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Gastrointestinal Helminths from Six Species of Frogs and Three

Species of Lizards, Sympatric in Pará State, Brazil

Author(s): Stephen R. Goldberg, Charles R. Bursey, Janalee P. Caldwell, Laurie

J. Vitt, and Gabriel C. Costa

Source: Comparative Parasitology, 74(2):327-342. 2007.

Published By: The Helminthological Society of Washington

DOI: 10.1654/4268.1URL: http://www.bioone.org/doi/full/10.1654/4268.1

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Gastrointestinal Helminths from Six Species of Frogs and Three Species of Lizards, Sympatric in Para State, Brazil

STEPHEN R. GOLDBERG,1,4 CHARLES R. BURSEY,2 JANALEE P. CALDWELL,3 LAURIE J. VITT,3 AND

GABRIEL C. COSTA3

1 Department of Biology, Whittier College, Whittier, California 90608, U.S.A. (e-mail: [email protected]),2 Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A.

(e-mail: [email protected]), and3 Sam Noble Oklahoma Museum of Natural History and Zoology Department, University of Oklahoma, Norman,

Oklahoma 73072, U.S.A. (e-mails: [email protected], [email protected], [email protected]).

ABSTRACT: Forty-three frogs representing 6 species ( Dendropsophus cachimbo, Scinax fuscomarginatus, Leptodactylus

fuscus, Leptodactylus martinezi, Leptodactylus mystaceus, and Leptodactylus rhodomystax ) and 35 lizards representing 3

species ( Kentropyx calcarata, Leposoma osvaldoi, and Potamites ecpleopus) collected in the Brazilian state of Para were

examined for helminths. One species of Trematoda, Brachycoelium salamandrae, and 12 species of Nematoda, adults of Capillaria recondita, Cosmocerca brasiliense, Cosmocerca podicipinus, Falcaustra belemensis, Falcaustra maculata,

Kentropyxia sauria, Oswaldocruzia vaucheri, Physaloptera retusa, Schrankiana formulosa, Schrankiana fuscus,

Schrankiana schranki, and juveniles of Acuariidae gen. sp., were found. Only B. salamandrae occurred in both frogs and

lizards. There were 1.53 6 0.13 (x 6 1 SE) helminth species/infected frogs and 28.52 6 11.7 helminth individuals/infected

frogs and 1.12 6 0.08 helminth species/infected lizards and 6.47 6 1.58 helminth individuals/infected lizards. Thirteen new

host records and 2 new locality records are reported.

KEY WORDS: Trematoda, Nematoda, Anura, Sauria, Para, Brazil.

The Amazonian-Guiana region of South America

has a diverse anuran fauna of at least 305 amphibian

species (Duellman, 1999) and at least 89 lizard species

(Avila-Pires, 1995). The purpose of this study is toreport gastrointestinal helminths from 6 species of

frogs and 3 species of lizards from the Brazilian

state of Para, the northeastern portion of the Amazo-

nian Guiana biogeographical region: Cachimbo tree-

frog, Dendropsophus cachimbo; brown-bordered

snouted treefrog, Scinax fuscomarginatus; whistling

frog, Leptodactylus fuscus; Martinez’s thin-toed frog,

Leptodactylus martinezi; rose-lipped thin-toed frog,

Leptodactylus rhodomystax ; and Spix’s kentropyx,

Kentropyx calcarata; Oswald’s leposoma, Leposoma

osvaldoi; and the common stream lizard, Potamitesecpleopus.

MATERIALS AND METHODS

Forty-three frogs ( Dendropsopus cachimbo, n ¼ 10),(S. fuscomarginatus, n ¼ 12), ( L. fuscus, n ¼ 12),( L. martinezi, n ¼ 3), ( Leptodactylus mystaceus, n ¼ 1),( L. rhodomystax , n¼ 5) and 35 lizards ( L. osvaldoi, n¼ 5),( P. ecpleopus, n ¼ 13), ( K. calcarata, n ¼ 17) were col-lected in Para, Novo Progresso Municipality, Brazil inOctober 2004 and initially deposited in the herpetologycollection of the University of Brasılia. These specimens

were later transferred to the Sam Noble Oklahoma Museumof Natural History (OMNH), University of Oklahoma,Norman, Oklahoma. Specimens were fixed in 10% neutralbuffered formalin and preserved in 70% ethanol. The

abdominal cavity of each specimen was opened and thedigestive tract was removed. The digestive tract was openedand examined for helminths under a dissecting microscope.Trematodes were regressively stained in hematoxylin,cleared in xylene, and mounted in Canada balsam for identification. Nematodes were placed on a glass slide,cleared in a drop of lactophenol solution, covered witha cover slip, and identified. Helminths were deposited inthe United States National Parasite Collection (USNPC),Beltsville, Maryland. Amphibian taxonomy is in accordancewith Frost et al. (2006); lizard taxonomy is in accordancewith Avila-Pires (1995) and Doan and Castoe (2005),however, we have indicated the taxonomic assignment of a host at the time of its collection.

Hylidae

Dendropsophus cachimbo

(Napoli and Caramaschi, 1999)

Ten specimens were collected 21 October 2004 at

Campo de Provas Brigadeiro Veloso, Novo Progresso

Municipality, Para, Brazil (9821928.80S, 54854920.20W)

and deposited in OMNH as 40195–40204.

Cosmocerca brasiliense Travassos, 1925

Prevalence and intensity: One of 10 hosts infected

(10%, 1).4 Corresponding author.

Comp. Parasitol.74(2), 2007, pp. 327–342

327



Temporal distribution: 21 October 2004, 1 host with

1 male nematode.

Site of infection: Large intestine.

Type host and type locality: Chaunaus crucifer

(¼ Bufo crucifer ), Angra do Reis, Brazil (Travassos,

1925).

Additional hosts from Brazil: Guenther’s robber

frog, Eleutherodactylus guentheri, (Travassos, 1925);

smooth horned frog, Proceratophrys appendiculata

(Boquimpani-Freitas et al., 2001); rock river frog,

Thoropa miliaris (Travassos, 1925).

Other reported hosts: Lowland thin-toed frog, Lep-

todactylus andreae (¼ Adenomera andreae; Bursey

et al., 2001); Napo’s thin-toed frog, Leptodactylus

hylaedactylus (¼ Adenomera hylaedactyla; Burseyet al., 2001); marine toad, Chaunus marinus (¼ Bufo

marinus; Bursey et al., 2001); ruby poison frog,

Epipedobates parvulus (¼ Dendrobates parvulus;

Dyer and Altig, 1976, 1977); Perez’s snouted frog,

Edalorhina perezi (Bursey et al., 2001); Amazon

robber frog, Eleutherodactylus altamazonicus (Dyer

and Altig, 1976, 1977); La Paz robber frog,

Eleutherodactylus cruralis (Bursey et al., 2001);

Rıo Mamore robber frog, Eleutherodactylus fenes-

tratus (Bursey et al., 2001); metallic robber frog,

Eleutherodactylus lanthanites (Dyer and Altig, 1976,1977); Peru robber frog, Eleutherodactylus peruvia-

nus (Bursey et al., 2001); Pachitea robber frog,

Eleutherodactylus toftae (Bursey et al., 2001); bleat-

ing frog, Hamptophryne boliviana (Dyer and Altig,

1976, 1977); giant gladiator treefrog, Hypsiboas

boans (¼ Hyla boans; Dyer and Altig, 1976, 1977;

Bursey et al., 2001); short-headed treefrog, Dendrop-

sophus brevifrons (¼ Hyla brevifrons; Bursey et al.,

2001); Troeschel’s treefrog, Hypsiboas calcaratus

(¼ Hyla calcarata; Bursey et al., 2001); banded

treefrog, Hypsiboas fasciatus (¼

Hyla fasciata; Burseyet al., 2001); map treefrog, Hypsiboas geographicus

(¼ Hyla geographica; Dyer and Altig, 1976, 1977);

Demerara Falls treefrog, Hypsiboas granosus (¼ Hyla

granosa; Bursey et al., 2001); Koechlin’s treefrog,

Dendroposphus koechlini (¼ Hyla koechlini; Bursey

et al., 2001); basin treefrog, Hypsiboas lanciformis

(¼ Hyla lanciformis; Dyer and Altig, 1976, 1977);

Leal’s treefrog, Dendropsophus leali (¼ Hyla leali;

Bursey et al., 2001); Beireis’ treefrog, Dendropso-

phus leucophyllatus (¼ Hyla leucophyllata; Dyer and

Altig, 1976, 1977; Bursey et al., 2001); marbled

treefrog, Dendropsophus marmoratus (¼ Hyla mar-

morata; Dyer and Altig, 1976, 1977; Bursey et al.,

2001); small-headed treefrog, Dendropsophus parvi-

ceps (¼ Hyla parviceps; Bursey et al., 2001); red-

robed treefrog, Dendropsophus rhodopeplus (¼ Hyla

rhodopepla; Bursey et al., 2001); Sarayacu treefrog,

Dendropsophus sarayacuensis (¼ Hyla sarayacuen-

sis; Dyer and Altig, 1976, 1977); Schubart’s treefrog,

Dendropsophus schubarti (¼ Hyla schubarti; Bursey

et al., 2001); common big-headed frog, Ischnocnema

quixensis (Dyer and Altig, 1976, 1977); Bolivian

white-lipped frog, Leptodactylus bolivianus (Bursey

et al., 2001); South American bullfrog, Leptodactylus

pentadactylus (Bursey et al., 2001); Peter’s frog,

Leptodactylus petersii (Bursey et al. 2001); Manaus

slender-legged treefrog, Osteocephalus taurinus

(Bursey et al., 2001); Surinam golden-eyed treefrog,

Trachycephalus coriaceus (¼ Phrynohyas coriacea;

Bursey et al., 2001); toady leaf frog, Phyllomedusa

atelopoides (Bursey et al., 2001); jaguar leaf frog, Phyllomedusa palliata (Bursey et al., 2001); brown-

belly leaf frog, Phyllomedusa tarsius (Dyer and

Altig, 1976, 1977); tiger-striped leaf frog, Phyllome-

dusa tomopterna (Bursey et al., 2001); white-lined

leaf frog, Phyllomedusa vaillanti (Bursey et al.,

2001); Tarauaca snouted treefrog, Scarthyla goino-

rum (¼Scarthyla ostinodactyla; Bursey et al., 2001);

Eirunepe snouted treefrog, Scinax garbei (Bursey

et al., 2001); yellow-snouted treefrog, Scinax icter-

icus (Bursey et al., 2001); Henle’s snouted treefrog,

Scinax pedromedinai (Bursey et al., 2001); Orinocolime treefrog, Sphaenorhynchus lacteus (Bursey

et al., 2001).

Geographic range: Brazil (Travassos, 1925), Ecua-

dor (Dyer and Altig, 1976, 1977), Peru (Bursey et al.,

2001).

Specimen deposited: USNPC 98183; 1 vial.

Remarks: Dendropsophus cachimbo represents a new

host record for Cosmocerca brasiliense.

Scinax fuscomarginatus

(Lutz, 1925)

Twelve specimens were collected from 21–31

October 2004. Seven collected on 31 October were

from 46.9 km W Campo de Provas Brigadeiro

Veloso, Novo Progresso Municipality, Para, Brazil

(9822950.10S, 55820925.70W) and deposited in OMNH

as 40216–40222. Four collected on 21 October were

from Campo de Provas Brigadeiro Veloso, Novo

Progresso Municipality, Para, Brazil (9821928.80S,

54854921.80W) OMNH 40211–40214. One collected

on 24 October was from 3 km W, 6.5 km N Campo

de Provas Brigadeiro Veloso, Novo Progresso

328 COMPARATIVE PARASITOLOGY, 74(2), JULY 2007



Municipality, Para, Brazil (9821928.80S,54856936.00W)

OMNH 40215.

Cosmocerca podicipinus Baker and

Vaucher, 1984 Prevalence and intensity: One of 12 hosts infected

(8%, 2).


1 male and 1 female nematodes.


Type host and type locality: Dark-bellied frog,

Leptodactylus podicipinus, Capitan Bado, Amambay

Province, Paraguay (Baker and Vaucher, 1984).

Additional hosts from Brazil: None.

Other reported hosts: Pacific fat sleeper, Dormitator

latifrons (Garrido-Olvera et al., 2004); Condoto

stubfoot toad, Atelopus spurrelli (Goldberg and

Bursey, 2003); South American leaf toad, Rhinella

margaritifer (¼ Bufo typhonius; Bursey et al., 2001);

dull rocket frog, Colostethus marchesianus (Bursey

et al., 2001); harlequin poison frog, Dendrobates

histrionicus (Goldberg and Bursey, 2003); imitator

robber frog, Eleutherodactylus imitatrix (Bursey

et al., 2001); polymorphic robber frog, Eleutherodacty-

lus rhodopis (Goldberg et al., 2002); brillant-thighed

poison frog, Allobates femoralis (¼ Epipedobates femo-

ralis; Bursey et al., 2001); marbled white-lipped frog,

Leptodactylus elenae (Baker and Vaucher, 1984);

whistling frog, Leptodactylus fuscus (Baker and

Vaucher, 1984); stripe-thighed frog, Leptodactylus

leptodactyloides (Bursey et al., 2001); Miranda’s

white-lipped frog, Leptodactylus macrosternum (Baker

and Vaucher, 1984); Sabinal frog, Leptodactylus

melanonotus (Goldberg and Bursey, 2002; Goldberg

et al., 2002); Forrer’s grass frog, Lithobates forreri

(¼ Ranaforreri; Goldberg andBursey, 2002); Lithobatescf. forreri (¼ Rana cf. forreri; Bursey and Goldberg,

2005); blue-spotted Mexican treefrog, Smilisca cyanos-

tica (Goldberg et al., 2002).

Geographic range: Brazil (this study), Colombia

(Goldberg and Bursey, 2003), Costa Rica (Bursey

and Goldberg, 2005), Mexico (Goldberg and Bursey,

2002), Paraguay (Baker and Vaucher, 1984), Peru

(Bursey et al., 2001).

Specimens deposited: USNPC 98194; 1 vial.

Remarks: Scinax fuscomarginatus represents a new

host record for Cosmocerca podicipinus. Brazil is

a new locality record.

Acuariidae gen. sp. (juveniles in cysts)


(8%, 2).


2 cysts.

Site of infection: Stomach wall.

Type host and type locality: Adult acuariid nematodes

are parasites of aquatic birds (Anderson, 2000).

Additional hosts of juveniles from Brazil: agile

mabuya, Mabuya agilis (Vrcibradic et al., 2000;

Vrcibradic et al., 2002); Caissara mabuya, Mabuya

caissara (Rocha and Vrcibradic, 2003); Hoge’s

mabuya, Mabuya macrorhyncha (Vrcibradic et al.,

2000; Vrcibradic et al., 2002); Amazon lava lizard,

Tropidurus torquatus (Vrcibradic et al., 2000).

Other reported hosts of juveniles: Edalorhina perezi

(Bursey et al., 2001); Hyla rhodopepla (Bursey et al.,

2001). Acuariid juveniles have also been reported

from the border anole, Norops limifrons from Costa

Rica (Bursey and Goldberg, 2003) and from the

western fence lizard, Sceloporus occidentalis from

California (Goldberg et al., 1998).

Geographic range: The family Acuariidae has

cosmopolitan distribution (Yamaguti, 1961).


Remarks: Acuariid nematodes typically mature in

aquatic birds; they require an arthropod intermediate

host to complete the life cycle; frogs may serve as para-

tenic hosts (Anderson, 2000). Scinax fuscomarginatus

represents a new host record for acuariid juveniles.

Leptodactylidae

Leptodactylus fuscus

(Schneider, 1799)

Twelve specimens were collected from 21 October– 11 November 2004. Eleven were collected on 29, 31

October and 1–3, 9, 11 November from 46.9 km W

Campo de Provas Brigadeiro Veloso, Novo Progresso

Municipality, Para, Brazil (9822950.10S,55820925.70W)

and deposited in OMNH as 40229–40239. One

was collected on 21 October at Campo de Provas

Brigadeiro Veloso, Novo Progresso Munici-

pality, Para, Brazil (9821928.80S, 54854921.80W)

and deposited as OMNH 40228.

Oswaldocruzia vaucheri Ben Slimane and

Durette-Desset, 1993


(8%, 2).

GOLDBERG ET AL.—HELMINTHS OF FROGS AND LIZARDS IN BRAZIL 329



Temporal distribution: 31 October, 2004, 1 host

with 2.

Site of infection: Small intestine.

Type host and type locality: common big-headedfrog, Ischnocnema quixensis, San Pablo, Napo Prov-

ince, Ecuador (Ben Slimane and Durette-Desset,

1993).


Other reported hosts: None.

Geographic range: Ecuador (Ben Slimane and

Durette-Desset, 1993), Brazil (this study).


Remarks: Leptodactylus fuscus represents a new host

record for Oswaldocruzia vaucheri. Brazil is a new

locality record for O. vaucheri.

Schrankiana formulosa Freitas, 1959

Prevalence, mean intensity, and range: Eight of 12

hosts infected (67%, 12.8 6 13.9 SD, 2–38).

Temporal distribution: 29, 31 October. 2, 3, 9, 11

November 2004, 8 hosts with 2, 2, 3, 8, 9, 9, 31, 38,

respectively.Site of infection: Large intestine.

Type host and type locality: Leptodactylus fuscus,

Itaguai, Rio de Janeiro, Brazil (Freitas, 1959).


Other reported hosts: Leptodactylus elenae (Baker

and Vaucher, 1988).

Geographic range: Brazil (Freitas, 1959), Paraguay

(Baker and Vaucher, 1988).


Remarks: This is the third study to report Schranki-

ana formulosa in Leptodactylus fuscus (Freitas, 1959;

Baker and Vaucher, 1988; this study).

Schrankiana fuscus Baker and

Vaucher, 1988

Prevalence, mean intensity, and range: Nine of 12


Temporal distribution: 31 October, 2, 3, 9, 11

November 2004, 9 hosts with 2, 4, 5, 9, 13, 16, 17,

17, 19, respectively.


Type host and type locality: Leptodactylus fuscus,

Alto Parana Province, Paraguay (Baker and Vaucher,

1988).



Geographic range: Brazil (this study); Paraguay

(Baker and Vaucher, 1988).


Remarks: Brazil is a new locality record for S. fuscus.

Leptodactylus martinezi

(Bokermann, 1956)

Three specimens were collected on 9 November

2004 from 46.9 km W Campo de Provas Brigadeiro


(9822950.10S, 55820925.70W) and deposited in OMNH

as 40225–40227.

Brachycoelium salamandrae (Frolich, 1789)

Dujardin, 1845

Synonymy: Fasciola salamandrae Froelich, 1789;

Distoma crassicolle Rudolphi, 1809; Distomum flavocinctum Linstow, 1879; Brachycoelium hospi-

tale Stafford, 1903; Brachycoelium obesum Nicoll,

1914; Brachycoelium trituri Holl, 1928; Brachycoe-

lium daviesi Harwood, 1932; Brachycoelium mer-

idionalis Harwood, 1932; Brachycoelium storeriae

Harwood, 1932; Brachycoelium dorsale Byrd, 1937;

Brachycoelium georgianum Byrd, 1937; Brachycoe-

lium louisianae Byrd, 1937; Brachycoelium mesor-

chium Byrd, 1937; Brachycoelium ovale Byrd, 1937;

Brachycoelium elongatum Cheng, 1958; Brachycoe-

lium mesocoeliiformis Freitas, 1961; Brachycoeliumambystomae Couch, 1966.

Prevalence, mean intensity, and range: Three of 3


Temporal distribution: 9 November 2004, 3 hosts

with 1, 5, 5, respectively.


Type host and type locality: alpine salamander,

Salamandra atra, Germany (Frolich, 1789).

Additional hosts from Brazil: goldenscale anole,

Anolis nitens (¼ A. chrysolepis but reported as Anolis

scypheus; Freitas, 1961); Leptodactylus rhodomystax




(this study); Leptodactylus rhodomystax (this study);

Leposoma osvaldoi (this study).

Other reported hosts: Nearctic Realm: northern cricket

frog, Acris crepitans (¼ Acris gryllus; Najarian, 1955);

American toad, Anaxyrus americanus (¼ Bufo amer-

icanus; Rosen and Manis, 1976); western toad,

Anaxyrus boreas (¼ Bufo boreas; Lehmann, 1965);

Fowler’s toad, Anaxyrus fowleri (¼ Bufo fowleri;

Brandt, 1936; Rankin, 1938; Hering and Murad,

1969); southern toad, Anaxyrus terrestris (¼ Bufo

terrestris; Manter, 1938); green treefrog, Hyla

cinerea (Harwood, 1932); crawfish frog, Lithobates

areolatus (¼ Rana areolata; Manter, 1938); pickerel

frog, Lithobates palustris (¼ Rana palustris; Rankin,

1938, 1945); southern leopard frog, Lithobates

sphenocephalus (¼ Rana sphenocephala; Harwood,1932; Brandt, 1936; Byrd, 1937; Manter, 1938;

Rankin, 1938; Parker, 1941); wood frog, Lithobates

sylvaticus (¼ Rana sylvatica; Odlaug, 1954; Najarian,

1955; Prudhoe and Bray, 1982; McAllister, Trauth

and Bursey, 1995a; McAllister, Upton et al., 1995);

Brimley’s chorus frog, Pseudacris brimleyi (Brandt,

1936; Rankin, 1938); spring peeper, Pseudacris

crucifer (Brandt, 1936; Rankin, 1938); ornate chorus

frog, Pseudacris ornata (¼ Pseudacris occidentalis;

Byrd, 1937); Pacific chorus frog, Pseudacris regilla

(¼ Hyla crucifer ; Lehmann, 1965); western chorusfrog, Pseudacris triseriata (Harwood, 1932; Odlaug,

1954); American bullfrog, Lithobates catesbeianus

(¼ Rana catesebiana; Parker, 1941; Prudhoe and

Bray, 1982); green frog, Lithobates clamitans

(¼ Rana clamitans; Fantham and Porter, 1948);

northern leopard frog, Lithobates pipiens (¼ Rana

pipiens; Lehmann, 1965; Rosen and Manis, 1976);

red-legged frog, Rana aurora (Lehmann, 1965);

spotted frog, Rana pretiosa (Lehmann, 1965);

Jefferson salamander, Ambystoma jeffersonianum

(Rankin, 1938, 1945); blue-spotted salamander, Ambystoma laterale (Muzzall and Schinderle, 1992);

long-toed salamander, Ambystoma macrodactylum

(Waitz, 1961); spotted salamander, Ambystoma

maculatum (Rankin, 1937, 1938, 1945; Rabalais,

1970; Dyer and Brandon, 1973; Rosen and Manis,

1976; Coggins and Sajdak, 1982; Bolek and Coggins,

1998); marbled salamander, Ambystoma opacum

(Byrd, 1937; Rankin, 1937, 1938; Parker, 1941;

Couch, 1966; Rabalais, 1970; Dyer and Brandon,

1973; Joy and Mills, 1975); mole salamander,

Ambystoma talpoideum (Dyer and Brandon, 1973);

small-mouthed salamander, Ambystoma texanum

(Harwood, 1932; Rosen and Manis, 1976); three-

toed amphiuma, Amphiuma tridactylum (Bennett and

Humes, 1938); southern dusky salamander, Des-

mognathus auriculatus (Bennett, 1938); Ouachita

dusky salamander, Desmognathus brimleyorum

(McAllister, Bursey et al., 1995); northern dusky

salamander, Desmognathus fuscus (Byrd, 1937;

Rankin, 1937, 1938, 1945; Parker, 1941; Odlaug,

1954; Fischthal, 1955a, b; Cheng, 1958; Cheng and

Chase, 1961; Rabalais, 1970; Rosen and Manis,

1976; Bogitsh and Ryckman, 1982); seal salamander,

Desmognathus monticola (Rankin, 1937; Cheng,

1958); Allegheny Mountain dusky salamander,

Desmognathus ochrophaeus (Rankin, 1937); black-

bellied salamander, Desmognathus quadramaculatus

(Rankin, 1937; Cheng, 1958); California giant sala-

mander, Dicamptodon ensatus (Pratt and McCauley,

1961); ensatina, Ensatina eschscholtzii (Lehmann,

1954; Pratt and McCauley, 1961); northern two-linedsalamander, Eurycea bislineata (Rankin, 1937, 1945;

Parker, 1941; Fischthal, 1955a, b; Rabalais, 1970);

long-tailed salamander, Eurycea longicauda (Rankin,

1937; Dyer and Brandon, 1973); cave salamander,

Eurycea lucifuga (Dyer and Brandon, 1973; Dyer and

Peck, 1975; Castle et al., 1987); many-ribbed

salamander, Eurycea multiplicata (McAllister, Trauth

and Bursey, 1995b); grotto salamander, Eurycea

spelaea (¼Typhlotriton spelaeus; Dyer, 1975); spring

salamander, Gyrinophilus porphyriticus (Fischthal,

1955b; Catalano et al., 1982); four-toed salamander, Hemidactylum scutatum (Rankin, 1938); black-spotted

newt, Notophthalmus meridionalis (Harwood,

1932; Rankin, 1937; Manter, 1938); eastern newt,

Notophthalmus viridescens (Stafford, 1900, 1903;

Holl, 1928; Rankin, 1937, 1938; Russell, 1951;

Fischthal, 1955b; Cheng, 1958; Jackson and Beau-

doin, 1967; Rabalais, 1970); Red Hills salamander,

Phaeognathus hubrichti (Brandon, 1965); eastern

red-backed Salamander, Plethodon cinereus (Stafford,

1903, 1905; Rankin, 1937, 1938, 1945; Fischthal,

1955a, b; Cheng, 1958; Cheng and Chase, 1961;Coggins and Sajdak, 1982; Muzzall, 1990; Bursey

and Schibli, 1995; Bolek and Coggins, 1998);

northern zigzag salamander, Plethodon dorsalis

(Dyer and Brandon, 1973); northern slimy salaman-

der, Plethodon glutinosus (Byrd, 1937; Rankin, 1937,

1938; Parker, 1941; Fischthal, 1955b; Cheng, 1958;

Cheng and Chase, 1961; Brandon, 1965; Rabalais,

1970; Dyer and Brandon, 1973; Brooks, 1979);

Jordan’s salamander, Plethodon jordani (Rankin,

1937, 1938; Dyer, 1983); western red-backed sala-

mander, Plethodon vehiculum (Panitz, 1969); Yo-

nahlossee salamander, Plethodon yonahlossee

(Rankin, 1937); red salamander, Pseudotriton ruber

(Rankin, 1937, 1938; Parker, 1941; Catalano et al.,




1982); rough-skinned newt, Taricha granulosa

(Lehmann, 1954; Pratt and McCauley, 1961; Weath-

erly and Canaris, 1961; Moravec, 1984); Texas alli-

gator lizard, Gerrhonotus liocephalus (¼Gerrhonotus

ophiurus; Goldberg et al., 1999); eastern glass

lizard, Ophisaurus ventralis (Byrd, 1937); little brown

skink, Scincella lateralis, (Harwood,1932; Byrd, 1937);

rough green snake, Opheodrys aestivus (Nicoll, 1911);

DeKay’s brown snake, Storeria dekayi (Harwood,

1932); eastern box turtle, Terrapene carolina (Rausch,

1947; Stuart and Miller, 1993).

Palaearctic Realm: common European toad, Bufo

bufo (Luhe, 1909; Andre, 1912, 1917; Freund, 1934);

European common frog, Rana temporaria (Dujardin,

1845; Freund, 1934); Japanese firebelly newt,

Cynops pyrrhoghaster (Rankin, 1938); Corsican

Mountain salamander, Euproctus montanus (Combesand Knoepffler, 1968; Timon-David and Giudicelli,

1969); Caucasian salamander, Mertensiella caucasica

(Yildirimhan et al., 2005); alpine salamander, Sala-

mandra atra (Frolich, 1789; Stossich, 1889);

European fire salamander, Salamandra salamandra

(Rudolphi, 1819; Dujardin, 1845; Creplin, 1846;

Stossich, 1889; Luhe, 1909; Andre, 1912, 1917;

Lopez-Neyra, 1916; Freund, 1934; Grabda and

Grabda, 1953; Szabo, 1961; Barus et al., 1963;

Vojtkova and Vojtek, 1972; Prokopic and Krivanec,

1975; Prudhoe and Bray, 1982; Bertman, 1986);spectacled salamander, Salamandrina terdigitata

(Parona, 1896; Sonsiono, 1896); Laurenti’s alpine

newt, Triturus alpestris (Stossich, 1889; Luhe, 1909;

Freund, 1934; Callot, 1946; Buttner, 1951; Barus and

Groschaft, 1962; Vojtkova and Vojtek, 1972); north-

ern crested newt, Triturus cristatus (Luhe, 1909;

Parona, 1896; Freund, 1934; Vojtkova and Vojtek,

1972); palmate newt, Triturus helveticus (Andre,

1917; Buttner, 1951); marbled newt, Triturus mar-

moratus (Pontallie, 1852; Stossich, 1889); smooth

newt, Triturus vulgaris (Diesing, 1850; Luhe, 1909;Freund, 1934; Barus et al., 1963; Frandsen, 1974);

slowworm, Anguis fragilis (Baylis, 1928; Callot,

1946).

Geographic range: Europe, North America, South

America (Prudhoe and Bray, 1982).

Specimens deposited: USPNC 98186; 2 slides.

Remarks: There are considerable differences of opin-

ion concerning the number of species assigned to the

genus Brachycoelium. Rankin (1938) concluded that

heavy infections produce many small flukes, and light

infections produce large individuals and reduce all

known species to B. salamandrae, the type species.

However, Parker (1941) and Cheng (1958) did not

accept the synonym and recognized 7 and 10 species,

respectively, and later, Cheng and Chase (1960) and

Couch (1966) described additional species to bring

the number of species assigned to the genus to 13.

Rabalais (1970) preferred the single species ap-

proach; McAllister et al. (1995) proposed the use

of a conservative approach suggesting that until an

exhaustive revision of the genus is completed,

B. salamandrae should represent all individuals as-

signed to the genus. Bolek and Coggins (1998) and

Goldberg et al. (1999) followed the conservative ap-

proach and assigned their specimens to B. salaman-

drae. Rankin (1938) reported the egg size range of

B. salamandrae as 39–55 lm 3 27–37 lm.

Moravec and Huffman (2000) described Brachy-

coelium longleyi from specimens fixed in situ inendemic plethodontid salamanders collected in cen-

tral Texas and differentiated it from B. salamandrae

by egg size, 21–27 lm 3 12–15 lm in B. longleyi,

39–55 lm 3 27–37 lm in B. salamandrae. Species

assigned to the genus Langeronia, a species also

infecting amphibians, have an egg size similar to

that reported for B. longleyi: Langeronia burseyi (23–

28 lm 3 12–15 lm); Langeronia macrocirra (18–

26 lm 3 7–11 lm); Langeronia parva (14–16 lm 3

9–10 lm); Langeronia provitellaria (19–22 lm 3

10–11 lm) (data from Caballero and Bravo Hollis,1949; Sacks, 1952; Christian, 1970; Dailey and

Goldberg, 2000). Brachycoelium and Langeronia

have similar morphological characters, but a major

difference is the position of the genital pore: median

in Brachycoelium and to the left of midline in

Langeronia. Fig. 1C of Moravec and Huffman (2000)

suggests the possibility of a nonmidline position for

the genital pore of B. longleyi. Given the small egg

size and the position of the genital pore we are hes-

itant to assign Moravec and Huffman’s specimen to

Brachycoelium, although we would agree that it is not synonymous with B. salamandrae. We will continue

the conservative approach and assign our specimens

to B. salamandrae.

Leptodactylus martinezi represents a new host rec-

ord for B. salamandrae. Four South American hosts

are now known for B. salamandrae: L. martinezi,

L. rhodomystax , Anolis nitens, L. osvaldoi. (Freitas,

1961; this study).

Leptodactylus mystaceus

(Spix, 1824)

One specimen was collected on 24 October 2004

at 3 km W, 6.5 km N Campo de Provas Brigadeiro





(9817959.30S, 54856936.00W) and deposited as

OMNH 40240.

Schrankiana freitasi Baker, 1982 Prevalence and intensity: One of 1 host infected

(100%, 234).


234 nematodes.


Type host and type locality: Leptodactylus pentadac-

tylus, Exu, Pernambuco, Brazil (Baker, 1982).



Geographical range: Brazil (Baker, 1982).

Specimen deposited: USNPC 98190; 1 vial.

Remarks: Leptodactylus mystaceus represents a new

host record for Schrankiana freitasi.

Leptodactylus rhodomystax

Boulenger, 1884

Five specimens were collected from 28 October to

7 November 2004 at 46.9 km W Campo de ProvasBrigadeiro Veloso, Novo Progresso Municipality,

Para, Brazil (9822950.10S, 55820925.70W) and de-

posited as OMNH 40241–40245.

Brachycoelium salamandrae (Fro lich, 1789)


(20%, 1).


with 1.


Specimens deposited: USNPC 98191, 1 host with 1.

Remarks: General information and additional remarks

are reported under L. martinezi. Leptodactylus

rhodomystax represents a new host record for B.

salamandrae.

Capillaria recondita Freitas and Lent, 1942


(20%, 4).


with 4.


Type host and type locality: Brazilian spiny-thumbed

frog, Crossodactylus gaudichaudii, Brazil (Freitas

and Lent, 1942).



Geographic range: Brazil (Freitas and Lent, 1942).


Remarks: Leptodactylus rhodomystax represents a

new host record for Capillaria recondita.

Falcaustra maculata (Rudolphi, 1819)

Freitas and Lent, 1941

(Syn. Ascaris mascula Rudolphi, 1819; Floren-

cioia nitida Travassos, 1920).

Prevalence, mean intensity, and range: Three of 5


Temporal distribution: 28, 29 October, 2004, 3 hosts

with 1, 2, 8, respectively.


Type host and type locality: Rio tropical racer, Mastigodryas bifossatus (¼Coluber lichtensteini),

Brazil (Rudolphi, 1819).

Additional hosts from Brazil: yellow Cucuru toad,

Chaunus ictericus (¼ Bufo ictericus; Rodrigues et al.,

1982); Crossodactylus gaudichaudii (Gomes and

Vicente, 1966); blacksmith gladiator frog, Hypiboas

faber (¼ Hyla faber ; Freitas and Lent, 1941); Criolla

frog, Leptodactylus ocellatus (¼ Leptodactylus caligi-

nosus; Freitas and Lent, 1941; Vicente and Santos,

1976; Fabio, 1982; Rodrigues et al., 1982); Lep-

todactylus pentadactylus (Freitas, 1955; Guimaraeset al., 1976; Rodrigues et al., 1982); Santa Catarina

frog, Hylodes nasus (¼ Elosia nasus; Freitas and Lent,

1941); Wied’s groundsnake, Liophis poecilogyrus

(¼ Leimadophis poecilogyrus; Freitas, 1955).

Other reported hosts: Schneider’s toad, Chaunus

schneideri (¼ Bufo paracnemis; Lent et al., 1946).

Geographic range: Brazil (Rudolphi, 1819), Para-

guay (Lent et al., 1946).


Remarks: Leptodactylus rhodomystax represents

a new host record for Falcaustra maculata.




Schrankiana schranki (Travassos, 1925)

Strand, 1942

(Syn. Schrankia schranki Travassos, 1925).

Prevalence, mean intensity, and range: Two of 5



with 16, 111, respectively.


Type host and type locality: Leptodactylus pentadac-

tylus, Brazil (Travassos, 1925)

Additional hosts from Brazil: Leptodactylus labyrin-

thicus (Fahel, 1952).

Other reported hosts: Leptodactylus mystaceus(Dyer, 1990).

Geographic range: Brazil (Travassos, 1925), Ecua-

dor (Dyer and Altig, 1977).


Remarks: Leptodactylus rhodomystax represents

a new host record for Schrankiana schranki.

Teiidae

Kentropyx calcarata Spix, 1825Seventeen specimens were collected 3–12 Novem-

ber 2004 at 46.9 km W Campo de Provas Brigadeiro



OMNH 41812–41828.

Kentropyxia sauria Baker, 1982

Prevalence, mean intensity, and range: Seven of 17


Temporal distribution: 3, 4, 6, 8, 11, 12 November,2004, 7 hosts with 1, 3, 3, 4, 5, 7, 11, respectively.


Type host and type locality: Kentropyx calcarata,

Belem, Brazil (Baker, 1982).



Geographic range: Brazil (Baker, 1982).


Remarks: This is only the second report of Kentro-

pyxia sauria.

Physaloptera retusa Rudolphi, 1819

(Syn. Spiroptera retusa [Rudolphi, 1819] Dujar-

din, 1845; Physaloptera mucronata Leidy, 1856;

Physaloptera largada Sprehn, 1932).

Prevalence, mean intensity, and range: Two of 17



with 1, 2, respectively.

Site of infection: Stomach.

Type host and type locality: golden tegu, Tupinambis

teguixin, Brazil (Rudolphi, 1819).

Additional hosts from Brazil: common lesser toad,

Chaunus granulosus (¼

Bufo granulosus; Goncalveset al., 2002); South American leaf toad, Rhinella

margaritifer (¼ Bufo typhonius; Goncalves et al.,

2002); giant ameiva, Ameiva ameiva (Poinar and

Vaucher, 1972; Cristofaro et al., 1976; Ribas, Rocha

et al., 1998); red worm lizard, Amphisbaena alba,

(Molin, 1860); sand dune lizard, Cnemidophorus

abaetensis (Dias et al., 2005); green tailed lizard,

Cnemidophorus littoralis (Vrcibradic et al., 2000); no

common name, Cnemidophorus nativo (Menezes

et al., 2004); Spix’s whiptail, Cnemidophorus

ocellifer (Ribas et al., 1995); Lutz’s tree iguana,

Liolaemus lutzae (Rocha, 1995); agile mabuya,

Mabuya agilis (Ribas, Teixeira-Filho et al., 1998);

two-striped mabuya, Mabuya bistriata (recorded as

Scincus spixii by Molin, 1860); Paraguay mabuya,

Mabuya dorsivittata (Rocha et al., 2003); spiny lava

lizard, Tropidurus spinulosus (Vicente, 1981); Am-

azon lava lizard, Tropidurus torquatus (Vicente and

Santos, 1967; Cristofaro et al., 1976; Vicente, 1981;

Ribas, Rocha et al., 1998; Vrcibradic et al., 2000);

golden tegu, Tupinambis teguixin (¼Tupinambis

nigropunctatus; Diesing, 1851; Rudolphi, 1819;

Molin, 1860; Ortlepp, 1922).

Other reported hosts: rainbow ameiva, Ameiva

undulata (Caballero, 1951); spotted anole, Anolis

punctatus (Bursey et al., 2005); canyon spotted

whiptail, Aspidoscelis burti (¼Cnemidophorus burti;

Goldberg and Bursey, 1989a); imbricate alligator

lizard, Barisia imbricata (Goldberg et al., 1999);

zebra-tailed lizard, Callisaurus draconoides (Telford,

1970); rainbow whiptail, Cnemidophorus lemniscatus

(Caballero and Vogelsang, 1947; Diaz-Ungria, 1964;

Diaz-Ungria and Gallardo, 1968); Laurent’s whiptail,

Cnemidophorus murinus (Specian and Whittaker,

1980); southern alligator lizard, Elgaria multicari-

nata (Telford, 1970); San Lucan alligator lizard,




Elgaria paucicarinata (Goldberg, Bursey and

Beaman, 2004); western skink, Pleistodon skiltonia-

nus (¼ Eumeces skiltonianus), (Telford, 1970); long-

nosed leopard lizard, Gambelia wislizenii (Telford,

1970); Gerrhonotus liocephalus (¼Gerrhonotus

ophiurus; Goldberg et al., 1999); common green

iguana, Iguana iguana (Diaz-Ungria and Gallardo,

1968); Amazon kentropyx, Kentropyx altamazonica

(Bursey et al., 2005); forest kentropyx, Kentropyx

pelviceps (Bursey et al., 2005); Bell’s anole,

Leiosaurus bellii (Goldberg et al., 2004); no common

name, Leiosaurus catamarcensis (Goldberg et al.,

2004); no common name, Liolaemus neuquensis

(Goldberg et al., 2004); copper anole, Norops

cupreus (Goldberg and Bursey, 2004); tree runner,

Plica plica (Bursey et al., 2005); harlequin race-

runner, Plica umbra (Bursey et al., 2005); Bocourt’sspiny lizard, Sceloporus acanthinus (Caballero,

1951); Clark’s spiny lizard, Sceloporus clarkii

(Goldberg et al., 1994); blue spiny lizard, Sceloporus

cyanogenys (Goldberg et al., 1995); emerald spiny

lizard, Sceloporus formosus (Goldberg et al., 2003);

common sagebrush lizard, Sceloporus graciosus

(Woodbury, 1934), mesquite lizard, Sceloporus

grammicus (Goldberg et al., 2003); Yarrow’s spiny

lizard, Sceloporus jarrovii (Goldberg and Bursey,

1990; Bursey and Goldberg, 1991, 1994; Goldberg

et al., 1995; Goldberg et al., 1996); desert spinylizard, Sceloporus magister (Pearce and Tanner,

1973; Walker and Matthias, 1973; Goldberg et al.,

1994; Goldberg et al., 1995); canyon lizard, Scelo-

porus merriami (Goldberg et al., 1995); southern

crevice spiny lizard, Sceloporus mucronatus (Gold-

berg et al., 2003); Sceloporus occidentalis (Grund-

mann, 1959; Telford, 1970; Pearce and Tanner, 1973;

Walker and Matthias, 1973; Goldberg et al., 1998);

Texas spiny lizard, Sceloporus olivaceus (Goldberg

et al., 1995); granite spiny lizard, Sceloporus orcutti

(Telford, 1970); blue-bellied lizard, Sceloporus parvus (Goldberg et al., 2003); crevice spiny lizard,

Sceloporus poinsettii (Goldberg et al., 1993; Gold-

berg et al., 1995); blue spiny lizard, Sceloporus

serrifer (Goldberg et al., 1995); crevice swift lizard,

Sceloporus torquatus (Goldberg et al., 2003); eastern

fence lizard, Sceloporus undulatus (Morgan, 1943;

Pearce and Tanner, 1973; Goldberg et al., 1994;

Goldberg et al., 1995); southern sagebrush lizard,

Sceloporus vandenburgianus (¼Sceloporus gracio-

sus; Goldberg and Bursey, 1989b; Goldberg et al.,

1997); rose-bellied lizard, Sceloporus variabilis

(Goldberg et al., 1995; Goldberg et al., 2003); striped

plateau lizard, Sceloporus virgatus (Goldberg et al.,

1994); rose whorl-tailed iguana, Stenocercus rose-

iventris (Bursey et al., 2005); turnip-tailed gecko,

Thecadactylus rapicauda (Bursey et al., 2005); black

lava lizard, Tropidurus melanopleurus (Roca, 1997);

red tegu, Tupinambis rufescens (Sprehn, 1932); com-

mon side-blotched lizard, Uta stansburiana (Telford,

1970); eastern hog-nosed snake, Heterodon platirhi-

nos (Ortlepp, 1922); American alligator, Alligator

mississippiensis (Walton, 1927).

Geographic range: Argentina, Brazil, Mexico, United

States, Venezuela, West Indies (Baker 1987); Peru

(Bursey et al., 2005).


Remarks: Kentropyx calcarata represents a new host

record for P. retusa.

Gymnopthalmidae

Leposoma osvaldoi Avila-Pires, 1995

Five specimens were collected from 2–6 Novem-

ber 2004 at 46.9 km W Campo de Provas Brigadeiro

Veloso, Novo Progresso Municipality, Para, Brazil,


OMNH 41787–41791.

Brachycoelium salamandrae (Fro lich, 1789)


(20%, 1).

Temporal distribution: 5 November 2004, 1 host

with 1.


Specimens deposited: USNPC 98195; 1 slide.


are reported under L. martinezi. Leposoma osvaldoi

represents a new host record for B. salamandrae.

Potamites ecpleopus

(Cope 1876)

Thirteen specimens were collected on 23 October

2004 at Cachoeira do Curua, Altamira Municipality,

Para, Brazil (884498.20S, 54857949.00W). Lizards

were deposited in OMNH as 41792–41804.

Falcaustra belemensis Baker and Bain, 1981

Prevalence, mean intensity, and range: Eight of 13

hosts infected (62%, 8.968.2 SD, 2–26).

Temporal distribution: 23 October, 2004, 8 hosts

with 2, 3, 4, 5, 6, 9, 16, 26. respectively.





Type host and type locality: Two-faced neusticurus,

Neusticurus bicarinatus (Baker and Bain, 1981).

Additional hosts from Brazil: None.Other reported hosts: None.

Geographic range: Brazil (Baker and Bain, 1981).


Remarks: Potamites ecpleopus represents a new host

record for Falcaustra belemensis.

Physaloptera retusa Rudolphi, 1819


(7%, 1).

Temporal distribution: 23 October 2004, 1 host with 1.

Site of infection: Stomach.



are reported under K. calcarata. Potamites ecpleopus

represents a new host record for P. retusa.

DISCUSSIONA total of 709 helminths was collected: 599 from

21 (49%) of 43 frogs and 110 from 17 (49%) of 35

lizards. Of these, 2 nematodes (0.2%) from frogs

were juveniles not capable of reaching maturity in

herptiles. There were 14 helminth species represented

in the sample, but no individual host harbored more

than 3 species. Only one helminth species, B.

salamandrae, occurred in both frogs and lizards. Of

the infected frogs, 11 (50%) harbored 1 species of

helminth; 9 (45%) harbored 2 species; and 1 (5%)

harbored 3 species. There were 1.536

0.13 (x6

1SE) helminth species/infected frogs and 28.52 6 11.7

helminth individuals/infected frogs. No frog species

harbored more than 4 helminth species: 3 frog species

harbored 1 helminth species; 1 frog species harbored

2 helminth species; 1 frog species harbored 3

helminth species; and one frog species harbored 4

helminth species. There were 2.0 6 0.52 helminth

species/frog host species. Of the infected lizards, 15

(88%) harbored 1 species of helminth; 2 (12%)

harbored 2 species. There were 1.12 6 0.08 helminth

species/infected lizard and 6.47 6 1.58 helminth

individuals/infected lizard. Aho (1990) compiled

distributional patterns for frog and lizard helminths

in general and reported the mean 6 1 SE total

number of helminth species per host frog species as

3.54 6 0.24 (range 0–9) and per lizard species as

2.06 6 0.13 (range 0–5). The values reported here

are within the ranges reported by Aho (1990),

although the mean infective species for frogs is lower

than that reported. This may be a reflection of

regional differences in frog infection rates. The values

for lizards are similar to the values of Aho (1990).

Of the 14 species of helminths only B. salaman-

drae infected both frog and lizard species, L. fuscus,

L. rhodomystax , and L. osvaldoi. The remaining 13

helminth species, with the exception of P. retusa,

were found to infect a single host species.

ACKNOWLEDGMENTS

We thank Jessica Carlson, Cassie Ho, and SeanKark for assistance with dissections. Reptile and

amphibian specimens were collected as part of

a collaborative project between the Sam Noble

Oklahoma Museum of Natural History and the

University of Brasılia (through Dr. Guarino R. Colli)

approved by the Ministerio de Ciencia e Tecnologia

(Portaria No. 649) and the Instituto Brasileiro do

Meio Ambiente e dos Recorsos Naturaıs Renovaveis

(IBAMA, Permit No. 0217/2004-CGFAU/LIC).

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