Research Article |
Corresponding author: Nadine Dupérré ( nadine.duperre@uni-hamburg.de ) Academic editor: Martin Husemann
© 2018 Nadine Dupérré, Danilo Harms.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Dupérré N, Harms D (2018) Raising the Dead: Rediscovery and redescription of some lost spider types (Araneae) described by Eugène Simon. Evolutionary Systematics 2: 1-20. https://doi.org/10.3897/evolsyst.2.24122
|
In this paper, we are redescribing type material from the Zoological Museum in Hamburg that was thought to be lost. These specimens were described in 1902 by Eugène Simon from material collected in Southern Patagonia and Fireland but the species were subsequently considered nomina dubia, or simply not considered at all. The rediscovery of this material leads to the revalidation of two genera and four species. The genera Clitistes and Zilephus are reinstated and the species Clitistes velutinus Simon, 1902 (Dictynidae), Zilephus granulosus Simon, 1902, Minyriolus australis Simon, 1902 (both Linyphiidae), and Lycosa michaelseni Simon, 1902 (Lycosidae) are redescribed. To avoid further confusion, we designate lectotypes for: Linyphiidae: Minyriolus australis Simon, 1902, Gongylidiellum uschuaiense Simon, 1902, Neriene fuegiana Simon, 1902, Clitistes velutinus Simon, 1902, Zilephus granulosus Simon, 1902; Amphinectidae: Rubrius radulifer Simon, 1902; Hahniidae: Hahnia michaelseni Simon, 1902, Bigois antarctica Simon, 1902 and Lycosidae: Lycosa michaelseni Simon, 1902. For all prior nomina dubia and newly designated lectotypes, the type specimens are re-described and properly illustrated for the first time.
Dictynidae , Linyphiidae , Lycosidae , nomen dubium, type catalogue, Wilhelm Michaelsen
Eugène Simon (1848–1924) is considered by many arachnologists as the prime father of systematic spider research and still widely praised as the most prolific spider taxonomist of all times. Simon worked at the Muséum national d´Histoire Naturelle in Paris where most of his type specimens are deposited. Determination of type material by subsequent researchers has often been problematic, mainly because Simon did not declare type material in his original descriptions. Simon also described material from other collections but rarely stated the type depository, which was common practice back then but is an essential element of taxonomic descriptions today. Simon is not to be blamed for his approach because taxonomic standards were very different back then. It is perhaps surprising that type material described by Eugène Simon was recently rediscovered at the Zoological Museum in Hamburg (
In 1890, a commission was set up to plan an expedition to Southern Patagonia and Fireland, but the expedition was delayed due to political instability in Chile. Wilhelm Michaelsen left Hamburg in late July 1892 and returned to Hamburg on the 10th September 1893 with great success (
After the return of the expedition, the arachnid specimens were integrated into the invertebrate collections of the Zoological Museum in Hamburg and forwarded to the taxonomic authorities for identification and description. It was the then-director of the museum Karl Kraepelin who invited Eugène Simon to participate in the analysis of the the specimens collected of the expedition. Eugène Simon was already a well-established arachnologist and had published on arachnids from Patagonia, Cape Horn and Terre-de-Feu (
Unfortunately, Simon did not mention the number of specimens he examined and refrained from illustrating the specimens. Consequently, it has been somewhat difficult for subsequent arachnologists to recognise the species but also to determine their status (holotype or lectotypes).
The present paper re-analyses part of the spider material presented by Simon in 1902; only the species that were designated as nomina dubia or species with problematic holotype designation are re-analysed. The present paper aims to i) re-establish “lost” genera and species that were wrongly declared to be invalid or nomina dubia; ii) designate lectotypes from the syntype series; and iii) illustrate and describe in necessary details the newly designated lectotypes.
All specimens are deposited at the Zoological Museum (
The taxonomy follows the
Minyriolus australis Simon, 1902: 15 (as Minyriolus (?) australis n. sp., description female)
Coll. Mich. 130. Süd-Feuerland, Uschuaia, Süßwassersee auf der Halbinsel, 19.XII.92.
A. Photo of Prof. J. Wilhem Michaelsen. B. Examples of labels with Eugène Simon numbering system. C. Examples of locality labels.
♀. long. 1 mm.
Minyriolus (?) australis n.sp., Nr. 16.
130. Uschuaia; Süsswasser-See auf d. Halbinsel. Coll. Michaelsen. 19.XI.92.
In 2007, Miller declared this species a nomen dubium: “The otherwise Palearctic genus Minyriolus was represented in the Neotropics by M. australis
Lectotype ♀ designated here (
Female (lectotype). Total length: 1.23; cephalothorax length 0.55; cephalothorax width: 0.38. COLORATION: (based on original description, translated from Latin): “cephalothorax bright olive-brown with thin black margin, ocular area blackish. Sternum black. Abdomen light brown. Legs pale yellow-reddish”. CEPHALOTHORAX: Longer than wide (Fig.
ABDOMEN: Oval (Fig.
Male. Unknown.
Argentina: Uschuaia.
The genus Minyriolus is composed of three Palearctic species (
Linyphiidae, Minyriolus australis Simon, 1902.
Gongylidiellum uschuaiense Simon, 1902: 16. (as Gongylidiellum (?) uschuaiense n. sp., description female).
♀. long. 0,8 mm.
Gongylidiellum (?) uschuaiense n. sp. Nr. 17.
142 Uschuaia, 14.XII.92.
In 2007, Miller synonymised Gongylidiellum uschuaiense
Lectotype ♀ designated here (
Female (lectotype). Total lenght: 1.31; cephalothorax length: 0.59; cephalothorax width: 0.42. COLORATION: “pale yellow-reddish, abdomen light gray”. CEPHALOTHORAX: Longer than wide (Fig.
Male. Unknown.
Argentina: Uschuaia.
Species from the genus Gongylidiellum are found in the Old World, except for Gongylidiellum uschuaiense. It is highly likely that this species does not belong in this genus, as already indicated by Simon when he gave the species name with a “?”. Further work on Argentinian Linyphiidae is necessary before the species can be placed more accurately.
Linyphiidae, Gongylidiellum uschuaiense Simon, 1902.
Neriene fuegiana Simon, 1902: 17 (as Neriene fuegiana n. sp., description female).
Oedothorax fuegianus Petrunkevitch, 1911: 262 (transferred female).
Oedothorax fuegianus Miller, 2007: 244, f. 186C (female illustration, misplaced in this genus).
♀. long. 2,7 mm.
Neriene fuegiana n. sp. Nr. 20.
187. Feuerld. S.K.wstl.v.Kp.S.Pio, Coll. Michaelsen. 27.XII.92.
This species was transferred by
Lectotype ♀ designated here (
Female (lectotype). Total length: 1.74; cephalothorax length: 0.59; cephalothorax width: 0.48. COLORATION: “cephalothorax pale yellow, eyes with thin black ring. Abdomen dorsally white, both sides with wide dark indistinct pattern, median line complete, with 4 or 5 slightly brownish arched transverse lines, ventrally pale reddish-brown.” CEPHALOTHORAX: Longer than wide (Fig.
Male. Unknown.
Chile, Kap San Pio.
Linyphiidae, Oedothorax fuegiana Simon, 1902.
Neriene michaelseni Simon, 1902: 18 (as Neriene michaelseni n. sp., description female).
Coll. Mich. 140. Süd-Feuerland, Uschuaia, Wald, unter vermodernden Baumstümpfen; 30.X.92. Coll. Mich. 141. Süd-Feuerland, Uschuaia, Land; 14.XI.92.
♀. long. 2,4 mm.
Neriene michaelseni n. sp. Nr. 22.
Locality label 1. 140. Uschuaia; Wald. Coll. Michaelsen 30.X.92.
Locality label 2. 141. Uschuaia. Coll. Michaelsen 14.XI.92.
Syntypes 1♂ penultimate (
Neriene michaelseni
Linyphiidae, Laminacauda plagiata (Tullgren, 1901).
Simon described the genus Clitistes and placed it in the family Linyphiidae. The description is rather short: he mentions that the genus is closely related to Clitolyna but differs by the eye arrangement, shorter clypeus, abdominal setae and shorter palps. The genus Clitolyna was synoymised by
Clitistes velutinus Simon, 1902: 20 (as Clitistes velutinus n. sp., description female).
♀. long. 2,5 mm.
Clitistes velutinus n. sp. Nr. 25.
178. Navarin, Puerto Toro, Wald, Coll. Michaelsen 19.XII.92.
The female specimen deposited in the
Female (lectotype). Total length: 2.89; cephalothorax length: 1.05; cephalothorax width: 0.96.
Lectotype ♀, designated here (
COLORATION (based on original description, translated from Latin): “cephalothorax dark brown, pars cephalica slightly paler, pars thoracica with thin black margin. Abdomen dorsally dark yellow-bluewish, medially with wide, darker band, anterior margin paler, in the middle part, three transverses, angular arches, apically with small spots; ventrally, dark yellow. Femur apically with brown ring”. CEPHALOTHORAX: Pyriform, longer than wide (Fig.
Male. Unknown.
Chile, Puerto Toro.
Transferred to Dictynidae, Clitistes velutinus Simon, 1902
Zilephus Simon, 1902: 22 (Gen. Zilephus nov.)
Here again, Simon´s description of the genus is rather short and he mentions that the genus resembles Microneta but differs by eye arrangment, clypeus and the granulation of the cephalothorax.
♂.long. 2 mm.
Zilephus granulosus n. sp. Nr. 27.
140. Uschuaia, Wald. Coll. Michaelsen. 30.X.92.
The data from the labels found with the
Lectotype ♀ designated here (
Female (lectotype). Total length: 2.32; cephalothorax length: 0.87; cephalothorax width: 0.71. COLORATION: (from original description, translated from Latin): “cephalothorax blackish or dark olive. Abdomen dorsally white, ornated with median broad band bluntely trilobate, and apically pointed, with black and white spots obliquely paired, ventrally reddish brown. Femur yellow, tibia and metatrsi apically with small brown ring; tibia IV with medially and apically small brown ring.” CEPHALOTHORAX: Pyriform, longer than wide (Fig.
Male. Unknown.
Argentina: Uschuaia
Linyphiidae, Zilephus granulosus Simon, 1902.
Theridion ventrosum Nicolet, 1849: 536 (description female).
Theridion recurvatum Tullgren, 1901: 191 (description juvenile).
Enoplognatha triangulifera Simon, 1902: 14 (as Enoplognatha triangulifera n. sp., female description).
Anelosimus recurvatus Levi, 1962: 12 (transferred female from Theridion, synonymy).
Anelosimus recurvatus Levi, 1963: 45, f. 49-52 (female, description male).
Anelosimus ventrosus Levi, 1967: 13 (transferred female from Theridion, synonymy male).
Anelosimus recurvatus Schiapelli & Gerschman, 1974: 86, f. 36–38 (male and female redescription).
Selkirkiella ventrosa Agnarsson, 2004: 476 (transferred male and female from Anelosimus).
Coll. Mich. 165. Süd-Feuerland, Harberton Harbour (Puerto Bridges); 10. I. 93. Coll. Mich. 187. Süd-Feuerland, Küstenstrich-Eben westlich von Kap San Pio; 27.xii.92.
♀ long. 4mm.
Enoplognatha triangulifera n. sp. Nr. 15.
Locality label 1 (with 2 ♀). 165. Puerto Bridges; Wald. Coll. Michaelsen. 10.I.93.
Locality label 2 (with 2 ♀). 187. Feuerld. S.K.wstl.v.Kp.S.Pio. Coll. Michaelsen. 27.xii.92.
Syntypes 4♀ (
Theridiidae, Selkirkiella ventrosa (Nicolet, 1849)
Coll. Mich. 75. Süd-Patagonien, Punta Arenas, unter Steinen und Baumstämmen; IX. 92.
♀. long. 13 mm.
Tomopisthes Kraepelini n. sp. Nr. 49.
75. Magelh. Str., Punta arenas; Coll. Michaelsen. IX.92.
Syntype ♀ (
This species was synonymised under Sanogasta approximata (
Anyphaenidae, Sanogasta approximata (Tullgren, 1901).
Coll. Mich. 81. Süd-Patagonien, Punta Arenas; X.-XII.92. (H. Michelsen leg.).
♀. long. 6–7 mm.
Tomopisthes conspersus n. sp. Nr. 52.
81. Magelh. Str., Punta Arenas; Coll. Michaels. Herbst 92 (Michaelsen l.).
Syntype ♀ (
Ramírez (2003:154) synonymised Tomopisthes conspersus based on a specimen from MHNP 21816, female holotype. The specimen in the MHNP is probably part of the type series but Ramírez did not formely designate a lectotype. The holotype assumption is invalid and the
Anyphaenidae, Sanogasta maculosa (Nicolet, 1849).
Tomopisthes
injucundus
Coll. Mich. 80. Süd-Patagonien, Punta Arenas; 25.II.93. Coll. Mich. 141. Süd-Feuerland, Uschuaia; 14.XI.92 Coll. Mich. 165. Süd-Feuerland, Harberton Harbour (Puerto Bridges), Wald, 10.I.93. Coll. Mich. 174. Süd-Feuerland, Arch. Isl. Picton, Banner Cove, 26.XII.92. Coll. Mich. 193. Süd-Feuerland, Puerto Pantalon, 2.I.93.
♀. long. 6 mm.
Tomopisthes injucundus n. sp. Nr. 53.
Locality label (with 2♀). 80. Punta Arenas, Mich. 25.II.93.
Locality label (with 2♀, 2♂). 141. Uschuaia, Coll. Michaelsen. 14.XI.92.
Locality label (with 2♀). 174. Isl. Picton, Coll. Michaelsen. 26.XII.92.
Locality label (with 5♀). 165. Puerto Bridges, Coll. Michaelsen. Wald, 9.I.93.
Locality label (with 1♀). 193. Feuerland, Puerto Pantalon; Coll. Michaelsen. 2.I.93.
Paralectotypes 2♂, 12♀ (
Ramírez (2003: 154) synonymised Tomopisthes injucundus under Sanogasta maculosa (Nicolet, 1849) based on female lectotype, three females and one male paralectotypes that he designated from specimens from “Tierre del Fuego, MHNP 21782 (the male paralectotype belongs to a different, presumably undescribed Sanogasta species)”. The specimen in the Paris Museum are probably part of the type series and were designated as such by Ramírez. Therefore, the
Anyphaenidae, Sanogasta maculosa (Nicolet, 1849).
Tomopisthes
modestus
Female, Coll. Mich. 76. Süd-Patagonien, Punta Arenas, unter Baustämmen; 15.III.93.
♀. long. 5 mm.
Tomopisthes modestus n. sp. Nr. 55.
76. Magelh. Str., Punta arenas; IX.92.
Syntype ♀ (
Ramírez (2003: 154) synonymised Tomopisthes modestus under Sanogasta maculosa (Nicolet, 1849) based on female holotype from Chile, Punta Arenas, IX.1892, Michaelsen coll., examined in MHNP. The specimen in the Paris Museum is probably part of the type series but Ramírez did not designate a lectotype. The holotype assumption is invalid and the
Anyphaenidae, Sanogasta maculosa (Nicolet, 1849).
Rubrius radulifer Simon, 1902: 36 (Rubrius radulifer n. sp., description female and male).
Calacadia radulifera Exline, 1960: 618 (Transferred female from Rubrius).
Coll. Mich. 30. Chile, Putabla bei Valdivia; 20.IV.93.
Coll. Mich. 38. Chile, Valdivia; 31.III.93.
Rubrius radulifer n. sp., Nr. 58.
Locality label 1 (with ♀). 30. Valdivia, Putabla; Coll. Michaelsen. 20.IV.93.
Locality label 2 (with ♂). 38. Valdivia. Coll. Michaelsen. 31.III.93.
♀. long. 12-15 mm. ♂. 10 mm
The
Lectotype ♂ designated here, paralectotype ♀ (juvenile) (
Male (Lectotype). Total length: 6.04; cephalothorax length: 3.36; cephalothorax width: 2.38. COLORATION (from original descrition, translated from Latin): “Cephalothorax reddish-yellow, frontal region of pars cephalica darker, with dark oblique broad band on both sides, with very intricate dentate V-form pattern, pars thoracica with marginal line slightly black, radiating line barely visible. Abdomen, yellow-reddish, with brown reticulate pattern, middle band pale with dentate pattern, border dark and sinuous. Legs yellow-reddish, femora and tibiae with two dark-olive rings, metatarsi and tarsi anteriorly darker”. CEPHALOTHORAX: Pear-shaped, longer than wide (Fig.
Chile, Valdivia.
Desidae, Calacadia radulifera Exline, 1960.
Rubrius paganus Simon, 1902: 37 (Rubrius paganus n. sp, description female and male).
Rubrius annulatus Lehtinen, 1967: 263, f. 159 (female illustration, female and male synonymy).
Rubrius annulatus Roth, 1967: 329, pl. 52, f. 13 (female redescription).
Rubrius paganus Roth, 1967: 332, pl. 52, f. 19-20 (female and male redescription).
Coll. Mich. 28. Chile, Chamil-chamil bei Valvidia; 23.IV.93.
♀. long. 10-12 mm. ♂. 8 mm.
Rubrius paganus n. sp. Nr. 59.
28. Valvidia, Chamilchamil. Coll. Michaelsen. 23.IV. 93.
Paralectotypes juveniles 1♂1♀ (
Rubrius paganus
Amaurobiidae, Rubrius annulatus F.O. Pickard-Cambridge, 1899.
Hahnia michaelseni Simon, 1902: 39 (Hahnia Michaelseni n. sp. description female).
Hahnia michaelseni Vellard, 1958: 136, f. 18-25 (female, description male, doubtful identification).
Hahnia michaelseni Lehtinen, 1967: 455, f. 376 (female).
Hahnia michaelseni Schiapelli & Gerschman, 1974: 89, f. 34–35 (female, doubtful identification).
Coll. Mich. 187. Süd-Feuerland, Küstenstrich-Eben westlich von Kap San Pio; 27.XII.92.
Hahnia Michaelseni n. sp., Nr. 61.
187. Feuerld.S.K. wstl.v. Kp. S. Pio. Coll. Michaelseni. 27.XII.92.
In 1958, Vellard described a female and a male based on specimens from Rusfin, that he believed was the species Hahnia michaelseni but he mentions (p.137) that he was not able to see the type and that there were some significant differences: “les différences de formule oculaires sont peut-être plus significatives. N’ayant pu comparer nos examplaires au type de Hambourg…. Il est bien difficile d’apprécier la valeur de ces différences”. Lethinen (1967: fig. 376) also illustrated a female but did not mention where the specimen came from. Finally, in 1974 Schiapelli and Gerschman redescribed the species based on a single female from Puerto San Carlos. Multiple authors (Vellard 1949, Lethinen 1967, Schiapelli and Gerschman 1974) have redrawn this species but nobody seems to have re-examined the type and the only illustration that matches with the specimens held at the
Lectotype ♀ designated here (
Female (lectotype). Total length: 2.26; cephalothorax length: 1.04; cephalothorax width: 0.78. COLORATION: (from original description, translated from Latin) “cephalothorax smooth, dark olive-brown, thoracic part with thin, barely distinct black border. Abdomen oval, dark gray, decorated anteriorly with longitudinal line, posteriorly with four transversal lines, slightly curved”. CEPHALOTHORAX: Pear-shaped, fovea longitudinal (Fig.
Male. Unknown.
Kap San Pio.
Hahniidae, Hahnia michaelseni Simon, 1902.
Bigois antarctica Simon, 1902: 40 (Bigois antarctica n. sp., description female).
Bigois antarctica Birabén, 1957: 4, f. 1-8 (female, description male).
Amaloxenops translata Schiapelli & Gerschman, 1959: 132 (new name for Birabén’s material, believed misidentified).
Intihuatana antarctica Lehtinen, 1967: 240, f. 372-373 (transferred male and female to Intihuatanan n. gen.).
Coll. Mich. 150. Süd-Feuerland, Uschuaia, unter Steinen und zwischen Steingeröll; 15.XI.92. Coll. Mich. 178. Süd-Feuerland, Arch., Isl. Navarin, Puerto Toro, Wald; 19.XII.92. Coll. 193. Süd-Feuerland, Puerto Pantalon; 2.I.93.
Bigois antarctica Simon, 1902. A. Female, habitus, dorsal view. B. Female habitus, ventral view. C. Male habitus, dorsal view. D. Epigynum, ventral view. E. Palp, dorsal view. F. Bulb and embolus, ventral view. Abbreviations: co: copulatory openings, e: embolus, pp: patellar process, tp: tibial basal process.
♀. long. 1,3 mm.
Bigois antarctica n. sp. Nr. 62.
Locality label vial 1 (with 1♀). 193. Feuerland, Puerto Pantalon; Coll. Michaelsen. 2.I.93.
Locality label vial 2 (with 1♂). 150. Uschuaia 15.XI.92.
Locality label vial 3 (with 1♀). 178. Navarin, Puerto Toro, Wald. Coll. Michaelsen. 19.XII.92.
Lectotype 1♀ designated here, paralectotypes 1♂, 1♀(
In 1957, Birabén redescribed the species based on a female topotype from Uschuaia and a male allotype from Bariloche, 1600 km away from Uschuaia. Schiapelli & Gerschman published a paper on the characters of the genus Bigois (1959) and did not agree with Birabén’s redescription. Based on his illustration and description, they suggested that his species should be placed in Amaloxenops and named it A. translata Schiapelli & Gerschman 1959.
Female (lectotype). Total length: 1.5; cephalothorax length: 0.68; cephalothorax width: 0.53. COLORATION: (from original description, translated from Latin): “pale yellow-reddish, abdomen white opaque”. CEPHALOTHORAX: Pear-shape, longer than wide (Fig.
Male. (paralectotype). Total length: 1.52; cephalothorax length: 0.73; cephalothorax width: 0.55.
CEPHALOTHORAX: As in female (Fig.
Argentina; Uschuaia, Chile; Puerto Toro and Puerto Pantalon.
Hahniidae, Intihuatana antarctica (Simon, 1902).
Lycosa michaelseni Simon, 1902: 42 (Lycosa Michaelseni n. sp. description female).
Alopecosa michaelseni Mello-Leitão, 1947: 263 (transferred to Alopecosa).
Alopecosa michaelseni Casanueva, 1980: 54 (nomina dubia).
Coll. Mich. 85. Süd-Patagonien, Punta Arenas, Wald; 18. X. 92. Coll. Mich. 179. Süd-Feuerländ. Archipel, Isl. Navarin, Puerto Toro, Wald; XI. 92 (F. Delfin leg.).
Lycosa Michaelseni n. sp. Nr. 68.
85. [Mag. Hb]. Punta Arenas, Wald. Coll. Michaelsen. 18. X. 92.
Lectotype ♀ designated here (
Lycosa michaelseni was transferred by
Female (lectotype). Total length: 10.97; cephalothorax length: 4.73; cephalothorax width: 3.48. COLORATION: (from original description, translated from Latin): “cephalothorax with black forehead, covered by yellow-grayish hairs, with a submarginal sinuous line on both sides. Abdomen black with dark brown hair, intermingled with a few white hairs, longitudinal lanceolate concolor band, posteriorly with spots in two rows, scarcely marked”.
CEPHALOTHORAX: Longer than wide, not elevated (Fig.
Male. Unknown.
Chile: Punta Arenas.
Aleopcosa is a large genus of wolf spiders with currently 161 desribed species that are distributed in Eurasia (75% of species), and a few (9%) with a Holarctic or Palearctic distribution (Blagoev & Dondale 2014). Only seven species occur in South America (Venezuela, Ecuador and Argentina) and probably do not belong to that genus but we retain this species in Alopectosa, emphasizing the need for revision.
Lycosidae, Alopecosa michaelseni (Simon, 1902).
Labelling and curatorial order of the
Of the 29 spider species described by Simon in his 1902 paper, some subsequent authors correctly assigned the types and mentioned that the specimens were deposited in the Hamburg Museum (
In the case of Tomopisthes, we refrained from designating lectotypes since Ramírez (pers. comm.) commented that Tomopisthes is under taxonomic revision and that there are numerous, closely related species, that could be under the same species name. Two species described by
The current study showcases – once more – the difficulties in working with very old type collections. Whilst the documentation of old and pale types that are locked away in large collections seems little adventurous, it provides the foundation for any work to come and taxonomic chaos and redundancy results if that study is not being done. We all have to face the problem of revising and looking for types all over the world and often consider this as wasted time but documenting, redescribing and illustrating types is of significant importance to ensure taxonomic stability, as well as offering valuable historical, biological and biogeographical data.
We sincerely thank Martina Mistera (CeNak) for helping us to find the picture of Wilhelm Michaelsen and Dr. Martin Ramírez for commenting on the taxonomic status of Tomopisthes. We also thank the referees for constructive criticism on an earlier draft of this manuscript. This study was made possible in part through the help of the “Society of the friends and donors of the Zoological Museum in Hamburg” who kindly provided financial support for a partial inventory of the