Chelis glaphyra group, with description of a new ...

12
Ecologica Montenegrina, 38, 2020, 51-62 https://zoobank.org/urn:lsid:zoobank.org:pub:4B85DA98-957E-4788-BC8B-25526047DC8B On the taxonomy of the Chelis glaphyra (Eversmann, 1843) species- group, with description of a new subspecies of Chelis gratiosa (Grum- Grshimailo, 1890) from Kyrgyzstan, brachipterous females of Chelis kashmirica (Ferguson, 1985) and Chelis golbecki (Dubatolov, 1996), and the checklist of the species-group (Lepidoptera: Erebidae: Arctiinae: Arctiini) AIDAS SALDAITIS 1, 4 & ANTON V. VOLYNKIN 2, 3 1 Nature Research Centre, Akademijos str., 2, LT-08412, Vilnius-21, Lithuania. E-mail: [email protected] 2 Altai State University, Lenina Avenue, 61, RF-656049, Barnaul, Russia. E-mails: [email protected]; [email protected] 3 National Research Tomsk State University, Lenina Avenue, 36, RF-634050, Tomsk, Russia 4 Corresponding author Received 27 November 2020 Accepted by V. Pešić: 30 November 2020 Published online 1 December 2020. Abstract In the present paper, the checklist of the Chelis glaphyra (Eversmann, 1843) species-group with 33 new combinations established is provided. A new subspecies of Chelis gratiosa (Grum-Grshimailo, 1890) is described from Kyrgyzstan. Chelis gratiosa kashmirica is upgraded to the species level: Chelis kashmirica (Ferguson, 1985), stat. & comb. nov., its brachipterous female is described and illustrated for the first time. Female genitalia of Chelis golbecki (Dubatolov, 1996), comb. nov. are described and illustrated for the first time. Key words: Palearctia, Central Asia, new combination. Introduction The genus Palearctia Ferguson, 1984 was revised by Dubatolov (1996) and Saldaitis & Ivinskis (2002) and the full checklist of the genus was provided by Dubatolov & de Vos (2010). Subsequently, Rönkä et al. (2016) synonymized Palearctia as well as six other genera with Chelis Rambur, 1866 based on the analysis of eight genetic markers. However, the authors did not establish new combinations for species comprised by the genera synonymized leaving the taxonomy of the enlarged genus Chelis quite messy. In the present paper, we provide the checklist of the species and subspecies previously assigned with Palearctia which we consider as Ch. glaphyra species-group, with establishing new combinations for them. Chelis gratiosa, comb. nov. is distributed in high mountain regions of Central Asia and is subdivided into 10 subspecies. During his entomological trip to Transalai Range in southern Kyrgyzstan, Mr. V. Ecologica Montenegrina 38: 51-62 (2020) This journal is available online at: www.biotaxa.org/em http://dx.doi.org/10.37828/em.2020.38.7

Transcript of Chelis glaphyra group, with description of a new ...

Ecologica Montenegrina, 38, 2020, 51-62

https://zoobank.org/urn:lsid:zoobank.org:pub:4B85DA98-957E-4788-BC8B-25526047DC8B

On the taxonomy of the Chelis glaphyra (Eversmann, 1843) species-

group, with description of a new subspecies of Chelis gratiosa (Grum-

Grshimailo, 1890) from Kyrgyzstan, brachipterous females of Chelis

kashmirica (Ferguson, 1985) and Chelis golbecki (Dubatolov, 1996),

and the checklist of the species-group (Lepidoptera: Erebidae:

Arctiinae: Arctiini)

AIDAS SALDAITIS1, 4 & ANTON V. VOLYNKIN2, 3

1 Nature Research Centre, Akademijos str., 2, LT-08412, Vilnius-21, Lithuania. E-mail: [email protected]

2 Altai State University, Lenina Avenue, 61, RF-656049, Barnaul, Russia. E-mails: [email protected];

[email protected] 3 National Research Tomsk State University, Lenina Avenue, 36, RF-634050, Tomsk, Russia

4 Corresponding author

Received 27 November 2020 │ Accepted by V. Pešić: 30 November 2020 │ Published online 1 December 2020.

Abstract

In the present paper, the checklist of the Chelis glaphyra (Eversmann, 1843) species-group with 33 new combinations

established is provided. A new subspecies of Chelis gratiosa (Grum-Grshimailo, 1890) is described from Kyrgyzstan.

Chelis gratiosa kashmirica is upgraded to the species level: Chelis kashmirica (Ferguson, 1985), stat. & comb. nov., its

brachipterous female is described and illustrated for the first time. Female genitalia of Chelis golbecki (Dubatolov,

1996), comb. nov. are described and illustrated for the first time.

Key words: Palearctia, Central Asia, new combination.

Introduction

The genus Palearctia Ferguson, 1984 was revised by Dubatolov (1996) and Saldaitis & Ivinskis (2002) and

the full checklist of the genus was provided by Dubatolov & de Vos (2010). Subsequently, Rönkä et al.

(2016) synonymized Palearctia as well as six other genera with Chelis Rambur, 1866 based on the analysis

of eight genetic markers. However, the authors did not establish new combinations for species comprised by

the genera synonymized leaving the taxonomy of the enlarged genus Chelis quite messy. In the present

paper, we provide the checklist of the species and subspecies previously assigned with Palearctia which we

consider as Ch. glaphyra species-group, with establishing new combinations for them.

Chelis gratiosa, comb. nov. is distributed in high mountain regions of Central Asia and is subdivided

into 10 subspecies. During his entomological trip to Transalai Range in southern Kyrgyzstan, Mr. V.

Ecologica Montenegrina 38: 51-62 (2020) This journal is available online at: www.biotaxa.org/em

http://dx.doi.org/10.37828/em.2020.38.7

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

52

Kartashov collected a single alive female of an unidentified subspecies of Ch. gratiosa and managed getting

eggs from it which made it possible to obtain a series of specimens of both sexes. All the specimens have

uniformly dark hindwing clearly different from those of all known subspecies of Ch. gratiosa and this

population represents an undescribed subspecies which is described below as new.

Dubatolov (1996) downgraded the taxon Ch. kashmirica to a subspecies of Ch. gratiosa due to its

external resemblance to Chelis gratiosa rupicola (Grum-Grshimailo, 1890), comb. nov. and the similar

valva shape. However, females of Ch. kashmirica are brachipterous and its male and female genitalia are

markedly different from those of Ch. gratiosa therefore we hereby upgrade it to the species level.

Additionally, Dubatolov (1996) described Chelis golbecki (Dubatolov, 1996), comb. nov. by two males and

one female but did not describe and illustrate its female genitalia. In the present paper, we describe and

illustrate the female genitalia of the species for the first time.

Material and methods

Abbreviations of the depositories used: ASV = private collection of Aidas Saldaitis (Vilnius, Lithuania);

MWM/ZSM = Museum Witt Munich in the Bavarian State Collection of Zoology (Museum Witt München /

Zoologische Staatssammlung München, Munich, Germany); SDM = private collection of Sergei Didenko

(Moscow, Russia); VKV = private collection of Vladimir Kartashov (Voronezh, Russia); WIGJ = World

Insect Gallery (Joniškis, Lithuania). Other abbreviations used: AV = genitalia slide prepared by A. Volynkin;

HT = holotype; PT = paratype.

The genitalia were dissected and mounted in euparal on microscope slides. The photos of adults were

taken using a Nikon D3100/AF-S camera equipped with a Nikkor, 18–55 mm lens. The photos of genitalia

were taken by the same camera attached to a microscope with an LM-scope adapter. All pictures were

processed using the Adobe Photoshop CC 2018® software.

Checklist of the Chelis glaphyra species-group

– glaphyra glaphyra (Eversmann, 1843), comb. nov.

– glaphyra aksuensis (O. Bang-Haas, 1927), comb. nov.

– glaphyra dublitzkyi (O. Bang-Haas, 1927), comb. nov.

– glaphyra manni (Staudinger, 1881), comb. nov.

= mannii Alpheraky, 1882

– golbecki (Dubatolov, 1996), comb. nov.

– gratiosa gratiosa (Grum-Grshimailo, 1890), comb. nov.

– gratiosa caroli (Dubatolov, 1996), comb. nov.

– gratiosa flavala (Dubatolov, 1996), comb. nov.

– gratiosa lochmatteri (Reich, 1933), comb. nov.

– gratiosa postflavida (Hampson, 1894), comb. nov.

– gratiosa rupicola (Grum-Grshimailo, 1890), comb. nov.

= glauca Staudinger, 1891

– gratiosa sarezica (Dubatolov, 2003), comb. nov.

– gratiosa sarycola (de Freina, 1997), comb. nov.

– gratiosa sergei (Dubatolov, 1996), comb. nov.

– kashmirica (Ferguson, 1985), stat. & comb. nov.

= ab. nebulosa Reich, 1932

– rasa (Saldaitis, Ivinskis & Churkin, 2000), comb. nov.

– wagneri (Püngeler, 1918), comb. nov.

– mira (Dubatolov & Tshistjakov, 1989), comb. nov.

– erschoffii erschoffii (Alpheraky, 1882), comb. nov.

– erschoffii issyka (Staudinger, 1889), comb. nov.

– erschoffii sarydzhasica (Plustsch & Dolin, 2000), comb. nov.

– erschoffii selmonsi (Böttcher, 1905), comb. nov.

– ferghana ferghana (Staudinger, 1887), comb. nov.

SALDAITIS & VOLYNKIN

Ecologica Montenegrina, 38, 2020, 51-62 53

– ferghana schottlaenderi (Strand, 1912), comb. nov.

– ferghana sussamyra (Dubatolov, 1996), comb. nov.

– gracilis gracilis (Dubatolov, 1996), comb. nov.

= miranda Plustsch & Dolin, 2000

– gracilis arcana (Plustsch & Dolin, 2000), comb. nov.

– marxi (O. Bang-Haas, 1927), comb. nov.

– mustangbhoti (Daniel, 1961), comb. nov.

– turkestana (Dubatolov, 1996), comb. nov.

– variabilis (Daniel, 1966), comb. nov.

– ammosovi (Dubatolov & Gurko, 2002), comb. nov.

– hauensteini (Kautt, 1996), comb. nov.

= altotibetana de Freina, 1997

Description of the new subspecies of Ch. gratiosa

Chelis gratiosa kartashovi ssp. nov.

(Figs 1–4, 9, 17, 24)

Type material. Holotype (Figs 1, 17): male, [Kyrgyzstan, Transalai Range] “KIRGIZIA S., Chon-Alai reg.,

Zaalaisky mountains range, Doroot-Korgon vill., Altyn-dara river, Aram-Kungei valley, H~3600m, original

female 15.vii.2018, ex. ovo 14–18.v.2019, leg. V. Kartashov”, gen. slide No.: AV5839 (Coll. ASV,

designated to WIGJ).

Paratypes: 9 males, 13 females, the same data as in the holotype, gen. slide No.: AV5840 (female) (Colls

ASV, VKV and WIGJ); 1 female, [Kyrgyzstan, Transalai Range] Kirgizia, Zaalaisky Ran., Chon-Alaisky

Distr., Altyn-Dara Riv., Aram-Kungei Val., 15–21.vii.2018, H-3600m, V. Kartashov leg. (Coll. SDM).

Etymology. The new subspecies is dedicated to prominent Russian Lepidoptera collector Vladimir

Kartashov.

Diagnosis. The new subspecies (Figs 1–4) is most similar externally to Ch. gratiosa rupicola (Fig. 5) and

also Ch. rasa (Figs 13, 14, 22, 28) and Ch. golbecki (Figs 15, 16, 23, 27), but is distinguishable by the

pinkish suffusion on the thorax and the forewing (it is absent in the three similar taxa), and the intense

blackish suffusion on hindwing making it almost uniformly dark with only a small pale spot in the

subterminal or terminal area medially (whereas in the three similar taxa, the hindwing has intense blackish or

dark brown suffusion in the subterminal area only, while the medial and basal areas are weakly suffused with

blackish scales and the discal spot is present). Additionally, the new subspecies differs from Ch. rasa and

Ch. golbecki by the longer rami of the male antenna, and from Ch. rasa by its less elongate forewing apex.

The male genital capsule of Ch. g. kartashovi (Fig. 17) is nearly identical to that of Ch. g. rupicola (Fig. 18)

and is also similar to those of Ch. g. gratiosa (Fig. 20) and Ch. g. flavala (Fig. 19), but differs from the two

latter subspecies by its more rectangular cucullus. The aedeagus of Ch. g. kartashovi differs from those of

the three similar subspecies by its markedly narrower dentate carinal plate. The vesicae of the four

subspecies display no remarkable differences. As the female genitalia of Ch. g. rupicola and Ch. g. flavala

are unknown therefore the female genitalia morphology of the new species (Fig. 24) is compared to that of

Ch. g. gratiosa (Fig. 25): Ch. g. kartashovi has a somewhat narrower ductus bursae and its appendix bursae

is longer than that of the nominate subspecies.

Description. External morphology of adults (Figs 1–4, 9). Forewing length 11-12 mm in males and 11-13

mm in females. Male antenna shortly bipectinate, female antenna serrate. Sexual dimorphism limited:

compared to males, females have forewing with more rounded apex and more reduced markings and dark

suffusion on hindwing less intense. Head dark brown with admixture of pinkish scales. Collar dark brown

with pinkish margin. Thorax blackish brown with two pinkish longitudinal stripes and pinkish margins of

tegulae. Upperside of abdomen blackish brown with ochreous brown triangular spots laterally, underside

ochreous brown; tip of abdomen densely covered in ochreous brown scales. Forewing ground color creamy

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

54

Figures 1–8. Chelis spp.: adults. Depositories of the specimens: 1, 3 in ASV; 2 in WIGJ; 4 in SDM; 5–8 in

MWM/ZSM.

SALDAITIS & VOLYNKIN

Ecologica Montenegrina, 38, 2020, 51-62 55

Figure 9. Chelis gratiosa kartashovi, paratypes. The specimens are deposited in VKV (photo by V. Kartashov).

with intense pinkish suffusion. Markings blackish brown. Subbasal spot asymmetrically V-shaped. Subbasal

area with large elongate triangular spot. Antemedial line strongly angled medially, consisting of two large

elliptical spots connected to each other by an oblique stroke and a costal dash. Postmedial line angled on the

Cu vein dilated in the cell and at costa and fused with antemedial line at anal wing margin. Subterminal line

interrupted, represented by four large triangular spots. Terminal area and anal margin blackish brown, inner

edge of terminal area strongly zigzagged. Cilia pinkish brown. Hindwing with intense dark suffusion making

it nearly uniformly blackish brown. Subterminal area with a small pale medial spot which may be protruding

to the terminal area. Cilia brown. Male genitalia (Fig. 17). Uncus large, broadly triangular with elongate and

tapered distal part pointed apically. Tuba analis broad, membranous. Tegumen short, with thick and heavily

sclerotized arms. Juxta large, X-shaped with wide arms, moderately sclerotized. Vinculum large, heavily

sclerotized, U-shaped. Valva short, its basal and medial sections wide, cucullus ca. 3 times narrower than

base of valva, rectangular with rounded corners. Sacculus narrow, heavily sclerotized. Aedeagus large,

elongate, its distal half somewhat curved dorsad; dorsal carinal plate narrow, triangular, covered in tiny

denticles. Vesica broad, C-like curved dorsad and anteriorly, with a small membranous subbasal

diverticulum dorsally, four short granulated diverticula outwardly and globular membranous subapical

diverticulum inwardly. Female genitalia (Fig. 24). Papillae anales wide, rectangular with rounded corners,

setose. Apophyses posteriores elongate, thin, with rounded tips. Apophyses anteriores fully reduced.

Postvaginal area with narrow pocket-like incision medially. Ostium bursae broad. Ductus bursae long, dorso-

ventrally flattened, somewhat tapered anteriorly, its posterior three fifth heavily sclerotized, while anterior

two fifth weakly sclerotized. Corpus bursae broad, elliptical, membranous, with area of weak granulation

posteriorly. Medial section of corpus bursae armed with two round signa bursae covered with tiny denticles.

Appendix bursae elongate with broad basal section, scobinated basally and membranous distally, situated

postero-ventrally, directed laterally to the right side.

Distribution and bionomics. The new subspecies is known from Transalai Range in southern Kyrgyzstan.

Single female was found siting on ground in an alpine grassland (at ca 3600 m altitude) where the vegetation

was damaged by grazing livestock.

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

56

Descriptions of brachipterous females

Chelis kashmirica (Ferguson, 1985), stat. & comb. nov.

(Figs 10–12, 21, 26)

Palearctia kashmirica Ferguson, 1985, Entomography, 3: 222, figs 44, 101, 102 (Type locality: [Pakistan, Azad

Kashmir] “Baltistan”).

Figures 10–16. Chelis spp.: adults. The specimens are deposited in MWM/ZSM.

SALDAITIS & VOLYNKIN

Ecologica Montenegrina, 38, 2020, 51-62 57

Figures 17–20. Chelis gratiosa sspp.: adults. Depositories of the specimens: 17 in ASV/WIGJ; 18–20 in MWM/ZSM.

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

58

Figures 21–23. Chelis spp.: adults. The specimens are deposited in MWM/ZSM.

SALDAITIS & VOLYNKIN

Ecologica Montenegrina, 38, 2020, 51-62 59

Figures 24–28. Chelis spp.: adults. Depositories of the specimens: 24 in ASV; 25–28 in MWM/ZSM.

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

60

Material examined. 26 males, 2 females, Pakistan, Kashmir, Himalaya Mts., Kashmir Deosai pass., 4250–

4800 m, 5.VII.2000, leg. Z. Varga & G. Ronkay, gen. slide Nos.: ZSM Arct. 2019–915 (male), ZSM Arct.

2019–916 (female) (prepared by Volynkin) (MWM/ZSM).

Diagnosis. Male of the species is smaller than those of subspecies of Ch. gratiosa. In the male genital

capsule of Ch. kashmirica, the uncus is narrower medially, the vinculum is rather V-shaped (while that of

Ch. gratiosa is U-shaped), the juxta is conspicuously shorter and narrower, the valva is less elongate and the

cucullus is noticeably smaller than those structures of Ch. gratiosa. The aedeagus of Ch. kashmirica is much

shorter than in Ch. gratiosa. In the aedeagus vesica of Ch. kashmirica, the medial diverticulum is C-like

curved (in Ch. gratiosa, it is globular), and the distal section is conspicuously shorter than that of Ch.

gratiosa. Female of Ch. kashmirica is brachipterous and has very short and narrow wings with strongly

elongate and pointed apex, while all the subspecies of Ch. gratiosa have normal winged females. In the

externally similar Ch. golbecki, the both female wings are much longer and broader than those of Ch.

kashmirica and the forewing apex is more rounded. The female genitalia of Ch. kashmirica differ from those

of Ch. gratiosa by the much narrower posterior section of the corpus bursae and the conspicuously smaller

signa bursae.

Figure 29. Kyrgyzstan, Transalai Range, Aram-Kungei valley, the type locality of Chelis gratiosa kartashovi (photo by

V. Kartashov).

Description of female. External morphology (Figs 11, 12). Forewing length 8 mm. Antenna short, thick,

shortly serrate whitish basally and medially and blackish apically. Body robust, brown. Forewing narrow,

elongate, with nearly parallel costal and anal margins and pointed apex. Forewing ground color and cilia pale

brown. Pattern reduced to two elongate blackish brown spots in the cell connected to each other by thin and

diffuse line, longitudinal blackish stroke in subbasal area, intense brown suffusion along the anal margin, and

two small diffuse dots in subterminal area. Hindwing pale brown with intense dark brown suffusion basally

SALDAITIS & VOLYNKIN

Ecologica Montenegrina, 38, 2020, 51-62 61

and along the anal margin, with three diffuse dark brown spots in terminal area. Cilia brown. Female

genitalia (Fig. 26). Papillae anales wide, trapezoid with rounded corners, setose. Apophyses posteriores

elongate, narrow. Apophyses anteriores fully reduced. Postvaginal area of 8 th sternite sclerotized laterally,

with narrow and short medial pocket-like incision. Ostium bursae broad. Ductus bursae long, narrow, dorso-

ventrally flattened, narrowed subposteriorly and dilated around ostium, sclerotized medially and posteriorly

and membranous anteriorly. Corpus bursae ovoid, membranous, with weak spinulose scobination at the base

of appendix bursae and two smalls round signa bursae medially. Appendix bursae broadly conical,

membranous, situated postero-ventrally, directed to the left side.

Distribution. The species is known from Pakistan (Azad Kashmir) (Ferguson 1985) and North India

(Himachal Pradesh) (Reich 1932, as Micrarctia glaphyra ab. nebulosa Reich, 1932).

Figure 30. Larvae of Chelis gratiosa kartashovi (photo by V. Kartashov).

Chelis golbecki (Dubatolov, 1996), comb. nov.

(Figs 15, 16, 23, 27)

Palearctia (Palearctia) golbecki Dubatolov, 1996; Neue entomologische Nachrichten, 37: 19, figs 10a 10b (Type

locality: Kyrgyzstan, Kyrgyz Ala-too Range] “Prov. Syr-Darja, Jug. Alexandrinum, Loc. Tshai-Sandyk”).

Material examined. 13 males, Kyrgyzstan, Kyrgyz Mts., Tjuz-Ashuu Pass, 3300m, 12–15.VII.2000, E.

Rut'jan leg. (MWM/ZSM); 4 males, 8 females, the same locality and collector, 3150–3300m, 27.VII.2000,

gen. slide Nos.: ZSM Arct. 2019–912 (male) and ZSM Arct. 2019–913 (female) (prepared by Volynkin)

(MWM/ZSM); 1 male, Suusamyr valley, Ala-Bel pass, 10–20.VII.2002, 3300m (MWM/ZSM); 2 males, 1

female, the same locality, 3200m, 20–30.VII.2002 (MWM/ZSM); 1 male, Kyrgyzstan, Otmek pass., 3000–

3300m, 5.VIII.1999 (MWM/ZSM).

Description of female genitalia (Fig. 27). Papillae anales wide, trapezoid with rounded corners, setose.

Apophyses posteriores elongate, thin. Apophyses anteriores fully reduced. Postvaginal area of 8th sternite

ON THE TAXONOMY OF THE CHELIS GLAPHYRA SPECIES-GROUP

62

sclerotized laterally, with short but broad medial pocket-like triangular incision. Ostium bursae broad, with

membranous margins. Ductus bursae elongate, narrow, dorso-ventrally flattened, sclerotized, with

membranous anterior end. Corpus bursae ovoid, membranous, with dense spinulose scobination at the base

of appendix bursae and two medium sized rounded signa bursae covered in tiny denticles. Appendix bursae

elongate, tubular, membranous, situated postero-ventrally, directed to the left side.

Distribution. The species is known from Kyrgyz Ala-too (Dubatolov 1996) and Talas Ala-too Ranges in

Kyrgyzstan.

Acknowledgements We express our sincere thanks to Dr. Axel Hausmann, Dr. Wolfgang Speidel and Mr. Ulf Buchsbaum (ZSM,

Munich, Germany) for their kind assistance provided during the second author’s studies at the MWM/ZSM

collection. We are also grateful to Mr. V. Kartashov (Voronezh, Russia) and Mr. S. Didenko (Moscow,

Russia) for multiple assistance in preparing this article.

References

Dubatolov, V.V. (1996) A review of the genus Palearctia Ferguson (on the systematics of the genus

Micrarctia Seitz s. l., Part 3) (Lepidoptera: Arctiidae). Neue entomologische Nachrichten, 37, 9–37.

Dubatolov, V.V. & de Vos, R. (2010) Tiger-moths of Eurasia (Lepidoptera, Arctiidae). Neue entomologische

Nachrichten, 65, 1–106.

Saldaitis, A. & Ivinskis, P. (2002) Some notes about species of the genus Palearctia Fergusson, 1984.

Atalanta, 33 (1/2), 157–172, 232–235.

Rönkä, K., Mappes, J., Kaila, L. & Wahlberg, N. (2016) Putting Parasemia in its phylogenetic place: a

molecular analysis of the subtribe Arctiina (Lepidoptera). Systematic Entomology, 41, 844–853.

https://doi.org/10.1111/syen.12194

Ferguson, D.C. (1985) Contributions toward reclassification of the world genera of the tribe Arctiini, Part 1 –

Introductrion and a revision of the Neoarctia-Grammia group (Lepidoptera: Arctiidae; Arctiinae). The

Entomography. An Annual Review for Biosystematics, 3, 181–275.

Reich, P. (1932) Neue Arctiiden. Internationale entomologische Zeitschrift, 26 (22), 233–238. [in German]