma 164-189;

deutonymphs – length of the idiosoma 342-367, width of the idiosoma 160-180; protonymph

– length of the idiosoma 296-332, width of the idiosoma 152-183; larva – length of the

idiosoma 250-263 and width of the idiosoma 181-197.

Protogamasellopsis granulosus Karg, 1994

Protogamasellopsis granulosus Karg, 1994b: 208.

Protogamasellopsis granulosus.— Karg, 2007: 124.

Specimen examined: Adult female holotype collected in Cueva Bella Vista, Bella Vista,

Santa Cruz, Galapagos Islands, 15 May 1985, in trap with manure. Type deposited at

Arachnologische Sammlung des Museums für Naturkunde, Berlin, Germany.

Diagnosis (adult female): Margin of epistome denticulate except for a spine-like


467

respectively [according Karg (1994b)]; unsclerotised integument laterad to opisthonotal shield

with six pairs of setae (R1-R5 and UR5) [according Karg (1994b)]; without presternal plates;

unsclerotized integument between genital and ventrianal shields without accessory plates

[according Karg (1994b)]; opisthogastric region with three pairs of round plates laterad to

ventrianal shield [according Karg (1994b)].

Adult female (Fig. 5.1 A-D) (holotype).

Gnathosoma: Fixed cheliceral digit 69 long, with six teeth in addition to apical tooth and a

setiform pilus dentilis (Fig. 5.1A); movable cheliceral digit 67 long, with two teeth in addition

to apical tooth. Margin of epistome denticulate, except for a spine-like anterocentral

extension, smooth (Fig. 5.1B). Deutosternal denticles in eight straight transverse rows, with 7-

12 denticles each (Fig. 5.1C). Measurements of setae: h1 32, h2 15, h3 24, sc broken.

Figure 5.1 - Protogamasellopsis granulosus Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome;

C. Hypostome; D. Tritosternum

Dorsal idiosoma: All structures damaged and inadequate for redescription.

Ventral idiosoma: Base of tritosternum 21 long and 14 wide proximally (Fig. 5.1D); laciniae

92, separated for about 70% of their total length, pilose. Without presternal plates. Sternal

shield smooth and with anterior margin indistinct; region anterior to the first pair of

468

lyrifissures (iv1) lightly sclerotized and punctate; posterior margin concave; approximately

103 long, including the lightly sclerotized and punctate region, and 95 wide at widest level;

with four pairs of setae and three pairs of lyrifissures. Peritreme extending anteriorly to mid-

level of coxa II. All other structures of ventral idiosoma damaged and inadequate for

redescription. Measurements of setae: st1 26, st2 26, st3 broken, st4 25.

Legs: Lengths: I broken; II: broken; III: 195; IV: 295.

Remarks

This species was described only on the basis of the adult female holotype, which is

very damaged. The only structures possible to be observed in detail are the gnathosoma,

sternal shield, peritreme and legs III and IV.

The measurements provided for adult female in the original description were length of

the idiosoma 420, width of idiosoma 170, length of setae j1 (cited as i1) 30, j2 (cited as s1)

20, j3 (cited as i2) 17, j4 (cited as i3) 22, j5 (cited as i4) 22, j6 (cited as i5) 20, z1 (cited as r1)

17, z6 (cited as z3) 25, r2 (cited as r4) 25, J1 (cited as I1) 20, Z4 30, Z5 42, S1 25, range of

other dorsal setae 20-25, Jv3 (cited as V3) 30, Jv4 (cited as V8) 45, post-anal 45, range of

other ventral setae 20-25, length of legs I 340, II 250, III 210 and IV 310.

Protogamasellopsis leptosomae Karg, 1994

Protogamasellopsis leptosomae Karg, 1994b: 210.

Protogamasellopsis leptosomae.— Karg, 2007: 123.

Specimen examined: Adult female holotype collected in Puntudo, Santa Cruz, Galapagos

Islands, 10 March 1985, in litter of fern [Pteridophyta] and wood chips. Type deposited at

Arachnologische Sammlung des Museums für Naturkunde, Berlin, Germany.

Diagnosis (adult female): Margin of epistome smooth; setae j3 and j4 not reaching the base

of j4 and j5, respectively; unsclerotised integument laterad to opisthonotal shield with eight

pairs of setae (R1-R5 and UR3-UR5); with four transverse groups of presternal plates, each

consisting of 1-2 platelets; unsclerotized integument between genital and ventrianal shields

without accessory plates; opisthogastric region without plates laterad to ventrianal shield.

469

Adult female (Fig. 5.2 A-F) (holotype): Dorsal and ventral idiosomal, hypostomal, subcoxal

and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 95 long, with 7-8 teeth in addition to apical tooth and a

setiform pilus dentilis (Fig. 5.2A); movable cheliceral digit 91 long, with two teeth in addition

to apical tooth. Margin of epistome smooth (Fig. 5.2B). Deutosternal denticles in nine

straight and transverse rows, each with 9-13 denticles (Fig. 5.2C). Measurements of setae: h1

52, h2 22, h3 broken, sc 29.

Dorsal idiosoma (Fig. 5.2D): Podonotal shield smooth, except for a punctate band along

posterior margin; 209 long and 151 wide at widest level; with 16 pairs of setae (setae s1 and

r1 absent) and with five pairs of distinguishable lyrifissures. Unsclerotised integument laterad

to podonotal shield with five pairs of setae (s2-s4, r2 and r4) and an elongate plate running

from the level of z1 to the level of z4, bearing seta r3. Opisthonotal shield smooth, except for

a punctate band along anterior margin; 229 long and 81 wide at widest level; with 15 pairs of

setae and five pairs of distinguishable lyrifissures. Unsclerotised integument laterad to

opisthonotal shield with eight pairs of setae (R1-R5 and UR3-UR5). Measurements of setae: j1

broken, j2 broken, j3 broken, j4 broken, j5 27, j6 27, z1 broken, z2 33, z3 29, z4 30, z5 30, z6

38, s2 32, s3 36, s4 35, s5 34, s6 33, r2 31, r3 42, r4 21, r5 34, r6 39, J1 27, J2 28, J3 26, J4

26, J5 16, Z1 27, Z2 28, Z3 35, Z4 42, Z5 59, S1 37, S2 26, S3 30, S4 45, S5 41, R1 27, R2 26,

R3 25, R4 20, R5 50, UR3 25, UR4 18, UR5 24.


5.2F); laciniae 115 long, separated for about 70% of their total length, pilose. With four

transverse groups of presternal plates, each group consisting of 1-2 platelets. Sternal shield


lightly sclerotized and punctate; posterior margin concave; approximately 99 long, including

the lightly sclerotized punctate region, 117 wide at widest level; with four pairs of setae and

three pairs of lyrifissures. Genital shield smooth, except for a punctate band along straight

posterior margin; sclerotized region slightly longer than wide; extending posteriorly behind

coxae IV; distance between st5-st5 52. Unsclerotised integument between genital and

ventrianal shields with setae Jv1 and Zv1. Ventrianal shield smooth; 174 long and 46 wide at

widest level, not fused to dorsal shield; with one pair of setae (Jv2) in addition to circum-anal

setae and one pair of distinguishable lyrifissures; post-anal seta longer than para-anal setae.

470

Figure 5.2 - Protogamasellopsis leptosomae Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome;

C. Hypostome; D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures enlarged for

improved visibility

471

Unsclerotised integument laterad to ventrianal shield with four pairs of setae (Jv3, Jv4, Zv2

and Zv3) and without plates. Peritreme extending anteriorly to mid-level of coxa II.


st2 33, st3 33, st4 31, st5 23, Jv1 26, Jv2 28, Jv3 30, Jv4 32, Zv1 25, Zv2 25, Zv3 20, para-anal

35, post-anal 71.


Remarks

This species was described on the basis of the adult female holotype. The following


agree with our observations of the female holotype: epistome denticulate; opisthonotal shield

without a punctate band along anterior margin and with 13 pairs of setae; unsclerotised

integument laterad to opisthonotal shield with six pairs of setae; with three pairs of elongate

presternal plates (plates of each side not subdivided); unsclerotised integument laterad to

ventrianal shield with seven pairs of setae; peritreme extending anteriorly to mid-level of coxa

III. No information was provided about the hypostome, tritosternum, lyrifissures and leg setal

counts. The measurements provided in the original description were: length of the idiosoma

400, width of idiosoma 140, range of length of dorsal setae 25-40, length of setae j1 (cited as

i1) 38, j5 (cited as i4) 25, r3 (cited as r5) 43, J1 (cited as I1) 25, J2 (cited as I2) 25, J3 (cited

as I3) 33, J4 (cited as I4) 25, J5 (cited as I5) 13, Z4 40, Z5 63, distance between J4 and Z5 55,

range of ventral setae 20-25, para-anal (cited as V4) 34, R5 (cited as V8) 22, length of legs I

430, II 310, III 250 and IV 380.

Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, 1991

Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, 1991: 189.

Protogamasellopsis posnaniensis.— Karg, 1993: 369; Karg, 1994b: 208; Karg, 2007: 123;

Castilho et al., 2009: 165.

Rhodacarella cavernicola Moraza, 2004: 4 (junior synonymy, following M.L. Moraza,

personal communication, 2007).

Specimens examined: Three females collected in Piracicaba, São Paulo State, Brazil, 20

October 2011, from a colony established at Departamento de Entomologia e Acarologia,

Escola Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de São Paulo

472

(USP), Piracicaba, São Paulo State, Brazil. Specimens deposited at Departamento de

Entomologia e Acarologia, ESALQ/USP.




(R1-R5 and UR1-UR5); with three transverse groups of presternal plates, each consisting of 1-

2 platelets; unsclerotized integument between genital and ventrianal shields with four

transversely elongate accessory plates; opisthogastric region with one pair of elongate and

three pairs of round plates laterad to ventrianal shield.

Adult female

Gnathosoma: Fixed cheliceral digit 91 (90-92) long, movable digit 84 (81-86) long.



level. Opisthonotal shield 252 (243-259) long and 98 (96-100) wide at widest level.


26), j6 23 (22-25), z1 22 (21-23), z2 28 (26-30), z3 28 (27-30), z4 27 (26-28), z5 28 (26-30),

z6 34 (32-35), s2 27 (25-29), s3 25 (24-26), s4 32 (31-33), s5 30 (28-31), s6 38 (36-40), r2 33

(31-35), r3 46 (43-49), r4 24 (23-25), r5 31 (30-33), r6 31 (29-32), J1 26 (25-28), J2 24 (21-

26), J3 25 (23-26), J4 25 (25-27), J5 15 (14-15), Z1 28 (27-30), Z2 27 (25-29), Z3 32 (30-35),

Z4 38 (35-41), Z5 57 (24-29), S1 34 (33-36), S2 32 (29-34), S3 30 (28-31), S4 31 (28-34) , S5

31 (29-33), R1 26 (24-27), R2 26 (24-27), R3 26 (24-29), R4 26 (24-29), R5 44 (41-46), UR1

20 (20-21), UR3 21 (20-21), UR3 28 (26-30), UR4 20 (20-21), UR5 32 (31-32).




118 (114-121) wide at widest level. Distance between st5-st5 51 (49-52). Ventrianal shield

169 (166-173) long and 69 (64-73) wide at widest level. Measurements of setae: st1 31 (30-

31), st2 31 (30-32), st3 31 (30-31), st4 29 (27-31), st5 23 (21-24), Jv1 25 (23-26), Jv2 29 (27-

473

31), Jv3 30 (28-32), Jv4 22 (21-24), Zv1 22 (21-23), Zv2 24 (22-25), Zv3 15 (15-16), para-

anal 36 (33-38), post-anal 70 (66-73).


Remarks

This species was described on the basis of the adult female holotype, an unspecified

number of adult female, deutonymph and protonymph paratypes collected from Poznań,

Poland, July to August 1989, in litter of Phoenix sp. [Aracaceae] in a greenhouse. Holotype is

deposited at Department of Forest Protection, University of Life Sciences, Poznań, Poland.

The original description of this species is very detailed, but the only measurements

provided were the length and width of the idiosoma of adult female (respectively 485-545 and

190-260), deutonymph (415-460 and 185-225) and protonymph (265-335 and 140-150).

Protogamasellopsis praeendopodalis Karg, 1994

Protogamasellopsis praeendopodalis Karg, 1994a: 123.

Protogamasellopsis praeendopodalis.— Karg, 1994b: 208; Karg, 2007: 123.

Specimens examined: Adult female holotype and one adult male paratype collected in

Fernandina, Galapagos Islands, 18 March 1985, in litter of grass and sand in coastal area.

Type deposited at Arachnologische Sammlung des Museums für Naturkunde, Berlin,

Germany.




(R1-R5 and UR1-UR5); with four pairs of transverse groups of presternal plates, each group

consisting of 1-2 platelets; unsclerotized integument between genital and ventrianal shields

without accessory plates; opisthogastric region with three pairs of round plates laterad to

ventrianal shield.

Adult female (Fig. 5.3 A-D) (holotype): Dorsal and ventral idiosomal, hypostomal, subcoxal

and leg setae smooth.

474

Gnathosoma: Chelicera broken. Margin of epistome denticulate except for a spine-like

anterocentral extension, smooth (Fig. 5.3A). Deutosternal denticles in eight rows, each with 9-

13 denticles; second row inverted V-shaped, other rows straight (Fig. 5.3B). Measurements of

setae: h1 39, h2 18, h3 27, sc 23.

Dorsal idiosoma (Fig. 5.3C): Podonotal shield smooth, except for a punctate band along

posterior margin; 219 long and 141 wide at widest level; with 16 pairs of setae (s1 and r1

absent) and with six pairs of distinguishable lyrifissures. Unsclerotised integument laterad to

podonotal shield with six pairs of setae (s2-s4 and r2-r4) and with an elongate plate running

from the level of z1 to the level of s4. Opisthonotal shield smooth; 217 long and 88 wide at

widest level; with 15 pairs of setae and eight pairs of distinguishable lyrifissures.

Unsclerotised integument laterad to opisthonotal shield with 10 pairs of setae (R1-R5 and

UR1-UR5). Measurements of setae: j1 36, j2 28, j3 29, j4 28, j5 30, j6 28, z1 23, z2 30, z3 31,

z4 31, z5 31, z6 36, s2 26, s3 24, s4 31, s5 26, s6 31, r2 32, r3 41, r4 22, r5 29, r6 36, J1 27,

J2 27, J3 26, J4 28, J5 17, Z1 28, Z2 31, Z3 35, Z4 40, Z5 55, S1 32, S2 34, S3 33, S4 35, S5

33, R1 26, R2 26, R3 27, R4 30, R5 44, UR1 21, UR2 21, UR3 29, UR4 20, UR5 26.

Ventral idiosoma (Fig. 5.3D): Tritosternum broken off. With four pairs of transverse groups

of presternal plates, each group consisting of 1-2 platelets. Sternal shield smooth and with

indistinct anterior margin; region anterior to the first pair of lyrifissures (iv1) lightly

sclerotized and punctate; posterior margin concave; approximately 97 long, including the

lightly sclerotized and punctate region, and 102 wide at widest level; with four pairs of setae

and three pairs of lyrifissures. Genital shield smooth, except for a punctate band along straight

posterior margin; with a pair of distinguishable lyrifissures posterior to st5; sclerotized region

longer than wide; extending posteriorly behind coxae IV; distance between st5-st5 56.

Unsclerotised integument between genital and ventrianal shields with setae Jv1 and Zv1.

Ventrianal shield smooth; 126 long and 71 wide at widest level, not fused to dorsal shield;

with one pair of setae (Jv2) in addition to circum-anal setae and without distinguishable

lyrifissures or pores; post-anal seta longer than para-anal setae. Unsclerotised integument

laterad to ventrianal shield with four pairs of setae (Jv3, Jv4, Zv2 and Zv3) and with three

pairs of round plates. Peritreme extending anteriorly to mid-level of coxa II. Peritrematal

shield indistinct. Measurements of setae: st1 31, st2 32, st3 32, st4 31, st5 27, Jv1 27, Jv2 27,

Jv3 33, Jv4 32, Zv1 26, Zv2 17, Zv3 26, para-anal 36, post-anal 64.

475

Figure 5.3 - Protogamasellopsis praeendopodalis Karg. Female. A. Epistome; B. Hypostome; C. Dorsal

idiosoma; D. Ventral idiosoma. Lyrifissures enlarged for improved visibility

476


Adult male (Fig. 5.4 A-C) (one paratype): Shape of setae as in adult female, except for a

spur-like ventral seta on tarsus IV (Fig. 5.4A).


setiform pilus dentilis (Fig. 5.4B); movable cheliceral digit 64 long, with one tooth in addition

to apical tooth. Spermatodactyl 72 long, at slight angle with axis of movable cheliceral digit

and abruptly bent distally. Epistome and deutosternum as in adult female. Measurements of

setae: h1, h2 broken, h3 broken, sc 21.

Dorsal idiosoma: Dorsal idiosoma similar to that of adult female, except for usclerotised

integument laterad to opisthonotal shield with five pairs of setae (R1-R5). Podonotal shield

204 long and 134 wide at widest level. Opisthonotal shield 209 long and 88 wide at widest

level. Measurements of setae: j1 broken, j2 29, j3 broken, j4 broken, j5 29, j6 28, z1 25, z2

and z3 broken, z4 30, z5 broken, z6 37, s2 25, s3 broken, s4 33, s5 broken, s6 34, r2 , r3 and

r4 broken, r5 27, r6 39, J1 26, J2 26, J3 27, J4 28, J5 16, Z1 30, Z2 33, Z3 38, Z4 39, Z5 63,

S1 35, S2 34, S3 36, S4 38, S5 35, R1 25, R2 26, R3 26, R4 28, R5 43.

Ventral idiosoma (Fig. 5.4C): Tritosternum broken. With four pairs of transverse groups of

presternal plates, each group consisting of 1-2 plates. Sternogenital shield smooth, with

indistinct anterior margin and a punctate band along straight posterior margin; region anterior

to the first pair of lyrifissures (iv1) lightly sclerotized and punctate; approximately 181 long,

including the lightly sclerotized and punctate region, and 106 wide at widest level; with five

pairs of setae and four pairs of lyrifissures; distance between st5-st5 43; genital opening on

anterior margin of shield. Ventrianal shield smooth; 150 long and 151 wide at widest level,

not fused to dorsal shield; with seven pairs of setae (Jv1-Jv4 and Zv1-Zv3) in addition to

circum-anal setae and two pairs of distinguishable lyrifissures; post-anal seta longer than

para-anal setae. Peritreme extending anteriorly to mid-level of coxa II. Peritrematal shield

narrow, not extending beyond peritreme. Measurements of setae: st1 31, st2 36, st3 35, st4 32,

st5 25, Jv1 25, Jv2 31, Jv3 37, Jv4 33, Zv1 24, Zv2 26, Zv3 37, para-anal 37, post-anal 72.

Legs: Lengths: I: 415; II: 291; III: 255; IV: 403. Leg setal counts and pretarsi as in adult

female.

477

Figure 5.4 - Protogamasellopsis praeendopodalis Karg. Male. A. Anterolateral view of femur, genu and tibia of

leg II; B. Lateral (antiaxial) view of chelicerae, with detail of spermatodactyl; C. Ventral idiosoma.

Lyrifissures enlarged for improved visibility

Remarks

This species was described on the basis of the adult female holotype, one adult female

paratype and one adult male paratype. The following characteristics appear in the original


female holotype: unsclerotised integument laterad to podonotal shield with five pairs of setae;

unsclerotised integument laterad to opisthonotal shield with nine pairs of setae; with four pairs

of elongate presternal plates (plates of each side not subdivided); unsclerotised integument

laterad to ventrianal shield with six pairs of setae; lyrifissures present only on podonotal

shield (a pair of lyrifissures), opisthonotal shield (three pairs) and sternal shield (two pairs).

No information was provided about the leg setal counts. The measurements provided for adult

females were: length of the idiosoma 390-410, width of idiosoma 160-180, range of podonotal

setae 30-32, length of setae j1 (cited as i1) 35, Z3 38, Z4 25, Z5 50, other opisthonotal setae

35, st1-st4 32, st5 25, Jv3 and para-anal (cited as ventrianal setae) 33, post-anal 60, length of

ventrianal shield 140, width of ventrianal shield 60, length of peritrematal shield 80, length of

478

legs I 400, II 300, III 250 and IV 400. For the adult male, the only characteristics provided in

the original description referred to details of the spermatodactyl, sternogenital shield and a

spur-like ventral setae of tarsus IV (length 17), as well as measurements for length of the

idiosoma 400, width of the idiosoma 170 and length of legs I 400, II 300, III 250 and IV 400.

Protogamasellopsis transversus Karg, 2000

Protogamasellopsis transversus Karg, 2000: 252.

Protogamasellopsis transversus.— Karg, 2007: 123.

Specimens examined: Adult female holotype and one female paratype collected in Cotopaxi

Province, Ecuador, 1973, in paramo [= páramo, Spanish term for a Neotropical habitat of high

altitude]. Type deposited at Arachnologische Sammlung des Museums für Naturkunde,

Berlin, Germany.


anterocentral extension, smooth; setae j3 and j4 reaching the base of j4 and j5, respectively;

unsclerotised integument laterad to opisthonotal shield with 10 pairs of setae (R1-R5 and

UR1-UR5); without presternal plates; unsclerotized integument between genital and ventrianal

shields without accessory plates; opisthogastric region with three pairs of round plates

posterior to coxa IV, and two pairs of round and two pairs of subquadrate plates laterad to

ventrianal shield.

Adult female (Fig. 5.5 A-F) (holotype and one paratype): Dorsal and ventral idiosomal,

hypostomal, subcoxal and leg setae smooth.

Gnathosoma: Fixed cheliceral digit 89-91 long, with seven teeth in addition to apical tooth

and a setiform pilus dentilis (Fig. 5.5A); movable cheliceral digit 87-89 long, with two teeth

in addition to apical tooth. Margin of epistome denticulate except for spine-like anterocentral

extension, smooth (Fig. 5.5B). Deutosternal denticles in eight rows, each with 9-13 denticles

(Fig. 5C); most anterior row anteriorly inverted V-shaped, other rows straight. Measurements

of setae: h1 37-38, h2 19-20, h3 34-36, sc 26-27.

479

Figure 5.5 - Protogamasellopsis transversus Karg. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome;

C. Hypostome; D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures enlarged for

improved visibility

480

Dorsal idiosoma (Fig. 5.5D): Podonotal shield smooth, except for a punctate band along

posterior margin; 217-234 long and 176-192 wide at widest level; with 16 pairs of setae (setae

s1 and r1 absent) and with six pairs of distinguishable lyrifissures. Unsclerotised integument

laterad to podonotal shield with five pairs of setae (s2-s4, r2 and r4) and with an elongate

plate running from the level of z1 to the level of z4, bearing seta r3. Opisthonotal shield

smooth, except for a punctate band along anterior margin; 221-248 long and 102-125 wide at

widest level; with 15 pairs of setae and eight pairs of distinguishable lyrifissures.

Unsclerotised integument laterad to opisthonotal shield with 10 pairs of setae (R1-R5 and

UR1-UR5). Measurements of setae: j1 43, j2 35-36, j3 36, j4 35, j5 36, j6 30, z1 26-27, z2 45-

46, z3 37, z4 32, z5 32, z6 46, s2 34-35, s3 32-33, s4 47-49, s5 40, s6 43, r2 44-46, r3 52-53,

r4 30-32, r5 39-40, r6 48-50, J1 33, J2 33, J3 28, J4 31, J5 14-16, Z1 36, Z2 38, Z3 45, Z4 52,

Z5 66, S1 42, S2 42, S3 44, S4 48, S5 44-46, R1 33-35, R2 32, R3 32, R4 35, R5 54-55, UR1

28-30, UR2 28-30, UR3 40-41, UR4 26-27, UR5 33-35.

Ventral idiosoma (Fig. 5.5E): Base of tritosternum 25-29 long and 16-17 wide proximally

(Fig. 5.5F); laciniae 93-95 long, separated for about 70% of their total length, pilose. Without

presternal plates. Sternal shield smooth and with indistinct anterior margin; region anterior to

the first pair of lyrifissures (iv1) lightly sclerotized and punctate; posterior margin concave;

approximately 125-131 long, including the lightly sclerotized and punctate region, and 129-

136 wide at widest level; with four pairs of setae and three pairs of lyrifissures. Genital shield

smooth; sclerotized region slightly longer than wide; extending posteriorly behind coxae IV;

distance between st5-st5 60-61. Unsclerotised integument between genital and ventrianal

shields with setae Jv1 and Zv1. Ventrianal shield smooth; 158-160 long and 79-87 wide at

widest level, not fused to dorsal shield; with one pair of setae (Jv2) in addition to circum-anal

setae; post-anal seta longer than para-anal setae. Unsclerotised integument laterad to

ventrianal shield with four pairs of setae (Jv2, Jv4, Zv2 and Zv3) and with two pairs of round

and two pairs of subquadrate plates. Opisthogastric region with three pairs of small, round

plates posterior to coxa IV. Peritreme extending anteriorly to mid-level of coxa II.

Peritrematal shield extending posteriorly to posterior margin of coxa IV. Measurements of

setae: st1 broken, st2 broken, st3 39, st4 36-37, st5 28, Jv1 31-32, Jv2 38, Jv3 37-40, Jv4 37-

40, Zv1 26, Zv2 33-34, Zv3 22-25, para-anal 47-48, post-anal 75.

Legs: Lengths: I: 410; II: 302; III: 258-290; IV: 425-431.

481

Remarks

This species was described on the basis of the adult female holotype and two adult

female paratypes. The following characteristics appear in the original description and

illustrations of the species, but do not agree with our observations of the adult female

holotype and an adult female paratype: podonotal shield with 19 pairs of setae; unsclerotised

integument laterad to podonotal shield with five pairs of setae; with three pairs of round

metapodal plates; opisthogastric region without plates posterior to coxa IV, and with three

pairs of round plates laterad to ventrianal shield; lyrifissures present only on podonotal shield

(a pair of lyrifissures) and sternal shield (three pairs). No information was provided about the

chelicera, hypostome, tritosternum and leg setal counts. The measurements provided in the

original description were: length of the idiosoma 420-450, width of idiosoma 150-180, length

of setae j1 (cited as i1) 40, j4 (cited as i3) 35, j5 (cited as i4) 35, r3 40, r5 50, J1-J2 (cited as

I1-I2) 25, J4 (cited as I4) 25, J5 (cited as I5) 10, Z3-Z4 25, Z5 60, R5 55, Jv3 (cited as V3) 40,

para-anal (cited as V4) 40, post-anal 70, length of legs I 420, II 310, III 280 and IV 400.

Key to Protogamasellopsis species (known adult females)

1. With presternal plates ........................................................................................................ 2

- Without presternal plates................................................................................................... 4

2. Margin of espistome totally smooth; with three pairs of UR setae (UR3-UR5)..................

. .................................................................... Protogamasellopsis leptosomae Karg, 1994a

- Margin of epistome denticulate except for a spine-like anterocentral extension, smooth;

with five pairs of UR setae (UR1-UR5) ............................................................................ 3

3. Unsclerotized integument between genital and ventrianal shields with four transversely

elongate accessory plates; opisthogastric region with one pair of elongate and three pairs

of round plates laterad to ventrianal shield………………………………………………..

. ............................. Protogamasellopsis posnaniensis Wiśniewski and Hirschmann, 1991

- Unsclerotized integument between genital and ventrianal shields without plates;

opisthogastric region with three pairs of round plates laterad to ventrianal shield .............

……………………………………….Protogamasellopsis praeendopodalis Karg, 1994a

4. Setae j3 and j4 about as long as distance between their bases and bases of j4 and j5,

respectively ................................................... Protogamasellopsis transversus Karg, 2000

- Setae j3 and j4 at most 0.6 times as long as distance between their bases and bases of j4

and j5, respectively............................................................................................................ 5

482

5. Unsclerotized integument along lateral margins of opisthonotal shield with five pairs of

setae (R1-R5). .............................................. Protogamasellopsis granulosus Karg, 1994b

- Unsclerotized integument along lateral margins of opisthonotal shield with 10 pairs of

setae (R1-R5, UR1-UR5) ................................................................................................... 6

6. Seta j5 at most half as long as distance between its base and base of j6; anterior margin

of opisthonotal shield with a pair of ellipsoidal structures ..................................................

…………………………………………...Protogamasellopsis dioscorus (Manson, 1972)

- Seta j5 about as long as distance between its base and base of j6; anterior margin of

opisthonotal shield without ellipsoidal structures………………………………………...

. ...................................................... Protogamasellopsis corticalis Evans and Purvis, 1987

Key to Protogamasellopsis species (known adult males)

1. With presternal plates ......................... Protogamasellopsis praeendopodalis Karg, 1994a

- Without presternal plates ................................................................................................... 2

2. Seta j5 at most half as long as distance between its base and base of j6; anterior margin

of opisthonotal shield with a pair of ellipsoidal structures ..................................................

…………………………………………...Protogamasellopsis dioscorus (Manson, 1972)

- Seta j5 about as long as distance between its base and base of j6; anterior margin of

opisthonotal shield without ellipsoidal structures………………………………………...

. ...................................................... Protogamasellopsis corticalis Evans and Purvis, 1987

References




2010.




483






21, p. 855–861, 1987.








n. 2, p. 179-216, 1994b.




243–262, 2000.








Acarologia, Paris, v. 13, p. 437–445, 1972.

484

MORAZA, M.L. Rhodacarella, a new genus of Rhodacaridae mites from North America

(Acari: Mesostigmata: Rhodacaridae). Zootaxa, Auckland, v. 470, p. 1–10, 2004.

SHAW, M. Mites and ticks from wedge-tailed shearwater (Puffinus pacificus) burrows on

Masthead Island. Queensland Naturalist, Brisbane, v. 37, p. 43–47, 1999.

WIŚNIEWSKI, J.; HIRSCHMANN, W. Protogamasellopsis posnaniensis nov. spec.

(Acarina: Mesostigmata) aus Palmenhaus in Polen. Bulletin of the Polish Academy of

Sciences, Biological Sciences, Warsaw, v. 39, p. 189–194, 1991.

485

6 RHODACARIDAE MITES (ACARI: MESOSTIGMATA: RHODACAROIDEA)

FROM THE STATE OF SÃO PAULO, BRAZIL, WITH DESCRIPTIONS OF A NEW

GENUS AND THREE NEW SPECIES

Abstract

This paper reports the occurrence of five species of Rhodacaridae mites collected in

the State of São Paulo, Brazil. One of these corresponded to a new genus and a new species,

described as Binodacarus brasiliensis Castilho and Moraes, 2010; two correspond to new

species of known genera, described as Multidentorhodacarus paulista Castilho and Moraes,

2010 and Rhodacarus matatlanticae Castilho and Moraes, 2010; and two correspond to

described species, Afrogamasellus citri Loots, 1969 and Protogamasellopsis dioscorus

(Manson, 1972).

Keywords: Litter; Rhodacaroidea; Soil; Taxonomy

Resumo

Esse artigo reporta a ocorrência de cinco espécies de ácaros Rhodacaridae coletadas

no Estado de São Paulo, Brasil. Uma dessas correspondeu a um novo gênero e espécie,

descrita como Binodacarus brasiliensis Castilho e Moraes, 2010; dois corresponderam a

novas espécies de gêneros conhecidos, descritas como Multidentorhodacarus paulista

Castilho e Moraes, 2010 e Rhodacarus matatlanticae Castilho e Moraes, 2010; e dois

corresponderam a duas espécies descritas, Afrogamasellus citri Loots, 1969 e

Protogamasellopsis dioscorus (Manson, 1972).

Palavras-chave: Folhedo; Rhodacaroidea; Solo; Taxonomia

6.1 Introduction

Rhodacaridae are edaphic mites found mainly in the first few centimeters of the soil

surface. Despite the scarce information available on their feeding behavior (WALTER;

HUNT; ELLIOTT, 1988; CASTILHO et al., 2009), rhodacarids are commonly mentioned in

the literature as predators (LEE, 1970; LINDQUIST; KRANTZ; WALTER, 2009).

486

Mites of this family are mostly known from the northern hemisphere (EVANS;

SHEALS; MACFARLANE, 1968; WALLWORK, 1970; PRICE, 1973; EVANS; TILL,

1979; KRANTZ; AINSCOUGH, 1990; COLEMAN; CROSSLEY JR., 1996) and Africa

(RYKE, 1962; VAN DEN BERG; RYKE, 1967; LOOTS, 1969a; LOOTS, 1969b;

HURLBUTT, 1974). Recent studies mentioned the occurrence of these mites in Brazil

(GNASPINI-NETO, 1989; TRAJANO; GNASPINI-NETO, 1991; MINEIRO; MORAES,

2001; SILVA; MORAES; KRANTZ, 2004), although, in all of these studies the rhodacarids

have been mentioned only at the generic level.

At present, there is a growing interest towards the use of more sustainable methods to

control pest organisms in different parts of the world. One of the possible alternatives is the

use of biological control. The objective of this paper is to report the rhodacarid species found

in surveys conducted in southeastern Brazil, when searching for prospective predatory mites

to be evaluated as potential biological control agents of soil pests.

6.2 Materials and methods

Litter and soil samples were collected in different parts of the State of São Paulo and

taken to a laboratory where mites were extracted using a modified Berlese-Tullgren apparatus

(OLIVEIRA et al., 2001). Mites of the suborder Mesostigmata were mounted in Hoyer‘s

medium and later examined under a phase contrast microscope.

Rhodacarid mites were then identified to species, as subsequently reported in this

paper. In the following descriptions, setal nomenclature is based on Lindquist and Evans

(1965). All measurements are given in micrometres (µm); each measurement consists of the

average for the number of individuals indicated for each species, followed (in parentheses) by

the respective range.

6.3 Results

Binodacarus Castilho and Moraes, 2010

Type species: Binodacarus brasiliensis Castilho and Moraes, 2010

Diagnosis: Adult female and male characterized mainly by having: only 2 scleronoduli on the

podonotal shield, between setae j5 and j6; podonotal shield reduced laterally, so that setae z1,

487

s2-s4 and r2-r5 are inserted on cuticle; a pair of elongate shields laterad of r3 and r4; and

greatly reduced peritreme (not reaching anterior margin of coxa IV).

Adult female: Fixed cheliceral digit with 4 teeth and movable cheliceal digit with 3 teeth, in

addition to the apical tooth of each digit. Palp chaetotaxy: 2-5-6-14-15; apotele 3-tined.

Deutosternal groove with 7 rows of denticles; with 6 pairs of ellipsoidal structures, apparently

corresponding to insertions of the ventral dilator muscles of the pharynx. Podonotal shield

with 14 pairs of setae (setae s1 and r1 absent; setae z1, s2-s4, r2-r5 on cuticle) and a pair of

crescent-shaped scleronoduli between setae j5 and j6. Cuticle along lateral margins of

podonotal shield with 2 pairs of elongated shields. Opisthonotal shield with 14 pairs of setae

(setae S4, R1-R5 on cuticle). Sternal shield longer than wide, with 4 pairs of setae and 3 pairs

of lyrifissures; seta st1 inserted in anterior punctate region of sternal shield; third pair of

lyrifissures on posterior margin of the shield. Genital shield longer than the length of its

straight posterior margin, extending posteriorly behind coxae IV. Cuticle between genital and

ventrianal shields with a pair of accessories shields. Cuticle between ventrianal and accessory

shields with 2 pairs of setae (Jv1 and Zv1). Ventrianal shield approximately as long as wide;

with 5 pairs of setae (Jv2-Jv4, Zv2-Zv3) in addition to circumanal setae. Peritreme greatly

reduced, not reaching anterior margin of coxa IV. Chaetotaxy of legs I-IV: coxae: 2, 2, 2, 1;

trochanters: 6, 5, 5, 5; femora: 13, 10, 6, 6; genua: 13, 11, 9, 9; tibiae: 14, 10, 8, 9; tarsi: not

counted, 18, 18, 17. All legs with pretarsi, each with 3 rounded pulvillar lobes, elongate

ambulacral stalk and strongly sclerotized claws.

Adult male: Fixed cheliceral digit with 4 teeth and movable cheliceral digit with 1 tooth, in

addition to apical tooth of each digit. Spermatodactyl longer than movable cheliceral digit.

Pattern of podonotal and opisthonotal shields as in female, except that in some individuals

with a pair of contiguous scleronoduli mediad to those also observed in female. Sternogenital

shield with 5 pairs of setae and 3 pairs of lyrifissures. Ventrianal shield with 5 pairs of setae

(Jv1-Jv3, Zv1-Zv2) in addition to circumanal setae. Cuticle posterolaterad of ventrianal shield

with 2 pairs of setae (Jv4 and Zv4). Metapodal shields absent. Peritreme as in female.

Chaetotaxy of all legs as in female, but a ventral setae of each genu and tibia II thorn-like, and

a ventral seta of femur II rod-like. All legs with pretarsi.

488

Etymology

The name Binodacarus refers to the presence of only two scleronoduli on the

podonotal shield. Gender masculine.

Remarks

Similarly to most Rhodacaroidea [except for Halolaelapidae; see Lindquist; Krantz

and Walter (2009)], mites of this genus have 4 pairs of sternal shield setae. This new genus is

placed in Rhodacaridae because of the following characteristics (in parentheses,

characteristics of other Rhodacaroidea): apotele 3-tined (2-tined in Digamasellidae, 3-tined in

Halolaelapidae, Laelaptonyssidae and Ologamasidae); scleronoduli present (absent in

Halolaelapidae, Laelaptonyssidae, most Ologamasidae, some Digamasellidae and some

Rhodacaridae); with desclerotized bands of punctate cuticle along posterior margin of

podonotal, anterior margin of opisthonotal, anterior margin of sternometasternal, posterior

margin of genital and anterior margin of ventrianal shields (desclerotized and punctuated

bands of cuticle absent in Digamasellidae, Halolaelapidae, Laelaptonyssidae and

Ologamasidae); genu I with 13 setae (12 in Digamasellidae and Halolaelapidae, 11 to 13 in

Laelaptonyssidae, 13 in Ologamasidae); tibia I with 14 setae (12 in Digamasellidae and

Halolaelapidae, 9 to 11 in Laelaptonyssidae, 14 in Ologamasidae); genu IV with 9 or 10 setae

(7 or 8 in Digamasellidae, 9 or 10 in Halolaelapidae, 7 to 10 in Laelaptonyssidae, 8 to 10 in

Ologamasidae); tibia IV with 9 or 10 setae (7 in Digamasellidae, 8 to 10 in Halolaelapidae, 7

to 10 in Laelaptonyssidae, 9 or 10 in Ologamasidae).

Females of the type species of the new genus differ from species in all other genera of

Rhodacaridae by having 2 scleronoduli (0, 3 or 4 scleronoduli in other genera); in addition, it

differs from species in most other genera of Rhodacaridae by having the following

combination of characteristics: anterior region of epistome triangular and with anterior margin

serrated (rather variable within the family), seta j2 not inserted in transverse line with setae j1

and z1 (in most other genera, these setae about transversely aligned), cuticle along lateral

margins of podonotal shield with 8 pairs of setae (up to 6 in other genera), cuticle along

lateral margins of opisthonotal shield with 6 pairs of setae (0 to 10 pairs in other genera),

peritreme not reaching anterior margin of coxa IV (in most other genera longer, reaching up

to anterior margin of coxa II) and genu and tibia IV with 9 setae each (9 or 10 setae in other

genera); species of no other genera display this combination of characteristics.

Short peritremes have also been reported for some species of each family of

Rhodacaroidea as well as for families of other superfamilies of parasitic and free-living

489

Mesostigmata. In this superfamily, scleronoduli are known for most genera of Rhodacaridae

(absent in Pennarhodeus Karg and Protogamasellopsis Evans and Purvis), most genera of

Digamasellidae (absent in Digamasellus Berlese, Lindquistoseius Genis, Loots and Ryke,

Panteniphis Willmann and Pontiolaelaps Luxton) and are rarely present in Ologamasidae

(some Gamasellus Berlese and some Rhodacaroides Willmann). In those cases, 2 to 4

scleronoduli are found. Besides the Rhodacaroidea, scleronoduli have been reported only in

Protogamasellus (Ascoidea). Scleronoduli are absent in Halolaelapidae and Laelaptonyssidae.

Binodacarus brasiliensis Castilho and Moraes, 2010

Diagnosis: Adult female and male: This species can be distinguished from other rhodacarids

by the combination of characters given in the diagnosis of the genus.

Adult female (Fig. 6.1 A–E) (5 specimens measured): All setae of gnathosoma, idiosoma and

legs sharp-tipped.

Gnathosoma: Fixed cheliceral digit 25 (23-26) long; movable cheliceral digit 26 (25-27)

long, with basal most tooth distinctly stronger; pilus dentilis and dorsal cheliceral seta

indistinct. Anterior region of epistome with a triangular projection; margin serrated.

Measurements of setae: h1 7 (6-7), h2 5 (5-6), h3 5 (5-6), h4 6 (5-6).

Dorsum of podosomal region: Podonotal shield smooth, except for a punctate band along

posterior margin; 114 (106-121) long and 90 (83-101) wide at widest level; apparently with 5

pairs of lyrifissures (anterolaterad of j2, anteromediad of z3, slightly posterior and mediad of

z5, anterolaterad of s6 and slightly posterior and mediad of r6). Cuticle laterad of r2 and r3

with a pair of punctate bands. Cuticle immediately anterior to podonotal shield with a

punctate band. Cuticle posteromediad of s2 with a pair of lyrifissures. Measurements of setae:

j1 7 (6-9), j2 6 (5-7), j3 4 (4-5), j4 5 (4-5), j5 4 (3-5), j6 4 (3-5), z1 5 (5-6), z2 5 (4-6), z3 5 (4-

6), z4 4 (3-5), z5 4 (4-5), z6 4 (4-5), s2 5 (4-5), s3 5 (4-6), s4 5 (4-6), s5 5 (4-6), s6 4 (4-5), r2

5, r3 5 (5-6), r4 5 (4-5), r5 4 (3-5), r6 5 (4-5).

Dorsum of opisthosomal region: Opisthonotal shield smooth, except for a punctate band

along anterior margin and anterior third of lateral margins; 108 (98-118) long and 76 (66-88)

490

Figure 6.1 - Binodacarus brasiliensis Castilho and Moraes. Female. A Dorsum of idiosoma; B. Venter of

idiosoma; C. Chelicera; D. Epistome; E. Hypostome

491

wide at widest level; apparently with 7 pairs of lyrifissures (posterolaterad of J1, posterior to

and almost in longitudinal line with S1, posteromediad of J2, posterior to and almost in

longitudinal line with S3, slightly anterior to and laterad of J4, anterolaterad of J5, slightly

anterior to and laterad of Z5). Cuticle immediately anterior of opisthonotal shield,

anterolaterad of J1, with a pair of lyrifissures. Measurements of setae: J1 4 (3-4), J2 3, J3 3,

J4 3 (3-4), J5 4, Z1 4 (3-4), Z2 3 (3-4), Z3 3, Z4 5 (4-5), Z5 6 (5-6), S1 4 (3-4), S2 4, S3 5 (4-

6), S4 4 (4-5), S5 6, R1 4 (3-5), R2 4 (3-4), R3 5 (4-5), R4 5 (4-5), R5 6 (5-6).

Venter of idiosoma: Sternal shield smooth, except for a weakly sclerotized and punctate

region anterior to first pair of lyrifissures; approximately 75 (71-81) long, including the

weakly sclerotized and punctate region, and approximately 40 (34-43) wide at widest level;

anterior margin indistinct; posterior margin concave. Genital shield smooth, except for a

punctate band along straight posterior margin; distance between st5-st5 26 (24-28). Ventrianal

shield mostly smooth, except for a punctate anteromedian lobe and for few diagonal lateral

striae; 86 (81-93) long and 79 (71-89) wide at widest level; apparently with 4 pairs of

lyrifissures (posterior to and slightly laterad of Jv1, posterolaterad of Zv2, anteromediad of

Jv3 and anterior to and almost in longitudinal line with Jv4); cribrum just off posterior margin

of ventrianal shield (given that the ventrianal shield extends very far back, in most specimens

cribum indistinct for being at the transition between the dorsal and the ventral surfaces of the

idiosoma). Metapodal shield subdivided into a small anterior fragment and a larger posterior

fragment. Cuticle posterolaterad of st5 and mesad of metapodal shield with a pair of

lyrifissures. Measurements of setae: st1 8 (7-10), st2 8 (6-9), st3 7 (5-8), st4 7 (5-8), st5 5 (4-

6), Jv1 5 (5-6), Jv2 5 (4-6), Jv3 5 (4-6), Jv4 5 (5-6), Zv1 5 (4-6), Zv2 4 (3-5), Zv4 6 (5-7).

Peritreme: Peritrematal shield indistinct.

Spermatheca: Not distinguishable.

Legs: Lengths of legs: I – 181 (179-185), II – 118 (113-121), III – 89 (88-91), IV – 147 (145-

151).

Adult male (Fig. 6.2 A–C) (4 specimens measured). All setae of gnathosoma, idiosoma and

legs sharp-tipped.

492

Gnathosoma: Epistome as in female. Fixed cheliceral digit 28 (24–32) long and movable

cheliceral digit 27 (23–31) long. Spermatodactyl 40 (39-41) long, abruptly curved at the base

and distally directed backwards; sharp-tipped. Measurements of setae: h1 6 (6-6), h2 5 (4-5),

h3 5 (5-6), h4 6 (5-7).

Dorsum of podosomal region: Podonotal shield 120 (106-126) long and 62 (52-73) wide at

widest level. Scleronoduli variable; in 2 specimens, similar to female; in other 2 specimens,

with a pair of contiguous scleronoduli mediad to those also observed in female; one or both of

the median scletonoduli incompletely formed. Measurements of setae: j1 5 (4-5), j2 4 (3-5), j3

5 (4-5), j4 5 (4-5), j5 3 (3-4), j6 3 (3-4), z1 4 (3-5), z2 4 (4-5), z3 5 (4-5), z4 4 (3-4), z5 4 (4-5),

z6 5 (5-6), s2 4 (3-4), s3 5 (4-6), s4 4 (3-4), s5 4 (3-5), s6 4 (4-5), r2 4 (3-4), r3 5 (4-5), r4 3

(3-4), r5 3 (2-3), r6 6 (5-6).

Dorsum of opisthosomal region: Opisthonotal shield 108 (99-117) long and 78 (61-88) wide

at widest level. Measurements of setae: J1 3 (3-4), J2 3 (3-3), J3 2 (2-3), J4 4 (3-4), J5 4 (4-

5), Z1 3 (3-4), Z2 3 (3-4), Z3 4 (3-4), Z4 5 (4-5), Z5 5 (4-5), S1 3 (3-4), S2 4 (4-5), S3 5 (4-5),

S4 4 (3-4), S5 5 (4-6), R1 4 (3-4), R2 4 (3-4), R3 4 (3-4), R4 3 (2-3), R5 5 (4-5).

Figure 6.2 - Binodacarus brasiliensis Castilho and Moraes. Male. A. Venter of idiosoma; B. Chelicera

493

Venter of idiosoma: Sternogenital shield smooth, except for punctate and weakly sclerotized

regions anterior to first pair of lyrifissures and posterior to setae st5; approximately 114 (100-

122) long, including the weakly sclerotized and punctate regions, and approximately 36 (32-

39) wide at widest level. Cuticle posterolaterad of st5 with a pair of lyrifissures. Ventrianal

shield centrally smooth, with a partially punctate band along anterior margin and with

diagonal lateral striae; approximately 90 (79-98) long and approximately 79 (63-86) wide at

widest level; with 5 pairs of lyrifissures (slightly anterior to and mesad of Jv1, posterolaterad

of Zv1, anteromediad of Jv2, posterolaterad of Zv2 and posterior to and almost in longitudinal

line with Jv3); as in female, cribrum difficult to see for being at the transition between the

dorsal and the ventral surfaces of the idiosoma. Measurements of setae: st1 8 (6-10), st2 7 (5-

7), st3 6 (5-6), st4 5 (4-6), st5 4 (3-4), Jv1 4 (3-4), Jv2 3 (3-4), Jv3 4 (4-5), Jv4 4 (4-5), Zv1 3

(2-3), Zv2 3 (2-4), Zv4 5 (4-5).

Legs: Lengths of legs: I – 188 (164–200), II – 122 (101–136), III – 104 (86–114) and IV –

146 (115–159)..

Locality and type material

Holotype female and one paratype male from soil under Psidium guajava L.

(Myrtaceae) at Pirassununga, 3-V-2000; 2 paratype males from the same substrate and

locality, 27-VII-2000; 1 paratype female from soil under Campomanesia pubescens (DC.) O.

Berg (Myrtaceae) at Luis Antonio, 2-V-2000; 2 paratype males from soil under Myrcia

guianensis (Aublet) DC. (Myrtaceae) at Luis Antonio, 27-VIII-2000; 2 paratype females from

soil under M. guianensis at São Carlos, 25-VII-2000. All types collected by A.R. Oliveira and

deposited at Departamento de Entomologia e Acarologia, Escola Superior de Agricultura

―Luiz de Queiroz‖, Universidade de São Paulo (ESALQ/USP), Piracicaba-SP, Brazil.

Etymology

This species is named after Brazil, the country of origin of this species.

Multidentorhodacarus paulista Castilho and Moraes, 2010

Diagnosis: Adult female with podonotal shield containing a punctate band along posterior

margin and a pair of fissures laterad of setae s1, z2 and z3. Opisthonotal shield with a

punctate band along median third of anterior margin. Setae r5, R2 and R5 inserted on cuticle.

494

Adult female (Fig. 6.3 A–F) (5 specimens measured): All setae of gnathosoma, idiosoma and

legs sharp-tipped.

Gnathosoma: Fixed cheliceral digit 54 (52-55) long, with 11 teeth and a distinct pilus dentilis

in addition to apical tooth; 2 basal teeth distinctly stouter; movable cheliceral digit 49 (48-51)

long, with 5 teeth, in addition to apical tooth of each digit. Anterior margin of epistome with 3

extensions; median extension clavate, with distal half denticulate; lateral extensions triangular

to truncate, slightly less than half as long as median extension, each with 2-3 distal denticles;

margin of epistome laterad of lateral extensions denticulate. Deutosternal groove with 8 rows

of denticles. Measurements of setae: h1 11 (10-13), h2 10 (9-11), h3 11 (11-12), h4 11 (10-

12).

Dorsum of podosomal region: Podonotal shield smooth, except for a punctate band along

posterior margin; 138 (136-141) long and 147 (140-149) wide at widest level; with a V-

shaped suture adjacent and posterior to bases of setae j4 and s2; with a pair of fissures laterad

of setae s1, z2 and z3; with 22 pairs of setae (seta r1 absent; seta r5 on cuticle), 4 pairs of

which (j1, j2, z1, s1) next to anterior margin; with 3 crescent-shaped scleronoduli between

setae j5 and j6 and apparently with 6 pairs of lyrifissures (laterad of and about in transversal

line with j2, posterolaterad of j3, laterad of and about in transversal line with z3, anterior to

and about in longitudinal line with z4, anterolaterad of z5 and laterad of and about in

transversal line with j6). Measurements of setae: j1 13 (12-13), j2 14 (13-14), j3 17 (16-17), j4

16 (15-17), j5 16 (15-16), j6 16 (15-16), z1 7 (6-7), z2 11 (10-12), z3 19 (17-20), z4 18 (16-

19), z5 16 (15-16), z6 19 (18-20), s1 6 (6-7), s2 18 (17-19), s3 21 (19-22), s4 18 (17-19), s5

18 (17-18), s6 18 (18-19), r2 13 (12-15), r3 18 (17-19), r4 23 (22-24), r5 15 (14-16), r6 15

(14-16).


along median third of anterior margin; 129 (123-134) long and 94 (87-100) wide at widest

level; with 18 pairs of setae (setae R2 and R5 on cuticle) and apparently with 6 pairs of

lyrifissures (anterolaterad of J1, posteromesad of J1, posteromesad of J2, anterolaterad of S3,

posteromesad of S3 and posteromesad of J4). Measurements of setae: J1 17 (16-18), J2 16

(16-17), J3 17 (15-18), J4 17 (15-18), J5 13 (12-14), Z1 17 (16-18), Z2 17 (16-18), Z3 20 (18-

22), Z4 18 (17-20), Z5 22 (20-24), S1 18 (16-19), S2 18 (16-19), S3 17 (16-17), S4 18 (15-19),

495

Figure 6.3 - Multidentorhodacarus paulista Castilho and Moraes. Female. A. Dorsum of idiosoma; B. Venter of

idiosoma; C. Chelicera; D. Epistome; E. Hypostome; F. Variations of section of spermathecal

apparatus

496

S5 21 (20-21), R1 13 (12-14), R2 12 (11-13), R3 9 (8-9), R4 12 (11-13), R5 14 (13-16).

Venter of idiosoma: Sternal shield smooth, except for a weakly sclerotized and punctuated

region whose posterior margin is slightly posterior to st1; approximately 89 (87-92) long,

including the weakly sclerotized and punctate region, and approximately 48 (44-53) wide at

widest level; anterior margin indistinct; posterior margin with small and sharp median

projection; with 4 pairs of setae and 3 pairs of lyrifissures (the third pair on posterior margin

of the shield). Genital shield smooth, except for a punctate band along convex posterior

margin; longer than length of posterior margin; extending posteriorly well behind coxae IV;

distance between st5-st5 31 (28-33). Cuticle posterolaterad of st5 with a pair of lyrifissures.

Ventrianal shield smooth, except for a punctate band along most of anterior margin; 90 (85-

93) long and 74 (67-81) wide at widest level; with 5 pairs of setae (Jv1-Jv3, Sv1-Sv2) in

addition to circumanal setae and apparently with 3 pairs of lyrifissures (posterolaterad of Zv1,

anterior to and about in longitudinal line with Zv2 and about in transversal line with Jv3);

ventrianal shield not fused to dorsal shield. Cuticle along lateral margins of ventrianal shield

with 2 pairs of setae (Zv3 and UR). With a pair of elongated metapodal shields. Measurements

of setae: st1 16 (15-17), st2 16 (15-17), st3 15 (14-15), st4 16 (14-17), st5 12 (11-12), Jv1 14

(13-14), Jv2 15 (14-15), Jv3 17 (16-18), Zv1 12 (11-13), Zv2 13 (12-14), Zv3 16 (15-16), UR

6 (7-6).

Peritreme: Extending anteriorly almost to level of anterior margin of coxa III. Peritrematal

shield indistinct.

Spermatheca: Tubular section of what seems to be part of spermathecal apparatus

distinguishable near coxa IV.

Legs: Lengths of legs: I – 246 (241-250), II – 177 (171-182), III – 165 (163-169), IV – 234

(231-237). Chaetotaxy of legs I-IV: coxae: 2, 2, 2, 1; trochanters: 6, 5, 5, 5; femora: 13, 10, 6,

6; genua: 13, 11, 9, 10; tibiae: 14, 10, 8, 10; tarsi: not counted, 18, 18, 17. Pretarsi only on

legs II - IV.

Adult male: Not found.

497


Holotype female and 2 paratype females from soil under Syagrus oleracea (Mart.)

Becc. (Arecaceae) at Piracicaba, 11-XI-2000; 8 paratype females (17-V-2000) and 6 paratype

females (11-XI-2000) from soil under Euterpe edulis Mart. (Arecaceae) at Piracicaba; 9

paratype females (17-V-2000), 6 paratype females (11-VIII-2000) and 5 paratype females

(11-XI-2000) from soil under Syagrus romanzoffiana (Cham.) Glassman (Arecaceae) at

Piracicaba; 2 paratype females (16-II-2000), 1 paratype female (16-V-2000) and 1 paratype

female (10-VIII-2000) from soil under S. oleracea at São Pedro; 1 paratype female (11-VII-

2000) from litter under E. edulis at Pariquera-Açu; 3 paratype females (19-VII-2000) from

soil under Campomanesia pubescens (D.C.) O. Berg (Myrtaceae) at Luis Antônio; 1 paratype

female (2-V-2000) and 3 paratype females (26-VII-2000) from soil under Myrcia guianensis

(Aubletet) D.C. (Myrtaceae) at Luis Antônio; 2 paratype females (26-VII-2000) from soil

under Psidium guajava L. (Myrtaceae) at Luis Antônio; 2 paratype females (27-VII-2000)

from soil under C. pubescens at Pirassununga; 3 paratype females (27-VII-2000) from soil

under M. guianensis at Pirassununga; 3 paratype females (27-VII-2000) from soil under P.

guajava at Pirassununga; 2 paratype fêmeas (25-VII-2000) from soil under M. guianensis at

São Carlos. All types collected by A.R. Oliveira and deposited at Departamento de

Entomologia e Acarologia, Escola Superior de Agricultura ―Luiz de Queiroz‖, Universidade

de São Paulo (ESALQ/USP), Piracicaba-SP, Brazil.

Etymology

The term paulista means ―from the State of São Paulo‖, referring to the Brazilian state

where the type specimens were collected.

Remarks

As a member of Multidentorhodacarus Karg, the species here described has a large

number of teeth on the fixed cheliceral digit (other species generally only with up to 9 teeth),

a V-shaped suture adjacent and posterior to bases of setae j4 and s2, and setae j1, j2, z1 and s1

on anterior margin of podonotal shield, no pre-sternal shield and 3 scleronoduli.

Multidentorhodacarus paulista Castilho and Moraes, 2010 is similar to Multidentorhodacarus

ruwenzoriensis (Loots, 1969a), but the latter does not have the anterolateral fissures laterad of

setae s1, z2 and z3, lacks the punctate bands along the posterior margin of the podonotal and

the anterior margin of the opisthonotal shields, has the posterior margin of podonotal shield

rounded and setae r5 and R5 inserted on podonotal and opisthonotal shields, respectively.

498

Rhodacarus matatlanticae Castilho and Moraes, 2010

Diagnosis: Adult female with 6 pairs of setae in addition to circumanal setae on the ventrianal

shield, median extension of anterior margin of epistome spatulate and mostly smooth, except

for 3 apical denticles, and with 3 podonotal scleronoduli transversely aligned..

Adult female (Fig. 6.4 A–E) (2 specimens measured). All setae of gnathosoma, idiosoma and

legs sharp-tipped.

Gnathosoma: Fixed cheliceral digit 55-58 long, with 5 teeth in addition to apical tooth but

without distinct pilus dentilis; movable cheliceral digit 61-63 long, with 3 teeth, in addition to

apical tooth of each digit. Anterior margin of epistome with 3 smooth extensions; median

extension spatulate, with 3 apical denticles; lateral extensions triangular, slightly less than half

as long as median extension. Deutosternal groove with 8 rows of.denticles. Measurements of

setae: h1 8-9, h2 8, h3 8-9, h4 11.

Dorsum of podosomal region: Podonotal shield smooth, except for a narrow punctate band

along bases of setae z6, s6, r5 and r6; 134-144 long and 145-149 wide at widest level; with 21

pairs of setae (seta r1 absent; setae r3 and r4 on cuticle), 4 pairs (j1, j2, z1, s1) on anterior

margin; with 3 transversally aligned, crescent-shaped scleronoduli immediately posterior to

setae j5; apparently with 7 pairs of lyrifissures (posteromesad of z1, posteromesad of j3,

anteromesad of s3, posterolaterad of j4, anterolaterad of j6, anterolaterad of r5 and

anteromesad of z6). Measurements of setae: j1 9, j2 8-9, j3 10-11, j4 10-11, j5 10, j6 10, z1 6,

z2 8, z3 6-7, z4 9-10, z5 10-11, z6 11, s1 4, s2 6-7, s3 5-6, s4 9-10, s5 10-11, s6 11, r2 6-7, r3

16, r4 6, r5 7-8, r6 11-12.


along anterior margin; 143-151 long and 101-106 wide at widest level; with 19 pairs of setae

(seta R1 on cuticle); apparently with 7 pairs of lyrifissures (anteromesad of J1, posteromesad

of J2, posteromesad of Z2, posterolaterad of S3, laterad of and about in transversal line with

J3, anteromesad of S5 and anterior to and about in longitudinal line with J5). Measurements

of setae: J1 11, J2 10, J3 10-11, J4 14, J5 6-7, Z1 11, Z2 10, Z3 12, Z4 12, Z5 22-25, S1 8, S2

9, S3 8, S4 8, S5 9-11, R1 6-7, R2 6, R3 6, R4 7, R5 13-15.

499

Figure 6.4 - Rhodacarus matatlanticae Castilho and Moraes. Female. A. Dorsum of idiosoma; B. Venter of

idiosoma; C. Chelicera; D. Epistome; E. Hypostome

500

Venter of idiosoma: Sternal shield smooth, except for a weakly sclerotized and punctuated

region, whose posterior margin is slightly posterior to st1, and for a punctuated region around

st4; approximately 94-95 long, including the weakly sclerotized and punctate region, and

approximately 47-55 wide at widest level; anterior margin indistinct; posterior margin with

small and sharp median projection; with 4 pairs of setae and 3 pairs of lyrifissures (the third

pair on posterior margin of the shield). Genital shield smooth, except for a punctate band

along straight posterior margin; extending posteriorly well behind coxae IV; distance between

st5-st5 25-27; with 1 pair of pores posterolaterad of st5. Ventrianal shield smooth, except for a

punctate band along anterior margin; 92-102 long and 76-81 wide at widest level; with 6 pairs

of setae in addition to circumanal setae; apparently with 4 pairs of lyrifissures (posterior to

and about in longitudinal line with Zv1, posteromesad of Jv3, posteromesad of Zv3 and

anterolaterad of paraanals); ventrianal shield not fused to dorsal shield. Cuticle posterolaterad

of Zv2 apparently with a pair of lyrifissures. With a pair of ellipsoidal metapodal shields.

Cuticle anterolaterad of metapodal shields with a pair of lyrifissures. Measurements of setae:

st1 9-10, st2 9-10, st3 9-10, st4 9-10, st5 8-9, Jv1 10, Jv2 11-12, Jv3 11, Jv4 10, Zv1 8-9, Zv2

7-8.

Peritreme: Extending anteriorly almost to median level of coxa III. Peritrematal shield

indistinct.

Spermatheca: Not distinguishable.

Legs: Lengths of legs: I – 265-277, II – 177 189-192, III – 163-170, IV – 214-217.

Chaetotaxy of legs I-IV: coxae: 2, 2, 2, 1; trochanters: 6, 5, 5, 5; femora: 13, 10, 6, 6; genua:

13, 11, 9, 10; tibiae: 14, 10, 8, 10; tarsi: not counted, 18, 18, 17. Pretarsi only on legs II - IV.

Adult male: Not found.


Holotype female and 1 paratype female from soil under Bactris setosa (Arecaceae) at

Cananéia, 12-VII-2000. Types collected by A.R. Oliveira and deposited at Departamento de

Entomologia e Acarologia, Escola Superior de Agricultura ―Luiz de Queiroz‖, Universidade

de São Paulo (ESALQ/USP), Piracicaba-SP, Brazil.

501

Etymology

The term matatlanticae means ―from Mata Atlântica‖, referring to the Brazilian

vegetation type from which the type specimens were collected.

Remarks

As a member of the genus Rhodacarus Oudemans, the species here described has a

relatively small number of teeth on the fixed digit, setae j1, j2, z1 and s1 on anterior margin of

podonotal shield, no pre-sternal shield and 3 scleronoduli. Rhodacarus aequalis Karg, 1971,

Rhodacarus agrestis Karg, 1971 and Rhodacarus calcarulatus Berlese, 1920 are similar to

this new species. However, in those species the median extension of the anterior margin of the

epistome is thorn-like and distally serrated and slightly inflated; Zv3 is inserted on cuticle; and

there seems to be 1 (R. agrestis and R. calcarulatus) or 2 (R. aequalis) additional

opisthogastric setae posterolaterad of the ventrianal shield. Additionally, in R. calcarulatus

and R. agrestis the median podonotal scleronodulus is distinctly anterior to the lateral

scleronoduli.

Afrogamasellus citri Loots, 1969

Afrogamasellus citri Loots, 1969a: 75-77, Figs. 64-70; Hurlbutt, 1974: 588.

Remarks

The Brazilian specimen differs from the original description in that it has 3 instead of

4 scleronoduli. Otherwise, it fits well the original description of this species. Information not

given in the original description are subsequently provided, based on the specimen collected

in the present work.

Fixed cheliceral digit 24 long, with 5 teeth in addition to apical tooth; movable

cheliceral digit 25 long, with 3 teeth, in addition to apical tooth of each digit; h1 9, h2 7, h3 8,

h4 9. Podonotal shield 137 long and 121 wide at widest level; j1 9, j2 11, j3 12, j4 11, j5 10,

j6 11, z1 6, z2 6, z3 9, z4 12, z5 12, z6 13, s1 4, s2 6, s3 13, s4 16, s5 15, s6 15, r2 13, r3 17,

r4 7, r5 9, r6 16. Opisthonotal shield 129 long and 108 wide at widest level; J1 12, J2 12, J3

11, J4 14, J5 14, Z1 15, Z2 15, Z3 15, Z4 14, Z5 23, S1 15, S2 15, S3 14, S4 16, S5 17, R1 11,

R2 10, R3 10, R4 11. Sternal shield approximately 83 long, including the weakly sclerotized

and punctate region anterior to first pair of lyrifissures, and approximately 64 wide at widest

level. Distance st5-st5 44. Ventrianal shield 102 long and 89 wide at widest level; st1 13, st2

502

13, st3 12, st4 11, st5 12, Jv1 12, Jv2 13, Jv3 13, Jv4 14, Zv1 13, Zv2 10, Zv3 14, Zv4 16.

Lengths of legs: I – 261, II – 172, III – 153, IV – 218.

Material examined:

One female from soil under Psidium guajava L. (Myrtaceae) at Luis Antônio, 26-VII-

2000.

Protogamasellopsis dioscorus (Manson, 1972)

Protogamasellus dioscorus Manson, 1972: 437-444, Figs. 1-21.

Protogamasellopsis dioscorus, Evans and Purvis, 1987: 855.

Remarks

The specimens collected fit well the original description of this species. Information

not given in the original description are subsequently provided, based on the 3 female

specimens collected in the present work.

Fixed cheliceral digit 43 (37-48) long, with 5 teeth in addition to apical tooth; movable

cheliceral digit 42 (36-47) long, with 2 teeth, in addition to apical tooth of each digit; h1 29

(27-30), h2 12 (11-12), h3 28 (27-28), h4 22 (21-23). Podonotal shield 208 (206-209) long

and 134 (133-135) wide at widest level; j1 37 (36-37), j2 26 (25-27), j3 27 (26-27), j4 26 (25-

26), j5 22 (21-22), j6 21 (20-21), z1 20 (20-19), z2 35 (34-35), z3 26 (25-26), z4 20, z5 26, z6

34 (33-35), s1 17 (16-17), s2 22 (22-23), s3 25 (24-26), s4 39 (38-40), s5 38 (37-39), s6 39

(38-39), r2 34 (33-34), r3 43 (42-43), r4 26, r5 33 (32-33). Opisthonotal shield 226 (224-228)

long and 81 (80-82) wide at widest level; J1 24, J2 24 (23-24), J3 22 (21-22), J4 26 (25-26),

J5 13 (12-14), Z1 22 (21-23), Z2 25 (24-25), Z3 29 (29-30), Z4 36 (35-36), Z5 49 (48-49), S1

38 (36-38), S2 32, S3 32 (31-33), S4 37 (36-37), S5 32 (31-33), R1 33 (32-33), R2 29 (28-29),

R3 30, R4 33 (32-34), R5 37 (36-37), UR1 30 (28-31), UR2 30 (29-30), UR3 29 (27-30), UR4

39, UR5 43 (42-43). Sternal shield approximately 121 (119-122) long, including the weakly

sclerotized and punctate region anterior to first pair of lyrifissures, and approximately 98 (94-

101) wide at widest level. Distance st5-st5 49 (48-49). Ventrianal shield 155 (152-157) long

and 61 (59-62) wide at widest level; st1 23 (22-24), st2 31 (30-31), st3 28 (27-29), st4 24 (23-

24), st5 19 (18-19), Jv1 19 (18-19), Jv2 24 (23-24), Jv3 30, Jv4 30 (30-31), Zv1 23 (22-23),

Zv2 21, Zv3 35 (34-36). Lengths of legs: I – 360 (357-362), II – 252 (240-265), III – 240

(232-247), IV – 370 (367-372).

503

Material examined

Three females from soil under Psidium guajava L. (Myrtaceae) at Pirassununga, 01-

XI-2000.

6.4 Discussion

Before the publication of this paper, only 15 of the ca. 144 rhodacarid species known

worldwide were described from South America, more specifically from Argentina, Ecuador

and Venezuela. The present paper gives the first description of a genus of this family from

South America. At this stage, it is difficult to estimate the diversity of this group in Brazil,

given the relatively small effort that has been dedicated to the knowledge of these mites in

this country. However, the fact that about 65% of the known rhodacarid species were

described from tropical and subtropical countries (R.C. Castilho, G.J. de Moraes and R.B.

Halliday, unpublished) suggests to be high the diversity of rhodacarids in Brazil, a large

country consisting of tropical and subtropical regions.

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EVANS, G.O.; SHEALS, J.G.; MACFARLANE, D. The terrestrial Acari of the British

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507

7 TWO NEW SPECIES OF RHODACARIDAE (MESOSTIGMATA:

RHODACAROIDEA) FROM IRAN

Abstract

Multidentorhodacarus n. sp. 1 and Rhodacarellus n. sp. 2 (Acari: Mesostigmata:

Rhodacaroidea: Rhodacaridae) are described and figured. These are the first descriptions of

Rhodacaridae mites from Iran.

Keywords: Multidentorhodacarus; Rhodacarellus; Soil mites; Taxonomy

Resumo

Multidentorhodacarus n. sp. 1 e Rhodacarellus n. sp. 2 (Acari: Mesostigmata:

Rhodacaroidea: Rhodacaridae) são descritos e ilustrados. Estas são as primeiras descrições de

ácaros Rhodacaridae do Irã.

Palavras-chave: Multidentorhodacarus; Rhodacarellus; Ácaros de solo; Taxonomia.

7.1 Introduction

Rhodacaridae Oudemans is a family of widespread free living mites found in the soil

and in accumulations of decaying organic material such as compost, manure and tidal debris

(LINDQUIST; KRANTZ; WALTER, 2009). They have been found to feed on insect larvae,

springtails, nematodes and mites (KARG, 1971; LEE, 1974; SARDAR; MURPHY, 1987;

WALTER; HUNT; ELLIOTT, 1988; LINDQUIST; KRANTZ; WALTER, 2009). The

biology of mites of this group is poorly known, but at least one species has been reported as

potentially useful as a biological control agent of insect and mite pests in the soil

(CASTILHO et al., 2009).

The world-wide catalogue of Rhodacaridae included 148 species in 15 genus

(CASTILHO; MORAES; HALLIDAY, 2012). Multidentorhodacarus Karg and

Rhodacarellus Willmann are two of the most diverse genera of Rhodacaridae, with 16 and 20

described species, respectively (CASTILHO; MORAES; HALLIDAY, 2012). The objective

508

of this paper is to provide taxonomic descriptions of two new species of Rhodacaridae found

in surveys conducted in Iran.


Orchard soil samples were collected in different parts of Isfahan and Kerman

Provinces of center and south-east of Iran, respectively, and taken to a laboratory where mites

were extracted using a Berlese funnel. Mesostigmatid mites were mounted in Hoyer‘s

medium and later separated into families. Rhodacarid mites were separated into

morphospecies, and representative samples of each morphospecies were sent for examination

under a phase contrast microscope at Escola Superior de Agricultura ―Luiz de Queiroz‖,

Universidade de São Paulo, Piracicaba, State of São Paulo, Brazil.

The mites were identified to genera and determined to correspond to two new species.

They were illustrated with the use of a camera lucida and measurements were taken of

structures considered taxonomically important. In the following descriptions, setal

nomenclature is based on Lindquist and Evans (1965) as adapted to rhodacarid species by

Castilho and Moraes (2010), with modification in relation to the identification of s and r setae

for Multidentorhodacarus, considered necessary after our examination of most species of this

genus (unpublished). Measurements of each structure are given in micrometres, with the

average measurement for the individuals examined followed (in parentheses) by the

respective range (for variable measurement).

7.3 Results

Multidentorhodacarus Karg

Multidentorhodacarus Karg, 2000b: 144 (described in Rhodacaridae).

Rhodacarus (Multidentorhodacarus) Shcherbak, 1980: 72 (nomen nudum).

Rhodacarus (Multidentorhodacarus).— Karg, 1996: 170 (nomen nudum).

Rhodacarus (Multidentrhodacarus) [sic].— Karg, 1998: 186 (nomen nudum).

Multidentorhodacarus.— Karg, 2000a: 259 (nomen nudum).

Multidentorhodacarus.— Castilho et al., 2011: in press.

Type species: Rhodacarus denticulatus Berlese, 1920, by original designation.

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NOTE: According to Castilho; Moraes and Halliday (2012), Multidentorhodacarus was

described as a subgenus of Rhodacarus by Shcherbak (1980), but the name was not made

available because the type species of the subgenus was not fixed (International Code of

Zoological Nomenclature, Article 13.3). The name only became available when Karg (2000b)

published a redescription of Multidentorhodacarus, now at the genus level, fixing Rhodacarus

denticulatus Berlese, 1920 as type species. The names of species described in the genus

Multidentorhodacarus before 2000 are available, even though the genus name was not

available at the time (International Code of Zoological Nomenclature, Article 11.9.3.1).

Multidentorhodacarus sp. n. 1

Diagnosis of adult females: As a member of Multidentorhodacarus, this species has a relatively

large number of teeth (more than eight) on the fixed cheliceral digit; arthrodial process of

chelicera shaped as a coronet-like fringe; setae j1, j2, z1 and s1 along anterior margin of

podonotal shield; V-shaped line immediately behind j4 and s2; three scleronoduli near j5; no

presternal shield.

Multidentorhodacarus sp. n. 1 has fixed and movable cheliceral digits with 10 and 4-5

teeth in addition to the apical tooth of each respective digit; epistome with anterior margin serrate

and provided with three extensions; pair of anterolateral extensions of epistome about a third the

length of anterocentral extension, all serrate; dorsum of podosomal region with 23 pairs of setae,

all of which, except r4, on podonotal shield; podonotal shield without fissures laterad of s1, z2

and z3; posterior margin of podonotal shield truncate and with a narrow punctate band; dorsum of

opisthosomal region with 20 pairs of setae, all of which, except R2 and R5, on opisthonotal

shield; anterior margin of opisthonotal shield convex and with a narrow punctate band; seta J4

about half as long as the distance between its base and the base of J5; seta Z3 twice as long as Z4;

setae S1 and S2 not reaching the bases of S2 and S3, respectively; seta Zv1 on unsclerotised

integument along lateral margin of ventrianal shield; peritrematal shield narrow, restricted to

region along peritreme.

Adult female (Fig. 7.1 A-F) (five specimens measured): All setae smooth.

Gnathosoma: Fixed cheliceral digit 55 (54-56) long, with ten teeth in addition to apical tooth

and a setiform pilus dentilis (Fig. 7.1A); movable cheliceral digit 54 (53-55) long, with 4-5

teeth in addition to apical tooth, the proximal considerably larger than others. Epistome with

510

Figure 7.1 - Multidentorhodacarus sp. n. 1 Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C.


improved visibility

511

anterior margin serrate and provided with three extensions; pair of anterolateral extensions

about a third the length of anterocentral extension, all serrate (Fig. 7.1B). Deutosternal

denticles in eight roughly transverse rows of about uniform lengths, each with 8-13 denticles

(Fig. 7.1C). Seta h3 directly posterior to h2. Measurements of setae: h1 14 (14-15), h2 12 (12-

13), h3 13 (12-13), sc 13 (13-14).

Dorsal idiosoma (Fig. 7.1D): Podonotal shield smooth, except for a narrow punctate band

along the straight posterior margin; 144 (142-145) long and 135 (134-136) wide at widest

level; with 22 pairs of setae (r1 absent) and seven pairs of distinguishable lyrifissures.

Unsclerotised integument along lateral margins of podonotal shield with a pair of setae (r4).

Opisthonotal shield smooth, except for a narrow punctate band along convex anterior margin;

139 (138-140) long and 90 (89-91) wide at widest level; with 18 pairs of setae and eight pairs

of distinguishable lyrifissures. Unsclerotised integument along lateral margins of opisthonotal

shield with two pairs of setae (R2 and R5). Measurements of setae: j1 13 (13-14), j2 16 (15-

17), j3 17 (16-17), j4 18 (17-19), j5 16 (15-16), j6 16 (15-17), z1 8 (8-9), z2 15 (15-16), z3 17

(16-17), z4 18 (17-19), z5 17 (16-18), z6 20 (19-20), s1 8 (8-9), s2 19 (18-20), s3 15 (14-15),

s4 21 (20-21), s5 20 (19-21), s6 18 (18-19), r2 19 (18-19), r3 30 (28-31), r4 16 (14-18), r5 22

(20-24), r6 20 (19-22), J1 18 (17-19), J2 17 (17-18), J3 18 (17-20), J4 19 (18-19), J5 15 (15-

16), Z1 19 (18-19), Z2 18 (17-19), Z3 20 (20-21), Z4 13 (10-15), Z5 37 (36-38), S1 17 (16-

17), S2 18 (17-18), S3 17 (17-18), S4 18 (17-19), S5 19 (17-21), R1 11 (10-12), R2 11 (10-11),

R3 10 (9-10), R4 12 (11-12), R5 19 (19-20).

Ventral idiosoma (Fig. 7.1E): Base of tritosternum 12 (12-13) long and 9 (8-9) wide

proximally (Fig. 7.1F); laciniae 46 (43-50), separated for about 95% of their total length,

slightly pilose. Sternal shield smooth and with anterior margin indistinct; region anterior to

the first pair of sternal setae (st1) lightly sclerotised and punctate; posterior margin with short

central spine-like projection; approximately 92 (91-93) long, from anterior margin of lightly

sclerotised punctate region to tip of medial projection of posterior margin, 63 (61-64) wide at

widest level; with four pairs of setae and three pairs of lyrifissures. Genital shield smooth,

except for a punctate ellipsoidal region near convex posterior margin; longer than wide;

extending posteriorly well behind coxae IV; distance between st5-st5 31 (30-33).

Unsclerotised integument posterolaterad of st5 with a pair of lyrifissures. Ventrianal shield

smooth, except for a narrow punctate band along central region of convex anterior margin;

100 (98-103) long and 64 (63-66) wide at widest level, not fused to dorsal shield; with four

512

pairs of setae (Jv1-Jv3 and Zv2) in addition to circum-anal setae and four pairs of

distinguishable lyrifissures; post-anal seta about 1.6 times as long as para-anal setae.

Unsclerotised integument along margins of ventrianal shield with three pairs of setae (Jv4,

Zv1 and Zv3). With a pair of elongate metapodal plates well behind insertion of coxa IV.

Peritreme extending anteriorly to level of median region of coxa III. Peritrematal shield

narrow, restricted to region along peritreme, sharp-tipped anteriorly. Measurements of setae:

st1 18, st2 18 (18-19), st3 17 (16-17), st4 16 (15-16), st5 13 (12-14), Jv1 16 (15-16), Jv2 18

(17-19), Jv3 18, Jv4 11 (10-12), Zv1 13 (13-14), Zv2 14, Zv3 11 (9-12), para-anal 24 (23-25),

post-anal 41 (40-42).

Legs: lengths: I: 245 (243-248); II: 183 (181-185); III: 173 (171-174); IV: 227 (226-229).

Numbers of setae on legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6;

genu: 13, 11, 9, 10; tibia: 14, 10, 8, 10; tarsi II-IV: 18, 18, 17. Pretarsus I absent; pretarsi II-IV

similar in shape and length, each consisting of an elongate ambulacral stalk, a pair of strongly

sclerotised claws and three rounded pulvillar lobes.

Adult male: Unknown.

Material examined

Holotype female and one paratype female from soil of an orchard at Khomani Shahr

(32°41‘N, 51°32‘E, alt. 1602 m), Isfahan Province, Iran, 7 July 2002; one paratype female

from soil of an orchard at Najafabad (32°38‘N, 51°23‘E, alt. 1754 m), Isfahan Province, Iran,

11 September 2003; one paratype female from soil of an orchard at Dorcheh (32°38‘N,

51°39‘E, alt. 1570 m), Isfahan, Isfahan Province, Iran, 20 August 2003; one paratype female

from soil of an orchard at Jiroft (28°40‘N, 57°44‘E, alt. 690 m), Kerman Province, Iran, 30

July 1998. All types collected by M. Jalaeian. The holotype and two paratype females are

deposited in the Acarological Collection, Zoological Museum, College of Agriculture, Tehran

University, Karaj, Iran; two paratypes female deposited at Departamento de Entomologia e

Acarologia, Escola Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de

São Paulo (USP), Piracicaba, State of São Paulo, Brazil.

Remarks

Multidentorhodacarus n. sp. 1 is most similar to Multidentorhodacarus sogdianus

(Shcherbak, 1980), but the latter has fixed and movable cheliceral digits with 13 and six teeth

513

in addition to apical tooth, respectively; the punctate band of the podonotal shield removed

from the posterior margin of the shield; without punctate bands along anterior margin of the

opisthonotal shield, posterior margin of genital shield and anterior margin of ventrianal shield;

with R5 inserted on opisthonotal shield; peritrematal shield indistinct.

Rhodacarellus Willmann

Rhodacarellus Willmann, 1935: 429 (described in Rhodacaridae).

Rhodacarellus.— Evans, 1957: 221; Sheals, 1958: 302; Lee, 1970: 36; Karg, 1971: 323;

Bregetova and Shcherbak, 1977: 272; Evans and Till, 1979: 206; Shcherbak, 1980:

76; Fouly, 1992: 436; Karg, 1993: 336; Castilho et al., 2011: in press.

Type species: Rhodacarellus subterraneus Willmann, 1935, by original designation.

Rhodacarellus sp. n. 1

Diagnosis of adult females: As a member of Rhodacarellus, this species has the arthrodial

process of chelicera shaped as a coronet-like fringe; podonotal shield mostly smooth, with j1,

j2 and z1 along anterior margin, without transverse line between j4 and j5 and with four

scleronoduli near j5; no presternal shield; basitarsus IV with three setae (pl4 absent).

Rhodacarellus sp. n. 1 has epistome with anterior margin smooth and provided with

three extensions and a pair of short spines; pair of spatulate anterolateral extensions of

epistome about two thirds the length of the aciculate anterocentral extension, all smooth;

podonotal shield with 19 pairs of setae; opisthonotal shield smooth, except for a narrow

punctate band along slightly convex anterior margin; seta Z5 longer than Z3; unsclerotised

integument posterolaterad of st5 with two pairs of lyrifissures; setae Jv1 and Zv1 on

unsclerotised integument along anterior margin of ventrianal shield; ventrianal shield mostly

smooth, with inverted U-shaped punctate band, and with lateral margins straight to slightly

convex; seta Jv4 present; post-anal seta about 1.7 times as long as para-anal seta; peritreme

extending anteriorly to level of posterior margin of coxa II.

Adult female (Fig. 7.2 A-F) (Five specimens measured): All setae smooth.

Gnathosoma: Fixed cheliceral digit 33 (33-34) long, with five teeth in addition to apical tooth

and a setiform pilus dentilis (Fig. 7.2A); movable cheliceral digit 35 (34-35) long, with three

teeth in addition to apical tooth. Epistome with anterior margin smooth and provided with

514

Figure 7.2 - Rhodacarellus sp. n. 1 Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C. Hypostome;

D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures enlarged for improved

visibility

515

three extensions and a pair of short spines; pair of spatulate anterolateral extensions about two

thirds the length of the aciculate anterocentral extension, all smooth (Fig. 7.2B). Deutosternal

denticles in seven roughly transverse rows of about uniform lengths, each with 8-14 denticles

(Fig. 7.2C). Seta h3 directly posterior to h1 and lateral to h2. Measurements of setae: h1 15

(15-16), h2 8 (8-9), h3 14 (13-14), sc 13 (13-14).

Dorsal idiosoma (Fig. 7.2D). Podonotal shield smooth, except for a narrow punctate band

next to the insertions of j1, j2 and z1, and along straight posterior margin; 142 (139-144) long

and 134 (132-135) wide at widest level; with 19 pairs of setae (s1 and r1 absent) and seven

pairs of distinguishable lyrifissures. Unsclerotised integument along lateral margins of

podonotal shield with three pairs of setae (s2, s3 and r4). Opisthonotal shield smooth, except

for a narrow punctate band along slightly convex anterior margin; 159 (157-160) long and

104 (101-106) wide at widest level; with 15 pairs of setae, nine pairs of distinguishable

lyrifissures and a pair of pore anterolaterad of J5; with a pair of ellipsoidal structure posterior

to J3 and a pair of ellipsoidal and a pair of rounded structures posterior to J4, apparently

corresponding to muscles insertions. Unsclerotised integument along lateral margins of

opisthonotal shield with four pairs of setae (R1-R4); seta R5 absent. Measurements of setae: j1

12, j2 15 (14-15), j3 13 (12-13), j4 16 (15-16), j5 13 (13-14), j6 14 (13-15), z1 6 (5-7), z2 14

(13-14), z3 15 (14-15), z4 14 (14-15), z5 15 (14-15), z6 18 (18-19), s2 5 (5-6), s3 6, s4 21 (20-

22), s5 21 (21-22), s6 16 (16-17), r2 5 (5-6), r3 27 (26-28), r4 8 (7-8), r5 9 (8-10), r6 23 (22-

24), J1 13 (12-13), J2 13, J3 12 (12-13), J4 17 (17-18), J5 15 (14-15), Z1 17 (17-18), Z2 17

(17-18), Z3 33 (32-33), Z4 21 (21-22), Z5 42 (41-42), S1 16 (16-17), S2 15 (15-16), S3 18

(17-18), S4 23 (23-24), S5 24 (23-24), R1 9 (8-9), R2 7 (6-8), R3 7 (7-8), R4 11 (11-12).



pilose. Sternal shield smooth and with anterior margin indistinct; region anterior to the first

pair of lyrifissures lightly sclerotised and punctate; posterior margin concave; approximately

89 (87-92) long, including the lightly sclerotised punctate region, 69 (68-70) wide at widest

level; with four pairs of setae and three pairs of lyrifissures. With a pair of ellipsoidal

structure between sternal and genital shields, apparently corresponding to muscles insertions.

Genital shield smooth, except for a punctate ellipsoidal region near convex posterior margin;

longer than wide; extending posteriorly behind coxae IV; distance between st5-st5 34 (32-36).

Unsclerotised integument posterolaterad of st5 with two pairs of lyrifissures. Unsclerotised

516

integument along anterior margin of ventrianal shield with two pairs of setae (Jv1 and Zv1);

distinct ellipsoidal structure anterior to ventrianal shield might correspond to a transversal

fold of the cuticle of this region. Ventrianal shield mostly smooth, with inverted U-shaped

punctate band; 107 (106-108) long and 93 (90-95) wide at widest level, not fused to dorsal

shield; with lateral margins straight to slightly convex; with five pairs of setae (Jv2-Jv4, Zv2

and Zv3) in addition to circum-anal setae, and three pairs of distinguishable lyrifissures; post-

anal seta about 1.7 times as long as para-anal setae; with a pair of ellipsoidal and a pair of

rounded structures posterolaterad and posteromesad of Jv3, respectively, apparently

corresponding to muscles insertions. With a pair of elongate metapodal plates, whose anterior

end is round and contains a pore-like structure, located well behind insertion of coxa IV.

Peritreme extending anteriorly to level of posterior margin of coxa II. Peritrematal shield not

much wider than peritreme in the region next to it, constrict behind stigma and fused to

exopodal and endopodal shields next to coxa IV, constrict anterior to peritreme, but

expanding beyond s3 and extending to level between z1 and z2; distally approaching but not

fusing to dorsal shields. Measurements of setae: st1 15 (14-15), st2 15 (15-16), st3 14 (14-15),

st4 15 (14-15), st5 12 (11-13), Jv1 9 (9-10), Jv2 14 (13-14), Jv3 16 (15-16), Jv4 30 (28-31),

Zv1 12 (11-12), Zv2 11 (11-12), Zv3 16 (15-17), para-anal 25 (23-26), post-anal 44 (42-45).


Numbers of setae on legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6;

genu: 13, 11, 9, 10; tibia: 14, 10, 8, 10; tarsi II-IV: 18, 18, 17. Pretarsi I-IV similar in shape,

each consisting of an elongate ambulacral stalk, a pair of strongly sclerotised claws, and three

rounded pulvillar lobes; pretarsus I about half as long as other pretarsi.

Adult male: Unknown.

Material examined

Holotype female and one paratype female from soil of an orchard at Najafabad

(32°38‘N, 51°23‘E, alt. 1754 m), Isfahan Province, Iran, 12 April 2003; three paratype

females from soil of an orchard at Borkhar (32°81‘N, 51°54‘E, alt. 1590 m), Isfahan

Province, Iran, 22 July 2003; three paratype females from soil of an orchard at Falavarjan

(30°51‘N, 51°32‘E, alt. 1600 m), Isfahan Province, Iran, 27 Jun 2003; one paratype female

from soil of an orchard at Khomani Shahr (32°41N, 51°32‘E, alt. 1602 m), Isfahan Province,

Iran, 7 July 2002. All types collected by M. Jalaeian. The holotype and five paratype females

517

are deposited in the Acarological Collection, Zoological Museum, College of Agriculture,

Tehran University, Karaj, Iran; three paratype females deposited at Departamento de

Entomologia e Acarologia, Escola Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ),

Universidade de São Paulo (USP), Piracicaba, State of São Paulo, Brazil.

Remarks

Rhodacarellus n. sp. 1 is most similar to Rhodacarellus tebeenus Hafez and Nasr,

1979, but the latter has genital shield with posterior margin truncate and without punctate

band; a pair of metapodal plates of uniform width and without pore-like structure;

peritrematal shield not fused to exopodal and endopodal shields next to coxa IV; and genua III

and IV and tibia IV with eight, eight and nine setae, respectively.

7.4 Discussion

Before the publication of this paper, only three rhodacarid species had been recorded

from Iran, Rhodacarellus epigynialis Sheals, Rhodacarellus silesiacus Willmann and

Rhodacarus sp. (KAMALI; OSTOVAN; ATAMEHR, 2001; HADDAD IRANINEZHAD;

HAJI GHANBAR; TALEBI CHAICHI, 2003). Dendroseius amoliensis Faraji, Sakenin-

Chelav and Karg, 2006 (Digamasellidae, described as Rhodacaridae) was the only species of

Rhodacaroidea, to which Rhodacaridae belong, described from Iran. Multidentorhodacarus n.

sp. 1 and Rhodacarellus sp. n. 1 are the first species of Rhodacaridae described from Iran.

Only nine of the 148 rhodacarid species known worldwide were described from Asia; three

species from mainland China, two from Tajikistan, two from Turkmenistan, one from Hong

Kong and one from Indonesia (CASTILHO; MORAES; HALLIDAY, 2012). At this stage, it

is difficult to estimate the diversity of this group in Iran, given the relatively small effort that

has been dedicated to its knowledge. The very diverse climatic conditions prevailing in that

country, from very hot and dry in the center and south to very cold (winter) in northwest,

suggests that a diversity of rhodacarid species could be found in Iran.

References

BERLESE, A. Centuria quinta di Acari nuovi. Redia, Florence, v. 14, p. 143-195, 1920.

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Mesostigmata. Leningrad: Nauka. 1977. p. 256-307. [in Russian]




2010.












FARAJI, F.; SAKENIN-CHELAV, H.; KARG, W. A new species of Dendroseius Karg from

Iran (Acari: Rhodacaridae), with a key to the known species. Zootaxa, Auckland, v. 1221, p.

63-68, 2006.

FOULY, A.H. Rhodacarellus citri, a new mite species from Egypt (Acari: Rhodacaridae).

Egyptian Journal of Applied Science, Cairo, v. 7, n. 3, p. 434–441, 1992.

HADDAD IRANINEZHAD, K.; HAJI GHANBAR H.R.; TALEBI CHAICHI P. Introduction

of some mesostigmatic mites of sugar beet fields in Miandoab plain. Journal of Agricultural

Sciences and Natural Resources, Gorgan, v. 2, n. 10, p. 147-157, 2003.

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77-81, 1979.

KAMALI, K., OSTOVAN, H.; ATAMEHR, A. A catalog of mites and ticks (Acari) of

Iran. Tehran: Islamic Azad University Scientific Publication Center, 2001. 192 p.



475 p.


Leach, Raubmilben. Teil 59. Jena: Gustav Fischer Verlag, 1993. 523 p.










243–262, 2000a.



72, p. 207–213, 2000b.

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219, 1970.









Press, 2009. p. 124-232.

SARDAR, M.A.; MURPHY, P.W. Feeding tests of grassland soil-inhabiting gamasine

predators. Acarologia, Paris, v. 28, p. 117-121, 1987.





13, v. 1, p. 298-304, 1958.


predatory arthropods from a shortgrass steppe soil. Pedobiologia, Jena, v. 31, p. 247-260,

1988.

WILLMANN, C. Ueber eine eigenartige Milbenfauna im Küstengrundwasser der Kieler

Bucht. Schriften des Naturwissenschaftlichen Vereins fur Schleswig–Holstein, Kiel, v. 20,

p. 422–434, 1935.

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8 A NEW SPECIES OF Gamasiphis (ACARI: OLOGAMASIDAE) FROM BRAZIL,

WITH A KEY TO SPECIES FROM THE NEOTROPICAL REGION

Abstract

Gamasiphis paulista Castilho, Moraes and Narita, 2010 was described based on

specimens representing all postembryonic stages, collected from litter and soil in Piracicaba,

State of São Paulo. This is the first species of Gamasiphis described from Brazil. A key is

provided for the separation of species of this genus known from the Neotropical Region.

Keywords: Mesostigmata; Rhodacaroidea; Edaphic mite; Litter; Taxonomy

Resumo

Gamasiphis paulista Castilho, Moraes e Narita, 2010 foi descrito com base em

espécimes representando todas os estágios pós-embriônicos, coletados em folhedo e solo em

Piracicaba, Estado de São Paulo. Essa é a primeira espécie de Gamasiphis descrita do Brasil.

Uma chave é construída para a separação das espécies descritas desse gênero da região

Neotropical.

Palavras-chave: Mesostigmata; Rhodacaroidea; Ácaros edáfico; Folhedo; Taxonomia

8.1 Introduction

Ologamasidae (Mesostigmata: Rhodacaroidea) is a large, widely distributed group of

predatory mites that inhabit soil, humus and compost (LINDQUIST; KRANTZ; WALTER,

2009). They are some of the most ubiquitous edaphic Mesostigmata in southeastern Brazil

(MINEIRO; MORAES, 2001; SILVA; MORAES; KRANTZ, 2004).

Gamasiphis Berlese is one of the most diverse genera of Ologamasidae, with 68

described species (CASTILHO; SILVA; MORAES, 2009a). According to Karg and

Schorlemmer (2009), Gamasiphis species occur mainly in tropical and subtropical areas.

Seventeen species of this genus have been reported from the Neotropical Region (FOX, 1949;

KARG, 1990, 1994a, 1994b, 1998, 2003, 2007; KARG; SCHORLEMMER, 2009).

522

Undetermined species also have been reported from Brazil (MINEIRO; MORAES, 2001;

SILVA; MORAES; KRANTZ, 2004).

Gamasiphis species have been considered as predators of soil mites (KARG;

SCHORLEMMER, 2009; LINDQUIST; KRANTZ; WALTER, 2009), although only a single

detailed biological study concerning a species of this genus has been conducted (LEE, 1974).

Given the high frequency in which ologamasids are commonly found in southeastern Brazil,

an effort has been made to evaluate their possible role as biological control agents of pest

organisms in this region. As part of this process, we were recently able to establish a

laboratory culture of an undescribed species of Gamasiphis, whose biology is presently under

investigation. The objective of this paper is to provide a taxonomic description of the post-

embryonic stages of this new species and a taxonomic key to separate the species of this

genus known from the Neotropical Region.

8.2 Materials and methods

Litter and soil samples were collected on the campus of Escola Superior de

Agricultura ―Luiz de Queiroz‖, Universidade de São Paulo, Piracicaba, State of São Paulo,

and taken to a laboratory where mites were extracted with a Berlese funnel. Ologamasids

collected were then sorted under dissecting microscope, and a culture of the most numerous

species was established according to the procedure described by Castilho et al. (2009b). After

three months of the establishment of the colony, samples of each developmental stage of the

mite were taken from the colony, cleared in Nesbitt‘s medium and mounted in Hoyer´s

medium for examination under phase-contrast microscopy. Samples of adult males and

females were also taken from the colony, coated with gold for scanning electron microscopy;

most of the photos obtained were not suitable for publication, but they were very useful in the

determination of several of the characteristics mentioned in this paper.

In the following description, dorsal idiosomal setal nomenclature is based on Lindquist

and Evans (1965). Measurements of each structure are given in micrometres, with the average

measurement for the individuals examined followed (in parentheses) by the respective range

(for variable measurement). The identification key to the Gamasiphis species from the

Americas was prepared based on the original descriptions and available redescriptions of the

species concerned.

523

8.3 Results

Gamasiphis Berlese

Gamasiphis Berlese, 1904: 261. Type species: Gamasus pulchellus Berlese, 1887, by original

designation.

Gamasiphis.— Berlese, 1906: 101; Berlese, 1914: 137; Bregetova, 1977: 308; Lee, 1970: 42;

Karg, 1990: 321; Karg, 1993: 169.

Ologamasellus (Micriphis) Berlese, 1914: 140. Type species: Gamasiphis gamasellus

Berlese, 1913, by monotypy. Synonymy by Lee (1970).

Ologamasus (Micriphis).— Baker and Wharton, 1952: 73.

Micriphis.— Ryke, 1962: 160.

Gamasiphis (Heteroiphis) Trägårdh, 1952: 55. Type species: Gamasiphis (Heteroiphis)

arcuatus Trägårdh, 1952, by original designation. Synonymy by Lee (1970).

Heteroiphis.— Ryke, 1962: 160; Bhattacharyya, 1968: 530.

Neogamasiphis Trägårdh, 1952: 57. Type species: Neogamasiphis hamifer Trägårdh, 1952, by

original designation. Synonymy by Lee (1970).

Diagnosis (adult female and male)

Species of this genus are characterised by having the anterior margin of the epistome

with at least one projection; the idiosoma dorsally convex and extensively covered by

sclerotised shields; podonotal and opisthonotal shields fused to form a holonotal shield

without a distinct line of fusion; setae j3 and r3 not inserted on tubercles; the holonotal shield

fused with the ventri-anal shield; the exopodal shield undivided adjacent to coxa III; the

peritrematal shield and part of the exopodal shield adjacent to coxa IV fused into a single unit,

which is either fused to the ventri-anal shield or separated from it by a narrow line of

unsclerotised cuticle; metapodal shield typically indistinct, either fused to neighbouring

shields or, often, apparently longitudinally elongate and located externally to and along the

posterior end of the fused peritrematal and exopodal shields.

Gamasiphis paulista Castilho, Moraes and Narita, 2010

Diagnosis of adult

524

Fixed and movable cheliceral digits with seven and four teeth respectively in the

female and seven and one teeth respectively in the male, in addition to the apical tooth of each

digit. Podosomal region of dorsal shield with 23 pairs of setae and opisthosomal region with

10 pairs of setae in both sexes; setae acicular, except j3-j6, z3-z5, s4, s5 and r5 lanceolate,

with constricted base. Ventral idiosoma with two pairs of presternal shields; ventri-anal shield

with eight pairs of setae in addition to circumanal setae. Female with seta st3 mesad and

slightly posterior to seta st2; male with st3 about in longitudinal line with st2 and st4.

Adult female (Fig. 8.1 A–F) (5 specimens measured): Setae of gnathosoma, idiosoma and

legs acicular, except j3-j6, z3-z5, s4, s5 and r5 lanceolate, with constricted base.

Gnathosoma: Fixed cheliceral digit 72 (72-73) long, with seven teeth in addition to apical

tooth and a setiform pilus dentilis (Fig. 8.1A); movable cheliceral digit 76 (75-77) long, with

four teeth in addition to apical tooth. Antiaxial lyrifissure distinct. Palp chaetotaxy 2-5-6-14-

15; apotele 3-tined. Anterior margin of epistome with an elongate, acute central projection

flanked by a pair of short, acute lateral projections (Fig. 8.1B). Deutosternal denticles in six

rows, with 10-14 denticles each (Fig. 8.1C); most anterior row ―V‖ shaped, followed by an

inverted ―V‖ shaped row; subsequent rows roughly transverse. Seta h2 shorter than h1 and h3.

Measurements of setae as in Table 8.1.

Dorsal idiosoma (Fig. 8.1D): Dorsum totally covered by a large, smooth dorsal shield

extending latero-ventrally to fuse or abut adjacent shields; 537 (530-550) long and 388 (363-

385) wide at widest point; apparently with two pairs of pores and eight pairs of lyrifissures.

Podosomal region with 23 pairs of setae (j1-j6, z1-z6, s1-s6 and r2-r6). Opisthosomal region

with 10 pairs of setae (J3-J5, Z1, Z3-Z5, S2-S4). Measurements of dorsal setae as in Table 8.1.

Ventral idiosoma (Fig. 8.1E): Base of tritosternum partially covered by the distal end of a

partially subdivided, apparently retractable lobe; distal margin of each half of the lobe with 3-

7 spine-like structures; trapezoidal base of tritosternum 14 (12-16) long and 13 (12-14) wide

proximally; laciniae 62 (58-66), totally separated from each other, pilose. Wide distinct

sections of exopodal shield present between coxae I-II and coxae II-IV; those sections

separated from peritrematal shield by a line of unsclerotised cuticle. Fused peritrematal shield

and section of exopodal shield next to leg IV extending posteriorly well beyond stigma, wider

525

Figure 8.1 - Gamasiphis paulista Castilho, Moraes and Narita, 2010. Female. A. Lateral (antiaxial) view of

chelicera; B. Epistome; C. Hypostome, D. Dorsal shield; E. Ventral idiosoma, F. Variations of

section of spermathecal duct near coxa IV. Pores and lyrifissures enlarged to allow their

differentiation

526

Table 8.1 - Measurements (µm) of the dorsal setae of each postembryonic stage of Gamasiphis paulista Castilho,

Moraes and Narita, 2010. (-) Absent

(continues)

Seta Adult female Adult male Deutonymph Protonymph Larva

j1 13 (12–15) 14 (13–15) 19 (18–20) 18 (15–20) 22 (20–24)

j2 26 (25–27) 24 (23–25) 25 (23–27) 39 (38–40) –

j3 51 (50–53) 45 (45–46) 46 (45–47) 38 26

j4 54 (53–55) 46 (45–48) 54 (53–55) 49 (48–50) 38 (37–39)

j5 43 41 (40–41) 36 (35–37) 34 (33–35) 24 (23–25)

j6 47 (45–50) 42 (40–45) 52 (50–55) 66 (65–68) 67 (65–70)

z1 8 (8–9) 7 (5–8) 10 (8–12) – –

z2 6 (5–6) 6 (5–6) 23 (20–25) 25 21 (20–22)

z3 15 (14–16) 14 (14–15) 35 – –

z4 43 (42–43) 37 (35–39) 52 (50–53) 46 (45–47) 36 (35–37)

z5 41 (40–42) 40 (40–41) 46 (45–47) 54 (53–55) 59 (58–60)

z6 45 (45–46) 45 (43–47) 14 (13–15) – –

s1 8 (7–8) 7 (7–8) 8 – –

s2 8 (7–8) 8 (7–8) 8 – –

s3 16 (15–17) 12 (12–13) 24 (23–25) – –

s4 53 (50–55) 51 (50–51) 43 (40–45) 41 (40–42) 66 (65–67)

s5 50 (48–52) 44 (43–44) 52 (50–53) 58 (55–60) –

s6 14 (13–15) 15 34 (33–35) 39 (38–40) –

r2 7 (6–7) 6 (6–7) 8 (7–8) 28 (26–30) –

r3 9 (8–10) 9 (9–10) 68 (65–70) 73 (70–75) –

r4 14 (13–14) 9 (8–9) 20 – –

r5 11 (10–12) 9 (8–10) 48 (45–50) 26 (25–27) –

r6 13 (12–15) 10 (9–10) 24 (23–25) – –

J3 11 (10–12) 9 (8–10) 12 (10–13) 66 (63–70) –

J4 9 (8–10) 7 (7–8) 9 (8–10) 69 (68–70) 50 (48–52)

J5 9 (8–10) 6 (5–7) 9 (8–10) 39 (38–40) 119 (115–123)

Z1 7 (7–8) 7 (7–8) 73 (72–73) 78 –

Z3 10 (8–12) 11 (10–12) 97 (97–98) 100 (98–102) 97 (93–100)

Z4 10 (8–11) 8 (7–8) 8 63 109 (105–112)

Z5 77 (75–80) 65 (63–67) 84 (80–87) 98 (95–100) 51 (50–52)

527

Table 8.1 - Measurements (µm) of the dorsal setae of each postembryonic stage of Gamasiphis paulista Castilho,


(conclusion)

Seta Adult female Adult male Deutonymph Protonymph Larva

S2 10 (9–10) 6 (5–7) 13 (10–15) 65 (63–67) –

S3 9 (8–9) 8 (7–8) 12 (10–13) 65 (63–67) 17 (16–18)

S4 9 (7–8) 6 (5–8) 10 62 (60–63) 21 (20–22)

at level of coxa IV and progressively narrowing posteriorly to a pointed tip [pars interior of

Karg (1990)]; separated from adjacent shields by lines of unsclerotised cuticle. Peritreme

extending anteriorly almost to level of anterior margin of coxa I. An elongate triangular shield

present laterad of posterior end of fused peritrematal shield and section of exopodal shield

next to leg IV [pars exterior of Karg (1990); perhaps corresponding to metapodal shield of

other Mesostigmata], flanked externally by a ribbon-shaped shield and separated from the

ventral extension of the posterior dorsal shield by a line of unsclerotised cuticle; this ribbon-

shaped shield fused anteriorly to dorsal shield and posteriorly to ventri-anal shield, bearing a

pore at the level of Zv1 (probably Rp of Lindquist and Evans, 1965). Endopodal shield totally

fused to sternal shield. Pre-sternal area with two pairs of smooth narrow presternal shields,

roughly parallel to each other and to the anterior margin of the sternal shield. Sternal shield

reticulate between st1 and st3 and smooth posteriorly; posterior margin deeply concave;

approximately 68 (63-72) long at mid-line and 92 (88-97) wide at widest point (endopodal

projection between coxae II and III); with four pairs of setae, st3 inserted posteromesad of st2,

and four pairs of lyrifissures; lyrifissure next to st2 oriented obliquely to longitudinal axis of

idiosoma. Genital shield with few diagonal striae; fitting into posterior concavity of sternal

shield; length shorter than width along posterior margin; posterior margin straight, slightly

anterior to posterior margin of coxa IV; distance between st5-st5 69 (67-72). Ventri-anal

shield transversely striated; separated from adjacent shields by lines of unsclerotised cuticle;

anal region of the shield separated from ventral region by an unsclerotised line, except for

area between setae Jv3, and totally fused to posterior end of ventral extension of dorsal shield;

ventri-anal shield 268 (245-288) long at mid-line (from anterior margin to post-anal seta), 280

(257-302) wide at widest point; with eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to

circumanal setae; Jv5 about level with anterior margin of anal opening and about 1.1 times as

long as para-anal seta; post-anal seta about 3.5 times as long as para-anal seta; shield with a

pair of pores and two pairs of lyrifissures. Measurements of ventral setae as in Table 8.2.

528

Spermatheca (Fig. 8.1F): Lightly sclerotised tubular section of part of spermathecal

apparatus distinguishable near coxa IV.

Table 8.2 - Measurements (µm) of ventral setae of each postembryonic stage of Gamasiphis paulista Castilho,


Seta Female Male Deutonymph Protonymph Larva

h1 31 (30–31) 29 (28–30) 23 22 (20–23) 21 (20–22)

h2 24 (24–25) 22 (22–23) 23 23 22

h3 25 (25–26) 24 (23–25) 25 24 (23–25) –

Sc 25 (23–26) 23 (23–24) 23 20 –

st1 16 (15–17) 11 (10–11) 21 (20–23) 21 (20–23) 19 (18–20)

st2 21 (20–22) 16 (15–17) 23 20 18

st3 13 14 (13–15) 23 (22–23) 20 20

st4 13 (12–13) 15 (14–17) 21 (20–23) – –

st5 15 (15–16) 16 (15–17) 20 8 –

Jv1 27 (27–28) 24 (24–25) 24 (23–25) 22 10

Jv2 23 (20–25) 24 22 22 24 (23–25)

Jv3 23 (22–23) 21 (20–22) 20 – –

Jv4 30 (28–32) 26 (25–27) 30 – –

Jv5 16 (15–17) 15 28 (25–30) 62 (60–63) 12 (11–13)

Zv1 20 (18–22) 23 (22–23) 23 (20–25) – –

Zv2 32 (30–33) 27 (27–28) 21 (20–23) 32 (30–33) 9 (8–10)

Zv3 23 (21–24) 19 (17–20) 26 (25–27) – –

Para-anal 15 (14–16) 16 (15–18) 15 (14–16) 26 (25–28) 49 (48–51)

Post-anal 53 (51 –54) 73 (71–75) 49 (48–50) 55 (53–56) 61 (64–59)


Chaetotaxy of legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6; genu: 13,

11, 9, 8; tibia: 14, 10, 8, 9; tarsus II-IV: 18, 18, 17. All legs with pretarsi, each with three

rounded pulvillar lobes, elongate ambulacral stalk and a pair of strongly sclerotised claws.

Adult male (Fig. 8.2 A-C) (5 specimens measured). Setae of gnathosoma, idiosoma and legs

similar to those of female.

529

Gnathosoma: Fixed cheliceral digit 56 (55-57) long, with seven teeth (pilus dentilis not

distinguishable) in addition to apical tooth (Fig. 8.2A); movable cheliceral digit 60 (59–60)

long, with one tooth in addition to apical tooth. Spermatodactyl 72 (70-75) long, curved,

apparently with an internal canal in proximal half, distal half spatulated, distally blunt.

Epistome and hypostome as in female. Seta h2 shorter than h1 and h3. Measurements of setae

as in Table 8.1.

Dorsal idiosoma: Dorsal shield similar to that of female. Measurements of setae as in Table

8.1.

Ventral idiosoma (Fig. 8.2B): Trapezoidal base of tritosternum 18 (17-19) long and 17 (16-

18) wide proximally; laciniae 66 (65-68), otherwise as in adult female. Except for the fusion

of sternal and genital shields, shape, pattern and fusions of ventral shields as in female.

Ribbon-shaped shield with a pore at level of Zv1 or slightly anterior. Sternogenital shield with

sparse reticulation between st1 and st2, smooth elsewhere; approximately 143 (142-145) long

and 101 (98-105) wide at widest point (at endopodal projection between coxae II and III);

with five pairs of setae and three pairs of lyrifissures; lyrifissure next to st2 oriented obliquely

to longitudinal axis of idiosoma; genital opening on anterior margin of shield, flanked by two

rounded flaps, each ending in a marginal row of elongated, pointed extensions. Ventri-anal

shield 270 (263-277) long at mid-line (from anterior margin to post-anal seta) and 293 (280-

312) wide at widest point, with eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to

circumanal setae; post-anal seta about 4.5 times as long as para-anal seta; with a pair of pores

and two pairs of lyrifissures. Measurements of ventral setae as in Table 8.2.


Chaetotaxy similar to that of female, but two ventral setae on femur II and genu II and one

ventral seta of tibia II spur-like (Fig. 8.2C). All legs with pretarsi, similar to those of adult

female

Deutonymph (Fig. 8.2 D-E) (3 specimens measured). Setae of gnathosoma, idiosoma and

legs similar to those of adult female.

530

Gnathosoma: Fixed cheliceral digit 50 long, with seven teeth in addition to apical tooth and a

setiform pilus dentilis; movable cheliceral digit 53 (52-54) long, with four teeth in addition to

apical tooth. Epistome as in female. Hypostomal setae of similar lengths. Measurements of

setae as in Table 8.1.

Dorsal idiosoma (Fig. 8.2D): Podonotal and opisthonotal shields separated. Podonotal shield

smooth; 223 (214-232) long and 368 (357-380) wide at widest point; with 21 pairs of setae

(j1-j6, z1-z6, s1-s6, r2, r3 and r5), three pairs of pores and four pairs of lyrifissures. Seta r4 on

unclerotised cuticle laterad of podosomal shield and seta r6 on unclerotised cuticle between

podonotal and opisthonotal shields. Opisthonotal shield reticulate, 187 (185-188) long and

322 (317-327) wide at widest point; with ten pairs of setae (J3-J5, Z1, Z3-Z5, S2-S4), two

pairs of pores, nine pairs of lyrifissures and a pair of pore-like structures anterolaterad of S2.


Ventral idiosoma (Fig. 8.2E): Trapezoidal base of tritosternum 25 (24-25) long and 19 (18-

20) wide proximally;laciniae 59 (56-62), otherwise as in adult female. Vestigial exopodal and

endopodal shields present between coxae II and III and between coxae III and IV.

Peritrematal shield distinct. Peritreme short, extending anteriorly only to median level of coxa

III. Presternal shields absent. Sternal shield reticulate, narrowed behind st4, with rounded

posterior margin; approximately 137 (135-140) long and 78 (75-80) wide at widest point (at

level of lyrifissure posterior to st2); with four pairs of setae (st1-st4), approximately aligned

longitudinally, and three pairs of lyrifissures. Setae st5 inserted on unsclerotised cuticle, at

level of posterior end of sternal shield. Two pairs of small, narrow platelets present posterior

to st5. Setae Jv1-Jv2, Jv4-Jv5 and Zv1-Zv3 and five pairs of lyrifissures present on

unsclerotised cuticle. Ventri-anal shield reticulate anterolaterally and smooth elsewhere; 67

(62-72) long along midline and 81 (78-85) wide at widest point; with seta Jv3 in addition to

circumanal setae; post-anal seta about 3.2 times as long as para-anal seta. Measurements of



Chaetotaxy of legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6; genu: 13,

11, 8, 8; tibia: 14, 10, 8, 9; tarsus II-IV 18, 18, 17. All legs with pretarsi similar to those of

adult female.

531

Figure 8.2 - Gamasiphis paulista Castilho, Moraes and Narita, 2010. A. Male, lateral (antiaxial) view of

chelicera; B. Male, ventral surface of idiosoma; C. Male, femur, genu and tibia of leg II; D.

Deutonymph, dorsal idiosoma, E. Deutonymph, ventral idiosoma. Pores and lyrifissures enlarged

to allow their differentiation

532

Protonymph (Fig. 8.3 A-B) (3 specimens measured): Setae of gnathosoma, idiosoma and

legs acicular, except j4, j5 and z5 lanceolate, with constricted base.


tooth and a setiform pilus dentilis; movable cheliceral digit 42 (41-43) long, with four teeth in

addition to apical tooth. Epistome as in female. Hypostomal setae of similar lengths.


Dorsal idiosoma (Fig. 8.3A): Podonotal and opisthonotal shields separated. Podonotal shield

smooth, 177 (170-185) long and 75 (74-76) wide at widest point; with 11 setae (j1-j6, z2, z4,

z5, s4 and s5) and two pairs of lyrifissures. Unclerotised cuticle laterad of podosomal shield

with four pairs of setae (s6, r2, r3 and r5), two pairs of lyrifissures (in one specimen, the

anterior most inserted at edge of podosomal shield), a pair of narrow platelets anterior to r2,

and a pair of smaller platelets bearing r5 and a pore-like structure. Unclerotised cuticle

between podonotal and opisthonotal shield with a pair of setae (Z1), three pairs of lyrifissures

(between j6 and J3) and three pairs of platelets. Opisthonotal shield reticulated; 100 (98-102)

long and 178 (175-182) wide at widest point; with six pairs of setae (J3-J5, Z3-Z5), a pair of

pores and three pairs of lyrifissures. Unsclerotised cuticle laterad of opisthonotal shield with

three pairs of setae (S2-S4) and a pair of lyrifissures. Measurements of setae as in Table 8.1.

Ventral idiosoma (Fig. 8.3B): Trapezoidal base of tritosternum 19 (18-20) long and 16 (15-

17) wide proximally; laciniae 48 (45-50), otherwise as in adult female. Peritrematal shield

slightly longer than peritreme, reaching anterior level of coxa IV. Sternal shield very lightly

sclerotised; 111 (107-115) long and 71 (70-72) wide at widest point (at level of lyrifissure

posterior to st2); with three pairs of setae (st1-st3) and two pairs of lyrifissures. Setae st5

inserted on unsclerotised cuticle, at level of posterior margin of coxa IV. Setae Jv1, Jv2, Jv5

and Zv2 and five pairs of lyrifissures on unsclerotised cuticle. Anal shield smooth; 53 (48-57)

long along midline and 69 (68-70) wide at widest point; post-anal seta about twice as long as

para-anal seta. Measurements of setae as in Table 8.2.


Chaetotaxy of legs I-IV: coxa: 2, 2, 2, 1; trochanter: 4, 4, 4, 4; femur: 10, 8, 5, 4; genu: 8, 6, 6,

5; tibia: 8, 7, 7, 7; tarsus of legs II-IV 17, 17, 17. All legs with pretarsi similar to those of

adult female.

533

Figure 8.3 - Gamasiphis paulista Castilho, Moraes and Narita, 2010. (Protonymph A-B) A. Protonymph, dorsal

idiosoma; B. Protonymph, ventral idiosoma; C. Larva, dorsal idiosoma; D. Larva, ventral idiosoma.

Pores and lyrifissures enlarged to allow their differentiation

534

Larva (Fig. 8.3 C-D) (3 specimens measured): All setae of gnathosoma, idiosoma and legs

slender.


tooth and a setiform pilus dentilis; movable cheliceral digit 35 (34-36) long, with four teeth in

addition to apical tooth. Epistome as in female. Measurements of setae as in Table 8.1.

Dorsal idiosoma (Fig. 8.3C): Podonotal and opisthonotal shields very lightly sclerotised,

separated. Podonotal shield smooth, with nine pairs of setae (j1, j3, j4, j5, j6, z2, z4, z5 and s4)

and four pairs of lyrifissures. Unclerotised cuticle laterad of podonotal shield with a pair of

lyrifissures and a pair of pore-like structures (posterolaterad of s4). Opisthonotal region

smooth; with six pairs of setae (J4, J5, Z3-Z5 and S4) and with a pair of lyrifissures.

Unsclerotised cuticle laterad of opisthonotal shield with a pair of seta (S3). Measurements of


Ventral idiosoma (Fig. 8.3D): Trapezoidal base of tritosternum 19 (18-20) long and 13 (13-

14) wide proximally; laciniae 42 (41-42), otherwise as in adult female. Only anal shield

distinguishable. Seven pairs of setae (st1-st3, Jv1, Jv2, Jv5 and Zv2) and a pair of lyrifissures

on unsclerotised cuticle. Anal shield smooth; post-anal seta about 1.2 times as long as para-

anal seta, but proportionally less so than in later stages. Measurements of setae as in Table

8.2.

Legs: Lengths: I: 285 (280-290); II: 232 (223-241); III: 227 (218-236). Chaetotaxy of legs I-

III: coxa: 2, 2, 2; trochanter: 4, 4, 4; femur: 10, 7, 5; genu: 8, 6, 6; tibia: 8, 7, 7; tarsus: II-III

16, 16. All legs with pretarsi similar to those of adult female.

Material examined

All type specimens were obtained from a laboratory culture initiated with specimens

collected in July 2008 by J.P.Z. Narita, from litter and soil from the campus of Escola

Superior de Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de São Paulo (USP),

Piracicaba, State of São Paulo, Brazil (22°42‘30‖ S, 47°38‘00‖ W).

535

Holotype female, 6 paratype females, 6 paratype males, 3 paratype deutonymphs, 3 paratype

protonymphs and 3 paratype larvae deposited at Departamento de Entomologia e Acarologia

(ESALQ, USP), Piracicaba-SP, Brazil.

Etymology

The name paulista refers to the type locality of the species, the State of São Paulo.

Remarks

Adult females of Gamasiphis paulista Castilho, Moraes and Narita, 2010 are similar to

adult females of Gamasiphis hemicapillus Karg; however, the latter have 20 pairs of setae in

the podosomal region (s1, r1, r2 and r3 absent) and 12 pairs of setae in the opisthosomal

region (J2 and Z2 present) of the dorsal shield, dorsal shield setae setaceous, unsclerotised

line between dorsal and ventri-anal shields reaching base of seta Zv3 [Vx7of Hirschmann

(1957), adopted in Karg´s publications] and seta Jv5 [V8 of Hirschmann (1957)] much longer

than para-anal seta.

Key to Gamasiphis species from the Neotropical Region (females)

The following key was prepared based on available descriptions and redescriptions of

mites of this genus known from the Neotropical region. Gamasiphis trituberosus Karg, 1990,

described from Saint Lucia, was not included because of insufficient details given for females

in the original description of the species, in which a male was designated as holotype.

1. With one pair of presternal shields................ Gamasiphis illotus Fox, 1949; Puerto Rico

- With two pairs of presternal shields ................................................................................. 2

2. Peritreme extending anteriorly at most to anterior margin of coxa II; central projection

of anterior margin of epistome distally expanded and denticulate .................................. 3

- Peritreme extending anteriorly to level of coxa I; central projection of anterior margin of

epistome aciculate and smooth ........................................................................................ 5

3. Seta Z4 about as long as seta Z5.......... Gamasiphis holocapillus Karg, 1990; Saint Lucia

- Seta Z4 shorter than seta Z5 ............................................................................................. 4

4. Central projection of anterior margin of epistome spatulate ..............................................

.....................................................................Gamasiphis furcatus Karg, 1990; Saint Lucia

- Central projection of anterior margin of epistome lanceolate.............................................

. .............................................. Gamasiphis plenosetosus Karg, 1994b; Galapagos Islands

536

5. At least Z5 and post-anal setae blunt or distally expanded.............................................. 6

- All idiosomal setae with sharp tips ................................................................................ 10

6. Ventri-anal shield with seven pairs of setae in addition to circumanal setae ....................

..................................................................Gamasiphis mediosetosus Karg, 2003; Ecuador

- Ventri-anal shield with eight pairs of setae in addition to circumanal setae ................... 7

7. Ventri-anal shield smooth; unsclerotised line between dorsal and ventri-anal shields

barely surpassing base of seta Zv3 ................. Gamasiphis pinnatus Karg, 1998; Ecuador

- Ventri-anal shield ornamented; unsclerotised line between dorsal and ventri-anal shields

reaching region between bases of setae Jv3 and Jv4 [V3 and V7 of Hirschmann (1957)]..

. ......................................................................................................................................... 8

8. Sternal and ventri-anal shields reticulate; most of the longer dorsal shield setae

distinctly expanded distally ........................... Gamasiphis silvestris Karg, 2007; Ecuador

- Sternal shield with sparse diagonal striae; ventri-anal shield transversely striated; dorsal

shield setae either slightly expanded distally or aciculate ............................................... 9

9. Anterior-most transverse striae of ventri-anal shield roughly straight; most of the longer

dorsal shield setae slightly expanded distally......................................................................

................................................................... Gamasiphis hamatellus Karg, 1998; Ecuador

- Anterior-most transverse striae of ventri-anal shield distinctly curved backward; except

for Z5, dorsal shield setae aciculate ...................................................................................

..........................................Gamasiphis undulatus Karg and Schorlemmer, 2009; Ecuador

10. Post-anal seta about as long as para-anal setae; dorsal shield reticulate; opisthonotal

region with two pairs of setae much longer than others......................................................

. ............................................................ Gamasiphis quadruplicis Karg, 1990; Saint Lucia

- Post-anal seta more than twice as long as para-anal setae; dorsal shield reticulate or

smooth; opisthonotal region with one or two pairs of setae much longer than others .. 11

11. Dorsal and ventri-anal shields extensively reticulate; seta j3 at most half as long as

distance between their bases and bases of the subsequent setae .......................................

....................................................................Gamasiphis adanalis Karg, 1990; Saint Lucia

- Dorsal shield smooth; ventri-anal shield only with transverse striate or with transverse

striae and a few diagonal striae connecting them; seta j3 at least two-thirds as long as

distance between its base and the base of j4 .................................................................. 12

12. Opisthonotal region with two pairs of setae (including Z5) much longer than others........

. ................................................................... Gamasiphis hyalinus Karg, 2003; Costa Rica

- Opisthonotal region with a single pair of setae (Z5) much longer than others .............. 13

537

13. Pre-anal setae distinctly anterior to anterior margin of anal opening ................................

......................................................................Gamasiphis pinguis Karg, 1990; Saint Lucia

- Posterior-most pre-anal seta (Jv5) about in level with anterior margin of anal opening14

14. Seta Jv5 about 1.1 times as long as para-anal seta. ............................................................

………............................Gamasiphis paulista Castilho, Moraes and Narita, 2010; Brazil

- Seta Jv5 at least twice as long as para-anal seta............................................................. 15

15. Unsclerotised line between dorsal and ventri-anal shields reaching base of seta Zv3; seta

Jv5 distinctly shorter than post-anal seta ...........................................................................

..............................................................Gamasiphis hemicapillus Karg, 1990; Saint Lucia

- Unsclerotised line between dorsal and ventri-anal shields reaching base of seta Jv3; seta

Jv5 as long as or longer than post-anal seta ................................................................... 16

16. Idiosomal setae aciculate; four pairs of J setae; seta Jv5 about twice as long as post-anal

seta............................................................... Gamasiphis decoris Karg, 1990; Saint Lucia

- Many dorsal shield setae as well Jv5 and post-anal seta lanceolate, with constricted

base; three pairs of J setae; seta Jv5 about 1.2 times as long as post-anal seta...................

. ........................................................ Gamasiphis vinculi Karg, 1994a; Galapagos Islands

References

BAKER, E.W.; WHARTON, G.W. An Introduction to Acarology. New York: The







BREGETOVA, N.G. (Eds.). Key to the soil-inhabiting mites Mesostigmata. Leningrad:

Nauka, 1977. p. 308-314. [in Russian]

538

CASTILHO, R.C.; SILVA, E.S.; MORAES, G.J. de. Mites GSDs. In: BISBY, F.A.;

ROSKOV, Y.R.; ORRELL, T.M.; NICOLSON, D.; PAGLINAWAN, L.E.; BAILLY, N.;

KIRK, P.M.; BOURGOIN, T.; BAILLARGEON, G. (Eds.). Species 2000 & ITIS Catalogue

of Life: 2009 Checklist. Reading: Species 2000, 2009a. Available from:

http://www.catalogueoflife.org. Date of access: 11 Nov. 2009.



Rhodacaridae). Biological Control, Orlando, v. 48, p. 164-167, 2009b.


1, p. 37-40, 1949.





Geographie der Tiere, Jena, v. 120, p. 169-188, 1993.






n. 2, p. 179-216, 1994b.




http://www.catalogueoflife.org/

539




2003.









219, 1970.

LEE, D.C. Rhodacaridae (Acari: Mesostigmata) from near Adelaide, Australian. III.

Behaviour and development. Acarologia, Paris, v. 16, p. 21–44, 1974.







Press, 2009. p. 124-232.






540




541

9 THREE NEW SPECIES OF Gamasiphis (ACARI: MESOSTIGMATA:

OLOGAMASIDAE) FROM BRAZIL, WITH COMPLEMENTARY INFORMATION

ABOUT Gamasiphis plenosetosus KARG AND A KEY TO THE WORLD SPECIES OF

THE GENUS

Abstract

Gamasiphis Berlese is one of the most diverse genera of Ologamasidae, with 68

species, corresponding to about 15% of the species of the family. Until now, a single species

of this genus was known from Brazil. Gamasiphis n. sp. 1, Gamasiphis n. sp. 2 and

Gamasiphis n. sp. 3 are described based on the morphology of adult females and males

collected from litter and soil in Piracicaba, São Paulo State, Brazil. The holotype of

Gamasiphis plenosetosus Karg, 1994 was examined, given its close similarity to the latter

species, and complementary morphological information about it is provided. A key for the

separation of females of the 60 recognizable world species of Gamasiphis is provided.

Keywords: Edaphic mites; Litter; Rhodacaroidea; Taxonomy

Resumo

Gamasiphis Berlese é um dos gêneros mais diversos de Ologamasidae, com 68

espécies, correspondendo a cerca de 15% das espécies da família. Até agora, uma única

espécie deste gênero era conhecida do Brasil. Gamasiphis n. sp. 1, Gamasiphis n. sp. 2 e

Gamasiphis n. sp. 3 são descritas com base na morfologia de fêmeas e machos adultos

coletados de folhedo e solo em Piracicaba, Estado de São Paulo. O holótipo de Gamasiphis

plenosetosus Karg, 1994 foi examinado devido à sua semelhança com a última espécie, e

informações morfológicas complementares foram fornecidas. Uma chave para a separação de

fêmeas adultas de 60 espécies conhecidas de Gamasiphis do foi fornecida.

Palavras-chave: Ácaros edáficos; Folhedo; Rhodacaroidea; Taxonomia

542

9. 1 Introduction

Ologamasidae Ryke (Mesostigmata: Rhodacaroidea) is a large and widely distributed

group of predatory mites encountered in soil, humus and compost (LINDQUIST; KRANTZ;

WALTER, 2009). These are some of the most common Mesostigmata in the soils of São

Paulo State, Brazil (MINEIRO; MORAES, 2001; SILVA; MORAES; KRANTZ, 2004), from

where 11 species have been described, namely Gamasiphis paulista Castilho, Moraes and

Narita, 2010; Gamasiphoides acanthioides Karg and Schorlemmer, 2011; Hydrogamasellus

iaculi Karg and Schorlemmer, 2009; Hydrogamasellus ubatubaensis (Hirschmann, 1966);

Neogamasellevans longipes Karg and Schorlemmer, 2009; Ologamasus brevidigitus Karg and

Schorlemmer, 2009; Ologamasus cananeiae Silva, Moraes and Krantz, 2007; Ologamasus

postpilus Karg and Schorlemmer, 2009; Ologamasus simplicitus Karg and Schorlemmer,

2009; Ologamasus trituberculatus Karg and Schorlemmer, 2009; Rykellus brevipellitus Karg

and Schorlemmer, 2009. In contrast, only two ologamasid species were described from the

remaining of the country, Ologamasus aberrans (Berlese, 1888) from Mato Grosso (a large

state now divided in Mato Grosso and Mato Grosso do Sul) and Periseius brasiliensis

Hirschmann, 1966 from Pernambuco State.

At this stage, it is difficult to estimate the diversity of Ologamasidae in Brazil, given

the relatively small effort that has been dedicated to its knowledge in the country. Ten species

were described from Brazil in the last five years, and certainly many more wait to be

described. Gamasiphis Berlese is one of the most diverse genera of Ologamasidae, with 68

described species, corresponding to about 15% of the species of the family (CASTILHO;

SILVA; MORAES, 2010). However, a single species of this genus, G. paulista, was

described from Brazil (CASTILHO; MORAES; NARITA, 2010). The objective of this paper

is to provide the description of three new species of Gamasiphis from Brazil, complementary

morphological information of Gamasiphis plenosetosus Karg, 1994, given its similarity with

one of the species described, and a key to the world species of this genus.


Soil and litter samples were collected from the campus of Escola Superior de

Agricultura ―Luiz de Queiroz‖ (ESALQ), Universidade de São Paulo (USP), Piracicaba, São

Paulo State, Brazil, and taken to a laboratory where mites were extracted using a Berlese

funnel. Mesostigmatids were mounted in Hoyer‘s medium and later separated into families.

543

Ologamasids were separated into morphospecies and examined under a phase contrast

microscope.

The specimens of Gamasiphis collected were determined to correspond to three new

species. Taxonomically relevant structures of these mites were illustrated with the use of a

camera lucida and measured with the use of a graded ocular. Descriptions of the new species

are here provided; setal nomenclature is based on Silva; Moraes and Krantz (2007) and

Castilho; Moraes and Narita (2010a). For each structure, the average measurement is given in

micrometres, followed (in parentheses) by the respective range (for variable measurements).

The key provided for the separation of the world species of Gamasiphis was prepared

based on the original descriptions and available redescriptions of the concerned species, as

well as on the examination of the holotypes of Gamasiphis adanalis Karg, 1990, Gamasiphis

anguis Karg, 1993, Gamasiphis appendicularis Karg, 1993, Gamasiphis ardor Karg, 1993,

Gamasiphis brevigenitalis Karg, 1993, Gamasiphis coniunctus Karg, 1995 and Gamasiphis

decoris Karg, 1990. In addition, we also examined pictures of particular structures of type

specimens of Gamasiphis elegantellus Berlese, 1910a, Gamasiphis elongatellus Berlese,

1910b, Gamasiphis pilosellus Berlese, 1913 and Gamasiphis productellus Berles, 1923,

provided by Istituto Sperimentale per la Zoologia Agraria, Florence, Italy.

9. 3 Results

Gamasiphis Berlese

Gamasiphis Berlese 1904: 261. Type species: Gamasus pulchellus Berlese, 1887, by

monoypy.

Gamasiphis.— Berlese (1906): 101; Berlese (1914): 137; Bregetova (1977): 308; Lee (1970):

42; Karg (1990): 321; Karg (1993): 169.

Ologamasellus (Micriphis) Berlese, 1914: 140. Type species: Gamasiphis gamasellus

Berlese, 1913, by monotypy. Sinonymy by Lee (1970).

Ologamasus (Micriphis).— Baker and Wharton (1952): 73.

Micriphis.— Ryke (1962): 160.

Gamasiphis (Heteroiphis) Tragårdh, 1952: 55. Type species: Gamasiphis (Heteroiphis)

arcuatus Tragårdh, 1952, by original designation. Sinonymy by Lee (1970).

Heteroiphis.— Ryke (1962): 160; Bhattacharyya (1968): 530.

Neogamasiphis Tragårdh, 1952: 57. Type species: Neogamasiphis hamifer Tragårdh, 1952, by

original designation. Sinonymy by Lee (1970).

544

Diagnosis of adults: Epistome of species of this genus with three anterior extensions, the

antero-medial aciculate, spatulate or lanceolate, smooth or serrate, usually longer than the

others (except Gamasiphis elegantellus Berlese, 1910a, Gamasiphis elongatellus Berlese,

1910b and Gamasiphis pilosellus Berlese, 1913, which have the three exensions of similar

lenghts). Arthrodial process of chelicera in the form of a short coronet-like fringe. Dorsum of

idiosoma totally covered by a large shield, extending latero-ventrally, abutting the peritremal

shield and fusing with ventrianal shield posteriorly. Endopodal shield totally fused to sternal

shield, composing a single unit which extends to level of posterior margin of coxa IV and

which has a deep posterior concavity fitting the tongue-shaped genital shield. Exopodal shield

distinctly represented by a subtriangular section between coxae I-II (not always clearly

distinct) and an elongated section running from middle of coxa II to middle of coxa IV, the

latter separated from the peritremal shield by a line of unsclerotized cuticle; fused peritremal

shield and section of exopodal shield around posterior half of coxa IV extending posteriorly

well beyond stigma, wider immediately behind coxa IV and progressively narrowing

posteriorly to a pointed tip; separated from adjacent shields by lines of unsclerotized cuticle.

Gamasiphis n. sp. 1

Diagnosis of adults: Antero-medial extension of epistome aciculate; larger idiosomal setae

(j2-j6, z4-z6, s2, s4, s5, Z5, Jv5 and post-anal) slightly expanded distally; seta j2 posterior and

slightly laterad to j1; seta j4 about as long as distance between its base and base of j5; seta z6

about as long as j6; seta s6 about 0.2 times as long as j6; seta j6 about 0.9 times as long as

distance between its base and base of J3; three pairs of J setae; two pairs of presternal shields;

sternal shield with four pairs of lyrifissures; ventrianal shield with eight pairs of setae in

addition to circumanal setae (Jv1-Jv5, Zv1-Zv3); seta Zv2 about 0.8 times as long as distance

between its base and base of Zv3.

Adult female (Fig. 9.1 A–F) (five specimens measured). Setae j2-j6, z4-z6, s2, s4, s5, Z5, Jv5

and post-anal slightly expanded distally; other setae aciculate.

Gnathosoma: Fixed cheliceral digit 67 (63-67) long, with six teeth in addition to apical tooth

and a setiform pilus dentilis (Fig. 9.1A); movable cheliceral digit 66 (65-68) long, with four

545

Figure 9.1 - Gamasiphis n. sp. 1. Female. A. Lateral (antiaxial) view of chelicera; B. Epistome; C. Hypostome;

D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures enlarged for improved

visibility

546

teeth in addition to apical tooth. Antiaxial lyrifissure and dorsal cheliceral seta distinct.

Antero-medial extension of epistome smooth and aciculate; antero-lateral extensions smooth,

aciculate and shorter than antero-medial extension; with two denticles between antero-medial

and each antero-lateral extension (Fig. 9.1B). Margins of deutosternum not distinct;

deutosternal denticles in eight rows, with 7-13 denticles each (Fig. 9.1C); anterior-most row

shaped as an inverted ―V‖ while subsequent rows roughly transverse. Seta h3 postero-mediad

to h1 and slightly anterior and mediad to h2. Measurements of setae: h1 35 (34-36), h2 30

(30-31), h3 40 (39-41), sc 32 (31-33).

Dorsal idiosoma (Fig. 9.1D): Idiosoma 569 (542-598) long and 506 (464-551) wide at widest

point; dorsal shield smooth. Podonotal region with 22 pairs of setae (s1 and r1 absent) and

nine pairs of distinguishable lyrifissures; seta j2 posterior and slightly laterad to j1; seta j4

about as long as distance between its base and base of j5; seta j6 about 0.9 times as long as

distance between its base and base of J3. Opisthonotal region with 12 pairs of setae (J1, J2,

S5 and R1-R5 absent) and eight pairs of distinguishable lyrifissures. Measurements of setae:

j1 14 (13-17), j2 44 (42-49), j3 56 (54-58), j4 67 (66-67), j5 53 (52-55), j6 91 (89-92), z1 9 (8-

11), z2 12 (11-12), z3 12 (11-12), z4 66 (65-67), z5 74 (73-76), z6 99 (96-101), s2 61 (59-63),

s3 14 (13-16), s4 75 (74-76), s5 81 (80-83), s6 15 (14-16), r2 22 (21-23), r3 19 (19-20), r4 14

(13-14), r5 30 (29-32), r6 15 (15-16), J3 8 (7-8), J4 8 (7-8), J5 11 (10-11), Z1 11 (10-11), Z2

8 (7-8), Z3 11 (10-11), Z4 7 (6-7), Z5 115 (113-117), S1 7 (7-8), S2 7 (7-8), S3 8 (7-8), S4 8.



pilose. With two pairs of presternal shields. Sternal shield reticulate between st1 and st2,

smooth posteriorly; approximately 80 (79-82) long at mid-line and 177 (173-182) wide

between tips of endopodal projection between coxae II and III; with four pairs of setae, st3

inserted slightly posterior and mediad to st2, and four pairs of lyrifissures. Genital shield

smooth; sclerotized section shorter than width along straight posterior margin, which is about

in line with posterior margin of coxa IV; distance between st5-st5 65 (63-67). Ventrianal

shield transversely striate anteriorly to Zv2 and smooth posteriorly; 279 (261-297) long at

mid-line (from anterior margin to post-anal seta), 278 (265-286) wide at widest point; with

eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to circumanal setae and three pairs of

lyrifissures; seta Zv2 about 0.8 times as long as distance between its base and base of Zv3;

anal region of the shield partially separated from ventral region by an unsclerotized line

547

(except for area between Jv3 and slightly laterad to it) that runs ventro-laterally forward to the

region next to S2; seta Jv5 about in level with posterior margin of anal opening and about four

times as long as para-anal seta; post-anal seta about five times as long as para-anal seta. A

narrow band of dorsal shield extending laterad and up to the posterior end of the fused

peritremal-exopodal shield. A narrow diagonal section of sclerotized cuticle laterad to

ventrianal shield connects the latter to the dorsal shield and bears a lyrifissure. Peritreme

extending anteriorly to mid-level of coxa I. Measurements of setae: st1 39 (37-41), st2 34 (33-

35), st3 27 (26-28), st4 30 (29-30), st5 21 (20-22), Jv1 29 (28-30), Jv2 24 (23-24), Jv3 23 (23-

24), Jv4 36 (35-37), Jv5 87 (83-91), Zv1 30 (29-31), Zv2 35 (33-36), Zv3 30 (29-32), para-

anal 21 (21-22), post-anal 102 (97-107).


Numbers of setae on segments of legs I-IV: coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13,

11, 6, 6; genu: 13, 11, 9, 8; tibia: 14, 10, 8, 9; tarsus II-IV: 18, 18, 17. All legs with pretarsi,

each with three rounded pulvillar lobes, elongate ambulacral stalk and a pair of strongly

sclerotized claws.

Adult male (Fig. 9.2 A-C) (five specimens measured). Shape of setae as in adult female,

except for two ventral setae on each of femur II and genu II and one ventral seta on tibia II,

spur-like (Fig. 9.2A).


tooth (Fig. 9.2B); pilus dentilis not distinguishable; movable cheliceral digit 52 (49–53) long,

with one tooth in addition to apical tooth. Antiaxial lyrifissure not distinct, but dorsal

cheliceral seta distinct. Spermatodactyl 68 (66-71) long, c-shaped, apparently with an internal

canal in proximal half and with distal half spatulated. Epistome, deutosternum and position of

hypostomal setae as in adult female. Measurements of setae: h1 33 (32-34), h2 28 (28-29), h3

35 (35-36), sc 30 (29-32).

Dorsal idiosoma: Idiosoma 458 (443-475) long and 369 (352-381) wide at widest point;

dorsal shield similar to that of adult female. Measurements of setae: j1 11 (11-12), j2 43 (41-

46), j3 40 (39-41), j4 65 (63-66), j5 45 (44-47), j6 84 (78-90), z1 7 (7-8), z2 11 (10-12), z3 11

(11-12), z4 62 (60-64), z5 69 (67-71), z6 89 (86-92), s2 55 (53-57), s3 11 (11-12), s4 69 (67-

71), s5 76 (74-78), s6 13 (12-14), r2 16 (15-18), r3 15 (14-16), r4 9 (8-11), r5 23 (22-25), r6

548

13 (12-13), J3 7 (6-7), J4 7 (6-8), J5 8 (7-9), Z1 9 (9-10), Z2 7 (7-8), Z3 11 (11-12), Z4 6 (6-

7), Z5 97 (93-101), S1 7 (6-7), S2 7 (7-8), S3 8 (7-8), S4 8 (7-8).

Figure 9.2 - Gamasiphis n. sp. 1. Male. A. Anterolateral view of femur, genu and tibia of leg II; B. Lateral

(antiaxial) view of chelicerae, with detail of spermatodactyl; C. Ventral idiosoma. Lyrifissures


Ventral idiosoma (Fig. 9.2C): Base of tritosternum 20 (18-21) long and 13 (12-15) wide

proximally; laciniae 67 (61-78), otherwise as in adult female. Except for the fusion of sternal

and genital shields (sternogenital shield), shape, pattern and fusions of ventral shields as in

adult female. With two pairs of presternal shields. Sternogenital shield reticulate between st1

and st2, smooth posteriorly; approximately 154 (152-156) long and 155 (150-161) wide

between tips of endopodal projections between coxae II and III; posterior margin slightly

concave; with five pairs of setae and four pairs of lyrifissures; genital opening on anterior

margin of shield. Ventrianal shield 220 (209-228) long at mid-line (from anterior margin to

post-anal seta) and 241 (237-246) wide at widest point, with eight pairs of setae (Jv1-Jv5,

Zv1-Zv3) in addition to circumanal setae and three pairs of lyrifissures; seta Jv5 about in level

with anterior margin of anal opening and about four times as long as para-anal seta; post-anal

seta about 4.5 to five times as long as para-anal seta. Measurements of setae: st1 29 (29-30),

549

st2 31 (30-31), st3 25 (24-26), st4 28 (27-29), st5 29 (28-31), Jv1 28 (26-30), Jv2 23 (22-24),

Jv3 21 (21-22), Jv4 32 (30-33), Jv5 80 (76-85), Zv1 27 (26-28), Zv2 32 (31-33), Zv3 26 (25-

28), para-anal 19 (18-19), post-anal 95 (88-101).


Numbers of setae of leg segments similar to those of adult female. All legs with pretarsi,

similar to those of adult female.

Material examined

All specimens collected by J.P.Z. Narita from soil and litter of a garden by ―Salvador

de Toledo Piza Junior‖ building, campus of Escola Superior de Agricultura ―Luiz de Queiroz‖

(ESALQ), Universidade de São Paulo (USP), Piracicaba, São Paulo State, Brazil (22º42‘30‖

S, 47º38‘00‖ W). Adult female holotype collected on October 7, 2008; three adult male

paratypes collected on June 15, 2009; three adult female paratypes and three adult male

paratypes collected on September 7, 2009; three adult female paratypes collected on

September 12, 2011; three adult female paratypes and three adult male paratypes collected on

October 14, 2011. All types deposited at Departamento de Entomologia e Acarologia,

ESALQ/USP.

Remarks

Gamasiphis n. sp. 1 is most similar to Gamasiphis hamatellus Karg, 1998, but the

latter has podonotal region with 17 pairs of setae (z2, s1, and r1-r5 absent); seta j4 about 0.8

times as long as distance between its base and base of j5; seta z6 about 0.2 times as long as j6;

seta s6 as long as j6; opisthonotal region with 10 pairs of setae (J1, J2, S1, S2, S5 and R1-R5

absent).

Gamasiphis n. sp. 2

Diagnosis of adults: Antero-medial extension of epistome aciculate; larger idiosomal setae

(j2-j6, z4-z6, s2, s4, s5, Z5, Jv5 and post-anal) slightly expanded distally; seta j2 laterad to j1;

seta j4 as long as distance between its base and base of j5; seta z6 about as long as j6; seta s6

about 0.3 times as long as j6; seta j6 about 0.6 times as long as distance between its base and

base of J2; four pairs of J setae; two pairs of presternal shields; sternal shield with four pairs

550

of lyrifissures; ventrianal shield with eight pairs of setae in addition to circumanal setae; seta

Zv2 as long as distance between its base and base of Zv3.

Adult female (Fig. 9.3 A–F) (five specimens measured): Setae j2-j6, z4-z6, s2, s4, s5, Z5, Jv5

and post-anal slightly expanded distally; other setae aciculate.


tooth and a setiform pilus dentilis (Fig. 9.3A); movable cheliceral digit 59 (57-61) long, with

four teeth in addition to apical tooth. Antiaxial lyrifissure and dorsal cheliceral seta distinct.

Antero-medial extension of epistome smooth and aciculate; antero-lateral extensions smooth,

aciculate and shorter than antero-medial extension (Fig. 9.3B). Margins of deutosternum not

distinct; deutosternal denticles in eight rows, with 7-14 denticles each (Fig. 9.3C); anterior-

most row shaped as an inverted ―V‖ while subsequent rows roughly transverse. Seta h3

posterior and about in longitudinal line with or slightly laterad to h1, and anterior and mediad

to h2. Measurements of setae: h1 31 (30-32), h2 21 (20-22), h3 31 (30-32), sc 27 (26-28).


point; dorsal shield smooth. Podonotal region with 21 pairs of setae (s1, r1 and r2 absent),

seven pairs of distinguishable lyrifissures and a pair of pores, the latter anterolaterad to z5;

seta j2 laterad to j1; seta j4 as long as distance between its base and base of j5; seta j6 about

0.6 times as long as distance between its base and base of J2. Opisthonotal region with 14

pairs of setae (J1 and R1-R5 absent), eight pairs of distinguishable lyrifissures and a pair of

pores, the latter antero-mediad to S2. Measurements of setae: j1 11 (10-11), j2 37 (36-38), j3

41 (41-42), j4 49 (47-52), j5 35 (33-37), j6 40 (37-43), z1 7 (6-8), z2 8 (7-8), z3 7 (6-7), z4 41

(39-43), z5 41 (40-43), z6 46 (45-47), s2 42 (41-43), s3 15 (14-16), s4 46 (45-47), s5 44 (40-

47), s6 12 (11-12), r3 8 (8-9), r4 6 (5-6), r5 15 (15-16), r6 7 (7-8), J2 12 (11-13), J3 11 (10-

11), J4 14 (13-14), J5 11 (11-12), Z1 15 (14-15), Z2 12 (11-12), Z3 11 (11-12), Z4 11 (10-11),

Z5 74 (73-75), S1 13 (12-14), S2 11 (10-11), S3 12 (12-13), S4 12 (11-12), S5 11 (11-12).





between tips of endopodal projections between coxae II and III; with four pairs of setae, st3

551


D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures and pores enlarged for

improved visibility

552

inserted slightly posterior and mediad to st2, and four pairs of lyrifissures. Genital shield

smooth; sclerotized section shorter than width along straight posterior margin, which is about

in line with posterior margin of coxa IV; distance between st5-st5 54 (51-56). Ventrianal

shield transversely striate anteriorly to Zv2 and smooth posteriorly; 220 (219-224) long at

mid-line (from anterior margin to post-anal seta), 230 (227-234) wide at widest point; with

eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to circumanal setae and three pairs of

lyrifissures; seta Zv2 as long as distance between its base and base of Zv3; anal region of the

shield partially separated from ventral region by an unsclerotized line (except for area

between Jv3 and slightly laterad to it) that runs ventro-laterally forward to the region next to

S3; seta Jv5 about slightly anterior to anterior margin of anal opening and about 3-4 times as

long as para-anal seta; post-anal seta about four times as long as para-anal seta. A narrow

band of dorsal shield extending laterad and up to the posterior end of the fused peritremal-

exopodal shield. A narrow diagonal section of sclerotized cuticle laterad to ventrianal shield

connects the latter to the dorsal shield. Peritreme extending anteriorly to posterior margin of

coxa I. Measurements of setae: st1 28 (27-29), st2 27 (26-27), st3 19 (18-20), st4 22 (21-22),

st5 22 (21-23), Jv1 29 (27-30), Jv2 30 (28-31), Jv3 25 (24-26), Jv4 39 (37-41), Jv5 62 (61-

64), Zv1 30 (29-31), Zv2 36 (34-37), Zv3 26 (24-27), para-anal 17 (16-19), post-anal 72 (71-

73).





sclerotized claws.

Adult male (Fig. 9.4 A-C) (five specimens measured): Shape of setae as in adult female,

except for two ventral setae on each of femur II and genu II and one ventral seta on tibia II,

spur-like (Fig. 9.4A).


tooth (Fig. 9.4B); pilus dentilis not distinguishable; movable cheliceral digit 47 (45-50) long,

with one tooth in addition to apical tooth. Antiaxial lyrifissure not distinct, but dorsal

cheliceral seta distinct. Spermatodactyl 59 (57-62) long, sigmoid, apparently with an internal

canal in proximal half and with distal half spatulated. Epistome, deutosternum and position of

553

hypostomal setae as in adult female. Measurements of setae: h1 30 (29-30), h2 19 (18-20), h3

31 (30-32), sc 27 (26-27).

Dorsal idiosoma: Idiosoma 406 (386-421) long and 305 (295-317) wide at widest point;

dorsal shield similar to that of adult female. Measurements of setae: j1 10 (9-10), j2 35 (35-

36), j3 42 (41-43), j4 48 (47-50), j5 35 (34-37), j6 36 (35-38), z1 7 (7-8), z2 8 (7-8), z3 8 (7-

8), z4 39 (38-41), z5 40 (39-41), z6 45 (44-46), s2 39 (37-40), s3 13 (12-13), s4 43 (42-44), s5

47 (45-48), s6 10 (10-11), r3 7 (7-8), r4 7 (7-8), r5 11 (11-12), r6 7 (7-8), J2 11 (11-12), J3

11 (10-12), J4 11 (10-11), J5 11 (10-12), Z1 11 (11-12), Z2 9 (9-10), Z3 9 (9-10), Z4 9 (9-10),

Z5 71 (69-73), S1 10 (10-11), S2 10 (9-11), S3 11 (10-12), S4 10 (10-11), S5 10 (10-11).




Ventral idiosoma (Fig. 9.4C): Base of tritosternum 15 (15-16) long and 12 (12-13) wide

proximally; laciniae 62 (59-65), otherwise as in adult female. Except for the fusion of sternal

and genital shields (sternogenital shield), shape, pattern and fusions of ventral shields as in

adult female. With two pairs of presternal shields. Sternogenital shield reticulate between st1

and st2, smooth posteriorly; approximately 123 (117-127) long and 132 (128-135) wide

554

between tips of endopodal projections between coxae II and III; posterior margin slightly

concave; with five pairs of setae and four pairs of lyrifissures; genital opening on anterior

margin of shield. Ventrianal shield 207 (201-216) long at mid-line (from anterior margin to

post-anal seta) and 216 (207-225) wide at widest point, with eight pairs of setae (Jv1-Jv5,

Zv1-Zv3) in addition to circumanal setae and three pairs of lyrifissures; seta Jv5 about in level

with posterior margin of anal opening and about 3-4 times as long as para-anal seta; post-anal

seta about 4-5 times as long as para-anal seta. Measurements of setae: st1 27 (27-28), st2 29

(29-30), st3 23 (21-25), st4 27 (26-28), st5 25 (23-27), Jv1 30 (29-30), Jv2 25 (24-27), Jv3 26

(25-26), Jv4 34 (33-36), Jv5 55 (53-57), Zv1 30 (30-31), Zv2 34 (33-35), Zv3 24 (23-26),

para-anal 15 (14-16), post-anal 70 (68-72).


Numbers of setae of leg segments similar to those of adult female. All legs with pretarsi,

similar to those of adult female.

Material examined



(ESALQ), Universidade de São Paulo (USP), Piracicaba, São Paulo State, Brazil (22º42‘30‖

S, 47º38‘00‖ W). Adult female holotype, one adult female paratype and two adult male

paratypes collected on October 14, 2011; one adult female paratype and one adult male

paratype collected on July 15, 2011; two adult female paratypes and three adult male

paratypes collected on August 21, 2011. All types deposited at Departamento de Entomologia

e Acarologia, ESALQ/USP.

Remarks

Gamasiphis n. sp. 2 is most similar to Gamasiphis lanceolatus Karg, 1987, but the

latter has podonotal region with 22 pairs of setae (s1 and r1 absent); seta j2 posterior to j1;

seta j4 about 1.2 times as long as distance between its base and base of j5; opisthonotal region

with 13 pairs of setae (J1, S5 and R1-R5 absent); peritreme extending anteriorly beyond coxa

I. It is also similar to Gamasiphis n. sp. 1 but the latter has podonotal region with 22 pairs of

setae (s1 and r1 absent); seta j2 posterior to j1; opisthonotal region with 12 pairs of setae (J1,

J2, S5 and R1-R5 absent); seta Zv2 about 0.8 times as long as distance between its base and

base of Zv3.

555

Gamasiphis n. sp. 3

Diagnosis of adults: Antero-medial extension of epistome lanceolate and serrate; all

idiosomal setae aciculate; seta j2 about as long as distance between its base and base of j3;

seta j3 about 0.8 times as long as distance between its base and base of j4; five pairs of J

setae; seta J2 as long as distance between its base and base of J3; seta J4 about 0.5 times as

long as distance between its base and base of J5; seta Z4 slightly mediad to Z5 and about 1.3

times as long as distance between its base and base of Z5; two pairs of presternal shields;

sternal shield with four pairs of lyrifissures; seta Zv1 about 1.3 times as long as Jv1; seta Zv2

as long as distance between its base and base of Zv3.

Adult female (Fig. 9.5 A–F) (five specimens measured): All setae aciculate.

Gnathosoma: Fixed cheliceral digit 45 (44-47) long, with six teeth in addition to apical tooth

and a setiform pilus dentilis (Fig. 9.5A); movable cheliceral digit 44 (42-45) long, with three

teeth in addition to apical tooth. Antiaxial lyrifissure and dorsal cheliceral seta distinct.

Antero-medial extension of epistome lanceolate and serrate; antero-lateral extensions smooth,

aciculate and shorter than antero-medial extension (Fig. 9.5B). Margins of deutosternum not

distinct; deutosternal denticles in eight rows, with 8-15 denticles each (Fig. 9.5C); anterior-

most row shaped as an inverted ―V‖ while subsequent rows roughly transverse. Seta h3

directly posterior to h1 and slightly anterior and mediad to h2. Measurements of setae: h1 14

(13-15), h2 11 (11-12), h3 14 (13-14), sc 13 (13-14).


point. Podonotal region reticulate postero-laterally and immediately behind j6, smooth

elsewhere; with 23 pairs of setae (r1 absent) and ten pairs of distinguishable lyrifissures; seta

j2 about as long as distance between its base and base of j3; seta j3 about 0.8 times as long as

distance between its base and base of j4. Opisthonotal region imbricate; with 18 pairs of setae

(S5 and R5 absent), 13 pairs of distinguishable lyrifissures and one pair of pores, the latter

slightly posterior and mediad to Z1; seta J2 as long as distance between its base and base of

J3; seta J4 about 0.5 times as long as distance between its base and base of J5; seta Z4 slightly

mediad to Z5 and about 1.3 times as long as distance between its base and base of Z5.

556


D. Dorsal idiosoma; E. Ventral idiosoma; F. Tritosternum. Lyrifissures and pores enlarged for

improved visibility

557


28), j6 26 (25-27), z1 8 (7-8), z2 24 (23-25), z3 29 (27-30), z4 30 (30-31), z5 29 (27-30), z6 32

(32-33), s1 22 (21-22), s2 22 (18-25), s3 26 (26-27), s4 30 (29-32), s5 32 (31-33), s6 35 (34-

36), r2 25 (23-26), r3 22 (20-23), r4 31 (30-32), r5 15 (13-16), r6 34 (32-35), J1 29 (28-30),

J2 35 (34-37), J3 35 (34-36), J4 26 (25-27), J5 21 (20-22), Z1 36 (34-37), Z2 37 (35-38), Z3

38 (37-40), Z4 30 (28-32), Z5 46 (45-46), S1 35 (35-36), S2 36 (35-38), S3 32 (32-33), S4 29

(28-30), R1 35 (33-37), R2 37 (36-37), R3 37 (36-38), R4 37 (36-38).





between tips of endopodal projections between coxae II and III; with four pairs of setae, st3

inserted well posterior and mediad to st2, and four pairs of lyrifissures. Genital shield mostly

smooth, with a single curved line delimiting each posterior corner; sclerotized section shorter

than width along straight posterior margin, which is about in line with posterior margin of

coxa IV; distance between st5-st5 34 (32-37). Ventrianal shield reticulate anteriorly to Zv3

and smooth posteriorly; 182 (178-186) long at mid-line (from anterior margin to post-anal

seta), 172 (169-174) wide at widest point; with eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in

addition to circumanal setae and three pairs of lyrifissures; seta Zv2 as long as distance

between its base and base of Zv3; anal region of the shield partially separated from ventral

region by an unsclerotized line (except for area between Jv3 and slightly laterad to it) that

runs diagonally to dorsum of idiosoma, reaching region next and posterior to Z1; seta Jv5

about in level with anterior margin of anal opening and about 2.5-3 times as long as para-anal

seta; post-anal seta about 1.5 times as long as para-anal seta. A narrow band of dorsal shield

extending laterad and up to the posterior end of the fused peritremal-exopodal shield.

Diagonal section of sclerotized cuticle laterad to ventrianal shield and connecting it to the

dorsal shield wide. Peritreme extending anteriorly almost to anterior margin of coxa II.

Measurements of setae: st1 19 (18-21), st2 15 (14-16), st3 11 (10-11), st4 17 (16-17), st5 15

(14-15), Jv1 19 (19-20), Jv2 21 (21-22), Jv3 26 (25-26), Jv4 36 (35-37), Jv5 27 (26-27), Zv1

26 (26-27), Zv2 31 (30-33), Zv3 35 (35-37), para-anal 10 (10-11), post-anal 15 (14-15).



558



sclerotized claws.

Adult male (Fig. 9.6 A–C) (two specimens measured): Shape of setae as in adult female,

except two ventral setae on femur II, one ventral seta on genu II and one ventral seta on tibia

II, spur-like (Fig. 9.6A).

Gnathosoma: Fixed cheliceral digit 38-42 long, with five teeth in addition to apical tooth and

a setiform pilus dentilis (Fig. 9.6B). Movable cheliceral digit 37-41 long, with one tooth in

addition to apical tooth. Antiaxial lyrifissure not distinct, but dorsal cheliceral seta distinct.

Spermatodactyl 44-50 long, c-chaped, apparently with an internal canal in proximal half and

with distal half spatulated. Epistome, deutosternum and position of hypostomal setae as in

adult female. Measurements of setae: h1 17, h2 14, h3 16, sc 16-17.

Dorsal idiosoma: Idiosoma 374-376 long and 257-284 wide at widest point; dorsal shield

similar to that of adult female. Measurements of setae: j1 10-11, j2 27, j3 26-28, j4 28-29, j5

23-24, j6 23-25, z1 6-7, z2 26-27, z3 27, z4 27-29, z5 25, z6 29-30, s1 21-22, s2 22-23, s3 25-

26, s4 27-30, s5 28-31, s6 29-32, r2 23-24, r3 19-21, r4 31-33, r5 12-13, r6 31-32, J1 26-27,

J2 30-31, J3 30-31, J4 26-27, J5 18-19, Z1 31-32, Z2 32, Z3 29-31, Z4 22-23, S1 32, S2 32-

33, S3 30-31, S4 24-25, R2 31-32, R3 34-35, R4 35-36, R5 34-36.

Ventral idiosoma (Fig. 9.6C): Base of tritosternum 16-18 long and 12-13 wide proximally,

laciniae 46-51 long, otherwise as in adult female. Except for the fusion of sternal and genital

shields (sternogenital shield), shape, pattern and fusions of ventral shields as in adult female.

With two pairs of presternal shields. Sternogenital shield reticulate between st1 and st2 and

with imbricated patches posteriorly to st2; 123-125 long and 114-116 between tips of

endopodal projections between coxae II and III; posterior margin slightly concave; with five

pairs of setae and four pairs of lyrifissures; genital opening on anterior margin of shield.

Ventrianal shield 169-172 long (from anterior margin to post-anal seta) and 159-167 wide at

widest point; with eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to circum-anal setae and

three pairs of lyrifissures; seta Jv5 about in level with anterior margin of anal opening and

about 2.5 times as long as para-anal seta; post-anal seta 1.3 times as long as para-anal seta.

Peritreme similar to adult female. Measurements of setae: st1 20-21, st2 19, st3 14-15, st4 14-

559

15, st5 15-16, Jv1 22-23, Jv2 24-25, Jv3 24, Jv4 32-33, Jv5 25-26, Zv1 26, Zv2 31-33, Zv3 31-

32, para-anal 10-11, post-anal 13.

Legs: Lengths: I: 331-336; II: 246-273; III: 207-217; IV: 279-291. Numbers of setae of legs

similar to adult female. All legs with pretarsi similar to those of adult female.




Material examined



(ESALQ), Universidade de São Paulo (USP), Piracicaba, State of São Paulo, Brazil

(22º42‘30‖ S, 47º38‘00‖ W). Adult female holotype, two adult female paratypes and one adult

male paratype collected on April 4, 2010; one adult female paratype collected on June 1,

2009; one adult female paratype collected on July 8, 2009; one adult male paratype collected

on November 16, 2009. All types deposited at Departamento de Entomologia e Acarologia,

ESALQ/USP.

560

Remarks

Gamasiphis n. sp. 3 is most similar to Gamasiphis plenosetosus Karg, 1994b, but the

latter has seta j4 about 0.6 times as long as distance between its base and base of j5; seta J2

about 1.8 times as long as distance between its base and base of J3; seta J4 as long as distance

between its base and base of J5; seta Z4 slightly laterad to Z5; seta Z4 about as long as

distance between its base and base of Z5. Gamasiphis furcatus Karg, 1990 differs from the

new species here described by having antero-medial extension of epistome spatulate;

podonotal region with 22 pairs of setae (s1 and r1 absent); seta j2 about 0.4 times as long as

distance between its base and base of j3; opisthonotal region with 19 pairs of setae (R5

absent); seta Zv2 about 0.8 times as long as distance between its base and base of Zv3. This

new species is also similar to Gamasiphis trituberosus Karg, 1990, but the latter has dorsal

shield totally imbricate; podonotal region with 21 pairs of setae (s1, r1 and r2 absent); seta j2

about 0.6 times as long as distance between its base and base of j3; opisthonotal region with

17 pairs of setae (S5, R3 and R5 absent); seta Zv1 about 0.7 times as long as Jv1; seta Zv2

about 0.7 times as long as distance between its base and base of Zv3.

Gamasiphis plenosetosus Karg

Gamasiphis plenosetosus Karg, 1994b: 210.

Diagnosis of adults: Antero-medial extension of epistome lanceolate and serrated; all

idiosomal setae aciculate; five pairs of J setae; seta J2 about 1.8 times as long as distance

between its base and base of J3; seta J4 as long as distance between its base and base of J5;

seta Z4 slightly laterad to Z5; seta Z4 about as long as distance between its base and base of

Z5; two pairs of presternal shields; sternal shield with four pairs of lyrifissures; seta Zv2 as

long as distance between its base and base of Zv3.

Adult female (holotype): All setae aciculate.


setiform pilus dentilis; movable cheliceral digit 42 long, with three teeth in addition to apical

tooth. Antiaxial lyrifissure and dorsal cheliceral seta not distinct. Antero-medial extension of

epistome lanceolate and serrated; antero-lateral extensions smooth, aciculate and shorter than

561

antero-medial extension. Deutosternum not distinguishable. Seta h3 directly posterior to h1

and slightly anterior and mediad to h2. Measurements of setae: h1 15, h2 12, h3 13, sc broken.

Dorsal idiosoma: Idiosoma 361 long and 284 wide at widest point. Podonotal region

reticulated postero-laterally and immediately behind j6, smooth elsewhere; with 23 pairs of

setae (r1 absent); lyrifissures and pores not distinguishable. Opisthonotal region imbricate;

with 18 pairs of setae (S5 and R5 absent); lyrifissures and pores not distinguishable; seta J2

about 1.8 times as long as distance between its base and base of J3; seta J4 as long as distance

between its base and base of J5; seta Z4 slightly laterad to Z5; seta Z4 about as long as

distance between its base and base of Z5. Measurements of setae: j1 broken, j2 broken, j3

broken, j4 broken, j5 broken, j6 27, z1 broken, z2 22, z3 broken, z4 28, z5 broken, z6 31, s1

broken, s2 broken, s3 25, s4 30, s5 31, s6 35, r2 23, r3 21, r4 29, r5 13, r6 33, J1 28, J2 33, J3

35, J4 33, J5 18, Z1 36, Z2 36, Z3 39, Z4 32, Z5 47, S1 36, S2 37, S3 33, S4 26, R1 36, R2 37,

R3 36, R4 35.

Ventral idiosoma: Tritosternum broken. With two pairs of presternal shields. Sternal shield

reticulate between st1 and st2, smooth posteriorly; approximately 75 long at mid-line and 112

wide between tips of endopodal projections between coxae II and III; with four pairs of setae,

st3 inserted well anterior and mediad to st2, and four pairs of lyrifissures. Genital shield

mostly smooth, with a single curved line delimiting each posterior corner; sclerotized section

shorter than width along straight posterior margin, which is slightly posterior to posterior

margin of coxa IV; distance between st5-st5 34. Ventrianal shield reticulate anteriorly to Zv3

and smooth posteriorly; 157 long at mid-line (from anterior margin to post-anal seta), 168

wide at widest point; with eight pairs of setae (Jv1-Jv5, Zv1-Zv3) in addition to circumanal

setae and with three pairs of lyrifissures; seta Zv2 as long as distance between its base and

base of Zv3; anal region of the shield partially separated from ventral region by an

unsclerotized line (except for area between Jv3 and slightly laterad to it) that runs diagonally

to dorsum of idiosoma, reaching region next and posterior to Z1; seta Jv5 about in level with

anterior margin of anal opening and about 2.6 times as long as para-anal seta. A narrow band

of dorsal shield extending laterad and up to the posterior end of the fused peritremal-exopodal

shield. Diagonal section of sclerotized cuticle laterad to ventrianal shield and connecting it to

the dorsal shield wide. Peritreme extending anteriorly almost to anterior margin of coxa II.

Measurements of setae: st1 21, st2 16, st3 9, st4 17, st5 15, Jv1 broken, Jv2 21, Jv3 25, Jv4

36, Jv5 29, Zv1 broken, Zv2 31, Zv3 36, para-anal 11, post-anal broken.

562

Legs: lengths: I: 282; II: 229; III: 195; IV: 261. Numbers of setae on segments of legs I-IV:

coxa: 2, 2, 2, 1; trochanter: 6, 5, 5, 5; femur: 13, 11, 6, 6; genu: 13, 11, 9, 8; tibia: 14, 10, 8, 9;

tarsus II-IV: 18, 18, 17. All legs with pretarsi, each with three rounded pulvillar lobes,

elongate ambulacral stalk and a pair of strongly sclerotized claws.

Material examined

Holotype female collected by W. Karg from litter, on roots and wood pieces in a

Miconia sp. (Melastomataceae) area, near Media Luna, Santa Cruz Island, Galapagos Islands,

on February 6, 1985. Holotype deposited at Arachnologischen Sammlung des Museums für

Naturkunde, Berlin, Germany.

Remarks

This species is known only from the adult female holotype, which is not illustrated in

this paper because of its poor condition. In the original description, Karg (1994b) neither

illustrated nor mentioned the presence of r6; in our interpretation this seta is present in the

holotype and thus the podonotal region has 23 instead of 22 pairs of setae. In addition, only

two pairs of lyrifissures (both on sternal shield) were shown in the illustration of the original

description; we could not see lyrifissures on the dorsum of the idiosoma either, because of the

poor condition of the holotype; however, it was possible to distinguish four pairs of

lyrifissures on the sternal shield and three pairs on the ventrianal shield. No information was

provided in the original description about the hypostome, tritosternum and leg setal counts.

The following measurements were provided in the original description: idiosoma 370 long,

220 wide, dosum podonotal setae 30–35 [except j1 (cited as i1) 15], dosum opisthonotal setae

30–40 [except J5 (cited as I5) 25 and Z5 53], st1 16, st2 13, st3 8, ventrianal setae 30 [except

Jv4 and Zv3 (cited as ―Kaudalrande‖) 40–43], length of legs I 310, II 240, III 200 and IV 270.

Key to world genera of Gamasiphis based on adult females

Sixty-eight species were mentioned by Castilho; Silva and Moraes (2010) in

Gamasiphis. Sufficient information is presently available in the literature to allow the

recognition of the females of 60 species, including the three new species here described. The

following key was elaborated to help the separation of those species.

Gamasiphis gamasellus Berlese, 1913 from Indonesia, Gamasiphis benoiti Loots,

1980 from Seychelles and Gamasiphis erinaceus Karg, 1993, Gamasiphis macrorbis Karg,

563

1993, Gamasiphis minoris Karg, 1996 and Gamasiphis superardor Karg, 1993 from New

Caledonia were not included in the key because they were described only based on adult

males. Gamasiphis uncifer Trägårdh, 1931 from Juan Fernandez Islands and Gamasiphis

illotus Fox, 1949 from Puerto Rico, described based on adult females, were also not included

because their descriptions are not sufficiently detailed to allow their separation from other

similar species. Gamasiphis elegantellus Berlese, 1910a, Gamasiphis elongatellus Berlese,

1910b and Gamasiphis pilosellus Berlese, 1913 from Indonesia were also not included

because the insufficient details provided in the corresponding original descripitions, and

because the poor conditions of the type specimens did not allow their appropriate

examination. In addition, the inclusion of Gamasiphis productellus Berlese, 1923 should be

considered as tentative; in the type specimen, the antero-median extension of the epistome

coincides with the margin of one of the chelicera, hampering a conclusive visualization of the

shape of the first structure; we assumed it to be aciculate, reasoning that it would be

distinguishable if distally expanded.

1. Without presternal shields ................. Gamasiphis ellipticus Karg, 1996, New Caledonia

- With 1-3 pairs of presternal shields ................................................................................. 2

2. With three pairs of presternal shields, arranged as the margins of a triangle ....................

.........................................................Gamasiphis holocapillus Karg, 1990, Lesser Antilles

- With 1-2 pairs of presternal shields ................................................................................. 3

3. With one pair of presternal shields................................................................................... 4

- With two pairs of presternal shields ............................................................................... 28

4. Antero-medial extension of epistome distally expanded ................................................. 5

- Antero-medial extension of epistome aciculate. ............................................................ 17

5. Seta Jv1 about 1.2 times as long as distance between its base and base of Jv2

...................................................................... Gamasiphis fornicatus Lee, 1970, Australia

- Seta Jv1 at most 0.7 times as long as distance between its base and base of Jv2 ............ 6

6. Seta st3 mediad and distinctly anterior to st2.....................................................................

......................................................Gamasiphis conciliator Berlese, 1916, New Caledonia

- Seta st3 mediad and in horizontal line or posterior to st2 ................................................ 7

7. Seta J4 at least 1.5 times as long as distance between its base and base of J5. ............... 8

- Seta J4 at most as long as distance between its base and base of J5 ............................. 10

564

8. Seta j6 about 0.8 times as long as distance between its base and base of J1; seta J1 as

long as distance between its base and base of J2................................................................

. ............................................................... Gamasiphis lenifornicatus Lee, 1973, Australia

- Seta j6 at least 1.2 times as long as distance between its base and base of J1; seta J1 at

least 1.7 times as long as distance between its base and base of J2 ................................ 9

9. Antero-medial extension of epistome club-shaped; seta j2 [indicated as s1 by Karg

(1993)] about 1.6 times as long as distance between its base and base of j3......................

................................................................Gamasiphis foliatus Karg, 1993, New Caledonia

- Antero-medial extension of epistome spatulate; seta j2 [unnamed seta between i1 and i2

of Karg (1995)] about five times as long as distance between its base and base of j3........

...........................................................Gamasiphis spinulosus Karg, 1995, New Caledonia

10. Seta j2 about six times as long as distance between its base and base of j3........................

. ....................................................Gamasiphis brevigenitalis Karg, 1993, New Caledonia

- Seta j2 at most as long as distance between its base and base of j3 .............................. 11

11. Seta Z5 at most as long as J5 ......................................................................................... 12

- Seta Z5 at least twice as long as J5 ................................................................................ 14

12. Seta Jv2 as long as distance between its base and base of Jv3............................................

. ........................................................................... Gamasiphis saccus Lee, 1973, Australia

- Seta Jv2 at most half as long as distance between its base and base of Jv3 .................. 13

13. Seta st3 directly posterior to st2 ........ Gamasiphis euincisus Karg, 1996, New Caledonia

- Seta st3 in horizontal line and mediad to st2 .....................................................................

................................................................Gamasiphis ovoides Karg, 1993, New Caledonia

14. Seta J1 at least as long as distance between its base and base of J2. ............................ 15

- Seta J1 at most 0.6 times as long as distance between its base and base of J2. ............ 16

15. Seta J1 about twice as long as distance between its base and base of J2; peritreme

extending anteriorly to mid-level of coxa II........................................................................

. .................................................. Gamasiphis arcuatus Trägårdh, 1952, French Polynesia

- Seta J1 about as long as distance between its base and base of J2; peritreme extending

anteriorly almost to posterior margin of coxa II ................................................................

.................................................................Gamasiphis anguis Karg, 1993, New Caledonia

16. Seta J4 about half as long as distance between its base and base of J5; with a short

latero-diagonal fissure running from level of Jv5 (V8 in Karg´s publications) to level of

S3 [same terminology of Karg (1993)]. .. Gamasiphis ardor Karg, 1993, New Caledonia

565

- Seta J4 about as long as distance between its base and base of J5; with a long latero-

diagonal fissure running from Jv5 to level of r6 [indicated as S7 by Karg (1997)]

................................................... Gamasiphis longiorsetosus Karg, 1997, New Caledonia

17. With a distinct line of fusion between podonotal and opisthonotal shields ................... 18

- Line of fusion between podonotal and opisthonotal shields not distinct ....................... 19

18. Setae Zv1 and Zv2 about as long as distance between their bases and bases of Zv2 and

Zv3, respectively ............................ Gamasiphis australicus Womersley, 1956, Australia

- Setae Zv1 and Zv2 about half as long as distance between their bases and bases of Zv2

and Zv3, respectively ........................ Gamasiphis gandensius Van Daele, 1975, Belgium

19. Opisthonotal region imbricate; seta J4 about twice as long as distance between its base

and base of J5 ................................................................................................................. 20

- Opisthonotal region smooth; seta J4 at most as long as distance between its base and

base of J5........................................................................................................................ 22

20. Seta j2 [indicated as s1 by Karg (1995)] about 0.7 times as long as distance between its

base and base of j3; seta j6 about half as long as distance between its base and base of

J1 .......................................................... Gamasiphis incudis Karg, 1993, New Caledonia

- Seta j2 at least twice as long as distance between its base and base of j3; seta j6 at least

1.2 times as long as distance between its base and base of J1 ....................................... 21

21. Antero-medial extension of epistome distally serrated; seta j5 [indicated as i4 by Karg

(1995)] about 0.6 times as long as distance between its base and base of j6; ventrianal

shield with transverse striae and a few diagonal striae connecting them ...........................

.................................................................. Gamasiphis caper Karg, 1995, New Caledonia

- Antero-medial extension of epistome smooth; seta j5 about as long as distance between

its base and base of j6; ventrianal shield imbricate.............................................................

. ........................................................ Gamasiphis coniunctus Karg, 1995, New Caledonia

22. Seta Z5 at most 1.2 times as long as J5. ......................................................................... 23

- Seta Z5 at least 2.7 times as long as J5 .......................................................................... 25

23. With a long fissure running antero-dorsally from level of Jv5 almost to the level of z6

[seta inserted laterally to seta indicated as i5 by Karg (1997)]; seta st3 mediad and

posterior to st2; peritreme extending anteriorly almost to anterior margin of coxa II........

. ....................................................... Gamasiphis longirimae Karg, 1997, New Caledonia

- With a short fissure running antero-dorsally from level of Jv5 (V8 in Karg´s

publications) to level of Z2 [seta inserted antero-laterally to seta indicated as J2 by Karg

566

(1993)]; seta st3 mediad to and in horizontal line with st2; peritreme extending

anteriorly almost to mid-level of coxa I ........................................................................ 24

24. Seta j5 [indicated as i4 by Karg (1993)] about half as long as distance between its base

and base of j6 [indicated as i5 by Karg (1993)]; ventrianal shield smooth.........................

................................................................ Gamasiphis incisus Karg, 1993, New Caledonia

- Seta j5 [indicated as i4 by Karg (1996)] about as long as distance between its base and

base of j6; ventrianal shield with transverse striae and a few diagonal striae connecting

them...................................................Gamasiphis eumagnus Karg, 1996, New Caledonia

25. Seta J4 about 1.2 times as long as distance between its base and base of J5......................

. ...................................................Gamasiphis appendicularis Karg 1993, New Caledonia

- Seta J4 at most 0.7 times as long as distance between its base and base of J5 ............. 26

26. Opisthonotal region without latero-diagonal fissure...........................................................

. .............................................................. Gamasiphis setosus Womersley, 1956, Australia

- Opisthonotal region with a latero-diagonal fissure running from Jv5 to level of Z2 [seta

immediately posterior to Z1 in Karg (1996)] ................................................................. 27

27. Seta j2 about twice as long as distance between its base and base of j3; seta Jv1 about

half as long as distance between its base and base of Jv2...................................................

. .............................................................. Gamasiphis flagelli Karg, 1993, New Caledonia

- Seta j2 about half as long as distance between its base and base of j3; seta Jv1 about 0.2

times as long as half the distance between its base and base of Jv2 ..................................

……………………………………Gamasiphis breviflagelli Karg, 1996, New Caledonia

28. Antero-medial extension of epistome distally expanded ............................................... 29

- Antero-medial extension of epistome aciculate ............................................................. 33

29. Seta J2 about 1.8 times as long as distance between its base and base of J3. ...................

.................................................Gamasiphis plenosetosus Karg, 1994b, Galapagos Islands

- Seta J2 at most as long as distance between its base and base of J3. ............................ 30

30. Seta j3 about 1.4 times as long as distance between its base and base of j4.......................

. ...........................................................Gamasiphis denticus Hafez and Nasr, 1979, Egypt

- Seta j3 at most 0.8 times as long as distance between its base and base of j4 .............. 31

31. Antero-medial extension of epistome laceolate; seta j2 about as long as distance

between its base and base of j3; seta Zv2 as long as distance between its base and base

of Zv3. ..................................................................................... Gamasiphis n. sp. 3, Brazil

567

- Antero-medial extension of epistome laceolate or spatulate; seta j2 at most 0.7 times as

long as distance between its base and base of j3; seta Zv2 0.8 times as long as distance

between its base and base of Zv3 ................................................................................... 32

32. Antero-medial extension of epistome spatulate; seta Zv1 slightly longer than distance

between its base and base of Zv2 .........Gamasiphis furcatus Karg, 1990, Lesser Antilles

- Antero-medial extension of epistome lanceolate; seta Zv1 about 0.8 times as long as

distance between its base and base of Zv2 .........................................................................

.........................................................Gamasiphis trituberosus Karg, 1990, Lesser Antilles

33. Distance between level of posterior-most pair of ventrianal setae (excluding circumanal

setae) and anterior margin of anus corresponding to about the length of the anal opening

........................................................................................................................................ 34

- Insertion of posterior-most pair of ventrianal setae (excluding circumanal setae) varying

from slightly anterior to posterior to anal opening......................................................... 37

34. Seta st3 mediad and in horizontal line to st2.................................................................. 35

- Seta st3 mediad and posterior to st2 ............................................................................... 36

35. Band of dorsal shield extending laterad to the fused peritremal-exopodal shield ending

broadly; seta Jv5 posterior to unsclerotized line that partially separates anal and ventral

regions ..................................................... Gamasiphis productellus Berlese, 1923, China

- Band of dorsal shield extending laterad to the fused peritremal-exopodal shield ending

sharply; seta Jv5 anterior to unsclerotized line that partially separates anal and ventral

regions. ............................................ Gamasiphis bengalensis Bhattacharyya, 1966, India

36. Seta j2 [indicated as s1 by Karg (1990)] laterad to j1; opisthonotal region smooth and

with a single pair of setae (Z5) much longer than others.....................................................

. .............................................................. Gamasiphis pinguis Karg, 1990, Lesser Antilles

- Seta j2 distinctly posterior to j1; opisthonotal region imbricate and with two pairs of

setae (including Z5) much longer than others ....................................................................

........................................................Gamasiphis quadruplicis Karg, 1990, Lesser Antilles

37. At least Z5 and post-anal setae blunt or distally expanded ............................................ 38

- All idiosomal setae with sharp tips ................................................................................ 47

38. With three pairs of J setae. ............................................................................................. 39

- With four pairs of J setae ............................................................................................... 45

39. Ventrianal shield with seven pairs of setae in addition to circumanal setae (Zv3 absent)...

. ............................................................... Gamasiphis mediosetosus Karg, 2003, Ecuador

- Ventrianal shield with eight pairs of setae in addition to circumanal setae ................... 40

568

40. Seta Jv5 about 1.4 times as long as post-anal seta. ........................................................ 41

- Seta Jv5 at most 0.9 times as long as post-anal seta ...................................................... 42

41. Seta j3 about as long as distance between its base and base of j4; ventrianal shield

smooth. .......................................................... Gamasiphis pinnatus Karg, 1998, Ecuador

- Seta j3 [unnamed seta between z1 and i3 of Karg and Schorlemmer (2009)] about twice

as long as distance between its base and base of j4 [indicated as i3 by Karg and

Schorlemmer (2009)]; ventrianal shield with transverse striae .........................................

..........................................Gamasiphis undulatus Karg and Schorlemmer, 2009, Ecuador

42. Many dorsal shield setae expanded distally .................................................................. 43

- Except for Z5, dorsal shield setae aciculate ................................................................... 44

43. Opisthonotal region with two pairs of setae (Z2 and Z5; same designations in Karg´s

publictions) much longer than others; ventrianal shield imbricate anteriorly to Jv3 and

smooth elsewhere. ......................................... Gamasiphis silvestris Karg, 2007, Ecuador

- Opisthonotal region with a single pair of setae (Z5) much longer than others; ventrianal

shield transversely striate anteriorly to Zv2 and smooth elsewhere....................................

. ................................................................................................ Gamasiphis n. sp. 1, Brazil

44. Seta Zv2 about 1.2 times as long as distance between its base and base of Zv3

.................................................................... Gamasiphis pulchellus (Berlese, 1887), Italy

- Seta Zv2 about 0.5 times as long as distance between its base and base of Zv3 ...............

......................................................................Gamasiphis hamatellus Karg, 1998, Ecuador

45. Ventrianal shield with seven pairs of setae in addition to circumanal setae (Zv2 absent).

................................................ Gamasiphis parpulchellus Nasr and Mersal, 1986, Egypt

- Ventrianal shield with eight pairs of setae in addition to circumanal setae. ................. 46

46. Seta j2 [indicated as s1 by Karg (1987)] distinctly posterior to j1; seta j4 [indicated as i3

by Karg (1987)] about 1.2 times as long as distance between its base and base of j5

[indicated as i4 by Karg (1987)]. ............. Gamasiphis lanceolatus Karg, 1987, Germany

- Seta j2 laterad to j1; seta j4 about 0.9 times as long as distance between its base and

base of j5 ................................................................................. Gamasiphis n. sp. 2, Brazil

47. Opisthonotal region with two pairs of setae (including Z5) much longer than others.

................................................................... Gamasiphis hyalinus Karg, 2003, Costa Rica

- Opisthonotal region with a single pair of setae (Z5) much longer than others .............. 48

48. Setae j6 [indicated as i5 by Karg (1994a)] at least 1.2 times as long as distance between

their bases ...................................................................................................................... 49

- Setae j6 at most 0.7 times as long as distance between their bases ............................... 50

569

49. Setae Jv5 about 0.7 times as long as post-anal seta. ..........................................................

.............................................................Gamasiphis sextus Vitzthum, 1921 from Germany

- Setae Jv5 [indicated as V8 by Karg (1994a)] about 1.3 times as long as post-anal seta

......................................................... Gamasiphis vinculi Karg, 1994a, Galapagos Islands

50. Ventrianal shield with seven pairs of setae in addition to circumanal setae (Jv5 absent)

..................................................................... Gamasiphis turgicalcareus Ma, 2009, China

- Ventrianal shield with eight pairs of setae in addition to circumanal setae. .................. 51

51. Seta Jv5 about as long as para-anal seta..............................................................................

. ...................................... Gamasiphis paulista Castilho, Moraes and Narita, 2010, Brazil

- Seta Jv5 at least twice as long as para-anal seta............................................................. 52

52. Dorsal and ventrianal shields mostly imbricate .................................................................

...............................................................Gamasiphis adanalis Karg, 1990, Lesser Antilles

- Dorsal shield smooth and ventrianal shield smooth or transversely striate anteriorly to

Zv2 and smooth elsewhere. ............................................................................................ 53

53. Seta Jv5 at least twice as long as post-anal seta. ............................................................ 54

- Seta Jv5 at most 1.3 times as long as post-anal seta ...................................................... 55

54. With three pairs of J setae; ventrianal shield smooth..........................................................

. ........................................ Gamasiphis hamifer (Trägårdh, 1952), Flint and Rapa Islands

- With four pairs of J setae; ventrianal shield transversely striate anteriorly to Zv2 and

smooth elsewhere .................................. Gamasiphis decoris Karg, 1990, Lesser Antilles

55. Seta st5 about 2.3 times as long as Jv1 ..............................................................................

........................................................Gamasiphis hemicapillus Karg, 1990, Lesser Antilles

- Seta st5 at most as long as Jv1 ....................................................................................... 56

56. Seta st5 at most 0.6 times as long as Jv1; seta Jv2 about as long as distance between its

base and base of Jv4. ...................................................................................................... 57

- Seta st5 about as long as Jv1; seta Jv2 at most 0.7 times as long as distance between its

base and base of Jv4 ....................................................................................................... 58

57. Seta j3 about 0.8 times as long as distance between its base and base of j4; seta j4 about

0.7 times as long as distance between its base and base of j5.............................................

. .................... Gamasiphis krieli Van Driel, Loots and Marais, 1977, Saint Helena Island

- Seta j3 about 1.7 times as long as distance between its base and base of j4; seta j4 about

1.3 times as long as distance between its base and base of j5 ............................................

..................................................................Gamasiphis indicus Bhattacharyya, 1978, India

570

58. Seta Zv1 about twice as long as Jv1; seta Jv5 about 0.7 times as long as post-anal seta....

. ............................................................... Gamasiphis maheensis Loots, 1980, Seychelles

- Seta Zv1 about as long as Jv1; seta Jv5 about as long as post-anal seta ........................ 59

59. Seta Zv2 about as long as distance between its base and base of Zv3; seta Jv2 about half

as long as distance between its base and base of Jv3 ........................................................

.........................................................Gamasiphis novipulchellus Ma and Yin, 1998, China

- Seta Zv2 about half as long as distance between its base and base of Zv3; seta Jv2 about

0.3 times as long as distance between its base and base of Jv3 .........................................

................................................................................Gamasiphis aduncus Ma, 2004, China

9. 4 Discussion

In addition to the characteristics mentioned in the diagnosis of Gamasiphis, other

characteristics were found in the three species described in this paper, and could also

correspond to characteristics common for Gamasiphis species. However, most of the

descriptions published so far make no reference to most of these characteristics, namely: fixed

cheliceral digit with a setiform pilus dentilis (also shown for G. adanalis, G. australicus, G.

bengalensis, G. brevigenitalis, G. coniunctus, G. fornicatus, G. furcatus, G. gandensius, G.

hamifer, G. hemicapillus, G. holocapillus, G. krieli, G. lanceolatus, G. maheensis, G.

paulista, G. plenosetosus, G. pinguis, G. quadruplicis, G. saccus, G. setosus, G. spinulosus,

G. uncifer, and G. vinculi); antiaxial lyrifissure of chelicera (also in G. paulista); margins of

deutosternum not distinct (also in G. benoiti, G. lanceolatus, G. maheensis, G. parpulchellus

and G. paulista); anterior-most row of deutosternal denticles arranged as in an inverted ―V‖ or

―U‖ (also in G. benoiti, G. decoris, G. fornicatus, G. lanceolatus, G. paulista and G.

pulchellus); seta h3 inserted between h1 and h2, from mediad to laterad to h1, and mediad to

h2 (also in G. australicus, G. benoiti, G. decoris, G. fornicatus, G. hamifer, G. krieli, G.

lanceolatus, G. maheensis, G. parpulchellus, G. paulista, G. pulchellus, G. saccus and G.

sextus). The antiaxial lyrifissure is subparallel to the adjacent dorsal surface of the chelicar or

slightly tilted downward anteriorly in adult females (not distinguishable in adult males).

Gamasiphis fornicatus, G. lenifornicatus, G. australicus are illustrated as having distinct

margins of deutosternum; however, this could have been done in the descriptions or

redescriptions just to indicate the limits of the structure. A single exception in relation to the

relative position of h3 refers to G. lenifornicatus; h3 is inserted postero-mediad to h2 in that

species. In addition, two and one pair of gland opening (pores) were distinguishable in

571

Gamasiphis n. sp. 2 and Gamasiphis n. sp. 3, respectively, whereas no pores were

distinguished in Gamasiphis n. sp. 1 and G. plenosetosus. Scarcity of pores seems also tobe

common in species of this genus.

Also relevant seems to be the presence of four pairs of lyryfissures on sternal (female)

or sternogenital (male) shields of the species described or redescribed in this paper. Gamasida

females and males usually have three pairs of lyryfissures (often mentioned as pores in the

literature) in the sternal area. These are located next and posterior to st1 and st2 and next and

antero-mediad to st4. The latter can either be located on the sternal or metasternal shields, or

on the unsclerotized membrane. In some groups of Gamasida [eg. Mucroseius and some

Proctolaelaps (both Mellicharidae) and Anystipalpus and some Antennoseius (both Ascidae)],

the lyryfissure next to st4 may be indistinguishable.

Despite the fact that this unusual characteristic had not been highlighted in previous

works, it is shared by at least two other species of the same genus, G. hamifer and G. paulista,

although in the latter species males have only three pairs of lyryfissures on the sternogenital

shield (nothing is known about this aspect for males of the former species). Four pairs of

lyryfissures in the sternal shield of adult females have also been clearly shown in the

illustrations of the original descriptions of 20 other ologamasid species, although nothing was

mentioned in this regard in the corresponding original descriptions; these species are

Acugamasus watsoni (Hirschmann), Athiasella dentata (Womersley), Cymiphis cymosus

(Lee), Cymiphis mansoni (Lee), Cymiphis nucilis (Lee), Desectophis magnosimilis Karg,

Euepicrius caesariatus Lee, Euepicrius lootsi Lee, Gamasellus morogoroensis Hurlbutt,

Gamasellus muscosus Hurlbutt, Gamasellus plumatilis Karg, Gamasellus tragardhi

(Womersley), Gamasellus uluguruensis Hurlbutt, Gamasellus virgosus (Lee), Gamasiphoides

costai Lee and Hunter, Heydeniella womersleyi Lee and Hunter, Hydrogamasellus antarcticus

(Trägårdh), Ologamasus cananeiae Silva, Moraes and Krantz, Ologamasus striolatus

(Berlese) and Pilellus rugipellis Lee and Hunter. In 13 of these species, males are also known

to have four pairs of lyrifissures in the sternogenital shield; these are A. dentata, C. cymosus,

D. magnosimilis, E. caesariatus, E. lootsi, G. morogoroensis, G. tragardhi, G. uluguruensis,

G. costai, H womersleyi, O. cananeiae, O. striolatus, P. rugipellis. Further studies should

show whether four pairs of sternal lyrifissures is a common feature of ologamasid females.

In relation to distribution, the vast majority of the species of Gamasiphis (43 species)

were described from tropical islands around he world (without including Australia), especially

from New Caledonia (23 species). This may be in large part related to differences in the

sampling efforts dedicated in each region. However, extensive surveys have been conducted

572

in Europe and in southern Africa, and few Gamasiphis species were described from those

regions. Of the three species described from Europe, two were originally collected from

greenhouses and one from an outdoor nursery. The four Gamasiphis species described so far

from Brazil were collected in a small (about 250 m2) internal garden, where soil humidity is

usually kept high.

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New Guinea. Journal of the Linnean Society, Zoology, London, v. 42, p. 505-599, 1956.

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10 FINAL CONSIDERATIONS

The results of this work are considered to represent a significant contribution to those

interested on the taxonomy of the Rhodacaroidea. One of the main results refers to the

updating and complementation of the species list within each of the families composing this

superfamily. The revision of four genera, the description of 5 new species and the

redescription in total or in part of 17 species are also considered relevant. Also of high interest

was the preparation of the diagnosis of the families Digamasellidae Evans, Laelaptonyssidae

Womersley, Ologamasidae Ryke, Rhodacaridae Oudemans and Teranyssidae Halliday and of

the genera of Rhodacaridae, as well as the preparation of dichotomous keys for the genera of

all families and for the world species of Gamasiphis, one of the large genera of

Ologamasidae.

Taxonomy of Rhodacaroidea mites (Acari: Mesostigmata) Raphael de Campos … · 2012. 3. 23. ·...

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