Proposal for the Synonymy of Some South-East Asian Whip Scorpion Genera (Arachnida, Uropygi, Thelyphonida)

ARTÍCULO:

Proposal for the Synonymy of some South-East Asian Whip Scorpion Genera (Arachnida, Uropygi, Thelyphonida)

Joachim Haupt

Abstract: The South-East Asian whip scorpion fauna has never been systematically re- vised, with numerous new species and genera added. There are several opi- nions what kind of character may serve to distinguish different genera, but it is relatively sure that certain characters cannot serve for this purpose, for exam- ARTÍCULO: ple the number of 'ommatoids' on the metasoma. Therefore genera based on this character are proposed to be removed. Two other genera (Minbosius Proposal for the Synonymy of some Speijer, 1933a, b and Ginosigma Speijer, 1933a, 1936) are also critically re- South-East Asian Whip Scorpion viewed. Genera (Arachnida, Uropygi, Key words: Uropygi, Abaliella n. syn., Chajnus n. syn., Ginosigma, Minbosius n. syn., Thelyphonida) Tetrabalius n. syn., Thelyphonus

Taxonomy: Thelyphonus ambonensis (Speijer, 1933) n. comb. for Tetrabalius am- Joachim Haupt boensis Speijer, 1933. Thelyphonus borneensis (Speijer, 1933) n. comb. for Institut of Ecology, Tetrabalius borneensis Speijer, 1933. Thelyphonus borneonus Haupt, 2009 n. nov (replacement name for Tetrabalius borneensis (Speijer, 1933b: 72-73)) n. Technical University Berlin, Germany. comb. Thelyphonus dammermanni (Speijer, 1933b) n. comb. for Tetrabalius Current address: dammermanni Speijer, 1933. Thelyphonus dicranotarsalis (Rowland, 1973a) n. Gluckweg 6, D-12247 Berlin. comb. for Abaliella dicranotarsalis Rowland, 1973, Thelyphonus florensis (Spei- [email protected] jer, 1933) n. comb. for Tetrabalius florensis Speijer, 1933. Thelyphonus gertschi (Rowland, 1973) n. comb. for Abaliella gertschi Rowland, 1973. Thelyphonus kopsteini (Speijer, 1933b) n. comb. for Minbosius kopsteini Speijer, 1933. Thely- phonus luzonicus Haupt, 2009 nomen novum (replacement name for Abaliella manilana (Kraepelin, 1900) n. comb. Thelyphonus manilanus C.L. Koch, 1843 n. comb. for Minbosius manilanus Speijer. Thelyphonus nasutus (Thorell, 1888) n. comb. for Tetrabalius nasutus Thorell, 1888.Thelyphonus rohdei (Kraepelin, 1897) n. comb. for Abaliella rohdei (Kraepelin, 1897). Thelyphonus samoanus (Kraepelin, 1897) n. comb. for Abaliella samoana (Kraepelin, 1897). Thelyphonus seticauda Doleschall, 1857 n. comb. for Tetrabalius seticauda (Doleschall, 1857). Thelyphonus willeyi (Pocock, 1898) n. comb. for Abaliella willeyi (Pocock, 1898).

Revista Ibérica de Aracnología ISSN: 1576 - 9518. Propuesta de sinonimia de algunos géneros de vinagrillos del Sudeste Dep. Legal: Z-2656-2000. Vol. 17 Asiático (Arachnida, Uropygi, Thelyphonida) Sección: Artículos y Notas. Pp: 13 − 20. Resumen: Fecha publicación: 30 Noviembre 2009 La fauna de vinagrillos del Sudeste Asiático, a la que se han añadido numero- sas nuevas especies y géneros, nunca ha sido revisada sistemáticamente. Hay Edita: diversas opiniones sobre qué clase de carácter puede ser utilizados para dis- Grupo Ibérico de Aracnología (GIA) tinguir diferentes géneros, pero es relativamente seguro que ciertos caracteres Grupo de trabajo en Aracnología no pueden ser utilizado para este propósito, por ejemplo el número de 'omma- de la Sociedad Entomológica toides' en el metasoma. Por lo tanto, los géneros basados en ese carácter son Aragonesa (SEA) propuestos para ser eliminados. Otros dos géneros (Minbosius Speijer, 1933a, Avda. Radio Juventud, 37 b y Ginosigma Speijer, 1933a, 1936) son también objeto de una revisión crítica. 50012 Zaragoza (ESPAÑA) Palabras clave: Uropygi, Abaliella n. syn., Chajnus n. syn., Ginosigma, Minbosius n. Tef. 976 324415 syn., Tetrabalius n. syn., Thelyphonus Fax. 976 535697 Taxonomy: Thelyphonus ambonensis (Speijer, 1933) n. comb. for Tetrabalius am- C-elect.: [email protected] boensis Speijer, 1933. Thelyphonus borneensis (Speijer, 1933) n. comb. for Tetrabalius borneensis Speijer, 1933. Thelyphonus borneonus Haupt, 2009 n. Director: Carles Ribera nov (replacement name for Tetrabalius borneensis (Speijer, 1933b: 72-73)) n. C-elect.: [email protected] comb. Thelyphonus dammermanni (Speijer, 1933b) n. comb. for Tetrabalius dammermanni Speijer, 1933. Thelyphonus dicranotarsalis (Rowland, 1973a) n. Indice, resúmenes, abstracts comb. for Abaliella dicranotarsalis Rowland, 1973, Thelyphonus florensis (Spei- vols. publicados: jer, 1933) n. comb. for Tetrabalius florensis Speijer, 1933. Thelyphonus gertschi http://entomologia.rediris.es/sea/ (Rowland, 1973) n. comb. for Abaliella gertschi Rowland, 1973. Thelyphonus publicaciones/ria/index.htm kopsteini (Speijer, 1933b) n. comb. for Minbosius kopsteini Speijer, 1933. Thely- phonus luzonicus Haupt, 2009 nomen novum (replacement name for Abaliella Página web GIA: manilana (Kraepelin, 1900) n. comb. Thelyphonus manilanus C.L. Koch, 1843 n. http://entomologia.rediris.es/gia comb. for Minbosius manilanus Speijer. Thelyphonus nasutus (Thorell, 1888) n. comb. for Tetrabalius nasutus Thorell, 1888.Thelyphonus rohdei (Kraepelin, Página web SEA: 1897) n. comb. for Abaliella rohdei (Kraepelin, 1897). Thelyphonus samoanus http://entomologia.rediris.es/sea (Kraepelin, 1897) n. comb. for Abaliella samoana (Kraepelin, 1897). Thelyphonus seticauda Doleschall, 1857 n. comb. for Tetrabalius seticauda (Doleschall, 1857). Thelyphonus willeyi (Pocock, 1898) n. comb. for Abaliella willeyi (Pocock, 1898).

14 Joachim Haupt

Introduction Papers Mus. Texas Tech Univ. 10, 3. AMNH. Abaliella rohdei (Kraepelin, 1897) syntypes. Abh. Geb. As the human population increases, whip scorpions Naturwiss. 25, 16-17. ZMB. become more and more rare, because habitats are dimin- Chajnus renschi Speijer, 1936 ♂ holotype. Mitt. zool. ished, although some species like Mastigoproctus gigan- Mus. Berlin 21, 258-259. ZMB. teus in USA and Ginosigma schimkewitschi in South- Ginosigma schimkewitschi (Tarnani, 1894) ♂. Horae East Asia come along very well in the surroundings of Soc. Ent. Ross. 29, 116-120. ZMB. human beings. But the possibility cannot be excluded Ginosigma lombokensis Speijer, 1936 ♀ holotype. Mitt. that certain species even became extinct. Many land- zool. Mus. Berlin 21, 256-257. ZMB. scapes become drier, because much water is used for Glyptogluteus augustus Rowland, 1973b ♂ holotype. other purposes, e.g. drinking water for human beings. Of Occ. Papers Mus. Texas Tech Univ. 16, 2-6. AMNH. course this is true for more developed countries like Mayacentrum guatemalae Víquez & Armas, 2006 ♀ Japan, but there may be also such developments in other paratype. Bol. Soc. Entomol. Aragonesa 38, 37-41. heavy populated areas like Indonesia. This is the domi- INBio. nant reason why this revision was carried out. Mayacentrum pijol Víquez & Armas, 2006 ♂ paratype, f The Asian whip scorpion fauna currently consists of 83 holotype. Bol. Soc. Entomol. Aragonesa 38, 37-41. species (Harvey, 2003), making it the most diverse re- INBio. gion of the world. In contrast, only a single species is Mimoscorpius pugnator (Butler, 1872) ♂ holotype. known from Africa, and 21 species are known from the Ann. Mag. Nat. Hist. Ser. 4 10, 204. BM America region. The Asian fauna is currently included Mimoscorpius pugnator (Butler, 1872), ♂, ♀. INBio. within 11 genera, with another genus in Africa and six Minbosius manilanus (C.L. Koch, 1843) ♂ syntype. Die genera in America. Arachniden 10, 28. ZMB Certain genera of Thelyphonida are defined on Minbosius kopsteini Speijer, 1933b ♂ allotype. Zool. the basis of their 'ommatoids'. Tetrabalius Thorell, 1888 Mededeelingen 16, 68-69. ML has two pairs of 'ommatidia' instead of one pair, but Ravilops wetherbeei (Armas, 2002) ♀ paratype. Bol. Abaliella Strand, 1928 has none at all (Rowland & Soc. Entomol. Aragonesa 37, 95-98. INBio. Cooke, 1973). Additionally, Chajnus Speijer, 1936 was Tetrabalius ambonensis Speijer, 1933b ♀ holotype. created for a species which has a cuticular thickening in Zool. Mededeelingen 16, 71-72. ML the otherwise thin cuticle of the 'ommatoids', but so far it Tetrabalius dammermanni Speijer, 1933b ♀ holotype, is only represented by the holotype from the Indonesian ♂, ♀. Zool. Mededeelingen 16, 73-74. ML island of Lombok. Tetrabalius seticauda (Doleschall, 1857) ♂, ♀. Natuurk. Moreover, two genera of whip scorpions are tijdschr. Nederl. Indie 13, 404. ML critically assessed, Minbosius Speijer, 1933b and Gi- Tetrabalius seticauda (Doleschall, 1857) ♂, ♀. Natuurk. nosigma Speijer, 1936. A third genus name (Teltus Spei- tijdschr. Nederl. Indie 13, 404. ZMB jer, 1936) was synonymized with Typopeltis (Haupt & Thelyphonus billitonensis Speijer, 1931 ♂ holotype. Song, 1996), and even a fourth name appeared (Gi- Zool. Mededeelingen 14, 85. ML popeltis, Speijer, 1934). Rowland & Cooke (1973) al- Valeriophonus nara (Valerio, 1981) ♂, ♀. Bol. Soc. ready cast some doubt on the creation of genera by Spei- Entomol. Aragonesa 37, 95-98. INBio. jer and made some critical remarks: 'the four genera added by Speijer (1933, 1936) have gone virtually unno- Results ticed, which is fortunate, for his inadequate understand- ing of the Uropygida adds only confusion to an already In this section diagnoses of the genera in alphabetic questionable classification'. Although, this fact may be order are provided, a proposition for synonymy includ- due to the low interest in these arachnids, even by arach- ing new combinations of species is made and a new key nologists. for Thelyphonida is established.

Material (Abaliella Strand, 1928: 4. 'Ommatoids' absent. Keel present. At least posterior opisthosomal tergites divided. Abbreviations: Some species with modifications on distal female tactile AMNH: American Museum of Natural History, New leg.) York.BM: British Museum of Natural History, London. (Chajnus Speijer, 1936: 258. Sclerotized spot within the INBio: Instituto Nacional de Biodiversidad, Heredia, one pair of 'ommatoids'. Keel present. At least posterior Costa Rica.ML: Natuurlijke Museum, Leiden. SMF: opisthosomal tergites divided. Female unknown.) Senckenberg Museum, Frankfurt. ZMB: Naturkundemu- Etienneus Heurtault, 1984, 122. One pair of 'ommatoids' seum, Berlin present. Keel absent. Pedipalpal patellar apophysis of male unmodified, like female. The following species were studied: Ginosigma Speijer, 1936, 256. One pair of 'ommatoids' Abaliella dicranotarsalis Rowland, 1973a, ♀ holotype. present. Keel present. Last joint of female tactile leg Occ. Papers Mus. Texas Tech Univ. 10, 5. AMNH. bayonet-shaped, some others modified. G. schim- Abaliella gertschi Rowland, 1973a, ♀ holotype. Occ. kewitschi: male with round spot on sternite V, filled Proposal for the Synonymy of some South-East Asian Whip Scorpion Genera (Arachnida, Uropygi, Thelyphonida) 15

Figures 1-6. 1. The 'ommatoid' (arrow) on the 3rd segment of the metasoma of Mastigoproctus giganteus. Bar: 2 mm. 2. Dupli- cation of the 'ommatoid' in Thelyphonus seticauda Doleschall, 1857 (arrows). 3. An area of thickened cuticula in the cover of the 'ommatoid' of Thelyphonus renschi holotype (arrow). Bar (Figs 2-4): 1.0 mm. 4. Whip organs (light spots) of Mastigoproctus giganteus. 5. Male of Ginosigma schimkewitschi terminal sexual organ. arrow: cuticular clasp with angle. Bar: 1.0 mm. 6. Male of Thelyphonus sucki for comparison: cuticular clasp without angle. Bar: 1.2 mm.

with hair sensilla. modified. Accessory tooth on inner margin of anterior Glyptogluteus Rowland, 1973b, 2. 'Ommatoids' absent. process of pedipalpal coxae. Keel present. Posterior opisthosomal tergites divided. Mastigoproctus Pocock, 1894, 129. One pair of 'omma- Male pedipalpal patellar apophysis unmodified. Male toids' present. Keel present. Male pedipalpal patellar sternites 8 and 9 modified. Female unknown. apophysis unmodified, similar to female. Posterior op- Hypoctonus Thorell, 1888, 360. One pair of 'ommatoids' isthosomal tergites undivided. present. Keel absent. Male pedipalpal patellar apophysis Mayacentrum Víquez & Armas, 2006, 37. One pair of modified. 'ommatoids' present. Keel present, but solely in posterior Labochirus Pocock, 1894, 132. One pair of 'ommatoids' part. All or part of the opisthosomal tergites divided, present. Keel absent. Male pedipalpal patellar apophysis whip organs absent. 16 Joachim Haupt

Figures 7-12. Female of Thelyphonus caudatus with receptacula seminis attached to the uterus externus laterally (plesiomorphic situation). Bar: 1.9 mm. 8. Female of Thelyphonus seticauda: Bar: 0.8 mm. 9. Female of Thelyphonus rohdei. Bar: 1.5 mm. The arrows point to the receptacula seminis. 10. Female of Mastigoproctus giganteus. Bar: 1.2 mm. The arrows point to the receptacula seminis. 11. Female of Typopeltis crucifer with anterior insertion of receptaculum seminis (arrows)(apomorphic situation). Bar: 1.3 mm. 12. Female of Typopeltis guangxiensis: same.

Mimoscorpius Pocock, 1894, 132. One pair of 'omma- Ravilops Víquez & Armas, 2006, 97. One pair of 'om- toids' present. Keel present. Male pedipalpal patellar matoids' present. Keel absent. Opisthosomal tergites I- apophysis modified, very long, pedipalpal femur very IV divided (IV only partially). Whip organs absent. long. ♂, ♀ with setae on inner surface of pedipalpal Male pedipalpal patellar apophysis longer and more patellar apophysis, tibia and tarsus slender than in female. (Minbosius Speijer, 1933b, 68. One pair of 'ommatoids' (Tetrabalius Thorell, 1888, 361. Two more or less di- present. Keel present. At least posterior opisthosomal vided pairs of 'ommatoids' present. Keel present. At tergites divided.) least posterior opisthosomal tergites divided. Distal Proposal for the Synonymy of some South-East Asian Whip Scorpion Genera (Arachnida, Uropygi, Thelyphonida) 17

joints of tactile leg in females modified.) Thelyphonus borneensis (Speijer, 1933 b) n. comb. (= Thelyphonellus Pocock, 1894, 133. 'Ommatoids' absent Tetrabalius borneensis Speijer, 1933 b: 72-73) or one pair present. Keel absent. Male pedipalpal patel- Thelyphonus borneonus Haupt, 2009 nomen novum lar apophysis unmodified. (replacement name for Tetrabalius borneensis (Speijer, Thelyphonus Latreille, 1802. One pair of 'ommatoids' 1933b: 72-73)) n. comb. Etymology: T. borneonus present. Keel present. Male pedipalpal patellar apophy- originates from the island of Borneo. sis unmodified, but in some species longer than in fe- Thelyphonus dammermanni (Speijer, 1933b) n. comb. (= male. At least posterior opisthosomal tergites divided. Tetrabalius dammermanni Speijer, 1933b: 73-74). Distal joints of tactile leg in females modified. Thelyphonus dicranotarsalis (Rowland, 1973a) n. comb. Typopeltis Pocock, 1894, 126. One pair of 'ommatoids' (= Abaliella dicranotarsalis Rowland, 1973a: 5-7). present. Keel present. Male pedipalpal patellar apophy- Thelyphonus florensis (Speijer, 1933b) n. comb. (= sis modified. Posterior opisthosomal tergites undivided. Tetrabalius florensis Speijer, 1933b: 74-75). Uroproctus Pocock, 1894, 129. One pair of 'ommatoids' Thelyphonus gertschi (Rowland, 1973a) n. comb. (= present. Keel present. Posterior opisthosomal tergites Abaliella gertschi Rowland, 1973a: 3-5). undivided. Male pedipalpal patellar apophysis modified. Thelyphonus kopsteini (Speijer, 1933b) n. comb. (= Accessory tooth on inner margin of anterior process of Minbosius kopsteini Speijer, 1933, b: 68-69). pedipalpal coxae. Thelyphonus luzonicus Haupt, 2009 nomen novum (re- Valeriophonus Víquez & Armas, 2005, 95. One pair of placement name for Abaliella manilana (Kraepelin, 'ommatoids' present. Keel present. Whip organs absent. 1900: 7-8)) n. comb. Etymology: T. luzonicus is derived With stridulation organ between pedipalpal coxae and from the island of Luzon. chelicerae. Thelyphonus manilanus C.L. Koch, 1843 (= Minbosius manilanus Speijer, 1936: 258). Thus the following genera are proposed to be syn- Thelyphonus nasutus (Thorell, 1888) n. comb. (= Tetra- onymized with Thelyphonus for the following reasons: balius nasutus Thorell, 1888: 401-405). The presence or absence of 'ommatoids' is obvious but it Thelyphonus rohdei (Kraepelin, 1897) n. comb. (= has nothing to do with taxonomy. Certainly, 'ommatoids' Abaliella rohdei (Kraepelin, 1897): 16-17). like whip organs are recognizable as light spots in the Thelyphonus samoanus (Kraepelin, 1897) n. comb. (= cuticle and serve an osmoregulative function (Haupt et Abaliella samoana (Kraepelin, 1897): 17). al., 1980) which is related to ecological conditions. Thelyphonus seticauda Doleschall, 1857 n. comb. (= Abaliella Strand, 1928, Archiv Naturgesch. 92 (8), 42. Tetrabalius seticauda (Doleschall, 1857): 404). Minbosius Speijer, 1933a, Tijdschr. Ent. 76, V. Thelyphonus willeyi (Pocock, 1898) n. comb. (= Tetrabalius Thorell, 1888, Annali Mus. Civ. Stor. Natur. Abaliella willeyi (Pocock, 1898): 98). Giacomo Doria (2) 6, 327-428. In the case of Minbosius no difference to Thelyphonus In consequence of this synonymy, the following was found. key is proposed, without distinguishing further families In Chajnus Speijer, 1936, Mitt. zool. Mus. Berlin 21, except Thelyphonidae. 258, the single specimen is a subadult male lacking genitalia, therefore it can only be treated as sp. dubium. 1. Stridulation organ present between coxa of pedi palp and base of chelicera……..Valeriophonus MORPHOLOGY OF GENITAL ORGANS - Stridulation organ absent …………………….2 2. ♂♂…………………………….…………….. 3 Studying the genial organs have revealed the following - ♀♀ (♂♂ included)...... 4 results: 3. Sternite III with median tooth (Asian spe- The Ginosigma male has an angle on the genital attach- cies)...... 4 ment, therefore it is definitely different from various - Sternite III without median tooth (American and species of Thelyphonus males (Fig. 5, 6). African species)…………….....….…….….. 11 Nevertheless, the females of Thelyphonus, Abaliella, 4. Pedipalpal coxa with median tooth ...... 5 Tetrabalius (Fig. 7, 8, 9) have a common character, the - Pedipalpal coxa without median tooth ...... 6 receptacula seminis are located laterally on the uterus 5. Keel present between lateral and median eyes .. externus, while further species of other south east asian ...... …………………………… Uroproctus genera (e.g. Typopeltis) have receptacula seminis in - Keel absent between lateral and median eyes direct continuation of the uterus externus in anterior ...... Labochirus, (Hypoctonus stoliczkae?) position (Fig. 11, 12). In the case of Minbosius 6. Asian species...... 7 manilanus the only 'adult' male has no genital organs. - American and African species ...... 11 As the second species (kopsteini) could not be studied, it 7. Keel present between lateral and median eyes…8 remains an open question. - Keel absent between lateral and median eyes

...... ……………………………… Hypoctonus NEW COMBINATIONS 8. Male pedipalpal patellar apophysis clearly modi- Thelyphonus ambonensis (Speijer, 1933b) n. comb. (= fied or very slender, female genital plate sculptu- Tetrabalius amboensis Speijer, 1933b: 71-72). tured, at least its posterior part differentiat...... Typopeltis 18 Joachim Haupt

- Male palpal apophysis nearly the same as in receptacula seminis to the uterus externus (in Ty- female ,,,,,,,,……………………………… 9, 10 popeltis). In fact, we will have to deal with three types: 9. Male sternites VIII and IX modified ………. 1. Thelyphonus, 2. Mastigoproctus – Thelyphonellus, 3. …………….. Glyptogluteus (female unknown) Typopeltis. Thelyphonus will be considered the most - Male sternites VIII and IX not modified, female plesiomorphic, Mastigoproctus – Thelyphonellus inter- genital plate unmodified…...... Thelyphonus mediary and Typopeltis apomorphic. (including female Ginosigma) Some female Thelyphonids have marks on the 10. Male with round structure on sternite V filled distal joints of the first leg. In order to be not misled one with hair sensilla, pedipalpal tibia enlarged but should distinguish whether these are cuts, black marks palpal apophysis similar to female… Ginosigma or really morphological distinctions. In at least one spe- - Male without any differentiation on sternite V.... cies of Typopeltis cuts are inflicted by the male, they are ………,…………………………………..…. 11 typically not symmetrical (Haupt, 1997). The unmated 11. Keel present between lateral and median eyes, at females are devoid of it and also devoid of darkenings least in posterior part …...... 12 on the distal joints of the tactile leg. The same could be - Keel absent between median and lateral eyes ..... true for the black marks of Thelyphonus – they are in ,,…………………………………...….… 14, 15 many cases also not symmetrical. In the genus Abaliella, 12. Male pedipalpal femur very long, tibia extremely there are morphological differentiations, but not in all flat. ♂, ♀ with long setae on inner surface of pe- species (e.g. A. rohdei). What they are good for? This dipalpal patellar apophysis, tibia and tarsus ...... question can only be solved by observing living female ...... Mimoscorpius specimens. - Male pedipalpal femur normal, no setae present Thorell (1888) and Kraepelin (1897) agree about on inner surface of pedipalps ...... ………….. 13 the monotony of Thelyphonida (...erweisen sich die 13. Opisthosomal tergites undivided, keel between Beschreibungen und namentlich die Abbildungen der anterior and lateral eyes in full length ……. älteren Autoren als vollkommen unzulänglich...[und so …………………..……………..Mastigoproctus ist] vielfach nur mit Hilfe des Originalexemplars ein - Opisthosomal tergites at least partly di- sicheres Urteil über ihre Identität...zu gewinnen.) Unfor- vided………..……………………………..… 14 tunately, this situation has not changed much – espe- 14. Keel only present in posterior part ……………. cially in South East Asia. Many details were described …………………………………... Mayacentrum which nowadays seem to be 'traditional', while certain - Opisthosomal tergites I-IV divided (IV only other information much needed (e.g. on genital organs) partly), with neither whip organs nor keel ….. is lacking...... Ravilops While Kraepelin (1899) only knew one single 15. Opisthosomal tergites II and III divided, the fol- family – Thelyphonidae - several authors tried to divide lowing one partially divided, median eyes more up Thelyphonida into separate subfamilies or families anterior, without keel, American species…... (Pocock, 1899, Speijer 1936, Rowland & Cook, 1973), ,,,,,………………………………Thelyphonellus but apparently it is too early to do that (Weygoldt, Median eyes more posterior than in Thelyphonel- 1979). In addition, the biology of several genera is not lus, without keel, African species ...... Etienneus fully understood – or not understood at all. Börner (1904) mainly studied Thelyphonus cau- Discussion datus and species of Mastigoproctus, i.e. plesiomorphic species. This resulted in the fact that he did not recog- One of the main questions today is on the phy- nize differences in apomorphic species. Moreover, he logeny of Thelyphonids. At least it has been possible to stated that 'ocelli or ommatoids' do not have the slightest distinguish plesiomorphic (i.e. Thelyphonus), and apo- similarity with real ocelli and he also did not find a morphic (i.e. Typopeltis) genera, even though a phy- similarity to luminary organs of other land arthopods. logenetic tree could not be constructed. Kraepelin (1900) expressed doubt concerning the impor- Weygoldt (1988, cf. discussion) distinguished two types tance of 'ommatoids' at the generic level. according to spermatophore function and morphology: Weygoldt (1979) studied Thelyphonellus ama- Thelyphonus which does not deal with the sperm zonicus (Butler, 1872) and found 'ommatoids' in some packages, at all. specimens, although small, while other specimens did Mastigoproctus, Thelyphonellus and Typopeltis not have such organs. The genera Abaliella Strand, 1928 which at the end of mating embrace the opisthosoma of and Glyptogluteus Rowland, 1973b totally lack such the female in order to empty the sperm packages. organs. He subdivided the second type in Mastigoproctus – Apparently, 'ommatoids' were overestimated in Thelyphonellus and Typopeltis which emptied the sperm the past concerning their taxonomic importance. Their packages entirely (2a & 2b). function was not understood. Once it became known I am inclined to raise this second type to inde- that they are related to the whip organs and contain a pendence in order to mirror the morphology of sexual transporting epithelium (Haupt et al., 1980) engaged in organs, lateral attachment of receptalula seminis to the the resorption of solutes and/or water, their presence or uterus externus (in Thelyphonus, Thelyphonellus, Masti- absence is certainly related to ecological conditions in goproctus etc.) and direct, anterior attachment of the the surrounding atmosphere. Proposal for the Synonymy of some South-East Asian Whip Scorpion Genera (Arachnida, Uropygi, Thelyphonida) 19

Therefore, the following genera are synonymized bok of the widely distributed G. schimkewitschi – once with Thelyphonus: Tetrabalius and Abaliella. the male will be found with the same typical structure on Some genera like Minbosius and Ginosigma sternite V. A bayonet-shaped last joint of the female contain two species, each. In the case of Minbosius tactile leg is fairly common among species of the genus species, they are from the Northern part of the Philip- Thelyphonus. It occurs in Thelyphonus leucurus Pocock, pines (T. luzonicus = Minbosius manilanus) and from an 1898 (T. X), T. asperatus Thorell, 1888 (as the holotype island (Noesa Laoet) near Ambo(i)n(a) in Indonesia happens to be a male, see depiction by Kraepelin, 1897), (kopsteini). The holotype of Minbosius kopsteini is a T. burchardi Kraepelin 1911, T. lawrenci Rowland, female only known from the literature, in the same vial 1973a (although tooth more basally). This shows that are a very small male (inspected) and a subadult female. Ginosigma is rather closely related to Thelyphonus. No wonder, why a generic definition is lacking: apparently, there is no difference to Thelyphonus and there- Acknowledgments fore I synonymize the genus Minbosius to Thelyphonus. In the case of Ginosigma the male has a round I am gratefully indebted to Jason Dunlop and A. Friederichs spot on sternite V with hair sensilla and the genital at- (Zoological Museum Berlin), the late J. Heurtault and C. Rollard (Muséum National d' Histoire Naturelle, (Paris), Janet tachment shows an angle as compared with Thelyphonus Beccaloni (Museum of Natural History, London), Lorenzo (Fig. 5). The last joint of the female tactile leg is bayo- Prendini and J.C. Huff (American Museum of Natural His- net-shaped and the penultimate joint shows three proc- tory, New York), P. Jaeger and J. Altmann (Senckenberg esses. Speijer (1936) included two species in this genus, Museum Frankfurt), E.J. van Nieukerken, C. Pepermans and J. G. schimkewitschi and G. lombokensis. As the second Smit (Museum Leiden) and C. Víquez (INBio, Costa Rica). species, of which only a female is known, has a bayonet Three unnamed referees helped largely to improve the manu- structure on the last joint of the tactile leg as well and script. only smaller processes on the penultimate one joint, it could well be considered as an island form from Lom-

References

BÖRNER, C. 1904. Beiträge zur Morphologie der Arachnida of the orders Pedipalpi and Solifugae. Arthropoden. I. Ein Beitrag zur Kenntnis der Natural Science London, 14: 213-231. Pedipalpen. Zoologica, 42: 1-174. ROWLAND, J.M. 1973a. New whipscorpions from New HARVEY, M.S. 2003. Catalogue of the smaller arachnid Guinea and the Solomon Islands (Thelyphonida, orders of the world. CSIRO publishing. Colling- Arachnida). Occasional Papers the Museum Texas wood Victoria, Australia. 385 pp. Tech University, 10: 1-8. ROWLAND, J.M. 1973b. HAUPT, J. & SONG, D. 1996. Revision of East Asian Uropygida (Arachnida) from the Philippine Is- whip scorpions (Arachnida Uropygi Thely- lands, with description of a new genus and spe- phonida). I. China and Japan. Arthropoda Selecta, cies. Occasional Papers the Museum Texas Tech 5: 43-52. University, 16: 1-11. HAUPT, J., EMDE, W. & WEYGOLDT, P. 1980. Die ROWLAND, J.M. 2002. Review of the South American Geißelorgane des Geißelskorpions Mastigoproctus whip scorpions (Thelyphonida: Arachnida). Ama- giganteus (Lucas) (Arachnida: Uropygi): Trans- zoniana, 17: 187-204. portepithel. Zoomorphologie, 96: 205-213. ROWLAND, J.M. & COOKE, J.A.L. 1973. Systematics of KRAEPELIN, K. 1897. Revision der Uropygi Thor. the Arachnid order Uropygida (=Thelyphonida). Abhandlungen aus dem Gebiete der Journal of Arachnology, 1: 55-71. Naturwissenschaften, 25: 3-60. SPEIJER, E.A.M. 1933a. (Bemerkungen über Pedipalpi KRAEPELIN, K. 1899. Pedipalpi. In Das Tierreich. 8. IV) Verslag 66ste Wintervergadering der Neder- Lieferung, Arachnoidea. Deutsche Zoologische landsche Entomologische Vereeniging. Tijdschrift Gesellschaft ed. pp. Friedländer, Berlin. pp.201- voor Entomologie, 76: 4-5. 265. SPEIJER, E.A.M. 1933b. Die Pedipalpi des Zoologischen KRAEPELIN, K. 1900. Über einige neue Gliederspinnen. Museums in Buitenzorg und die der Sammlung Abhandlungen aus dem Gebiete der Dr. F. Kopstein. Zoologische Mededeelingen, 16: Naturwissenschaften, 16: 1-8. 67-76. KRAEPELIN, K. 1911. Neue Beiträge zur Systematik die SPEIJER, E.A.M. 1933c. A new Pedipalp from Mt. Gliederspinnen. Mitteilungen des Kinabalu, Noth Borneo, 13,455 ft. Bull. Raffl. Naturhistorischen Museums Hamburg, 28: 59- Mus. Singapore, 8: 106-109. 108. SPEIJER, E.A.M. 1934. Note préliminaire sur le nouveau POCOCK, R.I. 1898. Scorpions, Pedipalpi and spiders genre Gipopeltis (Arach., Pedipalpi). Bulletin du collected by Dr. Willey in New Britain, the Muséum National d'Histoire Naturelle, (2) 6: 421- Solomon Islands, Loyalty Islands, etc. Annual 422. Magazine of Natural History, 7(1): 95-100. SPEIJER, E.A.M. 1936. Die orientalischen Pedipalpen POCOCK, R.I. 1899. The geographical distribution of the des Zoologischen Museums der Universität Berlin. 20 Joachim Haupt

Mitteilungen des Zoologischen Museums Berlin, VIQUEZ, C. & ARMAS L.F. DE. 2006. Un nuevo género y 21: 249-263. dos nuevas especies de vinagrillos centroamerica- THORELL, T. 1888. Pedipalpi e scorpioni dell' nos (Arachnida: Thelyphonida). Boletin Sociedad Arcipelago malese conservati nel Museo civico di Entomológica Aragonesa, 38: 37-41. Storia naturale di Genova. Annali del Museo WEYGOLDT, P. 1979. Thelyphonellus ruschii n.sp. und Civico di Storia Naturale Giacomo Doria, (2) 6: die taxonomische Stellung von Thelyphonellus 327-428. Pocock 1894 (Arachnida: Uropygi: Thelyphoni- VIQUEZ, C. & ARMAS L.F. DE. 2005. Dos nuevos gene- da). Senckenbergiana biologica, 60: 109-114. ros de vinagrillos de Centroamérica y las Antillas (Arachnida: Thelyphonida). Boletin Sociedad En- tomológica Aragonesa, 37: 95-98.