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Full text of "Revue suisse de zoologie"

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REVUE SUISSE DE ZOOLOGIE 



TOME 110— FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des- universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 230.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P 6434, CH-1211 Genève 6, Suisse 



ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

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MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



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Jël M EVE MARS 2003 ISSN 0035 -418 X <•) ■■ 



REVUE SUISSE DE ZOOLOGIE 



TOME 1 10— FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 

Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités suis- 
ses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-gra- 
phie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 
SUISSE Fr. 225.— UNION POSTALE Fr. 230.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 



Revue suisse de Zoologie 1 10 (1): 3-43; mars 2003 



Catalogue of type-material of stick-insects housed in the Muséum 
d'histoire naturelle, Geneva, with descriptions of some new taxa 
(Insecta: Phasmatodea) 

Oliver ZOMPRO' & Paul D. BROCKS 

1 Max-Planck-Institut für Limnologie, Arbeitsgruppe Tropenökologie, 
August-Thienemannstraße 2, D-24306 Plön, Germany. 

E-mail: zompro@mpil-ploen.mpg.de 
Website: www.sungaya.de 

2 „Papillon", 40, Thorndike Road, Slough, SL2 1SR, England. 

Catalogue of type-material of stick-insects housed in the Muséum d'his- 
toire naturelle, Geneva, with descriptions of some new taxa (Insecta: 
Phasmatodea). - Type specimens for 157 species-group names of Phasmida 
have been identified in the collection of the Muséum d'histoire naturelle, 
Geneva. The names are listed alphabetically, with the number of specimens, 
sex and locality data. The material includes most species described by 
Saussure and the majority of those described by Carl; in addition there is 
material of Bolivar, Brock, Brunner von Wattenwyl, Fritzsche & Gitsaga, 
Günther, Harz, Kirby, Redtenbacher, Werner and Zompro. Clitumnus hum- 
berti Carl, 1913 is re-named Ramulus carli Zompro nom. n. Lectotypes are 
designated for Xera debilis Redtenbacher, 1906, Clonistria guatemalensis 
Redtenbacher, 1908, Ramulus humberti (Saussure, 1861) and Anophelepis 
poeyi Saussure, 1868. One genus and four species are described as new. 
Parapodacanthus Brock gen. n., with the type-species P. hasenpuschorum 
Brock sp. n., is described from Queensland, Australia. Three Philippine spe- 
cies are provisionally placed in the genus Lonchodes Gray, 1835: L. puting- 
mantsa Zompro sp. n., L. magayon Zompro sp. n. and L. dalawangsungay 
Zompro sp. n.; this complex urgently needs a generic revision. The eggs of 
Alienobostra godmani (Redtenbacher, 1908) and Lonchodes palawanicus 
(Carl, 1913) are described and figured for the first time. Xylodus Saussure, 
1859, formerly synonymized with Cladomorphus Gray, 1835, and 
Micrarchus Carl, 1913 formerly synonymized with Pachymorpha Gray, 
1835, are re-established as valid genera. Bacteria divergens Redtenbacher, 
1 908 is found to be a new synonym of Phanocloidea muricata (Burmeister, 
1838), and Ocnophila adulterina Brunner v. Wattenwyl, 1907 of Lamponius 
guerini (Saussure, 1868). Prisomera palawanica Carl, 1915 is transferred to 
Stheneboea Stai, 1875, and Myronides trilineatus Carl, 1913 to Lopaphus 
Westwood, 1859. 

Key-words: Phasmatodea - type material - Geneva Museum of Natural 
History - new genus - new species - eggs - catalogue. 



Manuscript accepted 09.10.2002 



O. ZOMPRO & P. D. BROCK 



INTRODUCTION 

The Phasmatodea collection in the Muséum d'histoire naturelle, Geneva, 
Switzerland (MHNG) contains type specimens for 157 species-group names. The col- 
lection is very important for the taxonomist, as it includes most of Saussure 's type ma- 
terial (mid 1800's), which was published after Westwood's major catalogue of the 
order (1859). The vast majority of Carl's (1913, 1915) type specimens are also present, 
the remainder is in Naturhistorisches Museum, Basel, Switzerland (NHMB). Brunner 
von Wattenwyl and Redtenbacher (1906-08) had already selected a number of spe- 
cimens in MHNG as type material. In addition, there are single species described by 
Bolivar (1888), Günther (1937), Kirby (1904) and Werner (1918) and an invalid neo- 
type designation by Harz (1976). Recently Brock, Fritzsche & Gitsaga and Zompro 
have added material. The collection is considered to be one of the most important for 
the taxonomist (Brock, 1999). 

The phasmid collection is housed in 93 cabinet drawers with dimensions of 46 
x 36 x 6cm, with type material included in the general collection. Additionally, there 
are three drawers in the separate collection of Harz, which contain phasmids. Although 
mainly containing specimens named and labelled by curators, the collection includes 
some unidentified specimens, such as phasmids from the Philippines, only identified by 
K. Günther to genus level, housed in drawers labelled "Dépôt". There are also a few 
phasmid specimens in alcohol. Although little research has been made on the collec- 
tion since Carl's papers (1913, 1915), it has been examined by the following, who have 
added various labels to specimens: P. D. Brock, M. Hebard, C. F. Moxey and O. 
Zompro. Moxey placed various iectotype' labels on specimens; however, these are not 
valid, as his 1972 thesis on phasmids from the West Indies was not published. 
Hennemann also placed various notes, including Iectotype labels. These are also not 
valid before a publication. As both data are invalid at the present state, they are om- 
mitted from this catalogue. 

LIST OF TYPE-SPECIMENS 

The following is a complete list of type material of Phasmatodea known to be 
in the MHNG (up to March 2001), according to detailed, independent checks of 
specimens and literature by both authors. 

The systematics of this order are overworked by Zompro recently; therefore a 
systematic arrangement is pointless and names are arranged alphabetically by species 
as originally published. The cabinet drawer number is given in square brackets. Data 
are as recorded on data labels. If they do not agree with the data in the original publi- 
cation, comments are provided. Several species have not been found in the cabinet 
drawers, nor by reference to the museum's card index to species, despite detailed 
searches. Reference is made where specimens have been traced in other collections. 
The actual combination of each species is mentioned, if possible corresponding to the 
present state of research. 

Museum-codes are as follows: 

ANIC Australian National Insect Collection, CSIRO, Canberra, Australia. 

ANSP Academy of Natural Sciences, Philadelphia, USA. 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 5 

BMNH The Natural History Museum, London, England. 

CUMZ University Museum of Zoology Insect Collection, Cambridge, England. 

DEIC Deutsches Entomologisches Institut, Eberswalde, Germany. 

ETHZ Eidgenössische Technische Hochschule, Zurich, Switzerland. 

HNHM Hungarian Natural History Museum, Budapest, Hungary. 

ISNB Institut Royal des Sciences Naturelles, Brussels, Belgium. 

MCSN Museo Civico di Storia Naturale „Giacomo Doria", Genua, Italy. 

MHNG Muséum d'histoire naturelle, Geneva, Switzerland. 

MNHN Muséum National d'Histoire Naturelle, Paris, France. 

MNMS Museo Nacional de Ciencias Naturales, Madrid, Spain. 

NHMB Naturhistorisches Museum, Basel, Switzerland. 

NHMW Naturhistorisches Museum, Vienna, Austria. 

SMFD Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, 

Germany. 
SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany. 
SMTD Staatliches Museum für Tierkunde, Dresden, Germany. 
QMBA Queensland Museum, South Brisbane, Queensland, Australia. 
UMBB Übersee-Museum, Bremen, Germany. 

ZMAS Zoological Museum, Academy of Science, St. Petersburg, Russia. 
ZMHB Zoologisches Museum der Humboldt-Universität, Berlin, Germany. 
ZMUH Zoologisches Museum der Universität Hamburg, Germany. 
ZMUK Zoologisches Museum der Christian-Albrechts-Universität, Kiel, Germany. 
ZSMC Zoologische Staatssammlung, München, Germany. 

Other abbreviations used: 

HT / AT / PT / ST / LT / PLT: Holo- / Alio- / Para- / Syn- / Lecto- / Paralectotype. 

Type-information is listed in the following format: 

species-group name author, year of publication: page [original genus] 

[Number and sexes of types and localities as recorded in original description] 

Museum [Drawer-number]: Number of specimens. 

List of types including inscriptions of labels. Different labels are separated by ( ; ). 

[Valid name] 

(Name in collection) 

Comment. 

adulterina Brunner v. Watten wyl, 1907: 316 [Ocnophila] 

[â . Guadeloupe (Mus. Genav.).] 

MHNG [41]: 1 Ô. 

HT 6 : Guadeloupe, M. Schramm. 

[Lamponius guerini (Saussure, 1868) syn. n.] 

(In collection as Ocnophila adulterina Brunner v. Wattenwyl.) 
adumbratus Saussure, 1859: 62 [Xylodus] Fig. 1 

[9. Portorico.] 

MHNG [64]: 1 9. 

HT: 9, Porto Rico, Brésil, M. T. J. Gidet, 189/53 



6 O. ZOMPRO & P. D. BROCK 

[Xylodus adumbratus Saussure, 1859] 

(In collection as Phibalosoma ceratocephalum Gray.) 

Comment. Xylodus Saussure, 1859, with the type-species Xylodus adumbratus 

Saussure, 1859, has been synonymized in error with Phibalosoma Gray, 1835, 

by Redtenbacher (1908: 425). Phibalosoma is a synonym of Cladomorphus 

Gray, 1835, synonymized by Kirby (1904: 356). The genitalia of Xylodus differ 

strikingly from those of Cladomorphus, therefore Xylodus is re-established as a 

valid genus. Further research is in progress by Zompro. 
aestuans Saussure, 1861a: 474, pi. 11: 2 [Bacteria (Bacunculus)] 

[ 9 . Du vieux Calabar.] 

MHNG [?]: Not traced. 
annulata Carl, 1913: 32 [Dyme] 

[16. Guyana. Muséum de Genève.] 

MHNG [44]: 1 6. 

HT S : Guyana, Saussure. 

[Globocalynda annulata (Cari, 1913)] 

(In collection as Dyme annulata Cari.) 
antillarum Saussure, 1868: 65 [Bacteria] 

[Insula Gadalupe.] 

MHNG [42, 43]: 3 6, 4 9, 1 egg ex ovipositor. 

ST: 1 ó" , 1 9,1 egg ex ovipositor: Guadeloupe, Schramm; 2 <5 , 2 9 : 

Guadeloupe, H. d. Saussure; 9 : Guadeloupe, coll. Guerin. 

Further material in ETHZ. 

[Bacteria ferula (Fabricius, 1793)] 

(In collection as Dyme antillarum Saussure.) 

Comment. Bacteria ferula (Fabricius, 1793), was synonymized with Bacteria 

crudelis Westwood, 1859, by Kirby, 1904c: 346, and the latter was syn- 
onymized with Bacteria ferula (Fabricius, 1793) by Brock, 1998: 33. 
atropurpurea Carl, 1913: 33 [Dyme] 

[19. Espirito-Santo, Brasilia. Muséum de Genève.] 

MHNG [42]: 1 6. 

HT 6 : Espirito Santo, Brasil, ex coll. Fruhstorfer. [Recorded by Carl as 9 ] 

[Dyme atropurpurea Carl, 1913] 
augusta Carl, 1913: 17 [Cuniculina] 

[19. Than-Moi, Tonkin. Muséum de Genève.] 

MHNG [28]: 1 9. 

HT 9 : Tonkin, Than-Moi, Jan.-Juli, Fruhstorfer. 

[Ramulus augusta (Carl, 1913)] 
azteca Saussure, 1859: 62 [Bacteria] 

[ 9 . Littus mexicanum.] 

MHNG [?]: Not traced. 

[Pseudosermyle azteca (Saussure, 1859)] 
baculus Saussure. 1859: 62 [Bacteria] 

[9. America.] 

MHNG [45]: 1 9. 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 7 

HT 9 : Amer, mer, 475/20. 

(In collection as Bacunculus baculus Saussure.) 
bedoti Redtenbacher, 1908: 388, pi. 19: 1 [Rhaphiderus] 

[8. Tonking (Mus. Genf, Coll. m.)] 

MHNG[54]: 1 6. 

ST: 1 8, Tonkin. 

Further type-material in NHMW. 

[Rhaphiderus bedoti Redtenbacher, 1908] 
biceps Redtenbacher, 1908:451 [Pharnacia] 

[8 9. Assam (Mus. Stuttgart); Tonking (Mus. Paris); Java (Mus. Genf, Mus. 

Budapest, Mus. Berlin, Coll. m., Mus. Ak. Petersburg, Mus. Brüssel, Mus. 

Paris, Mus. Hamburg, Hofmus. Wien)] 

MHNG[69]:2 6,2 9. 

ST: 2 S, Java, Or., Passeroean 622/20; 2 9, Java. 

Further type-material present in ISNB, NHMW, SMNS, ZMAS, ZMHB, 

ZMUH. 

[Pharnacia biceps Redtenbacher, 1908] 
bicolor Car\, 1913: 14 [Clitumnus] 

[2 d.Than-Moi, Tonkin.] 

MHNG[25]:2 8. 

ST: 2 S : Tonkin, Than-Moi, Jan.-Juli, Fruhstorfer. 

[Ramulus bicolor (Carl, 1913)] 

(In collection as Clitumnus bicolor Carl.) 
bicolor Günther, 1937a: 90, pi. 1:3 [Neopromachus] 

[9 6 8, Torricelli-Gebirge, 700-900m] 

Type-series: 9 8 8. 

MHNG[40]:2 6. 

ST: 1 6 , Kais. Wilhelmsland, Torricelli Gebirge, Dr. Schlaginhaufen, 780 m, 

1910 1; 1 6 , Kais. Wilhelmsland, Torricelli Gebirge, Dr. Schlaginhaufen, 720 m, 

1910 1. 

Further type-material in SMTD. 

[Neopromachus pachynotus bicolor Günther, 1937] 
bifasciatus Redtenbacher, 1906: 102 [Eudes] 

[9. Costa-Rica (Mus. Genf).] 

MHNG[10]: 1 9. 

HT: 9 , Costa Rica, Amer. Centn, Biolley. 

Also a non-type 6 described by Carl (1913: 7) present. 

[Eudes bifasciatus Redtenbacher, 1906] 
biolleyi Carl, 1913: 9 [Prisopus] 

[1 ó\ La Laguna, Cardio, 1000m, Costarica. (P. Biolley leg.).] 

MHNG[16]: 18. 

HT 6 : Costa Rica, La Laguna, Chemin de Cardio, 1000 m, Biolley. 

[Prisopus biolleyi Cari, 1913] 
borneensis Cari, 1913: 44, pi. 1: 4, 5 [Echinoclonia] Fig. 15 

[1 9 . Brunei, Borneo sept. Muséum de Genève.] 



8 O. ZOMPRO & P. D. BROCK 

MHNG[76]: 1 9. 

HT 9 : Borneo, Staudinger. 

[Parastheneboea laetior (Brunner v. Watten wyl, 1907), synonymized by 

Günther, 1933: 260.] 

(In collection as Acanthocleon bomeense Carl.) 

Comment. "Acanthocleon" is an unpublished name and therefore invalid. 
brevipes Redtenbacher, 1908: 441, pi. 21: 4a, b [Stephanacris] 

[6 9. Neu-Guinea (Mus. Budapest; Coll. m., Coll. Staudinger).] 

MHNG[67]: 1 6, 1 9. 

ST (?): 1 6,1 9, N. Guinea, Biro 1899, Sattelberg, Huon Golf. 

Possibly syntypes from the Staudinger-collection, the status is unclear. The data 

are the same as for part of the type-material in NHMW and may have originated 

from HNHM. Brunner and Redtenbacher did not label all material, sometimes 

labels have been taken off or replaced subsequently. Material also present in 

ZMUH. 

[Stephanacris brevipes Redtenbacher, 1908] 
brevitarsus Carl, 1913: 38, pi. 1:6 [Antillophilus] 

[1 9 . Guadeloupe. Muséum de Genève.] 

MHNG[49]: 1 9. 

HT 9 : Guadeloupe, Saussure. 

[Lamponius guerini (Saussure, 1868), synonymized by Langlois & Lelong, 

1997: 49.] 
burkartii Saussure, 1868: 65 [Bacteria] 

[Mexico.] 

MHNG [?]: Not traced. 

[Phanocles burkartii (Saussure, 1868)] 
capitata Brunner v. Wattenwyl, 1907: 315 [Ocnophila] 

[6 . Peru (Mus. Genav.).] 

MHNG [41]: 5 6. 

ST: 5 6 , Lima, Pérou, Ane. coll. 

Also undescribed 9 are present, with same data, not treated in original 

publication. 

[Ocnophila capitata Brunner v. Wattenwyl, 1907] 
carinulatus Saussure, 1868: 63 [Bacillus] Figs 2-4 

[Ceylon ?] 

MHNG [30]: 2 9,29 nymphs. 

ST: 2 9,29 nymphs, Malabar. 

[Gratidia carinulata (Saussure, 1868)] 
ceylonica Saussure, 1868: 67 [Anophelepis] Fig. 5 

[Ceylon.] 

MHNG [76]: 1 9. 

HT: 9 , Ceylon, M. H. de Saussure. 

[Paramenexenus ceylonicus (Saussure, 1868)] 
ceylonica Saussure, 1868: 69 [Necroscia] 

[Ceylon.] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 9 

MHNG[90]:8 Ô, 1 9. 

ST: 8 (?, 1 9, Ceylon, Mr. Brunner d. W., 609/56. 

[In collection as Sipyloidea ceylonica (Saussure)] 

[Necroscia ceylonica Saussure, 1868] 
ceylonicus Saussure, 1868: 66 [Lonchodes] 

[Ceylon.] 

MHNG[25]:2 6. 

ST: 2 6 , Ceylon, Nitner. 

[Ramulus ceylonicus (Saussure, 1868)] 

[In collection as Clitumnus ceylonicus (Saussure)] 
circe Redtenbacher, 1908: 347 [Canachus] 

[o* 9. Neu-Caledonien (Mus. Genf); Singapore (?) (Hofmus. Wien).] 

MHNG[47]: 1 9. 

ST: 1 9 , Nov. Caledonie, Ozeanie. 

[Paracanachus circe (Redtenbacher, 1908)] 

Further type-material in NHMW. 
claraziana Saussure, 1868: 64 [Anisomorpha] 

[ 9 . Ager Argentinus (la Plata)] 

MHNG[64]: 1 <?. 

HT et: Rio Chulat, Rep. Arg., S. Claraza. [Saussure's 9 (nymph ?) is a S] 

[Agathemera claraziana (Saussure, 1868)] 

[In collection as Agathemerea pardalina (Westwood, 1859)] 
composita Carl, 1913: 42 [Achrioptera] 

[1 ó* „Madagascar" (?). Muséum de Genève.] 

MHNG[66]: 1 S. 

HT S : No label. 

[Achrioptera composita Carl, 1913] 
continentalis Carl, 1913: 18 [Entoria] 

[19. Tonkin (Than-Moi). Muséum de Genève.] 

MHNG[29]: 1 9. 

HT 9 : Tonkin, Than-Moi, Jan. -Juli, Fruhstorfer. 

[Entoria continentalis Carl, 1913] 
cornuta Saussure, 1861b: 128. [Bacteria] 

[Mexico calida.] 

MHNG [90]: Not traced. 
coronata Carl, 1913: 34 [Calynda] 

[1 9, San José, Costarica (1161m); 1 9, San Carlos, Costarica (250m). (P. 

Biolley leg.). Muséum de Genève.] 

MHNG [44]: 2 9. 

ST: 9 : Costa Rica, San Jose, 1161m; 9 : San Carlos, 250 m, Biolley. 

[Calynda coronata Carl, 1913] 
coulonianum Saussure, 1868: 66 [Bactridium] 

[Nova Hollendia.- (Specimen e Chile?)] 

MHNG [9]: 1 9. 

HT 9 : Nova Holl. 



10 O. ZOMPRO & P. D. BROCK 

[Heteronemia mexicana Gray, 1835, synonymized by Zompro, 2001b.] 

(In collection as Donusa prolixa Stài.) 
cubaensis Saussure, 1869: 159 [Bacteria] 

[Ç.L'illedeCuba.] 

MHNG [?]: Not traced. 

[Bacteria cubaensis Saussure, 1869] 
cubensis Saussure, 1868: 68 [Haplopus] 

[9. Cuba.] 

MHNG [64]: 1 9. 

HT 9 : Cuba. 

[Aplopus cubensis Saussure, 1868] 
cubensis Saussure, 1868: 69 [Phasma] Figs 6, 59 

[No locality mentioned.] 

MHNG [14]: 4 o\ 

ST: 2 S : Cuba, H. d. Saussure; 2 S : Cuba, acheté à M. Poey. 

[Alloephasma poeyi (Saussure, 1868), synonymized by Redtenbacher, 1906: 

127] 
cubensis Saussure, 1868: 67 [Phybalosoma] 

[d.Cuba.] 

MHNG [61]: 1 6. 

HT â : Cuba, H. d. Saussure. 

[In collection as Bacteria cubensis (Saussure)] 
cultratolobatus B runner v. Watten wyl, 1907: 273 [Carausius] 

[Brunei in ins. Borneo (Mus. Genav.).] 

MHNG [36]: 1 9. 

HT: 9 , Brunei, E Borneo, Staudinger. 

[Carausius cultratolobatus Brunner v. Wattenwyl, 1907] 
dalawangsungay Zompro sp. n. [Lonchodes] Figs 43-45, 57 

MHNG [39bis]: 1 9. 

HT: 9 , Philippinen, Luzon, Imugan. 

Description below. 
debilis Redtenbacher, 1906: 146 [Xera] 

[6 9 . Merida, Venezuela (Coll. m., Mus. Budapest): Columbien (Mus. Berlin).] 

MHNG [16]: 1 6, 1 9. 

PLT: 1 6,1 9 , Venezuela. Briceno, Merida; ex Mus. Hungar. 

These specimens originate from the HNHM-collection before it was destroyed 

by fire in 1956. 

MNHU[73/5]: 1 9. 

PLT: 1 9, Xera inermis Redt.* n. sp. Redtenbacher / Brunner det.; 788; 

Columb. Mor. 

NHMW[7/34]:5 o\ 4 9. 

LT, by present designation: 6 , Coll. Br.v.W. Merida Venezuela; det. Redtenb. 

Xera debilis; PLT: 9 , 5 eggs ex abdomen. Coll. Br.v.W. Merida Venezuela; det. 

Redtenb. Xera debilis. 

[Xera debilis Redtenbacher, 1906] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 1 1 

The LT designation is part of a revision of the genera of Phasmatodea: Areolatae 

by O. Zompro and will support nomenclatural stability. 
deplanata Redtenbacher, 1908: 411 [Bostra] 

[?. Cuba (Mus. Genf).] 

MHNG[60]: 1 9. 

HT: 9 , Cuba, M. Poey, 620/88. 

[Phanocloidea (?) deplanata (Redtenbacher, 1908)] 
divergens Redtenbacher, 1908: 418 [Bacteria] 

[Cayenne (Mus. Genf. Mus. Paris).] 

MHNG[61]: 1 9. 

ST: 9 , 2 eggs, Cayenne, Prudhomme, 620/85. 

Further type-material in MNHN. 

[Phanocloidea muricata (Burmeister, 1838), n. syn.] 
dohertyi Carl, 1913: 46 [Paroxyartes] Fig. 7 

[1 S . Jobi. (Doherty leg.). Muséum de Genève.] 

MHNG[76]: 1 S. 

HT S : Jobi, Doherty, ex coll. Fruhstorfer. 

[Paroxyartes dohertyi Cari, 1913] 
dubius Cari, 1915: 185 [Asprenas] 

[Neu-Caledonien. Mt. Ignambi, 29.IV.1911, 1 S, 1 9; Mt. Panie, 26.VI.1911, 

1 ó\] 

MHNG[49]: 1 S. 

ST S: 26.6.11, Panie Wald. 

Further material in NHMB. 

[Asprenas dubius Carl, 1915] 
effeminatus Carl, 1913: 37 [Asprenas] 

[1 S . Nova Caledonia. Muséum de Genève.] 

MHNG[49]: 1 6. 

HT: S : Nile. Caledonie, Mr. Rolle. 

[Asprenas effeminatus Cari, 1913] 
emortualis Saussure, 1859: 61 [Bacteria] 

[ 9 . Bahia.] 

MHNG[59]: 1 9. 

HT 9 : Bahia, Brésil. 

[In collection as Bactridium emortuale (Saussure)] 
exornata Redtenbacher, 1908: 405 [Clonistria] 

[c? 9. Cuba. (Mus. Genf).] 

MHNG[60]:2 o\ 2 9. 

ST: 2 <?, Cuba, Mr. H. d. Saussure; 2 9, Cuba. 

[Clonistria exornata Redtenbacher, 1908] 
fecundus Cari, 1915: 189, figs 3, 4 [Brachyrhamphus] Figs 8-9 

[Neu-Caledonien: Mt. Ignambi, 29.IV1911, 9 ad., S ad.; Mt. Panié (Wald), ca. 

500m, 26.VI.1911, 9 ad.; Tao, 25.VI.1911, 9 ad., 6 ad.; Tiouaca-Tal, 23. VI- 

11.191 1, 9 juv.; Ciu, oberh. Canala, ca. 300m, 3.II.1912, S ad.; Mt. Canala, ca. 

750m, 4.XI.1911, 9 juv.; Ngoi-Tal, 15.IX.1911, 6 ad.] 



12 O. ZOMPRO & P. D. BROCK 

MHNG[48]: 1 6, 1 2 nymph. 

ST: <?: Ngoyi, 15.9.11; 9, nymph: Mt. Kanala, 4.11.11. [A second 9 is pres- 
ent, not recorded by Carl: „Ilegrope, 3.3.12"] 

Further material in NHMB. 

[Carlius fecundus Carl, 1915] 
freygessneri Redtenbacher, 1908: 421 [Bacteria] 

[ 9 . Guatemala (Mus. Genf).] 

MHNG[61]: 1 9. 

HT: 9, Guatemala, Mr. H. d. Saussure, 2/14. 

[Phanocloidea freygessneri (Redtenbacher, 1908)] 
fritzschei Zompro, 2000: 53, figs 9-21 [Gratidia] 

[Holotypus (o*): Nakhon Ratchasima, Amphoe Pak Chong, Thanon Thannarat 

km 7, 300m, 101 Q 24'50" E 14 Q 36'50" N, 18.X.1997]. Data for paratypes, see 

Zompro (2000). 

MHNG[30]: 1 6,1 9. 

PT: 1 6 , Nakhon Ratchasima, Amphoe Pak Chong, Thanon Thannarat km 7, 

300m, 101 Q 24'50" E 14 Q 36'50" N, auf Acalypha wilkesiana Muell., 

17.XII.1997, leg. I.Fritzsche; 358-63 Coll. O. Zompro; 1 9, Nakhon 

Ratchasima, Amphoe Pak Chong, Thanon Thannarat km 6, 325m, 101-21 ' E 

14 Q 32' N, 16.1.1998, leg. I. Fritzsche; 358-117 Coll. O. Zompro. 

Further type-material in BMNH, CUMZ, DEIC, MCSN, MNHN, NHMW, OX- 

UM, SMFD, SMTD, UMBB, ZMHB, ZMUH, Coll. I. Fritzsche, Coll. F 

Hennemann, Coll. O. Zompro, affiliated with ZMUK. 

[Gratidia fritzschei Zompro, 2000] 
fruhstorferi Carl, 1913: 29 [Dixippus] 

[2 9 . Insula Bawean (H. Fruhstorfer leg.). Muséum de Genève.] 

MHNG[38]:2 9. 

ST: 2 9 : Bawean, Regenzeit, Fruhstorfer. 

[Carausius fruhstorferi (Carl, 1913)] 

[In collection as Phasgania fruhstorferi (Carl)] 
fulvescens Saussure, 1868: 67 [Anophelepis] 

[Insula Nouka-Hiva.] 

MHNG [?]: Not traced. 

[Graeffea fulvescens (Saussure, 1868)] 
glaber Redtenbacher, 1906: 167 [Leocrates] 

[Sumatra (Mus. Genf).] 

MHNG [19]: 1 9. 

HT: 9 , Sumatra, Schneider. 

[Haaniella glaber (Redtenbacher, 1906)] 
godeffroyi Redtenbacher, 1908: 360, pi. 16: 12 [Nisyrus] 

[ 9 . Tongatabu, Freundschaftsinseln (Mus. Genf).] 

MHNG [49]: 1 9. 

HT: 9 , Tongatabu, Oceanie, H. d. Saussure. 

[Cotylosoma godeffroyi (Redtenbacher, 1908)] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 13 

godmani Redtenbacher, 1908: 41 1 [Bostra] Figs 33-34 

[9. Nicaragua (Coll. m.), Costarica (Mus. Genf), Guatemala (Coll. Godman).] 

MHNG [60]: 1 9 

ST: 1 9,1 egg: Costa Rica, Amer, cent., Mr. P. Biolley, 620/76. 

Further type-material in BMNH and NHMW. 

[Alienobostra godmani (Redtenbacher, 1908)] 
guadeloupensis Redtenbacher, 1908: 404 [Clonistria] 

[cT. Guadeloupe (Coll. m.), Guatemala (Mus. Stuttgart). 9: Guadeloupe (Mus. 

Genf).] 

MHNG [60]: 2 9. 

ST: 2 9 , Guadeloupe, Mr. H. d. Saussure. 

Further type-material in NHMW and SMNS and MNHN. 

[Clonistria guadeloupensis Redtenbacher, 1908] 
guatemalensis Redtenbacher, 1908: 405 [Clonistria] Figs 10-11 

[S . Guatemala (Mus. Genf).] 

MHNG [60]: 

LT, by present designation: S, Guatemala, Mr. H. d. Saussure, 2/14. 

PLT: S, Guatemala, Mr. H. d. Saussure, 3/28. 

Further type-material in ZMHB. 

[Clonistria guatemalensis Redtenbacher, 1908] 

The two specimens belong to different genera. The LT designation is part of a 

revision of South American genera of Phasmatodea by O. Zompro and will 

support nomenclatural stability. 
guerini Saussure, 1868: 64 [Pygirhynchus] 

[Insula Guadalupe.] 

MHNG [50]: 2 o\ 1 6 nymph, 4 9. 

ST: 2 â , 1 3 nymph, 4 9 : Guadeloupe, H. d. Saussure. 

[Lamponius guerini (Saussure, 1868)] 
hasenpuschorum Brock gen. n., sp. n. [Parapodacanthus] 

MHNG : 1 3 . 

PT: Cairns, "Nov. Holl". 

HT and further PT in QMBA, ANIC and Coll. P. D. Brock. 

Description below. 
horridum Carl, 1913: 3 [Acanthoclonia] 

[1 cT. Banos, Ecuador. Muséum de Genève.] 

MHNG [4]: 1 S. 

HT S : Banos, Ecuador. 

[Acanthoclonia horrida Cari, 1913] 
humberti Saussure, 1861a: 472 [Bacillus (Ramulus)] 

[â 9 . Lille de Ceylan, par M. A. Humbert.] 

MHNG [29]: 1 6,2 9. 

LT, by present designation: 1 â , PLT : 2 9 : Trincomalie, Ceylon, Voyage 

Humbert. 

[Ramulus humberti (Saussure, 1861)] 

[In collection as Dubreulia humbertiana (Saussure).] 



14 O. ZOMPRO & P. D. BROCK 

Comment. This is the type-species of Ramulus Saussure, 1861. Most of the 

species belonging to this genus erroneously have been treated as members of 

Baculum Saussure, 1861 (Type: Baculum ramosum Saussure, 1861), a mono- 

typic genus from Brazil. See also comments under ramosum. To support stabil- 
ity in the use of the names in the future, a lectotype is here designated for R. 

humberti. 
humberti Carl, 1913: 13 [Clitumnus] 

[1 o\Ceylan.] 

MHNG[25]: 1 6. 

HT S : Ceylon, Schilling, ex coll. Fruhstorfer. 

[Ramulus carli Zompro, nom. n.] 

Comment. This species is a member of Ramulus Saussure, 1861, type-species: 

Ramulus humberti (Saussure, 1861), and therefore requires a new name. 
humbertiana Saussure, 1868: 68 [Necroscia] 

[Ceylon.] 

MHNG[87]: 2 9. 

ST: 2 9 : Ceylon, Humbert. 

[Sosibia humbertiana (Saussure, 1868)] 
imbellis Brunner v. Wattenwyl, 1907: 312 [Ocnophila] 

[9. ? (Mus. Genav.)] 

MHNG[41]: 3 9. 

ST: 3 9:0 Amérique, coll. Guerin. 

Comment. The specimens are labelled as S , but all three type specimens are in 

fact 9 . 
impennis Carl, 1913: 36 [Asprenas] 

[19. Nova Caledonia. Muséum de Genève.] 

MHNG[49]: 1 9. 

HT 9 : Nile. Caledonie, Mr. Rolle. 

[Asprenas impennis Carl, 1913] 

Comment. A â with same data also present, neither recorded nor described by 

Carl. 
incerta Carl, 1913: 54 [Aruanoidea] 

[3 â . Than-Moi, Tonkin. Muséum de Genève.] 

MHNG[86]: 3 ó\ 

ST: 3 â: Tonkin,. Than-Moi, Jan.-Juli, Fruhstorfer. 

[Necroscia incerta (Carl, 1913)] 
inconspicuus Brunner v. Wattenwyl, 1907: 334 [Bacunculus] 

[6 . Costa Rica (Mus. Genav., Coll. m.), Mexico (Mus. Paris).] 

MHNG[45]:2 ö\ 

ST: 2 o\ Costa Rica, Amer. Centr., P. Biolley. 

Further type-material in NHMW and MNHN. 

[Pseudosermyle inconspicua (Brunner v. Wattenwyl, 1907)] 
ingens Redtenbacher, 1908: 453 [Pharnacia] 

[9. Malabar, Meppadi (Mus. Genf); Obes-Tenasserim 4000-6000' (Wood- 
Mas.).] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 15 

MHNG[69]: 1 9. 

ST: 9 , Meppadi, Malabar, Mr. Atzenwiller. 

Further type-material in the Wood-Mason collection, which is now possibly in 

New Delhi, India. 

[Pharnacia ingens Redtenbacher, 1908] 
ingerens Brunner v. Wattenwyl, 1907: 206 [Cuniculina] 

[ 9 . Java (Mus. Genav.).] 

MHNG[26]: 1 9. 

HT: 9 , Java, Fruhstorfer. 

[Ramulus ingerens (Brunner v. Wattenwyl, 1907)] 
insignis Redtenbacher, 1906: 90 [Paranisomorpha] Fig. 14 

[c?9. Costa Rica, Zentral- Amerika (Mus. Genf [<?], Coll. m, Coll Goudot 

[9]).] 

MHNG[8]: 1 9. 

9, syntype: Costa Rica, Amer. Cent, Biolley. 

Not ó* , as recorded by Redtenbacher. Further type-material in BMNH, NHMW. 

[Paranisomorpha insignis Redtenbacher, 1906] 
intermedia Carl, 1913: 40 [B ostro] 

[19. Guatemala. Muséum de Genève.] 

MHNG[60]: 1 9. 

HT 9 : Guatemala, H. d. Saussure. 

[Phanocles intermedins (Carl, 1913)] 
jamaicana Zompro, 2001a: 214, figs 48-49, 96-97, 132-133 [Caribbiopheromera] 

[HT ô , Jamaica, PSG-culture, origin: Jamaica. Culture O. Zompro VII. 1995 

(MHNG, ex coll. O. Zompro 118-2); PT: 1 9,2 eggs (MHNG, ex coll. O. 

Zompro 118-5); 2 6, 3 9, several eggs: as HT (OZ 118-1, 3-4, 6-7); 1 c?, 3 9 

(Coll. F. Hennemann 61-1-4).] 

MHNG: 1 6, 1 9, 2 eggs. 

HT: Ó, Zucht O. Zompro VII.1995; 118-2 Coll. O. Zompro; 

PT: 1 9 , Zucht O. Zompro VII. 1995; 118-5 Coll. O. Zompro. 

Further type-material in Coll. O. Zompro, affiliated with ZMUK, and Coll. F. 

Hennemann. 

[Caribbiopheromera jamaicana Zompro, 2001] 
jurinei Saussure, 1868: 69 [Metriotes] 

[No locality mentioned.] 

MHNG [15]: 1 9. 

HT: 9 : Amasm, Jurinei. 3 6 , not referred to by Saussure, also present. 

[I sagoras jurinei (Saussure, 1868)] 
linearis Saussure, 1868: 65 [Ceroys] 

[America meridionalis.] 

MHNG [?]: Not traced. 
lobulatus Carl, 1913: 12 [Eucarcharus] 

[19. Ceylon. Muséum de Genève.] 

MHNG [24]: 1 9. 



16 O. ZOMPRO & P. D. BROCK 

HT 9 : Ceylon. 

[Eucarcharus lobulatus Carl, 1913] 
lobulatus Redtenbacher, 1908: 351 [Laciphorus] 

[ 9 . Lima, Peru (Mus. Genf).] 

MHNG [?]: Not traced. 

[Laciphorus lobulatus Redtenbacher, 1908] 
longimana Saussure, 1859: 61 [Bacteria] 

[Bahia.] 

MHNG [59]: 2 S. 

ST: 2 ô : Amer, merid. 

[In collection as Bactridium longimanum (Saussure).] 
longiscapha Redtenbacher, 1908: 435 [Diapherodes] 

[ 9 . Portorico (Mus. Berlin, Mus. Genf).] 

MHNG [65]: 1 9. 

ST: Porto-Rico, Antilles, Mr. H. de Saussure. 

Further type-material in ZMHB. 

[Diapherodes longiscapha Redtenbacher, 1908] 
magayon Zompro sp. n. [Lonchodes] Figs 41-42, 56 

MHNG[39bis]: 1 6. 

HT: 9 , Philippinen, Luzon, Mt. Banahao. 

Description below. 
malaccae Saussure, 1868: 69 [Necroscia] Fig. 16 

[Malacca.] 

MHNG [81]: 1 9. 

HT 9 : Indes or., Ane. coll. 

[Asceles malaccae (Saussure, 1868)] 
melanocephala Cari, 1913: 18 [Cuniculina] 

[2 S . Lompa-Battau, Celebes merid. 3000'; 1 â . Bua-Kraeng, Celebes merid. 

5000'. Muséum de Genève.] 

MHNG [26]: 3 S. 

ST: 2 â: S. Celebes: Lompa-Battau, 3000 ft., III. 1896, Fruhstorfer; 1 o": Bua- 
Kraeng, 5000 ft., III. 1896, Fruhstorfer. 

[Ramiti us melanocephalus (Carl, 1913)] 
melanocephala Carl, 1913: 5 [Orobia] 

[5 S . Madagascar. Muséum de Genève.] 

MHNG [6]: 5 ó\ 

ST: 5 â : Madagascar, H. d. Saussure. 

[Leiophasma melanocephala (Cari, 1913)] 
mexicanus Saussure, 1859: 62 [Acanthoderus] 

[Mexico.] 

MHNG [41]: 1 9. 

HT 9 : Mexico, Potrero. 

[Sermyle mexicana (Saussure, 1859)] 

[In collection as Ocnophila mexicana (Saussure)] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 17 

mexicanus Saussure, 1859: 63 [Prisopus] 

[Mexico.] 

MHNG[16]: 1 9. 

HT 9 : Mirador, Mexique. 

[Prisopus mexicanus Saussure, 1859] 

(In collection as Prisopus berosus Westwood.) 
minor Carl, 1913: 26 [Pseudostheneboea] 

[2 S . Khasi-Hills, Assam. Muséum de Genève.] 

MHNG[33]:2 S. 

ST: 2 c?:Khasi Hills. 

[Pseudostheneboea minor Carl, 1913] 
minuta Redtenbacher, 1908: 480 [Parasipyloidea] 

[C.Ceylon (Mus. Genf).] 

MHNG: Not traced. 

[Parasipyloidea minuta Redtenbacher, 1908] 
modesta Brunner v. Wattenwyl, 1907: 206 [Cuniculina] 

[9 . Java (Mus. Genav., Mus. Hamb., Mus. Budap.), Java orient. (Mus. Berol.)] 

MHNG [26]: 1 9. 

ST: 1 9 , Java, Fruhstorfer. 

Further type-material in ZMHB and ZMUH. HNHM material was lost in fire in 

1956. 

[Ramulus modestus (Brunner v. Wattenwyl, 1907)] 
modesta Carl, 1913: 6 [Orobia] 

[Id. Madagascar. Muséum de Genève.] 

MHNG [5]: 1 6. 

HT â : Madagascar, H. d. Saussure. 

[Leiophasma modesta (Carl, 1913)] 
monticola Carl, 1913: 41 [Bacteria] 

[19. Costa-Rica (?) (W. Pittier leg.). Muséum de Genève.] 

MHNG [61]: 1 9. 

HT $ : Costa Rica, Buenos Aires a Calamanca, W. Pittier, 12.IV.97. 

[Phanocloidea monticola (Carl, 1913)] 
moricula Redtenbacher, 1908: 545 [Sipyloidea] 

[6 . Kina-Balu, Brunei, Borneo (Coll. m., Mus. Genf).] 

MHNG [90]: 2 6. 

ST: 1 S, Borneo, Iles de Sonde, Mr. Fruhstorfer, 620/27; 1 â, Brunei, E 

Borneo, Staudinger. 

Further type-material in NHMW. 

[Sipyloidea moricula Redtenbacher, 1908] 
munda Redtenbacher, 1908: 525 [Aruanoidea] 

[6 9. Philippinen (Coll. m., Mus. Genf).] 

MHNG [84]: 1 9. 

ST: 9 , Philippines. 

Further type-material in NHMW. 



18 O. ZOMPRO & P. D. BROCK 

[Necroscia munda (Redtenbacher, 1908)] 

[In collection as Necroscia munda (Redtenbacher)]. 
mystica Brunner v. Wattenwyl, 1907: 233, pi. 9: 8 [Zehntneria] 

[9. Promont. Bonae Spei (Mus. Genav.).] 

MHNG[31]: 1 9. 

HT: 9 , Cap. b. sp., Peringuey. 

[Zehntneria mystica Brunner v. Wattenwyl, 1907] 

Comment. Cap. b. sp. refers to Cape of Good Hope, Republic of South Africa. 
nanus Carl, 1913: 52, pi. 1: 9 [Paramenexenus] 

[1 â , 1 9 . Tonkin. Muséum de Genève.] 

Type-series: 1 6,1 9 . 

MHNG[76]: 1 ó\ 9. 

ST: 1 6 , 9 , syntypes: Tonkin, Baudet. 

[Paramenexenus nanus Carl, 1913] 
nuptialis Redtenbacher, 1908: 410 [Bostra] 

[6 , Guyana (Mus. Genf).] 

MHNG [?]: Not traced. 
obtusecornuta Redtenbacher, 1908: 409 [Bostra] 

[6 9. Guatemala (Coll. m. Alk.), Costarica (Mus. Berlin, Mus. Genf, Coll. m.).] 

MHNG [60]: 1 6. 

ST: 1 6, Costa Rica, Amer, merid., Mr. P. Biolley. 

Further type-material in NHMW and ZMHB. 

[Paracalynda (?) obtusecornuta (Redtenbacher, 1908)] 
occipitalis Carl, 1913: 4 [Orobia] 

[99. Madagascar. Muséum de Genève.] 

MHNG [6]: 5 9. 

ST: 5 9 : Madagascar, H. d. Saussure. [Ail 9 seem to be types; Carl did not refer 

to a definite number] 

[Leiophasma occipitalis (Carl, 1913)] 
olmeca Saussure, 1869: 156 [Bacteria] 

[6 . Le Mexique. Cordillère orientale.] 

MHNG [45]: 1 6. 

HT 9 : Mexique. 

[Pseudosermyle olmeca (Saussure, 1859)] 

[In collection as Bacunculus olmecus (Saussure)] 
palawanica Cari, 1913: 30 [Prisomera] Figs 35-36 

[1 9 . Palawan. Muséum de Genève.] 

MHNG [39]: 19,1 egg ex ovipositor. 

HT 9, 1 egg ex ovipositor: Philippines, Palawan, 1898, Doherty, ex coll. 

Fruhstorfer. 

[Stheneboea palawanica (Cari, 1913) comb, n.] 

[In collection as Lonchode s palawanica (Cari)]. 

Comment. The egg is described below. 
parvulus Carl, 1913: 27 [Acacus] 

[6 6. Java (Dr. L. Zehntner leg.). Muséum de Genève.] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 19 

MHNG[35]: 8 6. 

ST: 8 â : Java, Zehntner. 

Further material in ANSP. 

[Parasipyloidea aenea (Carl, 1913), synonymised by Günther, 1934: 529.] 
parvulus Carl, 1913: 35 [Bacunculus] 

[1 â . San Pedro Sula, Honduras. Muséum de Genève.] 

MHNG[45]: 1 S. 

HT â : Honduras, San Pedro Sula, ex coll. Fruhstorfer. 

[Pseudosermyle parvula (Cari, 1913)] 
parvulus Cari, 1913: 24, pi. 1: 12, 13 [Micrarchus] Figs 12-13 

[1 â Greymouth, 1 S Heteraunga (Nova Seelandia). Muséum de Genève.] 

MHNG[29]:2 Ô. 

ST: S : Greymouth, Nile. Zealand, Helms; S : Nov. Zealand, Heteraunga. 

[Micrarchus hystriculeus (Westwood, 1859), synonymized by Salmon, 1999: 

88.] 

Comment. Salmon (1991: 88) synonymized Micrarchus Carl, 1913 (not of 

Kaup) (Type-species: Micrarchus parvulus Carl, 1913: 24, by monotypy) with 

Pachymorpha Gray, 1835 (Type-species: Bacillus squalidus Gray, 1833: 18, by 

monotypy). The genitalia of these genera show striking differences, therefore 

Micrarchus is re-established as a valid genus. The terminal abdominal segments 

of a pair of Pachymorpha squalida (Gray, 1835) from New South Wales 

[MHNG] are figured in figs 31-32. 
pedestris Redtenbacher, 1908: 540 [Candaules] 

[ 9 . Java (Mus. Genf.).] 

MHNG [87]: 1 S. 

HT: S , Java, H. d. Saussure. 

[Lopaphus pedestris (Redtenbacher, 1908)] 
peruana Saussure, 1868: 65 [Bacteria] 

[Peru.] 

MHNG [42]: 2 ó\ 

LT: S: Iquitos, Pérou, 600/81, Ane. coll. PLT: 6: Pérou. 

[Oreophoetes peruana (Saussure, 1868)] 

[In collection as Allophyllus peruanus (Saussure)] 

Comment. Also 1 S and 1 9 from Banos, Ecuador present, but not referred to 

by Saussure. 

LT designated by Zompro, 2002: 146. 
pictetì Redtenbacher, 1908: 522 [Aruanoidea] 

[e?, Borneo (Mus. Genf).] 

MHNG [84]: 1 S. 

HT: â , Borneo, Iles de Sonde, Mr. C. Pictet. 

[Scionecra picteti (Redtenbacher, 1908)] 
poeyi Saussure, 1868: 67 [Anophelepis] Figs 17, 58 

[Insula Cuba.] 

MHNG [14]: 2 9. 

LT, by present designation: 9 : Cuba, acheté a M. Poey; PLT: 19,9: Cuba, 



20 O. ZOMPRO & P. D. BROCK 

acheté a M. Poey; 3 9 : Cuba, M. H. de Saussure. 

[Alloeophasma poeyi (Saussure, 1868)] 

Comment. This lectotype designation is part of a revision of the Phasmatodea: 

Areolatae by O. Zompro and will support nomenclatural stability. 
poeyi Saussure, 1868: 68 [Creoxylus] 

[Larva. Cuba.] 

MHNG [16]: 1 6 nymph. 

HT 6 , nymph: Cuba, Antilles, acheté a M. Poey. 

[Creoxylus poeyi Saussure, 1868] 
prima Zompro, 1999: 124, figs 1-3 [Microphasma] 

[Ceylan (Central), Horton plains, 2100m, 15.11.1970, leg. C. Besuchet / 1. Lobi] 

MHNG [33]: 1 $ 

HT: 9, Ceylan (Central), Horton plains, 2100m, 15.11.1970, leg. C. Besuchet / 

I. Lobi. 

[Microphasma prima Zompro, 1999] 
proximus Carl, 1913: 28 [Carausius] 

[1 ô . Java, Dr. L. Zehntner. Muséum de Genève.] 

MHNG [36]: 1 ó\ 

HT: â : Java, Zehntner. 
pulchrepicta Cari, 1913: 22 [Gratidia] Figs 18-19 

[1 S . Promont. Bonae Spei. Muséum de Genève.] 

MHNG [30]: 1 o\ 

HT S: Cap b. Sp., Peringuey. 

[Gratidia pulchrepicta Cari, 1913] 

Cap. b. sp. refers to Cape of Good Hope, Republic of South Africa. 
pusillus Redtenbacher, 1906: 61 [Ceroys] 

[<?. Peru (Mus. Genf)] 

MHNG [4]: 1 S. 

HT: 6 , Peru. 

[Ceroys pusillus Redtenbacher, 1906] 
putingmantsa Zompro sp. n. [Lonchodes] Figs 37-40, 54-55 

MHNG [39bis]: 1 ó\ 1 $. 

HT: â , Philippinen, Aroroy. 

PT: 1 $ , Philippinen. Aroroy. 

Description below. 
quitensis Saussure, 1868: 69 [Phasma] 

[Aequator.] 

MHNG [13]: 1 9. 

HT 9 : Quito, Pérou. H. de Saussure. 

[Pseudophasma quitensis (Saussure, 1868)] 
rachis Saussure, 1868: 64 [Acanthoderus] Figs 20-21 

[Nova Caledonia.] 

MHNG [49]: 4 o\ 1 9. 

ST: 4 6, 1 $ : Nile. Caledonie, Oceanie, Mr. Deyrolle. 

[Cnipsus rachis (Saussure, 1868)] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 21 

ramosum Saussure, 1861b: 127 [Bacillus (Baculum)] 

[ 9 . Brasilia] 

MHNG [?]: Not traced. 

[Baculum ramosum (Saussure, 1868)] 

Comment. This species is the type of the problematic genus Baculum Saussure, 

1861. Baculum ramosum originates from Brazil, while all other species 

described in "Baculum' are from Asia. Furthermore, Baculum ramosum shows 

a character not present in any species descibed in this genus subsequently. All 

these species actually belong in Romulus Saussure, 1861, and, to avoid further 

confusion, are listed within this genus. A revision of the whole genera around 

Baculum and Ramulus is in progress by Zompro. 
reclusa Brunner v. Watten wyl, 1907: 327 [Dyme] 

[Terra caliente in Mexico. (Sauss., Mus. Genav., Coll.m.)] 

MHNG [44]: 1 9. 

ST: 1 9 , Mexico. 

Further type-material in NHMW. 
rossia Rossi, 1788: 259, pi. 8: 1 [Pseudomantis] 

MHNG [coll. Harz 1A]: 1 9. 

NT: 9, Spanien, Port Bon, 28.5.66, Dr. Eckerlein leg. 

Invalid neotype designation for a species described from Pisa, Italy - correct 

neotype designated by Capra & Poggi (1980). Specimen deposited in MCSN. 

[Bacillus rossius Rossi, 1788] 
rouxi Carl, 1915: 174, 187, fig. 1 [Labidiophasma] Fig. 22 

[Ngoi-Tal, 15.X.1911, 2 9.] 

MHNG [48]: 1 9. 

ST: 9,Ngoye, 15.9.11. 

Further material in NHMB. 

[Labidiophasma rouxi Carl, 1915] 
rubescens Saussure, 1868: 68 [Necroscia] Fig. 23 

[Malacca.] 

MHNG [83]: 1 9. 

HT 9 : Malacca, Indes, Guerin. 

[Marmessoidea rubescens (Saussure, 1868)] 

[In collection as Trigonophasma rubescens (Saussure)] 
rubus Saussure, 1861 [Cladoxerus] 

[9. Brasilia. (Bahia.)] 

MHNG [63]: 3 9 9. 

ST: 2 9 : Brésil; 9 : without label. 

[Cladomorphus rubus (Saussure, 1861)] 

[In collection as Phibalosoma rubus (Saussure)] 
rufopectus Redtenbacher, 1908: 416 [Bacteria] 

[o\ Cayenne (Mus. Genf).] 

MHNG [61]: 1 S. 

HT: S, Cayenne, Prudhomme, 620/85. 

[Phanocloidea rufopectus (Redtenbacher, 1908)] 



22 O. ZOMPRO & P. D. BROCK 

saginata Redtenbacher, 1906: 86, pi. 4:10 [Dina] 

[â 2 . Borneo, Brunnei, Kina ] 

MHNG [8]: 

ST: S , Brunei, E Borneo, Staudinger. 

Further type-material in NHMW. 

[Dinophasma saginata (Redtenbacher, 1906)] 
saussurei Kirby, 1904: 397 [Haaniella] 

[nom. n.] 

MHNG. [19]: 1 9. 

HT: 9 , Coll. Guerin. 

[Haaniella saussurei Kirby, 1904] 

(In collection as Heteropteryx saussurei Redtenbacher.). 

Comment. Introduced as a name for Heteropteryx grayi var. Saussure, 1869: 

310, which subsequently was also named saussurei by Redtenbacher, see below. 
saussurei Redtenbacher, 1906: 171 [Heteropteryx] 

[ 9 . Sunda-Inseln (Mus. Genf); Borneo (Hofmus. Wien, Mus. Budapest, Mus. 

Genua.] 

MHNG [19]: 1 9. 

ST: 9 , Coll. Guerin. 

Further type-material in HNHM, NHMW and MCSN. 

[Haaniella saussurei Kirby, 1904] 

[In collection as Heteropteiyx saussurei Redtenbacher] 

Comment. The same specimen as the above mentioned holotype of Haaniella 

saussurei Kirby, 1904, being type-specimen of both authors. 
saussurei Bolivar, 1888: 136 [Phantasis] 

[ 9 . Cuba. No. 104 9 de la coll. Gundlach.] 

MHNG [49]: 1 9. 

ST: 9 , Cuba, Saussure. 

Further type-material in MNMS. 

[Hesperophasma saussurei (Bolivar, 1888)] 

[In collection as Phantasis planula (Westwood)] 
schauinslandi Brunner v. Wattenwyl, 1907: 238, pi. 10: 3a, b [Argosarchus] 

[9. Ins. Chatam prope N. Seelandiam (Mus. Bremense), N. Seelandia (Mus. 

Berol., Mus. Genav.)] 

MHNG [32]: 1 9. 

ST: 1 9 , New Zealand, Greymouth, Helms. 

Further type-material in UMBB and ZMHB. 

[Argosarchus horridus (White, 1846), synonymized by Salmon, 1991: 48.] 
schneiden Werner, 1918: 268 [Battacus] 

[Bintang Mariah, Montes Battale] 

MHNG [82]: 1 9. 

HT: 9 , Sumatra, Binthang Mariah, Mt. Battak. 

[Tagesoidea nigrofasciata Redtenbacher, 1908, synonymized by Brock, 1990: 

14.] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 23 

schulthessi Carl, 1913: 21, pi. 1: 7, 8 [Paraleptinia] Figs 24-27 

[1 9 . Promont. Bonae Spei. Muséum de Genève.] 

MHNG [30]: 

HT 9 : Cap b. Sp., Peringuey. 

[Paraleptinia schulthessi (Carl, 1913)] 

Cap. b. sp. refers to Cape of Good Hope, Republic of South Africa. 
scutatus Carl, 1913: 49, pi. 1: 1,3 [Andropromachus] Fig. 28 

[1 $ . Tonkin. Muséum de Genève.] 

MHNG [77]: 1 9. 

HT 9 : Tonkin, Baudet. 

[Andropromachus scutatus Carl, 1913] 
secunda Carl, 1913: 31 [Libethra] 

[16. Merida, Venezuela. Muséum de Genève.] 

MHNG [41]: 1 6. 

HT 6 : Venezuela, Merida. 

[Rugosolibethra secunda (Cari, 1913)] 
segregata Carl, 1913: 26, pi. 1: 2 [Pseudostheneboea] Fig. 29 

[1 6 . Khasi-Hills, Assam. Muséum de Genève.] 

MHNG [33]: 1 6. 

HT 6 , holotype: Khasi Hills. [A second 6 with same data is present, but not 

referred to by Carl] 

[Pseudostheneboea segregata Carl, 1913)] 
soror Carl, 1913: 32 [Libethra] 

[1 $ . Merida, Venezuela. Muséum de Genève.] 

MHNG [41]: 1 $. 

HT 9 : Merida. 

[Rugosolibethra soror (Carl, 1913)] 
speisen Carl, 1915: 193 [Megacrania] 

[Malo, XI.1911, 8 9 ad., 2 9 juv.] 

MHNG [51]: 1 9. 

ST 9 : Malo (N. Hebr.), Dr. Speiser. 

Further material in NHMB. 

[Megacrania bate sii spei seri Carl, 1915] 

(In collection as Megacrania batesi var. speiseri Carl.) 
speisen Carl, 1915: 193 [Hermarchus] 

[Mal, XI.1911, 2 9;Ambrym, 111.1911,2 9.] 

MHNG [68]: 1 9. 

ST: 9 : Malekula, Neue Hebriden, Mai 191 1, D. Speiser. 

Further material in NHMB. 

[Hermarchus inermis speiseri Carl, 1915] 
spinigera Saussure, 1859: 62 [Bacteria] 

[6 . Brasilia.] 

MHNG [?]: Not traced. 

[Canuleius euterpinus (Westwood, 1859), synonymized by Kirby, 1904: 353.] 



24 O. ZOMPRO & P. D. BROCK 

spinipennis Carl, 1913: 47 [Oxyaries] 

[1 S . Tonkin. Muséum de Genève.] 

MHNG[76]: 1 ó\ 

HT S : Tonkin, Baudet. 

[Oxyaries spinipennis Cari, 1913] 
spinosissimus Cari, 1913: 46 [Oxyartes] 

[ 1 S . Tonkin. Muséum de Genève.] 

MHNG[76]: 1 6. 

HT â: Tonkin, Baudet. 

[Oxyartes spinosissimus Cari, 1913] 
spinosus Saussure, 1868: 68 [Lopaphus] Fig. 30 

[Malacca.] 

MHNG[79]: 1 ó\ 

HT ó* : Malacca, Indes, Guerin. 

[Centrophasma spinosum (Saussure, 1868)] 
subintegra Carl, 1913: 20 [Medaura] 

[1 $ . Khasi-Hills, Assam. Muséum de Genève.] 

MHNG[30]: 1 $. 

HT ? : Khasi Hills, Assam. 

[Medaura scabriuscula (Wood-Mason, 1873), synonymized by Brock & 

Cliquennois, 2000: 15.] 
submarginata Redtenbacher, 1908: 532 [Diacanthoidea] 

[â 9. Sumatra (Mus. Genf); Perak (Mus. Hamburg, Westw.).] 

MHNG[86]:2 6. 

ST: 2 S , Sumatra, lies de Sonde, Dr. H. Dohrn. 

Further type-material in NHMW and ZMUH. 

[Diacanthoidea diacanthos (de Haan, 1842), synonymized by Brock, 1995: 88.] 
sumatranus Brunner v. Wattenwyl, 1907: 184 [Phobaeticus] 

[ 9 . Sumatra (Mus. Genav.), Indraviri in ins. Sumatra (Mus. Basil.).] 

MHNG[23]: 1 9. 

ST: 9 , Sumatra, Lakat, acheté d. M. Schneider. 

Further type-material in NHMB. 

[Pharnacia sumatrana (Brunner v. Wattenwyl, 1907)] 
superbus Carl, 1913: 15 [Clitummts (?)] 

[2 S . Than-Moi, Tonkin. Muséum de Genève.] 

MHNG[25]:2 6. 

ST: 2 S : Tonkin, Than-Moi, Jan.-Juli, Fruhstorfer. 

[Ramni us superbus (Carl, 1913)] 
suspecta Carl, 1913: 55 [Calvisia] 

[19. Bornéo sept. Muséum de Genève.] 

MHNG[93]: 1 9. 

HT 9 : Nord Borneo, ex coll. Fruhstorfer. 

[Calvisia suspecta Carl, 1913] 
tarsatus Carl, 1913: 23 [Mimarchus] 

[3 9 . Greymouth, Blue Cliffs, Nova Seelandia. Muséum de Genève.] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 25 

MHNG[31]: 3 6. 

ST: S : Nov. Zealand; 6 : Nile. Zealand, Marcet; S : Greymouth, Blue Cliffs, 

Nile. Zealand, Helms. 

[Mimarchus tarsatus Carl, 1913] 

Comment. Jewel & Brock (in press) have corrected the present classification. 
thomae Saussure, 1868: 64 [Pygirhynchus] 

[Insula Saint-Thomas.] 

MHNG[?]: Not traced. 

[Agamemnon thomae (Saussure, 1868)] 
tolteca Saussure, 1859: 62 [Bacteria] 

[S . Montes mexicani.] 

MHNG [?]: Not traced. 
transiens Redtenbacher, 1908: 539, pi. 26: 7 [Candaules] 

[6 9 . Java (Coll. m., Mus. Genf, Mus. Hamburg, Mus. Berlin, Mus. Budapest).] 

MHNG [87, 88]: 

ST: [Drawer 87] 5 ó\ 9 5 , [Drawer 88] 13 d, Zehntner, Java. 

Further type-material in HNHM, NHMW, ZMHB and ZMUH. 

[Lopaphus transiens (Redtenbacher, 1908)] 
triangulifera Redtenbacher, 1906: 124, pi. 5: 2 [Oestrophora] 

[8 9 . Surinam, Cayenne (Mus. Paris, Mus. Genf, Coll. m.)] 

MHNG [14]: 1 ó\ 

ST: 1 d, Surinam, Amer, mer., coll. Jurin. 

Further type-material in MNHN and NHMW. 

[Oestrophora triangulifera Redtenbacher, 1906] 
tricolor Redtenbacher, 1908: 570 [Calvisia] 

[9. Sumatra (Coll. m.); Brunei, Borneo (Mus. Genf).] 

MHNG [93]: 1 9. 

PLT: 1 9 , Brunei, E Borneo, Staudinger. 

LT in NHMW. 

[Calvisia hemus (Westwood, 1859), synonymized by Brock, 1995: 87.] 

Comment. LT designated by Brock, 1995: 87. 
trilineatus Carl, 1913: 28 [Myronides] 

[ô S . Silhet, Assam. Muséum de Genève.] 

MHNG [35]: 5 S. 

ST: 5 ô : Silhet, Assam. 

[Lopaphus trilineatus (Carl, 1913) comb, n.] 
unicolor Carl, 1913: 8, pl. 1: 10 [Eudes (?)] 

[1 ô , 1 9 . La Palma, 1600m, Costarica. (P Biolley leg.). Muséum de Genève.] 

MHNG [10]: 1 6, 1 9. 

ST: 1 <?, 1 9 : Costa Rica, La Palma, 1600m, Biolley. 

[Eucles unicolor Carl, 1913] 
unispinosa Carl, 1913: 1 [Antongilia] 

[là, Madagascar. Muséum de Genève.] 

MHNG[1]: 1 S. 



26 O. ZOMPRO & P. D. BROCK 

HT â : Madagascar, V. Heyden. 

[Antongilia unispinosa Carl, 1913] 
verruculosa Brunner v. Wattenwyl, 1907: 248 [Stheneboea] 

[6. Java (Coll. m., Mus. Genav.), Perak (Coll. m.)] 

MHNG[33]:9 o\ 

PLT: 9 6 , Java, Zehntner. 

Also 5 undescribed 9 with same data. Further type-material in NHMW. 

[Stheneboea verruculosa (Brunner v. Wattenwyl)] 

LT in NHMW designated by Brock (1995: 86). The Java material is considered 

to represent a different species by Brock (1995: 86). 
virescens Redtenbacher, 1906: 83 [Abrosoma] 

[a 9 . Ceylon (Coll. m.); Andamanen (Hofmus. Wien); Sumatra (Mus. Genf).] 

MHNG [?]: Not traced. 

A specimen that is possibly a type in NHMB. Further type-material in NHMW. 

[Abrosoma virescens Redtenbacher, 1906] 

LT designated by Brock, 1998a: 64. 
viridis Zompro, 1998: 163, fig. 10 [Breviphetes] 

[Holotypus: S , (Neuguinea) Kaiser Wilhelmsland, Torricelli-Gebirge, 1910-, Dr. 

Schlaginhaufen (MHNG). - Allotypus: 9 , Daten wie Holotypus (SMTD).] 

MHNG [33] : 1 o\ 

HT: â , Kais. Wilhelmsland, Toricelli Gebirge, 780m, Dr. Schlaginhaufen. 

Further material in SMTD. 

[Breviphetes viridis Zompro, 1998] 
yersiniana Saussure, 1868: 65 [Bacteria] 

[Porto-Rico.] 

MHNG [43]: 1 6. 

HT: â , Puerto Rico, Yersin. 

[Bacteria yersiniana (Saussure, 1868)] 

[In collection as Dyme yersiniana (Saussure)] 
zehntneri Redtenbacher, 1908:421 [Bacteria] 

[9 . Guatemala (Mus. Genf); Vera Cruz (Coli. Godman).] 

MHNG [?]: Not traced. 

Type-material in BMNH. 

[Phanocles zehntneri (Redtenbacher, 1908)] 
zehntneri Carl, 1913: 53 [Centema] 

[4 S . Simpar, in ins. Java. (Dr. L. Zehntner.). Muséum de Genève.] 

MHNG [80]: 4 6. 

ST: 4 o* : Java, Zehntner. 

[Centrophasma zehntneri (Carl, 1913)] 
zehntneri Redtenbacher. 1908: 479 [Parasipyloidea] 

[9 . Kandy, Ceylon (Mus. Genf).] 

MHNG [?]: Not traced. 

ST 9 present in NHMW. 

[Parasipyloidea zehntneri Redtenbacher, 1908] 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 27 

zomproi Fritzsche & Gitsaga, 2000: 11-14, 2 figs [Parapachymorpha] 

[Holotyp 6: Thailand, Nakhon Ratchasima, S Khao Mai Pok, 900- 100m, 

101°19'E 14°31'N, 19.-25.X.1997, leg. I. Fritzsche (Coll. O. Zompro)]. Data 

for Paratypes, see Fritzsche & Gitsaga (2000). 

MHNG[30]: 1 S, 1 9. 

PT: 1 o\ 1 9, Zucht O. Zompro 1.1999; 349-64, 100 Coll. O. Zompro. 

Further type-material in DEIC, ISNB, MCSN, NHMW, SMTD, ZMHB, 

ZMUH, ZSMC, Coll. I. Fritzsche, Coll. F. Hennemann, Coll. O. Zompro, 

affiliated with ZMUK. 

[Parapachymorpha zomproi Fritzsche & Gitsaga, 2000] 

DESCRIPTIONS 

Alienobostra godmani (Redtenbacher, 1908) Figs 33-34 

Material examined. 1 egg, ex ovipositor ST: 1 5 , Costa Rica, Amer, cent., Mr. P. Biolley, 
620/76 [MHNG, drawer 60]. In collection as Bostra godmani Redtenbacher. 

Description. Capsule brown, with flat ridges of lighter colour. Micropylar plate 
positioned in the capsule in an angle, elongated cordiform in shape, marginated. 
Capitulum yellow brown, with lateral impressions, in the middle deeply impressed. 

Measurements (mm): Total: 3.3; length: 2.8; width: 1.8; height: 1.4. 

Stheneboea palawanica (Carl, 1913) Figs 35-36 

Material examined. 1 egg, ex ovipositor HT: Philippines, Palawan, 1898, Doherty, ex 
coll. Fruhstorfer [MHNG, drawer 39]. In collection as Lonchodes palawanica (Carl). 

Description. Capsule brownish, depressed laterally, asymetrical dorsoventrally, 
capsule covered with white and brown spots, swollen ventroposteriorly. Capitulum on 
a stalk, mushroom-shaped, orange brown. Capsule marginated anteriorly. Posterior end 
projecting. Micropylar plate strongly angled, whitish brown. 

Measurements (mm): Total: 3.5; length: 3.0; width: 2.7; height: 1.8. 

Lonchodes dalawangsungay Zompro sp. n. Figs 43-44 

Material examined. Holotype, 9: Philippinen, Luzon, Imugan.[MHNG, drawer 39bis]. 
Left midleg broken off and missing, also left metatarsus. Fifth tarsomere missing in right 
metatarsus. 

Diagnosis. Lonchodini. Differing from all species known by the two very large 
spines between the eyes and the ventroposterolateral lobes of abdominal segment VII. 

Description. General colour midbrown. Body covered with lighter granulae, not 
shiny. 

Head slightly narrowed posteriad, with indistinct median line. Eyes projecting 
hemispherically, with two prominent, acute horns between them, these directed 
anteriad. Vertex with several small tubercles, posterior margin with four short, longitu- 
dinal impressions. Gula present. Scapus rectangular, flat, ventrally impressed. Pedi- 
cellus cylindrical, two thirds as wide and slightly less than half as long as scapus. 
Antennae projecting beyond middle of abdominal segment II. 

Prothorax with distinct anterior, lateral and posterior margins. With two 
tubercles anteromedially, and transverse impression medially. This impression also 



28 O. ZOMPRO & P. D. BROCK 

with two tubercles submedially. Anterior half with two longitudinal impressions 
sublaterally. Mesonotum strongly elongated, about 7.5 times as long as pronotum, irre- 
gularly granulated, with fine, sometimes indistinct, median line, and some separated 
larger tubercles laterally. Segment slightly dilating near the coxae. Mesosternum 
granulated. Metathorax structured as mesothorax, but about one third shorter. 

Profemora curved and compressed basally, triangular in cross-section, edges 
lamellate. Protibiae also triangular in cross-section, longer than profemora, edges also 
lamellate. Probasitarsus slightly longer than following three tarsomeres combined, 
with prominent crest in the anterior half. Second to fourth segment decreasing in 
length, also with dorsal crest. Fifth tarsomere curved, without dorsal crest, as long as 
third and fourth segment combined. 

Meso- and metafemora trapezoidal in cross-section, with pointed edges and 
ventromedian carina. With a broad, single spine ventrolaterally before the knee. Meso- 
and metatibiae quadrate in cross-section. Mesotibiae shorter, metatibiae longer than 
their appertaining femora. In meso- and metatarsi basitarsi carinated, but without 
prominent crest in the anterior half. Following tarsomeres also with dorsal carina, 
decreasing in length. Fifth tarsomere curved, as long as previous two segments. 

Median segment quadrate, with broad posterior margin, this margin lighter and 
less granulated, less than one fourth as long as metanotum. 

Abdominal segment II slightly more than twice as long as median segment, as 
long as III and IV. V longest segment. VI as II. VII slightly shorter than II, with ventro- 
posterolateral lobes. Sternites II to VI with a very flat, black impression before 
posterior margin. VII in distal half depress laterally, carinated medially. VIII longer 
than IX and X combined, carinated dorsally. X with a v-shaped incision posteriorly. 
Supranal plate slightly projecting, triangular, with median carina. Subgenital plate with 
median keel in the posterior half, slightly broadened in the distal half, depressed 
ventrolaterally. Cerci short, acute, bristled. 

Measurements (mm): Body: 124.5; head: 5.6; antennae: 67.5; pronotum: 3.9; 
mesonotum: 29.0; metanotum: 17.5; mediansegment: 3.9; profemora: 29.6; protibiae: 
29.9; mesofemora: 23.0; mesotibiae: 21.7; metafemora: 26.8; metatibiae: 28.8; pro- 
tarsi: 9.3; mesotarsi: 7.6; metatarsi: 7+. 

Male unknown. 

Name, "dalawang sungay", Tagalog for "two horns", refering to the two promi- 
nent spines between the eyes. 

Lonchodes magayon Zompro sp. n. Figs 41-42, 56 

Material examined. Holotype, 6: Philippinen, Luzon, Mt. Banahao [MHNG, drawer 
39bis]. Left foreleg, fifth tarsomere of left metatarsus and claws of right metatarsus broken off 
and missing. 

Diagnosis. Lonchodini. The colouration of this species resembles strikingly 
Periphetes granifer (Westwood, 1859), but magayon n. sp. differs in the form of the 
genitalia. This colouration is unique in the whole Lonchodini. 

Description. Whole body granulated, head, prothorax, posterior parts of tho- 
racic segments, knees and tibiae green, abdomen greenish red, remaining parts of body 
orange brown. 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 29 

Head elongated trapezoid, with a small, triangular impression between the eyes, 
vertex with several small tubercles. Clypeus light brown. Eyes brown, projecting hemi- 
spherically. Antennae projecting beyond abdominal tergite III. Scapus flattened rectan- 
gular. Pedicellus cylindrical, one third as long and two thirds as wide as scapus. Third 
antennomere considerably narrower and slightly longer than pedicellus. 

Pronotum slightly shorter than head, green, with distinct anterior margin and 
deep median and transverse impression; slightly granulated. Mesonotum strongly 
granulated, brown, with green median line, broadened and green in the coxal area, 
epimera also green. Mesosternum strongly granulated. Metanotum structured as 
previous segment, one third shorter. 

Profemora curved and compressed basally, four-edged, edges lamellate and 
bristled; with two green, lateral teeth anterolaterally, the exterior one prominent, the 
interior one indistinct. Probasitarsus as long as following tarsomeres combined, 
following segments decreasing in length, all tarsomeres carinated dorsally. Meso- and 
metafemora trapezoid in cross-section, edges distinct, but not lamellate, bearing very 
short, transparent brisles; ventroapically with large lateral spines. 

Abdominal segments with fine, raised median line. Mediansegment quadrate, 
abdominal segment II 2.5 times as long. II to VI of similar length and width. VII 2/3 as 
long as VI, dilated posteriad, as long as IX. Lateral margins of IX parallel, median line 
deeply impressed in the posterior third. X 1 .5 times as long as IX, posterolateral edges 
elongated triangular and projecting, these edges interiorly with long, black teeth. Cerci 
small, short, curved inwards, bristled. Subgenital plate comparably flat, slightly 
swollen. 

Measurements (mm): Body: 72.0; head: 3.2; antennae: 52.4; pronotum: 2.2; 
mesonotum: 19.5; metanotum: 12.3; mediansegment: 2.2; profemora: 19.8; protibiae: 
20.0; mesofemora: 16.4; mesotibiae: 16.6; metafemora: 18.5; metatibiae: 19.3; pro- 
tarsi: 7.9; mesotarsi: 5.6; metatarsi: 5.7+. 

Female unknown. 

Name. "magayon", Bicolano, the language of the people inhabiting the 
collecting site, for "beautiful", refering to the colouration. 

Lonchodes putingmantsa Zompro sp. n. Figs 37-40, 54-55 

Material examined. HT, S: Philippinen, Aroroy. [MHNG, drawer 39bis]. Right meta- 
tarsus and claw of left metatarsus broken off and missing. 

PT: 1 ? : Philippinen, Aroroy. [MHNG]. The abdomen is damaged by evisceration, the 
right antenna is broken off. but glued to the head. 1 S , Philippinen, Aroroy. [ZSMC]. 

Diagnosis. Lonchodini. Characterized by the white spots on genae, thorax and 
the male's terminal abdominal segments. These features distinguish putingmantsa dis- 
tinctly from all other species in Lonchodini. Body strongly granulated. 

Description S . Body brown, slightly shiny. Pro- and mesofemora green, their 
knees black. Metafemora brown, their knees also black. Tibiae greenish brown. 

Head slightly longer than wide, narrowed distad. Eyes projecting hemsipheri- 
cally, between them with two flat, but acute tubercles. Colour of clypeus lighter. Genae 
white, with black margin dorsally. Tentorium also white. Antennae projecting beyond 
abdominal tergite III. Scapus flattened rectangular, two times as wide and three times 



30 O. ZOMPRO & P. D. BROCK 

as long as pedicellus. Pedicellus cylindrical. Third antennomere half as wide and 1.5 
times as long as pedicellus. 

Pronotum with median and transverse impression, anterior margin broad, with 
two tubercles submedially. The white spots are positioned mediolaterally, also cover- 
ing part of the epimera. 

Prosternum white. Mesonotum strongly elongated and granulated, with fine, 
shiny median carina and widely separated small, transparent bristles (lOx); dilating 
slightly before coxae, this area with white spots laterally, also covering epimera. 
Mesosternum structured as mesotergum, between coxae white spotted. Metathorax as 
mesothorax, but one third broader and one third shorter. 

Profemora green, curved and compressed basally, four-edged, edges bristled; 
ventroapically with two submedian spines. Protibiae almost quadrate in cross-section, 
edges also bristled. First four tarsomeres with distinct dorsal carina. Basitarsus slightly 
longer than following four tarsomeres combined; second segment longer than third and 
fourth combined, third longer than fourth, fifth segment curved, slightly broadened 
apicad. Mesofemora and tibiae also green, trapezoid in cross-section. Mesofemora with 
two prominent, black lateral spines ventroapically. Metafemora and tibiae brown, 
structured as midlegs. Meso- and metatarsus as protarsus. 

All abdominal segments with fine, median carina. Mediansegment slightly 
longer than wide, strongly granulated and with white spots laterally. Abdominal seg- 
ments II to VII with a small (sometimes indistinct), white spot antero- and postero- 
laterally. II 2.5 times as long as median segment, II to VI of similar length, decreasing 
in width. VIII half as long as VII, dilating posteriad. Lateral margins of IX almost 
circular, whitish. X almost divided medially, white laterally, posterior ends strongly 
elongated and projecting, with striking black teeth interiorly. External vomer not pro- 
duced, cerei strongly elongated, bristled. Sternite II with two white lateral spots 
anteromedially, III to VII with a small impression posteromedially. 

Measurements â (mm): Body: 96.2; head: 4.0; antennae: 64.8; pronotum: 2.8; 
mesonotum: 24.5; metanotum: 16.1; mediansegment: 3.2; profemora: 25.0; protibiae: 
27.2; mesofemora: 18.5; mesotibiae: 19.0; metafemora: 21.9; metatibiae: 25.3; pro- 
tarsi: 7.9; mesotarsi: 6.2; metatarsi: 6.7+. 

Description 9 . More robust than 6 , without white spots, body and extremities 
uniformly midbrown. 

Head from dorsal aspect oval in shape, between the hemispherically projecting 
eyes with two small, acute spines. Clypeus lighter brown. Antennae projecting dis- 
tinctly beyond abdominal tergite II. Scapus strikingly flattened, rectangular, pedicellus 
cylindrical. Third antennomere half as wide and 1.5 times as long as pedicellus. 

Pronotum slightly shorter and narrower than head, strongly granulated, with 
deep, blackish median and transverse impression and broad margin anteriorly. 
Mesonotum strongly elongated, with light median carina and irregular, small, dark 
spots and transparent bristles (lOx), these are also present on the abdominal segments. 
Mesonotum with same structure, about 3/5 as long. 

Profemora curved and compressed basally, four edged in corss-section, edges 
lamellate and bristled. Protibiae quadrate in cross-section, edges bristled. Probasitarsi 
carinated dorsally, as long as following four tarsomeres without claws combined, 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 3 1 

following three segments decreasing in length, fifth tarsomere broadened apicad. 
Meso- and metafemora straight basally, trapezoid in cross-section, edges slightly 
lamellate and bristled. Tibiae as protibiae, meso- and metatarsi as protarsi. All knees 
black. 

Abdominal segment II twice as long as median segment, II to VII of similar 
length, VIII about 2/3 as long as VII, slightly dilating posteriad, VIII as long as IX and 
X combined, X slightly longer than IX, posteriorly with a notch. Supra anal plate pro- 
jecting, triangular. Sternites II to VII with small, black impression posteromedially. 
Subgenital plate boat-shaped, with an sharp keel ventromedially, acute posteriorly, 
projecting X by half of X's length. Cerci very short, small, elongated triangular. 

Measurements 2 (mm): Body: 124.1; head: 5.6; antennae: 71.0; pronotum: 5.0; 
mesonotum: 29.0; metanotum: 16.7; mediansegment: 4.3; profemora: 28.4; protibiae: 
30.1; mesofemora: 20.9; mesotibiae: 18.8; metafemora: 22.2; metatibiae: 24.0; pro- 
tarsi: 9.0; mesotarsi: 8.0; metatarsi: 8.8. 

Name, "puting mantsa", Tagalog for "white spotted", refers to the characteristic 
white spots on the genae, thorax and the terminal abdominal segments of the male. 

Comment. The white spots consist of a dust-like material. This kind of 
colouration was previously unknown in phasmids. The female paratype's abdomen is 
damaged by evisceration. 

Parapodacanthus Brock gen. n. Figs 46-53 

Type-species: Parapodacanthus hasenpuschorum Brock sp. n., by present 
designation. 

Diagnosis. Typical Tropidoderini. Closely related to Podacanthus Gray, 1835, 
but easily distinguished by the presence of large spine-like tubercles on the meso- 
notum, in addition to differences in egg structure. Eggs are small in this new genus, 
oval with a small capitulum; capsule slightly hairy and not ridged. In Podacanthus 
Gray, eggs are large, oval or elongate with large capitulum; capsule not hairy, with 
numerous raised ridges. 

Description. Medium sized, large winged Tropidoderini. Head slightly longer 
than wide. Eyes large. Three ocelli present. Antennae long, but shorter than length of 
fore leg. Scapus slightly broader than remaining segments. 

Pronotum slightly longer than head, with central impression. Mesonotum short, 
just over 1.5 times the length of pronotum. The other spines slanting outwards, pos- 
terior pair shorter than preceding pairs. Metanotum slightly longer than mesonotum. 
Lateral margins of meso- and metathorax with series of small tubercles. Metathorax 
with small tubercles ventrally. Tegmina long and slender, slightly shorter in â than in 
9, tapered to a round tip. Alae long with darker veins. Legs moderately long and 
slender, slightly hairy. Profemora smooth, except for apical spine. Meso- and 
metafemora with eight fairly even serrations ventrolaterally, and an apical spine. Tarsi 
of modest length. 

Abdomen slender, in S anal segment twice as long as abdominal segment IX, 
split into large lobes, broad on dorsal part, then considerably narrowed, ending in a 
club posteriorly. Lobes forming an expanded, circular arch-like structure. Subgenital 



32 O. ZOMPRO & P. D. BROCK 

plate swollen, raised in centre, then tapered to slightly rounded tip, slightly projecting 
beyond posterior margin of sternite IX. In 9 , posterior margin of X bold triangular 
incised; supra-anal plate visible, rounded at tip. Operculum long and narrow, rounded 
at tip, extending beyond end X. Cerci much longer than X, broadened and leaf-like, 
slightly pointed at tip; overlapping or incurved. 

Distribution. Australia, North Queensland. 

Species included. Parapodacanthus hasenpuschorum Brock sp. n. 

Parapodacanthus hasenpuschorum Brock sp. n. Figs 46-53, 60-62 

Material examined. HT: S; Stone Creek, Garradunga, Innisfail, N. Qld., 19.1.1996, J. 
Hasenpusch [QMBA]. 

PT: 3 o\ 10 9 ; Stone Creek, Garradunga, Innisfail, N. Qld. (2 S, 1 9 , 16.1.1997, leg. P. 
Hasenpusch; 1 o\ 2 9 9, 17.1.1997, leg. J. Hasenpusch; 1 9, 09.XI.1995, leg. P. Hasenpusch; 2 
9, 20.XII.1995, leg. J. Hasenpusch; 1 9, 19.1.1996, leg. J. Hasenpusch; 1 9, 14.1.1997, leg. S. 
Hasenpusch; 1 9, 18.1.1997, leg. J. Hasenpusch; 1 9, 20.1.1997, leg. J. Hasenpusch) [QMBA]; 
3 o* , 3 9 ; North Queensland: Stone Creek, Garradunga, Innisfail, N. Qld. (1 â • , with abdomen 
broken off, 1 9, XI.1992, leg. J. Hasenpusch; 1 Ô, 1 9, 29.X.1997, leg. J. Hasenpusch; 1 6, 1 
9 , and eggs, 25.X.1997, leg. J. Hasenpusch & P. D. Brock) [Coll. P. D. Brock]; 1 o\ 3 9 , Stone 
Creek, Garradunga, Innisfail, N. Qld. (1 o\ 21.1.1997, leg. J. Hasenpusch; 2 9 9, 21.1.1997, leg. 
S. Hasenpusch; 1 9, 20.1.1997, leg. P. Hasenpusch) [ANIC]; 1 S; Cairns, "Nov. Holl" [MHNG]. 

Adults 

Attractive leaf green and brown, medium sized insect of glossy appearance, 
with four pairs of large spine-like tubercles on mesonotum and dark pink alae. 

Description â . Head slightly longer than wide. Genae green with a narrow dark 
brown band. Eyes large, brown. Three ocelli present. Antennae long, black with scapus 
and following segments dark brown. 

Pronotum green, slightly longer than head, with central impression. Mesonotum 
more than 1.5 times as long as pronotum, with four pairs of large brown spine-like 
tubercles; the front pair are much longer, rather horn-like, black tipped, curved anteriad 
and laterad. The other spines slant outwards, the posterior pair shorter than the 
preceding pairs. Remainder of mesonotum green. Metanotum slightly longer than 
mesonotum; green. Lateral margins of meso- and metathorax with series of small 
tubercles. Metasternum with small tubercles. Tegmina long and slender, tapered to a 
rounded tip, green with broad cream margin followed by two thick central dark mauve 
lines, with a narrow white line between; then green area, which includes three cream 
stripes. Pre-anal part of alae the same colour, although margin is a narrower cream 
band, followed by two mauve lines and remainder green. Inner margin transparent / 
reddish. Alae dark pink with darker pink veins. Legs elongate, slightly hairy, fore legs 
yellowish brown, mid and hind legs, particularly femora dark reddish green; profemora 
smooth, except for apical spine. Meso- and metafemora with eight fairly evenly spaced 
serrations on ventrolateral carinae, and an apical spine. Tibiae smooth, except inner 
margin of metatibiae slightly serrate. Tarsi of modest length. Apices of leg segments 
darker. 

Abdomen slender, abdominal segment VII reduced in size compared with pre- 
vious segments, VIII shorter, broadened anteriorly; IX shorter again and narrow, slight- 
ly expanded at base. Anal segment twice as long as IX, split into two large lobes, broad 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 33 

on upper part, then considerably narrowed, ending in a club at tip. The lobes form an 
expanded, circular arch-like structuure. Subgenital plate swollen, raised in centre, then 
tapered to slightly rounded tip, just exceeding end of IX. Cerci much longer than X, 
broadened and leaf-shaped, slightly pointed at tip; overlapping. 

Measurements (mm), HT, â: Body: 83.0; head: 3.5; antennae: 39.0; pronotum: 
3.8; mesonotum: 6.5; metanotum: 9.0; tegmina: 16.0; alae: 58.0; median segment: 6.0; 
profemora: 20.0; protibiae: 17.0; mesofemora: 17.0; mesotibiae: 13.0; metafemora: 
20.0; metatibiae: 18.0; protarsus: 11.0; mesotarsus: 7.0; metatarsus: 9.0; cerei: 6.0. 

Measurements (mm), PT, â: Body: 83.0-88.0; head: 3.5-3.7; antennae: 39.0- 
40.0 (tips often broken off); pronotum: 3.8-4.0; mesonotum: 6.5-7.0; metanotum: 9.5; 
tegmina: 16.0-17.0; alae: 58.0-60.0; median segment: 5.5; profemora: 20.0-21.0; 
protibiae: 17.0-19.0; mesofemora: 16.0-17.0; mesotibiae: 12.0-13.0; metafemora: 
20.0-21.0; metatibiae: 17.5-18.0; protarsus: 11.0; mesotarsus: 7.0; metatarsus: 9.0- 
10.0; cerei: 6.0-6.5. 

Description 9 . Broader than male, but description similar except insect is 
nearly all leaf-green; tegmina and pre-anal part of alae are green, often with broad 
cream band of varying magnitude. The mesonotum is brown on the spine-shaped 
tubercles as in male, and there are brownish bands on the head. The legs are greenish, 
apices of femora and tibiae with slight brown tinge. Interolateral margin of metatibiae 
with 14 small serrations. 

Metathorax with series of ten small tubercles laterally; much smaller series on 
metathorax. Metasternum with white margins and blotch on posterior part; tubercles 
also present. 

Abdomen green, with middle part broad reddish brown. End of anal segment 
boldly triangularly incised, supra-anal plate visible, rounded at tip, which extends 
beyond end of anal segment. Cerci long, as long as IX and X combined; leaf-shaped, 
broader at base, incurved and slightly pointed at tip. 

Measurements (mm), PT, ?: Body: 100.0-117.0; head: 5.0-6.0; antennae: 42.0- 
50 (tips often broken off); pronotum: 4.7-5.5; mesonotum: 7.0-8.0; metanotum: 15.0- 
16.0; tegmina: 25.0-31.0; alae: 75.0-86.0; median segment: 7.0; profemora: 20.5-24.0; 
protibiae: 19.0-22.0; mesofemora: 16.0-22.0; mesotibiae: 12.0-14.0; metafemora: 
21.0-27.0; metatibiae: 17.0-21.0; protarsus: 12.0-14.0; mesotarsus: 8.0-9.5; metatar- 
sus: 9.0-10.5; cerei: 5.0-6.5. 

Nymphs 

The first pair of spines on the mesonotum are black, followed by a central black 
longitudinal band, not reaching end of mesonotum. In the adult, this colouration 
changes and all spines are brown. 

Egg 

Small, dark brown (orange when freshly laid), capsule oval, slightly hairy with 
small lighter brown knog-like capitulum on a stalk. Capsule with numerous small 
cristae. Micropylar plate small, cordiforme, its center dark brown to black, with lighter 
brown median line reaching posterior pole. 

Measurements (mm): Capsule length: 2.7; width: 1.5; height: 1.5. 



34 o. zompro & p. d. brock 

Name 

Named Parapodacanthus in view of its apparant affinity with Podacanthus 
Gray. The species is named after the Hasenpusch family, who collected the majority of 
the type-series and have a passion for insects; few individuals can boast of such a beau- 
tiful range of insects in and around the vicinity of their rainforest home. 

Distribution 

Endemic for North Queensland, Australia. This species was only found at the 
lower slopes of Innisfail and Garradunga, Cairns, Kuranda, at Black Mt. Road and Mt. 
Lewis. 

Comments 

This rainforest species was found feeding on Melicope elleryanas (Rutaceae) 
and Acronychia acidula (Rutaceae) in the wild, whereas the closely related 
Podacanthus species feed on Eucalyptus spp. (Myrtaceae). Parapodacanthus hasen- 
puschorum sp. n. refuse to feed on various Eucalyptus spp. or Tetradium daniellii 
(Rutaceae). The male of an unidentified species in the collection of Mr T Hiller (Mt. 
Glorious, South-East Queensland) was found on Tetradium sp. Both sexes can fly well. 
Adults and nymphs emit an unpleasant smelling odour from their prothoracic glands. 
Newly hatched nymphs are brick red with green leg joints. 

ACKNOWLEDGEMENTS 

The authors wish to thank Prof. Dr V. Mahnert, Dr P. Schwendinger and Dr C. 
Lienhard [MHNG] for their support, and especially Ms A. Teschke (Berlin, Germany), 
who kindly arranged and digitalized the figures. Prof. Dr Victor P. Gapud and Orlando 
L. Eusebio (University of the Philippines, Los Banos, Philippines) supported naming 
the three species described from the Philippines. Dr P. E. Bragg (Nottinghamshire, 
England), Prof. Dr J. Adis and Prof. Dr W. Junk (both Max-Planck-Institute for 
Limnology, Ploen, Germany) provided their support in every respect. 

J. Hasenpusch (Garradunga, North Queensland, Australia) kindly provided 
notes and foodplant details of Parapodacanthus hasenpuschorum. 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 



35 




Figs 1-15. Terminal abdominal segments, lateral aspect. To various scales. Xylodus adumbratus 
Saussure, 1859. 1. HT, 9. Gratidia carinulata (Saussure, 1868). 2. ST, 9. 3. Dorsal aspect. 4. 
Ventral aspect. Paramenexenus ceylonicus (Saussure, 1868) 5. HT, 9. Phantasca (?) cubensis 
(Saussure, 1868). 6. HT, S . Paroxyartes dohertyi Carl, 1913. 7. HT, S . Brachyrhamphus fecun- 
dus Carl, 1915. 8. ST, 8. 9. ST, Ç.Clonistria guatemalensis Redtenbacher, 1908. 10. LT, c\ 11. 
PLT, 3 . Micrarchus hystriculeus (Westwood, 1859). 12. HT of Micrarchus parvulus Carl, 1913. 
ST, 3. 13. ST, 8. Dorsal aspect. Paranisomorpha insignis Redtenbacher, 1906. 14. ST, 5. 
Parastheneboea laetior (Brunner v. Wattenwyl, 1907). 15. HT of Echinoclonia bomeensis Carl, 
1913, 9. 



36 



O. ZOMPRO & P. D. BROCK 




Figs 16-29. Terminal abdominal segments, lateral aspect. To various scales. Asceles malaccae 
(Saussure, 1868). 16. HT, 9. Alloeophasma poeyi (Saussure, 1868). 17. HT, 9. Gratidia pul- 
chrepicta Carl, 1913. 18. HT, S . 19. Head, dorsal aspect. Cnipsus rhachis (Saussure, 1868). 20. 
ST, 3. 21. ST, 9. Labidiophasma rouxi Carl, 1915. 22. ST, 9. Marmessoidea rubescens 
(Saussure, 1868). 23. HT, 9 . Paraleptinia schulthessi (Carl, 1913). 24. HT, 9 . 25. Dorsal aspect. 
26. Ventral aspect. 27. Head, dorsal aspect. Andropromachus scutatus Carl, 1913. 28. HT, 9. 
Pseudostheneboea segregata Carl, 1913). 29. HT, â . 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 



37 




Figs 30-36. Terminal abdominal segments, lateral aspect. To various scales. Centwphasma 
spinosum (Saussure, 1868). 30. HT, S . Pachymorpha squalida (Gray, 1833). 31. (5.32. 9. Eggs. 
Scale: 1 mm. Alienobostra godmani (Redtenbacher, 1908), egg. 33. Lateral aspect. 34. Dorsal 
aspect. Stheneboea palawanica (Carl, 1913), egg. 35. Lateral, aspect. 36. Dorsal aspect. 




Figs 37-45. Lonchodes putingmantsa Zompro sp. n. Right mesofemur, apex, lateral aspect. 37. 
HT, 6. 38. PT, 9. Terminal abdominal segments, lateral aspect. 39. HT, â. 40. PT, 9. Scale: 
5 mm. Lonchodes magayon Zompro sp. n., HT, 6 . 41. Right mesofemur, apex, lateral aspect. 42. 
Terminal abdominal segments, lateral aspect. Scale: 5 mm. Lonchodes dalawangsungay Zompro 
sp. n., HT, 9. 43. Head, lateral aspect. Scale: 1 mm. 44. Left mesofemur, apex, lateral aspect. 
45. Terminal abdominal segments, lateral aspect. Scale: 5 mm. 



38 



O. ZOMPRO & P. D. BROCK 






Figs 46-53 
Parapodacanthus hasenpuschorum Brock gen. n., sp. n. Terminal abdominal segments. 46. S , 
lateral aspect. 47. 6 , dorsal aspect. 48. 9 , lateral aspect. 49. 9 , dorsal aspect. Scale: 5 mm. Egg. 
50. Lateral aspect. 51. Dorsal aspect. 52. 6 . 53. 9. Scale: 1 mm. 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 



39 






z>i>Gc~. *>'" 



56 




Figs 54-57 
Lonchodes putingmantsa Zompro sp. n. 54. HT, S. 55. FT, 9. Lonchodes magayon Zompro sp. 
n. 56. HT, <? . Lonchodes dalawangsungay Zompro sp. n. 57. HT, $ . 



40 



O. ZOMPRO & P. D. BROCK 





^| 



58 




59 



Figs 58-59 
Alloeophasma poeyi (Saussure, 1868). 58. LT, 9. 59. S , ST of Phasma cubense Saussure, li 



PHASMATODEA TYPES IN THE GENEVA MUSEUM 



41 




Figs 60-62 
Parapodacanthus hasenpuschorum Brock gen. n., sp. n. 60. S . 61. 9. 62. Mating 6 9 . 



42 O. ZOMPRO & P. D. BROCK 



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Revue suisse de Zoologie 110(1): 45-89; mars 2003 



Ground spiders (Gnaphosidae; Araneae) from Crete and adjacent 
areas of Greece. Taxonomy and distribution. Ill: Zelotes and allied 
genera 

Maria CHATZAKI 12 , Konrad THALER 3 & Moysis MYLONAS 12 

1 Dept of Biology, University of Crete, 71 100, Irakleio, Crete, Greece. 

2 Natural History Museum of the University of Crete, Knossou Av., 
71409 Irakleio, Crete, Greece, PO box 2208. 

3 Institut für Zoologie und Limnologie der Universität, Technikerstraße 25, 
A-6020 Innsbruck, Austria. 



Ground spiders (Gnaphosidae; Araneae) from Crete and adjacent 
areas of Greece. Taxonomy and distribution. Ill: Zelotes and allied 
genera. - The Gnaphosidae zelotine genera Camillina, Drassyllus, 
Tr achy zelotes, Setaphis and Zelotes from Crete and the adjacent islands of 
Gavdos, Gavdopoula, Dia, Kos, Karpathos and Antikythira are investigated. 
Taxonomic details are presented for 20 species. Three species (Drassyllus 
pumiloides sp.n., Zelotes daidalus sp.n., and Z. minous sp. n.) are new to 
science. A new combination {Camillina metellus [from Zelotes}) and two 
new synonymies (Trachy zelotes stubbsi = T. adriaticus, Zelotes bucharensis 
= Z. scrutatus) are proposed. Two species (Z. aerosus and Z. solstitialis) are 
recorded for the first time in Europe, whilst four species are new records for 
Greece (Drassyllus praeficus, Trachyzelotes adriaticus, Setaphis carmeli 
and Z. subterraneus). Another two species are new records for Crete (Z. la- 
bi lis and Z. ni li cola). 

Key-words: Araneae - Gnaphosidae - Zelotes - Greece - Crete - South 
Aegean - taxonomy - distribution. 

INTRODUCTION 

Zelotes Gistel, 1848 is a highly speciose genus with worldwide distribution, 
included in the zelotine group of the Gnaphosidae. These genera possess a charac- 
teristic preening comb on metatarsi III and IV. The status of the genus Zelotes is pro- 
blematic due to the vague limits of the taxonomical characters defining it, and to the 
great variety of genital characters among its species (Platnick & Shadab, 1983). Since 
there is no contemporary, comparative revision of Zelotes in the Old World, the study 
of this genus is very difficult, especially in the Mediterranean, where its diversity 
becomes very high. 

Up to now, 357 Zelotes species have been identified in the whole world, of 
which, about 200 occur in the Old World (see Platnick, 2001). On a regional level, 



Manuscript accepted 31.05.2002 



46 M. CHATZAKI ET AL. 

species catalogues for Central Europe list 30 zelotine species (Grimm, 1985), for 
Czechoslovakia 19 (Buchar, 1992), for Romania 24 (Weiss & Petrisor, 1999), for 
Bulgaria 35 (Deltshev & Blagoev, 2001), and for Italy 57 (Pesarini, 1994). The recent 
revision by Levy (1998) revealed 36 species in Israel, making this group the most 
species-rich among gnaphosids and '...probably the largest of all indigenous spider 
genera'. These data are indicative of the high diversity of the genus in the Medi- 
terranean region. It has also been observed in American Zelotes, that their species 
numbers are very high in California. 

In Greece, as most records have been based on occasional hand collections, the 
number of species of these fast moving ground spiders has been greatly under- 
estimated. Hadjissarantos (1940) recorded 10 zelotines out of 32 gnaphosids from 
Attiki. Research into the literature on Crete reveals 1 1 zelotines out of 34 Gnaphosidae. 

In this study we present the results of an extensive survey along the island of 
Crete and the adjacent islands Gavdos, Gavdopoula and Dia, as well as additional 
material from the Aegean islands Kos, Karpathos and Antikythira and from mainland 
Peloponnisos. At all localities, pitfall traps have been used. This is the most suitable 
method for collecting these spiders, which are mostly nocturnal and very active on the 
ground (Levy, 1998). In total, 20 species have been identified. These belong to the 
genera Camillina [1 sp.], Drassyllus [2 spp.], Trachyzelotes [4 spp.], Setaphis [1 sp.] 
and Zelotes [12 spp.]. These data correspond to 9 of the previously recorded species 
(excluding Z. clivicolus and Z. oblongus, which probably are misidentifications of 
allied species). Consequently our results double the number of zelotines recorded for 
Crete. There are a few more species in our collection, which have not yet been satis- 
factorily identified, so we expect the actual number of zelotine species to be higher. 

MATERIAL AND METHODS 

Sampling strategy, exact localities and habitat type of each site are given in 
Chatzaki et al. (2002a). In total, 59 sites were selected along the length of the island of 
Crete, 1 1 on the island group Gavdos - Gavdopoula and 3 on the island Dia (Fig. 1). 
Sampling sites cover Crete from north to south, west to east and along the altitudinal 
gradients of the three mountain massifs of the island, namely, Lefka Ori Mts., 
Psiloreitis Mt. and Lasithiotika Ori Mts. Most of the habitats selected on Crete are 
phrygana (plant communities which include dwarf, aromatic, thorny shrubs) and 
maquis. Few of the sites are pine forests or are situated close to permanent or tempo- 
rary water reservoirs. Some records of material collected from other areas of mainland 
Greece and the Aegean islands (Fig. 2) have been added, but are not shown in the dis- 
tribution maps given for the species. 

Spiders were collected using pitfall traps (12 cm height, 9.5 cm in diameter). 
The killing preservative was ethylene glycole. At each site, 15-20 traps were set and 
changed in two-months intervals. In most cases only material from the period of high 
activity of Gnaphosidae, e.g., late spring to early autumn (Chatzaki et al., 1998; 
Chatzaki, 1998), has been analyzed and is presented here. 

The collection of material was financially supported by scientific projects of the 
EEC concerning biodiversity, i.e. TERRA, INTEREG II (ARCHIMED), or by the 



GROUND SPIDERS OF CRETE. III. 



47 




Crete 



^T^ 



Dia 

^52-54 



104&8 5»\Rethymno ^ 2 < .„Irakleio ^ " 4g 



Gavdopoula 

♦23 

n 

Gavdos 
13-22 



*Làoj£ì 



V 59 63^6|^ Q i- 
^7*58 62*67>? 9 



66**59 63 i_ Ä ^asithi 



7^2 



ì«l: 



♦47 <oW 8 S 6 *¥X- 73*/ 

4.Q ^ u " -—64 — -^*^-^ - — vJ^ 

'46 <e?~^~- 



66 



Fig. 1 
Map of sampling sites on Crete and the surrounding islands Gavdos, Gavdopoula and Dia. 







, Antikythira 






Crete 






«Ü,. 




Karpathos <K 



Gavdos 



Fig. 2 
Map of sampling sites on Attiki, Peloponnisos and Aegean islands. 



Ministry of Environment, Physical Planning and Public Works (Project: "Gavdos, an 
island on the edge of Crete"), undertaken by the Natural History Museum of Crete 
(NHMC) and by the Biological Department of the University of Crete. 

Identifications were carried out at the Natural History Museum of Crete 
(NHMC) and at the Zoological Institute of Innsbruck, Austria. Most of the material 
presented here is used for the Ph.D. thesis of the first author and, if not stated other- 
wise, it is deposited at the NHMC. Material from the collection of Dr Hadjissarantos, 



48 M. CHATZAKI ET AL. 

deposited at the Zoological Museum of the Biological Department of Athens (ZMUA), 
and material from the collection of Roewer, deposited at the Senckenberg Museum of 
Natural History, Frankfurt am Main (SMF), as well as from the collection of the second 
author (CTh) and of Dr J. Levy (Hebrew University, Jerusalem, HUJ), has also been 
examined. Voucher specimens have been deposited at the Natural History Museum of 
Geneva (MHNG). 

The following abbreviations are used in the text: TL: total length, PL: prosoma 
length, PW: prosoma width, OL: opisthosoma length, Cy: cymbium, Ta: tarsus, Me: 
metatarsus, Ti: tibia, Pa: patella, Fe: femur, d: dorsal, v: ventral, AME: anterior median 
eyes, ALE: anterior lateral eyes, PME: posterior median eyes, PLE: posterior lateral 
eyes. All measurements are given in mm. All drawings presented here are by the first 
author. 

RESULTS 

Camillina metellus (Roewer, 1928) comb. n. Figs 3-9, 115 

Zelotes metellus Roewer, 1928 (p. 1 10, Fig. 15), CRETE: Rethymno, Aptera (type locality). 

Etymology. As Roewer named this species after the Roman conqueror 'Metellus 
Creticus', the specific name is a noun in apposition, therefore invariable. 

Material. C. metellus : CRETE: Georgioupoli, Phragmites stand at spring rivulet close 
to sandy seashore, E of the village (1 9, 19/5/2001. NHMC leg. & don. Kronestedt). 

C. metellus (?): CRETE: CHANIA: Site l(a6<?c?;b3ö*oM9); Site 2(a8ö*c?399; 
b 6 â S 1 9 ); Site 4 (b 1 8) (leg. Lymberakis); Site 23 (c 2 ? 9 ) (leg. Paragamian); RETHYM- 
NO: Site 27 (a 16 S 6 1 9) (leg. Chatzaki); Site 28 (a 1 o*; b 1 S 1 9); Site 29 (a 2 S â); Site 
39 (al S); Site 41(a299;b4ó*cM9) (leg. Nikolakakis); IRAKLEIO: Site 46 (a 2 9 9 ; b 
1 9) (leg. Papadimitrakis); Site 52 (b 4 S S); Site 53 (b 1 6 (leg. Nikolakakis); LASITHI: Site 
55 (a 1 6; b 2 o*o* [CTh]); Site 73 (a 2 dd) (leg. Chatzaki); Site 68 (a 1 <J) (leg. 
Papadimitrakis); Site 63 (b 5 â â; b 1 â 1 9 [MHNG]) (leg. Stathi). 

C. metellus (?): KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt 
field, (1 ó\ pitfall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

Comparative material examined: Zelotes metellus: CRETE: 1 9 holotype, SMF: CR 
612/62. 

Taxonomy. Zelotes metellus was described by Roewer (1928) on the basis of a 
single female collected on Crete. Examination of the type verified the identification of 
a specimen, collected by Dr. Kronestedt in Georgioupoli (Fig. 8-9). In our collection 
there are females which are very similar to the type, but not identical (see Fig. 6-7). 
These females are matched with male specimens (Figs 3-5), both of them bearing the 
diagnostic characters of the genus Camillina: embolar base (EB) recessed, situated 
medially, terminal apophysis (TA) bifid, epigyne with median plate (MP) (sensu 
Platnick & Shadab, 1982a). On the basis of these characters we conclude that this 
species belongs to Camillina. Considering the similarity of the females of this species 
with the 'true' Z. metellus, we presume that the latter also belongs to the same genus, 
therefore the transfer is proposed. In the absence of male specimens, matching the 



Figs 3-9 
Camillina metellus (8-9), C. metellus (?) (3-5, 6-7). Palp of S , ventral view (3), dorsal view (4), 
retrolateral view (5), epigyne (6, 8). vulva (7, 9). EB: embolar base, E: embolus, 1: lamella, TA: 
terminal apophysis. MP: median plate. Scale lines 0.1mm. 



GROUND SPIDERS OF CRETE. III. 



49 




50 M. CHATZAKI ET AL. 

single 'true' C. metellus female, we dare not suggest that a second C amili ina occurs on 
Crete. Therefore we reluctantly place all these specimens in one species, C. metellus. 

Camillina was established by Tullgren (1910, sub Camilla) for a species from 
Tanzania. There are about 20 species known from the Old World (see Platnick & 
Murphy, 1987), mainly from Africa, and another 40 species from the New World (see 
Platnick & Shadab, 1982a). C. europaea Dalmas, 1922 is the only species of the genus 
recorded in Europe (southern Italy, Di Franco, 1997). This is the first record of the 
genus in Greece. 

Measurements 6 (9), n = 8 (7): TL: 2.9-4.5 (3.5-4.5), PL: 1.4-1.9 (1.5-1.7), 
PW: 1.1-1.6(1.1-1.5), OL: 1.5-2.4 (1.5-2.4), PL/PW: 1.14-1.41 (1.13-1.45). 

Ô 2 : Small spiders with colour varying from yellow to light brown or grey. Ta 
and Me of contrasting light colour. Habitus as in Zelotes. AME very close to each oth- 
er and to ALE, forming a continous row, straight or slightly recurved. PME oval, closer 
to PLE than to each other, all set in a straight or slightly procurved line. Opisthosoma 
oval, anterior spinnerets cylindrical and well separated. 

Legs: Ta and Me I-II with scopula hairs. Ta III-IV with stripes of dense, fine 
spines. Preening combs present on Me III and IV. Spination: Fe: I-II d 2-3 ; HI-IV d 
6-7. Pa: I-II, IV - ; III r 1. Ti: I-II - ; III-IV spinose. Me: I-II - ; III-IV spinose. 

6 Pedipalp (Figs 3-5): Tibial apophysis long and tapering. Sperm duct follow- 
ing the tegulum from retro- to prolateral side, then turning inwards. Embolus (E) 
strongly developed, rising from the prolateral, dorsal part of the tegulum, transverse, 
distally broadened, its surface rough. Terminal apophysis (TA) bifid, with tips crossed 
(Fig. 4). A membranous lamella (1), originating from the dorsal part of the tegulum, 
also present. Retinaculum large, with anterior edge pointed. 

9 Epigyne (Fig. 6): Anterior margin divided, posteriorly prolonged. Median 
plate (MP) oval, longer than wide. Introductory orifices situated laterally and near the 
posterior rim of the median plate. 

Vulva (Fig. 7): Introductory ducts short and sclerotized. Glandular heads situat- 
ed anteriorly, often with a sclerotized ventral rim. Spermathecae globular and laterally 
connected to the introductory ducts. Fertilisation ducts situated at the anterior border 
of spermathecae. 

Ecology. C. metellus has been mainly collected on phrygana near the Cretan 
coasts and inland, only on the southwestern slopes of Mt. Psiloreitis. The species 
reaches altitudes of about 1400m (Fig. 1 15). 

Distribution. Crete, Karpathos (Greek endemic?). 

Drassyllus praeficus (L. Koch, 1866) Figs 17-19, 116 

Identification: Grimm (1985, p. 267, Figs 317, 319-320). 

Material. CRETE: CHANIA: Site 4 (a 3 SS); Site 5(a3o*o*3 99;bl9) (all leg. 
Lymberakis): Site 1 1 (b 2 S S) (leg. Stathi): RETHYMNO: Site 24 (a 1 2) (leg. Stathi); Site 28 
(a 7 2 9); Site 29 (a 2 â S 6 9 9); Site 29 (b 1 9); Site 32 (a 22 S S 49 9 9); Site 39 (a 5 S S 
7 9 9); Site 40 (a 1 1 S â ; b 14 9 9 ) Site 41 (a 1 9 ) (all leg. Nikolakakis); Site 30 (a 1 9 ); Site 
31 (a 1 2) (all leg. Trichas); Site 34 (a 10 S3 24 9 9; b 1 9; e 27 SS 3 9 9) (all leg. 
Chatzaki); IRAKLEIO: Site 42 (e 5 S S 1 9) (leg. Chatzaki); Site 43 (a 51 S S 84 9 9; b 18 
9 9); Site 44 (a 9 SS 6 9 9); Site 47 (a 24 SS 11 9 9; b 10 9 9; e 6 S S 2 9 9) (all leg. 
Nikolakakis); Site 46 (al 9); Site 49 (a 9 S S 23 9 9; b 2 9 9); Site 51 (a 24 9 9) (all leg. 



GROUND SPIDERS OF CRETE. III. 5 1 

Papadimitrakis); Site 48 (a 3 6 3 3 9 9 ) (leg. Trichas); LASITHI: Site 55 (b 1 9 ; e 2 3 3); Site 
56(a8c?<?999;b299;dll 3 3) (all leg. Chatzaki); Site 58 (a 1 6 22 9 9 ); Site 62 (a 1 
9) (all leg. Papadimitrakis); Site 59 (c 1 3 2 9 9 [MHNG]); Site 70 (a 1 3) (all leg. Trichas). 

Ecology. D. praeficus is common on Crete, occurring from the coastal plains of 
the island up to 1650m. However, it is less common in the district of Chania and absent 
from the Lefka Ori Mts., whereas it is abundant on the other two mountains of Crete 
(Fig. 1 16). The peak of activity in both males and females is from spring to autumn. 

Distribution. Europe to Central Asia (first record for Greece). 

Drassyllus pumiloides Chatzaki sp. n. Figs 10, 12, 14-15, 116 

Etymology: The name of this species indicates its close relationship to D. pumilus (C. L. 
Koch, 1839). 

Material. Type material: Site 63a (Istro) (1 S holotype [NHMC]; Site 58a (2 9 9 
paratypes [NHMC]; 1 9 paratype [MHNG]). 

CRETE: CHANIA: Site 1 (a 1 3); Site 2 (a 3 9 9); Site 6 (d 1 3); Site 7 (c 1 3) (all 
leg. Lymberakis); RETHYMNO: Site 28 (b 1 9); Site 29 (a 2 o* 3 2 9 9 ); Site 39 (a 1 9 ; b 1 
3 [MHNG]); Site 40 (b 3 3 3 2 9 9 ) (all leg. Nikolakakis); Site 34 (a 1 3 1 9 ) (leg. Chatzaki); 
IPvAKLEIO: Site 43 (a3 33 1 9; b 1 9); Site 45 (a 1 <J); Site 47 (e 1 3 1 9); Site 50 (b 1 9) 
(all leg. Nikolakakis); Site 49 (a 3 3 3 ; a 1 9 [CThl; b 1 9 ) (all leg. Papadimitrakis); LASITHI: 
Site 55 (a 1 3) (leg. Chatzaki); Site 63 (a 1 3) (leg. Stathi); Site 58 (a 3 9 9) (leg. 
Papadimitrakis). 

Comparative material examined: D. pumilus (C. L. Koch, 1839): Austria, N. Tirol, Ötz- 
tal (1 3, 5/5/1962, CTh); Italy, Trentino, V. Ledro, Bezzecca (1 9, 27/5/1963, CTh). 

D.jubatopalpis Levy, 1998: Israel, Golan, Odern forest, 950m (lo* 5/1997, leg. Sharon, 
HUJ 15222). Israel, same locality (19, 5/1996, leg. Sharon, Col. Levy, 15223). 

Taxonomy. Measurements 6 (9), n = 6 (6): TL: 2.6-3.2 (2.6-3.4), CL: 1-1.2 
(1.1-1.3), PW: 1.4-1.8 (0.8-0.9), OL: 1.4-1.8 (0.8-0.9), PL/PW: 1.2-1.37 (1.2-1.62). 

6 9 : Small spiders of light to dark brown colour. Prosoma longer than wide. 
Eyes round, except for oval PME. AME closer to ALE than to each other. Posterior row 
of eyes recurved, anterior row procurved or straight. 

Legs: Co, Tr, Me and Ta yellow, the rest brown. Ta and Me I-II with stripes of 
scopula hairs, Ta and Me III-IV with dense spiny hairs. Me III-IV with apical preening 
comb. Spination: Fe: I-IV d 1-2. Pa: I-IV - . Ti: I-II - ; III-IV spinose. Me: I-II - ; III d 
4;rVv2p 1. 

S Pedipalp (Figs 10, 12): Tibial apophysis pointed, sharply bent dorsally. 
Embolus filiform, retrolateral embolar projection (EP) small. Terminal apophysis (TA) 
(sensu Platnick & Shadab, 1982b) prominent, rounded, almost covering the base of the 
embolus. Retinaculum retrolateral and very broad, with a very small projection pro- 
ximal to it. 

Epigyne (Fig. 14): Anterior margin continuous and almost touching the intro- 
ductory ducts. Lateral margins forming loops alongside the introductory ducts. 

Vulva (Fig. 15): Introductory ducts broad, curved, with small glandular heads 
on the anterior arch. Spermathecae two-lobed, with a large, globular chamber, and a 
smaller median chamber leading to the fertilisation ducts. 

Comments. This species is very close to D. pumilus (C. L. Koch, 1839), as can 
be seen from Figs 11, 13 and 16 (see also Grimm 1985, p. 271, Figs 313, 323-324). 
Apart from different size, D. pumilus being much larger (3 9 3. 5-4. 8mm, according to 



52 



M. CHATZAKI ET AL. 




FlGS 10-19 
Drassyllus pumiloides sp. n. (10, 12, 14-15), Drassyllus pumilus (11, 13, 16), Drassyllus prae- 
ficus (\7-\9). Palp of ó\ ventral view (10-11), retrolateral view (12-13, 17), epigyne (14, 16, 18), 
vulva (15, 19). EP: embolar projection. Scale lines 0.1mm. 



GROUND SPIDERS OF CRETE. III. 53 

Grimm), the new species also differs in the following characters: terminal apophysis 
less prominent; embolus longer; embolar projection broader; anterior margin of 
epigyne much closer to the introductory ducts; lateral margins almost parallel and 
straight, then turning sharply; introductory ducts more slender, with longer glandular 
heads; spermathecae pear-like. D. villicoides (Giltay, 1932) from Peloponnisos clearly 
differs in size (TL 7.5mm, according to Giltay, 1932). Females of D. jubatopalpis 
Levy, 1998 are also very similar, but different (in our specimens introductory ducts and 
median plate shorter, no ducts in the spermathecae, see Levy, 1998, p. 154, Figs 
136-137), the males easily distinguishable by the shape of the tibial apophysis and by 
details of the palpal organ. 

Ecology. D. pumiloides occurs almost all over Crete, being absent only from the 
easternmost areas, apparently avoiding the driest habitats (Fig. 116). It reaches alti- 
tudes up to 1650m. 

Distribution. Crete (endemic?). 

Trachyzelotes barbatus (L. Koch, 1866) Figs 20-21, 26-27, 1 17 

Identification: Platnick & Murphy (1984, p. 15 Figs 27-30). 

Material. CRETE: CHANIA: Site l(al5c?c?699;b3Jc?3 99;cl 9); Site 2 (a 
29<?<?ll99;b3<?c?4 9 9); Site 4 (a 6 6 6 1 9;b 1 6 5 9 9);Site6(dl 9;e2c?<?;fl 
9) (all leg. Lymberakis); RETHYMNO: Site 26 (a 1 9) (leg. Lymberakis). 

ANTIKYTHIRA: Potamos, 700m W: sparse phrygana on sandy soil close to the village, 
(7 6 6 4 9 9 , pitfall traps 27/3/2001 - 5/8/2001, leg. Chatzaki). 

Ecology. T. barbatus is not very common on Crete (Fig. 1 17). It has been found 
on sites at the western periphery of Crete and on Antikythira, always in phrygana. 

Distribution. Mediterranean to Central Asia, USA. In our study area this species 
has been recorded only from western Crete and from Antikythira, suggesting an 
immigration from the Balkan peninsula. 

Trachyzelotes malkini Platnick, 1984 Figs 22-23, 28, 117 

Identification: Platnick & Murphy (1984, p. 22, Figs 51-54). 

Material. CRETE: CHANIA: Site 5(a7c?c?4 99); Site 6 (h 1 6; i 1 6) (all leg. 
Lymberakis); Site 13 (c\2 66399 ;d366399 [MHNG]); Site 14 (c 3 0*0* 1 9; d 1 9); 
Site 15 (c 3 6 6; d 2 9 9); Site 16 (c 2 66); Site 20 (a 1 6); Site 21 (d 1 9) (all leg. 
Paragamian); RETHYMNO: Site 24 (a 1 6 1 9) (leg. Stathi); Site 25 (a 2 6 6); Site 26 (a 4 
66 3 9 9;b2 9 9 ) (all leg. Lymberakis); Site 27 (a 4 66 2 99;f5 6 6); Site 55 (b 2 66) 
(all leg. Chatzaki); Site 28 (a 4 6 6 3 9 9 ; b 1 6); Site 29 (a 4 6 6 1 9 ); Site 39 (a 8 6 6 1 9 ; 
b 1 9); Site 40 (a 1 6 1 9; b 27 6 6 9 9 9; c 1 9); Site 41 (a 5 6 6 2 9 9) (all leg. 
Nikolakakis); IRAKLEIO: Site 42 (e 1 6) (leg. Chatzaki); Site 43 (a 1 6 1 9; b 1 9); Site 44 
(a 3 6 6 2 9 9;b 1 9); Site 45 (a 3 6 6 3 9 9; b 2 9 9); Site 47 (a I 6;b2 6 6 5 9 9; e I 
6); Site 50 (b 7 6 6 1 9; c 1 9) (all leg. Nikolakakis); Site 46 (a 3 <î c? 7 9 9; b 2 <? c? 1 9); 
Site 49 (a 22 6 6 7 9 9; b 8 9 9); Site 51 (a 2 9 9) (all leg. Papadimitrakis); LASITHI: Site 
55 (a 6 6 6); Site 56(a3ÓÓ*299;b2c?ó*l9); Site 73 (e 4 6 6) (all leg. Chatzaki); Site 
58 (a 8 6 6 1 9 ); Site 64 (d 2 6 6 1 9 ); Site 65 (a 2 6 6 3 9 9 ) (all leg. Papadimitrakis); Site 
63 (a 26 6 6 2 9 9;b6 99); Site 71 (a 2 6 6 ) (all leg. Stathi); Site 59 (a 4 99;c 10 66); 
Site 61 (b 1 6) (all leg. Trichas). 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (1 6 2 9 9 , 
pitfall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

Ecology. This is the commonest Trachyzelotes species on Crete. It is 
widespread on the whole island, as well as on Gavdos and on Karpathos. It occurs in 



54 M. CHATZAKI ET AL. 

phrygana, from the periphery of Crete into the mainland and up to 1450m. The peak of 
activity of males is in late spring to early summer and that of females in spring and 
summer. 

Distribution. Russia, Turkey, GREECE: Crete: Chania: Kalathas, Akrotiri; 
Lasithi: Agios Nikolaos; Mallia (Platnick & Murphy, 1984: 23). 

Trachyzelotes lyonneti (Audouin, 1826) Figs 24-25, 29, 117 

Identification: Platnick & Murphy (1984, p. 6, Figs 7-10), Levy (1998, p. 105, Figs 
19-22). 

Material. CRETE: CHANIA: Site 1 (a 1 <J); Site 4 (a 1 S; b 1 S) (all leg. Lymberakis); 
RETHYMNO: Site 27 (a 1 6) (leg. Chatzaki); Site 41 (a 1 9) (leg. Nikolakakis); IRAKLEIO: 
Site 47(alo*;b5c?o*3 99;e299); Site 50 (b 23 S S 10 9 9) (leg. Nikolakakis); Site 50 
(c 1 S 4 9 9) (leg. Papadimitrakis); Site 48 (a 11 SS 4 9 9) (leg. Trichas); LASITHI: Site 55 
(ale?) (leg. Chatzaki); Site 60 (a 2 SS 1 9) (leg. Stathi); Site 64 (c 3 S S) (leg. Nikolakakis); 
Site 64 (d 12 S S 10 9 9; e 1 S) (all leg. Papadimitrakis); Site 61 (b 1 S); Site 66 (a 2 SS) 
(all leg. Trichas). 

Ecology. This species occurs in phrygana of the lowlands of Crete, below 600m 
(Fig. 117). It is absent from western Crete, which might be due to interaction with 
T. adriaticus, as will be pointed out later. Adults are found from spring to autumn. 

Distribution. Mediterranean to Central Asia, USA, Brazil, Peru, GREECE: 
Crete: Irakleio, Gortyna (Platnick & Murphy, 1984: 23). T. lyonneti has a Medi- 
terranean centre of distribution and was imported to USA, as commented by Platnick 
& Murphy (1984). 

Trachyzelotes adriaticus (Caporiacco, 1953) Figs 30-31, 32-39, 117 

T. stubbsi Platnick & Murphy, 1984 (p. 9, Figs 15-16), Cyprus, Cape Kiti, near Larnaka airport 

(type locality). Syn. n. 
T. stubbsi: Levy (1998, p. 106, Figs 23-24), Israel. 

Identification: Platnick & Murphy (1984, p. 9, Figs 1 1-14), Levy (1998, sub T. stubbsi). 

Material. CRETE: CHANIA: Site 1 (a 1 S); Site 2 (a 15 S S 5 9 9); Site 5 (a 2 S S) 
(all leg. Lymberakis); Site 21 (b 1 9); Site 23(a299;c8o*c?3 99) (all leg. Paragamian); 
RETHYMNO: Site 34 (a 3 9 9) (leg. Chatzaki); Site 25 (a 10 S 6 1 9) (all leg. Lymberakis); 
Site 32 (a 2 S S 1 9 ) (leg. Nikolakakis). 

ANTIKYTHIRA: Potamos, 700m W: sparse phrygana on sandy soil close to the village, 
(10 S S 3 9 9 , pitfall traps 27/3/2001 - 5/8/2001, leg. Chatzaki). 

Taxonomy. Males of this species have been collected from three islands: Crete, 
Gavdopoula and Antikythira. On all three islands, male specimens differ in size and in 
the prolateral extension of the terminal apophysis (sensu Platnick & Murphy, 1984). In 
males from Gavdopoula, which are the smallest, the tibial apophysis of some 
specimens is not bifid and is relatively more straight, whilst the extension of the 
terminal apophysis is rather small and has an extra angle below (Figs 34-35). Males 
from Antikythira are the largest. The tibial apophysis is almost the same as that of 
T. adriaticus from Crete, but the extension of the terminal apophysis has a characteris- 
tic hammer-like shape (Figs 36-37). Interestingly, these features fit the description of 
T. stubbsi Platnick & Murphy (1984, p. 9, Figs 15-16), known from a single male 
caught on Cyprus, and from a later record from Israel (Levy, 1998, p. 106, Figs 23-24). 
The female of T. stubbsi was unknown. All the female Trachyzelotes caught in the same 



GROUND SPIDERS OF CRETE. III. 



55 




Figs 20-25 
Trachyzelotes barbatus (20-21), T. malkini (22-23), T. lyonneti (24-25). Palp of S , ventral view 
(20, 22, 24), retrolateral view (21, 23, 25). Scale lines 0.1mm. 



traps along with "T. stubbsi" males, are indistinguishable from those of T. adhaticus 
(see Figs 30-31). We suggest that the differences mentioned above are due to intra- 
specific variability, and therefore we place T. stubbsi in the synonymy of T. adriaticus. 

Ecology. T. adriaticus has been found on few sites of western Crete and on 
Gavdos, where it occurs along with T. malkini. Apparently the distributions of T. adria- 
ticus and T. lyonneti do not overlap on Crete (Fig. 117). We presume that there must be 
some kind of competition between these two species, especially when taking into 
consideration the dispersal capacities of these spiders. 

Distribution. Italy, Balkans, China (first record for Greece). 



Setaphis carmeli (O. P.-Cambridge, 1872) 



Figs 40-41, 118 



Identification: Levy (1998, p. 96, Figs 2-5), Platnick and Murphy (1996, p. 6, Figs 5-8). 

Material. CRETE: CHANIA: Site 6 (c 2 S 3) (leg. Lymberakis); RETHYMNO: Site 25 
(a 3 S â) (leg. Lymberakis); IRAKLEIO: Site 44 (al S); Site 47 (a 1 S) (all leg. Nikolakakis); 
LASITHI: Site 63 (a 1 6) (leg. Stathi). 

Comparative material examined: S. carmeli: Israel, Golan heights, Odern forest (Id, 
5/1997, leg. Sharon, HUJ 15221). 



56 



M. CHATZAKI ET AL. 









Figs 26-31 



Trachyzelotes barbants (26-27), T. malkini (28), T. lyonneti (29). T. adriaticus (30-31). Epigyne 
(26, 28-29, 30), vulva (27, 31). Scale lines 0.1mm. " 



GROUND SPIDERS OF CRETE. III. 



57 






Figs 32-39 
Trachyzelotes adriaticus. Specimens from Crete (32-33, 38-39), Gavdopoula (34-35) and 
Antikythira (36-37). Palp of S, ventral view (32, 34, 36), retrolateral view (33, 35, 37, 39), tibial 
apophysis, dorsal view (38). Scale lines 0.1mm. 



58 M. CHATZAKI ET AL. 

Taxonomy. The characters of our specimens fit the descriptions and drawings by 
both Levy (1998) and Platnick & Murphy (1996). In our specimens, the embolus is 
long, forming two coils (Fig. 40). The examination of a male from Israel verified the 
identity of our specimens. Females have not yet been captured on Crete. 

Ecology. S. carmeli was found only in the lowlands of Crete and is not very 
common (Fig. 118). 

Distribution. Mediterranean (first record for Greece). 

Zelotes aerosus Charitonov, 1946 Figs 42-43, 118 

Identification: Charitonov (1946, p. 26, Fig. 41). 

Material. CRETE: CHANIA: Site 2 (a 1 S) (leg. Lymberakis). 

Taxonomy. This species was described by Charitonov (1946) from Uzbekistan 
and has not been recorded in the literature ever since. Although the type was not 
available to us, we feel sure that our specimen belongs to Z. aerosus, because of the 
shape of the tibial apophysis and the characteristic embolus, transversing the tegulum 
in its middle. Only one male prosoma was found in our collection. 

6 Pedipalp (Figs 42-43): Tibial apophysis long, with tapering end. Embolus 
starting at anterior part of tegulum with a broad base, then abruptly narrowed, filiform, 
forming a loop at the retrolateral side of the tegulum. Apical part of the tegulum with 
a conductor-like apophysis (A), composed of a tapering sclerotized process and a 
membrane. 

Comments. Two distinct characters of the palpal organ, i.e. the embolar base and 
the tegular process, as well as the shape of the female epigyne (see Charitonov 1946, 
Fig. 42), separate Z. aerosus from other zelotines. Its generic placement should be re- 
evaluated when more material is available. 

Ecology. Z. aerosus has been collected only at one locality, in phrygana close to 
the sea (Site 2). 

Distribution. Uzbekistan, Crete (first record for Europe). 

Zelotes caucasius (L. Koch, 1866) Figs 44-45, 48-49, 1 19 

Identification: Levy (1998, p. 137, Figs 92-96), Grimm (1985, p. 281, Figs 231, 
234-235). 

Material. CRETE: CHANIA: Site 1 (a 14 cî cî 8 9 9 ; b 2 cî cî 14 9 9 ; c 1 (5299); 
Site 2 (a 14 cîcî599;bl8cîcî23 9 9); Site 3 (a 10 6 6 6 9 9;b5 9 9); Site 4 (a 2 99;b 
2 6 6 1 9;c 1 9); Site 5 (a 3 6 6 1 9;b4 9 9); Site 6 (f 1 9; g 1 9); Site 7 (e 1 6\f4 99 
g 3 9 9);Site8(f8 cîcî599;g499;hl 9) (all leg. Lymberakis); Site 13 (a 2 6 6 16 9 9) 
Site 13 (c 10 6 6 6 9 9 ; d 12 6 6 19 9 9 ); Site 14 (a 3 cî cî 1 1 99;b399;c27ó*ó*14 99 
dl9 cîcî 44 9 9); Site 15 (c 5 cî cî 5 9 9 [MHNG]; d 2 6 6 5 9 9); Site 16 (a 1 6 5 9 9; c 5 
cîcî899;d6cî<îl 9); Site 17 (c 1 cî); Site 18(a999;c2cîcî799;d24cîcî 11 99); 
Site 19(c2cîcîl9;dlcî2 9 9 ); Site 20 (a 2 cîcî;b4 cîcî 7 9 9); Site 22 (a 1 cî 1 9;b2 
cî cî); Site 23 (a 1 cî; c 22 cî cî 19 9 9) (all leg. Paragamian); RETHYMNO: Site 35 (al 9); 
Site 36 (a 19 cî cî 7 9 9) (all leg. Trichas); Site 27 (a 9 cî cî 4 9 9); Site 34 (a 3 cî cî 1 9; b 2 
9 9); Site 37 (a 15cî<îl0 99;b999;c3 99;e6 cîcî); Site 55 (b 9 cî cî 42 9 9) (all leg. 
Chatzaki); Site 25(a23cîcî8 99;b5cîcî499; Site 26 (b 1 9 ) (all leg. Lymberakis); Site 
28 (a 5 cîcî 1 9; b 9 cîcî 15 9 9); Site 29 (b 1 cî 1 9); Site 32 (a 14 cîcî 10 9 9;b 15 9 9); 
Site 39 (a 2 cî cî; b 6 cî cî 21 9 9 ); Site 40 (b 29 cîcî 17 9 9; c 20 cîcî 22 9 9); Site 41 (a 43 
cî<î22 99;b4cî<î57 99) (all leg. Nikolakakis): IRAKLEIO: Site 43 (a 4 cî cî 2 9 9 ; b 12 
cîcî 26 9 9; c 6 9 9 ); Site 44 (a 5 cîcî;b8cîcî999;c499); Site 45 (a 4 cîcî 9 9 9; b 5 



GROUND SPIDERS OF CRETE. III. 



59 







Figs 40-43 
Setaphis carmeli (40-41), Zelotes aerosus (42-43). Palp of S, ventral view (40, 42), retrolateral 
view (41, 43). A: conductor-like apophysis. Scale lines 0.1mm. 



60 M. CHATZAKI ET AL. 



99); Site 47 (b 12 SS 13 99;c 1 9); Site 52 (b 6 SS 7 9 9 ); Site 53 (b 5 SS 3 99)(all 
leg. Nikolakakis);Site46(a21 de? 38 9 9; b 17 9 9 ); Site 49 (a 6 S S\b \\ S S 1 9 9;c \ 
9); Site 51(a4dc?2 99;bl9) (all leg. Papadimitrakis); Site 48 (a 2 9 9) (leg. Trichas); 
LASITHI: Site 27 (b 2 S S); Site 55 (a 7 S S 4 9 9 [CTh]); Site 56 (a 5 S S 2 9 9); Site 57 
(a 59 S S 62 9 9 ; b 4 9 9 ); Site 73(a8c?<?27 99;b2c?Q* 1 9 ; e 1 S) (all leg. Chatzaki); 
Site 58 (a 4 S S 13 9 9; b 2 9 9); Site 62 (a 2 9 9); Site 64 (d 3 S S 4 9 9); Site 65 (a 1 S 
3 9 9); Site 68 (a 2 9 9) (all leg. Papadimitrakis); Site 63 (a 1 S; b 1 S 4 9 9); Site 71 (a 6 
c?<?l9;b51<?c?4l99;c79 9) (all leg. Stathi); Site 61 (a 1 S); Site 70 (b 1 S); Site 72 
(a 1 9) (all leg. Trichas). 

KOS: Kefalos - Ag. Ioannis, 1 km S, phrygana and adjacent pine forest, (1 S 5 9 9, pit- 
fall traps 26/6/2001 - 9/9/2001, leg. Chatzaki). 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (1 S 3 9 5 , 
pitfall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

ANTIKYTHIRA: Potamos, 700m W: sparse phrygana on sandy soil close to the village, 
(38 S S 34 9 9 , pitfall traps 27/3/2001 - 5/8/2001, leg. Chatzaki). 

Taxonomy. Z. caucasius is a well-defined species, occurring in the whole 
Mediterranean region, its range extending to Central Europe, with northernmost local- 
ities in Slovakia and eastern Austria. Males are characterised by a prolateral tegular 
apophysis (TA) and a very long, curved embolus with a broad embolar base which 
forms a pointed retrolateral tip. In our specimens the tegular apophysis is lying close 
to the tegulum, whereas in the drawings of Levy (1998, Figs 92-93) it is directed to one 
side. The epigyne of females is characterised by an entire, transverse anterior margin, 
followed by a posterior one, leading into a large atrium. The introductory channels are 
situated between two large pouches, a smaller ventral pouch and a larger dorsal one, 
which underly almost the entire surface of the epigyne. As stated by Levy (1998), these 
characters clearly separate Z. caucasius from all other Zelotes. 

Ecology. This species has been captured very frequently in pitfall traps, as was 
also the case in Israel (Levy, 1998). It is very common on Crete, having been collected 
from most of the sites investigated (Fig. 1 19) and in great numbers. It is also present 
on all south Aegean islands investigated in this study. Adults occur during the whole 
year, with a peak of activity of females in mid-summer and of males in late spring and 
in early autumn. 

Distribution. Europe to Central Asia. 

Zelotes solstitialis Levy, 1 998 Figs 46-47, 50-51,118 

Identification: Levy (1998, p. 139, Figs 97-101). 

Material. CRETE: RETHYMNO: Site 39 (b 1 9 ); Site 40(c22 99;d2c?(?499 
[CTh]); Site 41 (b 1 9) (all leg. Nikolakakis); IRAKLEIO: Site 47(c7o*cM0 99;d299) 
(all leg. Nikolakakis); Site 46 (b 2 9 9); Site 50 (d 1 S 2 9 9 ) (all leg. Papadimitrakis); 
LASITHI: Site 27 (b 1 9) (leg. Chatzaki); Site 64 (d 1 S; e 16 SS 16 99) (all leg. 
Papadimitrakis); Site 64 (f 5 S S 6 9 9) (leg. Nikolakakis); Site 63 (c 1 S) (leg. Stathi). 

KOS: Kefalos - Ag. Ioannis, 1 km S, phrygana and adjacent pine forest, (1 ó* 5 9 9 , pit- 
fall traps 26/6/2001 - 9/9/2001, leg. Chatzaki). 

PELOPONNISOS: Mainalo Mt., alpine phrygana (2 SS, pitfall traps 9/7/1997 - 
12/10/1997, leg. Anastasiou). 

Taxonomy. This species is very close to Z. caucasius. Apart from Z. solstitialis 
being larger, the two species differ in the following characters: in males of Z. solsti- 
tialis there is no prolateral tegular apophysis, the retrolateral tip of the embolar base is 
more pronounced and the embolus itself is longer, reaching almost to the tip of the 



GROUND SPIDERS OF CRETE. III. 



61 







Figs 44-47 
Zelotes caucasius (44-45), Z. solstitialis (46-47). Palp of o\ ventral view (44, 46), retrolateral 
view (45, 47). TA: tegular apophysis. Scale lines 0.1mm. 



62 



M. CHATZAKI ET AL. 







\ 51 



Figs 48-51 
Zelotes caucasius (48-49), Z. solstitialis (50-51). Epigyne (48, 50), vulva (49, 51). Scale lines 
0.1mm. 



cymbium. The tibial apophysis in Z. solstitialis has an additional broad, ventral lobe. 
Female genitalia differ in the shape of the central opening, in the dimensions of the two 
pouches, and in the course of the introductory ducts. A small gland exists on the lateral 
side of each dorsal pouch. 

Ecology. As already stated by Levy (1998), adults of this species occur only 
during the dry months of summer and early autumn. 

Distribution. Israel, Syria? (first record for Europe). 



GROUND SPIDERS OF CRETE. III. 63 



Zelotes subterraneus (C. L. Koch, 1833) Figs 52-56, 120 

Identification: Grimm (1985, p. 227, Figs 282, 303-304). 

Material. CRETE: CHANIA: Site 2 (al 9 ; c 26 6 6 1 2 ; d 1 9;e2 99;f8 99 
[CTh]); Site 3 (c 33 Ó*ó*299;dl7 99;e3 9 9); Site 5 (al 9;c 12 6 6 5 9 9); Site 6 (a 4 
99;b299;c399;h38c?d 1 9;i2o*c?999;j499;kl 9;1499;ml 9); Site 7 
(i9 66) (all leg. Lymberakis); Site 14 (a 1 6; b 1 6 1 9; c 1 9); Site 15 (c 3 66 1 9); Site 
16(a4ó*ó*;b799;c6 9 9); Site 18 (a 2 6 6 1 9;b 1 6); Site 19 (b 1 9); Site 21 (a 2 6 6) 
(all leg. Paragamian); RETHYMNO: Site 27(a299;bl5(?cM9;c24c?<M8 99;d2c?c? 
999;el<?499;fl4 9 9); Site 34 (b 10 o A <?;c3o*c?299;d4 9 9); Site 37 (b 12 6 6; 
cl8 99;dl4 99;e299); Site 38(alcMl99b499) (all leg. Chatzaki); Site 25 (c 16 
66 3 9 9) (leg. Lymberakis); Site 24 (a 1 9 ) (leg. Stathi); Site 28 (a 2 9 9 ); Site 29 (b 6 6 6); 
Site 32 (b 9 6 6); Site 39 (b 5 9 9); Site 40 (b 1 9 ; d 37 6 6 9 9 9 ; e 1 9 ); Site 41 (a 4 9 9 ) 
(all leg. Nikolakakis); Site 30 (al 9); Site 33 (a 2 6 6 2 9 9); Site 35 (al 6); Site 36 (a 1 9) 
(all leg. Trichas); IRAKLEIO: Site 42(el2 99;f299;h299;i53<?c? 11 2 9; ] 20 66 
15 99;hl5c?d;k8 9 9 ) (all leg. Chatzaki); Site 43 (a 3 9 9;c2 66 1 9); Site 44 (a 4 99; 
cl 66); Site 45 (a 2 9 9); Site 47 (a2 ?2;bl 6 6 \0 2 2; c 9 6 6; d U 6 6 6 2 2; e 
2 9 9); Site 50 (a 4 9 9; d 4 6 6; e 31 6 6 8 9 9 ); Site 52 (a 3 9 9;b 1 9; c 44 6 6 6 9 9); 
Site 53 (a 1 9 ; c 20 6 6 12 9 9 ); Site 54(a299;cl7c5c?399) (all leg. Nikolakakis); Site 
46 (b 23 6 6); Site 49 (a 1 9 ; b 1 9 ; c 15 6 6); Site 51 (b 3 6 6) (all leg. Papadimitrakis); Site 
48 (a 1 (?) (leg. Trichas); LASITHI: Site 55 (a 5 9 9 ; b 36 ö* c? 4 9 9 ; c 21 66 1 2 2; d\ 2; 
e 1 9); Site 56 (a 1 9; b 16 6 6 2 2 2; c 1 9); Site 57 (a 1 6 [CTh]; b 5 6 6 1 9;c2 9 9 
[MHNG]); Site 73 (a 3 9 9; b 8 6 6 [MHNG]; c 26 6 6 12 9 9; d 2 9 9; e 4 9 9) (all leg. 
Chatzaki); Site 71 (a 8 9 9 ; c 1 9 ; d 14 6 6) (all leg. Stathi); Site 64 (a 2 6 6 1 9;c3 99;e 
5<?o A ;f28c?c?7 99) (all leg. Nikolakakis); Site 58 (b 10 6 6 1 9 ); Site 62 (a 4 9 9 ; b 16 
66 12); Site 65 (a 2 9 9); Site 68 (b 5 66) (all leg. Papadimitrakis); Site 63 (a 6 9 9; c 38 
6 6 8 9 9); Site 60 (a 2 9 9); Site 71 (c 31 6 6; d 1 9; e 3 9 9) (all leg. Stathi); Site 59 (a 
I 6;b2 66 I 2; cl 6); Site 61 (aU 6 6 3 2 2;b \4 6 6 4 9 9 ); Site 66 (a 2 9 9); Site 67 
(a 2 6 6 3 2 2); Site 70 (b 3 6 6 1 2 2); Site 72 (a 2 9 9; b 12 6 6) (all leg. Trichas). 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (599, pitfall 
traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

PELOPONNISOS: Mainalo Mt., alpine phrygana (2 9 9, pitfall traps 9/7/1997 - 
12/10/1997, leg. Anastasiou). 

Comparative material examined: Z. subterraneus: Austria, N. Tirol, Halltal 1000- 1200m 
(39 9, 13/6/1992, leg. Knoflach); Austria, N. Tirol, Ötztal, Längenfeld (16, 8/8/1992, leg. 
Knoflach). 

Taxonomy. This species clearly belongs to the subterraneus group. Among the 
central-European species of the group recorded by Grimm (1985), Z. subterraneus is 
closest to our specimens, albeit rather larger in size (TL 6 = 5.7-7.8, 9 = 5.8-9.2, ac- 
cording to Grimm versus TL 6 = 3.8-6.8, 9 = 6.4-9.2) and deviating slightly in the 
shape of the vulva. However, male characters fit well the description by Grimm (see 
Figs 52-56). A decision about the status of these populations is postponed until more 
material from Northern Greece is available. 

Ecology. Z. subterraneus is common all over Crete, Gavdos and Dia (Fig. 120). 
Adult females are present throughout the year, males are very rare in the dry period. 
The peak of activity of both males and females occurs in autumn to early winter. Z. 
subterraneus is one of the very few zelotines collected in the pitfall traps during win- 
tertime. 

Distribution. Trans-Palearctic, in the temperate and boreal zones, recently 
recorded from Bulgaria (Deltshev & Blagoev, 2001) (first record for Greece). 



64 



M. CHATZAKI ETAL. 





55 



r' 



56 





Figs 52-56 
Zelotes subterraneus. Palp of S, ventral view (52), dorsal view (53), retrolateral view (54), 
epigyne (55), vulva (56). Scale lines 0.1mm. 



Zelotes creticus (Kulczynski, 1903) Figs 57-61, 121 

Identification: Kulczynski (1903, p. 42, Fig. 4). 

Material. CRETE: CHANIA: Site 5(a5 99;bl9;c23dd499); Site 6 (a 1 6 ; b 

1 9;c2 9 9;d 1 9; e 8 9 5; h 5 S8\il Sa [CTh]; 1 1 â 1 9); Site 7 (a 1 <5;b8 9 9;c2 



GROUND SPIDERS OF CRETE. III. 



65 






Figs 57-61 
Zelotes creticus. Palp of 3 , ventral view (57), dorsal view (58), retrolateral view (59), epigyne 
(60), vulva (61). Scale lines 0.1mm. 

99;d999;el3 99;f399;gl8c?d;h38dc?;h599;il<?299;j3 9 9); Site 8 (a 
3 6â;b4 ââ;d\ 9;fl 9;h4SS 1 9;ill SS 1 9;j2 c?c5;kl 9); Site 9 (a 8 SS I 9; 
b7 SS 19 ; c399;d399; e7 SS ;f4 SS; gl S 2 9 9 ) (all leg. Lymberakis); Site 11 (a 
1 9 ; b 3 9 9 ); Site 12 (e 1 9 ) (all leg. Stathi); RETHYMNO: Site 25 (a 7 9 9 ; c 25 c? S 4 9 9 ); 
Site 26(a499;bl9;cl5<?c?3 99) (all leg. Lymberakis); Site 32 (a 12 9 9 ; b 10 S S 5 
9 9); Site 40 (a 7 9 9 [CTh]; b 6 9 9; c 2 S S; d47 S S 7 9 9 ; d 3 c? <? 3 99 [MHNG]; e 
1 9 ) (all leg. Nikolakakis). 



66 M. CHATZAKI ET AL. 

Taxonomy. Zelotes creticus was described by Kulczynski (1903) based on two 
female specimens and the species has not been revised since that time. We collected fe- 
males of this species together with their males, which we present here for the first time. 

Measurements 6(9), n = 13 (12): TL: 5.6-7.4 (6-9.1), PL: 2.4-3.4 (2.6-3.6), 
PW: 1.8-2.6 (2-2.8), OL: 2.8-3.7 (2.6-5.1), PL/PW: 1.19-1.66 (1.18-1.4). 

â 2 : Large spiders of dark colour. Eyes round. Posterior and anterior rows of 
eyes straight. Males with large, brown scutum on the opisthsoma. 

Legs: Me III-IV with apical preening combs. Ta and Me I-II with stripes of ven- 
tral scopula hairs, Ta and Me III-IV with dense spiny hairs. Spination: Fe: I d 2; II d 3; 
III d 3-5; IV d 2-4. Pa: I-II, IV - ; III r 1. Ti: I-II - ; III-IV spinose. Me: I-II - ; III-IV 
spinose. 

â Pedipalp (Figs 57-59): Tibial apophysis slender, straight, longer than l / 2 of 
the length of the cymbium. Tegulum robust, with a sharp projection at its anterior/pro- 
lateral edge, probably corresponding to terminal apophysis. Embolus short, filiform, 
transverse, starting from dorsal/retrolateral front of tegulum, retinaculum at its 
anterior/retrolateral corner. 

Epigyne (Fig. 60): Anterior margins small, cap-like, far in front of the areola. 
Lateral margins relatively short, curved; posterior margin faint, biconvex. Introductory 
orifices at the edge of lateral and posterior margins. 

Vulva (Fig. 61): Introductory ducts short and sclerotized, transverse, with 
glandular heads at their anterior ends. Spermathecae globular, with fertilisation ducts 
situated at their anterior part. 

Ecology. Z. creticus is common in the middle to high altitudes of the mountains 
of western Crete, but absent from eastern Crete. Also the type material of Kulczynski 
was collected from middle altitudes of western Crete (Chania, Askifou, 1000m). The 
species has also been caught three times in phrygana of the lowland (Sites 25, 26 and 
32) (Fig. 121). Together with Callilepis eretica, Drassodes lapidosus, Drassyllus 
praeficus and others (see also Chatzaki et al. 2002 a, b), Z. creticus belongs to the few 
gnaphosid species which become more numerous with increasing altitude. The peak of 
activity of males occurs in late summer and of females in late spring and early autumn. 

Distribution. Crete (endemic). 

Zelotes cf. ilotarum (Simon, 1884) Figs 62-64, 68-69, 121 

Material. CRETE: CHANIA: Site 4 (c 2 3 3 1 9 ) (leg. Lymberakis); IRAKLEIO: Site 
44 (a 9 3 3 6 9 9 ; b 1 9 ); Site 47(a999;b699;d27ó*ó*299d2ó*d299 [MHNG] 
e 6 9 9 [CTh]) (all leg. Nikolakakis); Site 49 (a 1 3 [CTh]; a5 99;b2 99;c5<?o*3 99) 
Site 51 (a2l99;b29c?<?l9) (all leg. Papadimitrakis); LASITHI: Site 59 (a 59 3 3 12 9 9 
bS 33 \ 9; c 38 33 20 99) (all leg. Trichas). 

Comparative material examined: Z. ilotarum: Peloponnisos, Aigio (233, 3/4/1939, Col. 
Hadjissarantos, 2780, ZMUA); Attiki, Kato Souli (4 9 9, 14/4/1939, Col. Hadjissarantos, 2781, 
ZMUA). 

Taxonomy. Z. ilotarum was described by Simon (1884) on the basis of females 
from Naxos and Athens. It was rediscovered in Peloponnisos and Attiki by 
Hadjissarantos (1940), who presented the first illustration of the epigyne and described 
the male for the first time (p. 82, Figs 26-27). This species is characterised by the 
distinct shape of the embolic division (EB and E) of the male palp (Figs 65-67) and by 



GROUND SPIDERS OF CRETE. III. 



67 




Figs 62-67 
Zelotes cf. ilotarum (62-64), Z. ilotarum (65-67). Palp of 3 , ventral view (62, 65), dorsal view 
(63, 66), retrolateral view (64, 67). TA: terminal apophysis, p: prolateral process, EB: embolar 
base, E: embolus X: dorsal extension. Scale lines 0.1mm. 



68 M. CHATZAKI ET AL. 

the female epigyne, which is narrow compared to its length (Figs 70-71). The pro- 
minent edge of the tegulum has a prolateral process (p), a dorsal extension (X) of the 
intercalary sclerite (sensu Platnick & Shadab, 1983) and a transverse terminal apo- 
physis (TA), which are also dinstinctive characters. We have found males and females 
of a species from Crete, which is very close to Z. ilotarum, but not identical, as will be 
discussed later. 

Measurements 6(9), n = 11 (12): TL: 5.2-7.1 (5.7-9.2), PL: 2.4-3 (2.5-3.5), 
PW: 1.8-2.4 (2-2.7), OL: 2.6-3.8 (3-4.9), PL/PW: 1.21-1.37 (1.19-1.45). 

Ô 9 : Dark brown spiders of medium to large size. Eyes round, posterior and 
anterior rows straight. Males with large scutum on opisthosoma. 

Legs: Me III-IV with apical preening combs. Ta and Me I-II with scopula hairs, 
Ta and Me III-IV with spiny hairs. Spination: Fe: I-II d 2; III-IV d 3-6. Pa: I-II - ; III- 
IV r 1. Ti: I-II - ; III-IV spinose. Me: I-II v 2; III-IV spinose. 

ó* Pedipalp (Figs 62-64): Tibial apophysis long, tapering. Tegulum robust, 
strongly sclerotized, with a prolateral process situated at its apical angle (p). Intercalary 
sclerite of tegulum continuing to dorsal side by a transverse extension (X). Embolic 
division on retrolateral side of tegulum, with a strong, S- shaped base (EB) bearing 
microdenticles on its surface and a small sclerotized embolus (E), visible only from the 
dorsal side of the bulbus. Terminal apophysis (TA) triangular, dorsal. Retinaculum 
transverse, at apical end of tegulum. 

Epigyne (Fig. 68): Anterior margin with two caps, well separated. Lateral 
margins narrow, parallel to each other, with introductory orifices at their rims. 

Vulva (Fig. 69): Introductory ducts short, coiled, leading to the centre, to 
globular spermathecae. Glandular heads located below lateral margins, anterior to the 
introductory ducts. Dorsal surface of the introductory channels fused to an oval plate 
(OP). 

Comments. Z. ilotarum and our species apparently are very similar. The same 
structures are present in males as well as in females. However, the 'true' Z. ilotarum 
(sensu Hadjissarantos) differs in the following characters: in males (compare Figs 
62-64 and 65-67) the prolateral tegular process (p) is broader; the S-shaped sclerotized 
embolar base is stronger, its end wider and bifid; the terminal apophysis is more trans- 
verse. In females (compare Figs 68-69 and 70-71) the anterior caps are further apart 
than the width of the lateral margins; the spermathecae are larger and closer together; 
the glandular heads are at the same level as the introductory ducts. Differences in the 
oval plate of the females examined are probably due to a difference in age. 

When a more thorough knowledge of the gnaphosid fauna of Greece has been 
obtained and the distributions of the species has become more precisely known, it will 
be possible to decide whether there already exists a species name available in the 
literature, or whether this form should be regarded as a variation of Z. ilotarum only. 

Ecology. Z. cf. ilotarum is not very common on Crete (Fig. 121). It has been 
found at several sites in central and eastern Crete, mainly in the lowlands. It has not 
been found above 1000m. The peak of activity in males occurs in early autumn and in 
females in late spring. 

Distribution. GREECE: Attiki ?; Peloponnisos ? (Hadjissarantos, 1940); Naxos 
? (Simon, 1884); Crete. 



GROUND SPIDERS OF CRETE. III. 



69 




68 



r\ r^ 69 





70 



71 





Figs 68-71 
Zelotes cf. ilotarum (68-69), Z. ilotamm (70-71). Epigyne (68, 70), vulva (69, 71). OP: oval 
plate. Scale lines 0.1mm. 



Zelotes scrutatus (O. P.-Cambridge, 1872) 



Figs 72-76, 122 



Drassodes cofiniotes: Roewer, 1928 (p. 105, Fig. 10), CRETE: Rethymno, Topolia (type loca- 
lity); Roewer 1954 (p. 385); Bonnet, 1956 (p. 1563); Levy, 1998. 

Zelotes bucharensis Charitonov, 1946: p. 26, Figs 39-40, UZBEKISTAN, Yakkabagh (type 
locality). Syn. n. 



70 M. CHATZAKI ET AL. 



Identification: Levy (1998, p. 125, Figs 62-68). 

Material. CRETE: CHANIA: Site 1 (a 3 S S 6 9 9 ; b 4 S S;c24 3 S); Site 2 (a 9 S S 
3 9 9;bl S 2 9 9); Site 3 (a 3 cîc?;bl <î); Site 4 (al 9;b2 SS 5 9 9); Site 5 (al o*;bl 
S) (all leg. Lymberakis); Site 13 (a 2 S S 109 9; c 1 S; d 2 <î Ó* 4 9 9); Site 14 (a 1 S 1 2; 
c499;d3o*c?2 9 9); Site 15 (d 2 SS); Site 16 (d4 SS 3 9 9); Site 18 (d 1 9); Site 19 (a 
l(?599;clc?399;d5(?<?299) (all leg. Paragamian); RETHYMNO: Site 25 (a 8 S S; 
a299;b499) (all leg. Lymberakis); Site 27(a22o*ó*399;b8ó*ó*299;cl 9 ; f 1 S); 
Site 34 (a 1 S ) (all leg. Chatzaki); Site 28 (a 4 9 9 ; b 1 S 2 9 9 ); Site 32 (a 3 S S 3 9 9 ); Site 
39(a5c?<?299;b2c?<?2 9 9); Site 40 (b l\ S S \ 9; c 4 S S 4 9 9; d l 9); Site 41 (a 
llc?c?2 99;b21c?<?2 99) (all leg. Nikolakakis); IRAKLEIO: Site 42 (e 2 S S 1 9 ; g 1 
9;f2 9 9) (all leg. Chatzaki); Site 44 (a 5 SS [CTh];a2 9 9;b7 SS 10 9 9 ); Site 47 (a 4 
SS 1 9\b25 SS; b 8 99 [CTh]; c 2 S S 5 9 9 [MHNG]; e 3 SS) (all leg. Nikolakakis); 
Site 46 (a 3 SS 2 9 9;b 1 9); Site 49 (b 2 9 9); Site 50 (c 2 SS 2 9 9;d 1 9); Site 51 (al 
9 ) (all leg. Papadimitrakis); LASITHI: Site 55 (a4 SS 1 9;b5 SS 1 9;cl 9); Site 73 (a 1 
S; b 1 9) (all leg. Chatzaki); Site 58 (a 1 S); Site 62 (a 1 S; b 1 S); Site 64 (d 2 S S 1 9; e 1 
S 1 2) (all leg. Papadimitrakis); Site 63 (b 2 c? cî); Site 71 (a3 S S;b3 S S 1 9; c 1 S) (all 
leg. Stathi); Site 66 (a 3 S S 3 9 9 ); Site 70 (b 1 9 ) (all leg. Trichas). 

PELOPONNISOS: Mainalo Mt., alpine phrygana (1 S, pitfall traps 9/7/1997 - 
12/10/1997, leg. Anastasiou). 

Comparative material examined: Z. scrutatus: Israel, Hatira ridge near Sede Boger (1 S 
1 9 , 8/1991, Col. Lubin, 14628). 

Taxonomy. The taxonomical characters of this species fit well the description of 
Levy (1998) and with the specimens received for comparison from Israel (see terminal 
apophysis (TA), embolar base and tip of embolus in Figs 72-73). In our material, these 
characters and the shape of the median plate of the females are constant, while the shape 
of the introductory channel varies, as reported by Levy (1998). Z. bucharensis 
Charitonov, 1946 is proposed here as a new synonym, on the basis of congruence in the 
shape of the median plate and the tibial apophysis (see Charitonov 1946, Figs 39-40). 

Ecology. Z. scrutatus is common and widespread all over Crete (Fig. 122), 
occurring also on the surrounding islands of Gavdos and Dia and on mainland Pelo- 
ponnisos. On Crete, it reaches altitudes up to 1200m, but seems to prefer the lowlands. 
Adults' peaks of activity occur from late spring to early autumn. 

Distribution. North Africa, Israel, Syria, Uzbekistan; GREECE: Attiki, 
Kokkinos Mylos (Hadjissarantos, 1940, sub D. cofiniotes). 

Zelotes minous Chatzaki sp. n. Figs 77-81, 122 

Etymology: The name of this species is derived from Minos, king of Crete during the 
Minoan era; adjective. 

Material. Type material: Site 27a (Eksantis, Agios Kyprianos) (1 S holotype, 4 9 9 
paratypes, [NHMC]; 1 6 1 9 paratypes [MHNG]). 

CRETE: CHANIA: Site 1 (b 1 9); Site 6 (g 1 9) (all leg. Lymberakis); Site 16 (a 1 9; 
d 3 9 9); Site 17 (c 1 S); Site 19 (d 3 S S) (all leg. Paragamian); RETHYMNO: Site 27 (a 2 
SS 5 9 9 ; b 1 9 ) (all leg. Chatzaki); IRAKLEIO: Site 52 (b 1 9 ); Site 53 (b 1 9 ); Site 54 (b 
1 S)\ (all leg. Nikolakakis); LASITHI: Site 55 (a9 S S;b2 S S [CTh]); Site 73 (a 2 9 9) (all 
leg. Chatzaki); Site 63 (b 1 9) (leg. Stathi); Site 65 (a 1 <?) (leg. Papadimitrakis). 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (4 S S 1 9 . 
pitfall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

Comparative material examined: Z. puritanus Chamberlin, 1922: Austria, N. Tirol, 
Innsbruck, Martinswand (3 S S 1 9, pitfall traps 23/4/1963 - 7/6/1963, CTh). 

Taxonomy. Measurements <?($), n = 7 (8): TL: 2.9-4.8 (4.4-4.9), PL: 1.5-2.1 

(1.7-1.9), PW: 0.9-1.6 (1.3-1.6), OL: 1.4-2.5 (2.3-2.8), PL/PW: 1.2-1.66 (1.13-1.3). 



GROUND SPIDERS OF CRETE. III. 



71 





76 




Figs 72-76 
Zelotes scrutatus. Palp of S , ventral view (72), dorsal view (73), retrolateral view (74), epigyne 
(75), vulva (76). Scale lines 0.1mm. 

S 9 : Small spiders of light brown colour. PME closer to PLE than to each other, 
smaller than the rest of eyes, round. 

Legs: Me III-IV with apical preening combs. All Ta, Me, Ti full of long hairs. 
All Co, Me, Ta, Pa and Ti III-IV yellow, other articles brown. Spination: Fe: I-IV d 1- 
3. Pa: I-IV - . Ti: I-II - ; III v 3 r 1; IV v 5-6 r 1. Me: III - ; III p 3 r 2; IV v 2 p 1 r 2. 



72 



M. CHATZAKI ET AL. 






Figs 77-81 
Zelotes minous sp. n. Palp of S, ventral view (77), dorsal view (78), retrolateral view (79), 
epigyne (80), vulva (81). EB: embolar base. Scale lines 0.1mm. 



â Pedipalp (Figs 77-79): Tibial apophysis short. Tegulum broadly membra- 
nous. Embolar base (EB) bulging, with a sclerotized dorsal process. Embolus short, 
directed along retrolateral side of tegulum, rising from dorsal end of embolar base. 



GROUND SPIDERS OF CRETE. III. 73 

Epigyne (Fig. 80): Anterior margin continuous. Median plate tongue-like, its 
anterior ends curved. 

Vulva (Fig. 81): Introductory orifices probably situated at curved part of median 
plate, leading to a large pouch, with lateral glandular heads. Introductory ducts short. 
Spermathecae large, globular. 

Comments. This species is closely related to Z. puritanus Chamberlin, 1922 (see 
Grimm 1985, Figs 247c, 248, 270-271). Apart from being much larger, Z. puritanus 
also differs in details of the embolic division of the male palp and in the shape of the 
median plate of the epigyne. 

Ecology. Z. minous sp. n. is widespread on the island of Crete, though not very 
common (Fig. 122). It also occurs on the islands of Gavdos and Dia. It prefers 
phryganic habitats close to the sea and was never collected in mountainous areas. 

Distribution. Crete, Karpathos (Greek endemic?). 

Zelotes nilicola (O. P.-Cambridge, 1874) Figs 82-86, 118 

Identification: Platnick & Shadab (1983, p. 186, Figs 263-266), Levy (1998, p. 142, Figs 
104-107). 

Material. CRETE: CHANIA: Site 18(clc?l9;d3c?tf299) (all leg. Paragamian); 
IRAKLEIO: Site 48 (a 5 8 8 1 9 ) (leg. Trichas); LASITHI: Site 64 (d 4 8 8 3 9 ? ) (leg. 
Papadimitrakis). 

Taxonomy. The taxonomical characters of this species fit well the descriptions 
of Platnick & Shadab (1983) and of Levy (1998). The palpal organ possesses a strange, 
saw-like, dorsal process (p) originating from the embolar base, as already reported by 
Levy (1998). 

Ecology. This species is rare on Crete (Fig. 118). It has been collected at three 
sites on Crete and at one on Gavdos, all of them close to the coast and to fresh water, 
in areas with sandy soil. Adults occur in late summer. 

Distribution. Circum-Mediterranean: Egypt, Tunisia, Corsica, Giglio Isl., 
Cyprus, Israel, Canary Isis., GREECE: Attiki: Athens (Platnick & Shadab, 1983), 
introduced to the USA (according to Platnick & Shadab, 1983). 

Zelotes labilis Simon, 1914 Figs 87-90, 93-96, 98-101, 104, 123 

Identification: Hadjissarantos (1940, p. 82, Figs 28-29). 

Material. CRETE: CHANIA: Site 3 (a 10 c? c? 5 99;b2<?c?299;cl 9); Site 4 (b 
1 9); Site 6 (d 1 8; e 4 SS 1 9; f 1 8); Site 7 (e 1 S 1 9; f 1 9; g 1 9); Site 8(eH;f 
2dc?l9;g299;kl 9); Site 9 (a 8 99;c2c?c?l9;d799;g3 9 9); Site 10 (a 1 S 4 
9 9); (all leg. Lymberakis); Site 14 (c 15 8 8 2 9 9; d 5 8 8 5 9 9); Site 15 (c4 83; d2 3 3 
1 9); Site 16(c21 33 1 9;d 15 33 11 9 9; 1 8 [CTh]); Site 17 (c 7 3 8; d 2 8 8 3 9 9); 
Site 18 (al 9;c5 8 8;c299 [CTh]; d 2 83 5 9 9); Site 19 (c 5 3 8 1 9; d 6 8 8 2 9 9); 
Site 20 (a 1 8; b 1 8 1 9); Site 21 (d 3 9 9); Site 22 (b 4 8 8); Site 23 (c 7 3 8 6 9 9) (all 
leg. Paragamian); RETHYMNO: Site 25 (a 7 9 9 ; b 1 9 ) (all leg. Lymberakis); Site 27 (a 27 
3 8 6 9 9;bl 3 2 9 9 ) (all leg. Chatzaki); Site 34 (a 2 33 3 9 9; b 1 3 7 9 9; e 1 9); Site 
37(al8c?c?1399;blc?499;el<?l 9) (all leg. Chatzaki); Site 35 (a 1 9 ) (leg. Trichas); 
Site 28 (al c?299;bl0c?<?9 9 9); Site 29 (b 1 9); Site 32 (a 3 9 9;b 1 9); Site 39 (a 2 
88 4 9 9); Site 40 (b 8 3 3 10 9 9; c 9 9 9); Site 41 (a 18 3 8 11 9 9; b 1 8 9 9 9) (all 
leg. Nikolakakis); IRAKLEIO: Site 42(tS 8 8 4 9 9;fS 8 8 1 9;g299) (all leg. Chatzaki); 
Site 43 (al 8;bl 8 2 9 9 ); Site 44 (b 5 8 3 1 9); Site 45 (a 3 88 1 99;bl 8 6 9 9); Site 
46 (a 17 8 8 11 9 9; b 7 9 9); Site 47 (b 12 3 8 16 9 9; e 2 3 8 1 9); Site 49 (b 2 33 



74 



M. CHATZAKI ET AL. 




Figs 82-86 
Zelotes nilicola. Palp of 6, ventral view (82), dorsal view (83), retrolateral view (84), epigyne 
(85), vulva (86). p: dorsal process. Scale lines 0.1mm. 



GROUND SPIDERS OF CRETE. III. 



75 









Figs 87-92 
Zelotes labilis, specimens from the lowlands (87-88), specimens from high altitudes (89-90), Z. 
daidalus sp. n. (91-92). Palp of S, ventral view (87, 89, 91), retrolateral view (88, 90, 92). TA: 
terminal apophysis. Scale lines 0.1mm. 



5 9 9); Site 50 (c U6??;dl 9); Site 51 (a 1 S 1 2; b 1 2); Site 52 (b 1 2); Site 53 (b 1 
9 ) (all leg. Nikolakakis); LASITHI: Site 55(a8cîc?222;b2c?c?422); Site 56 (a 6 3 S 
4 9 2; b 1 S 1 2); Site 57 (a 7 SS 4 9 9); Site 73 (a 1 S 13 9 9; b 1 S 1 9) (all leg. 
Chatzaki); Site 58 (a 2 SS 4 9 2;b4 9 9 ); Site 62 (a 1 Ó* 3 99;bl 9);Site64(dl Ó* 1 2); 
Site 65 (al 9); Site 68 (a 1 9) (all leg. Nikolakakis); Site 63 (b 1 S 1 9); Site 71 (b 3 o* S 5 
9 9 [MHNG]) (all leg. Stathi); Site 59 (a 1 S) (leg. Trichas). 



76 M. CHATZAKI ET AL. 



ANTIKYTHIRA: Potamos, 700m W: sparse phrygana on sandy soil close to the village, 
(1 S 3 2 9, pitfall traps 27/3/2001 - 5/8/2001, leg. Chatzaki). 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (4 2 9, pit- 
fall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

KOS: Kefalos - Ag. Ioannis, 1 km S, phrygana and adjacent pine forest, (6 2 2, pitfall 
traps 26/6/2001 - 9/9/2001, leg. Chatzaki). 

PELOPONNISOS: Mainalo Mt., alpine phrygana (1 6 2 2 2, pitfall traps 9/7/1997 - 
12/10/1997, leg. Anastasiou). 

Comparative material examined (vidit M. Chatzaki): Z. labilis: Attiki, Ekali (IS, 
10/5/1935, Col. Hadjissarantos, ZMUA); Samos, Koumaradaioi (12, 1/8/1937, Col. 
Hadjissarantos, ZMUA). 

Taxonomy. Z. labilis was first described by Simon (1914) based on female spec- 
imens from the Pyrenees. Later Hadjissarantos (1940) identified one female from 
Samos by using the key of Simon, and one male from Attiki, which he described for 
the first time. We are therefore uncertain about the true identity of the species collect- 
ed by Hadjissarantos and about its relationship to Z. labilis (sensu Simon). However, 
as our specimens fit the description of Hadjissarantos (see shape of tibial apophysis, 
terminal apophysis and median plate of the epigyne), we follow his opinion. This 
species shows a high variability in genital characters, as will be discussed later. A more 
detailed study, covering the whole range of their distribution, may prove that these dif- 
ferent forms belong to more than one species. 

Measurements 0^(2), n= 142 (200): TL: 2.75-5.35 (2.9-7.4), PL: 1.3-2.25 (1.3- 
2.25), PW: 1-1.8 (0.95-2.05), PL/PW: 1.17-1.63 (1.1-1.52). 

Pedipalp 6 (Figs 87-90, 93-96): Tibial apophysis small, conical. Tegulum oval, 
partly membranous, with sperm duct following its periphery. Terminal apophysis (TA) 
triangular, situated at the apical end of the tegulum. Position of retinaculum retrolater- 
al, apical. Embolus small, situated dorsally to the terminal apophysis. Embolar base 
bulging, rounded, forming a median hole with strongly sclerotized rim (H). The shape 
of this hole changes when seen from different angles (compare Figs 93 and 94) and it 
also differs among specimens. Here we give the two extreme forms (compare dorsal 
view Figs 93, 95 and dorsal/prolateral view 94, 96). In specimens from the lowlands 
(Figs 87-88 and 93-94) the rim is rounded, with a small tip; in specimens from Lefka 
Ori, at altitudes above 1650m (Figs 89-90 and 95-96), it is more prominent, forming a 
triangular process (p). In specimens from the other mountain regions of Crete the rim 
has two pointed tips (Lasithi, Limnakaro plateau) or is provided with some small tips 
(Psiloreitis Mt.). 

Epigyne (Fig. 98): Anterior margin divided, with two well-separated caps. 
Median plate U-shaped, often V-shaped, and longer than wide. In specimens from high 
altitudes the median plate is as long as wide (see Fig. 100). 

Vulva (Fig. 99): Introductory channels starting at lateral sides of the median 
plate with a pouch, then leading through transverse ducts to globular spermathecae. 
Glandular heads of variable size, attached to the pouches. In specimens from high al- 
titudes (Fig. 101) the pouches are larger and the distance between the posterior rim of 
the median plate and the spermathecae is greater. 

Comments. This species presents great variability in size and in morphological 
characters of the genitalia. Out of 142 male and 200 female specimens examined, on- 
ly 22 and 9, respectively, are considered as large (TLó* > 4.5mm and TL 2 > 6mm), 



GROUND SPIDERS OF CRETE. III. 77 

while the majority are of smaller size. This large form, mostly present in the high alti- 
tudes of Crete, shows some differences in the genitalia, as mentioned above. Also spec- 
imens from Mt. Mainalo in Peloponnisos belong to a special type. As there are inter- 
mediate forms between these extremes (see also Fig. 104), we place all these speci- 
mens into one species. A further analysis of the genitalia and in a wider geographical 
range must be conducted, in order to achieve a better understanding of the taxonomic 
structure of this 'species'. 

Ecology. Z. labilis is very common on Crete and surrounding islands (Fig. 123), 
as well as on Peloponnisos. The small lowland form occurs up to 1200m, while the 
large form was found from 1650m upwards. The two extremes are apparently inter- 
connected by a zone of transition. Adults occur from spring to late autumn, with a peak 
of activity from late spring to early summer. 

Distribution. France, Sicily, Greece. 

Zelotes daidalus Chatzaki sp. n. Figs 91-92, 97, 102-103, 123 

Etymology: The name is taken from Greek mythology: Daidalos was the architect of the 
Minoan palace of Knossos; noun in apposition, hence invariable. 

Material. Type material: Site 27a (Exantis, leg. Chatzaki) (1 6 holotype, 2 9 9 
paratypes [NHMC]; 1 S 1 2 paratypes [MHNG]). 

CRETE: CHANIA: Site 2 (a 7 6 o* ; b 3 3 <?); Site 3 (a 2 9 9 [CTh]; b 1 6); Site 6 (f 1 
(?) (all leg. Lymberakis); Site 16 (c 1 S [CTh]; d 5 Ô S) (all leg. Paragamian); RETHYMNO: 
Site 39 (al S); Site 40 (b 1 3); Site 41 (a 2 S S) (all leg. Nikolakakis); IRAKLEIO: Site 47 (b 
1 S) (leg. Nikolakakis); LASITHI: Site 55 (a 3 99;b4 99) (all leg. Chatzaki); Site 63 (b 1 
8); Site 71 (b 3 S â) (all leg. Stathi). 

Taxonomy. This species is close to Z. labilis. Both species have a similar habi- 
tus (Z. daidalus sp. n. is rather smaller), and the general structure of the epigyne is sim- 
ilar as well. However, Z. daidalus sp. n. can be easily distinguished by the shape of the 
tibial apophysis, the terminal apophysis and the transverse embolus. In Z. daidalus sp. 
n., the median plate of the epigyne is wider than in Z. labilis, the glandular heads are 
in a different position and the distance between posterior rim of median plate and sper- 
mathecae is greater. 

Measurements <? (9), n= 16 (8): TL: 3-3.7 (3.5-6), PL: 1.4-1.6 (1.4-2), PW: 1- 
1.25 (1.1-1.6), OL: 1-1.25 (1.8-3.5), PL/PW: 1.26-1.39 (1.2-1.4). 

â 9 : Habitus as in Zelotes labilis. Dark brown spiders of small size. Prosoma 
equally wide along its whole length. Anterior and posterior rows of eyes straight. 

Legs: Me III-IV with apical preening combs. All Co, Me and Ta yellow, other 
articles brown. All Ta and Me with spiny hairs. Spination: Fe: I-II d 0-2; III-IV d 2-3. 
Pa: I-IV - . Ti: I-II - ; III-IV spinose ventrally. Me: I-II - ; III-IV spinose ventrally. 

â Pedipalp (Figs 91-92, 97): Tibial apophysis relatively short, rounded. 
Terminal apophysis (TA) bulging, rounded, hiding the embolus and the retinaculum, 
seen only from the dorsal view. Embolus short, tranverse, with distinct embolar base 
(EB). 

Epigyne (Fig. 102): Anterior margin wide, almost continuous. Median plate 
wider than long, its posterior margin broadly curved. 

Vulva (Fig. 103): Introductory channels starting with pouches, glandular heads 
attached to posterior angles. Introductory ducts transverse, spermathecae globular, as 



78 



M. CHATZAKI ET AL. 




Figs 93-97 
Zelotes labilis, specimens from lowlands (93- 94), specimens from high altitudes (95-96), Z. 
daidalus sp. n. (97). Palp of S , dorsal view (93-97). H: sclerotized rim, p: triangular process, 
EB: embolar base. Scale lines 0.1mm. 



GROUND SPIDERS OF CRETE. III. 



79 




102 




103 



Figs 9-8-103 
Zelotes labilis, specimens from lowlands (98-99), specimens from high altitudes (100-101). Z. 
daidalus sp. n. (102-103). Epigyne (98, 100, 102), vulva (99, 101, 103). Scale lines 0.1mm. 



80 



M. CHATZAKI ET AL. 



2,5 

Ê 
E 2 



e 

O 1,5 

E 

I 1 
o 

0,5 



♦ ♦ 






♦7£ÌFT: 




ri 



0,5 



1 1,5 2 

Prosoma Length (mm) 



2,5 



Fig. 104 
Scatter diagram of ó" Z. labilis morphometric measurements: cymbium / prosoma length. 

in Z. labilis. Distance between the posterior rim of the median plate and the spermath- 
ecae greater than in Z. labilis. 

Ecology. Z. daidalus sp. n. occurs mainly at the periphery of Crete (Fig. 123), 
probably indicating an immigration from other regions rather than an endogenous ori- 
gin. It is not very common either on Crete or on Gavdos. 

Distribution. Crete, Gavdos. 



Zelotes tenuis (L. Koch, 1 866) 



Figs 105-114, 124 



Identification: Levy (1998, p.131, Figs 78-81), Platnick & Shadab (1983, p.185, Figs. 
259-262 [sub Z. pallidas, see Platnick, 1989:489]). 

Material. CRETE: CHANIA: Site 2 (b 2 S 6 1 9 ); Site 3(alc?;b9ó*oM9;c3ó*c?); 
Site 4 (al 6 \b5 6 6 11 9 9 ); Site 5 (b 25 ó*ó*27 99;c2ó*oM 9); Site 6 (e 5 6 6 1 9;f 
22 6 6 4 9 9; g 16 9 9; h 3 9 9); Site 7 (e 1 6; f 2 9 9) (all leg. Lymberakis); Site 15 (d 2 
<?ó*); Site 16 (e 3 6 6;d2 9 9); Site 17 (e 3 6 6; d2 66 2 9 9); Site 18 (a 2 9 9; e 1 6; d 
2 e? ó*); Site 19 (a 1 9; e 1 6; d 2 6 6); Site 20 (b 1 e?); Site 21 (a 1 9; e 1 Ó*;d6 9 9); Site 
22 (b 3 9 9 ) (all leg. Paragamian); RETHYMNO: Site 25 (a 3 6 6); Site 26 (a 6 6 6 1 9 ; b 16 
6 6 8 9 9; e 1 9) (all leg. Lymberakis); Site 27(a6ó*óM9;b4ó*<?19 99) (all leg. 
Chatzaki);Site28(bl3 6 6 23 9 9); Site 29 (a 3 66\bS 66 6 9 9 ); Site 39 (a 2 66); Site 
40 (b 35 Wl5;c 10 66 46 99; d 3 99); Site 41 (a 1 6; b 5 66 6 9 9) (all leg. 
Nikolakakis); IRAKLEIO: Site 42(f9ó*ó*;g6ó*ó*799;h799 (all leg. Chatzaki); Site 43 
(b2 6 6)\ Site 44 (a 2 Ó*c?;bl5c?ó*699;b3ó*<? [MHNG]; e 2 9 9; e 2 9 9 [MHNG]); 
Site 45 (a 55 Ó*c?899;b6c?<?26 9 9 ); Site 47 (b 14 6 6 7 9 9; e 7 9 9 ); Site 50 (e 8 9 9); 
Site 52 (a 1 6; b 25 6 6 4 9 9; e 2 9 9) Site 54 (b 7 6 6 3 9 9) (all leg. Nikolakakis); Site 46 
(a 34 6 6 20 9 9;b35 9 9 ); Site 49 (a 32 6 6 4 9 9; b 20 6 6 36 9 9; e 5 9 9); Site 50 (e 



GROUND SPIDERS OF CRETE. III. 



81 





Figs 105-110 
Zelotes tenuis, rare form (105-107), common form (108-110). Palp of S , ventral view (105, 
108), dorsal view (106, 109), retrolateral view (107, 110). f: subterminal fringe. Scale lines 
0.1mm. 



82 



M. CHATZAKI ET AL. 






113 




N 114 



FIGS 111-114 
Zelotes tenuis, rare form (111-112), common form (113-114). Epigyne (111, 113), vulva (112, 
114). Scale lines 0.1mm. 

35 SO 6 99;cl S var.; d 17 S S 27 9 9) (all leg. Papadimitrakis); LASITHI: Site 55 (a 2 
SS 1 9;b 1 S 4 9 9); Site 73 (a 5 SS 4 9 9;b 1 9 ) (all leg. Chatzaki); Site 58 (a 11 SS 
[CTh];a3 9 9; b 13 9 9 [CTh]); Site 62 (a 10 SS 8 9 9;b24 9 9); Site 64 (d 10 SS 2 9 9; 
d2 SS 3 9 9 var.; e 3 9 9); Site 65 (a 14 SS 3 9 9; b 2 9 9 ); Site 68 (a 7 SS 6 9 9;b 1 
9) (all leg. Papadimitrakis); Site 60 (a 2 SS); Site 63 (b 6 d <J 2 9 9; c 10 9 9); Site 71 (b 17 
SS 8 9 9;c6 9 9) (all leg. Stathi); Site 61 (a30*cT3 99;b4ddl 9); Site 66 (a 3 SS 1 
9); Site 67 (a 6 SS 2 9 9) Site 69 (a 1 S 3 9 9); Site 70 (b 1 9); Site 72 (a 3 SS 1 9;b7 
c? o* 2 9 9 ) (all leg. Trichas). 



GROUND SPIDERS OF CRETE. III. 



83 




<! 







• Camillina metellus 
200m 
800m 



Fig. 115 

Camillina metellus, distribution on Crete. 



■-■-■: 




-.- ■ -. z •', . -V. --"^'-^P ~,%? 



r Drassyllus pumiloides sp.n. 
• Drassylus praeficus 

200m 

1200m 






Fig. 116 
Drassyllus praeficus, D. pumiloides sp. n., distribution on Crete. 



<r 






'*• » 



© 

© 



© 



©/ 



p Trachyzelotes adriaticus 

* Trachyzelotes barbatus 

• Trachyzelotes lyonnetti Ann 

s Trachyzelotes malkini ouum 

Fig. 117 
Trachyzelotes adriaticus, T. barbatus, T. lyonneti, T. malkini, distribution on Crete. 



84 



M. CHATZAKI ET AL. 




• Setaphis carmeli 
A Zelotes aerosus 

F Zelotes solstitialis 

* Zelotes nilicola 
800m 

FlG. 118 
Setaphis carmeli, Zelotes aerosus, Z. solstitialis, Z. nilicola, distribution on Crete. 




• Zelotes caucasius 

400m 
1000m 



FlG. 119 
Zelotes caucasius, distribution on Crete. 



xf' 



^L 



«. 




Zelotes subterraneus 

1000m 
1800m 



FlG. 120 
Zelotes subterraneus, distribution on Crete. 



GROUND SPIDERS OF CRETE. III. 85 



- • * 

* * 

* 

* Zelotes cf. ilotarum 

• Zelotes creticus 

200m 
800m 

Fig. 121 



Zelotes creticus, Z. cf. ilotarum, distribution on Crete. 






**\ 



# Zelotes minons sp.n. 

• Zelotes scrutants 

200m 
1200m 



Fig. 122 
Zelotes scrutatus, Z. minous sp. n., distribution on Crete. 



4« 



..-. p4 I 



© Zelotes daidalus sp.n. 400m 
-77, ; a;- 1000m 

• Zelotes labilis 2000m 



Fig. 123 
Zelotes labilis, Z. daidalus sp. n., distribution on Crete. 



86 M. CHATZAKI ET AL. 




m Zelotes tenuis 

400m 

Fig. 124 
Zelotes tenuis, distribution on Crete. 

KARPATHOS: Pyles - Volada, 1 km E, phrygana on an earlier burnt field, (14 S 6 6 
9 9, pitfall traps 12/5/2001 - 23/8/2001, leg. Chatzaki). 

KOS: Kefalos - Ag. Ioannis, 1 km S, phrygana and adjacent pine forest, (21 cT 6 34 9 9 , 
pitfall traps 26/6/2001 - 9/9/2001, leg. Chatzaki). 

Taxonomy. Measurements £(9): n = 14 (16): TL: 3.4-7.2 (3.5-9.6), PL: 1.6-3 
(1.5-3.4), PW: 1.2-2.5 (1.1-2.4), OL: 1.6-4.1 (1.6-5.7), PL/PW: 1.17-1.52 (1.19-1.47). 

a 9 : Medium sized spiders of reddish brown colour. Posterior row of eyes 
straight or slightly recurved, anterior row straight or slightly procurved. PME round or 
oval. 

Legs: Me III-IV with apical preening combs. Ta I-II with scopula hairs, III-IV 
with spiny hairs. Spination: Fe: I-II d 3; III-IV d 5-6. Pa: I-II, IV - ; III r 1. Ti: I - ; II 
v 0-1; III-IV spinose. Me: I v 0-2; II v 3; III-IV spinose. 

6 Pedipalp (Figs 105-110): Tibial apophysis almost straight. Cymbium oval. 
Embolar base broad, transverse, its end sharply turning to the dorsal side. Embolus 
short, dorsal, often with a subterminal fringe (f). 

Epigyne (Fig. Ill, 113): Anterior margin wide, continuous. Median plate di- 
vided posteriorly by a median septum, introductory orifices situated at its lateral sides. 

Vulva (Fig. 112, 114): Introductory channels starting with pouches, well-sepa- 
rated from each other, then leading as narrow, parallel ducts to the spermathecae. 
Glandular heads posterior. 

Comments. In the material collected from Crete, two forms of this species are 
present; a rare one (males found at Sites 50 and 68, Figs 105-107 and females found at 
Sites 2, 4, 5, 6, 7, 26, 27, 52, 54, Figs 1 1 1-1 12), and a very common one (Figs 108-1 10 
and 1 13-114). In males, the two types differ in the shape of the embolar base (compare 
Figs 105 with 108 and 107 with 110) and in details in the embolic division (compare 
Figs 106 with 109). In females, differences are seen in the width of the median septum 
of the epigyne (compare Figs 111 with 113) and in the introductory channels (com- 
pare Figs 112 with 1 14). As males and females of the two types are mixed at the same 
localities, we cannot suggest that they belong to different species. Therefore, we place 
them into one species, accepting that the differences presented in male and female 
genitalia reflect intraspecific variation. 



GROUND SPIDERS OF CRETE. III. 87 

Ecology. Z. tenuis is very common on Crete, Gavdos and Dia, as well as on 
Karpathos and Kos. It is widespread in the lowlands, but does not occur above 1000m. 
Two peaks of activity were observed in males, one in late spring and one in early 
autumn; in females one in mid-summer. 

Distribution. Mediterranean: Spain, France, Corsica, Italy, Croatia, Egypt, 
GREECE: Crete: Akrotiri, Kalathas Pond (Platnick & Shadab, 1983); Karpathos; Kos, 
introduced to the USA (Platnick & Shadab, 1983). 

CONCLUDING REMARKS 

In this paper we revise the gnaphosid zelotine group of Crete and adjacent areas. 
In total, 20 species are reported, belonging to five genera {Camillina, Drassyllus, 
Trachyzelotes, Setaphis and Zelotes). Three species are new to science (Drassyllus 
pumiloides sp. n., Zelotes daidalus sp. n., and Z. minous sp. n.). Camillina metellus is 
a new combination for Zelotes metellus Roewer, 1928 and the first record of Camillina 
for Greece. Trachyzelotes stubbsi Platnick & Murphy, 1984 and Zelotes bucharensis 
Charitonov, 1946 are proposed as new synonyms of T. adriaticus (Caporiacco, 1953) 
and Z. scrutatus (O. P. Cambridge, 1872), respectively. Two species are recorded for 
the first time from Europe (Z. aerosus and Z. solstitialis), four are recorded for the first 
time from Greece (Drassyllus praeficus, Trachyzelotes adriaticus, Setaphis carmeli 
and Z. subterraneus) and two are recorded for the first time from Crete (Z. labilis and 
Z. nilicola). 

Most of the species examined have a Mediterranean distribution (T. barbatus, T. 
lyonneti, S. carmeli, Z. labilis, Z. subterraneus, Z. nilicola) or an East Mediterranean 
distribution (Z. solstitialis), whilst few have a wider distribution (D. praeficus, T 

malkini, Z. aerosus, Z. scrutatus). Apart from the newly described species, another 
three are still considered endemic to Greece (C. metellus) or endemic to Crete (Z. creti- 
cus and Z. cf. ilotarum). However, until a more thorough study of the East 
Mediterranean region is carried out, knowledge of the true distributions of the zelotine 
species cannot be considered to be precise. For example, a more detailed analysis and 
further material are needed to clarify the taxonomic status of the dubious species Z. 
ilotarum and Z. labilis on the Balkan Peninsula, the Greek islands and in Asia Minor. 

Many zelotines are common on Crete, show non-specific ecological preferences 
and are abundant in most types of lowland habitats. Z. caucasius, Z. tenuis, Z. labilis, 
Z. scrutatus and T. malkini are among the commonest and most abundant spiders on 
the island. However, some species are more localized, i.e. are present either in the west- 
ern part (T barbatus, T adriaticus, Z. creticus) or in the eastern part of Crete (D. praefi- 
cus, T. lyonneti, Z. cf. ilotarum). This may be due to either competitive interaction be- 
tween congeners (as mentioned in the case of the Trachyzelotes group) or to different 
limits of tolerance to dryness, which probably makes the less tolerant species avoid the 
driest parts in the east of the island. The greater abundance of D. praeficus and Z. creti- 
cus recorded in the higher altitudes of Crete, indicate some preference of these species 
for mountainous regions, at least in the Mediterranean. 

There are 1 1 and 6 zelotines recorded on Gavdos and Dia, respectively. These 
species belong to the commonest Cretan species of the group. Exceptions are C. 



88 M. CHATZAKI ETAL. 

meîellus and the new species Z. minous sp. n. and Z. daidalus sp. n. (only on Gavdos), 
which occur on these islands, but are not very common on Crete. 

Most of the species reported here present similar phenologies, being active from 
spring to autumn. Males have two peaks of activity, one in late spring and one in early 
autumn and the females have one peak in late spring or in the mid-summer. The only 
exception is Z. subterraneus, which is more active during the autumn and the winter 
months. 

ACKNOWLEDGEMENTS 

We are very grateful to M. Nikolakakis, M. Papadimitrakis and S. Roberts, who 
collected and sorted most of the material presented in this paper, and to J. Murphy for 
linguistic revision of the text. We also thank Dr G. Anastasiou for loan of important 
comparative material from Peloponnisos, Prof. A. Legakis of the University of Athens, 
for loan of the material from Hadjissarantos' collection, Dr M. Grasshoff for loan of 
specimens from Roewer's collection and Dr T. Blick and Dr G. Levy for loan of further 
material, Dr J. Gruber for providing important literature and Dr B. Knoflach for advice 
on the drawings. Financial support has been given by the Biology Department of the 
University of Crete, the Natural History Museum of Crete, the University of Innsbruck 
and the Onassis Foundation. 

REFERENCES 

Bonnet, P. 1956. Bibliografia araneorum. Toulouse, 2 (2): 919-1926. 

Buchar, J. 1992. Kommentierte Artenliste der Spinnen Böhmens (Araneida). Acta Universitatis 

Carolinae Biologica 36: 383-428. 
Charitonov, E. 1946. [Neue Formen aus der Spinnenfauna der SSSR]. [Mitteilungen des natur- 
wissenschaftliches Instituts der staatlichen Molotov Universität M. Gorkij] Novye formy 

paukov fauny SSSR. Izv. est.-nauchn. Inst. Molotovsk. Univ. 12: 19-32 (in Russian and 

Latin). 
Chatzaki, M. 1998. Systematics and phenology of ground living spiders of the island group 

Gavdos - Gavdopoula. Master Thesis, Univerity of Crete, Irakleio, 150 pp. (in Greek). 
Chatzaki, M., Trichas, A., Markakis, G. & Mylonas, M. 1998. Seasonal activity of the 

ground spider fauna in a Mediterranean ecosystem (Mt. Youchtas, Crete, Greece). 

Proceedings of the 1 7th European Colloquium of Arachnologw Edinburgh 1997: 

235-243. 
Chatzaki, M., Thaler, K. & Mylonas, M. (2002a). Ground spiders (Gnaphosidae; Araneae) of 

Crete (Greece). Taxonomy and distribution. I. Revue suisse de Zoologie 109(3): 559-601. 
Chatzaki, M., Thaler, K. & Mylonas, M. (2002b). Ground spiders (Gnaphosidae, Araneae) of 

Crete and adjacent areas of Greece. Taxonomy and distribution. II. Revue suisse de 

Zoologie 109(3): 603-633. 
Deltshev, C. & Blagoev, G. 2001. A critical check list of Bulgarian spiders (Araneae). Bulletin 

of the British Arachnological Society 12: 1 10-138. 
Di Franco. F 1997. New considerations about the gnaphosid fauna of Italy (Araneae: 

Gnaphosidae). Bulletin of the British arachnological Society 10: 242-246. 
Giltay, L. 1932. Arachnides recueillis par M. d'Orchymont au cours de ses voyages aux Balkans 

et en Asie Mineure en 1929, 1930 et 1931. Bulletin du Musée royal d'Histoire naturelle 

de Belgique 8 (22): 1-40. 
Grimm, U. 1985. Die Gnaphosidae Mitteleuropas (Arachnida, Araneae). Abhandlungen des 

Natwwissenschaftlichen Vereins in Hamburg (NE) 26: 1-317. 



GROUND SPIDERS OF CRETE. III. 89 



Hadjissarantos, H. 1940. The spiders of Attiki. Thesis, University of Athens, Athens, 132 pp. 

(in Greek). 
Kulczynski, V. 1903. Aranearum et Opilionum species in insula Creta a Comité Dre Carolo 

Attems collectae. Bulletin de l' Académie des Sciences de Cracovie 1903: 32-58. 
Levy, G. 1998. The ground-spider genera Setaphis, Trachyzelotes, Zelotes, and Drassyllus 

(Araneae:Gnaphosidae) in Israel. Israel Journal of Zoology 44: 93-158. 
Pesarini, C. 1994. Arachnida Araneae. In: Minelli A., Ruffo, S. & La Posta, S. (eds). Checklist 

delle specie della fauna italiana, 23: 1-42. Calderini, Bologna. 
Platnick, N. I. 1989. Advances in spider taxonomy 1981-1987. Manchester University Press, 

Manchester & New York, 673 pp. 
Platnick, N. I. & Shadab, M. U. 1982a. A revision of the American spiders of the genus 

Camillina (Araneae, Gnaphosidae). American Museum Novitates 2748: 1-38. 
Platnick, N. I. & Shadab, M. U. 1982b. A revision of the American spiders of the genus 

Drassyllus (Araneae, Gnaphosidae). Bulletin of the American Museum of Natural 

History 173: 1-97. 
Platnick, N. I. & Shadab, M. U. 1983. A revision of the American spiders of the genus Zelotes 

(Araneae, Gnaphosidae). Bulletin of the American Museum of Natural Histoiy 174: 

99-191. 
Platnick, N. I. & Murphy, J. A. 1984. A revision of the spider genera Trachyzelotes and 

Urozelotes (Araneae, Gnaphosidae). American Museum Novitates 2792: 1-30. 
Platnick, N. I. & Murphy, J. A. 1987. Studies on Malagasy spiders, 3. The zelotine Gnapho- 
sidae (Araneae, Gnaphosoidea), with a review of the genus Camillina. American 

Museum Novitates 2874: 1-33. 
Platnick, N. I. & Murphy, J. A. 1996. A review of the zelotine ground spider genus Setaphis 

(Araneae, Gnaphosidae). American Museum Novitates 3162: 1-23. 
Platnick, N. I. 2001. The World Spider Catalog, Version 2.0: 

http://research.amnh. org./entomology 
Roewer, C. F. 1928. XI. Araneae. In: Zoologische Streifzüge in Attika, Morea und besonders auf 

der Insel Kreta. Abhandlungen des naturwissenschaftlichen Vereins Bremen 27: 92-123. 
Roewer, C. F., 1954. Katalog der Araneae. Bremen, 2(1), 923 pp. 
Simon, E. 1884. Matériaux pour servir à la faune des Arachnides de la Grèce. Annales de la 

Société entomologique de France (6) 4: 305-356. 
Simon, E. 1914. Les Arachnides de France. Roret, Paris, 6/1, 308 pp. 
Tullgren, A. 1910. Araneae. In: Wissenschaftliche Ergebnisse der Schwedischen Zoologischen 

Expedition nach dem Kilimandjaro, dem Meru und dem Umgebenden Massaisteppen 

Deutsch-Ostafricas 1905-1906 unter Leitung von Prof. Dr. Yngve Sjöstedt, 20: 85-172, 

pis. 1-4. 
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du Muséum d' Histoire naturelle „Grigore Antipa" 16: 79-107. 



Revue suisse de Zoologie 1 10 (1): 91-109; mars 2003 



A review of the Chinese jumping spiders studied by Dr E. 
Schenkel (Araneae: Salticidae) 

Xian-Jin PENG 1 , Shu-Qiang LH* & Christine ROLLARD2 

1 Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, China. 

2 Muséum National d'Histoire Naturelle, Laboratoire de Zoologie (Arthropodes), 
F-75005 Paris, France. 



A review of the Chinese jumping spiders studied by Dr E. Schenkel 
(Araneae: Salticidae). - 41 species of Chinese jumping spiders described 
by Dr E. Schenkel as new species from 1936 to 1963 are reviewed in part 
and listed in the present paper. 13 of these species are still valid in their 
original combination, 17 were placed in synonymy, 9 were before trans- 
ferred, and Menemerus yunnanensis Schenkel, 1963 is here transferred to 
Pseudicius. Illustrations of the body of Asianellus potanini (Schenkel, 
1963), Heliophanus potanini (Schenkel, 1963) and Talavera trivittata 
(Schenkel, 1963) and of diagnostic structures of Pseudicius yunnanensis 
(Schenkel, 1963), Thiania cavaleriei Schenkel, 1963 and Thiania luteo- 
brachialis Schenkel, 1963 are given. 

Key-words: Araneae - Salticidae - taxonomy - China - Schenkel. 

INTRODUCTION 

In recent years, due to large-scale habitat destruction and fragmentation, almost 
all ecosystems in China have experienced severe disturbances. Only a few isolated 
areas still posses their original flora and fauna. Because of this rapid habitat destruc- 
tion, many species had disappeared long before they could be collected and described. 
One of the ways to retrieve information on species that are currently difficult to find 
but were once quite abundant is to examine specimens collected when their natural 
habitats were more intact. 

Most early studies on Chinese jumping spiders were made by arachnologists 
from overseas. Among of them, Dr Ehrenfried Schenkel, former curator of the Natural 
History Museum of Basel, contributed most publications (Schenkel, 1936, 1938a, 
1938b, 1953, 1963), but illustrations and other information in these papers are some- 
times not sufficient for correct identification. Recently, we checked some salticid 
specimens studied by E. Schenkel and deposited at the Muséum d'Histoire Naturelle, 
Paris. Dr Ambros Haenggi (NMB) provided information on Chinese salticid specimens 
studied by E. Schenkel and deposited at the Naturhistorisches Museum Basel. Based 
on the results of our work along with work documented by other colleagues, 41 species 



; All correspondence should be addressed to Shu-Qiang Li, e-mail: lisq@panda.ioz.ac.cn 
Manuscript accepted 01.07.2002 



92 X.-J. PENG ETAL. 

of Chinese jumping spiders described as new species by E. Schenkel are reviewed and 
listed in the present paper. 13 of these species remain in their original combination, 17 
are junior synonyms, 9 were transferred to other and Menemerus yunnanensis 
Schenkel, 1963 is here transferred to Pseudicius. 

MATERIAL AND METHODS 

The materials used for this study are deposited in the following institutions and 
universities (abbreviations are used in the text): 

HNU Hunan Normal University, Changsha, China 

IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China 

JLU Jilin University, Changchun, China 

MHNG Muséum d'histoire naturelle, Genève, Switzerland 

MNHNP Muséum National d'Histoire Naturelle, Paris, France 

NMB Naturhistorisches Museum Basel 

For each species, only the following synonyms and references are given in the 
text: References to Schenkel's papers; References to révélant papers by Chinese 
colleagues, and synonyms not listed in Platnick's spider catalogue (Platnick, 2002). 

Updated information on the distribution in China of each species is provided at 
the provincal level. The names of localities and distribution data are given according to 
current Chinese standard, as shown in Map 1 . 

Other abbreviations used in the text: D = Description, T = Transferred. 

TAXONOMY 

1. Asianellus festivus (C. L. Koch, 1834) 

Aelwillus festivus (C. L. Koch): Zhu & Shi, 1983: 199, figs 180a-e (Ó* 9); Chen & Zhang, 1991: 
318, fig. 338(9). 

Phlegra pichoni Schenkel, 1963: 438, figs 251a-b (8); Yin & Wang, 1979: 36, figs 20A-E {S 9 ); 
Hu, 1984: 383, figs 398.1-3 (<J 9); Chen & Gao, 1990: 191, figs 244a-b ( S 9 , misiden- 
tified). 

Phlegra festiva (C. L. Koch): Harm, 1977: 69, figs 2a-b, 10-12, 14a-b (To* 9 from Aelwillus, re- 
jected); Weiss, 1979: 246, figs 21-25 (d 9); Zhang, 1987: 249, figs 221.1-4 (S 9); Peng 
et al., 1993: 168, figs 590-597 (Ó* 9); Song, Chen & Zhu, 1997: 1739, figs 51a-b (9). 

Asianellus festivus (C. L. Koch): Logunov & Heciak, 1996: 106, figs 1-5, 8, 10, 17-19, 23-28, 
39 (T<? 9 from Phlegra); Song, Zhu & Chen, 1999: 505, figs 288M-0, 289B-C (<J 9). 

Material examined: Id (holotype of Phlegra pichoni Schenkel, 1963), Hangzhou City, 
Zhejiang Province, 1925 (MNHNP); 6ó*ó*5 9 9, Laishui District, Hebei Province, 12.- 
13.V.2001, leg. X. J. Peng (MHNG); lo*, Zhangjiajie City, Hunan Province, 25.VIII.1984, leg. 
Y J. Zhang (HNU); 1 9, Mt. Yuelu, Changsha City, Hunan Province, 24. VI. 1986, leg. L. P. Xie 
(HNU); 1 9, Fuping District, Shaanxi Province, 25.Vn.1998, leg. J. Chen (IZCAS); 1 9, Bikou 
Township, Wenxin District, Gansu Province, 25. VI. 1998, leg. J. Chen (IZCAS). 

Distribution. China (Anhui, Beijing, Gansu, Guangxi, Guizhou, Hebei, 

Heilongjiang, Hubei, Hunan, Jilin, Shaanxi, Shandong, Shanxi, Sichuan, Zhejiang), 

Palearctic Region. 

2. Asianellus potanini (Schenkel, 1963) Fig. 1 

Phlegra potanini Schenkel, 1963: 436, figs 250a-b (6); Wesolowska, 1981b: 153, figs 72-76 
(<?); Proszynski, 1990: 283 (To* 9 from Aelwillus). 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 



93 



\A 



r J 



Xinjiang 



x.A 



- ^^ 




y-v (Heilongjiang/ 

! ^W JHin/^ 

. ^rUtaonin^ 
BEIJING ^/ 



v.. 



Tibet 



7 v-> Inner Mongolia a, 

~ x -Ç a D5H\ . ,~, Ä ^ 1 fffiarfpn 

V Z, ft ./c Hebel ^r-- 7 

•-X <A|ingxtd ) / y P 3 

Qinghai 1 1 ^ Shan$ Shandong 

:■ iL sÄanx^^^V^ 95 " 



\! Sichuan "5 H ubeì^ hui ^ Shanghaì 
C ^u •-"' ^.-^^-^"^"^Zheifeng 
;A ,t;,'. î cnorjcjqjng'sj Jjangxi-t^.j ' 

" 1 "V .-'f' J:^-. JHunans / / 
J \ y .. Guizhoü(, ,-, > Fùjianl ^ 



£ Yunnan | 



GuangxP , V" f /Taiwan 

^c, Guangdong V 

**hong Kong 



Hainanf^ / 



Map 1. Political map of China (source: http://nfgis.nsdi.gov.cn). 



Asianellus potanini (Schenkel): Logunov & Heciak, 1996: 113, figs 7, 12, 31-32, 36, 49-56 
(To* 9 from Phlegm); Song, Zhu & Chen, 1999: 506, figs 288P-Q, 289D (<J ?). 

Material examined: 1 o* (holotype of Phlegm potanini), collection data missing (accord- 
ing to Schenkel, 1963: possibly in "Kloster Dschoni, Kansu", 24 V. ...label missing) (MNHNP). 
Distribution. Kazakhstan to China (Gansu). 

3. Bianor hotingchiehi Schenkel, 1963 

Bianor hotingchiehi Schenkel, 1963: 434, figs 249a-f (o*); Yin & Wang, 1979: 27, figs 1A-E 
(o* 9); Yin & Wang, 1981: 268, figs 1A-H (S 9); Hu, 1984: 354, figs 368.1-5 (<? 9); 
Song, 1987: 286, fig. 243 (<J 9); Feng, 1990: 198, figs 173.1-6 {6 9); Chen & Gao, 
1990: 180, figs 229a-c (<J 9); Chen & Zhang, 1991: 288, figs 301.1-5, 302.1-6 (Ó* 9); 
Song, Zhu & Li, 1993: 883, figs 58A-D S 9); Peng et ai, 1993: 26, figs 34-42 (Ó* 9); 
Zhao, 1993: 391, figs 195a-c (Ó* 9); Song, Zhu & Chen, 1999: 506, figs 289J, 290A, 
324M(o*9). 

Material examined: 1 o*2 9 9 , Xiangyin District, Hunan Province, XL 1980, leg. Z. Q. Hu 
(HNU); 16, Mt. Longqi, Jiangle District, Fujian Province, 19.VIII.1991 (MHNG); 19, Mt. 
Tianmu, Zhejiang Province, 5.IX.1989 (MHNG); lo\ Cili District, Hunan Province, VII.1986, 
leg. X. J. Peng (HNU); 1 9 , Xiangyin District, Hunan Province, X.1986, leg. X. J. Peng (HNU). 



94 X.-J. PENG ET AL. 

Distribution. China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Henan, 
Hubei, Hunan, Jiangsu, Jiangxi, Shandong, Sichuan, Yunnan, Zhejiang), Bangladesh, 
Myanmar, Vietnam. 

4. Carrhotus xanthogramma (Latreille, 1819) 

Carrhotus xanthogramma (Latreille): Lessert, 1910: 592; Prószyfiski, 1973b: 100, figs 8-14 
(3 9); Guo, 1985: 175, figs 2-99.1-4 (3 9); Song, 1987: 287, figs 244 (3 9); Zhang, 
1987: 236, figs 207.1-4 (0*9); Chen & Gao, 1990: 181, figs 230a-c (0*9); Chen & 
Zhang, 1991: 295, figs 310.1-3 (3 9); Peng et al., 1993: 38, figs 84-91 (3 9); Zhao, 
1993: 392, figs 196a-c (c? 9); Song, Zhu & Chen, 1999: 507, figs 290K, 291C (Ó* 9). 

Carrhotus pichoni Schenkel, 1963: 444, figs 254a-c (3); Yin & Wang, 1979: 28, figs 3A-E 
(3 9); Yin & Wang, 1981: 269, figs 2A-F (3 9); Hu, 1984: 356, figs 370.1-4 (Ó* 9). 

Carrhotus bicolor (Walckenaer): Peng, 1989: 158, figs 2A-F (Ó* 9). 

Material examined: IS , (paratype of Carrhotus pichoni), Zhejiang Province (on locality 
label: Hangtscheu Tschekiang), leg. Dr Pichon, 1925 (NMB2162a); lo" 19, Dadou District, 
Guangxi Zhuang Autonomous Region, alt. 280m, 29.III.1988, leg. M. Wu (MHNG); 1 9, Daxin 
District, Guangxi Zhuang Autonomous Region, alt. 280m, 30.III.1988, leg. M. Wu (MHNG); 
1 9, Nanping, Shangsi District, Guangxi Zhuang Autonomous Region, alt. 350m, 11. VI. 2000, 
leg. J. Yao (MHNG); 1 3 , Xialei Town, Daxin District, Guangxi Zhuang Autonomous Region, 
alt. 320m, 31.III.1998, leg. M. Wu (MHNG); 19, Dongzhong Town, Fangchenggang City, 
Guangxi Zhuang Autonomous Region, alt. 270m, 5. IV 1998, leg. J. Chen (MHNG); 1 9, Yaogou, 
Fuping District, Shaanxi Provinces, alt. 870-1000m, 25.VII. 1998, J. Chen (IZCAS). 

Distribution. China (Fujian, Guizhou, Guangdong, Guangxi, Hunan, Hubei, 
Jilin, Liaoning, Shaanxi, Shandong, Sichuan, Taiwan, Zhejiang), Palearctic Region. 

5. Davidina magnidens (Schenkel, 1963) 
Davidia magnidens Schenkel, 1963: 465, figs 263a-f (9). 

No specimens examined. 
Distribution. China (Zhejiang). 

Remarks: This species was described by Schenkel (1963) from an immature 
specimen, which could not be found in the MNHNP. 

6. Evarcha albana (L. Koch, 1878) 

Evarcha albaria (L. Koch): Simon, 1903: 697, figs 837 (Ó*); Schenkel, 1963: 459, figs 260a-c 
(cT 9); Yin & Wang, 1979: 30, figs 6A-C (3 9); Wang, 1981: 137, figs 77A-C (<J 9); Hu, 
1984: 361, figs 376.1-4 (3 9); Guo, 1985: 176, figs 2-100.1-4 (3 9); Zhu & Shi, 1983: 
212, figs 184a-d (cî 9); Song, 1987: 290, figs 246 (Ó* 9); Zhang, 1987: 238, figs 209.1- 
4 (3 9); Feng, 1990: 199, figs 174.1-6 (3 9); Chen & Gao, 1990: 183, figs 232a-b 
(3 9); Chen & Zhang, 1991: 312, figs 331.1-4 (Ó* 9); Song, Zhu & Li, 1993: 884, figs 
59A-D (3 9); Peng et al., 1993: 61, figs 159-165 (3 9); Zhao, 1993: 395, figs 198a-c 
(3 9); Song, Chen & Zhu, 1997: 1734. figs 44a-d (3 9); Song, Zhu & Chen, 1999: 509, 
figs292P-Q, 294B-C(o*9). 

Evarcha pichoni Schenkel, 1963: 459, figs 261a-b (3). 

Material examined: \3 (holotype of Evarcha pichoni Schenkel, 1963), Hangzhou City, 
Zhejiang Province, 1925 (MNHNP); lo*59 9, Miaotaizi Town, Liuba District, Shaanxi 
Province, alt. 1350-1730m, 21VII.1998, leg. J. Chen (MHNG); 19, Yaogou, Fuping District, 
Shaanxi Provinces, alt. 870-1000m, 25VII.1998 (MHNG); 19, Kangxian District, Gansu 
Province, alt. 1400-1680m, 15VII.1998, leg. J. Chen (MHNG); 23 3, Bifenggou, Bikou Town, 
Wenxian District, Gansu Province, alt. 900-1500m, 25VI.1998, leg. J. Chen (MHNG); lo\ 
Kangxian District, Gansu Province, alt. 1470-1610m, 13.VTL1998, leg. H. J. Wang (MHNG); 
Id 19, Mt. Baiyun. Kangxian District, Gansu Province, alt. 1250-1750m, 12VII.1998, leg. J. 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 95 



Chen (MHNG); 19, Mt. Baiyun, Kangxian District, Gansu Province, alt. 1300-1750m, 
12.VII.1998, leg. J. Chen (MHNG); 12c? S 16 9 9, Cili District, Hunan Province, 27.VII.1986, 
leg. X. J. Peng (HNU); 1 9 , Hangzhou City, Zhejiang Province (on locality label: Hangtscheu 
Tschekiang), leg. Dr Pichon, 1925 (NMB2162a). 

Distribution. China (Anhui, Gansu, Hubei, Hebei, Henan, Hunan, Fujian, 
Guangdong, Guangxi, Jiangsu, Jilin, Liaoning, Shaanxi, Shandong, Shanxi, Sichuan, 
Xinjiang, Yunnan, Zhejiang), Russia, Korea, Japan. 

7. Hasarina contortospinosa Schenkel, 1963 

Hasarina contortospinosa Schenkel, 1963: 462, figs 262a-g (3); Wesolowska, 1981b: 132, figs 
10-13 (3); Xiao, 1991: 383, figs 1-4 (9); Peng et al, 1993: 83, figs 254-263 (3 9); 
Song, Zhu & Chen, 1999: 513, figs 299F, M (3 9). 

Material examined: 1 9 , Xuanen District, Hubei Province, 24.V1989 (MHNG). 

Distribution. China (Fujian, Gansu, Hubei, Hunan, Sichuan). 

8. Heliophanus curvidens (O. P.-Cambridge, 1872) 

Heliophanus berlandi Schenkel, 1963: 399, figs 229a-c (o"9; preoccupied by H. berlandi 
Lawrence, 1937); Prószynski & Zochowska, 1981: 18, figs 7-11 (3 2); Song, Zhu & 
Chen, 1999: 513, figs 299P-R, 300B-C (3 9). 

Heliophanus curvidens (O. P.-Cambridge): Simon, 1876: 165; Song, Zhu & Chen, 1999: 514, 
figs300D-E,J-L(c?9). 

Material examined: 1 3 (holotype of Heliophanus berlandi), collection data missing (ac- 
cording to Schenkel, 1963: "Lau-wa-sja (Louwacheng) am Siningho", Gansu, 19.IV 1885) 
(MNHNP); 1 3 (paratype of Heliophanus berlandi), Gansu Province (locality given on label: 
"Chautschuan a. Siningho"), leg. Potanin, 22.IV1885 (NMB2176a). 

Distribution. Israel to China (Gansu, Xinjiang). 

9. Heliophanus potanini Schenkel, 1963 Figs 2-3 

Heliophanus potanini Schenkel, 1963: 397, figs 228a-b {3 9); Zhou & Song, 1988: 3, figs 4a-e 
(3 9); Hu & Wu, 1989: 373, figs 282.4-7 (3 9); Song, Zhu & Chen, 1999: 514, figs 
300R-S, 301A(c?9). 

Menemerus fagei Schenkel, 1963: 422, figs 242a-f (9). 

Material examined: 1 9 (holotype of Menemerus fagei, epigynum missing), collection 
data missing (according to Schenkel, 1963: "Zwischen Tschintasy und Suwanko, Su, Suchow", 
1.-5. VI. 1886) (MNHNP); 1 9 (paratype of Heliophanus potanini), Inner Mongolia (locality giv- 
en on label: "Grenze Chara-Sucha, Etsin-Gol"), leg. Dr Potanin 23.-29.VII. 1886 (NMB2180a); 
23 619, Bohu District, Xinjiang Uygur Autonomous Region, 20.V1985, leg. N.L. Zhou (IZ- 
CAS). 

Distribution. China (Inner Mongolia, Jiangsu, Xinjiang, Zhejiang), Afghanistan, 
Central Asia, Mongolia. 

10. Mendoza canestrinii (Ninni, 1868) 

Mithion pichoni Schenkel, 1963: 414, figs 238a-b (3). 

Mithion tschekiangensis Schenkel, 1963: 418, figs 240a-f (9). 

Marpissa obscura Kroneberg, 1875: 46, pi. 5, fig. 33 (3 9); Song, Zhu & Chen, 1999: 533, figs 

302P, 303G(c?9). 
Marpissa magister Prószynski, 1973b: 116; 1976: 155, fig. 251, 258 (T3 9 from Mithion = 

Thyene); Yin & Wang, 1979: 33, figs 14A-D (3 9); Song, 1980: 205, figs 114a-f (3 9); 

Hu, 1984: 372, figs 389.1-4 (Ó* 9); Guo, 1985: 180, figs 2-103.1-3 (3 9); Zhu & Shi, 

1983: 205, figs 187a-e (3 9); Song, 1987: 296, figs 252 (3 9); Zhang, 1987: 241, figs 



96 X.-J. PENG ETAL. 



213.1-4 (S 2); Feng, 1990: 208, figs 183.1-6 (3 9); Chen & Gao, 1990: 185, figs 235a- 
b (S 2); Chen & Zhang, 1991: 307, figs 326.1-5 (3 9); Peng et ai, 1993: 118, figs 394- 
402 (6 9); Zhao, 1993: 405, figs 206a-c (3 9); Song, Zhu & Chen, 1999: 533, figs 
302N, 303E(c?2). 
Mendoza canestrinii (Ninni): Logunov, 1999: 49, figs 14-15, 24-25, 29, 37, 39, 44, 78-79, 98- 
104, 107-108, 114, 119, 123-124, 131-132. 

Material examined: 1 9 (holotype of Mithion tschekiangensis), Zhejiang Province 
(locality given on label: W. Tschekiang) (MNHNP); 1 3 (paratype of Mithion pienoni), 
Hangzhou City, Zhejiang Province (locality on label: Hangtscheu, Tschekiang), leg. A. Pichon, 
1925 (NMB2194a); 23 3, Xinguo District, Jiangxi Province, VIII.1979, leg. C. M. Yin (HNU); 
73 3599, Guizhou Province, 1979, leg. J. Li (HNU); 433299, Chongan District, Fujian 
Province, VII. 1986, leg. X. Q. Xiao (HNU); 43 3, Suining District, Hunan Province, VIII. 1984, 
M. Y. Liu (HNU); 13, Yangjiang District, Guangdong Province, 16. V. 1976, leg. W. N. Wu 
(MHNG); 13, Leishan District, Guizhuou Province, 27.VI. 1988 (IZCAS). 

Distribution. China (Anhui, Beijing, Fujian, Guangdong, Guangxi, Guizhou, 
Hebei, Henan, Hunan, Hubei, Jiangsu, Jilin, Shaanxi, Shandong, Shanxi, Sichuan, 
Zhejiang), North Africa, Palearctic Region. 

11. Mendoza elongata (Karsch, 1879) 

Mithion hotingchiehi Schenkel, 1963: 416, figs 239a-e (3). 

Marpissa elongata (Karsch): Prószyhski, 1973b: 1 14, figs 47-50 (3); Yin & Wang, 1979: 34, figs 

15A-B(c?); Hu, 1984: 372, figs 388.1-8 (Ó* 9); Guo, 1985: 178, figs 2-102.1-4 (3 9); 

Feng, 1990: 207, figs 182.1-4 (3 9); Chen &Zhang, 1991: 306, figs 325.1-3 (3 9); Peng 

et al., 1993: 116, figs 387-393 (Ó* 9); Zhao, 1993: 403, figs 205a-c (3 9); Song, Zhu & 

Chen, 1999: 533, figs 302M, 303D, 327C-D (3 9). 
Mendoza elongata (Karsch): Logunov, 1999: 53, figs 45, 122 (To* 2 from Marpissa). 

Material examined: Id, Bikou town, Wenxian District, Gansu Province, alt. 7000m, 
24.VI. 1998, leg. J. Chen (MHNG); 13, Mt. Baiyun, Kangxian District, Gansu Province, alt. 
1300-1750m, 12.VII.1998, leg. J. Chen (MHNG); 2ó*ó*32 2, Sangang, Chongan District, 
Fujian Province, VII. 1986, leg. X. J. Peng (HNU); 1 2 , Shangping Town, Hefeng District, Hubei 
Province, 30. V 1989 (IZCAS). 

Distribution. China (Beijing, Fujian, Gansu, Guizhou, Heilongjiang, Hubei, 
Hunan, Jiangsu, Shaanxi, Shanxi, Sichuan, Taiwan, Zhejiang), Russia, Korea, Japan. 

12. Menemerus bivittatus (Dufour, 1831) 

Menemerus bonneti Schenkel, 1963: 430, figs 248a-e (3). Peng et al., 1993: 125, figs 418-421 

(3); Song, Zhu & Chen, 1999: 534, fig. 303K (Ó*). 
Menemerus bivittatus (Dufour): Peckham & Peckham, 1886: 292; Zabka, 1985: 240, figs 283- 

292 (0*2); Peng et al., 1998: 37, figs 1-3(2). 

Material examined: 1 3 (holotype of Menemerus bonneti Schenkel, 1963), collection da- 
ta missing (according to Schenkel, 1963: Vinhi, 7.III.1925) (MNHNP); 12, Goulufeng, 
Hengyang City, Hunan Province, 2.VIII.1997, leg. X. J. Peng (HNU); 1 3, Haikou City, Hainan 
Province, X.1980, leg. Z. Q. Feng (HNU); 23 3, Mengla District, Yunnan Province, VII.1983, 
leg. J. F. Wang (HNU); 1 3 1 9 , Tianchi, Jiangfeng Rain Forest, Hainan Province, XII. 1989 (MH- 
NG); 5o* o\ Jiangfeng Rain Forest, Hainan Province, 20.V1990 (IZCAS). 

Distribution. China (Guangdong, Hainan, Hunan. Yunan), pantropical distri- 
bution. 

13. Menemerus fulvus (L. Koch, 1878) 

Menemerus sinensis Schenkel, 1963: 427, figs 246a-b (2). 
Menemerus schensiensis Schenkel. 1963: 429, figs 247a-d (2). 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 97 



Menemerus confusus Bösenberg & Strand, 1906: 350, pi. 9, fig. 146, pi. 14, fig. 370 (6 2); Yin 
& Wang, 1979: 35, figs 17A-D (<? 2); Hu, 1984: 376, figs 392.1-2 (6 9); Guo, 1985: 
181, figs 2-104.1-3 (6 9); Song, 1987: 297, figs 253 (6 9); Zhang, 1987: 242, figs 
214.1-5 (6 9); Feng, 1990: 209, figs 184.1-6 (6 9); Chen & Gao, 1990: 186, figs 236a- 
b (6 9); Chen & Zhang, 1991: 308, figs 327.1-3 (8 9); Zhao, 1993: 406, figs 207a-c 
(0*5). 

Menemerus fulvus (L. Koch): Prószynski, 1987: 155 (Tó* 9 from Hasarius). 

Material examined: 1 5 (holotype of Menemerus schensiensis), Zhejiang Province (lo- 
cality given on label: W. Tschekiang) (MNHNP); 1 2 (paratype of Menemerus schensiensis, 
epigynum missing), Hanzhou, Zhejiang Province (locality given on label: Hantschongfu) 
(MNHNP); 1 9 (paratype of sinensis), Nanjing City, Jiangsu Province (locality on label: Nan 
King), leg. G. de Joannis, 1908 (NMB2187a); 2 2 9 (paratypes of Menemerus sinensis Schenkel 
1963, one specimen without epigynum), Nanjing, Jiangsu Province, 1908 (locality given on 
label: Nan Kin) (MNHNP); 16 619 9, Changsha City, Hunan Province, VIII.1986, leg. X. J. 
Peng (HNU); 42 2 , Shangrao City, Jiangxi Province, VI. 1987, leg. X. J. Xie (HNU); 3o* 0*3 2 2 , 
Xichang City, Sichuan Province, X.1975 (HNU); 3 2 2, Jiangfeng Rain Forest, Hainan Province, 
20.V.1990 (MHNG). 

Distribution. China (Anhui, Beijing, Fujian, Guangxi, Guizhou, Hainan, Hebei, 
Hunan, Hubei, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang), India to Japan. 

14. Menemerus legendrei Schenkel, 1963 

Menemerus legendrei Schenkel, 1963: 423, figs 243a-e (2). 

No specimens examined. 
Distribution. China (Yunnan). 

15. Menemerus wuchangensis Schenkel, 1963 
Menemerus wuchangensis Schenkel, 1963: 424, figs 245a-e (2). 

No specimens examined. 
Distribution. China (Hubei). 

16. Myrmarachne schenkeli Peng & Li, 2002 

Myrmarachne lesserti Schenkel, 1963: 391, figs 226a-b (preoccupied by M. tesserti Lawrence, 

1938, 6); Song, Zhu & Chen, 1999: 535, figs 305D, N (o\ 2mislabeled). 
Myrmarachne schenkeli Peng & Li, 2002: 139. 

No specimens examined. 

Distribution. China (Hong Kong) (?) 

Remarks: This species was described from one male collected at "Castle Peak". 
But it is not clear whether "Castle Peak" is in China (Hong Kong) or in USA 
(California) (Schenkel, 1963). 

17. Pellene s denisi Schenkel, 1963 

Pellenes denisi Schenkel, 1963: 440, figs 252a-b (2); Wesolowska, 1981b: 151, figs 68-69 (2); 

Logunov, Marusik & Rakov, 1999: 122. 
Pellenese albomaculatus Peng & Xie, 1993: 80, figs 1-4(2); Song, Zhu & Chen, 1999: 537, figs 

306F-G, 327Q(2). 

Material examined: 1 2 (holotype of Pellenes denisi, epigynum missing), collection data 
missing (according to Schenkel, 1963: "Etsingol oberhalb Kau-taeh", Innere Mongolei, 
20.VI.1886) (MNHNP); 12, Qinshui District, Gansu Province, VII.1986, leg. H. Bei (HNU); 



98 X.-J. PENG ETAL. 

2 2 2, Bohu, Xinjiang Uygur Autonomous Region, 2.V.1982, leg. C. D. Zhu (JLU). 
Distribution. China (Gansu, Inner Mongolia, Xinjiang), Tajikistan. 

18. Pellenes gobiensis Schenkel, 1936 

Pellenes gobiensis Schenkel, 1936: 307, fig. 108 (2); Wesolowska, 1981b: 152, figs 70-71 (2); 
Logunov, 1992: 60, figs 5a-h (3 9); Song, Zhu & Chen, 1999: 537, figs 306H-J, 307 A 

(39). 

No specimens examined. 

Distribution. China (Inner Mongolia), Russia, Mongolia. 

19. Phintella arenicolor (Grube, 1861) 

Jotus difficilis Bösenberg & Strand, 1906: 336, pi. 14, fig. 379 (2); Yin & Wang, 1979: 31, figs 

11A-D (0*2); Hu, 1984: 369, figs 384.1-4 (3 2). 
Dexippus tesserti Schenkel, 1963: 451, f. 257a-k {3 9). 
Phintella melloteei (Simon): Prószyhski, 1983b: 6, fig. 14 (T3 9 from Telamonia); Song, 1987: 

294, fig. 250 (2); Feng, 1990: 204, figs 179.1-6 {3 9) Chen & Gao, 1990: 190, figs 

242a-b (3 2); Chen & Zhang, 1991: 293, figs 308.1-2 (2); Peng et al., 1993: 156, figs 

540-547 (3 9); Zhao, 1993: 414, figs 214a-c (3 9); Song, Chen & Zhu, 1997: 1737, figs 

49a-c(c?). 
Phintella arenicolor (Grube): Logunov & Wesolowska, 1992: 135, figs 24A-C, 25A-B, 26A-C, 

27A, C, E (removed 3 from P. castriesiana, contra Prószyhski, 1979: 310, sub Icius); 

Song, Zhu & Chen, 1999: 538, figs 307J, 308A-B (3 9). 

Material examined: 1 2 , Dongzhong Town, Fangchenggang City, Guangxi Zhuang 
Autonomous Region, alt. 270m, 5.IV.1998, leg. J. Chen (MHNG); 1 2, Xuanen District, Hubei 
Province, 24. V. 1989 (IZCAS). 

Distribution. China (Beijing, Gansu, Hubei, Hunan, Jilin, Shanxi, Yunnan, 
Zhejiang), Russia, Korea, Japan. 

20. Phintella cavaleriei (Schenkel, 1963) 

Dexippus cavaleriei Schenkel, 1963: 454, figs 258a-e (2). 

Icius cavaleriei (Schenkel): Wesolowska, 1981b: 134, figs 18-21 (T2 from Dexippus); Song, Yu 
& Yang, 1982: 210, figs 8-12 (2); Hu, 1984: 366, figs 382.6-10 (3 9). 

Phintella cavaleriei (Schenkel): Prószyhski, 1983b: 6 (T2 from Icius); Song, 1987: 293, figs 
249 (3 9); Feng, 1990: 201, figs 176.1-4 (3); Chen & Gao, 1990: 189, figs 241a-b 
(<î 2); Chen & Zhang, 1991: 293, figs 307.1-4 (3 9); Peng et a!., 1993: 154, figs 532- 
539 (3 2); Song, Chen & Zhu, 1997: 1736, figs 48a-c (2); Song, Zhu & Chen, 1999: 
538, figs 307M, 308E-F, 328C (Ó* 2). 

Material examined: 1 2 . Bifengaou, Bikou Town, Wenxian District, Gansu Province, alt. 
900-1500m, 25.VI.1998, leg. J. Chen (MHNG); 12, Badong District, Hubei Province, 
22. V. 1989 (MHNG). 2 c? 3 5 2 2 , Mt. Xingdoushan, Maoba Town, Lichuan City, Hubei Province, 
6.VI.1989 (IZCAS); 23 3, Chayuan Town, Hefeng District, Hubei Province, 31.V.1989 (IZ- 
CAS). 

Distribution. China (Fujian, Gansu, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, 
Sichuan, Zhejiang), Korea. 

21. Phintella versicolor (C. L. Koch, 1846) 

Jotus munitus Bösenberg & Strand, 1906: 334, pi. 14, figs 374. 392 (Ó* 2); Yin & Wang, 1979: 

32, figs 13A-E (<J 2): Hu, 1984: 370, figs 386.1-6 (3 9). 
Dexippus davidi Schenkel. 1963: 446, f. 255a-e (3). 
Phintella versicolor (C. L. Koch): Prószyhski. 1983a: 44, figs 3 (T3 9 from Chiysilla); Song, 

1987: 288, fig. 245 (3 9); Feng, 1990: 202, figs 177.1-4 (3 9); Chen & Gao, 1990: 191, 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 99 



figs 243a-b (3 9); Chen & Zhang, 1991: 290, figs 303.1-5 (3 9); Peng et al, 1993: 162, 
figs 569-576 (3 9); Zhao, 1993: 411, figs 212a-c (3 9); Song, Chen & Zhu, 1997: 1738, 
figs 50a-c (9); Song, Zhu & Chen, 1999: 539, figs 308O-P, 309F-G, 328E-F (3 9). 

Material examined: 1 9 , Funong Town, Fangchenggang City, Guangxi Zhuang 
Autonomous Region, alt. 220m, 19.IV.1998, leg. M. Wu (MHNG); 19, Longsheng District, 
Guangxi Zhuang Autonomous Region, 6.III.1998, leg. M. Wu (MHNG); 1 3 1 9 , Maheba Town, 
Xianfeng District, Hubei Province (IZCAS); 19, Jinghong, Yunnan Province, 2.XI.1988 (IZ- 
CAS); 49 9, Xinzhuang Town, Xiushan District, Hubei Province, 11. VI. 1989 (MHNG). 

Distribution. China (Anhui, Guangdong, Guangxi, Hubei, Hunan, Jiangxi, 
Taiwan, Yunnan, Zhejiang), Hawaii, Japan, Korea, Sumatra. 

22. Plexippoides discifer (Schenkel, 1953) 

Plexippus discifer Schenkel, 1953b: 88, figs 41 (3); Yin & Wang, 1979: 38, figs 23A-D (3 9); 

Yin & Wang, 1981: 271, figs 3A-G (3, D9); Hu, 1984: 384, figs 400.1-5 (3 9). 
Plexippoides discifer (Schenkel): Prószynski, 1976: 156, fig. 293 (Tó* into generic nomen 

nudum); Prószynski, 1984b: 400 (To" from Plexippus); Zhu & Shi, 1983: 210, figs 192a- 

b (3); Zhang, 1987: 253, figs 225.1-3 (3); Chen & Gao, 1990: 192, figs 245a-b (3 9); 

Pengetal., 1993: 173, figs 606-611 (3 9); Zhao, 1993: 417, figs 216a-b (3); Song, Zhu 

& Chen, 1999: 540, figs 309P-Q, 310G-H, 328 J (3 9). 

Material examined: lo", Qiqi, Yunnan Province, 9.-14.VII.2000 (HNU). 
Distribution. China (Beijing, Hebei, Hunan, Shandong, Shanxi, Yunnan, 
Zhejiang). 

23. Plexippus setipes Karsch, 1879 

Plexippus setipes Karsch, 1879: 89 (9); Yin & Wang, 1979: 37, figs 22A-E (3 9); Song, 1980: 
202, figs 112a-e (3 9); Wang, 1981: 136, figs 76A-C (3 9); Hu, 1984: 387, figs 403.1- 
2(3 9); Guo, 1985: 183, figs 2-106.1-3 (3 9); Zhu & Shi, 1983: 213, figs 195a-c (9) 
Song, 1987: 301, figs 257 (3 9); Zhang, 1987: 251, figs 223.1-3 (9); Feng, 1990: 214 
figs 189.1-5 (3 9); Chen & Gao, 1990: 194, figs 248a-c (3 9); Chen & Zhang, 1991 
297, figs 313.1-4 (3 9); Song, Zhu & Li, 1993: 886, figs 63A-D (3 9 ); Peng et ai, 1993 
185, figs 646-652 (3 9); Zhao, 1993: 419, figs 218a-c (3 9); Song, Zhu & Chen, 1999 
541, figs 3111, 312D, 328N (3 9). 

Dexippus berlandi Schenkel, 1963: 456, figs 259a-e (9). 

Material examined: 19 (holotype of Dexippus berlandi Schenkel, 1963), West Zhejiang, 
IV. 1873 (MNHNP); 23 319, Liuku suburb, Yunnan Province, 25.VI.2000, leg. D Kavanaugh & 
H. M. Yan (HNU); 2c?c?l9, Liuku along Nujiang, Yunnan Province, 26.VI.2000, leg. D. 
Kavanaugh, C. Griswold & H. M. Yan (HNU); lo", Gongshan suburb, Yunnan Province, 
26.VI.2000, leg. H. M. Yan (HNU); la, Liuku along Yongping, Yunnan Province, 26.VII.2000, 
leg. D Kavanaugh & H. M. Yan (HNU); 1 9 , Fulong Town, Fangchenggang City, Guangxi 
Zhuang Autonomous Region, alt. 200m, 23.IV 1999, leg. G. Q. Zhang (MHNG); 19, Nanjing, 
Zhejiang Province, 28.IV 1925, leg. Prof. G. Ping (IZCAS); 19, Xuanen District, Hubei 
Province, 24.V1989 (MHNG); 13, Yingjiang, Dehong District, Yunnan Province, 10. VII 
(MHNG); 1 3 , Mt. Longqi, Jiangle District, Fujian Province, 1 1. IX. 1990, leg. S. Q. Li (IZCAS). 

Distribution. China (Anhui, Fujian, Hebei, Gansu, Guangdong, Guangxi, 
Hubei, Hunan, Jiangsu, Jiangxi, Shanghai, Sichuan, Shaanxi, Shangdong, Shanxi, 
Yunan, Zhejiang), Japan, Turkmenistan, Vietnam. 

24. Pseudeuophrys obsoleta (Simon, 1868) 

Euophrys bacelari Schenkel, 1938: 18, figs 7 (o*). 

Euophrys obsoleta Simon, 1876: 196; Hu & Li, 1987b: 328, figs 48.3-4 (Ó*). 



100 X.-J. PENG ET AL. 

Pseudeuophrys obsoleta (Simon): Zabka, 1997: 78, figs. 272-277 (T 3 9 from Euophjys). 
No specimens examined. 
Distribution. Palearctic Region. 

25. Pseudicius yunnanensis (Schenkel, 1963) comb. nov. Figs 4-7 

Menemerusl yunnanensis Schenkel, 1963: 426, fig. 245 (9); Wesolowska, 1981b: 150, figs 64- 
67 ( 9 ); Song, Zhu & Chen, 1999: 534, fig. 304D ( 9 ). 

Material examined: 1 9 (holotype of Menemerusl yunnanensis, epigynum missing), 
Yunnan Province, 1913 (locality given on label: Yunnan-Fou, alt. 1850-2000m) (MNHNP). 

Remarks. This species clearly belongs to the genus Pseudicius because of the 
presence of a row of peculiar stridulatory spines on tubercles situated below the eye lat- 
erally (arrow in fig. 5) and of corresponding microscopic spines prolaterally on femur 
I (arrow in fig. 6). 

Distribution. China (Yunnan). 

26. Pseudoheliophanus similis Schenkel, 1963 

Pseudoheliophanus similis Schenkel, 1963: 433 (description of juvenile). 

Material examined: 1 9 (holotype, immature), collection data missing (according to 
Schenkel, 1963: "Fluss Barchany im N. von Borobalgassun, Ordos", 12.IX.1884) (MNHNP). 

Distribution. China (Inner Mongolia). 

27. Rhene plana (Schenkel, 1936) 

Ballus planus Schenkel, 1936: 244, fig. 80 (9). 

Rhene planus Logunov, 1993: 51, figs 3A-B (T9 from Ballus); Song, Zhu & Chen, 1999: 558, 
figs315A-B(9). 

Material examined. 1 9 (paratype of Ballus planus), Gansu Province (locality given on 
label: S. Kansu, K13, Hummen coll.), K13 Hedius Exp. (NMB2161a). 

Distribution. China (Gansu). 

28. Salticus potanini Schenkel, 1963 

Salticus potanini Schenkel, 1963: 410, figs 236a-e (9); Wesolowska, 1981a: 79, figs 106-107 
(9); Prószyhski, 1982: 288, fig. 43 (9); Tu & Zhu, 1986: 94, figs 34-38 (3 9); Zhang 
& Zhu, 1987: 33, figs 3A-C (3 9); Zhang, 1987: 254, figs 226.1-4 (3 9); Zhou & Song, 
1988: 8, figs lla-c (9); Hu & Wu, 1989: 383, figs 300.1-7 (3 9); Chen & Gao, 1990: 
197, figs 251a-b (3 9); Peng et al, 1993: 206, figs 727-734 (3 9); Song, Zhu & Chen, 
1999: 558, figs 315F-G, I, 329G (c? 9). 

Material examined: 3 3 3 3 9 9 , Qintongxia Forest Farm, Ningxia Hui Autonomous 
Region, 16.III.1982, leg. L. D. Jia (HNU); lc?l9, Yangcheng District, Hebei Province, 
12.VII.1980, leg. M. S. Zhu (HNU); 1 3, Luguantai, Zhouzhi District, Shaanxi Province, 
10.X.1992, leg. X. J. Pens (HNU); 19, Bohu District, Xinjiang Uygur Autonomous Region, 
7.VI. 1982, leg. N. L. Zhou (MHNG). 

Distribution. China (Hebei, Hubei, Hunan, Jiangsu, Ningxia, Shaanxi, Shanxi, 
Sichuan, Taiwan, Xinjiang, Zhejiang); Korea, Mongolia. 

29. Sitticus avocator (O. P.-Cambridge, 1885) 

Sitticus paraviduus Schenkel. 1963: 402, figs 232a-c (o* 9); Zhu & Shi, 1983: 215, figs 198a-c 

(3). 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 101 



Sitticiis avocator (O. P.-Cambridge): Prószynski & Zochowska, 1981: 26, figs 25-26 (Te? from 
Attulus); Zhou & Song, 1988: 9, figs 12a-f (c? 9); Hu & Wu, 1989: 386, figs 302.6-10 
(c? 9); Tang & Song, 1990: 52, figs 4A-C (9 ); Song, Zhu & Chen, 1999: 559, figs 315Q- 
R, 3 16F-G (<?$). 
Material examined: le? (holotype of Sitticus paraviduus), collection data missing 

(according to Schenkel, 1963: "Grenze Chara su-Cha , linkes Ufer des Etsingol", 23. -29. VII, 

1886) (MNHNP). 

Distribution. China (Inner Mongolia, Shanxi, Tibet, Xinjiang), Russia, Central 
Asia to Japan. 

30. Sitticus clavator Schenkel, 1936 

Sitticus clavator Schenkel, 1936: 247, fig. 81 (c?); Song, 1987: 303, fig. 259 (<J); Logunov, 
1993: 9, figs 7, 24-25 (<J); Song et al., 1996: 108, figs 4A-C (9); Song, Zhu & Chen, 
1999: 559, figs 316H-J, 317A (c? 9). 

Sitticus penicillatus (Simon): Peng et al., 1993: 218, figs 772-777 (c? 9, misidentified). 

Material examined: 1 Ô 1 9 , Mt. Huang, Anhui Province, 23.X.1974 (HNU); 2 c? â 1 9 , 
Longjing, Jilin Province, 7.-8.VI.1971 (JLU); lc?, Sanhe District, Jilin Province, V. 1971 (JLU); 
3 c? o\ Yesanpo, Laishui District, Hebei Province, 12.-13.V.2001, leg. X. J. Peng (MHNG). 

Distribution. China (Anhui, Gansu, Hebei, Jilin). 

31. Sitticus sinensis Schenkel, 1963 

Sitticus sinensis Schenkel, 1963: 404, figs 233a-d (<J 9); Yin & Wang, 1979: 39, figs 25A-E 
(c? 9); Hu, 1984: 391, figs 408.1-3 (c? 9); Zhu & Shi, 1983: 217, figs 200a-i (<? 9); 
Zhang, 1987: 258, figs 230.1-4 (<? 9); Hu & Wu, 1989: 392, figs 306.1-2 (9); Chen & 
Zhang, 1991: 305, figs 323.1-3 (6 9); Peng et ai, 1993: 219, figs 778-786 (c? 9); Zhao, 
1993: 426, figs 224a-d (c? 9); Song, Zhu & Chen, 1999: 559, figs 316P, 317D. 329M 
(0*9). 

Material examined: lc?39 9, Yesanpo Town, Laishui District, Hebei Province, 12.- 
13.V.2001, leg. X. J. Peng (MHNG); lo", Huairou District, Beijing, 1.VI.2001, leg. X. J. Peng 
(IZCAS); 16c? 0*8 9 9 , Jilin Province, 1955-1984 (JLU); 7<? c?5 9 9 , Beijing City, 6-8.VII.1974 
(HNU); 2c? â, Shijiazhuang, Hebei Province, V.1985 (HNU); 19, Manasi District, Xinjiang 
Uygur Autonomous Region (IZCAS); 29 9, Ürümqi, Xinjiang Uygur Autonomous Region, 
11.V.1984, leg. W. H. Zhen (MHNG); lc?l9, Manasi District, Xinjiang Uygur Autonomous 
Region, 12.V.1984 (IZCAS); lc?29 9, Bohu District, Xinjiang Uygur Autonomous Region, 
15.IV.1981 (IZCAS); lc?, Luochuan, Shaanxi Province, 10.VI.1981 (IZCAS). 

Distribution. China (Beijing, Gansu, Hebei, Hunan, Jilin, Liaoning, Shaanxi, 
Shandong, Shanxi, Qinghai, Xinjiang). 

32. Synagelides cavaleriei (Schenkel, 1963) 

Tagoria cavaleriei Schenkel, 1963: 394, figs 227a-l (c? 9 ). 

Synagelides cavaleriei (Schenkel): Bohdanowicz & Heciak, 1980: 248, figs 1-9 (c?9); Song, 

1987: 306, figs 262 (c? 9); Bohdanowicz, 1987: 66, figs 1-4 (c? 9); Hu & Li, 1987b: 334, 

figs 49.3-4 (9); Song, Zhu & Chen, 1999: 560, figs 318L-M, 319D-E (c? 9). 

Material examined: 1 c? 1 9 (syntypes of Tagoria cavaleriei), today's name of locality not 
known (locality given on label: Anschunfou) (MNHNP); 3c?c?, Mt. Qijiemeishan, Xuanen 
District, Hubei Province, 25.V1989, leg. S. Q. Li (IZCAS). 

Distribution. China (Guizhou, Hubei, Tibet). 

33. Talavera trivittata (Schenkel, 1963) Fig. 8 

Euoph-ys trivittata Schenkel, 1963: 401, figs 231a-b (9); Wesolowska, 1981b: 130, figs 8-9 (9); 
Hu, 1984:359, figs 374.1-2 (9); Paik, 1986: 20, figs 1-10 (c? 9); Hu & Wu, 1989: 363, 
figs 285.3-4 (9). 



102 



X.-J. PENG ET AL. 




Figs 1-8 
1. Body of Asianellus potanini (Schenkel, 1963), male. 2-3. Heliophanus potanini (Schenkel, 
1963): 2 body of female; 3 cheliceral teeth. 4-7. Pseudicius yunnanensis (Schenkel, 1963) comb. 
nov.: 4 body of female; 5 carapace, lateral view; 6 femur I, prolateral view; 7 cheliceral teeth. 8. 
Body of Talavera trixittata (Schenkel. 1963), male. 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 



103 



0.5 mm 




Figs 9-14 
9-11. Thiania cavaleriei Schenkel, 1963: 9 body of male; 10 left palpal organ, ventral view; 
11 left palpal organ, retrolateral view. 12-14. Thiania luteobrachialis Schenkel, 1963: 12 body 
of female; 13 epigynum; 14 vulva. 



104 X.-J. PENG ET AL. 

Talavera trivittata (Schenkel): Logunov, 1992: 78 (Te? 2 from Euophrys); Song, Zhu & Chen, 
1999: 561, fig. 321F (2). 

Material examined: 1 2 (holotype of Euophrys trivittata), collection data missing (ac- 
cording to Schenkel, 1963: "Kloster Schine, Sume, Ordos", 9.IX.1884) (MNHNP). 

Distribution. China (Inner Mongolia), Korea, Japan. 

34. Tasa davidi (Schenkel, 1963) 

Thianella davidi Schenkel, 1963: 412, figs 237a-e (6). 

Tasa davidi (Schenkel): Wesolowska, 1981b: 157, figs 88-92 (6); Peng et al., 1993: 230, figs 

819-823 (Ó*); Song, Zhu & Chen, 1999: 561, figs 319L-M, 330B (<?); Peng, Gong & 

Kim, 2000: 13, figs 15-18(2). 

Material examined: 30*0*, Daoxian District, Hunan Province, 24.IV.1988, leg. L. S. 
Gong (MHNG); 1 o*3 2 2 , Ganxia Town, Daoxian District, Hunan Province, 3.VII.1987, leg. L. 
S. Gong (HNU); 32 2, Bajia Town, Daoxian District, Hunan Province, 31.IX.1987, leg. L. S. 
Gong (HNU); 1 2, Shuangqiao Town, Daoxian District, Hunan Province, 10. VIII. 1987, leg. L. 
S. Gong (HNU); 1 3 , Ningxiang District, Changsha City, VIII. 1986, leg. X. Q. Xiao (HNU); 1 2 , 
Mt. Lushan, Jiangxi Province, VI. 1987, leg. L. P. Xie (HNU); 36 6, Daoxian District, Hunan 
Province, 24. IX. 1988, leg. L.S. Gong (HNU). 

Distribution. China (Hunan, Jiangxi, Shaanxi). 

35. Thiania cavaleriei Schenkel, 1963 Figs 9-11 

Thiania cavaleriei Schenkel, 1963: 406, figs 234a-g (6); Prószynski, 1976: 150, fig. 94 (<?); 
Prószynski, 1984a: 147 (3); Song, Zhu & Chen, 1999: 562, figs 320L-M (<J). 

Material examined: Id (holotype of Thiania cavaleriei), today's name of locality not 
known (locality given on label: Anschunfu) (MNHNP). 
Distribution. China (Gansu). 

36. Thiania luteobrachialis Schenkel, 1963 Figs 12-14 

Thiania luteobrachialis Schenkel, 1963: 408, fig. 235 (2). 

Material examined: 1 2 (holotype of Thiania luteobrachialis), today's name of locality 
not known (locality given on label: "Lo Thoei-Tong") (MNHNP). 
Distribution. China. 

37. Thyene imperialis (Rossi, 1846) 

Thyene imperialis (Rossi): Peckham & Peckham, 1901: 307, pi. 25, fig. 4; Wesolowska, 1981a: 
75, figs 94-101 (3 2); Prószynski, 1987: 109-110 (ó*); Peng et al., 1993: 242, figs 862- 
868 (3 2); Xie & Peng, 1995: 107, figs 5A-F (3 2). 

Thyene sinensis Schenkel, 1963: 441, figs 253a-f (2). 

Material examined: 233, Chongan District, Fujian Province, VII.1986 (HNU); 16, 
Wuning District, Guangxi Zhuang Autonomous Region, VIII. 1981 (HNU); 16, Tianlin District, 
Guangxi Zhuang Autonomous Region, IX. 1981 (HNU); 26 6, lianfeng, Ledong District, 
Hainan Province, XII. 1989 (MHNG). 

Distribution. China (Fujian, Guangdong, Guangxi, Hainan, Hubei), Old World. 

RESULTS AND DISCUSSION 

Dr E. Schenkel described 41 new salticids species from China between 1936 
and 1963. 13 of them remain in the original combination: i.e. Bianor hotingchiehi 
Schenkel, 1963; Hasarina contortospinosa Schenkel, 1963; Heliophanus potanini 
Schenkel, 1963; Menemerus legendrei Schenkel, 1963; Menemerus wuchangensis 



CHINESE JUMPING SPIDERS STUDIED BY SCHENKEL 105 

Schenkel, 1963; Pellenes denisi Schenkel, 1963; Pellenes gobiensis Schenkel, 1936; 
Pseudoheliophanus similis Schenkel, 1963; Salticus potanini Schenkel, 1963; Sitticus 
clavator Schenkel, 1936; Sitticus sinensis Schenkel, 1963; Thiania cavaleriei 
Schenkel, 1963; Thiania luteobrachialis Schenkel, 1963. 

9 species were earlier transferred to other genera; one is transferred in this 
paper: 

1. Ballus planus Schenkel, 1936 = Rhene plana (Schenkel, 1936) 

2. Davidia magnidens Schenkel, 1963 = Davidina magnidens (Schenkel, 1963) 

3. Dexippus cavaleriei Schenkel, 1963 = Phintella cavaleriei (Schenkel, 1963) 

4. Euophiys trivittata Schenkel, 1963 = Talavera trivittata (Schenkel, 1963) 

5. Menemerus fagei Schenkel, 1963 = Heliophanus potanini Schenkel, 1963 

6. Menemerus yunnanensis Schenkel, 1963 = Pseudicius yunnanensis (Schenkel, 
1963) comb. nov. 

7. Phlegra potanini Schenkel, 1963 = Asianellus potanini (Schenkel, 1963) 

8. Plexippus discifer Schenkel, 1953 = Plexippoides discifer (Schenkel, 1953) 

9. Tagoria cavaleriei Schenkel, 1963 = Synagelides cavaleriei (Schenkel, 1963) 
10. Thianella davidi Schenkel, 1963 = Tasa davidi (Schenkel, 1963) 

17 species were placed in synonymy: 

1 . Carrhotus pienoni Schenkel, 1 963 = Carrhotus xanthogramma (Latreille, 1819) 

2. Dexippus berlandi Schenkel, 1963 = Plexippus setipes Karsch, 1879 

3. Dexippus davidi Schenkel, 1963 = Phintella versicolor (C. L. Koch, 1846) 

4. Dexippus lesserti Schenkel, 1963 = Phintella arenicolor (Grube, 1861) 

5. Dexippus tschekiangensis Schenkel, 1963 = Phintella versicolor (C. L. Koch, 
1846) 

6. Euophrys bacelari Schenkel, 1938 = Pseudeuophrys obsoleta (Simon, 1868) 

7. Evarcha pichoni Schenkel, 1963 = Evarcha albaria (L. Koch, 1878) 

8. Heliophanus berlandi Schenkel, 1963 = Heliophanus curvidens (O. P.-Cam- 
bridge, 1872) 

9. Menemerus bonneti Schenkel, 1963 = Menemerus bivittatus (Dufour, 1831) 

10. Menemerus sinensis Schenkel, 1963 = Menemerus fulvus (L. Koch, 1878) 

11. Menemerus schensiensis Schenkel, 1963 = Menemerus fulvus (L. Koch, 1878) 

12. Mithion hotingchiehi Schenkel, 1963 = Mendoza elongata (Karsch, 1879) 

13. Mithion pichoni Schenkel, 1963 = Mendoza canestrinii (Ninni, 1868) 

14. Mithion tschekiangensis Schenkel, 1963 = Mendoza canestrinii (Ninni, 1868) 

15. Phlegra pichoni Schenkel, 1963 = Asianellus festivus (C. L. Koch, 1834) 

16. Sitticus paraviduus Schenkel, 1963 = Sitticus avocator (O. P.-Cambridge, 1885) 

17. Thyene sinensis Schenkel, 1963 = Thyene imperialis (Rossi, 1846) 

Myrmarachne lesserti Schenkel, 1963 is preoccupied by M. lesserti Lawrence, 
1938, and was renamed as Myrmarachne schenkeli Peng & Li, 2002. 



106 X.-J. PENG ET AL. 



ACKNOWLEDGMENTS 

We are very grateful to Dr Ambros Haenggi (NMB) for checking all the mate- 
rial studied by Dr E. Schenkel and deposited in the NMB. Special thanks are given to 
Dr Peter Schwendinger (Geneva) and Dr Peter Jäger (Mainz) for their continued sup- 
port during our study on Chinese jumping spiders. 

The present study was supported by the National Natural Sciences Foundation 
of China (NSFC-30270183, 39970102), National Science Fund for Fostering Talents in 
Basic Research (NSFC-J0030092), also partly supported by the project of Knowledge 
Innovation Program, Chinese Academy of Science (KSCX2-1-06A, KSCX3-IOZ-01) 
and the Life Science Special Fund of CAS supported by the Ministry of Finance 
(STZ-00-19). 

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Revue suisse de Zoologie 110 (1): 111-124; mars 2003 



New Sphaerochthonius species from the Neotropical region (Acari: 
Oribatida) 

Heinrich SCHATZ 

Institut für Zoologie und Limnologie, Leopold-Franzens-Universität, Technikerstr. 25, 

A-6020 Innsbruck, Austria. E-mail: heinrich.schatz@uibk.ac.at 



New Sphaerochthonius species from the Neotropical region (Acari: 
Oribatida). - Two new species of the family Sphaerochthoniidae (Acari: 
Oribatida) are described and illustrated. Sphaerochthonius litoralis sp. n. 
occurs in the littoral zone of several islands of the Galapagos archipelago 
(Ecuador), S. windsori sp. n. was collected in dry deciduous forests in 
Panama. Adult and juvenile instars of both species are described. A key of 
Sphaerochthonius species known from the Neotropical region is added. 

Key- words: Sphaerochthonius - Acari - Oribatida - taxonomy - new species 
- Galapagos Islands - Panama - Neotropical Region. 

INTRODUCTION 

Members of the oribatid mite genus Sphaerochthonius are inhabitants of 
organic litter. They are known from all Zoogeographie regions, but occur mainly in the 
tropics and subtropics. Up to now, four Sphaerochthonius species have been described 
and recorded from the Neotropical region, most of them from one site and single 
specimen: Sphaerochthonius fungifer Mahunka, 1983 (Guatemala, Tikal), S. phyllo- 
phorus Balogh & Mahunka, 1969 (Brazil, Rio de Janeiro; see also Balogh et al., 1969), 
S. Uruguay ensis Pérez-Inigo & Sarasola, 1998 (Uruguay, San Miguel mountain forest) 
and S. splendidus (Berlese, 1904) (Galapagos Islands, see Schatz, 1998, 1999). An 
addititional undescribed species was reported from the Galapagos Islands in the 
Eastern Pacific Ocean (Schatz, 1998, 1999). The latter species is described in this 
paper, along with another new Sphaerochthonius species from Panama. 

The notogastral setal nomenclature follows that of Grandjean (1934) as sum- 
marized by Hunt et al. (1998). All measurements are given in micrometers (urn). 

DESCRIPTIONS AND REMARKS 

Sphaerochthonius litoralis sp. n. Figs 1-14 

Sphaerochthonius sp. Schatz, 1998: 379; Schatz, 1999: 504 

Type material: The adult holotype (type locality: Ecuador, Galapagos Islands, Isla 
Sombrero Chino southeast off Isla Santiago, near Cape Trenton, 0° 22' S, 90° 35' W) and 2 adult 
paratypes from the same locality are preserved in alcohol and will be deposited in: Muséum 
d'histoire naturelle, Genève, Switzerland, other specimens in the author's collection in Inns- 
bruck, Austria. 



Manuscript accepted 27.08.2002 



112 H.SCHATZ 



Other material examined: Sphaerochthonius litoralis sp. n. is known from 56 specimens 
(54 adults, 2 tritonymphs) collected on eight islands of the Galapagos Archipelago. No morpho- 
logical differences between the populations on the different islands were observed. All records 
are from the eastern and central islands of the Galapagos. 

Specific collection data (all collections by the author): Isla Bartolomé, near Pinnacle 
Rock, littoral zone: sea level, in dry to moist leaf litter under May tenus octogona (11 .ii. 1985: 1 
adult, 26.xii.1986: 6 adults), in moist mangrove leaf litter under Laguncularia racemosa 
(26.xii.1986: 1 adult). Isla Floreana (Santa Maria), at flamingo lagoon in Punta Cormoran, 
littoral zone: at sea level, in moist, decayed mangrove leaf litter under Laguncularia racemosa 
(6.iv.l985: 6 adults). Isla Santa Fé, northeastern part, littoral to arid zone: above beach, 5 m a.s.l., 
in dry to moist leaf litter under Cordia lutea (14.iii.1983: 1 adult). Isla Genovesa, Darwin Bay, 
littoral zone: at sea level, lava crevice, in dry to moist decayed leaf litter under Rhizophora 
mangle (17.ii.1985: 3 adults); around Lake Arcturus, littoral zone: 20 m a.s.l., in moist, decayed 
leaf litter and black humus under Rhizophora mangle (16.ii.1985: 3 adults). Isla Rabida, littoral 
zone: at flamingo lagoon, 5 m a.s.l., in moist, decayed mangrove leaf litter under Rhizophora 
mangle (14.iii.1988: 3 adults). Isla San Cristobal, littoral zone: south of Wreck Bay, behind 
rocky coast, at sea level, in moist leaf litter under Conocarpus erecta and Laguncularia race- 
mosa (4.Ì.1987: 1 adult); in moist and salty, well-decayed leaf litter, humus and rotten wood un- 
der Avicennia germinans (4.Ì.1987: 5 adults, 1 tritonymph). Isla Santa Cruz, near Charles Darwin 
Research Station, littoral zone: 5 m a.s.l., in moist leaf litter and humus under Sesuvium portula- 
castrum (3.iii.l985: 1 adult); ibid., behind coast, 5 m a.s.l., in dry to moist, well-decayed leaf 
litter and pieces of wood under Hippomane manzinella (22. i. 1987: 1 adult); Puerto Ayora, near 
Hotel "Fragata", littoral zone: at sea level, in moist, decayed mangrove leaf litter under Avicennia 
germinans (30.xii.1986: 3 adults); ibid., in moist, decayed leaf litter under Sesuvium portula- 
castrum (30.xii.1986: 4 adults). South of Puerto Ayora, "'Divine's Bay", littoral zone: at sea lev- 
el, in moist, partially decayed mangrove leaf litter under Rhizophora mangle (29.xii.1986: 8 
adults). West of Canal de Itabaca, littoral zone: 5 m a.s.l., in moist, partially decayed mangrove 
leaf litter and sand under Avicennia germinans (13.i. 1987: 1 adult). Transition zone near 
"Caseta": 180 m a.s.l., in dry old dropping of giant tortoise, containing digested grass 
(26. ii. 1985: 1 adult). Moist zone: Scalesia forest near Cerro Crocker, 700 m a.s.l., in dry to moist 
bark, lichens and moss on a Scalesia pedunculata trunk, arboricolous (6.iii.l987: 1 adult). Isla 
Sombrero Chino, northern part, near lava flow, littoral zone, 5 m a.s.l., in moist, decayed leaf lit- 
ter and red soil under Sesuvium edmonstonei (21.ii.1987: 3 adults, 1 tritonymph); ibid., in dry to 
moist, decayed leaf litter and humus under Heliotropium angiospermum (21. ii. 1987: 1 adult). 

Diagnosis: Adults of the new species differs from its congeners by the follow- 
ing combination of character states: surface structure on prodorsum and on major part 
of notogaster consisting of a polygonal network with triangular thickenings in its 
angles; nose-shaped protuberance on prodorsum present; except setae exa, exp, d, all 
prodorsal and notogastral setae biramous, papillate; adorai, subcapitular, epimeral, 
genital and anal setae distally almost flagelliform; 8 pairs of genital, 9-11 pairs of anal 
and 5 pairs of adanal setae present, the latter uniramous with branching papillae, setae 
(ul) on palptarsus fused, bifid. Tritonymph with biramous and ciliate setae d and 
asymmetric biramous setae e. 

Adult (Figs 1-12): Total body size (n=38): 321 (range: 280-350) x 192 (170- 
215), length of notogaster 217 (195-225, 66-70% of total body length). Color yellow 
to light brown. Prodorsal and major part of notogastral surface (plates Na, Py and Is) 
covered with polygonal network, margins and especially angles of areolae irregularly 
thickened, surface granulate (Fig. 6). Reticulation of cuticle accentuated by cero- 
tegument. Lateral plates of notogaster, mentum, epimeral, genital and ano-adanal 
plates with granulate surface. 



NEW NEOTROPICAL SPHAEROCHTHONIUS 1 13 

Prodorsum (Fig. 1): Shape elongate, lateral edges slightly converging anterior- 
ly. Rostrum medially with a thickened, nose-shaped protuberance, slightly protruding 
anteriorly beyond anterior margin of prodorsal shield. Surface structure polygonal to 
hexagonal as on notogaster, area between interlamellar setae with larger polygons. 
Rostral, lamellar and interlamellar setae biramous, papillate ("T-shaped" in the sense 
of Wallwork, 1960), both rami of ro, le almost equal in length, il with longer anterior 
rami, extension of both rami ro 40-50, le 50-55, il 30-35 long. Anterior (exa) and pos- 
terior (exp) exobothridial setae inserted close to each other anterolateral to bothridia; 
both uniramous, papillate, exa -12, exp 25 long. Bothridia directed laterad, head of 
sensilli ramose with branching papillae, rami directed dorsad (hence sensilli appearing 
narrow in dorsal view, see Fig. 1, broader in lateral view, see Fig. 2), total length of sen- 
silli 50, head 40. Between bothridia a small elevated ridge present on prodorsum. 

Notogaster (Figs 1-3): Shape almost round. Small angle on each side of noto- 
gaster on broadest part, formed by lateral angle of triangular lateral plates L. Broad 
transverse scissure (of type "L" in the sense of Grandjean, 1947; see Norton, 2001) 
present between plates Na and Py, originating from lateral angles of plates L, curved 
anteriad, covered by hypertrophied posterior part of plate Na, hiding insertions of se- 
tal row d on scissure. Indistinct thickened suture present on plate Py, also originating 
from lateral angles of plates L, almost transverse, passing near insertions of setal row 
e. On ventral side single transverse scissure anterior to plate Py and straight longitudi- 
nal scissure present on each side between plates Is and L. 

Sixteen pairs of notogastral setae present. Setae d short, setiform, smooth, d l 
4-5, d 2 7 long. Other notogastral setae biramous, papillate. Rami of setal row c aligned 
in almost longitudinal direction, slightly asymmetric with shorter anterior rami, 
extension of both rami: c 1 75, c 2 65-70, c 3 65-70, c p 60-70, rami of setae e aligned trans- 
versely, extension e 1 70-75, e 2 60, rami of/ 7 aligned longitudinally, extension 70, rami 
off 2 aligned transversely, extension 60, setae h r h 3 on edge of notogaster, extension of 
both rami 60, setae ps I -ps 3 visible in lateral and ventral view, extension of both rami: 
psj 50-60, ps 2 , ps 3 40-50, setae ps 2 and ps 3 inserting on plates Is lateral to ano-adanal 
plates. 

Gnathosoma (Figs 3-5): Mentum small (35 x 30), subcapitular setae setiform, 
basally slightly thickened, distally very thin, almost fiagelliform, h, m l7 m 2 10-12, a 
16-20 long. Three pairs of adorai setae present, same shape as subcapitular setae, -12 
long. Chelicerae (Fig. 4) of chelate-dentate type, narrow, size (n=6): 45-60 x 20-25, 
movable digit 20-25 long, both setae inserted near dorsal margin, directed anteriad, cha 
setiform, ciliate, 20-22 long, chb strongly thickened, palmate, 10-12 long. Palps (Fig. 
5) with 5 short articles, attenuating from trochanter to tibia, tarsus distally prolonged. 
Setal formula (trochanter to tarsus, solenidion in parentheses) - 2 - 1 - 3 - 11(1), setae 
cm 40 long, acm fiagelliform, 35 long, with a distal loop, other tarsal setae relatively 
long (12-15). Setae (ul) and sul eupathids, (ul) fused, bifid with long curved seta and 
short spine. 

Epimeral region (Fig. 3): Epimeral plates I and II separate, plates III and IV 
almost completely fused on each side. Epimeral setal formula 3-2-3-4, setae 3c 
inserting on anteriorly directed apophyses, setae 4b, 4c, 4d in one row on posterior part 
of plate. Setae relatively long, setiform, basally slightly thickened, distally very thin, 



1 14 H. SCHATZ 

almost flagelliform, la, lc, 2a, 2b ciliate, others smooth, lc, 2a, 2b 25-30, others 
15-20 long. 

Genito-anal region (Figs 3, 7): Genital plates small, elongate (70-75 x 20), 
shape oval, surface granulate. Eight pairs of genital setae present, shape as epimeral se- 
tae, 20-27 long. Anal and adanal plates fused. Ten pairs of anal setae present (some 
specimens with 9 or 1 1 pairs), basally thickened, distally very thin, almost flagelliform, 
curved mediad and posteriad, 12-15 long. Five pairs of adanal setae present (Fig. 7), 
uniramous with branching papillae, 20-30 long, bases connected by a sclerotized ridge. 

Legs (Figs 8-12): Legs of moderate length (without claws 30-37% of body 
length). Setal formulas of legs (trochanter to tarsus, solenidia in parentheses): leg I 
. 4 . 4 . 6(1) - 19(3), leg II 1 - 5 - 4 - 6(1) - 17(2), leg III 2 - 3 - 3 - 4(1) - 12, leg IV 
2- 3 - 3 - 4(1) - 11. All legs heterotridactylous, with a small, but strong median claw 
and two almost setiform lateral claws. Setae d and I" on femur I and setae d on femora 
II - IV thickened and densely ciliate. Solenidia (Oj on tarsus I curved anteriad, covered 
with cerotegument granulae, 25 long, solenidia qp ; on tibia I very long (85-90), inserted 
on large distal projection. 

Immatures (Figs 13,14): Tritonymph: Color pale. Size (n=2) 275-295 x 180, 
notogaster 200-210 long. 

Prodorsum (Fig. 13): Nose-shaped structure as in adult. Prodorsal setae as in 
adult, extension of both rami: ro 40, le 50, il 35, anterior ramus of il almost twice as 
long as posterior one, exa 10, exp 18-20. Total length of sensilli 50 urn, head 30. 

Gastronotic region (Figs 13,14): Broad dorsal scissure between plates Na and 
Py well-developed and deep, additional posterior scissure present along insertions of 
setal row e. Surface structure of plates Na, Py, Is with polygonal structure as in adults, 
lateral plates L triangular, surface granulate with radial striae, inner margin with a 
series of folds. Sixteen pairs of notogastral setae present, setae c r c p biramous, 
papillate, extension of both rami 55-60. Setae d on anterior slope of cavity, biramous, 
ciliate, small, rami directed transversely, extension of both rami: d, 17, d 2 8-10. Setae 
e biramous, papillate, asymmetric, with long rami directed posteriad, e 1 70-80, e 2 
40-50 long, and short second rami originating on base, directed mediad, 15-20 long, 
setae/, h, ps biramous, papillate, extension of both rami: /, h, psj 45-55, ps 2 , ps 3 35, 
latter inserting on plate Is, rami aligned almost transversely, rami of setal row /aligned 
longitudinally, others parallel to body edge. 

Gnathosoma (Fig. 14): Size of mentum 30x20, subcapitular setae 12-14 long. 
Palps: setal formula as in adult, setae acm 25-30 long, setae (v) short, thick, spiniform, 
(ul) fused, bifid. 

Ventral region (Fig. 14): Shape of epimeral plates and shape, position and num- 
ber of epimeral setae as in adult, setae of rows I and 2 12-15 long, others 6-9. Genital 
plates with 3 pairs of genital papillae, 7 pairs of genital setae, 10-12 long. Anal and 
adanal plates fused, 9 pairs of anal setae, 9-10 long, and 5 pairs of adanal setae present, 
the latter uniramous, papillate, 20 long. Shape of genital and anal setae as in adults. 

Legs: All legs monodactylous, setal formula as in adults, setae d and /" on 
femur I and setae d on femora II - IV thickened with short ciliae. Solenidion qp y on tibia 
I 70-75 Ions. 



NEW NEOTROPICAL SPHAEROCHTHONIUS 



115 




ta *mmxit»mitrii>mnnmtmtii!""' 




Figs 1-3 
Sphaerochthonius litoralis sp. n., adult - 1. Dorsal view; 2. Lateral view; 3. Ventral view (Scale: 
100 urn). 



Etymology: The name is derived from the Latin word "litoralis" = "belonging to 
the littoral" and refers to the known preferred habitat of the species. 

Remarks: In many aspects the new species morphologically corresponds with 
the original description of S. transversus Wallwork, 1960. Both species have a reti- 
culate surface structure, biramous notogastral setae with rami of almost equal length, 
especially in row e and /, and 5 pairs of adanal setae. Differences between the two 
species are found in: the presence of a nose-shaped structure on prodorsum in 



116 



H. SCHATZ 




Figs 4-12 
Sphaewchthonius litorali s sp. n., adult - 4. Chelicera; 5. Palp; 6. Surface structure on notogaster; 
7. Adanal setae adf, 8. Leg I; 9. Leg II; 10. Leg DI; 1 1. Leg IV, trochanter - tibia; 12. Leg IV, tar- 
sus. All appendages in abaxial aspect (Scale a - Figs 4-7, Scale b - Figs 8-12: 50 urn). 



NEW NEOTROPICAL SPHAEROCHTHON1US 



117 




Figs 13-14 
Sphaerochthonius litoralis sp. n., tritonymph - 13. Dorsal view; 14. Ventral view (Scale: 
100 urn). 



S. litoralis, the shape of interlamellar setae il (litoralis: biramous, transversus: unira- 
mous), details in surface structure (litoralis: triangles in angles of polygons, transver- 
sus: dense granulate secretion along edges of polygons), shape of adanal setae 
(litoralis: widely branching, transversus: biramous with very short inner rami), in 
setation of palptarsi (litoralis: setae (ut) fused, bifid with 1 long and 1 short seta, trans- 
versus: setae (ul) fused, bifid with 2 long setae), and in shape of setae d, e in 
tritonymphs (litoralis: dj, d 2 biramous, e h e 2 biramous, anterior ramus short, trans- 
versus: setae d not mentioned, probably very short, setae e uniramous). 

The number of anal setae varies in S. litoralis: 75% of the adult population on 
the Galapagos Islands bear 10 pairs, 7.5% 9 pairs, and 17.5% 11 pairs. No correlation 
between the anal setation and occurrence on different islands or sites could be found. 
This character is variable within several species of the family; different numbers of 
anal setae are known from Similochthonius australis (Mahunka, 1985): 5-7, S. decora- 
tus Mahunka, 1985: 4-5, Sphaerochthonius strinata Mahunka, 1982: 9-10, S. suzukii 
Aoki, 1977: 7-8, S. transversus Wallwork, 1960: 10 and more, 5. variesetosus 
Mahunka, 1997: 10-13 pairs of anal setae. 

The tritonymph of Sphaerochthonius litoralis sp. n. differs from the adult instar 
by the smaller posterior edge of plate Na, leaving the scissure and setae d uncovered; 



118 H. SCHATZ 

by the presence of an additional transverse scissure along setal row e, which is reduced 
to an indistinct suture in adults; by the shape of setae d and e; by the number of geni- 
tal and anal setae; and by the number of claws. The monodactyly and the number of 
dorsal scissures ("coupures") in juvenile instars of Sphaerochthoniidae are discussed in 
van der Hammen (1959). 

Ecology: The vast majority of specimens of Sphaerochthonius litoralis sp. n. 
was collected in the littoral zone. This species occurs mainly in moist, partially de- 
cayed to strongly decayed leaf litter under mangroves and other deciduous trees, as 
well as under Sesuvium and Heliotropium, also in deeper humus layers. Remarkable 
records are from the shore of Lake Arcturus, a remote saltwater crater lake in the inte- 
rior of Isla Genovesa (3 specimens); in an old giant tortoise dropping at the transition 
zone of Isla Santa Cruz (1 specimen); and in epiphytic lichens and moss in a highland 
cloud forest of the same island (1 specimen). 

Sphaerochthonius windsori sp. n. Figs 15-26 

Material examined: Rep. Panama: Madden Dam (type locality: 9°12,5' N, 79°37,2' W, 
45 m a.s.l.), dry deciduous forest, in decayed leaf litter and humus under Bursera simaruba 
(8.Ü.1996: 7 adults, 1 trito-, 1 protonymph, H. Schatz coll.). IslaTaboga, along Sendero Las Très 
Cruces (9°33,5' N, 8°37,L W, 100 m a.s.l.), dry deciduous forest, in decayed litter with pieces 
of wood (24. i. 1996: 2 adults, H. Schatz coll.). The adult holotype and 2 adult paratypes from the 
same locality are preserved in alcohol and will be deposited in: Muséum d'histoire naturelle, 
Genève, Switzerland, other specimens in the author's collection in Innsbruck, Austria. 

Diagnosis: Adults of the new species differs from its congeners by the follow- 
ing combination of character states: surface structure on prodorsum and on major part 
of notogaster consisting of polygonal network surrounded by relatively large round and 
elevated areolae; nose-shaped protuberance present on prodorsum; except setae exp, d, 
all prodorsal and notogastral setae biramous, both rami almost equal in length or 
slightly asymmetric, ciliate to papillate; rami of notogastral setae e,f aligned in trans- 
verse direction; 8 pairs of genital, 4 pairs of anal and 4 pairs of biramous adanal setae 
present; seta /' on palptarsus with ciliae at right angle, setae (ul) fused, bifid. Juvenile 
instars: setae d very short, setiform, setae e and/ biramous, rami aligned in transverse 
direction, adanal setae setiform in protonymph, biramous in tritonymph. 

Adult (Figs 15-24): Total body size (n=9): 270 (255-295) x 188 (180-190), 
length of notogaster 181 (170-200, 64-71% of total body length). Color dark yellow to 
light brown. Prodorsal and major part of notogastral surface (except lateral plates L) 
covered with irregular polygonal network surrounded by smaller and larger elevated 
areolae, surface granulate (Fig. 18). Lateral plates L granulate, with longitudinal striae 
(Fig. 17). Mentum, epimeral, genital and ano-adanal plates with granulate surface. 

Prodorsum (Fig. 15): Shape almost quadrangular, anterior edge rounded. 
Rostrum medially with a nose-shaped protuberance, slightly protruding anteriorly 
beyond anterior margin of prodorsal shield. Rostral, lamellar, interlamellar (Fig. 19) 
and anterior exobothridial setae biramous, papillate ("T-shaped" in the sense of 
Wallwork, 1960). both rami almost equal in length, extension of both rami: ro 40-45, 
le 45-50, il 30-35, exa 12-15. Papillae of w, le, il 3-5 long. Posterior exobothridial 



NEW NEOTROPICAL SPHAEROCHTHONIUS 1 19 

setae (exp) lateral to bothridia, curved, uniramous, papillate, directed laterad to antero- 
dorsad, 10 long. Bothridia directed laterad, head of sensilli with anterodorsad directed 
papillae, total length of sensilli 50, head 40. Posterior edge of prodorsum sclerotized. 

Notogaster (Fig 15): Shape almost round. Broad transverse scissure (of type "L" 
in the sense of Grandjean, 1947; see Norton, 2001) present between plates Na and Py, 
curved anteriad. Anterior part of plate Py covered by hypertrophied posterior part of 
plate Na (overlapping part 27-50 long*), hiding insertions of setal row d on scissure. 
Indistinct thickened transverse suture present anterior to setal row e, not visible in all 
studied specimens. On ventral side a single transverse scissure present anterior to plate 
Py towards ano-adanal plate, and a straight longitudinal scissure present on each side 
between plates Is and L. 

Sixteen pairs of notogastral setae present. Setae d very short, setiform, smooth, 
2-3 long, hardly visible under plate Na. Other notogastral setae biramous, papillate. 
Rami of setal row c aligned in diagonal to longitudinal direction, slightly asymmetric 
with shorter anterior rami, extension of both rami 30-45, rami of setae e (Fig. 20) and 
/aligned transversely, extension of both rami 20-30, setae h r h 3 on edge of notogaster, 
extension of both rami 30-35, stta&ps r ps 3 visible in lateral and ventral view, extension 
of both rami: psj 25-30, ps 2 , ps 3 20-25, ps 2 and/?s ? inserting on plates Is lateral to ano- 
adanal plates. 

Gnathosoma (Figs 17, 22, 23): Mentum small (25 x 20-25), subcapitular setae 
attenuating, setae h inserting on posterior part of mentum, directed posteriad, h, m } , m 2 
5-10, a 15-20 long. Three pairs of adorai setae present, -10 long. Chelicerae (Fig. 22) 
narrow, size (n=4): 45-55 x 16-20, movable digit 13-17 long, both setae directed 
anteriad, cha setiform, cibate, 15-20 long, chb strongly thickened, palmate, 12-15 long, 
with 4-6 fingers. Palps (Fig. 23) with 5 short articles, total length 40-50. Setal formula 
(trochanter to tarsus, solenidion in parentheses) - 2 - 1 - 3 - 1 1(1), cm 50 long, acm 
flagelliform, with a distal loop, 30-35 long, setae (It) 15-20 long, seta /' pectinate with 
ciliae at right angle. Setae (ul) and sul eupathids, (ul) fused, bifid with two short spines. 

Epimeral region (Fig. 17): Epimeral plates I and II separate, plates III and IV 
almost completely fused on each side. Epimeral setal formula 3-2-3-4, setae 3c insert- 
ing on anteriorly directed apophyses, 4b, 4c, 4d in one row on posterior part of plate. 
Setae attenuating, on plates I and II ciliate, 10-15 long, on plates III and IV smooth, 6- 
10 long. 

Genito-anal region (Figs 17, 21): Genital plates small, shape oval, surface 
granulate, almost spiculate. Eight pairs of genital setae present, attenuating, 8-12 long. 
Anal and adanal plates fused. Four pairs of anal setae present (Fig. 21), basally 
thickened, distally attenuating, curved mediad and posteriad, 8-12 long. Four pairs of 
adanal setae present (Fig. 21), biramous with ciliae, extension of both rami 12-15, 
ciliae 2-3 long. Bases of adanal setae connected by a sclerotized ridge. 

Legs (Fig. 24): Legs of moderate length (without claws 35-46% of body length). 

Setal formula of legs (trochanter to tarsus, solenidia in parentheses): leg I 

(Fig. 24) - 4 - 4 - 6(1) - 19(3), leg II 1 - 5 - 4 - 6(1) - 17(2), leg III 2 - 3 - 3 - 4(1) - 

* 
The degree of overlap of the hypertrophied posterior part of plate Na depends on the way of 

preparation. When squeezed, the plates move apart and the transverse scissure becomes un- 
covered. In this position setae d are well visible. 



120 H. SCHATZ 

12, leg IV 2- 3 - 3 - 4(1) - 1 1. All legs heterotridactylous, with a small, but strong me- 
dial claw and two weaker lateral claws, claws 20-30 long. Setae d and I" on femur I 
and setae d of femora II and III uniramous, papillate, setae d on femur IV thickened 
with short ciliae. Solenidia coj on tarsus I curved anteriad, 25 long, solenidia cp ; on tib- 
ia I very long (70-90), inserted on large distal projection. 

Immatures (Figs 25, 26): Color pale. Size: protonymph (n=l): 190 x 130, noto- 
gaster 130 long, tritonymph (n=l): 260 x 170, notogaster 170 long. Surface structure 
of both instars as in adult, in protonymph surrounding areolae very small. 

Prodorsum (Fig. 25): Nose-shaped structure as in adult in both instars. Shape of 
prodorsal setae as in adult, extension of both rami ro, le 30-35, exa 12-15, exp 10 long 
in tritonymph; extension of both rami ro, le 20, il 15, exa 12, exp 7 long in protonymph. 
Total length of sensilli 50, head 40 in tritonymph; length of sensilli 25, head 20 in 
protonymph. 

Gastronotic region (Figs 25, 26): Broad dorsal scissure between plates Na and 
Py present as a well-developed sulcus. Lateral plates L triangular, surface granulate 
with radial striae, inner margin with a series of folds. In both instars sixteen pairs of 
gastronotic setae present, setae c r c p biramous, papillate, slightly asymmetric, exten- 
sion of both rami 25-35 in tritonymph; setae c r c p biramous, ciliate, slightly asym- 
metric, extension of both rami 15-20 in protonymph. Setae d on anterior slope of 
cavity, very short, setiform, smooth, 2 long in tritonymph; 1-2 long in protonymph, 
hardly visible in between the surface microstructure. Pygidial plate Py with two trans- 
verse ridges, setal row e on anterior, row/on posterior ridge, rami of setae e, /directed 
transversely in both instars, setae h and/?5 7 on lateral edge of notogaster, setae ps 2 and 
ps 3 on plate Is. All posterior setae biramous. papillate, extension of both rami 15-20 in 
tritonymph; biramous, ciliate, extension of both rami 10-15 in protonymph. 

Gnathosoma: Number of subcapitular and adorai setae of both instars as in 
adult. Size of chelicera 50 x 17 in tritonymph, setae cha 15, chb 10 long, palmate, with 
4 fingers in tritonymph; chelicera of protonymph not studied. Palp: setal formula of 
both instars of both instars as in adult, setae cm 45 long in tritonymph, 22 in proto- 
nymph, setae acm without loop in both instars, length 27 in tritonymph, 15 in proto- 
nymph, eupathidial setae (»/) fused, bifid, setae /' pectinate with ciliae at right angle in 
both instars. 

Ventral region (Fig. 26): Shape of epimeral plates, position, number and shape 
of epimeral setae in tritonymph as in adults, setae 5 long. Epimeral setal formula 2-1- 
2?-2? (not well studied due to folded legs) in protonymph, setae 4-5 long. Genital 
plates with 6 pairs of setae, setiform, 7 long in tritonymph; 1 pair of genital setae, seti- 
form to spiniform, 4 long in protonymph. Anal and adanal plates fused, 4 pairs of seti- 
form anal setae and 4 pairs of biramous, ciliate adanal setae present, extension of both 
rami 6-8 in tritonymph; no anal setae. 4 pairs of setiform adanal setae, 4-5 long in 
protonymph. 

Legs: All legs monodactylous, setal formula not studied in detail, setae d and I" 
on femur I, setae d on femora II - IV thickened, papillate in tritonymph; setae d on 
femora I and II thickened and ciliate in protonymph. Solenidion qp 7 on tibia I 80 long 
in tritonymph. 55 long in protonymph. 



NEW NEOTROPICAL SPHAEROCHTHONIUS 



121 





Figs 15-17 
Sphaerochthonius windsori sp. n., adult - 15. Dorsal view; 16. Lateral view; 17. Ventral view 
(Scale: 100 um). 



Eggs: Several females observed with one or two large eggs. Dimensions of eggs 
(n=6) 107-125 x 70-100, shape of eggs oval to elliptic. Eggs mostly in longitudinal 
orientation, except in one female in transverse orientation in posterior part of noto- 
gaster. Surface and edge of eggs appearing smooth inside the notogaster, but not 
studied in detail. 



side. 



Variation: One adult specimen from Isla Taboga with 9 genital setae on one 



122 



H. SCHATZ 



18 



I ■ ^ 



mmm 



19 




Figs 18-24 
Sphaerochthonius windsori sp. n., adult - 18. Surface structure of prodorsum and notogastral 
plates Na, Py; 19. Seta il; 20. Seta ef, 21. Anal and posterior adanal setae; 22. Chelicera; 23. 
Pedipalp; 24. Leg I. All appendages in abaxial aspect (Scale: 50 pm). 



Etymology: The new species is gratefully dedicated to Dr Donald M. Windsor, 
Smithsonian Tropical Research Institution, Rep. Panama, for his hospitality and 
continuous support of our expeditions in Panama. 

Remarks: The new species is similar to studied specimens of Sphaerochthonius 
splendidus (Berlese, 1904) in having a polygonal surface structure, biramous setae on 
prodorsum and notogaster, and in possessing the same number of anal and adanal setae. 
Differences between the two species are found in: presence of a nose-shaped structure 
on prodorsum in S. windsori, shape of notogastral setae c 2 , c 2 , rows e and f (windsori: 
biramous, subequal in length, splendidus: biramous, strongly asymmetric with short 
anterior rami), alignment of rami of setal rows e and / (windsori: transverse, splen- 
didus: longitudinal), details in surface structure (windsori: large and round elevated 
areolae, forming edges of polygonal network, splendidus: clearly outlined polygons, 
areolae absent or small). Sphaerochtonius splendidus has a wide circumtropical and 
-subtropical distribution and was also frequently found in Panama by the author. 

The tritonymphs of Sphaerochthonius windsori sp. n. differ from the adult instar 
by the smaller posterior edge of plate Na, leaving the scissure uncovered; by the 
presence of additional transverse ridges along setal rows e and /(the anterior is present 
as an indistinct suture or totally reduced in adults); by the number of epimeral, genital, 
and anal setae; and by the number of claws (see also remarks on S. litoralis sp. n.). 



NEW NEOTROPICAL SPHAER0CHTH0N1US 



123 




Figs 25-26 
Sphaerochthonius windsori sp. n., protonymph - 25. Dorsal view; 26. Ventral view (Scale: 
50 urn). 

Ecology and distribution: The few finds of Sphaerochthonius windsori sp. n. 
indicate a preference for arid conditions. The species was not recorded in other sites, 
although more than 300 samples were taken by the author in different regions and 
habitats in Panama (Schatz, 1997). 



Key to known Neotropical Sphaerochthonius species 

1 Setae of rows e,f, h on posterior notogastral plate uniramous, 4 pairs of 

adanal setae 2 

Setae of rows e,f, h on posterior notogastral plate biramous 4 

2 Setae of rows e,f, h on posterior notogastral plate long, notogaster with 

reticulate surface structure 3 

Setae of rows e,f, h on posterior notogastral plate short, fungiform, no- 
togaster with foveolate surface structure S.fungifer Mahunka, 1983 

(Guatemala, Tikal) 

3 Setae of rows e, f, h on posterior notogastral plate broad, phylliform 

S. phyllophorus Balogh and Mahunka, 1969 

(Brazil, Rio de Janeiro) 

Setae of rows e,f, h on posterior notogastral plate long, small, papillate 

S. Uruguay ensis Pérez-Inigo and Sarasola, 1998 

(Uruguay) 

4 Anterior notogastral setae (row c) biramous, T-shaped, both rami of 

almost equal length 5 

Anterior notogastral setae (row c) biramous, T-shaped, strongly asym- 
metrical, anterior ramus much shorter than posterior one, setae of rows 



124 H. SCHATZ 

e, f, h on posterior notogastral plate also asymmetrical, 4 pairs of 

adanal setae S. splendidus (Berlese, 1904) 

(Galapagos Islands, Central America, Holarctic region, Australia) 

5 5 pairs of adanal setae 6 

4 pairs of adanal setae, notogaster with irregular reticulate surface struc- 
ture surrounded by elevated areolae S. windsori sp. n. 

(Panama) 

6 Prodorsum with nose-shaped protuberance, reticulate surface structure 
on notogaster with small triangles in angles, adanal setae widely 

branching S. litoralis sp. n. 

(Galapagos Islands) 

Prodorsum without nose-shaped protuberance, reticulate surface struc- 
ture on notogaster irregularly surrounded by areolae, adanal setae bira- 

mous, inner rami short S. transversus Wallwork, 1960 

(Ghana; records in Neotropical region doubtful) 

ACKNOWLEDGEMENTS 

Logistic support by the Charles Darwin Research Station, the Servicio Parque 
Nacionâl Galapagos, Ecuador, the Smithsonian Tropical Research Institution, Rep. 
Panama, and the Institute of Zoology and Limnology, Leopold-Franzens-University of 
Innsbruck, Austria, is gratefully acknowledged, as are the collecting permits from the 
authorities of Ecuador and Panama. Dr R.A. Norton, Syracuse, N.Y., USA, provided 
constructive criticism of the manuscript. 

REFERENCES 

Balogh, J., Mahunka, S. & Zicsi, A. 1969. The scientific results of the Hungarian Soil 
Zoological expedition to South America. 14. A report on the collectings of the second 
expedition. Folia Entomologica Hungarica (series nova) 22: 453-474. 

Grandjean, F. 1934. La notation des poils gastronotiques et des poils dorsaux du propodosoma 
chez les Oribates (Acariens). Bulletin de la Société zoologique de France 59: 12-44. 

Grandjean, F. 1947. Les Enarthronota (Acariens). Première série. Annales des Sciences na- 
turelles. Zoologie II e série, 8: 213-248. 

Hammen, L. van der 1959. Berlese 's primitive Oribatid mites. Zoologische Verhandelingen 
Leiden 40: 1-93. 

Hunt, G. S., Norton, R. A., Kelly, J. P. H., Colloff, M. J. & Lindsay, S. M. 1998. An interac- 
tive glossary of oribatid mites. CSIRO Publishing Melbourne, CD-ROM. 

Norton, R. A. 2001. Systematic relationships of Nothrolohmanniidae, and the evolutionary plas- 
ticity of body form in Enarthronota (Acari: Oribatida) (pp. 58-75). In: Halliday, R. B., 
Walter, D. E., Proctor, H. C, Norton, R. A. & Colloff, M. J. (eds). Acarology: 
Proceedings of the 10th International Congress. CSIRO Publishing, Melbourne. 

Schatz, H. 1997. Oribatid mites from the northern neotropical region - a survey of research, past 
and present. Abhandlungen und Berichte des Naturkundemuseums Görlitz 70: 61-70. 

Schatz, H. 1998. Oribatid mites (Acari: Oribatida) from the Galapagos Islands - Faunistics, 
Ecology and Speciation. Experimental and Applied Acarology 22: 373-409. 

Schatz, H. 1999. Oribatid mites of the Galapagos Islands: Faunistics, ecology and speciation 
(pp. 499-530). In: Bruin, J., van der Geest, L. P. S. & Sabelis, M. W. (eds). Ecology 
and evolution of the Acari. Kluwer Academic Publishers, Dordrecht. 

Wallwork, J. A. 1960. Some Oribatei from Ghana. I. Sampling localities. II. Some members of 
the Enarthronota Grandj. Acarologia 2: 368-388. 



Revue suisse de Zoologie 1 10 (1): 125-128; mars 2003 



Una nuova specie di Otiorhynchus Germar, 1822 del Pakistan 
(Coleoptera, Curculionidae, Polydrusinae) * 

Luigi MAGNANO 

Via Montenero, 53, 1-53036 Poggibonsi SI, Italia. 



A new species of Otiorhynchus Germar, 1822 from Pakistan (Coleo- 
ptera, Curculionidae, Polydrusinae). - Otiorhynchus (Melasemnus?) 
lepidus sp. n. is described, belonging to the section 3 of the system proposed 
recently by the author. The epipleura slightly bent at the level of the hind 
coxae do not make possible a satisfactory placement, since the new species 
has no affinity with any subgenus of this section. O. lepidus shares the above 
character with O. relicinus Arnoldi, 1975 from Altai. The new species may 
belong to a new genus which has not been described waiting for a more 
abundant material from the areas at the border of the known range of 
Otiorhynchus. 

Key- words: New species - Curculionidae - Polydrusinae - Otiorhynchus - 
Pakistan. 

INTRODUZIONE 

Fra il materiale recentemente studiato del Muséum d'histoire naturelle di 
Ginevra (Svizzera) (MHNG) ho trovato una nuova specie che provvisoriamente ascrivo 
al genere Otiorhynchus e qui di seguito descrivo. 

MATERIALI E METODI 

Il presente studio si basa sull'esame di cinque esemplari del Pakistan, conser- 
vati nella collezione entomologica del Museum d'histoire naturelle di Ginevra 
(MHNG). 

Otiorhynchus (Melasemnus?) lepidus sp. n. Figg. 1-5 

Materiale esaminato: Holotypus S, Pakistan, Swat, Kalam, 2100 m, 17. v. 1983, 
Besuchet-Löbl (MHNG). Paratypi: 1 ó\ Pakistan, Swat, Kalam, 2100 m, 17.V.1983, Besuchet- 
Löbl; 1 ó\ 1 9, Pakistan, Swat, vallée d'Ushu, 2300 m, 15.V.1983, Besuchet-Löbl, (MHNG). 1 
paratypus 6, Pakistan, Swat, Kalam, 2100 m, 17.V.1983, Besuchet-Löbl nella collezione 
dell'autore. 

Diagnosi. Un Otiorhynchus appartenente alla sezione 3 dell'attuale sistema 
(Magnano, 1998), ma con epipleure curve al livello delle anche posteriori e con 
nessuna affinità con i sottogeneri ad essa pertinenti. La specie è ascritta provvisoria- 
mente al sottogenere Melasemnus. 



* XLVI contributo alla conoscenza dei Curculionidi. 
Manoscritto accettato il 06. 1 1 .2002 



126 L. MAGNANO 

Descrizione dell'holotypus â. Lunghezza (protorace più elitre): 5,5 mm; 
larghezza massima delle elitre: 2,6 mm. Colore castano. Rostro, compresi gli pterigi 
arrotondati all'esterno, lungo quanto largo; epistoma subtriangolare, fronte ripiegata in 
avanti e appena distinta dall' epifronte, la quale ha i lati subparalleli ed è larga quanto 
il rostro al livello della inserzione delle antenne. Scrobe semiaperte in avanti, all'in- 
dietro terminanti alla metà della distanza fra gli occhi e l'inizio della dilatazione delle 
scrobe. Rostro separato dal capo da una lieve depressione trasversale davanti gli occhi. 
Capo due volte più largo che lungo, occhi grandi: il loro diametro maggiore è appena 
più stretto della distanza interoculare. Epifronte percorsa lungo la linea mediana da una 
carena appena accennata. Scultura del capo e del rostro costituita da granuli con 
un'area centrale che occupa la parte apicale, dov'è inserita una setola lunga tre volte il 
suo diametro. Scapo diritto, di grossezza pressoché uguale dalla base fino ai 2/3, poi 
ingrossato a clava; primo antennomero due volte più lungo che largo all'apice, clavi- 
forme; secondo 3,5 volte più lungo che largo all'apice e appena più ingrossato nella 
parte apicale; terzo 1,5 volte più lungo che largo all'apice; 4°-7° 1,3 volte più lunghi 
che larghi e tutti a tronco di cono. Clava fusiforme allungata, poco più lunga degli 
ultimi quattro antennomeri e 3,3 volte più lunga che larga, articolo basale a forma di 
cono. Protorace 1,2 volte più largo che lungo, regolarmente arrotondato ai lati, orlo 
anteriore appena più stretto dell'orlo basale, scultura con granuli ombelicati piccoli e 
fitti un po' più piccoli ai lati, sul disco misti ad aréole nella metà apicale. Pubescenza 
come sul capo e rostro, diretta verso la linea mediana. Elitre ovali, con la massima 
larghezza nel terzo basale; strie composte da una serie di aréole piccole ma ben evi- 
denti, separate da un granulo della metà più piccolo del loro diametro. Interstrie del 
doppio più larghe delle strie, con granuli un po' più grandi di quelli del pronoto e 
leggermente schiacciati sul disco; le interstrie laterali e le altre sulla declività posteriore 
con granuli convessi poco fitti e disposti in una serie. Epipleure appena smarginate al 
livello delle anche posteriori. Vestitura composta da setole all'interno delle aréole aven- 
ti la loro stessa lunghezza. Interstrie con due serie di setole curve, appena sollevate dal 
tegumento e appena più lunghe di quelle del protorace, del rostro e del capo. Femori 
clavati, con un piccolo dente spiniforme, nei profemori seguito da uno o due granuli 
minuti; orlo esterno delle protibie retto nei 2/3 basali, poi leggermente curvo all'inter- 
no, parte interna bisinuosa, la prima concavità termina nei 2/3 basali, quella apicale 
porta tre spine, i loro intervalli crenulati. Meso e metatibie come le protibie, ma le in- 
cisure e i denti meno evidenti. Metatibie leggermente curve all'interno. Tarsomeri 1 e 
2 di uguale lunghezza, terzo profondamente bilobo, onichio lungo il doppio del terzo 
tarsomero, unghie libere. Metasterno e urosterni con aréole fitte e profonde, piccole, 
distanti fra loro della metà del loro diametro sulle quali è inserita una setola 3 volte più 
lunga del loro diametro. Metasterno e primo urosterno apparente profondamente inca- 
vati. Edeago e habitus come in figg. 1, 2, 4, 5. 

I paratipi non si differenziano sostanzialmente dall'olotipo. La 9 ha le elitre più 
larghe. Spiculum ventrale come in fig. 3. 

Osservazioni. O. lepidus non presenta affinità con alcuna specie nota. Al taxon 
è quindi stata data una collocazione assolutamente provvisoria perché esso, pur 
presentando i caratteri della terza sezione dell'attuale sistema (Magnano, 1998), ha le 
epipleure leggermente curve al livello delle anche posteriori, carattere che appartiene 



UNA NUOVA SPECIE DI OTIORHYNCHUS 



127 









FiGG. 1-6. - 1-5: Otiorhynchus (Melasemnus? ) lepidus sp. n. 1: edeago in visione dorsale; 
2: edeago in visione laterale; 3: spiculum ventrale; 4: 6 , habitus, schematico; 5: habitus. - 6: 
Otiorhynchus relicinus habitus, schematico, da Arnoldi, 1975. 



128 L. MAGNANO 

agli altri generi del complesso Otiorhynchus Germar, 1822 e alle sezioni 1, 2 e 4. La 
nuova specie potrebbe appartenere ad un nuovo genere che per il momento, per 
scarsezza di materiale, si è preferito non descrivere. Anche Otiorhynchus relicinus 
Arnoldi, 1975 dell'Aitai ha le epipleure leggermente curve al livello delle anche pos- 
teriori. Si noti che Arnoldi (1975) non ha collocato la specie in nessun gruppo allora 
descritto. Vale la pena notare che queste specie sono presenti nella parte centro-meri- 
dionale dell'area di distribuzione del genere Otiorhynchus. La tribù Otiorhynchini, ed 
in particolare il genere Otiorhynchus, necessitano di una profonda revisione per fare un 
minimo di chiarezza nella loro sistematica. A questo riguardo la fauna delle regioni 
himalayana e centroasiatica meritano una particolare attenzione perché possono 
portare un notevole contributo alla conoscenza della tribù. Il carattere delle epipleure 
leggermente curve al livello delle anche posteriori condiviso da O. relicinus (fig. 6) e 
O. lepidus (fig. 4) è una pura convergenza. Non conosco in natura O. relicinus, ma il 
confronto delle figg. 4 e 6 rende evidente il fatto che si tratta di due entità completa- 
mente differenti appartenenti probabilmente a due generi diversi. 

Derivatio nominis. Lepidus = grazioso. 

Distribuzione geografica: Pakistan nord orientale. 

BIBLIOGRAFIA 

Arnoldi, L. V. 1975. Weevils of the genus Otiorhynchus Germar (Coleoptera, Curculionidae) of 
Mongolia and of adjacent regions of the USSR. Nasekomye Mongola 6(3): 254-284 (in 
russo). 

Magnano, L. 1998. Notes On The Otiorhynchus Germar, 1824 Complex. (Coleoptera: 
Curculionidae). In: Colonnelli, E., Low, S. & Osella, G. (eds). Taxonomy, ecology 
and distribution of Curculionoidea (Coleoptera: Polyphaga). Proceeding of a 
Symposium (22 August, 1996, Florence, Italy). XX International Congress of 
Entomology. Museo Regionale di Storia Naturale, Torino: 51-80. 



Revue suisse de Zoologie 1 10 (1): 129-132; mars 2003 



Two new species of the genus Quedius from Algeria and Pakistan 
(Coleoptera: Staphylinidae: Staphylinini: Quediina) 

Petr STOURAC 

Taussigova 1169, CZ-182 00 Praha 8-Kobylisy, Czech Republic. 



Two new species of the genus Quedius from Algeria and Pakistan 
(Coleoptera: Staphylinidae: Staphylinini: Quediina). - The new species 
Quedius (Micwsaurus) asphaltinus from Pakistan, and Quedius (Raphirus) 
boreoafricanus from Algeria are described, illustrated and compared to 
related species. 

Key-words: Taxonomy - new species - Coleoptera - Staphylinidae - Algeria 
- Pakistan. 

INTRODUCTION 

The genus Quedius Stephens, 1829 comprises almost 800 species occuring in 
all zoogeographical regions of the world. 

Palaearctic species (about 500) live usually in forest floor litter, leaf litter, 
around bases and in hollows of old trees, under hay heaps etc. Some species prefer 
caves, watersides, swamps, ant hills and underground nests of insects or various little 
mammals. 

Two new species were found among the material from the collection of the 
Muséum d'histoire naturelle Genève and from some private collections. The descrip- 
tions follow below. All measurements in the descriptions are given in millimeters. 

TAXONOMICAL PART 

Quedius (Microsaurus) asphaltinus sp. n. Figs 1-2 

Type material. Holotype $ : "Pakistan, NW Frontier Prov., Nathiagali (20 km N Murree), 
34.04 N/73.24 E, 2400 m, J. Kalâb leg., 9.-ll.viii.2000". In the collection of the National 
Museum Prague, Czech Republic. 

Description. Female. Length 12 mm. Black, abdomen inconspicuously irides- 
cent; front tibia, apex of tarsi as well as apex of abdomen somewhat paler. 

Head of obtusely quadrangular shape, wider than long (length/width= 
1,81/1,97), slightly arcuately narrowed to neck, posterior angles indistinct; frons 
behind antennal insertions with shallow triangular impression. Eyes small and rather 
flat, hardly protruding from lateral contours of head, tempora longer than eyes seen 
from above (0,81/0,66); no additional setiferous punctures between anterior frontal 
punctures; posterior frontal puncture situated closer to posterior margin of eye than to 
posterior margin of head, two smaller consecutive setiferous punctures between it and 



Manuscript accepted 24.07.2002 



130 P- STOURAC 

posterior margin of head; temporal puncture closer to posterior margin of head than to 
posterior margin of eye, no fine punctures behind temporal puncture; surface of head 
with very fine and dense microsculpture of mostly transverse waves and with scattered 
sparse micropunctulation. 

Antenna moderately long, segment 3 longer than segment 2 (0,34/0,25), 
segment 4 and 5 about as long as wide, segments 6-10 wider than long (gradually 
becoming wider and shorter), segment 1 1 slightly shorter than two preceding segments 
combined. 

Pronotum wider than long (length/width=2, 19/2,44) widest behind middle, 
basal margin broadly rounded, lateral portions not explanate, distinctly narrowed an- 
teriad; dorsal rows each with three punctures, sublateral rows each with two punctures, 
posterior puncture situated distinctly before level of large lateral puncture; surface of 
pronotum with microsculpture similar to that on head, micropunctulation finer. 

Scutellum impunctate, surface with very dense and fine microsculpture of trans- 
verse waves, apical half with some scattered micropunctures. 

Elytra about as long as wide (length/width=2,69/2,66), at base narrower than 
pronotum at widest point, slightly widened posteriad, at sides distinctly longer 
(2,56/2,19), at suture hardly longer (2,31/2,19) than pronotum at midline; punctation 
coarse, sparse and markedly unevenly dispersed, interspaces glossy with scattered 
points (Fig. 1). 

Abdomen with tergite 7 bearing a distinct, whitish apical seam of palisade setae; 
punctation of tergites finer and much denser than that on elytra, rather uniform toward 
apex of abdomen. Genital segment with tergite 10 short and wide, markedly narrowed 
toward apex (Fig. 2). 

Pubescence of elytra and abdomen piceous black, uniform. 

Front tarsi slightly dilated, segment 2 about as wide as apex of tibia. 

Male. Unknown. 

Comparison. Quedius asphaltinus belongs by the body shape, the chaetotaxy of 
the head and the pronotum and the dark colouration near Q. (Micro saurus) mesome- 
linus (Marsham, 1802). It can easily be distinguished by its larger size and by the 
conspicuous punctation of the elytra. 

In the Near East occurs another robust species with very sparse punctation of 
elytra - Q. fagelianus Coiffait, 1967. It belongs to the subgenus Distichalius Casey, 
1915 (elytra with three irregular longitudinal rows of punctures only). It differs from 
Q. asphaltinus apart from the subgeneric characters given above, mainly as follows: 
palpi, antennae and legs paler, mostly rufobrunneous; head more transverse (1/w ratio 
about 0,75); only one puncture between posterior frontal puncture and posterior margin 
of head. 

Etymology. The specific epithet refers to the deep dark colouration and the 
relatively shining surface of the body. 

Quedius (Raphirus) boreoafricanus sp. n. Figs 3-6 

Type material. Holotype 6: "Algérie: Atlas de Blida, Chréa, Les Glacières, 1100 m. 
3.V.1988. Besuchet-Löbl-Burckhardt". In the collection of Muséum d'histoire naturelle Genève, 
Switzerland. 



TWO NEW SPECIES OF QUEDIUS 



131 








Figs 1 -6 
1 - 2: Quedius (Microsaurus) asphaltinus sp. n. (holotype 9 ): 1 - punctation of left elytra, 2 - fe- 
male tergite 10. 3 - 6: Quedius (Raphirus) boreoafricanus sp. n. (holotype 6): 3 - median lobe 
in lateral view, 4 - median lobe in ventral view, 5 - underside of paramere, 6 - apical emargina- 
tion of male sternite 8. 



Description. Male. Length 6,5 mm. Head dark brown; pronotum piceous brown 
with lateral portions and anterior angles paler; elytra testaceous, suturai area narrowly 
darkened; abdomen dark brown, iridescent. Antennae and palpi testaceorufous; bases 
of antennae, legs and apical portions of abdominal tergites testaceous. 

Head rounded, about as long as wide (length/width=0,88/0,89); eyes large and 
convex, tempora much shorter than length of eyes seen from above (0,17/0,56); no 
additional setiferous punctures between anterior frontal punctures; posterior frontal 
puncture very close to posteromedian margin of eye, one puncture between it and 
posterior margin of head; temporal puncture touching posterior margin of eye, tempora 
with some fine punctures; surface of head with dense and very fine microsculpture of 



132 P- STOURAC 

transverse waves gradually becoming irregular toward clypeus and forming numerous 
meshes there. 

Antenna moderately long, segment 3 inconspicuously longer than segment 2 
(0,19/0,16), segments 4 and 5 somewhat longer than wide, following segments 
gradually becoming shorter, segments 7-10 about as long as wide, segment 11 slightly 
shorter than two preceding segments combined. 

Pronotum as long as wide (length/width=l,19/l,19), broadly rounded basally, 
slightly narrowed anteriorly; dorsal and sublateral rows each with three punctures, 
posterior sublateral puncture situated behind level of large lateral puncture; surface of 
pronotum with microsculpture similar to that on head. 

Scutellum impunctate, surface with dense and fine microsculpture of transverse 
waves. 

Elytra as long as wide (length/width= 1,44/1 ,44), at base hardly narrower than 
pronotum at widest point, slightly widened posteriad, at sides longer (1,38/1,19), at 
suture about as long as pronotum at midline; punctation fine and somewhat not 
uniform, longitudinal interspaces between punctures two times larger than diameters of 
punctures, transverse interspaces about as wide as diameters of punctures, surface 
between punctures without microsculpture. 

Abdomen from tergite 4 narrowed toward apex, tergite 7 with distinct, whitish 
apical seam of palisade setae; punctation of tergites finer and slightly denser than that 
on elytra. Sternite 8 with triangular emargination (Fig. 6), shape of tergite 10 and 
sternite 9 very similar to that of other species in the Q. nemoralis X humeralis-group. 

Pubescence of elytra and abdomen brownish and uniform. 

Front tarsi slightly dilated, segment 2 about as wide as apex of tibia. Right 
middle tibia with tarsi is missing. 

Aedeagus relatively narrow and elongate, his shape in Figs. 3-5. 

Female. Unknown. 

Comparison. The new species belongs to the Q. nemoralis X humeralis-group, 
but only two species of it occur in North Africa - Q. humeralis Stephens, 1832 and 
Q. pauliani Koch, 1941. The aedeagus of Q. pauliani is very similar to that of 
Q. nemoralis Baudi, 1848 (median lobe spoonshaped in ventral view). The aedeagus of 
Q. boreoafricanus resembles more that of Q. humeralis, but it differs mainly in the very 
long apical keel of the median lobe (about 2,4 times longer than that of Q. humeralis); 
the apical portion of median lobe is narrow, but not pointed in ventral and lateral views 
and the paramere is slightly wider and less pointed (see Coiffait 1978, Fagel 1968). 

Etymology. The specific epithet refers to the occurrence of the species in North 
Africa. 

REFERENCES 

Coiffait, H. 1978. Coléoptères Staphylinides de la région paléartique occidentale III. Sous- 
famille Staphylininae, Tribu Quediini. Sous famille Paederinae, Tribu Pinophilini. 
Supplément à la Nouvelle Revue d'Entomologie VIII (4): 364 pp. 

Fagel, G. 1968. Contribution à la connaissance des Staphylinidae. CHI. - Le complexe des 
Quedius oblitteratus Er. - nemoralis Baudi. Bulletin de l'Institut royal des Sciences 
naturelles de Belgique AA (24): 1-14. 



Revue suisse de Zoologie 110 (1): 133-139; mars 2003 



Zoogeography in southern Japan as revealed by ground-living 
arachnids 

Joachim HAUPT 

Institute of Ecology / Biology, Technical University Berlin, Franklinstr. 28/29, FR 1-1, 

D-10587 Berlin, Germany. 

E-mail: hptjeiic@sp.zrz.tu-berlin.de 



Zoogeography in southern Japan as revealed by ground-living 
arachnids. - In the past, primarily birds and butterflies, but also amphibians, 
reptiles and mammals, have been used for Zoogeographie studies in 
southern Japan. Meanwhile, taxonomic studies have also been carried out 
on arthropod groups living in and on the soil. Since it is quite unlikely that 
these ground-living organisms have crossed the sea, either actively or 
passively, they are ideal for Zoogeographie studies. The present invest- 
igations are based on four groups of ground-living arachnids: whipscorpions 
and spiders (Hexathelidae, Ctenizidae, Mesothelae). A close Zoogeogra- 
phie al connection was found between the Yaeyama Islands and nearby 
Taiwan. The central and northern parts of the Ryukyu Islands are faunisti- 
cally closest to Kyushu. No distinct border between an Oriental and a 
Palaearctic fauna can be detected. Instead, only climatic and other ecolo- 
gical factors appear to be responsible for distributional boundaries of 
various organisms observed in different regions of southern Japan. 
Similarities to the fauna of continental China can be explained by land 
connections between the Ryukyus and the continent during the Pleistocene. 

Key -words: Arachnida - zoogeography - Uropygi - Mesothelae - Hexathe- 
lidae - Ctenizidae - Japan - Taiwan. 

INTRODUCTION 

According to differences in the possibilities of distribution of species concer- 
ned, conclusions can be drawn about their origin from geographic réfugia during the 
ice ages. More easily distributing species can expand their ranges more quickly and 
effectively. With every typhoon, flying species like birds or butterflies are brought 
north, both welcomed by excited birdwatchers and butterfly collectors, but in general 
none of these stray species is able to establish permanent footholds farther north. This 
is certainly due to climatic and maybe also other ecological reasons. There may be no 
more suitable free ecological niches available. 

Several attempts have been made to establish Zoogeographie lines in southern 
Japan, mostly based on the distribution of insects (the Miyake Line south of Kyushu) 



Manuscript accepted 26.06.2002 



134 J- HAUPT 

(Ezaki, 1929), on the distribution of amphibians, reptiles and mammals (the Watase- or 
Aoki Line at the Tokara trench) (Okada & Koba, 1931; Hikida et al, 1992; Hikida & 
Ota, 1997) or on the distribution of insects and birds (the Hachisuga Line at the Kerama 
trench) (Paik, 1953). Such efforts date back to Watase (1912), who distinguished three 
Zoogeographie areas in the region: 1) Taiwan (and the Ryukyus), 2) the region of 
Satsuma extending from Kyushu up to central Honshu, and 3) northern Honshu and 
Hokkaido. 

While previous studies have concentrated on the groups of animals mentioned 
before, species living in and on the ground are much less likely to migrate or to be dis- 
persed passively. Consequently, they are better suited to mirror the prehistoric biogeo- 
graphic situation. For this reason, different groups of arachnids, which were revised 
taxonomically during recent years, were used for the current study. 

MATERIAL AND METHODS 

Based on taxonomic studies of Uropygi (see Haupt & Song, 1996), Mesothelae 
(see Haupt, 1983; Song & Haupt, 1984), the hexathelid genus Macrothele (see 
Shimojana & Haupt, 1998) and the ctenizid genus Latouchia (see Haupt & Shimojana, 
2001), distribution data have been compiled in order to elucidate the Zoogeographie 
situation in southern Japan. Specimens mentioned are partly deposited in the Muséum 
d'histoire naturelle of Geneva. 

RESULTS 

The following species* were used for the current study; their distribution as 
documented by the current knowledge is summarized in fig. 1 . 

Uropygi: Typopeltis stimpsoni (Wood, 1862) (Amamioshima) (Haupt & Song, 
1996), T. stimpsoni (Amamioshima) (Zoological Museum Moscow, unpubl.), T. stimp- 
soni (Kyushu) (Yoshikura, 1965), T. stimpsoni (Izena) (Ikehara & Shimojana, 1975). 

Typopeltis crucifer Pocock, 1894 (Okinawa and Iheya) (Ikehara & Shimojana, 
1975), T. crucifer (Ishigaki and Iriomote) (Haupt & Song, 1996), T. crucifer (Taiwan) 
(Haupt & Song, 1996 and Hungarian National Museum Budapest, unpubl.), T. crucifer 
(Kashoto = Ludao, Green Island) (Haupt & Song, 1996 and Hungarian National 
Museum Budapest, unpubl.), T. crucifer (Lanyu = Orchid Island) (J. Haupt, unpubl.). 

Mesothelae: Heptathela kimurai kimurai (Kyushu) (Haupt, 1983), H. kimurai 
amamiensis (Amamioshima) (Haupt, 1983), H. kimurai yanbaruensis (Yanbaru, north- 
ern Okinawa) (Haupt, 1983), H. kanenoi (Tokunoshima) (Ono, 1996), H. kikuyai, H. 
nishikawai, H. yaginumai (all Kyushu), H. yakushimaensis (Yakushima) (Ono, 1998). 

Ryuthela* nishihirai nishihirai (central and southern Okinawa) (Haupt, 1983), 
R. nishihirai ishigakiensis (Ishigaki and Iriomote) (Haupt, 1983), R. sasakii 
(Kumejima), R. secundaria (Kumejima), R. owadai (Tokashiki, Kerama), R. tanikawai 
(Iriomote) (Ono, 1997). 



* Taxonomic note: In the current catalogue of spiders (Platnick, 2002) Ryuthela is still 
synonymized with Heptathela. On the other hand I do not support the description of new species 
instead of subspecies from various islands in the region, for this reason names of Heptathela and 
Ryuthela have been used as published originally. 



ZOOGEOGRAPHY IN SOUTHERN JAPAN 



135 



Honshu 




Kyushu 
■MiyakeLine 



TO^Wafase Line 
Lsv"H Amamioshima 



^ 






•\ 




Ls°^?\ Okinawa 



-HachisugaLine 




Lf^x. Miyako 

Tc 

™& Yaeyama Isl. 



Taiwan 



"?9 Jc 

Lanyu 



Fig. 1 
Distribution of some ground-living arachnids in the Ryukyus and their neighbourhood. Bars 
indicate the distribution borders between the whipscorpions Typopeltis stimpsoni (Ts) and 
Typopeltis crucifer (Tc) (Uropygi: Thelyphonida), Heptathela (H) and Ryuthela (R) (Araneae: 
Mesothelae) and subspecies of the ctenizids Latouchia swinhoei (Ls) and subspecies of 
Latouchia formosensis (Lf). This border line coincides with the northern distribution of the hexa- 
thelid Macrothele gigas (Mg) which is also found in Taiwan. The position of Latouchia japon- 
ica (Lj) remains questionable as the male is still unknown. Note that none of the distribution 
boundaries coincides with zoogeographical lines established previously. 



136 J- HAUPT 

Ctenizidae: Latouchia formosensis formosensis (Taiwan) (Haupt & Shimojana, 
2001), L. formosensis hyla (Kuroshima, Iriomote) (Haupt & Shimojana, 2001), L. 
swinhoei swinhoei (Okinawa) (Haupt & Shimojana, 2001), L. swinhoei xena 
(Amamioshima, Tokunoshima) (Haupt & Shimojana, 2001), L. swinhoei aypta 
(Kyushu) (Haupt & Shimojana, 2001), L. swinhoei typica (Honshu) (Haupt & 
Shimojana, 2001). 

Hexathelidae: Macrothele gigas (Ishigaki, Iriomote, Taiwan) (Shimojana & 
Haupt, 1998; Tso, pers. comm.). 

DISCUSSION 

Populations isolated on different islands certainly make their own progress 
towards speciation. During recent years several new species, e.g., of Mesothelae, have 
been described from the Ryukyus (Ono, 1996, 1997), showing that Heptathela and 
Ryuthela are also present on additional islands such as Tokara, Kerama or Kumejima, 
respectively. In fact, the description of new island species or the upgrading of sub- 
species to species level does not affect our interpretation: In this context it is not 
important whether an insular population has already obtained independent species 
status or not, because we have to look at the phylogenetic relationships documented by 
common derived morphological (synapomorphic) characters. Here, we concentrate on 
genera as mesothelid spiders are concerned, as this clarifies the situation more 
distinctly. 

The continental Chinese mesothelid species are clearly separated from the 
Japanese and Ryukyuan ones (Song & Haupt, 1984). Although Ono & Nishikawa 
(1989) tried to link the Japanese Heptathela kimurai amanuensis Haupt, 1983 to the 
Chinese H. hangzhouensis Chen, Zhang et Zhu, 1981, the former is more closely 
related to neighbours in the north (Kyushu) and in the south (Okinawa: Yanbaru), as it 
shares synapomorphic characters with them. This evaluation is based on the mor- 
phology of the female genital plate with its ventrolateral depressions, and on the 
different parts of the male palpal organ (Haupt, 1983, Song & Haupt, 1984). Therefore 
the phylogenetic relationships between mesothelid populations on Kyushu and in the 
northern Ryukyus, including Northern Okinawa (Yanbaru), have to be considered as 
being closer within each other than with species on the continent (Haupt, 1990). This 
fact supports the hypothesis that the connection to the continent broke up earlier than 
the connection between the northern and central Ryukyu islands and Kyushu, which is 
in accordance with geological data (Kimura, 1996). 

Since mesothelid spiders have not yet been recorded from Taiwan (Chen, 1996), 
they cannot be used to determine the zoogeographical relationship between the 
Yaeyama Islands (especially Ishigaki and iriomote) and Taiwan. For this purpose 
mygalomorph spiders of the families Ctenizidae and Hexathelidae are suitable (fig. 1). 
They prove a closer zoogeographical connection between Yaeyama and Taiwan. Such 
relationship (between Yaeyama and eastern Taiwan) has also been reported for the frog 
Rana limnocharis Wiegmann, 1835 (Toda, 1999). In fact these observations are not 
surprising, as the Yaeyama islands are located rather close to Taiwan. It is known that 
populations on islands which are in close proximity are also more closely related to 



ZOOGEOGRAPHY IN SOUTHERN JAPAN 137 

each other than populations from islands farther apart (Mac Arthur & Wilson, 1963). 
In this context it is surprising that Kimoto (1982), who worked on Chrysomelid bee- 
tles, also included Yaeyama as the southernmost Ryukyus, when he stated a close con- 
nection between Ryukyus and Kyushu. 

While the trench between Okinawa and Miyako (Hachisuga Line) has often 
been considered as an important dividing line, our results show that this is obviously 
not the case as far as the arachnids examined are concerned (fig. 1). Ryuthela (Meso- 
thelae) and Typopehis crucifer (Uropygi) are transgressing this line concerning their 
distribution. The distribution of Typopehis stimpsoni and of Latouchia swinhoei clearly 
points to the fact that also the Watase Line (Tokara trench) as well as the Miyake Line 
have not played any role in the dispersal of these species. 

Paik (1953) tried to establish a special biogeographic delimination linked to the 
northern distribution of Heptathela (Aso Line on Kyushu), but Latouchia occurs as far 
north as Tokyo. As both genera, Heptathela and Latouchia, share the same lifestyle as 
trapdoor spiders, this clearly demonstrates the climatic reasons for differences in 
northern distribution. Latouchia may be slightly better adapted to colder climate than 
Heptathela. 

Furthermore, the northern and southern distribution boundaries of arachnid 
species studied do not coincide. The distribution border between Ryuthela and Hepta- 
thela is found in the northern part of Okinawa (Yanbaru) and not between separate 
islands, although some low-lying parts of Yanbaru were possibly submerged during 
certain interglacial periods. 

Obviously, climatic differences present on the continent are also manifested in 
the Ryukyu Islands. This is apparently the only possibility to explain the current 
distribution pattern in various groups of ground-living arachnids. Similarities in the 
species spectrum between the Ryukyus and the continent can be explained by the 
pleistocene land connection (Kimura, 1996). Since then the islands were cut off and its 
fauna undertook a separate development. 

CONCLUSION 

To summarise, all these data provide evidence for a transient zone and suggest 
that there is no distinct border line between the Oriental and the Palaearctic fauna in 
the Ryukyu Islands of southern Japan. Biogeographic lines which were established be- 
fore and partly mark geological trenches are not supported by the distribution of 
ground-living arachnids. Their distribution mirrors climatic factors. 

ACKNOWLEDGEMENTS 

I am gratefully indebted to Dr Jason Dunlop for looking through the English 
text, and to Dr P. Schwendinger for critical remarks. 



138 J- HAUPT 



REFERENCES 

Chen, S.H. 1996. A checklist of spiders in Taiwan (in Chinese). Annals of the Taiwan Museum 

39: 123-156. 
Esaki, T. 1929. Zoogeographische Beziehung der Insel Yakushima und Kyushu nach Verbreitung 

der Tagfalter (in Japanese). Bulletin of the Biogeographical Society of Japan 1 (2): 47- 

56. 
Haupt, J. 1983. Vergleichende Morphologie der Genitalorgane und Phylogenie der liphis- 

tiomorphen Webspinnen. I. Revision der bisher bekannten Arten. Zeitschrift für zoolo- 
gische Systematik & Evolutionsforschung 21: 275-293. 
Haupt, J. 1990. Comparative morphology and phylogeny of liphistiomorph spiders (Araneida: 

Mesothelae). III. Provisional diagram of relationships in Heptathelidae. Comptes rendus 

du Xllème Colloque européen d'Arachnologie (Paris): 134-140. 
Haupt, J. & Shimojana, M. 2001: The spider fauna of soil banks: The genus Latouchia 

(Arachnida: Araneae: Ctenizidae) in Southern Japan and Taiwan. Mitteilungen des 

Naturkunde mus e ums Berlin, Zoologische Reihe 77: 95-1 10. 
Haupt, J. & Song, D. 1996. Revision of East Asian whip scorpions (Arachnida: Uropygi). I. 

China and Japan. Arthropoda Selecta (Moskva) 5: 43-52. 
Hikida, T. & Ota, H. 1997. Biogeography of reptiles in the subtropical East Asian islands (pp. 

11-28). In: Lue, K.Y. & Chen, T.-H. (eds). Proceedings of the Symposium on the 

Phylogeny, Biogeography and Conservation of Fauna and Flora of East Asian Region. 

Taipei. 
Hikida, T., Ota, H. & Toyama, M. 1992. Herpetofauna of an encounter zone of oriental and 

palaearctic elements: Amphibians and reptiles of the Tokara group and adjacent islands 

in the northern Ryukyus, Japan. Biological Magazine of Okinawa 30: 29-43. 
Ikehara, S. & Shimojana, M .1975. Okinawa no riku no dobutsu. (Terrestrial Animals in 

Okinawa) (in Japanese). Fudokisha, Naha, 143 pp. 
Kimoto, S. 1982. Zoogeography and ecology of the Ryukyu Archipelago with special reference 

to leaf beetles (Coleoptera: Chrysomelidae). Entomologia Generalis 8: 51-58. 
KiMURA, M. 1996. Quarternary paleography of the Ryukyu Arc (in Japanese). Journal of 

Geography 105: 259-285. 
MacArthur, R. & Wilson, E. O. 1963. An equilibrium theory of insular zoogeography. Evo- 
lution 17: 373-387. 
Okada, Y. & Koba, K. 1931. Considerations on the distribution of animals in Japan (in 

Japanese). Dobutsugaku Zasshi 43 (508-5 10): 320-35 1 . 
Ono, H. 1996. Two new species of the families Liphistiidae and Thomisidae (Araneae) from the 

Ryukyu islands, Southwest Japan. Acta Arachnologica 45: 157-162. 
Ono, H. 1997. New species of the genera Ryuthela and Tmarus (Araneae, Liphistiidae and 

Thomisidae) from the Ryukyu islands, Southwest Japan. Memoirs of the National 

Science Museum A 23: 149-163. 
Ono, H. 1998. Spiders of the genus Heptathela (Araneae, Liphistiidae) from Kyushu. Memoirs 

of the National Science Museum 30: 13-27. 
Ono, H. 2000. Zoogeographie and taxonomic notes on spiders of the subfamily Heptathelinae 

(Araneae, Mesothelae, Liphistiidae). Memoirs of the National Science Museum 33: 

145-151. 
Ono, H. & Nishikawa, Y. 1989. Taxonomic revision of the Heptathelid spider (Araneae, 

Mesothelae) from Amamioshima island, the Ryukyus. Memoirs of the National Science 

Museum A 22: 119-125. 
Paik, K. 1953. A study on the geographical distribution of Heptathela kimurai (Kishida). Acta 

Arachnologica 13: 63-68. 
Platnick, N.I. 2002. The world spider catalog. The American Museum of Natural History, New 

York. 



ZOOGEOGRAPHY IN SOUTHERN JAPAN 1 39 



Shimojana, M. & Haupt, J. 1998. Taxonomy and natural history of the funnel-web spider genus 

Macwthele (Araneae: Hexathelidae: Macrothelinae) in the Ryukyu Islands (Japan) and 

Taiwan. Species Diversity (Sapporo) 3: 1-15. 
Song, D. & Haupt, J. 1984. Comparative Morphology and phylogeny of liphistiomorph spiders 

(Araneae: Mesothelae). II. Revision of new Chinese Heptathelid species. Verhandlungen 

des natwwissenschaftlichen Vereins zu Hamburg (NF) 27, 443-451. 
Toda, M. 1999. Historical biogeography of East Asian populations of Rana limnocharis 

(Amphibia: Anura): A review. In: Ota, H. (ed.). Tropical Island Herpetofauna. Elsevier, 

Amsterdam. 
Yoshikura, M. 1965. Postembryonic development of a whip scorpion, Typopeltis stimpsoni 

(Wood). Kumamoto Journal of Science, Series B, Section 2, 7(2): 21-51. 
Watase, S. 1912. Dobutsugakujo yori mitaru nihongunto to tairikutono kankei. (The relationship 

between Japan islands and Asian continent based on zoology) (in Japanese). 

Kagakusekai 5 (8): 62-65. 



Revue suisse de Zoologie 110(1): 141-154; mars 2003 



New taxonomic considerations on some species of the genus 
Grosphus Simon, with description of a new species (Scorpiones, 
Buthidae) 1 

Wilson R. LOURENÇO 

Laboratoire de Zoologie (Arthropodes), Muséum National d'Histoire Naturelle, 

61 rue de Buffon, F-75005 Paris, France. E-mail: arachne@mnhn.fr 



New taxonomic considerations on some species of the genus Grosphus 
Simon, with description of a new species (Scorpiones, Buthidae). - 

Revised redescriptions are proposed for two closely related species of 
Grosphus Simon, namely: Grosphus limbatus (Pocock) and Grosphus 
bistriatus Kraepelin. A new species, Grosphus ankarafantsika sp. n. also 
related with these, is described from the Ankarafantsika Reserve in the 
Province of Majunga, Madagascar. The morphology of the basal middle 
lamellae of the pectines is illustrated by using scanning electron micro- 
scopy. A revised key to the 1 1 known species of the genus is presented. 

Key- words: Scorpions - Grosphus - revision - new species - Madagascar - 
Ankarafantsika Reserve. 

INTRODUCTION 

As mentioned in recent papers (Lourenço, 1999, 2001), the first Grosphus 
species to be described was Scorpio (Androctonus) madagascariensis Gervais (1843) 
= Grosphus madagascariensis (Gervais, 1843). This was followed by another isolated 
description, of Buthus limbatus (= Grosphus limbatus), by Pocock (1889). Kraepelin 
(1900) contributed to the study of the genus Grosphus with a more important publi- 
cation in which several species were described. In his comprehensive monograph on 
the scorpions of Madagascar, Fage (1929) described a new variety of Grosphus lim- 
batus, which he named annulata. This variety has meanwhile been raised to specific 
rank as Grosphus annulatus (Lourenço, 1996). Several other taxa new to the fauna of 
Madagascar were described by Lourenço (1995, 1996). The genus Grosphus, however, 
remained more or less stable in its composition. After the publication of Fage's (1929) 
monograph only three new species were described (Lourenço, 1996, 1999, 2001). 

Even though the original descriptions of Grosphus species were in most cases 
quite clear and have been confirmed in subsequent diagnoses (Fage, 1929; Lourenço, 
1996), certain characters may require a re-examination. The taxonomy of Grosphus is 
based mainly on two major characters: the pattern of colouration, and the morphology 
of the basal middle lamellae of the female pectines. For some closely related species, 



1 This study was supported by the Department of cultural affairs, City of Geneva, Switzerland. 
Manuscript accepted 01.07.2002 



142 W. R. LOURENÇO 

such as Grosphus limbatus and Grosphus bistriatus recent studies (see Lourenço, 
1996) were largely based on old and faded material preserved in alcohol. In this 
material, the patterns of colouration and pigmentation could no longer be observed 
precisely. In the present study, redescriptions are given for Grosphus limbatus and 
Grosphus bistriatus. These redescriptions are based on newly collected material which 
allows precise observation of the patterns of pigmentation. In addition, the morphology 
of the basal middle lamella of the pectines is illustrated using scanning electron 
microscopy. A new species of Grosphus, also related with G. limbatus and G. bistria- 
tus, is described as well. 

TAXONOMIC TREATMENT 

Redescriptions of Grosphus limbatus (Pocock, 1889) and Grosphus bistriatus 
Kraepelin, 1900 

Grosphus limbatus (Pocock) Figs 1-2, 9 

Buthus limbatus Pocock, 1889: 346. 
Grosphus limbatus; Kraepelin, 1891: 72. 
Grosphus limbatus; Kraepelin, 1895: 84 
Grosphus limbatus; Kraepelin, 1899: 33. 
Grosphus limbatus; Fage, 1929: 653. 
Grosphus limbatus; Lourenço, 1996: 10. 

Material examined; Madagascar, Central region, "Massif de l'lbity", 40 km south of 
Antsirabe (2254 m), II/III/2000 (F. Glaw & M. Vences coll.): 6 females, 1 female with brood (34 
juveniles), 1 female with brood (72 juveniles); 11/2001 (G. Garcia Herrero coll): 2 females (bad- 
ly preserved). 

Coloration. Basically yellow to reddish-yellow. Prosoma: carapace dark 
yellowish, with an inverted triangular blackish spot stretching from the median eyes to 
the lateral eyes; a wide blackish central spot behind the median eyes, fusing with the 
central blackish band over the mesosomal tergites; lateral margins blackish; eyes sur- 
rounded by black pigment. Mesosoma: yellowish with a central longitudinal blackish 
band over tergites I- VII. Metasoma: segments I-IV reddish-yellow; segment V reddish 
with dark or blackish zones especially on the ventral aspect. All ventral and latero- 
ventral carinae marked with blackish spots. Vesicle reddish-yellow with some 
dispersed dark spots; aculeus reddish at its base and blackish at its extremity. Venter: 
coxapophysis, sternum, genital operculum and pectines yellowish; sternites yellow 
with some discrete reddish zones; sternite VII with blackish spots over the carinae. 
Chelicerae yellowish without any variegated pigmentation; teeth of fingers reddish to 
blackish. Pedipalps: yellowish with only two small blackish spots at the base of the 
movable fingers; rows of granules on the edges of the fingers dark to blackish. Legs 
yellowish, without spots. 

Morphology. Carapace moderately granular; anterior margin almost straight, 
with a weak median concavity. All carinae weak; furrows moderate to strong. Median 
ocular tubercle slightly anterior to the center of carapace; median eyes separated by one 
ocular diameter. Three pairs of lateral eyes. Sternum between sub-triangular and sub- 
pentagonal in shape. Mesosoma: tergites with thin, moderately intense granulation. 



ON SOME SPECIES OF GROSPHUS SIMON 143 

Median carina moderately developed on all tergites. Tergite VII pentacarinate. Venter: 
genital operculum consisting of two subtriangular plates. Pectines: pectina! teeth count 
(24 to 27); basal middle lamellae of each pecten elongated and weakly curved, widen- 
ing only in proximal third. Sternites smooth, with moderately elongated stigmata; VII 
with four vestigial carinae and a few thin granules. Metasoma: segments I to III with 
10 carinae, moderately crenulate. Segment IV with 8 carinae, moderately crenulate. 
Segment V with 5 carinae, the dorsal one being only weakly marked. Dorsal carinae on 
segments I-IV with weak posterior spinoid granules. Intercarinal spaces moderately to 
weakly granular. Telson with dispersed granules on lateral and ventral surfaces; dorsal 
surface smooth; aculeus moderately curved and shorter than the vesicle; subaculear 
tooth absent. Cheliceral dentition characteristic of the family Buthidae; two distinct 
basal teeth present on the movable finger, the more basal one being slightly reduced 
(Vachon, 1963); ventral aspect of both fingers and of manus with dense, long setae. 
Pedipalps: femur pentacarinate; patella with carinae represented by spinoid granules on 
the internal and dorsal faces; tibia without carinae, all faces weakly granular to smooth. 
Movable and fixed fingers with 13/12 oblique rows of granules. Trichobothriotaxy; or- 
thobothriotaxy A-a (Vachon, 1974, 1975). Legs: tarsus with numerous very short thin 
setae ventrally. Patellar spurs present on legs III and IV; pedal spurs present on legs I 
to IV; all spurs strong. 

Grosphus bistriatus Kraepelin Figs 3-4, 15, 17, 21 

Grosphus bhstriatus Kraepelin, 1900: 14. 
Grosphus bistriatus; Kraepelin, 1901: 267. 
Grosphus bistriatus var. pallicauda Strand, 1908: 485. 
Grosphus bistriatus; Fage, 1929: 651. 
Grosphus bistriatus; Lourenço, 1996: 13. 

Material examined: Madagascar, Western Region, Ambalanjanakomby, 11/2000 (O. 
Ramilijaona coll.): 1 male. Malainmbandy, VIII/2000 (O. Ramilijaona): 1 female, 1 juvenile (fe- 
male). 

Coloration. Basically yellowish. Prosoma: carapace pale yellow with two longi- 
tudinal blackish lines behind the median eyes; one small dark spot on the anterior 
margin and two on the posterior margin fusing with the longitudinal blackish bands on 
mesosomal tergites; eyes surrounded by black pigment. Mesosoma: yellowish with two 
longitudinal blackish bands over tergites I- VI, more densely marked in the posterior 
half of each tergite. Metasoma: all segments yellow with some dark pigmentation on 
their ventral aspect, most pronounced on segment V; two diffuse spots present on the 
dorsal face of segment V Vesicle yellow with dispersed light brownish spots laterally 
and ventrally; aculeus reddish yellow. Venter: coxapophysis, sternum, genital oper- 
culum and pectines yellowish; sternites yellow. Chelicerae yellowish, without any 
variegated pigmentation; only a few dark spots at the base of reddish fingers. Pedi- 
palps: yellowish throughout without any pigmentation. Legs yellowish with discrete 
brownish variegated pigmentation. 

Morphology. Carapace intensely granular; anterior margin almost straight with 
a weak median concavity. All carinae weak to moderately developed; furrows 
moderately developed. Median ocular tubercle anterior to the center of carapace; 
median eyes separated by a little more than one ocular diameter. Three pairs of lateral 



144 



W. R. LOURENÇO 



-. 







Figs 1-4 
1-2. Grosphus limbatus. female, ventral and dorsal aspects (photos CI. Ratton). 3-4. Grosphus 
bistriatus, male, ventral and dorsal aspects (photos CI. Ratton). 



ON SOME SPECIES OF GROSPHUS SIMON 145 

eyes. Sternum between sub-triangular and sub-pentagonal in shape. Mesosoma: 
tergites with a thin and intense granulation. Median carina moderate on all tergites. 
Tergite VII pentacarinate. Venter: genital operculum consisting of two subtriangular 
plates. Pectines: pectinal teeth count (28 in males; 25-26 in females); basal middle 
lamellae of each pecten not dilated in males, elongated and weakly curved in females, 
widening in proximal half. Sternites smooth, with moderately elongated stigmata; VII 
with four vestigial carinae and a few thin granules. Metasoma: segments I and II with 
10 carinae, moderately crenulate. Segments III and IV with 8 carinae, moderately 
crenulate. Segment V with 5 carinae, the dorsal ones being only weakly marked. Dorsal 
carinae on segments I-IV without any posterior spinoid granules. Intercarinal spaces 
moderately granular. Telson with very few granules on lateral and ventral surfaces; dor- 
sal surface smooth; aculeus moderately curved and shorter than the vesicle; subaculear 
tooth absent. Cheliceral dentition characteristic of the family Buthidae (Vachon, 1963); 
two distinct basal teeth present on the movable finger, the more basal one being slight- 
ly reduced; ventral aspect of both fingers and of manus with dense, long setae. 
Pedipalps: femur pentacarinate; patella with carinae represented by some spinoid gran- 
ules on the internal face; tibia without carinae, all faces weakly granular to smooth. 
Movable and fixed fingers with 12/1 1 oblique rows of granules. Trichobothriotaxy; or- 
thobothriotaxy A-a (Vachon, 1974, 1975). Legs: tarsus with numerous short thin setae 
ventrally. Patellar spurs present on legs III and IV, pedal spurs present on legs I to IV; 
all spurs strong. 

Description of a new species 

Grosphus ankarafantsika sp. n. Figs 5-8, 10-14, 16, 18-20, 22-25 

Grosphus bistriatus; Fage, 1929: 651-652 (in part, misidentification) 
Grosphus bistriatus; Lourenço, 1996: 13 (misidentification) 
Grosphus bistriatus; Lourenço, 2001: 459 (misidentification) 

Material examined: Madagascar, Majunga Province, Ankarafantsika Reserve, Ampijoroa 
Forest Station (16°18'S 46°48'E), sand area of Paquypodium (G. Garcia Herrero coll.), 27/11, 
01/III/2001 (pitfall traps): 1 female (holotype), 11 males (paratypes); "Jardin Botanique A" (G. 
Garcia Herrero coll.), 24-27/11/2001 (pitfall traps): 6 males, 2 females, 1 female with brood (45 
juveniles) (paratypes). Type material deposited in the Muséum d'histoire naturelle, Genève. 

Etymology: The name of the type locality (Ankarafantsika Reserve) is placed in 
apposition to the generic name. 

Diagnosis: The morphology of the new species shows that it is close to 
Grosphus bistriatus Kraepelin, but it can be readily distinguished from that species by 
the following characters: (i) carapace yellowish with an inverted triangular blackish 
spot; (ii) dark confluent zones on lateral sides of tergites, next to the blackish bands; 
(iii) spots on legs vestigial or absent; (iv) carapace moderately granular; (v) basal 
middle lamellae of the female pectines elongated and curved, constantly narrowing 
from the base to the apex. 

Description: It is based on female holotype and one male paratype. 
Measurements in Table I. 

Coloration. Basically yellowish with some dark zones on the body. Prosoma: 
carapace yellow with an inverted blackish triangular spot between median and lateral 



146 



W. R. LOURENÇO 





^T 



v . ; 



8 





Figs 5-8 
Grosphus ankarafant sika sp. n. 5-6. Female holotype. 7-8. Male paratype. Ventral and dorsal 
aspects (photos CI. Ratton). 

eyes; two dark spots on the posterior margin; eyes surrounded by black pigment. 
Mesosoma: dark yellow with confluent dark zones and with two longitudinal blackish 
strips over tergites I- VI running from the two spots on the posterior margin of the cara- 
pace. Metasoma: all segments yellowish with some vestigial dark pigmentation on the 
ventral carinae. Vesicle reddish yellow without spots; aculeus with yellowish base and 
dark reddish tip. Venter: coxapophysis, sternum, genital operculum and pectines 



ON SOME SPECIES OF GROSPHUS SIMON 147 



Table I. Morphometric values (in mm) of the male paratype and female holotype of Grosphus 
ankarafantsika sp. n. 



S -paratype 9 -holotype 



Total lenght 37.0 49.0 

Carapace: 

- length 4.6 6.0 

- anterior width 3.3 4.6 

- posterior width 4.8 7.5 
Metasomal segment I: 

- length 3.3 4.4 

- width 2.8 3.9 
Metasomal segment V: 

- length 5.7 7.3 

- width 3.0 3.8 

- depth 2.3 3.3 
Vesicle: 

- width 2.3 3.0 

- depth 1.9 2.8 
Pedipalp: 

- Femur length 3.5 5.2 

- Femur width 1.2 1.6 

- Patella length 5.0 6.0 

- Patella width 1.8 2.4 

- Tibia length 8.4 10.2 

- Tibia width 2.3 2.7 

- Tibia depth 2.2 2.6 
Movable finger: 

- length 4.6 6.2 



yellowish; sternites reddish yellow. Chelicerae yellowish with dark variegated 
pigmentation on the anterior third; fingers reddish. Pedipalps: yellowish with vestigial 
dark zones on femur and patella. Legs yellowish with vestigial brownish zones (absent 
in some specimens). 

Morphology. Carapace moderately granular; anterior margin almost straight, 
with a weak median concavity. All carinae weak; furrows moderately developed. 
Median ocular tubercle anterior to the center of carapace; median eyes separated by a 
little more than one ocular diameter. Three pairs of lateral eyes. Sternum between sub- 
triangular and sub-pentagonal in shape. Mesosoma: tergites with thin but moderately 
intense granulation. Median carina moderately developed in all tergites. Tergite VII 
pentacarinate. Venter: genital operculum consisting of two subtriangular plates. 
Pectines: pectinal teeth count 24-24 (female holotype; variation: 27 to 31 in males; 24 
to 27 in females); basal middle lamellae of each pecten not dilated in males; elongated 
and curved in females; constantly narrowing from the base to the apex. Sternites 
smooth, with elongated stigmata; VII with two vestigial carinae. Metasoma: segments 
I and II with 10 carinae, moderately crenulate. Segments III and IV with 8 carinae, 
moderately crenulate. Segment V with 5 carinae, the dorsal one being only weakly pro- 
nounced. Dorsal carinae on segments I-IV without any posterior spinoid granules. 
Intercarinal spaces weakly granular. Telson with granules scattered over latero-ventral 



148 



W. R. LOURENÇO 




Figs 9-14 
9. Grosphus limbatus. Detail of basal middle lamellae of the female pecten. 10-14. Grosphus 
ankarafantsika sp. n. 10. Metasomal segments IV-V and telson, lateral aspect (female holotype). 
11-14. Trichobothrial pattern. 11. Tibia (female holotype). 12. Femur (female holotype). 13. 
Femur (male paratype). 14. Patella (male paratype). 



ON SOME SPECIES OF GROSPHUS SIMON 



149 



À 


- 


tifo 


J: 

Mi 


/ /■ / / // . 








\ 


..—- 


w 


:\ 


\ \ \ 








r 8 


wa» 


15 




' 


K60 


l00Hm 




• - 




Figs 15-18 
15,17. Grosphus bistriatus. 16,18. Grosphus ankarafantsika sp. n. 15-16. Left pecten of female. 
17-18. Basal middle lamellae, in detail. 



and ventral surfaces; its dorsal surface smooth; aculeus moderately curved and shorter 
than the vesicle; subaculear tooth absent. Cheliceral dentition characteristic of the 
family Buthidae (Vachon, 1963); two distinct basal teeth present on the movable finger, 
the more basal of them being slightly reduced; ventral aspect of both fingers and of 
manus with dense, long setae. Pedipalps: femur pentacarinate; patella with a dorso- 
internal carina and with several spinoid granules on the internal face; tibia smooth, 
without carinae, all faces weakly granular to smooth. Movable and fixed fingers with 
11/10 (females) and 12/11 (males) oblique rows of granules. Trichobothriotaxy; 
orthobothriotaxy A-a (Vachon, 1974, 1975). Legs: tarsus with numerous short thin 
setae ventrally. Patellar spurs present on legs III and IV; pedal spurs present on legs 
I to IV; all spurs strong. 

Remarks: The new species was collected in the Reserve together with Grosphus 
madagascariensis (Gervais) and Grosphus gardai Lourenço. These specimens are also 
deposited in the Muséum d'histoire naturelle, Genève. 

Discussion 

Buîhus limbatus (= Grosphus limbatus) was described by Pocock (1889) on the 
basis of four specimens (one male and three females) collected by Rev. R. Baron in 



150 



W. R. LOURENÇO 




X2.000 10HHI 




Figs 19-21 
Pectines. 19. Male pecten of Grosphus ankarafantsika sp. n., global view. 20-21. Microstructure 
of peg sensilla on teeth. 20. Grosphus ankarafantsika sp. n. 21. Grosphus bistriatus. 



ON SOME SPECIES OF GROSPHUS SIMON 



151 




Figs 22-25 
Grosphus ankarafantsika sp. n. (male paratype). 22. Movable finger of tibia pedipalp. 23. Detail 
of distal extremity of the same. 24. Movable finger of chelicera. 25. Fixed finger of chelicera. 



Madagascar, but without giving any precise locality. The description by Pocock was 
very precise, and Fage (1929) was subsequently able to identify further material of this 
species (probably without examination of Pocock 's type material) and to demonstrate 
its presence on the Central Plateau of Madagascar. 

The morphology of the basal middle lamellae of the female pectines is a 
diagnostic character for Grosphus species. The description given by Pocock (1889) for 
this character is clear and accurate: "in the female the basal tooth is very much enlarged 
being about twice as long as the rest and flask-shaped, i. e. dilated proximally and 
abruptly narrowed and slender distally." This character was precisely illustrated by 
Pocock (1889: Fig. 7a), and also by Fage (1929: Fig. 6); its morphology was later 
confirmed by Lourenço (1996: Fig. 8). 

The precise identity of Grosphus bistriatus appears unclear. This species was 
described by Kraepelin (1900), on the basis of two specimens (syntypes) collected near 
Tulear in the south of Madagascar. Fage (1929), based his redescription on several 
specimens from different localities, including some from the "Massif d'Ambre" and 
"Maevatanana" in the northern range of the island. One of the syntypes, preserved in 
alcohol and deposited in the "Muséum national d'Histoire naturelle" in Paris, was 



152 



W. R. LOURENÇO 




Fig. 26 
Map showing the type locality of Grosphus ankarafantsika sp. n. (black circle) and the sites from 
where the new material of G. limbatus (black star) and G. bistriatus (black square) has been col- 
lected. 



probably already bleached when Fage examined it. Therefore, the colouration 
described by Fage was probably based on other non-type specimens. The morphology 
of the basal middle lamellae of the female pectines, illustrated by Fage (1929: Fig. 5), 
differs from that of the bleached syntype. In his diagnosis of G. bistriatus, Lourenço 
(1996) accepted the redescription by Fage (1929) and reported the presence of this 
species in the "Réserve naturelle intégrale n° 7 de F Ankarafantsika); a misiden- 
tification subsequently confirmed by Lourenço (2001). 



ON SOME SPECIES OF GROSPHUS SIMON 153 

An examination of newly collected material of G. bistriatus from close to the 
type locality (region of Tulear), provide clarification on coloration and patterns of 
pigmentation of this species. In addition, the precise morphology of the basal middle 
lamellae of female pectines is described in here. Moreover, the population present in 
the Ankarafantsika Reserve is described as a new species, i. e. G. ankarafantsika sp. n. 
Grosphus limbatus, G. bistriatus, G. intertidalis Lourenço and Grosphus ankara- 
fantsika sp. n. represent a species group which is isolated from the other species of the 
genus. 

Key to the species of the genus Grosphus 

1 Pectines with a maximum of 21 teeth 2 

(1) Pectines with more than 22 teeth 4 

2 Colouration yellowish to reddish yellow, with variegated brownish pig- 
mentation; body length about 30 mm G. garciai 

(2) Colouration dark, from reddish brown to dark brown; body length about 

50 mm 3 

3 Colouration reddish brown to dark brown, without light spots; metaso- 
mal segment I longer than wide; basal middle lamellae of female 
pectines oval in shape G. madagascarienis 

(3) Colouration reddish brown with lighter spots; metasomal segment I 
wider than long; basal middle lamellae of female pectines subquadran- 

gular in shape G. hirtus 

4 Colouration blackish throughout; pectines with 30 to 40 teeth; body 
length more than 90 mm G. grandidieri 

(4) Colouration reddish brown to yellowish, never blackish; body length 

less than 90 mm 5 

5 Mesosoma with homogenous colouration, reddish brown or yellowish 6 

(5) Mesosoma with a blackish median longitudinal band, or with two 
blackish lateral longitudinal bands 9 

6 Total length more than 70 mm; mesosoma reddish brown; basal middle 
lamellae of female pectines two times longer than wide at their base 
G.flavopiceus 

(6) Total length less than 60 mm; mesosoma yellowish; basal middle lamel- 
lae of female pectines three times longer than wide at the base 7 

7 Metasomal segment V and telson pale yellowish G. intertidalis 

(7) Metasomal segment V and telson with blackish spots or blackish 
throughout 8 

8 Metasomal segment V and telson with blackish spots G. annulatus 

(8) Metasomal segment V and telson blackish G.feti 

9 Mesosoma with a wide blackish median longitudinal band; basal middle 
lamellae of female pectines three times longer than wide at their base 

and covering the 4 proximal teeth G. limbatus 

(9) Mesosoma with two narrow blackish lateral longitudinal bands 10 



154 W. R. LOURENÇO 

10 Carapace without a blackish triangular spot; basal middle lamellae of 
female pectines weakly curved, widening in proximal half and covering 
the two proximal teeth G. bistriatus 

(10) Carapace with a blackish triangular spot; basal middle lamellae of the 
female pectines curved and constantly narrowing from the base to apex 
covering the four proximal teeth G. ankarafantsika sp. n. 

ACKNOWLEDGEMENTS 

I am very grateful to R. Cleva and L. Albenga (Muséum national d'Histoire 
naturelle, Paris), for technical help with the preparation of SEM photos. To Prof. John 
L. Cloudsley-Thompson, London, and to Dr Peter Schwendinger, Muséum d'histoire 
naturelle, Geneva for reviewing the manuscript. 

REFERENCES 

Fage, L. 1929. Les Scorpions de Madagascar. Faune des Colonies françaises 3. Société 

d'Editions Géographiques, Maritimes et Coloniales, Paris, pp. 637-694. 
Gervais, P. 1844. Remarques sur la famille des Scorpions. Archives du Muséum d'Histoire 

Naturelle, Paris 4: 201-240. 
Kraepelin, K. 1891. Revision der Skorpione. I. Die Familie der Androctonidae. Jahrbuch der 

Hamburgischen wissenschaftlichen Anstalten 8: 1-144. 
Kraepelin, K. 1895. Nachtrag zu Theil I der Revision der Scorpione. Jahrbuch der 

Hamburgischen wissenschaftlichen Anstalten 12: 73-96. 
Kraepelin, K. 1899. Scorpiones und Pedipalpi. In: F. Dahl (ed.). Das Tierreich. Herausgegeben 

von der Deutschen zoologischen Gesellschaft. 8 (Arachnoidea). R. Friedländer und Sohn 

Verlag, Berlin, 265pp. 
Kraepelin, K. 1900. Üeber einige neue Gliederspinnen. Abhandlungen aus dem Gebiete der 

Naturwissenschaften 16: 3-17. 
Kraepelin, K. 1901. Catalogue des scorpions des collections du Muséum d'histoire naturelle de 

Paris. Bulletin du Muséum national d'histoire naturelle, Paris 7: 263-265. 
Lourenço, W. R. 1995. Description de trois nouveaux genres et quatre nouvelles espèces de 

Scorpions Buthidae de Madagascar. Bulletin du Muséum National d'Histoire Naturelle, 

Paris, 4e sér. 17(1-2): 95-106. 
Lourenço, W. R. 1996. Scorpions (Chelicerata, Scorpiones). In: Faune de Madagascar N° 87. 

Muséum National d'Histoire Naturelle, Paris, 102 pp. 
Lourenço, W. R. 1999. A new species of Grosphus Simon (Scorpiones, Buthidae), the first 

record of an intertidal scorpion from Madagascar. Entomologische Mitteilungen aus dem 

Zoologischen Museum Hamburg 12 (158): 297-307. 
Lourenço, W. R. 2001. Another new species of Grosphus (Scorpiones, Buthidae) for 

Madagascar. Revue suisse de Zoologie 108 (3): 455-461. 
Pocock, R. I. 1889. Notes on some Buthidae, new and old. Annals and Magazine of Natural 

History, ser. 6, 3: 334-351. 
Strand, E. 1908. Arachniden aus Madagaskar, gesammelt von Herrn Walter Kaudern. 

Zoologische Jahrbücher, Abtheilung für Systematik 26: 453-488. 
Vachon, M. 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères 

chez les Scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2è sér. 35 

(2): 161-166. 
Vachon, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de 

Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux 

et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum National 

d'Histoire Naturelle, Paris, 3è sér., n° 140, Zool. 104: 857-958. 
Vachon, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des Scorpions 

(Arachnides) dans le classement des genres de la famille des Buthidae Simon. Comptes 

Rendus de ï Académie des Sciences. Paris, sér. D, 281: 1597-1599. 



Revue suisse de Zoologie 1 10 (1): 155-206; mars 2003 



Catalogue des types de Lamiinae (Coleoptera, Cerambycidae) 
conservés au Muséum d'histoire naturelle, Genève 

Jeannine MORATI 

27 chemin du pont, Grésin, F-01200 Léaz, France. 



A catalogue of the types of Lamiinae (Coleoptera, Cerambycidae) 
housed in the Muséum d'histoire naturelle, Geneva. - A list is provided 
of 218 holotypes, the syntypes of 15 species and subspecies, and paratypes 
of 129 species and subspecies of the Lamiinae presently housed in the 
Muséum d'histoire naturelle, Geneva. Most of the names were published by 
S. Breuning. As the taxonomic treatment of the Lamiinae was often inade- 
quate it was necessary to check the original descriptions for all names listed. 
The authenticity of several "types" is found doubtful. They are indicated by 
"?HP" or "?PT". Unpublished information from labels is given in square 
brackets. Illegible handwritten text is indicated by the sign □, When a 
description was published twice, a reference to the second description is 
given in parentheses. Separate lists are given for type material of varieties 
published before 1961, and for unavailable names published with designa- 
tion of type material. 

Key-words: Coleoptera - Cerambycidae - Lamiinae - type material. 

INTRODUCTION 

Le Muséum d'histoire naturelle de Genève (MHNG) possède une collection de 
Lamiinae qui compte quelques 5000 espèces. Elle consiste en une collection historique 
de A. Melly à laquelle se sont ajoutés les collections H. -A. Junot, R. Mussard, W. Siess, 
les récoltes faites par les entomologistes du Muséum ainsi que des lots importants 
cédés par S. Breuning. 

Cette collection contient de nombreux spécimens étiquetés holotype, paratype 
ou syntype. Pour la plupart, il s'agit de taxa décrits par Breuning qui, au cours de 
soixante ans consacrés à l'entomologie, a étudié la plupart des collections de Lamiinae 
des principaux musées européens et publié environ 630 articles dont plus de 600 sont 
consacrés aux Lamiinae. Les spécimens types des espèces qu'il a décrites se retrouvent 
dispersés dans ces différentes institutions et il paraissait utile d'établir une liste des 
noms de ces taxa, conformément à la recommandation 72F 4 du Code International de 
Nomenclature Zoologique (4 e éd., 1999), pour faciliter le travail des réviseurs poten- 
tiels qui, en plus des difficultés découlant des anciennes définitions sommaires des 
taxa, se trouvent souvent dans l'impossibilité de localiser le matériel typique indis- 
pensable à toute étude. 



Manuscrit accepté le 1 1 .06.2002 



156 J- MORATI 



MATERIEL ET METHODES 



Les sources principales ont été le Catalogue des Lamiaires du Monde 
(Breuning, 1958-1969), la bibliographie de Breuning (Breuning, 1984), les Zoological 
Records et les descriptions originales des différents auteurs. 

Les noms des taxa sont écrits suivant l'orthographe (éventuellement complétée 
par l'orthographe subséquente reconnue), la combinaison et le statut originaux. Ils sont 
répertoriés alphabétiquement, sans mention de sous-genre ni de tribu. La localité type 
est donnée pour chaque nom comme publiée. La page indiquée correspond à la pre- 
mière page sur laquelle le nom a été introduit. La date relevée est celle de la parution 
effective, suivie entre parenthèses de la date imprimée sur la publication, quand elles 
sont différentes. En cas d'homonymie, les références relevées concernent l'espèce en 
collection. Les références bibliographiques sont apportées in extenso en fin de cata- 
logue. Quand une même description «originale» est publiée deux fois, la deuxième 
publication est notée entre parenthèses. Breuning ayant diversement signé ses publica- 
tions: Stephan von Breuning, S. Breuning ou Etienne de Breuning, dans un souci 
d'homogénéité, l'orthographe «Breuning S.» a été choisie, suivie de (E.) quand la 
signature est francisée, la particule n'est pas mentionnée. Certaines informations d'éti- 
quettes manuscrites illisibles, sont notées ill.; quand seules certaines lettres sont illi- 
sibles, elles sont remplacées par □. Le terme «type» est transcrit holotype quand la 
description originale le précise ou quand elle ne s'appuie que sur un seul spécimen, 
syntype quand elle s'appuie sur plusieurs spécimens ou quand les spécimens sont notés 
cotypes (CT). Contrairement à ce qui a été écrit (Breuning, 1974), les types de la col- 
lection Chassot n'ont pas été déposés au MHNG (communication personnelle, 
Besuchet, 1999). 

Certaines incohérences ayant été décelées, il paraissait nécessaire de vérifier 
l'authenticité «en tant que type» de ces spécimens et la disponibilité de leur nom. La 
validité de ces noms n'a cependant pas été contrôlée. 

- Holotype s, syntype s. L'authenticité des spécimens en tant que porte-nom a été 
établie par le contrôle avec les descriptions originales. Il nous a semblé préférable de 
donner la priorité aux informations de ces dernières qui font référence, leur accord avec 
les informations des étiquettes présumant de l'authenticité du spécimen. Sont consi- 
dérés comme authentiques les «types» dont les données des étiquettes correspondent à 
celles de la description originale, ils sont notés HT ou ST. Quand les données des éti- 
quettes complètent l'information publiée, elles sont apportées entre crochets. Quand 
les données des étiquettes sont en contradiction avec celles de la description originale 
ou quand le lieu de dépôt mentionné n'est pas le MHNG, les «types» sont considérés 
comme douteux, ils sont notés ?HT ou ?ST, les informations contradictoires sont alors 
notées entre crochets. 

- Paratypes. Leur authenticité est difficilement verifiable. Ils ne sont pas tou- 
jours mentionnés dans les descriptions originales et quand ils le sont, leur nombre n'est 
que rarement indiqué, leur lieu de conservation a pu changer étant souvent utilisés pour 
des échanges par les collectionneurs privés. Conventionnellement, sont présumés 
authentiques les «paratypes» dont les données des étiquettes correspondent à celles de 
la description originale et dont il est fait mention dans celle-ci. Ils sont notés PT. Sont 



CATALOGUE DES TYPES DE LAMIINAE 157 

considérés comme douteux les «paratypes» qui ne sont pas mentionnés dans la 
description, ceux dont les informations des étiquettes et de la description sont contra- 
dictoires ou ceux qui se trouvent en «surnombre», ils sont notés ?PT. N'ont pas été 
mentionnés ceux portant une date de récolte postérieure à celle de la description! 

La disponibilité des noms a été établie selon les dispositions du Code Inter- 
national de Nomenclature Zoologique, 4 e éd., 1999 (ICZN). 

- Les noms du niveau espèce et sous-espèce publiés comme tels sont répertoriés 
dans la liste 1. Celle-ci compte 218 holotypes, 17 syntypes de 15 espèces et 203 
paratypes de 129 espèces pouvant être considérés comme authentiques. Il s'agit prin- 
cipalement de représentants de la tribu des Saperdini (Oberea: 50 holotypes, 1 syntype 
et 36 paratypes de 25 espèces; Phytoecia: 36 holotypes, 2 syntypes de 2 espèces et 14 
paratypes de 9 espèces). 

- Les noms publiés en tant que «variété» avant 1961, sont disponibles (ICZN: 
art. 45.6.4) et répertoriés dans la liste 2 (1 holotype et 2 syntypes d'une espèce). 

- Les noms infra-subspécifiques autres que ceux de «variété» ou «forme» 
publiés avant 1961, ne sont pas disponibles, sauf s'ils ont été utilisés au niveau sub- 
spécifique avant 1985 (ICZN: art. 45.6.4.1.) (ce que nous n'avons pas vérifié dans ce 
travail). Tous ces noms sont donc répertoriés dans la liste 3. Les noms du niveau infra- 
subspécifique publiés après 1960, bien que n'étant pas disponibles (ICZN: art. 45.6.2. 
et 45.6.3) sont également répertoriés dans la liste 3. 

- Les noms de collection n'ont pas été retenus pour éviter une interprétation 
erronée de leur disponibilité, mais pourraient être communiqués sur demande, sous 
forme d'une liste. Cette liste répertorie les noms du niveau espèce, sous-espèce ou 
infra-subspécifique pour lesquels aucune référence n'a été trouvée dans la littérature. Il 
pourrait s'agir d'espèces non décrites, d'espèces renommées ou corrigées par Breuning 
ou encore de noms ayant échappé à notre vigilance. Elle répertorie également des noms 
pour lesquels des notes inédites de Breuning ont été retrouvées. 



LISTE 1: NOMS DU NIVEAU ESPECE ET SOUS-ESPECE 

Abryna coenosa mindanaonis Breuning, 1980e: 169. 

2 ?PT: [Insel Mindanao]. 

Acalolepta australis orientalis Breuning, 1982: 18. 

PT: Papouasie, archipel de la Louisiade, île Sud-Est, IV. 1898, Meek coll. 
Acalolepta australis saintaignani Breuning, 1970a: 377 (Breuning, 1982: 18). 

3 PT: St. Aignan, 8.IX.1897 [Meek coll.]; 2 ?PT: [St Aignan, VIII-XI.97, 
Meek]. 

Acalolepta brunnescens Breuning, 1980e: 175. 

3 ?PT: [Insel Mindanao]. 
Acalolepta densefuscomarmorata Breuning, 1982: 19. 

?PT: [Woodlark, Meek, 95]. 
Acalolepta densemarmorata Breuning, 1970b: 475. 

PT: Neu Hannover, II-III.97, Webster. 



158 J- MORATI 

Acalolepta holobrunnea Breuning, 1980e: 174. 

3 PT: Insel Mindanao. 
Acalolepta lumawigi Breuning, 1980e: 175. 

?PT: [Philippines, Insel Mindanao]. 
Acalolepta meeki Breuning, 1982: 18. 

PT: Papouasie, archipel de la Louisiade, Ile Sud-Est, IV. 1898, Meek coll. 
Acalolepta paravariolaris Breuning, 1980e: 175. 

?PT: [Insel Mindanao]. 
Acalolepta saintaignani Breuning, 1982: 18. 

2 PT: Papouasie, archipel de la Louisiade, île Saint- Aignan, VIII-XI.1897, 

Meek coll. 
Acalolepta woodlarkiensis Breuning, 1970b: 476. 

?PT: [Woodlark, Meek, 95]. 

Acridoschema mbargai Breuning, 1972a: 282. 

PT: Cameroun, M'Balmayo, 11.72, leg. M'Barga. 

Agapanthia fallax Holzschuh, 1974 (1973): 95. 

PT: Anatolie, prov. Mus, Buglangeçidi, 1600m, 11-17.VI.1973, Holzschuh. 

Agelasta albesignata Breuning, 1980e: 158. 

?PT: [Philippines, Mindanao, leg. Lumawig]. 
Agelasta albostictica Breuning, 1980e: 158. 

4 ?PT: [Insel Mindanao]. 

Agelasta mediofasciatipennis Breuning, 1974g: 78. 

?PT: [Philippinen, Luzon, Prov. Benguet S. P., Baguio]. 

Amblymora obiensis Breuning, 1956d: 685. 

HT: Ile Obi, ex coll. Preiss, coll. Lepesme [Moluques]. 

Anauxesis rufipennis Breuning, 1976b: 738. 

HT: Kenya, Shimba Hills, près Kwale, 29.XI.1974, leg. V Mahnert et J. Perret 

[près Madabara, Pumping station, tamisage]. 
Anauxesis rufoscapa Breuning, 1950b: 7. 

HT: Transvaal, Shilouvane, leg. H. Junod. 

Ancylonotopsis pictoides Breuning. 1974d: 775. 
?PT: [Ht Zaïre, Kingansani]. 

Anexodus sarawakensis Sudre. 1997: 253. 

HT: â , E. Malaisie, Sarawak, confi. Sun Oyan, et Mujong river, E. Kapit, 500m, 
18.V.1994, Lobi et Burckhardt; 2 PT c? et 9 : idem. 

Annamanum fuscomaculatum Breuning, 1979b: 99. 

?PT: [Yunnan. Peyentsin]; 2 ?PT: [Yunnan, Djoukoula]. 



CATALOGUE DES TYPES DE LAMIINAE 159 

Aprìona yayeyamai Breuning, 1976b: 739. 

HT: ô , Japon, Ile Ishigakishima, Yayayama, leg. Ohbayashi. 

Arachneosomatidia beatricea Sudre, 2000: 83. 

HT: New Caledonia, Mt Khogi, prim, for., 400-500m, litter, 18-19.X.1998, 
Lobi; PT: idem. 

Astathes bimaculatoides Breuning, 1971a: 3. 

?PT: [Laos, Paksé, Coll. Rondon]; 2 ?PT: [Laos, Paksé, 31.V.67]; 2 ?PT: [idem 
mais 15.VI.67]. 

Australiorondonia australica Breuning, 1982: 25. 
3 PT: Australia. 

Bacchisa nigricornis Breuning, 1969a: 37. 

HT: S, Laos, Pa-Hia, 19.IV.50, leg. J. Romieux. 

Batrachorhina rodriguezi Breuning, 1948: 22. 
PT: Rodrigues, 1934, Dr Madge. 

Blapsilon baloghi Breuning, 1978a: 25. 

2 PT: New Caledonia, Mont Rembai, 19-21.1.1977, leg. Dr J. Balogh. 
Blapsilon kaszabi Breuning, 1978a: 26. 

PT: New Caledonia, Mont Rembai, 19-21.1.1977, leg. Dr J. Balogh. 

Blepisanis disconotaticollis Breuning, 1950c: 21. 

2 PT: Kongo Belg. [Beni]. 
Blepisanis flavovittata Breuning, 1950c: 23. 

?PT: [Congo, Elisabethville]. 
Blepisanis guineensis Kolbe, 1893: 78. 

ST: Togo, Bismarckburg [1.VI-15.VII.91, R. Büttner S.]. 
Blepisanis incallosa Breuning, 1950d: 273. 

?PT: [Angola, Bailundo]. 
Blepisanis lineata Aurivillius, 1913: 35. 

ST: Angola, Bailundo. 
Blepisanis orientis Aurivillius, 1910: 151. 

2 ST: Kilimandjaro, Kibonoto, Kulturzone [S. Jöstedt, 1905-6, 1300m]. 
Blepisanis subcallosa Breuning, 1950c: 22. 

2 ?PT: [Angola, Bailundo]. 
Blepisanis uniformis Breuning, 1950c: 19. 

2 ?PT: [Angola, Bailundo]. 

Brachychilus lituratus Blanchard, 1851: 500. 

ST: Chili, Santiago, Coquimbo, Illapel, Arquero, etc. [Chili, Santiago, Gay, 
1849]. 



160 J- MORATI 

Brachychilus scutellaris Blanchard, 1851: 500. 

ST: Chili, Santiago, Santa Rosa, etc. [Chili, Gay, 1849]. 

Brachyolene unicolor Breuning, 1974b: 372. 

HT: Côte d'Ivoire, Sassandra, XII. 1949, leg. R. Mussard. 

Cada grossepunctata Breuning, 1980e: 160. 
?PT: [Insel Mindanao]. 

Celosterna perakensis Breuning, 1976b: 739. 

HT: Malaisie, Perak, Taiping, 14.III.1972, leg. K.C. Liew. 

Cereopsius sexmaculatus immaculithorax Breuning, 1974f: 238. 

2 PT: Borneo, Maloewi [Coll. Besnard]. 
Cereopsius spilotoides Breuning, 1974a: 74. 

HT: Philippines, He Luzon, Manila, Zambo, IX. 1928, W. Parsons; PT: idem. 

Ceroplesis aestuans dakarensis Fiedler, 1938: 591. 

PT: Dakar [Afr. Occ, Dakar /det. H. Fiedler, 1937]; PT: Dakar [Sénégal, Dakar, 

Fr. West Africa / det. H. Fiedler, 1937]. 
Ceroplesis aestuans ubangiensis Fiedler, 1938: 592. 

PT: Uganda [Brit. Uganda, Graner / det. H. Fiedler, 1937]; PT: Fort Crampel 

[det. H. Fiedler, 1937]. 
Ceroplesis aulica jokoana Fiedler, 1938: 603. 

?PT: [Sd. Kamerun, Heyne, Berlin- Wilm. / det. H. Fiedler, 1937]; ?PT: 

[Tykargeb., ill., VIII. 1912 / det. H. Fiedler, 1937]. 
Ceroplesis calabarica lemekensis Fiedler, 1938: 597. 

PT: Nyanza Province, Salt Marsh [B.E.A]. 
Ceroplesis calabarica ubangica Fiedler, 1938: 594. 

PT: Fort Crampel [Long, F.2 / det. H. Fiedler. 1937]; PT: Lado [Brit. Afr., A 

Heyne, Berlin- Wilm. / det. H. Fiedler, 1937]; ?PT: [Bamenida, 7.VII.05, Sch. / 

det. H. Fiedler, 1937]; ?PT: [Congo français / det. H. Fiedler, 1937]. 
Ceroplesis capensis bamumica Fiedler, 1938: 577. 

PT: Bamum [Kamerun, Kuti, 1040m / det. H. Fiedler, 1937]. 
Ceroplesis capensis camerunensis Fiedler, 1938: 577. 

PT: Fort Crampel [Congo français / det. H. Fiedler, 1937]. 
Ceroplesis capensis usambarica Fiedler, 1938: 575. 

PT: Tanga [det. H. Fiedler, 1937]. 
Ceroplesis hauseri conjuncta Hintz, 1920 (1918): 172. 

HT: Deutsch-Ostafrica, Njassasee. 
Ceroplesis hottentotta natalensis Fiedler, 1938: 612. 

2 PT: Natal [det. H. Fiedler, 1937]. 
Ceroplesis orientalis natalensis Fiedler, 1938: 605. 

PT: Natal [det. H. Fiedler, 1937]. 
Ceroplesis semitrabeata nguruana Fiedler. 1938: 608. 

?PT: [Africa occid., Coll. Nonfried / det. H. Fiedler, 1937]. 



CATALOGUE DES TYPES DE LAMIINAE 161 

Ceylanoparmena loebli Breuning, 1971b: 306. 

HT: Ceylan Central, Hatton, montagne boisée à l'est de la ville, 9.II.1970, 
1400m, tamisage en forêt [Mussard, Besuchet et Lobi]. 

Chariesthes nigroapicipennis Breuning, 1977a: 266. 
PT: Doum, Akono [12.VII.70 / Test cacao]. 

Cleptometopus assamanus Breuning, 1967a: 32. 

HT: Inde, Manipur, 30. VIII. 1960, leg. Schmid [Assam-Manipur, Langdang, 

5300ft]; PT: idem [Assam-Manipur, Phaiphengmu, 5000ft]. 
Cleptometopus grossepunctatus Breuning, 1967a: 32. 

HT: Assam, leg. Schmid [Kameng]. 
Cleptometopus parolivaceus Breuning, 1972c (1971): 416. 

HT: Sikkim, Chumtang, 17.VII.1959, 1550m [Himalaya, Schmid]. 
Cleptometopus scutellatus Hüdepohl, 1996: 3. 

HT: S, Sumatra , Jambi, W Mt Tujuh Lake, 1400m, 14.XI.1989, Agasti, Löbl, 

Burckhardt. 
Cleptometopus sikkimensis Breuning, 1972c (1971): 416. 

HT: Sikkim, Manu, 10.V1959, 1500m [Himalaya, Schmid]. 
Cleptometopus subteraureus Breuning, 1967a: 33. 

HT: Western Bengale, 15.XII.1959, leg. Schmid [Git Dabling, 4900-5100ft]. 

Conizonia chehirense Breuning, 1943: 103. 

HT: Anatolie, Ak-Chéir [1900]; ?PT: [Anatolien, Ak-Chehir, 1900, Korb]. 

Coptops illicita rosacea Breuning, 1980e: 157. 
2?PT: [Insel Mindanao]. 

Cristoberea assamensis Breuning, 1954a: 531. 
2 ?PT: [Assam, Shillang]. 

Cristorhodopina mussardi Breuning, 1966c: 15. 

HT: Assam, 2.VI.1961, leg. Schmid, coll. Mussard [Kameng, Ihum La 2400m]. 

Cylindrepomus grammicus hecate Dillon et Dillon, 1948: 274. 
PT: I. Salomon, Bougainville. 

Daphisia formosana Schwarzer, 1925: 153. 

ST: Formosa, Taihorin, 1911, H. Sauter. 

Dasylinda javanica Vuillet, 1912: 20. 

ST: Java, Gounod Gedeh, Ledru 1898, coli. Oberthür. 

Declivocondyloides loebli Sudre, 2000: 82. 

HT: New Caledonia, Col des Roussettes, litter in ravine, 12.IX.1998, I. Lobi; 
PT: idem; PT: New Caledonia, La Trappe above St Louis, wood and leaf litter, 
22.X. 1998, 1. Lobi. 



162 J- MORATI 

Diadelia betschi Breuning, 1975: 260. 

PT: Madagascar Centre, Andringitra Est, Ambalamarovandana, 1500- 1600m, 
FDHMA, 15-25.1.1971 [15.1.1971]. 

Dolichoprosopus philippine nsis Breuning, 1980e: 171. 
?PT: [He Mindanao, coll. Lumawig]. 

Dorcadion blandulus Holzschuh, 1977: 131. 

PT: Anatolien, prov. Bingöl, östlich des Kuruca geçidi, 1700m, 29.IV. 1975, leg. 

C. Holzschuh et F. Ressi. 
Dorcadion brauni Breuning, 1979a: 92. 

6 PT: Anatolia oca, Tavas-ovasi ca. 1000m, 26.III.1978, Heinz leg., coll. Heinz; 

4 PT: Anatolie oca, Tavas, Heinz, 1000m, III. 1978; 2 PT: Anatolia Tavas, 

1000m, 111.78. 
Dorcadion calabricum Breuning, 1942: 127. 

?PT: [Calabrien, Antonimina, leg. Paganetti]. 
Dorcadion czipkai Breuning, 1973c: 54. 

3 PT: Iran, Azarbaijan, Savelan, 2600m, 29.VII.1972, leg. W. Heinz. 
Dorcadion divisum subdivisum Breuning, 1955b: 263. 

PT: Anatolien, Ankara, V.1937, leg. Dr N. Vasvary. 
Dorcadion laevepunctatum Breuning, 1944: 14. 

?PT: [Maced., Ali Botus Exp., Maran et Taborsky Igt.]. 
Dorcadion pedestre kaszabi Breuning, 1956a: 404. 

?PT: Hongrie centrale, Dömsöd, Apajpuszta, V1953, Kaszab [1952, szikes leg- 

elo / egyelve IV.29 / leg. I. Kovacs]. 
Dorcadion pseudinfernale Breuning, 1943: 94. 

?PT: [Mts Amanus]. 

Driopea schmidi Breuning, 1972c (1971): 418. 

HT: Himalaya, Kumaon, Pauri Garhwal, Tarsali, 6.V1958, 1900-2100m [F. 
Schmid]. 

Eczemotes saintaignani Breuning, 1982: 16. 

PT: Papouasie, archipel de la Louisiade, île Saint-Aignan, Meek coll., VIII- 
XI. 1897; ?PT: [St Aignan, Vili to XI.97, Meek]. 

Elongatohomelix mortoni Breuning, 1967a: 29. 

HT: S , Sumatra, Baros, ex coll. W. Morton. 

Enicodes kaszabi Breuning, 1978a: 23. 

PT: New-Caledonie, I. Lifou, Bjengen, 21.11.1977, leg. Dr J. Balogh 
[22.11.1977]; ?PT: [New-Caledonie, I. Lifou, Ouaméne, 22.11.1977, leg. Dr J. 
Balogh]. 

Entelopes fuscotarsalis Breuning, 1954a: 480. 

?PT: [Bornéo, Matang RD., 5.II.1915]; ?PT: [idem, mais 20.1.16]. 



CATALOGUE DES TYPES DE LAMIINAE 163 

Entelopes griseipennis Breuning, 1954a: 479. 

2 ?PT: [Bornéo occi, Pontianak]. 
Entelopes jonoptera sumatrana Breuning, 1950b: 15. 

2 ?PT: [Sumatra]. 
Entelopes subsimilis Breuning, 1968a: 37. 

?PT: [B. Van Heua, 30.IV.65]. 

Epaphra minor Heller, 1934: 283. 

ST: Luzon, Prov. Laguna, prope Paete, F. C. Hadden leg., ex coll. Dr Reinh. 
Meyer, Darmstadt [Pangil, Laguna, PI.]. 

Epectasis hiekei Breuning, 1974e: 157. 
PT: Mexico, Cerro de Plumas. 

Epepeotes spinosoides Breuning, 1980e: 170. 

?HT: lie Mindanao, coll. Lumawig, MNHN [ill., Wallace]. 

Eryssamena besucheti Breuning, 1972c (1971): 417. 

HT: Assam, Kameng, Talung Dzong, 12.V.1961, 2100m [Himalaya, F. Schmid]. 
Eryssamena schmidi Breuning, 1972c (1971): 417. 

HT: Himalaya, Kumaon, Pauri Garhwal, Pana, 10.VIII.1958, 2500m 

[F. Schmid]. 

Essisus vivesi Breuning, 1978c: 135. 

?PT: [Australie, Brookfield, J. Sedlacek leg., XII.76]. 

Eunidia albosignata Breuning, 1972a: 281. 

?PT: [Zaire, Mambasa, Ituri]. 
Eunidia boafoi Breuning, 1978d: 94. 

?PT: [Ghana, Tafo, LX-X.1967, Boafo]. 
Eunidia flavoapicata tchadensis Lepesme & Breuning, 1955: 854. 

2 PT: Air, Oued Eir, 14.VIII.1949. 
Eunidia fulvescens Breuning, 1976b: 737. 

HT: Kenya, Neu-Moschi, 800m, IV 1912; 2 PT: idem. 
Eunidia nigroapicaloides Breuning, 1976a: 207. 

2 ?PT: [Chota-Nagpore, Nowatoli, V-VI.1896, R.P Cardon]. 
Eunidia pseudosenilis Breuning, 1970c (1971): 664. 

?PT: [Botswana, Serowe, Swaneng Hill, 20.XI.79, U.V., Forchhammer leg., 

n°5123]. 
Eunidia vagevittipennis Breuning, 1981b: 56. 

?PT: [ill., Iringa D.O.A., coll. Plasoo]. 
Eunidia varicolor Breuning, 1971c: 381. 

PT: Ghana, Kumasi. 

Exocentrus assamensis Breuning, 1972c (1971): 419. 

HT: Assam, Kameng, Nyukmadong, 2.VIII.1961, 2000-2400m [Himalaya, 
F Schmid]. 



164 J- MORATI 

Exocentrus celebicus Breuning, 1956c: 249. 

?HT: Celebes, Kandelari, IV. 1874, leg. O. Beccari, MCSN [Celebes]. 
Exocentrus fuscipes Breuning, 1981a: 51. 

PT: Kenya, Arusha, XI. 1961, P.P. De Moor. 
Exocentrus holonigra Breuning, 1972c (1971): 419. 

HT: Assam, But, 23. VI. 1961, 1750m [Himalaya, F. Schmid]. 
Exocentrus laosicus Breuning, 1963b: 47. 

?PT: [Laos, Wong Tevada, Vientiane, 9.X.62, Coll. Rondon]; ?PT: [id. mais 

16.X.62]. 
Exocentrus misellomimus Breuning, 1963b: 48. 

?PT: [Laos, Wong Tevada, Vientiane, 18.X.62, Coll. Rondon / M]; ?PT: [id. 

mais 28.X.62]. 
Exocentrus par av arie gatus Breuning, 1981a: 51. 

?PT: [Tvl, Rustenburg, 1-2.XII.1957, D.W. Rorke]. 
Exocentrus parcus Holzschuh, 1984: 157. 

PT: Népal, Arun- Valley, Dhankuta, Khantbari-Arunthan, 11 00- 1300m, 

29.V1938, leg. Holzschuh. 
Exocentrus ravillus Holzschuh, 1984: 158. 

PT: E. Népal, Arun- Valley, Dhankuta, Lamobagar, 1400m, 8-14.VI.1983, 

C. Holzschuh. 
Exocentrus rondoni Breuning, 1963b: 47. 

?PT: [Laos, Phon Tion, 10.VI.63, Coll. J. A. Rondon]; ?PT: [id. mais 22.VI.63]. 
Exocentrus unialbovittatus Breuning, 1969a: 36. 

HT: Congo, Haut-Katanga, Panda, IV 1929, leg. J. Romieux. 
Exocentrus vitticollis Breuning, 1974c: 418. 

PT: Ethiopie, Prov. Kaffa, Mui, 700m, 10.IV1972, leg. R.O.S. Clarke [ref. 0550 

N/3545E]. 

Falsepilysta rossetti Breuning, 1982: 10. 
2 ?PT: [Rössel Is., I-III.98, Meek]. 

Frea mussardi Breuning, 1969a: 36. 

HT, PT (allotype): S, Côte-dTvoire, Adiopé, VII. 1948, leg. Mussard. 

(Lì allotype manque). 
Frea taverniersi Breuning, 1973a: 96. (orthographié tavernieri dans la collection.) 

?PT: [Zaïre, Kisangani]. 

Glenea assamana Breuning, 1967a: 34. 

HT: o\ Assam, leg. Schmid [Khasi-Faintia Hills, Mawpran, 900-2000ft]; 3 PT: 

idem. 
Glenea bedoci mussardi Breuning, 1967a: 35. 

HT: ó\ N.E.Assam, 23.V1961, leg. Schmid [Kameng, Amatulla, 1300-2000ft]. 
Glenea besucheti Breuning, 1974a: 74. 

HT: 9 , Philippines, He Luzon, Manila, Calo, VI 928, W Parsons. 
Glenea bougainvillei Breuning, 1958b: 315. 

HT: Bougainv. 



CATALOGUE DES TYPES DE LAMIINAE 165 

Glenea mira bernardii Breuning, 1977a: 298. 

PT: Gabon, mont Bengue, 6.X.1967, leg. G. Bernardi. 
Glenea nitidicollis rufina Breuning, 1976b: 740. 

HT: Indonésie. 
Glenea perakensis Breuning, 1956b: 126 (Breuning, 1956d: 699). 

3 ?PT: [Malaisie, Perak, Gounong Boubou, 1100m, W. Doherty]. 
Glenea peregoi sumatrensis Breuning, 1976b: 740. 

HT: Sumatra, Benkoelen, Moeara Tenam. 
Glenea pici schmidi Breuning, 1967a: 35. 

HT: Ô, N.E.Assam, 23.V.1961, leg. Schmid [Kameng, Amatulla, 1300-2000ft]. 
Glenea pseudogiraffa taverniersi Breuning, 1974d: 777. 

?PT: [Ht Zaïre, Kisangani]. 

Glenida cyaneo fasciata Breuning, 1952: 116. 
?PT: [British Bootang, J. Durel]. 

Gnoma blanchardi Breuning, 1945 : 533. 

?PT: [St Aignan, VIII-IX.97, Meek]. 
Gnoma? plumigera Westwood, 1 848: 1 1 . 

HT: Java, coll. A. Melly [Himilaya Mount, Melly]. 

Goephanes ruficornis Breuning, 1964a: 97. 

HT: Madagascar, Diego-Suarez [Coll. G. Junod]. 

H oplorana mus sardi Breuning, 1958 (1957): 177. 

HT: S , Madagascar Est, Antalaha, coll. Mussard [XI.56]. 

H oplothrix paramicator Breuning, 1966c: 14. 

HT: Inde, Manipur, 28.VIII.1960, leg. Schmid, coll. Mussard [Khaozang, 
3750ft]. 

Hyagnis indicus Breuning, 1969a: 35. 

HT: Inde, Madumalai, 19.11., voyage Carl et Escher. 

Hyllisia suturaloides Breuning, 1981a: 44. 

PT: Ost Transvaal, Trichardtsdal, 25.XI.1972, H.J. Botha. 

Inermoparmena besucheti Breuning, 1971b: 304. 

HT: Ceylan Central, Horton Plains, 2100m, 15.11.1970, tamisages en forêt 
[Mussard, Besuchet et Lobi]; 4 PT: idem; 3 PT: Hakgala, versant NE, 1700- 
1800m, 28.1.1970, tamisages en forêt; 3 PT: Pidurutalagala, versant SO, 2200m, 
tamisages en forêt. 

Jordanoleiopus mirei Breuning, 1977a: 294. 

PT: Cameroun, Nkom (Saa), 24.IX.1971, leg. Ph. de Miré. 



166 J- MORATI 

Lasiocercis hovanoides Breuning, 1964a: 96. 

HT: 9 , Madagascar, Diego Suarez, ex. coll. Junod. 
Lasiocercis truncatoides Breuning, 1975: 258. 

2 PT: Madagascar Centre, Adringitra Est, Anjavidilava, 2000m, FDSMP, sur 

Cussonia sp. (Araliacées), 18.XH-15.I.1971. 

Linda annamensis Breuning, 1954a: 556. 

?HT: 9, Annam, Caleu, ZMUH [Annam, Caleu]. 
Linda assamensis Breuning, 1954a: 555. 

?PT: [Assam]. 
Linda bimaculicollis Breuning, 1954a: 558. 

?PT: [Assam, rég. de Shillong] ; ?PT: [Assam, Khasi Hills]. 
Linda javaensis Breuning, 1954a: 557. 

HT: 9 , Java. 
Linda tonkinensis Breuning, 1959b: 172. 

?PT: [Tonkin, Mts Mauson, Fruhstorfer]; ?PT: [Tonkin, Montes Mauson, 

2-3000', IV- V., H. Fruhstorfer]. 

Mahenes denteiti Breuning, 1980d: 50. 

HT: Iles Seychelles, Beau Vallon, VII. 1978, leg. C.V. Demelt [Mahé]. 

Mallosia imperatrix kurdistana Breuning, 1970d: 30. 
HT: â , Kurdistan. 

Mecas albovitticollis Breuning, 1955c: 146. 
?PT: [Mexico]. 

Megalofrea transvaaliensis Breuning, 1981a: 48. 

PT: Transvaal, Warmbaths, 1.1917, G. Kobrow. 

Menesia bicoloricornis Breuning, 1963a: 6. 

HT: 9 , Sumatra, Bukoelen, Moeara-Tenam, leg. M.E. Walsh [1935]; PT: idem. 
Menesia gleneoides Breuning, 1965c: 39. 

?PT: [Laos, Parkading, 27.VI.64, Coll. Rondon]; ?PT: [Parkading, 20.V63]. 
Menesia laosensis Breuning, 1963c: 20. 

?PT: [Loc Wa, Km 17, Pa Sang, Vientiane, 1.IV.62]; ?PT: [Km 17, Pa Sang, 

Vientiane, 31. V.62]. 
Menesia latevittata Breuning, 1954a: 425. 

2 ?PT: [Borneo, Sandakan, Baker]. 
Menesia longitarsis Breuning. 1954a: 417. 

2 PT: Borneo. Sandakan [Baker]. 
Menesia ochreicollis Breuning, 1954a: 422. 

?PT: [Bornéo, Sandakan. Baker]. 
Menesia octoguttata Breuning, 1954a: 411. 

HT: 9 . China. Provinz Szetschuan, Tatsienlu, leg. Stötzner; ?PT: [Su-Tchuen, 

Mo-Sy-Mien]. 



CATALOGUE DES TYPES DE LAMIINAE 167 

Menesia sexvittata Breuning, 1962b: 14. 

HT: Java, Malang; ?PT: [Java occ, Mt Gedeh]. 
Menesia subguttata Breuning, 1954a: 414. 

HT: 9 , Borneo, Sandakan. 

Menesida flavipennis Breuning, 1954a: 446. 

?PT: [Sumatra, Medan]. 
Menesida rufula Breuning, 1954a: 448. 

HT: S , Java, Pengalengan, 1300m, leg. H. Fruhstorfer; ?PT: [Java, Preanger, 

Mts Djampangs]. 

Micromandibularia ruf a Breuning, 1954a: 528. 
HT: ? , Indien, Manipur. 

Microsomatidia reticulata Sudre, 2000: 80. 

HT: New Caledonia, Mt. Koghi, prim, for., 400-500m, litter, 18-19.X.1998, 
I. Lobi; PT: idem; PT: New Caledonia, Col de Mouirange, nr Table d'Union, 
260m, litter, 16.IX.1997, 1. Lobi. 

Microrhodopis albovittata Breuning, 1976b: 740. 

HT: Malaisie, Fräsers Hill, 17-30.XI.1974, leg. P. Pfanner. 

Mimeryssamena besucheti Breuning, 1972c (1971): 418. 

HT: Himalaya, Kumaon, Pauri Garhwal, Mandoli, 1. VI. 1958, 2300m 
[F. Schmid]. 

Mimeuseboides exacavatipennis Breuning, 1967a: 33. 

HT: N.E. Assam, 13.V.1961, leg. Schmid [Kameng, Jhum La, 7800ft]. 

Mimocrossotus rhodesianus Breuning, 1972a: 283. 
?PT: [Rhodésie, Salisbury, 17.X.1971]. 

Mimomorpha flavopunctata Breuning, 1980e: 178. 
2 ?PT: [Insel Mindanao]. 

Mimophaeopate assamensis Breuning, 1967a: 31. 

HT: N.E. Assam, 23.V.1961, leg. Schmid [Kameng, Amatulla, 1300-2000ft, 
23.V.61]. 

Mimostedes decellei Breuning, 1968b: 347. 

PT: Côte d'Ivoire, Bingerville, 16.11.1964, J. Decelle; PT: id. mais 23.11.1964; 
PT: id. mais 1-8.III.1964. 

Mispila tonkinensis Breuning, 1964a: 96. 

HT: Tonkin, Langson, Ban-danh [E. Autran, 1898 / P de Borre]. 

Moechotypa formosana Breuning, 1974b: 372. 

HT: Formose, Puli Taitung, V-VI.71, leg. K.H. Chen. 



168 J- MORATI 

Monochamus densepunctatus Breuning, 1980e: 173. 

?PT: [Insel Mindanao]. 
Monochamus kinabaluensis Hiidepohl, 1996: 5. 

HT: 9, Ost-Malaysia, Sabah, Mt Kinabalu, 1500m, 30.VI.1987, Burckhardt, 

Lobi leg. [30.IV. 1987]. 
Monochamus marmoratipennis Breuning, 1974a: 74. 

HT: Côte d'Ivoire, Sassandra, 1.1950, R. Mussard. 
Monochamus subtrangularis Breuning, 1972c (1971): 416. 

HT: Assam, Kameng, Bhairabkunda, 19.V1961, 200-300m [Himalaya, 

F. Schmid]. 

Monohammus bifasciatus Westwood, 1848: 59. 

HT: Himalayas, coll. A. Melly [Saunders Harrington]. 
Monohammus punctulatus Westwood, 1848: 12. 

HT: Himalayas, coll. A. Melly. 
Monohammus westwoodii Westwood, 1848: 12. 

HT: Himalayas, coll. A. Melly. 

Morimopsis assamensis Breuning, 1966c: 14. 

HT: 6 , Assam, North East Frontier Agency, Kaliktang, 14.V1961, leg. Dr F. 

Schmid, coll. Mussard; PT (allotype): 9, dito, Talung-Dzong, 2300m alt., 

4.VI.1961, Dr Schmid, dtlo., coli. Mussard. 
Morimopsis mussardi Breuning, 1966c: 14. 

HT: ô, Assam, North-East Frontier Agency, Dirang Dzong, 1800m alt., 

21. VII. 1961, leg. Dr Schmid, coll. Mussard. 

Mussardia griseoplagiata Breuning, 1959a: 5. 

HT: Congo belge, Uelé, coll. Mussard [Bili 1952, Verbeke]. 

Mutatocoptops lumawigi Breuning, 1980e: 157. 
?PT: [Insel Mindanao]. 

Nemaschema flavovittatum Breuning, 1976b: 739. 

HT: Nouvelle Calédonie, Sana Pirogue, 4.XII.1952. 
Nemaschema ochreovittatum Breuning, 1978a: 23. 

PT : Nouvelle Calédonie, Mont Rembai, 19-21.1.1977, leg. Dr J. Balogh. 

Neodorcadion altaicum Suvorov, 1909: 89. 

ST: Altai, village Causaque de Bolschenarim, 08. VI. 1906, ree. A.G. Jacobson 
[Semipalatinckaya obi. Altay Irtysh ct. Volsheparym. 8. VI. 1906. A.Yakobson 
(en cyrillique)]. 

Neonitocris rhodesica Breuning, 1950c: 29. 

?PT: [Kafakumba, XL 1933. FG. Overlaet]. 

Neoserixia pulchra Schwarzer, 1925: 148. 

ST: Formose, Kankau, V 1912, H. Sauter [Koshun]. 



CATALOGUE DES TYPES DE LAMIINAE 169 

Niphonatossa mussardi Breuning, 1967a: 29. 

HT: 9, Assam, Cachar District, 9. V. 1960, leg. Schmid [Bandarkhal, 500ft, 
9.V.60, Schmid]. 

Niphoparmena basilewskyi Breuning, 1960c: 336. 
PT: Tanganika. 

Nitocris funesta Fâhraeus, 1872: 57. 

?PT: [Caffraria]. 
Nitocris maculicollis Peringuey, 1888: 184. 

ST: ill., Stalbord, 11.87/9. 
Nitocris sanguinalis Kolbe, 1893 : 77. 

ST: Togo, Bismarckburg, VI. 1891. 

Nonyma attardi Breuning, 1972a: 284. 

HT: Zaïre, Ituri, Mambasa. 
Nonyma nigeriae Breuning, 1978d: 108. 

?PT: [Nigeria, Ibadan, 15.1.1963, D.C. Eidt]. 

Nupserha schmidi Breuning, 1966c: 16. 

HT: 9, North-East Assam, Biloda, 2000m alt., VI.1961, leg. Schmid, coll. 

Mussard [Assam- Kameng, Yhum-La, 16.IX.61]. 
Nupserha similis Breuning, 1978d: 113. 

?PT: [Tanganica, W. Usambara Mts, 2000m, Maganta, III. 1962]. 
Nupserha tanganjicae Breuning, 1978d: 112. 

PT: Tanzanie, Mts Rungwe, 2600m, XI. 1962 [5-10.XI.62]; 3 ?PT: [idem]. 
Nupserha tanganjicae uluguruensis Breuning, 1978d: 113. 

2 ?PT: [Tanganyika, Ulguru Mts, 1500-1 800m]. 
Nupserha ugandensis Breuning, 1978d: 111. 

?PT: [Uganda, Entebbe, II- VII. 1972, Exp. H. Falke]. 

Oberea andamana Breuning, 1962a: 152. 

?PT: [Andaman Islands]. 
Oberea andamanica Breuning, 1962a: 152. 

2 PT: lies Andamanes. 
Oberea angolana Breuning, 1961b: 62. 

HT: ô , Angola, Coconda [Bailundo]. 
Oberea angolensis Breuning, 1950d: 273. 

2 ?PT: [Angola, Bailundo]. 
Oberea annamensis Breuning, 1969a: 37. 

HT: S, Annam [Coll. Junod]; 2 ?PT: [Annam]. 
Oberea anterufa Breuning, 1962a: 142. 

HT: 9 , Philippines, He Dalapiri. 
Oberea apicenigrata Breuning, 1962a: 170. 

?PT: [Laos, Yuang-Prabang]. 



170 J- MORATI 

Oberea artocarpi andamanensis Breuning, 1962a: 147. 

?PT: [And. Is.]. 
Oberea aterrima Breuning, 1961b: 100. 

HT: o\ Celebes, Tombugu [H. Kuhn, '95]. 
Oberea auriventris Breuning, 1961b: 118. 

2 PT: Borneo, Sandakan, Baker. 
Oberea baliana Breuning, 1961b: 117. 

HT: 9, Ile Bali, IIMV.1896, leg. Doherty. 
Oberea bangueyensis Breuning, 1950b: 25. 

?PT: [Nord Borneo, Banguey]; ?PT: [Banguey I., Staudinger]. 
Oberea batoensis nigrata Breuning, 1961b: 101. 

PT: Sumatra, Medan; ?PT: [idem]. 
Oberea bicoloripennis Breuning, 1950c: 43. 

?PT: [Zambesi, Boroma]. 
Oberea bimaculicollis Breuning, 1962a: 191. 

HT: ô , Cambodge. 
Oberea bisbimaculata Breuning, 1962a: 190. 

HT: 9 , Tonkin, Chapa. 
Oberea bisbipunctata discoreducta Breuning, 1969a: 37. 

HT: 9 , Annam [coll. Junod]. 
Oberea bisbipunctulata Breuning, 1962a: 191. 

PT: British Bootang, 1898, L. Durel; PT: idem, mais Padong, 1914. 
Oberea bootangensis Breuning, 1962a: 177. 

HT: 6, Bootan [L. Durel]; PT: 9 : idem [Maria Basti]. 
Oberea callosicollis Breuning, 1962a: 171. 

?PT: [Bengal, Chapra]; ?PT: [Tenasserim, Tandong, 4000', Mai, Fruhstorfer leg.]. 
Oberea cariniscapus Breuning, 1956g: 23. 

?HT: Java, IRSN [Malacca, Perak]. 
Oberea circumscutellaris Breuning, 1961b: 96. 

HT: â , Ile Engano. 
Oberea davaoensis Breuning, 1961b: 139. 

?PT: [Ilayavo □□□, Birlig /Philipp., Sumpei); ?PT: [Philipp., Sumpei). 
Oberea densepilosa Breuning, 1955a: 76. 

?PT: [Abyssinia, Harrar). 
Oberea densepunctipennis Breuning, 1962a: 144. 

?PT: [Philippinen, Luzon, Prov. Laguna). 
Oberea elongaticollis Breuning. 1961b: 140. 

?PT: [Mindanao. Davao). 
Oberea erythrocephala palawanicola Breuning, 1964b: 6. 

HT: ó\ Iles Palawan; ?PT: [Mindoro). 
Oberea flava Breuning, 1961b: 122. 

HT: o\ Celebes, Toli-Toli; ?PT: [S. Celebes, Bua-Kraeng, 5000', 11.1896, 

H. Fruhstorfer). 
Oberea flavescens Breuning, 1947c (1946): 146. 

HT: 9 , Chine, province Sétchouen, Tatsienlou. 



CATALOGUE DES TYPES DE LAMIINAE 1 7 1 

Oberea flavoantennalis Breuning, 1961b: 125. 

HT: 9 , Celebes, Tomboegoe. 
Oberea flavoantennata Breuning, 1962a: 146. 

?PT: [Philippinen, Luzon?, Malacia?]. 
Oberea florensis Breuning, 1961b: 130. 

HT: 9 , South Flores, X.1896, leg. Everett; PT: o\ idem. 
Oberea floresica Breuning, 1961b: 131. 

?PT: [Flores]. 
Oberea fulviceps Breuning, 1950c: 38. 

?PT: [Cameroons, Bitye Ja River]; ?PT: [Kamerun]. 
Oberea fuscicollis Breuning, 1961b: 138. 

2 ?PT: [Philippinen, Mindanao, Surigao]. 
Oberea fuscipennis perakensis Breuning, 1962a: 179. 

?PT: [Annam, Quinhone]; ?PT: [Tonkin, Montes Manson, IV-V., 2-3000', 

H. Fruhstorfer]. 
Oberea gabunensis Breuning, 1950c: 39. 

2 ?PT: [Gabon, Bas Ogoué]. 
Oberea griseopennis Schwarzer, 1925: 154. 

ST: Formose, Kankau, Koshun, H. Sauter [V.1912]. 
Oberea grossepunctata Breuning, 1947c (1946): 145. 

HT: 9 , Chine, prov. du Yunnan, Fleuve Soling-ho [Vallis flumin.]; ?PT: [Chin., 

Yunnan, El. Solingho]. 
Oberea himalayana Breuning, 1972c (1971): 420. 

HT: 9, Assam, Kameng, But, 23.VI.1961, 1750m [Himalaya, Schmid]. 
Oberea holatripennis Breuning, 1982: 29. 

?PT: [Pekin / Ex. Musaeo, Arm. David, 1900]. 
Oberea holonigra Breuning, 1961b: 97. 

HT: â , Ile Engano. 
Oberea infranigra Breuning, 1962a: 181. 

HT: 9 , Cambodge. 
Oberea javana Breuning, 1961b: 105. 

HT: ò\ Java, Ganung, Tangkuban Prahu [1000-1800m, V.1933]. 
Oberea javanicola Breuning, 1950b: 28. 

?PT: [Java mend., 1500', 1891, H. Fruhstorfer]; ?PT: pas d'étiquette de localité. 
Oberea kanarensis Breuning, 1950b: 28. 

?HT: Inde, Kanara, IRSN [C.P. Ceylon, Pundaloya]; ?PT: [N. Kanara, T.R. 

Bell]; ?PT: [Kanara]. 
Oberea kandyana Breuning, 1962a: 153. 

PT: Ceylan, Kandy; ?PT: [W.P. Ceylon, Henaralgada, Bot. Gards]. 
Oberea keyensis Breuning, 1961b: 128. 

2 ?PT: [Little Kei., H. Kühn]. 
Oberea kunbirensis Breuning, 1953: 37. 

?PT: [India, Kunbir]. 
Oberea lepesmei Breuning, 1956d: 701. 

?PT: [Côte d'Ivoire, Dirubalu]. 



172 J- MORATI 

Oberea lepesmiana Breuning, 1956d: 700. 

?PT: [Malay Ar., Xulla-Mangoli, X-XI, Doherty]; ?PT: [Sula Mangoli, X-XI, 

Doherty]. 
Oberea luluensis Breuning, 1950c: 40. 

?PT: [Cameroons, Bitye Ja River, 3000ft]. 
Oberea lutea nilghirica Breuning, 1955a: 77. 

2 PT: Inde, Nilghiri Hills. 
Oberea manipurensis Breuning, 1961b: 98. 

HT: S , Inde, Manipur; PT: idem. 
Oberea matangensis Breuning, 1962a: 179. 

?PT: [Borneo, Baker]. 
Oberea medioflavoantennalis Breuning, 1961b: 124. 

HT: 9 , Celebes, Tomboegos. 
Oberea mediofusciventris Breuning, 1962a: 171. 

HT: ô , South East Borneo. 
Oberea mentaweiensis Breuning, 1961b: 108. 

HT: o\ Iles Mentawei, Ile Sipora, VI. 1924, leg. Karny. 
Oberea meridionalis Breuning, 1961b: 71. 

?PT: [Cap. B. Spei.]. 
Oberea nigerrima Breuning, 1950b: 25. 

?PT: [Java occident., Pengalengau, 4000, 1893, H. Fruhstorfer]; ?PT: pas d'éti- 
quette de localité. 
Oberea nigrescens Breuning, 1961b: 110. 

PT: Kuching, X.1906; ?PT: [idem mais V.1907, J.H]. 
Oberea nigripennis Breuning, 1950c: 38. 

PT: Congo B., Katanga, riv. Kawa; ?PT: [Gabon, Bas Ogoué]. 
Oberea nigripennis camerunica Breuning, 1961b: 92. 

?HT: $, Cameroun, Ja River, Bitye, ex. coll. Oberthür, MNHN [Cameroun, 

Batanga]; ?PT: [Cameroons, Bitye Ja River, 3000ft.]. 
Oberea nigripes Breuning, 1950b: 26. 

?PT: [Java, Soekaboemi, Coll. Le Moult]; ?PT: [Java occid., Sukabumi, 2000', 

1893, Fruhstorfer]. 
Oberea nigroapiciventris Breuning, 1962a: 167. 

?PT: [Borneo mer.]. 
Oberea nigrobasipennis Breuning, 1950c: 38. 

2 PT: Congo beige, Eala. 
Oberea nigrofemoralis Breuning, 1950c: 39. 

?PT: [Cameroons, Bitye Ja River, 3000ft]: ?PT: [Kamerun, Mukoja, Farm]. 
Oberea nigrolateralis Breuning, 1950b: 27. 

2 PT: Java, Soekaboemi. 
Oberea nigrolineatipennis Breuning, 1970e: 172. 

?PT: [Katanga, Zilo, IV.69]. 
Oberea nyassana Breuning, 1950b: 30. 

2 ?PT: [Mozambique, Zambesie]. 



CATALOGUE DES TYPES DE LAMIINAE 173 

Oberea occidentalis Lepesme & Breuning, 1953: 104. 

?PT: [Kumassi, Newberry]. 
Oberea octava kuchingensis Breuning, 1962a: 197. 

?PT: [Bornéo occ.]; ?PT: [Kuching, 11.1907, H.F.]. 
Oberea opacipennis Breuning, 1961b: 132. 

HT: 9 , Ile Sumbawa; PT: idem. 
Oberea ornativentris Breuning, 1961b: 70. 

HT: $ , Rhodésie; PT: 9 , Kenya. 
Oberea palawanensis Breuning, 1961b: 115. 

?PT: [N. Palawan, Binuluan, XI-XII.1913, leg. G.Boettcher]. 
Oberea parteflavoantennalis Breuning, 1961b: 101. 

HT: S, Bornéo, Sandakan, leg. Baker. 
Oberea partenigricollis Breuning, 1961b: 137. 

2 ?PT: [Philippin., Bilaran]. 
Oberea pedemontana koniensis Breuning, 1962a: 218. 

HT: 9, Anatolie, Eregli [Türkei, Konia, 1050m, 6.VII.1952, Eregli, leg. Erich 

Schmidt]. 
Oberea philippinensis Breuning, 1961b: 137. 

PT: Philippines, He Mindanao oriental; ?PT: [Mind.]. 
Oberea pontianakensis Breuning, 1962a: 176. 

HT: S , Bornéo, Pontianak; ?PT: [Borneo, Pontianak]. 
Oberea pseudobalineae Breuning, 1955a: 76. 

?PT: [Luzon, Manille]; ?PT: [Rio Baubo, 64]. 
Oberea pseudolacana Breuning, 1956d: 700. 

2 ?PT: [N. Borneo, Kina-Balu]. 
Oberea pseudonigrocincta Breuning, 1961b: 69. 

HT: o\Erithrée. 
Oberea pseudopascoei Breuning, 1950c: 41. 

2 ?PT: [Gabon, Bas Ozooné]. 
Oberea pseudoposticata Breuning, 1962a: 165. 

HT: Ô, Tenasserim, Meetan, IV. 1887. 
Oberea puncticollis Breuning, 1962a: 146. 

?PT: [Philippinen, Luzon, Prov. Apayao]. 
Oberea pupillatoides Breuning, 1947c (1946): 145. 

HT: 9, Chine, province du Chansi [26.VI.16]. 
Oberea ressli Demelt, 1963a: 150. 

3 PT: Asia minor sept., Kizilcahamam [62]. 
Oberea ruficeps muchei Breuning, 1981c: 26. 

HT: Pamir-Alai, Tadshikistan, environs de Siddi, 2000-2500m, I- VII. 1980, leg. 

Heinz Muche; 2 PT: idem, MHNG. {Les trois exemplaires n'ont pas été 

retrouvés). 
Oberea rufiniventris Breuning, 1968a: 4L 

?PT: [Pakkading, 15.IV66]; ?PT: [Bibu Ena, 15.IV65]. 
Oberea rufoantennalis Breuning, 1962a: 143. 

?PT: pas d'étiquette de localité. 



174 J. MORATI 

Oberea rufosternalis Breuning, 1961b: 95. 

PT: Yunnan, Peyentsin; ?PT: [Sud Yunnan, Tche-Ping-Tcheou]. 
Oberea sanghirica Breuning, 1961b: 100. 

HT: 9, Sanghir, Taroena, HI-IV, leg. Doherty [2000]. 
Oberea semifuscipennis Breuning, 1950b: 29. 

?PT: [Mai'ssour, Shimoga]; ?PT: [Nilgiri Hills, H.L. Audrewes, ill., Teppenkadu, 

V.l 1,2500']. 
Oberea semirubra Breuning, 1962a: 148. 

HT: 6 , Cochinchine, Cap St Jacques [Baria]. 
Oberea senegalensis Breuning, 1961b: 66. 

?PT: [Sénégal]. 
Oberea shibatai Hayashi, 1962: 40. 

PT: Amami-Oshima, Naze, 29.IV.1961, Y Susumu leg. 
Oberea shirahatai Ohbayashi, 1956: 762. 

2 PT: Nord Honshiu, Sakata, Yamagata Pref., 9. VII. 1952, leg. K. Shirahata. 
Oberea silhetica Breuning, 1961b: 117. 

HT: 9 , Inde, Silhet, Chandkiva. 
Oberea sobosana Ohbayashi, 1956: 761. 

PT: S, Kiushiu, Mt Sobo, Oita Pref., Kyushu, Mt Sobo, Oka, 23-26. VII. 1952, 

Y Miyake leg. [26. VII. 1952]. 
Oberea subabdominalis Breuning, 1962a: 166. 

2 ?PT: [Birmanie, Theinzelk, 1913, P. Loizeau / 167]. 
Oberea subbasalis Breuning, 1950c: 42. 

?PT: [P. Natal]. 
Oberea subferruginea Breuning, 1965a: 52. 

2 ?PT: [Kongsédone, 30.IV65]. 
Oberea subneavei Breuning, 1961b: 75. 

HT: â , Cameroun. Mukonje Farm; ?PT: [Cameroun, Yaoudé]. 
Oberea subnigrocincta Breuning, 1950c: 42. 

2 ?PT: [Cameroons, Bitye Ja River, 3000ft]. 
Oberea subsericea Breuning, 1962a: 180. 

HT: 9 , Chine, prov. Kwei-chow, Nanning. 
Oberea subsuturalis Breuning, 1954b: 19. 

?PT: [Kenya, Diatha]. 
Oberea subteraurea Breuning. 1961b: 118. 

HT: â, Bornéo. Sandakan. leg. Baker: ?PT: [Bornéo occ, Riv. Sambey près 

Noabang / 1897. J.B. Ledru]. 
Oberea subvaricornis Breuning. 1961b: 73. 

PT: Durban, XII.03: PT: Durban, Congella, G.F. Leigh. 
Oberea subviperina Breuning. 1960a: 15. 

?HT: Philippines. Mindanao, Surigao, Mus. Dresde [Mindanao, Surigao]. 
Oberea sumbana Breuning, 1961b: 131. 

PT: Waingapoe, 96, Everett; ?PT: [Sumba]. 
Oberea sumbawana Breuning. 1961b: 132. 

HT: 9 . Ile Sumbawa. 



CATALOGUE DES TYPES DE LAMIINAE 1 75 

Oberea sumbawanica Breuning, 1961b: 133. 

HT: â , Ile Sumbawa. 
Oberea sumbawensis Breuning, 1961b: 131. 

HT: â, Ile Sumbawa, Mt. Harunasa; ?PT: [Insel Sumbawa]. 
Oberea taihokuensis Breuning, 1962a: 168. 

HT: 9 , Formose, Taihoku [20.IV. 1932, Col. M. Chujo]. 
Oberea tatsienlui Breuning, 1947c (1946): 145. 

HT: S, Chine, prov. de Sétchouen, Tastsienlou [Tatsienlu Dsaschi / 

Bergschluchten, Em. Reitter]. 
Oberea tenggeriana Breuning, 1963a: 10. 

HT: 2 , Java, Mts Tengger, leg. Walsh. 
Oberea travancorensis Breuning, 1962a: 151. 

HT: S , Inde, Travancore. 
Oberea tricoloricornis Breuning, 1961c: 43. 

HT: ó\ Cameroun, Bitye, 1000m [3000ft]. 
Oberea trigonalis Breuning, 1950c: 42. 

?PT: [Transvaal, Hartmann]; ?PT: [ill., Zouberi]. 
Oberea truncatipennis Breuning, 1962a: 170. 

HT: 6 , South East Borneo. 
Oberea unimaculicollis Breuning, 1962a: 194. 

HT: 2 , Sumatra, Medan. 
Oberea wittei Breuning, 1954c: 78. 

2 PT: 9, Lusinga, 1760m, 28.XI-17.XII.1947 [Congo]. 
Oberea yunnanensis Breuning, 1947c (1946): 147. 

HT: 2 , Chine, province Yunnan, Fleuve Soling-ho; ?PT: [China, Prov. Yunnan, 

Vattis flumin, Soling-ho]. 

Obereopsis coimbatorana Breuning, 1974b: 374. 

HT: Inde, Madras, Coimbatore, 500m, XI. 1971, leg. T.R.S. Nathan; PT: idem. 
Obereopsis himalayana Breuning, 1972c (1971): 420. 

HT: 2, Himalaya, Kumaon, Pauri Garhwal, Binaik Chatti, 2.VII.1958, 2150- 

2250m [F Schmid]; PT: idem. 
Obereopsis nigriceps nigroabdominalis Breuning, 1972c (1971): 420. 

HT: Sikkim, Yugang, 24.VII.1959, 1750m [Himalaya, F. Schmid]. 
Obereopsis paratricollis Breuning, 1967a: 36. 

HT: 2 , N.E. Assam, leg. Schmid [Kameng]. 
Obereopsis trinotaticollis Breuning, 1967a: 36. 

HT: ó\ Inde, Manipur, 15.VI.1960, leg. Schmid; PT: idem [Assam, Chingsao, 

3800-5400ft]. 

Oeax collaris nigromaculatus Breuning, 1978d: 103. 
?PT: [Ghana, Tafo, VI. 1969, E.O. Boafo]. 

Ossonis modiglianii Breuning, 1950a: 200. 

2 ?PT: [Sumatra, Si-Rambé, XII.90-III.91, E. Modigliani]. 



176 J- MORATI 

Oxylamia vagemarmorata Breuning, 1974a: 74. 

HT: Ô, Côte d'Ivoire, Sassandra, R. Mussard, 1.1950. 

Oxylia androsensis Breuning, 1963a: 10. 

HT: <?, Grèce, Ile Andros, 27.IV. 1960 [24.IV.60]. 

Parabrimus ruficornis Breuning, 1981a: 46. 

PT: Transvaal, Mariepskop, 11.1962, N. Leleup. 

Paraclodia besucheti Breuning, 1974b: 374. 

HT: Inde, Kerala, Cardamon Hills, Periyar, 950m, 4.XI.1972, leg. Besuchet, 
Lobi, Mussard, attiré par les lumières de l'Hôtel Aranya Nivas. 

Paracondyloides flavofasciatus Breuning, 1978a: 25. 

?PT: [Nile Calédonie, Caavatch, 15.11.1977, leg. Dr J. Balogh]. 

Paradystus ceylonicus Breuning, 1954a: 460. 

HT: $ , Insel Ceylon. 
Paradystus infrarufus Breuning, 1954a: 460. 

HT: 9, Tenasserim, Tamdong, 1700m, mai, leg. Fruhstorfer. 
Paradystus innotatus Breuning, 1954a: 459. 

?PT: [SO Sumatra, Merang]; ?PT: [Bornéo, Sandakan, Baker]. 

Paraglenea latef asciata Breuning, 1952: 127 (Breuning, 1956d: 697). 

?PT: [Tonkin, Chapa, 1. VII. 1917, Jeanvoine]; ?PT: [Tonkin, Thanh-Moi, 24. Vi- 
li. 1917, Jeanvoine]. 

Paraglenea transversefasciata Breuning, 1952: 128. 
2 ?PT: [Laos, Lonang-Prahang]. 

Parahyllisia indica Breuning, 1974e: 163. 

PT: Indien, Kodaicanal [Indes Orientales, Mts. Kodaicanal, J. Castets]. 

Paralamiodorcadion schmidi Breuning, 1967a: 28. 

HT: $, Inde, Manipur, 9VI.1960, leg. Schmid [Assam]. 

Paramenesia nigrescens Breuning, 1966b: 53. 

HT: â, Tonkin, Monts Mauson, leg. Fruhstorfer [IV- V, 2-3000']. 

Paramimiculus pterolophioides Breuning, 1964a: 95. 

HT: 3 , Lourenço Marques, Makulane, leg. Dr G. Audeoud. 

Paraserixia flava Breuning, 1954a: 442. 
?PT: [Burma]. 

Parasmermus tonkinensis Breuning, 1969a: 36. 
HT: 6 , Tonkin, ach. Baudet. 



CATALOGUE DES TYPES DE LAMIINAE 177 

Parazygocera baloghi Breuning, 1978b: 69. 

2 PT: New Caledonie, Mt Mandjello, 17.11.1977, leg. Dr J. Balogh. 

Pareutetrapha nigrimaculata Breuning, 1952: 138. 

?PT: [Tonkin occ., Rég. de Hoa Binh]. 
Pareutetrapha olivacea Breuning, 1952: 138. 

HT: 9, China, Lou-tse-kiang [1901]; ?PT: [Tali-fr, Yunnan, China]. 

Parischnolea chiliensis Breuning, 1980b: 69. 

PT: Chili, El Nague, Coquimbo, 10.X.1976, J. Solervicens Ree. 

Phantasis nodulosa Sudre & Teocchi, 2000: 25. 
PT: Abyss., Harrar, coll. Plason / $ . 

Phelipara assamana Breuning, 1967a: 31. 

HT: Inde, Manipur, 15.IV. 1959, leg. Schmid [Assam, Chingsao, 3800-5400ft, 
15.VI.60]. 

Philotoceraeus descarpentriesi Breuning, 1975: 265. 

2 PT: Madagascar Centre, Adringitra Centre, plateau Andohariana 
[Soaindrana], 2000-2100m, HFAM 1, 9.XI-10.XII.1970. 

Phlyarus rufoscapus Breuning, 1976b: 740. 

HT: Tonkin, Mts Mauson, 700- 1000m, IV- V, leg. H. Fruhstorfer. 

Phryneta margaritifera Westwood, 1848: 11. 

ST: Nepaul, coll. A. Melly [Margaritifera Melly, Westd Or. Enty 5 th plate]. 

Phytoecia achillena Holzschuh, 1971: 68. 

PT: Anatolien, Amanusgebirge, Prov. Adana, Nurdagi gecidi, 1000m- 1200m, 

13-27.V1970, leg. C. Holzschuh [800-1300m]. 
Phytoecia adusta Reitter, 1889: 43. 

?PT: [Arménien]. 
Phytoecia akbesiana Pic, 1900: 20. 

?PT: [Syrie]; ?PT: [ill., Kiril Dagh]. 
Phytoecia albosuturalis Breuning, 1947c (1946): 143. 

HT: Shi-wan-tsze, Hiu-man-Kiang-keou [Hiu-mou-kiang-keou]. 
Phytoecia anteatra Breuning, 1966b: 53. 

HT: Angola, Loanda [Quela, Loanda, Fagenda Luhanga, Prinz Carolath S.G.]; 

PT: idem. 
Phytoecia apicefuscipennis Breuning, 1967b: 40. 

HT: 9, Cameroun, Kribi [Cameroun, Batanga]. 
Phytoecia argenteosuturalis Breuning, 1955e: 356. 

PT: Upper Sources, Olifants River, Ceres C. P. 
Phytoecia atricollis Breuning, 1960b: 23. 

HT: $ , Tanganyika, Lindi [D. O. Afr.]. 



178 J- MORATI 

Phytoecia atripennis Breuning, 1951: 450. 

HT: 9 , Congo belge, Beni. 
Phytoecia atrohumeralis Breuning, 1964b: 7. 

HT: 6 , Congo, Elisabethville. 
Phytoecia basilewskyi Breuning, 1950c: 46. 

?PT: [Tchad, Haut Chari, Fort Sibut, Collection Le Moult]; ?PT: [idem mais 

Congo Français]. 
Phytoecia bucharica Breuning, 1943: 101. 

HT: S, Boukhara, Tchitchantan [Ost Buchara, Tschitschantan, Nufswald, F. 

Hauser, 1898]; ?PT: [Ost Buchara, Tschitschantan, Nufswald, F. Hauser, 1898]. 
Phytoecia chinensis Breuning, 1943: 102. 

HT: 6 , Chine, province Pétchili. 
Phytoecia ciliciae Breuning, 1951: 406. 

HT: â , Ciliciae Adana; PT: idem. 
Phytoecia coeruleipennis Breuning, 1947a (1946): 22. 

HT: S , Cachemire, Kulu; PT: 9 , idem. 
Phytoecia coeruleomicans Breuning, 1947a (1946): 22. 

HT: 6 , Japon, île Hondo, Chiuzenji. 
Phytoecia coerulescens cretensis Breuning, 1947e: 60. 

HT: île Crète [Kreta, Hag. Dheka, Holtz / 5. Ill]; ?PT: [Kreta, Iraklion, IV.53]. 
Phytoecia comes szetschuanica Breuning, 1967b: 40. 

HT: S , Chine, Prov. de Szetschouan, Tatsienlu. 
Phytoecia fortic omis Breuning, 1947a (1946): 22. 

HT: 6 , Inde Britannique. Cachemire, Kulu; PT: idem; ?PT: [Kashmir, Kulu]. 
Phytoecia fuscolateralis Breuning, 1977b: 297. 

?PT: [Programma litorale 1, 10.XI.71, Trans. A, Ora 18.00, Zona 5, Direz N.]. 
Phytoecia grisea Breuning, 1947a (1946): 21. 

HT: 9, Inde Britannique, Spiti [Manikar]; PT: S, idem [Pulga]. 
Phytoecia hispanica {hispancia orthographe originale incorrecte) Breuning, 1951: 

364. 

HT: 9 , Espagne, province de Temei, Montes Universales, V. 1949. 
Phytoecia indica Breuning, 1951: 404. 

HT: 9 , Inde. Coimbatore. 
Phytoecia infranigra Breuning, 1951: 429. 

HT: 6, Kenya, Ikutha. 
Phytoecia kashmirica Breuning. 1943: 101. 

HT: S , Cachemire, Koulou [Manali Kulu]; ?PT: [Indes angl., Spiti, Manikarn]. 
Phytoecia kukunorensis Breuning, 1943: 102. 

HT: o\ Thibet, Koukounor, 3200m [1898, F. Hauser]. 
Phytoecia lahoulensis Breuning. 1951: 359. 

HT: ó\ Kashmir, Lahoul [Kelling / G. Babault, VII. 1914]; PT: idem [Gandla]. 
Phytoecia lateralis medionigriventris Breuning, 1961a: 306. 

2 ?PT: [Ouganda. Entebbe]. 
Phytoecia latesuturalis Breuning, 1951: 430. 

?PT: [Tang., Kilimandjaro). 



CATALOGUE DES TYPES DE LAMIINAE 179 

Phytoecia moreana Breuning, 1943: 102. 

HT: 6, Morée, Nauplie [Argos, 1935, Maran et Step., Coll. Barton]; ?PT: 

[Nauplia / Pelopon., Argos, 1935, Maran et Step., Coll. Barton]. 
Phytoecia neavei guineana Breuning, 1951: 428. 

?PT: [Togo, Bismarkburg]; ?PT: [W. Africa, Abutshi, R. Niger]. 
Phytoecia nigerrima Breuning, 1947a (1946): 21. 

HT: 9 , Japon, île Hondo, Chiuzenji. 
Phytoecia nigroapicalis Breuning, 1944: 16. 

HT: 6, Irak, Bagdad; ?PT: [Mesopotania, Mosul, V.09, Coll. Hauser]. 
Phytoecia nigrofemorata Breuning, 1947a (1946): 23. 

HT: 6 , Inde Britannique, Cachemire, Kulu. 
Phytoecia nigrohumeralis Breuning, 1950c: 46. 

?PT: [Belg. Kongo, Beni]. 
Phytoecia nigrovittata Breuning, 1951: 431. 

?PT: [Ubanghi]. 
Phytoecia orientis medionigripennis Breuning, 1961a: 306. 

2 ?PT: [ill., Ouganda, M.Rn □ □]. 
Phytoecia ornata Breuning, 1951: 438. 

HT: â , Angola, Bailundo. 
Phytoecia parvula Breuning, 1960b: 23. 

HT: Algérie, Teniet el haad [Schaumm]. 
Phytoecia pilipennis Reitter, 1895: 161. (note manuscrite de Hùdepohl: «ne peut être 

un PT»). 

?ST: Anatolie Kubischtek, Ordubad, Araxesthal [Armenien, Ordubad]. 
Phytoecia pseudocallosa Breuning, 1962b: 15. 

HT: 9, Angola. 
Phytoecia pseudofervida Breuning, 1951: 419. 

?PT: [Durban, X.1896, J.P Gregor]. 
Phytoecia pseudolateralis Breuning, 1954c: 82. 

?PT: [Congo, Lusinga]. 
Phytoecia pseudolatesuturalis Breuning, 1961c: 44. 

HT: S, Uganda. 
Phytoecia pseudoneavei Breuning, 1954c: 80. 

?PT: [Congo, Lusinga]. 
Phytoecia pseudorientis Breuning, 1962b: 15. 

HT: Uganda, Mt Ruwenzori. 
Phytoecia pseudosomereni Breuning, 1964a: 102. 

HT: 9 , Cameroun, Dsang. 
Phytoecia punctipennis Breuning, 1947c (1946): 143. 

HT: o\ Chine [Tchenkiang, 2 I.V. 18]. 
Phytoecia ruf a allardi Breuning, 1974d: 777. 

4 PT: Uganda, Entebbe, VII. 1973. 
Phytoecia rufovittipennis Breuning, 1972c (1971): 419. 

HT: o\ Himalaya, Kumaon, Pauri Garhwal, Gamsali, 11. VIL 1958, 3350m 

[F. Schmid]. 



180 J- MORATI 

Phytoecia rufulescens Breuning, 1951: 421. 

?PT: [Natal, Malv □ □ □, 1.XI.07]; ?PT: [Natal]. 
Phytoecia somereni Breuning, 1951: 449. 

2 ?PT: [Kenya, Ikatha]. 
Phytoecia stenostoloides Breuning, 1943: 101. 

HT: 9 , Transbaïkalie, Verchné-Oudinsk. 
Phytoecia subcoeruleata Breuning, 1951: 410. 

HT: 5, Togo, Bismarckburg; ?PT: [Togoland, 1892-1893, L. Conradt]. 
Phytoecia subdorsata Breuning, 196 Id: 256. 

?PT: [Congo Beige, Luluabourg]. 
Phytoecia suturevittata Breuning, 1951: 429. 

?PT: [E. Africa, Mombas]. 
Phytoecia suvorovi (suvorowi orthographe originale incorrecte) Pic, 1905: 38. 

2 ST: Caucase [Kaukasus, Olty, E. Koenig / Phytoecia suworowici Kòg / Phyt. 

suvorovi Pic, det.]. 
Phytoecia tenuilinea mateui Breuning, 1951: 365. 

2 PT: Rio de Oro, Mateu. 
Phytoecia vulneris paganettii Breuning, 1944: 16. 

HT: S, Italie méridionale, Apulie, San Basilio, Paganetti; ?PT: [Murgien, 

Grottaglie / Paganetti]. 

Planodema albosternalis Breuning, 1950b: 7 (1956e: 720). 
HT: 6, Congo belge, Elisabethville [XL 19 11]. 

Prosoplus costatus Hiidepohl, 1996: 4. 

HT: $, Ost-Malaysia, Bornéo, Sabah, Mt Kinabalu, 3300m, Panar Laban, 

IV-V.87, Burckhardt, Lobi leg. [4.V.1987]. 
Prosoplus laterialbus Breuning, 197 la: 2. 

2 ?PT: [Nouvelle Guinée, Tifalmin, 1.69]. 
Prosoplus major Breuning, 1964a: 96. 

HT: Celebes, Makassar, leg. A. Naville [602 Mr, 25]. 
Prosoplus rosselli Breuning, 1982: 16. 

PT: Papouasie, archipel de la Louisiade, île Rössel, Meek coll., I-III.1898. 

Prosopocera ferrierei Breuning, 1950b: 5. 

HT: o\ Congo belge, Elisabethville [X.1911]. 
Prosopocera forchhammeri Breuning, 1986: 46. 

?PT: [Botswana, Serowe, Swaneng Hill, 12.11.77, light, B 71]. 
Prosopocera forchhammeriana Breuning. 1986: 47. 

PT: Botswana. Serowe, Swaneng Hill, 23.XI.77, light, Harold von Zyl, B. 812. 
Prosopocera haemorrhoidaloides Breuning. 1978d: 94. 

PT: Ghana, Tafo, V-VIII.1966, leg. E.O. Boafo; 8 ?PT: [idem mais H. 1968, E.O. 

Boafo]. 
Prosopocera infragrisea Breuning, 1950b: 5. 

HT: S , Transvaal, Shilouvane, leg. H. Junod. 



CATALOGUE DES TYPES DE LAMIINAE 181 

Prosopocera mussardi Lepesme & Breuning, 1956: 660. 

HT: Côte d'Ivoire, Adzopé, III. 1949, ree. R. Mussard, coll. Lepesme. 
Prosopocera parapropinqua Breuning, 1986: 47. 

PT: Botswana, Serowe, Swaneng Hill, 3 .XII. 1977 (MVL), P. F. leg. [UV, B. 

1085]. 
Prosopocera rhodesica forchhammeri Breuning, 1986: 48. 

PT: Botswana, Serowe, Swaneng Hill, 20.XI.79, UV, P F. leg [Forchhammer 

leg., n° 5110]. 

Pseudepilysta loebli Hùdepohl, 1996: 3. 

HT: 2, Ost-Malaysia, Sabah, Mt. Kinabalu, 1550m, 27.IV.1987, Burckardt - 
Lobi leg. [23.IV. 1987]. 

Pseudopharsalia flavostictica Breuning, 1969b: 194. 

PT: Bornéo, Pontianak; ?PT: [Bornéo, Pontianak]. 

Pseudozorilispe celebensis Breuning, 1976a: 208. 

PT: Sud Celebes, Bua Kraeng, 1700m, 11.1896, H. Frùhstorfer. 

Pterolophia bituberculatoides Breuning, 1976b: 738. 

HT: Malaisie, Singapour. 
Pterolophia egumensis Breuning, 1973b: 659. 

PT: île Egum, Yanarba, XI. 1895, leg. Meek. 
Pterolophia flavicollis Breuning, 1977a: 280. 

?PT: [village N Fida, IV- 3.VII.70 / test cacao]. 
Pterolophia fusca Breuning, 1969a: 35. 

HT: Ceylan, Peradenya. 
Pterolophia fuscomarmorata Breuning, 1977a: 280. 

?PT: [Cameroun, Nkolbisson, 6.X.71]. 
Pterolophia inplagiata Breuning, 1980e: 166. 

?PT: [Insel Mindanao]. 
Pterolophia lineatipennis Breuning, 1974b: 371. 

HT: Formose, Puli, Taitung, V- VI. 1971, leg. K.H. Chen; PT: idem. 
Pterolophia ochreosignata Breuning, 1974a: 73. 

HT: Philippines, Ile Sibuyan, San Fernando, W. Parson, VII. 1928; 2 PT: idem. 
Pterolophia paraforticomis Breuning, 1976b: 738. 

HT: Malaisie, Fräsers Hill, 17-30.XI.1974, leg. P. Pfanner. 
Pterolophia paramulticarinata Breuning, 1977a: 280. 

?PT: [NLoubon, 2.VII.71]. 
Pterolophia pf anneri Breuning, 1976b: 738. 

HT: Malaisie, Fräsers Hill, 17-30.XI.1974, leg. P Pfanner; PT: idem. 

?HT: [Malaisie, Cameron Highland, V.75, P. Pfanner]. 
Pterolophia pluricarinipennis Breuning, 1969a: 34. 

HT: o\ Inde, Perumalinaloi, 21. IV 1927, voyage Carl et Escher [Inde méri- 
dionale, Palnis]. 



182 J- MORATI 

Pterolophia postscutellaris Breuning, 1967a: 30. 

HT: N.E. Assam, 4.VIII.1961, leg. Schmid [Kameng, Nakhu, 3800-4800ft, 

4.VII.61]. 
Pterolophia rondoniana Breuning, 1962e: 54. 

PT: Vientiane, K17, Tangone, 19.X.62; PT: Vientiane, K17, Palesane, 17.X.62. 
Pterolophia schmidi Breuning, 1967a: 30. 

HT: Inde, Manipur, 9. VI. 1960, leg. Schmid [Assam, Sirahi Kashana, 6000- 

7500ft]. 
Pterolophia tenebrie oide s Breuning, 1976b: 739. 

HT: Malaisie, Singapour. 
Pterolophia woodlarkiana Breuning, 1973b: 659. 

?PT: [Woodlark, Meek. 95]. 

Ramularius albosticticus Breuning, 1964a: 97. 

HT: â , Lourenco Marques, Makulane, leg. Dr Audeoud; 2 PT: idem. 

Rhodopina assamana Breuning, 1966c: 15. 

HT: 9, Assam, North-East Frontier Agency, VI. 1961, coll. Mussard [Kameng, 

Nafra, 24.VI.61, 3500-4000ft, Schmid]. 
Rhodopina manipurensis Breuning, 1972c (1971): 416. 

HT: et, Assam, Manipur, Hkayam Boum, 20.VI.1960, 2300-2600m [Inde, 

Schmid]. 

Rhytiphora corrhenoides Breuning, 1970a: 374. 

PT: Australie, Dawson Distr., coll. Barnard [Queensland, Monto, 10.1.1975 / 
8546]. 

Ropica assamensis Breuning, 1972c (1971): 415. 

HT: Assam, Kameng, Shergaon, 9.V1961, 1850-2000m [Himalaya, 9.V.61, 

Schmid]. 
Ropica ghanaensis Breuning, 1978d: 95. 

2 ?PT: [Ghana, Tafo, 11.1968, E.O. Boafo]; ?PT: [idem, mais 9.X.1967]. 
Ropica postmaculata Breuning, 1978d: 95. 

?PT: [Ghana, Tafo, 11.1968, E.O. Boafo]. 

R ucentra smetanai Hiidepohl, 1992: 179. 

4 PT: Bornéo, Sabah, Mt Kinabalu N.P., east base St John's Pk, 4000m, 08.VI- 
11.1988. Smetana; 2 PT: idem, mais 3900m. 

Saperda gleneoides Breuning, 1950b: 14. 

2 ?PT: [Tonkin, Montes Mauson, IV- V, 2-3000', H. Fruhstorfer]. 
Saperda mandschukuoensis Breuning, 1943: 104. 

HT: 6, Kharbine; ?PT: [Mandchuk □ □ □, Charbin]. 
Saperda scalaris algeriensis Breuning, 1952: 176. (Clerm. i. 1.). 

HT: 9, Algerien, Yakouren [VI.29, Ou □ □ □]; ?PT: [Kabylie, Yakouren, 

VI. 1902. Dr A. Chobaut]. 



CATALOGUE DES TYPES DE LAMIINAE 183 

Saperda subscalaris Breuning, 1952: 179. 

HT: 9 , China, Provinz Yunnan, Fluss Soling-ho. 

Schoenionta ichneumonoides Breuning, 1954a: 537. 

?PT: [Cochinchine, Cap St Jacques, Baria]. 
Schoenionta javanicola Breuning, 1954a: 538. 

HT: 9 , Java, Soekaboemi, Coll. Le Moult. 
Schoenionta merangensis Breuning, 1954a: 539. 

?PT: [Sumatra, Medan]; ?PT: [Sumatra, Merang]. 
Schoenionta philippinica Breuning, 1954a: 541. 

HT: S , Philippinen, Insel Mindanao, Zamboanga, leg. Baker. 

Serixia assamana Breuning, 1967a: 34. 

HT: Assam, Cachar District, 9.V.1960, leg. Schmid [Bandarkhal, 500ft]. 

Serixiomenesia flavosignata Breuning, 1959a: 12. 
?PT: [Malakka, Kouala Lumpur]. 

Serixiophytoecia vitticollis Breuning, 1950e: 380. 

?PT: [Kuching, Jan. 04, J.H.]; ?PT: [Kuching, 16.IX.1899]. 

Setoparmena mussardi Breuning, 1971b: 305. 

HT: Ceylan Central, au-dessus de Talatuoya, 27.1.1970, entre 850 et 1000m, 
tamisage de forêt [Mussard, Besuchet et Lobi]; PT: idem. 

Similosodus burckhardtì Hüdepohl, 1996: 4. 

HT: ó\ Ost-Malaysia, Bornéo, Sabah, Sepilok, IV-V.82, leg. Burckhardt. 

Sophronica attardi Breuning, 1974d: 777. 

?PT: [Ht Zaïre, Kisangani]. 
Sophronica ceylanica Breuning, 1971b: 306. 

HT: Ceylan, North central, Medawachchiya, 6. II. 1970, à la lumière du 

Resthouse [Mussard, Besuchet et Lobi]. 
Sophronica junodi Breuning, 1950b: 12. 

HT: Transvaal, Shilouvane, leg. Junod. 
Sophronica vittipennis Breuning, 1986: 55. 

PT: Botswana, Serowe, Swaneng Hill, 7.X.77, U.V. 

Spinogoephanes pauliani Breuning, 1975: 263. 

PT: Madagascar Centre, Adringitra Centre, plateau Andohariana (Soaindrana), 
2000-2 100m, HFAM 1, 9.XI-10.XII.1970. 

Spinosomatidia similis Breuning, 1981a: 46. 

PT: Cape Province, King Williamstown, XII. 1961, im Humus, N. Leleup. 

Stenellipsis albovittata Breuning, 1978a: 26. 

?PT: [New Caledonia, Nouméa, Mt. Koghi / 12-13.11.1977, leg. Dr J. Balogh]. 



184 J- MORATI 

Stenellipsis assimilis Breuning, 1978a: 29. 

?PT: [New Caledonia, I. Lifou, Ejengen, 21.11.1977, leg. Dr J. Balogh]. 
Stenellipsis basipustulata Breuning, 1978a: 27. 

?PT: [New Caledonia, Mt Mandjelia, 17.11.1977, leg. Dr J. Balogh]. 
Stenellipsis fuscolateralis Breuning, 1978a: 28. 

?PT: [New Caledonia, I. des Pins, 23-27.11.1977, leg. Dr J. Balogh]; ?PT: [New 

Caledonie, I. Lifou, Chépénéhé, 20.11.1977, Dr J. Balogh]. 
Stenellipsis kaszabi Breuning, 1978a: 28. 

?PT: [New Caledonia, Boursil, Col. D. Roussettes / 26-27.1.1977, leg. Dr J. 

Balogh]. 
Stenellipsis obscurithorax Breuning, 1978b: 69. 

?PT: [New Caledonia, Mt Panie, 8.X.1977, leg. Dr J. Balogh]. 
Stenellipsis postmaculata Breuning, 1978a: 28. 

?PT: [New Caledonia, I. des Pins, 23-27.11.1977, leg. Dr J. Balogh]. 
Stenellipsis subunicolor Breuning, 1978a: 27. 

PT: New Caledonia, I. Lifou, Bjengen, 21.11.1977, leg. Dr J. Balogh. 

Stenostola brunnescens Breuning, 1947c (1946): 144. 

HT: S , Japon, île Hondo, Gifu. 
Stenostola callosicollis Breuning, 1943: 100. 

HT: Mandchourie, Handaohetzy [VI. 193 8]; ?PT: [idem]. 
Stenostola trivittata Breuning, 1947c (1946): 143. 

HT: 9 , île de Hondo, Chiuzenji; ?PT: [Yunohana, Tateiwa V., Minami-Aizu, 

14.VI.1949, Y.K. / North of Nikko, Honshi]. 
Stenostola univittata Breuning, 1947c (1946): 144. 

HT: â, Hondo, Gifu; ?PT: [idem]. 

Stibara nigrovittata Breuning, 1954a: 467. 
2 ?PT: [Burma]. 

Sybra frasersi Breuning, 1976b: 737. 

HT: Malaisie, Fräsers Hill, 17-30.XI.1974, leg. P. Pfanner. 
Sybra pfanneri Breuning, 1976b: 737. 

HT: Malaisie, Fräsers Hill, 17-30.XI.1974, leg. P. Pfanner. 

?HT: [Malaisie, Cameron Highlands, V.1975, P. Pfanner]. 
Sybra postbasicristata Breuning, 1974b: 371. 

HT: Inde. Kerala, Cardamon Hills, entre Pambanar et Peermade, 950m, 

5.XI.1972, leg. Besuchet, Lobi, Mussard. 

Sybrinus albomarmoratus Breuning, 1978d: 107. 

2 ?PT: [Ghana, Tafo, IX-X.1967, E.O. Boafo]; ?PT: [idem mais 11.1968]. 

Sybrinus flavicans Breuning, 1978d: 107. 

PT: Ghana, Tafo, E.O. Boafo, IV. 1968; 2 ?PT: [idem]; ?PT: [idem mais IX- 
X.1967]; 8 ?PT: [idem mais 11.1968]; ?PT: [idem mais VI. 1968]; ?PT: [idem 
maisX.1968]. 



CATALOGUE DES TYPES DE LAMIINAE 185 

Tetraulax junodi Breuning, 1950b: 6. 

HT: $ , Transvaal, Shilouvane, leg. H. Junod; PT: idem. 

Thyestilla coerulea Breuning, 1943: 100. 

HT: ? , Chine, province Sétchouen, Tatsienlou [Stötzner]. 

Tmesisternus fergussoni Breuning, 1970b: 478. 

?PT: [He Fergusson, IX-XII.1924]. 
Tmesisternus rossetti Breuning, 1982: 20. 

PT: Rossell I., I-III.98, Meek. 

Trichauxa albovittata descarpentriesi Breuning, 1975: 262. 

HT: Madagascar Centre, Adringitra Est, Anjavidilava, 1850- 1950m, FDHM 2, 
18.XIL1970-15.I.1971. 

Trichohyllisia mussardi Breuning, 1957a: 835. 

HT: Congo Belge, Uélé, 1.1952, coll. Mussard [Bili, Verbeke]. 

Trichopterolophia schurmanni Breuning, 1980c: 49. 

PT: â , Ile de Bali, Bedugul, XII. 1979, leg. Schurmann, coll. Schurmann; PT: 
idem mais 1.80. 

Tricondyloides parinermis Breuning, 1978a: 25. 

PT: Nile Calédonie, Mt Rembai, 19-21.1.1977, leg. Dr J. Balogh. 

Tuberculetaxalus lumawigi Breuning, 1980e: 163. 
?PT: [Insel Mindanao]. 

Zosne matangensis Breuning, 1950e: 378. 

?PT: [Bornéo, Matang]; ?PT: [Matang, 3600ft] . 



LISTE 2: NOMS DU NIVEAU INFRA-SUBSPÉCIFIQUE DISPONIBLES 

Nitocris nigrocincta Aurivillius var. opaca Aurivillius, 1907: 38. 
2 ST (CT): Deutsch Ost- Africa, Kigonsera [1906]. 

Oberea consentanea Pascoe var. unicolor Breuning, 1956f: 236. 

?PT: [Fu Kien, Fochow]; ?PT: [Tonkin, Hoa-Binh]. 
Oberea euphorbiae Germar var. intermedia Breuning, 1947e: 59. 

HT: 9 , Hongrie, Uzghorod [Art. Vavra]; ?PT: [So Sur Sur, Fritsch]. 
Oberea formosana Pic var. ruficornis Breuning, 1956f: 235. 

?PT: [Chekiang, Tienmuschan]. 

Phytoecia gaubili Mulsant var. separata Pic, 1895: 70. 
?PT: [Teniet,Alg.]. 



186 J- MORATI 

LISTE 3: NOMS DU NIVEAU INFRA-SUBSPÉCIFIQUE NON DISPONIBLES 

Agelasta mindanaonis Breuning m. albescens Breuning, 1980e: 158. 

2 ?PT: [Insel Mindanao]. 
Agelasta mindanaonis Breuning m. nigerrima Breuning, 1980e: 158. 

?PT: [Insel Mindanao]. 

Blepisanis coerulea Aurivillius m. viridipennis Breuning, 1950b: 20. 

HT: 9, Congo belge, Elisabethville [XI. 19 11]; ?PT: [Kenya, Ikutha]; ?PT: 

[Vihl. Nyansa]. 
Blepisanis exilis Pascoe m. ochreovittata Breuning, 1950b: 21. 

HT: 9, Transvaal, Makanete, leg. H. Junod [Ricatla]; ?PT: [Transval, 

Lichtenburg, XI.1903]; ?PT: [Salisbury, O'Neil, 13.11.1914]. 
Blepisanis flavovittata Breuning m. albovittata Breuning, 1950c: 24. 

?PT: [Congo, Elisabethville]. 
Blepisanis insignis Aurivillius m. lateriflava Breuning, 1950c: 20. 

2 ?PT: [Belg. Kongo, Elisabethville]. 
Blepisanis neavei Aurivillius m. aurosternalis Breuning, 1950c: 20. 

?PT: [Congo, Elisabethville]; ?PT: [E'ville, XI.1911]. 
Blepisanis neavei Aurivillius m. rufosternalis Breuning, 1950b: 20. 

2 ?PT: [Kenya, Ikutha]. 

Cleptometopus mniszechi Lacordaire m. rufovittatus Breuning, 1966a: 109. 
HT: Nord-Ost-Assam, 3. VII, leg. Schmidt, coll. Mussard [Nakhu]. 

Chlorisanis viridis Pascoe m. violaceosuturalis Breuning, 1954a: 454. 
?PT: [Tenasserim, Tandong, 4000', V., Fruhstorfer leg.]. 

Conizonia compacta Ménétriés m. discovittata Breuning, 1947e: 59. 

HT: Arménie, Ordubad; ?PT: [Armenie, Ordubad]. 
Conizonia warnieri Lucas m. brunnea Breuning, 1954a: 495. 

HT: 9, Oran, Magenta [M. Rotrou, 1915]; ?PT: [Oran, Magenta]. 
Conizonia warnieri Lucas m. ruficornis Breuning, 1954a: 495. 

HT: S , Oran, Magenta. 

Dirphya nigricornis Olivier m. pseudonigricornis Breuning, 1950b: 23. 
HT: 9 , Transvaal, Shilouvane, leg. H. Junod. 

Dorcadion emgei Ganglbauer m. multivittatum Breuning, 1947b (1946): 104. 

HT: 9 , Grèce, Kyllène. 
Dorcadion tibiale Jakovlev m. accessorium Breuning, 1947d (1946): 166. 

HT: S . Turkestan. Vallée du Naryn. 
Dorcadion subsericatum Pic m. junctum Breuning, 1962c: 370. 

2 PT: Anatolie. Kastamuni, leg. Breuning [27.IV59]. 
Dorcadion veluchense Pic m. peristericum Breuning, 1947b (1946): 106. 

?PT: [Peristeri, Pindos]; ?PT: [Peristeri]. 



CATALOGUE DES TYPES DE LAMIINAE 187 

Entelopes glauca Guérin m. nigroreducta Breuning, 1954a: 477. 
?PT: [Borneo, Setipas]; ?PT: [id, mais Bornéo occ.]. 

Glenea diverselineata Pic m. yunnana Breuning, 1976b: 740. 

HT: Chine, Prov. Yunnan, Tche-Ping-Tcheou. 
Glenea luctuosa Pascoe m. rufoabdominalis Breuning, 1956b: 183. 

?HT: Bougainville, BMNH [Bougainv.]. 

Gnathoenia congoana Belon m. immaculipennis Breuning, 1966c: 16. 

HT: 9, Congo belge, Befori, coll. Mussard [équateur, XII.51]. 
Gnathoenia congoana Belon m. reticulata Breuning, 1937: 238. 

?PT: [Congo, Stanleyville]. 

Hyllisia leucosuturata Hunt & Breuning m. multialbovittata Breuning, 1981a: 44. 
?PT: [Transval, Kruger Park]. 

Linda annulicornis Matsushita m. ruficeps Breuning, 1954a: 553. 

HT: 2, China, Provinz Yunnan, Yunnanfu [Kouy-tchiou]. 
Linda apicalis Pic m. yunnana Breuning, 1976b: 741. 

HT: Chine, Prov. Yunnan, Talifu. 
Linda f emorata Chevrolat m. rufifrons Breuning, 1954a: 549. 

HT: $ , China, Provinz Kweichow, Nanning. 
Linda gracilicornis Pic m. rufofemorata Breuning, 1954a: 550. 

HT: S , China, Provinz Szetschuan, Tatsienlu, ex coll. Reitter. 
Linda gracilicornis Pic m. tatsienlui Breuning, 1954a: 550. 

HT: 9 , China, Provinz Szetschuan, Tatsienlu, ex coll. Reitter; ?PT: [Tatsienlu, 

Szetschuan]. 
Linda nigroscutata Fairmaire m. conjuncta Breuning, 1954a: 558. 

2 ?PT: [Yunnan, Yunnan sea]. 
Linda nigroscutata Fairmaire m. longeplagiata Breuning, 1954a: 558. 

HT: 6 , China, Provinz Yunnan, Fluss Solingho, ex coli. Hauser; ?PT: [Kuatun, 

2500m, 27. 40n. Br., 117.408 L., J. Klapperich, 31.V.1938, Vukien]. 
Linda rubescens Hope m. unicolor Breuning, 1954a: 556. 

HT: 6 , Sikkim, Gnatong. 
Linda vitalisi Vuillet m. nigroreducta Breuning, 1954a: 561. 

2 ?PT: [Tonkin, Chapa]. 

Loboberea pygidialis Gahan m.javaensis Breuning, 1954a: 533. 

?HT: Java, Preanger, ZMAN [W. Java, Sukabumi, 2000 ft, H. Fruhstorfer]; 

?PT: [Java occident., Sukabumi, 2000', 1893, H. Fruhstorfer]. 
Loboberea pygidialis Gahan m. nigroreducta (nigroteducta orthographe originale 

incorrecte) Breuning, 1954a: 533. 

HT: Ó , Sumatra, Palembang. 
Loboberea pygidialis Gahan m. trimaculata Breuning, 1950a: 200. 

?PT: [Java, Gounod gedeh]; ?PT: [Sumatra, Palembang]. 



188 J. MORATI 

Mecas laminata m. discopunctata Breuning, 1955c: 150. 

2 ?PT: [Mexico, Cordova]. 
Mecas laminata m. rufobasalis Breuning, 1955c: 150. 

?PT: [Mexico, Cordova]; ?PT: [Mexique]. 
Mecas laticeps m. mediopunctata Breuning, 1955c: 151. 

?PT: [N. Yucatan, Temax, Gaumer]. 
Mecas laticeps m. sutureflava Breuning, 1955c: 151. 

2 ?PT: [N. Yucatan, Temax, Gaumer]. 
Mecas rotundicollis m. mediomaculata Breuning, 1955c: 149. 

?PT: [Mexico, ill., Tehnucax □ □ □]. 
Mecas rubripes m. callosoreducta Breuning, 1955c: 143. 

?PT: [Mexico]. 

Menesia bimaculata Breuning m. latevitticollis Breuning, 1954a: 424. 

2 ?PT: [Borneo]. 
Menesia bimaculata Breuning m. prolongatevittata Breuning, 1954a: 424. 

?PT: [Borneo, Sandakan, Baker]. 
Menesia nigriceps Aurivillius m. inhumeralis Breuning, 1954a: 422. 

HT: 9, Borneo, Sandakan, leg. Baker; ?PT: [Malacca/ Staudinger]. 
Menesia sulphurata Gebier m. gifuensis Breuning, 1954a: 410. 

HT: 9, Japan, Insel Gifu, Hondo, Provinz Amagodani, 13.VIII.1947, leg. 

Kohbayashi. 

Menesida flavipennis Breuning m. basiflava Breuning, 1954a: 446. 

?PT: [Sumatra, Medan]. 
Menesida flavipennis Breuning m. mediofusca Breuning, 1954a: 446. 

2 ?PT: [Sumatra, Medan]. 

Nitocris schoutedeni Breuning m. verbekei Breuning, 1957b: 206. 

HT: ò\ Congo Belge, Uele, coll. Mussard [Bili, 1952, Verbeke]. 

Nupserha ugandensis Breuning m. nigrolateralis Breuning, 1978d: 112. 
?PT: [Uganda, Entebbe, JJ- VII. 1972, Exp. H. Falke]. 

Oberea angolensis Breuning var. bimaculaticeps Breuning, 1961b: 62. 

HT: 6 , Angola, Bailundo. 
Oberea angolensis Breuning var. nigritomaculata Breuning, 1961b: 62. 

HT: 9 , North West Rhodesia. 
Oberea annulicornis Pascoe var. craengiana Breuning, 1961b: 120. 

?HT: 9, Celebes, Bonthain, leg. Ribbe, MNHN [S.Celebes, Bua-Kraeng, 

5000', 11.1896, H. Fruhstorfer]; ?PT: [S.Celebes, Lompa-Battau, 3000ft, 111.96, 

H. Fruhstorfer]. 
Oberea annulicornis Pascoe var. umbrosior Breuning, 1961b: 120. 

?PT: [Celebes, Tamboegoe]. 
Oberea artocarpis Gardner m. trifusciventris Breuning, 1962a: 147. 

2 ?PT: [Inseln Andamanen]. 



CATALOGUE DES TYPES DE LAMIINAE 189 

Oberea atricilla Fairmaire m. apicenigrescens Breuning, 1950c: 41. 

?PT: [Kamerun, Mt Bronje-Farm]; ?PT: [Cameroons, Bitye-Ja River, 3000 feet]. 
Oberea atricilla Fairmaire var. asmarensis Breuning, 1961b: 82. 

HT: 6 , Erythrée, Asmara [VIII]. 
Oberea atricilla Fairmaire var. bimac ulive ntris Breuning, 1961b: 82. 

HT: 9 , Cameroun, Ja River [Bitye Ja River]. 
Oberea atricilla Fairmaire m. parteruficeps Breuning, 1950c: 41. 

?PT: [Congo français, N'Djolé, J. Arrighi]; ?PT: [Cameroons, Bitye Ja River, 

3000 ft]. 
Oberea atricilla Fairmaire m. rufopygidialis Breuning, 1950c: 41. 

?PT: [Cameroons, Bitye-Ja River, 3000 feet]; ?PT: [Kenya, Ikutha]. 
Oberea basalis Chevrolat m. flavicans Breuning, 1950c: 42. 

?PT: [Gabon]; ?PT: [Gabon, Bas Ozooné, Favarel]. 
Oberea bicoloricornis Pic var. rubroantennalis Breuning, 1962a: 186. 

HT: $ , Chine, province Szetschuan, Tatsienlu [Reitter]. 
Oberea cingulata Aurivillius m. parteflavosternalis Breuning, 1964a: 102. 

HT: Uganda [Afr. c, coll. Plason]. 
Oberea consentanea Pascoe m. mausoni Breuning, 1950d: 274. 

2 ?PT: [Tonkin, Montes Mauson, IV-V, 2000', H. Fruhstorfer]. 
Oberea curialis Pascoe m. nigroscapus Breuning, 1950b: 25. 

?PT: [Java Orient., Mt Ardjoeno]; ?PT: [L'Ile de Ceram, V.M. Duchon]. 
Oberea davaoensis Breuning var. antenigrocollis Breuning, 1961b: 139. 

PT: He Mindanao, Davao; ?PT: [idem]. 
Oberea depressa Gebier m. infradepressa Breuning, 1967b: 40. 

HT: 6, Mandchourie, He Askold [M. Jackowskie, 1880]; PT: idem [42,5° lat. 

N, 102° long., 1878]. 
Oberea depressa Gebier var. subdepressa Breuning, 1962a: 215. 

HT: $ , Chine, prov. Kiangi-si, Kiu-kiang. 
Oberea discoidalis Jordan var. maculisternalis Breuning, 1961b: 86. 

HT: Ô, Cameroun, Ja River [3000 feet]. 
Oberea erythrocephala Schrank m. macedonica Breuning, 1947e: 58. 

HT: 9, Macédoine, Usküb [2.6.68]; ?PT: [Bulgarien, Stanimaka, VII.33, 

Pfeiffer]. 
Oberea erythrostoma Heller var. antenigrescens Breuning, 1961b: 135. 

HT: 2, Ile Samar [VI-VII.96, J. Whithead]. 
Oberea erythrostoma Heller var. banahoana Breuning, 1961b: 135. 

HT: $ , Ile Luzon, Mt Banahao; PT: Insel Luzon, Laguna. 
Oberea erythrostoma Heller var. cebuensis Breuning, 1961b: 134. 

HT: S , Philippines, Ile Cebu; PT: idem. 
Oberea erythrostoma Heller var. medionigricollis Breuning, 1961b: 135. 

HT: S , Ile Luzon. 
Oberea erythrostoma Heller var. melanothorax Breuning, 1961b: 135. 

HT: 9 , Ile Luzon, Mt Isorong; PT: idem. 
Oberea erythrostoma Heller var. partenigripes Breuning, 1961b: 136. 

HT: S, Ile Luzon, Albay [Whithead, 94]. 



190 J- MORATI 

Oberea erythrostoma Heller var. pseudomakilingi Breuning, 1961b: 135. 

?PT: [Mindoro]; ?PT: [Philipp.]. 
Oberea erythrostoma Heller var. ruficuspis Breuning, 1961b: 135. 

HT: S , Ile Luzon, Laguna. 
Oberea erythrostoma Heller var. samarana Breuning, 1961b: 135. 

HT: 9 , Ile Samar [VI-VII.96, J. Whithead]. 
Oberea erythrostoma Heller var. samarensis Breuning, 1961b: 135. 

HT: 9, Ile Samar [VI-VII.96, J. Whithead]. 
Oberea formosana Pic var. spinipennis Breuning, 1962a: 163. 

?PT: [S.E. Borneo]. 
Oberea fuscicollis Breuning var. fuscolateralis Breuning, 1961b: 138. 

HT: cT, He Luzon, Cap Engano [Whitehead leg.]. 
Oberea fuscicollis Breuning var. infrarufescens Breuning, 1961b: 138. 

?PT: [Philippinen, Mindanao, Surigao]. 
Oberea fuscicollis Breuning var. mindanaonis Breuning, 1961b: 139. 

HT: ó\ Ile Mindanao; ?PT: [Luzon, mt B □ □ □]. 
Oberea griseopennis Schwarzer var. basimaculata Breuning, 1962a: 219. 

HT: ó\ Formosa, Sozan [30.IV 1933, Col. M. Chujo]; ?PT: [S. Formosa, 

Alikang, VI.09, Sauter S. V.]. 
Oberea herzi Ganglbauer var. longevittata (Pic i.l.) Breuning, 1962a: 205. 

HT: 9 , Corée, Pu Ryöng; ?PT: [Korea, Pu Ryong]. 
Oberea herzi Ganglbauer var. mediomaculata Breuning, 1962a: 205. 

HT: 6 , Mandchourie, Handaohstzy. 
Oberea herzi Ganglbauer var. rufipennis Breuning, 1962a: 205. 

HT: <?, Chine, Nanking. 
Oberea humeralis Gressitt var. fruhstorferi Breuning, 1962a: 180. 

HT: 9, Tonkin, Mts Mauson, 700- 1000m, IV- V, leg. Fruhstorfer. 
Oberea humilis Fairmaire var. togoensis Breuning, 1961b: 78. 

PT: Togo, Bismarckburg, leg. Conradt. 
Oberea humilis Fairmaire m. pseudohumilis Breuning, 1950c: 42. 

?PT: [Kamerun, Bitye River]. 
Oberea inclusa Pascoe var. partenigrescens Breuning, 1962a: 203. 

HT: S , Corée, Pu Ryong. 
Oberea insperans Pascoe m. celebesiana Breuning, 1950a: 199. 

?PT: [Celebes, Makassar]. 
Oberea japonica Thunberg m. infranigrescens Breuning, 1947e: 58. 

HT: 9 , Japon, Orni [Centn Japan, Orni, Zeze, VI.96, T. Lenz v., 12.IX.96]; ?PT: 

[Japon, Kioto, Villion]. 
Oberea lacana Pic m. mediofuscofemoralis Breuning, 1967b: 40. 

HT: 9 , Cambodge. 
Oberea lacana Pic var. orientalis Breuning, 1961b: 95. 

HT: â , Tonkin, Hoabinh; ?PT: [Tonkin occ, Rég. de Hoa Binh]. 
Oberea laetifica Pascoe var. toliensis Breuning, 1961b: 122. 

HT: S. Celebes. Toli-Toli; ?PT: [N. Celebes, Toli-Toli, Fruhstorfer]. 



CATALOGUE DES TYPES DE LAMIINAE 191 

Oberea lateapicalis Pic var. rufobasalis Breuning, 1962a: 151. 

HT: S , Inde, Anamalai Hills. 
Oberea lepesmei Breuning, var. haemorrhoidalis Breuning, 1961b: 91. 

?PT: [Togo, Bismarkburg]. 
Oberea longissima Aurivillius var. basirufa Breuning, 1961b: 91. 

?PT: [Cameroons, Bitye-Ja River]; ?PT: [Côte d'Ivoire, Adiopodoumé, V.32, 

J. Lepesme]. 
Oberea luluensis Breuning, var. atricollis Breuning, 1961b: 88. 

HT: â, Cameroun, Mukonje Farm; ?PT: [Belg. Congo, Yungumbi, Fraser]. 
Oberea lutea Thunberg var. nigroapicalis Breuning, 1962a: 154. 

?PT: [Maissour, Shiroga]; ?PT: [Darjoeling, Mus. Vaekenfurg]. 
Oberea macilenta Newman var. anterubripes Breuning, 1961b: 140. 

?PT: [Philippis]; ?PT: pas d'étiquette de localité. 
Oberea melanostoma Heller var. melas Breuning, 1961b: 136. 

HT: 9, Ile Luzon, Mt Isorog; ?PT: [Philipps.]. 
Oberea melanostoma Heller var. rufinithorax Breuning, 1961b: 136. 

?HT: [Pancan N □ □ □ ]; ?PT: [Ambulanga]. 
Oberea mimetica Heller var. boholensis Breuning, 1962a: 146. 

HT: 9 , Philippines, lie Bohol; ?PT: [Luzon / Mt Banahao, IV. 1914, Boettcher]. 
Oberea mimetica Heller var. pseudomimetica Breuning, 1962a: 145. 

HT: 9 , Philippines. 
Oberea mimetica Heller var. submimetica Breuning, 1962a: 145. 

?PT: [Luzon, Mt Banahao]. 
Oberea montivagans Fisher var. rubroscapus Breuning, 1962a: 158. 

HT: 9, Java, Pranger; ?PT: [Java]. 
Oberea neavei Aurivillius var. antennalis Breuning, 1961b: 75. 

?PT: [C. Beige, ill.]. 
Oberea nefasta Pascoe var. atroampliata Breuning, 1961b: 129. 

HT: 9 , Nouvelle Guinée, Geelwinck Bay, leg. Doherty. 
Oberea nefasta Pascoe var. flavoabdominalis Breuning, 1961b: 129. 

HT: â, Nouvelle Guinée, Andai, leg. Doherty; ?PT: [Fergusson I., IX-XII.94, 

A.S. Meek]. 
Oberea nefasta Pascoe var. rufuloides Breuning, 1961b: 129. 

HT: S, Ile Roon près Nouvelle Guinée; ?PT: [N. Guineen, Warao]. 
Oberea neptis Pascoe var. plurimaculiventris Breuning, 1962a: 161. 

?PT: [Sumatra's O.K., Brastagi, 1300m, 6.IV.1918, J.B. Corporaal]. 
Oberea nigerrima Breuning m. anterufofemoralis Breuning, 1964b: 6. 

HT: S , Java, Batavia. 
Oberea nigerrima Breuning var. discovittata Breuning, 1961b: 100. 

HT: S , Java; PT: Java, Soekaboemi. 
Oberea nigerrima Breuning var. rubricollis Breuning, 1961b: 100. 

HT: 9 , Java, Soekaboemi. 
Oberea nigrocincta Aurivillius m. camerunensis Breuning, 1964b: 6. 

HT: S , Cameroun, Kriby. 



192 J- MORATI 

Oberea nigriceps White m.flavipennis Breuning, 1950a: 199. 

?PT: [Formosa, Kosempo, Coll. Sauter]; ?PT: [China, Hong-Kong]. 
Oberea nigriceps White var. nigromaculicollis Breuning, 1962a: 183. 

HT: S , Chine, Ngan hwei. 
Oberea nigriceps White m. obscuripennis Breuning, 1967b: 40. 

HT: o*, Tonkin, Monts Mauson, V-VI, 700-1000m alt., leg. H. Fruhstorfer 

[IV- V, 2-3000']. 
Oberea nigripennis Breuning m. rufiniscapus Breuning, 1950c: 38. 

?PT: [Gabon, Bas Ozooné]. 
Oberea nigriventris Bates var. postrufofemoralis Breuning, 1961b: 94. 

HT: 6 , Chine, Prov. Fokien, Foochow. 
Oberea notata Pic var. rufoantennata Breuning, 1962a: 196. 

HT: 6 , Chine, prov. Chekiang, Kiukiang; ?PT: [Chekiang, Kiu Kiang]. 
Oberea nyassana Breuning m. kenyensis Breuning, 1955a: 76 ou Breuning, 1958a: 37. 

?PT: [Kenya, Ikutha]. 
Oberea nyassana Breuning var. laterinigra Breuning, 1961b: 74. 

?PT: [Mozambique, Zambesie]. 
Oberea occidentalis Lepesme & Breuning var. apicenigra Breuning, 1961b: 89. 

HT: 9 , Sierra Leone, Moyamba. 
Oberea pedemontana Chevrolat m. subtestacea Breuning, 1947e: 58. 

HT: S, Tyrol, Rovereto; ?PT: [Tirol, Rovereto]. 
Oberea pictipes Pascoe m. nigrof e morata Breuning, 1956g: 23. 

?HT: Java, ISNB [Java]; ?PT: [Java, Mile Badollet, 623-29]. 
Oberea pictipes Pascoe var. rubripes Breuning, 1961b: 108. 

HT: â , Bornéo. 
Oberea pictipes Pascoe var. tenasserimensis Breuning, 1961b: 108. 

2?PT: [Malacca, Perak]. 
Oberea posticata Gahan var. rubroantennata Breuning, 1962a: 165. 

HT: S , Assam, Suriga. 
Oberea praedita Pascoe var. antenigripes Breuning, 1961b: 116. 

?PT: [Borneo, Pontianak]. 
Oberea pseudopascoei Breuning m. propinqua Breuning, 1950c: 41. 

2 ?PT: [Gabon, Bas Ozooné]. 

Oberea pseudovaricornis Hunt & Breuning m. fuscoampliata Breuning, 1978d: 115. 

3 ?PT: [Prov. du Cap, Port St- John, 11.1963, leg. G. Heinrich]. 

Oberea pseudovaricornis Hunt et Breuning var. mossambicensis Breuning, 1961b: 73. 

HT: ô , Mossambique, Zambezi. 
Oberea puncticollis Breuning var. crassepuncticollis Breuning, 1962a: 146. 

HT: cT, He Luzon, prov. Tayabas, Infante; ?PT: [Philippinen, Luzon, Prov. 

Laguna]. 
Oberea pupillata Gyllenhal m. bimaculatoides Breuning, 1947e: 58. 

HT: <?, Moravie, Prostejov; ?PT: [Skuje, Vlach.]. 
Oberea pupillata Gyllenhal m. pseudodepressa Breuning, 1947e: 58. 

HT: 6 . Sibérie, Krasnojarsk; ?PT: [Sibir., Irtysch]. 



CATALOGUE DES TYPES DE LAMIINAE 193 

Oberea quianga Heller var. partefuscicollis Breuning, 1961b: 137. 

HT: ó\IleCebu. 
Oberea reimschi Breuning var. merangana Breuning, 1961b: 105. 

?HT: Sumatra, Merang, coll. Breuning [Sumatra, Medan]. 
Oberea rubetra Pascoe var. basinigricollis Breuning, 1961b: 113. 

HT: 6 , Bornéo. 
Oberea rubetra Pascoe var. mediorufinipes Breuning, 1961b: 113. 

?PT: [Sumatra, Medan]; ?PT: [Sumatra occident.]. 
Oberea rubetra Pascoe var. parterufipes Breuning, 1961b: 113. 

HT: 9, Bornéo, Sandakan, leg. Baker; ?PT: [Borneo, Sandakan, Baker]. 
Oberea ruficollis Fabricius var. rufolineata Breuning, 1962a: 227. 

HT: $ , Floride, Stanford [V et VI.95]. 
Oberea sansibarica Harold var. binigromaculata Breuning, 1961b: 63. 

HT: ó\ Kenya, Maguga [6.III.54]. 
Oberea schaumi Le Conte var. infrarufa Breuning, 1962a: 223. 

HT: ó\ Californie, Mariposa [VI.74]; ?PT: [U.S., California]. 
Oberea schaumi Le Conte var. vittipennis Breuning, 1962a: 224. 

2 ?PT: [U.S., Ohio]. 
Oberea scutellaroides Breuning m. rufithorax Breuning, 1947e: 58. 

HT: S , Chine, province Tchékiang, Tschenkiang [2.VI.18]; ?PT [China]. 
Oberea seminigra Chevrolat m. clareabdominalis Breuning, 1956d: 700. 

?PT: [Philipp., Semp □ □ □]; ?PT: [Philippines, Luzon, Bilucao, La Laguna]. 
Oberea sobrina Boisduval var. basifuscoabdominalis Breuning, 1961b: 127. 

?PT: [Insel Ceram]. 
Oberea subabdominalis Breuning var. momeitensis Breuning, 1962a: 167. 

HT: S , Birmanie, Momeit [Hte Birmanie, Etat de Momeit, 600m]. 
Oberea suturalis Jordan var. flaveola Breuning, 1961b: 79. 

?PT: [Kamerun, Mukonje Farm]; ?PT: [Ogowe]. 
Oberea taihokuensis Breuning var. flavosternalis Breuning, 1962a: 169. 

HT: S , Formose, Kosempo; ?PT : [ill., Kurau]. 
Oberea tricolor Aurivillius var. binaluanica Breuning, 1961b: 112. 

?PT: [Kinabalu]. 
Oberea tricolor Aurivillius var. discoclarior Breuning, 1961b: 112. 

HT: ó\ Ile de Bali, III-IV.1896, leg. Doherty. 
Oberea trigonifera Aurivillius m. mac ulive ntralis Breuning, 1958a: 37. 

?PT: [Kap Colony, East London]; ?PT: [ill., Peramiha, 1905]. 
Oberea tripunctata Swedenbord var. subdeficiens Breuning, 1962a: 229. 

HT: et, New York. 
Oberea tripunctata Swedenbord var. subexilis Breuning, 1962a: 228. 

HT: $, Canada, Montréal [Que., VI.]. 
Oberea tripunctata Swedenbord var. submandarina Breuning, 1962a: 229. 

HT: 9 , Missouri. 
Oberea unimaculicollis Breuning var. nigroampliata Breuning, 1962a: 194. 

HT: $ , Sumatra, Medan. 



194 J- MORATI 

Oberea varicornis Fâhraeus m. laterinigrescens Breuning, 1950c: 41. 

?PT: [Delazoa, Bai.]; ?PT: [Mozambique, Zambezi]. 
Oberea viperina Pascoe var. burmanensis Breuning, 1961b: 104. 

?PT: [Burma]. 
Oberea viperina Pascoe var. clariventris Breuning, 1961b: 103. 

HT: S , presqu'île de Malacca, Perak, Gounong-Boubou, leg. Doherty [1100m]. 
Oberea viperina Pascoe var. penangensis Breuning, 1961b: 104. 

PT: Malacca, Perak, Doherty. 
Oberea viperina Pascoe var. transitiva Breuning, 1961b: 103. 

HT: c? , Ile Engano. 
Oberea walkeri Gahan var. sikkimensis Breuning, 1961b: 185. 

HT: S , Sikkim, Darjeeling, leg. Andrewes. 
Oberea yunnanensis Breuning var. atripennis Breuning, 1961b: 98. 

HT: S , Chine, province Yunnan, Solingho. 

Obereopsis sericea Gahan m. subluteonigra Breuning, 1966c: 16. 

HT: S, Manipur, Poi, 1200m, 4. VII. 1960, coll. Mussard [Assam]. 
Obereopsis verticenigra Breuning f. flavo abdominalis Breuning, 1974b: 374 

iflvooabdominalis orthographe originale incorrecte). 

HT: Inde, Kerala, Trivandrum Dt., Poonmudi Range, 950m, V.1972, leg. T.R.S. 

Nathan. 

Ossonis clytomima Pascoe m. flavotibialis Breuning, 1954a: 430. 
HT: 6 , Borneo, Sandakan, leg. Baker. 

Paradystus notator Pascoe m. fuscoampliatus Breuning, 1954a: 459. 

HT: Soekaranda, 1.1894, Dohrn; ?PT: [Medan, Mjöb.]. 
Paradystus notator Pascoe m. sericeoprolongatus Breuning, 1954a: 459. 

?PT: [Borneo, Sandakan, Baker]; ?PT: [ill., Lawas, II-III.96, A. Everett]. 

Paraglenea fortunei Saunders m. clarevittata Breuning, 1952: 126. 

HT: 9 , China, Insel Chusan. 
Paraglenea fortunei Saunders m. conjunctefasciata Breuning, 1952: 126. 

HT: 6 , China, Provinz Szetschuan, Tatsienlu. 
Paraglenea fortunei Saunders m. unicoloripennis Breuning, 1952: 127. 

HT: 6\ China, Insel Chusan; ?PT: [idem]. 
Paraglenea fortunei Saunders m. viridicollis Breuning, 1952: 125. 

HT: S , China, Provinz Hupe, Ichang; ?PT: [idem]. 

Pardaloberea curviticeps Pic m. partenigrescens Breuning, 1954a: 534. 
?PT: [Tonkin, Hoa-Binh]. 

Phytoecia algerica Desbrochers des Loges m. nigroanalis Breuning, 1947e: 59. 

HT: Algérie; ?PT: [Oran]. 
Phytoecia analis Fabricius m. rufescens Breuning, 1951: 443. 

2 ?PT: [Côte d'Ivoire, Dimbokro]. 



CATALOGUE DES TYPES DE LAMIINAE 195 

Phytoecia aurivillii Breuning m. violaceipennis Breuning, 1951: 410. 

HT: 9, Angola, Bailundo; ?PT: [Kenya, Ikitha]. 
Phytoecia balcanica Frivaldszky m. subvitticollis Breuning, 1951: 92. 

HT: 8, Anatolie, Amasia, leg. Korb [1888]; ?PT: [Balkan]. 
Phytoecia bodemeyeri Reitter m. luristanica Breuning, 1947e: 59. 

HT: 8, Louristan [Persia]; ?PT: [Persia, Luristan]. 
Phytoecia bodemeyeri Reitter m. rufof emorata Breuning, 1947e: 59. 

HT: 2 , Louristan [Persien, v. Bodemeyer]. 
Phytoecia coerulescens Scopoli m. marthae Breuning, 1951: 100. 

HT: 9, Catalogne, San Feliù de Guixols, 18.V.1950; ?PT: [Colline Morières, 

28.V.1908]. 
Phytoecia collaris Pascoe m. bulbosa Breuning, 1951: 414. 

?PT: [P. Natal /M. Germ.]. 
Phytoecia comes Bates m. griseopubescens Breuning, 1951: 96. 

HT: 8 , Ile Hondo, Chiuzenji; ?PT: [Hondo, Chiuzenji]. 
Phytoecia comes Bates m. ohbayashii Heyrovsky, 1952: 13. 

?PT: [Oomor-Jap., coll. Dr Heyrovsky]. 
Phytoecia delagrangei Pic m. subgrisea Breuning, 1951: 354. 

HT: 8, Marasch [V.28, Taurus, leg. H. Kulzer]; ?PT: [Marasch, V.28, Taurus, 

leg. H. Kulzer]. 
Phytoecia erivanica Reitter m. rufipennis Breuning, 1947e: 59. 

HT: 8, Transcaucasie, Suchoj-Fontan [1910, H. Kulzer]; ?PT: [Suchoj Fontan, 

1910, H. Kulzer]. 
Phytoecia erythaca Pascoe m. nigritipennis Breuning, 1951: 412. 

?PT: [Zulu, Caffraria]. 
Phytoecia erythaca Pascoe m. parteruficeps Breuning, 1951: 412. 

HT: 8 , Zululand [Zulu, Caffraria]. 
Phytoecia exilis Pascoe m. vittulipennis Breuning, 1951: 422. 

HT: 2 , Transvaal. 
Phytoecia ferrugata Ganglbauer ab. houskai, Heyrovsky, 1948: 20. 

PT: Palestine, Jerusalem, Houska leg. 
Phytoecia fervida Pascoe m. nigriceps Breuning, 1951: 418. 

?PT: [Natal]. 
Phytoecia fervida Pascoe m. nigrodiscalis Breuning, 1951: 418. 

?PT: [ill., Maloen, 61 □ □ □ 99, JP Ce..]. 
Phytoecia fervida Pascoe m. rufinitibialis Breuning, 1951: 418. 

?PT: [Natal]. 
Phytoecia flavovittata Breuning m. latevittipennis Breuning, 1964b: 7. 

HT: 2 , Congo, Elisabethville. 
Phytoecia flavovittata Breuning m. latevitticollis Breuning, 1951: 424. 

2 ?PT: [Mozambique, Zambesi]. 
Phytoecia geniculata Kolbe m. flaviventris Breuning, 1950c: 44. 

?PT: [Belg. Kongo, Beni]. 
Phytoecia geniculata Kolbe m. fus cibasic omis Breuning, 1950b: 31. 

?PT: [Côte d'Ivoire, Dimbokro]; ?PT: [Niger, Ogruga]. 



196 J- MORATI 

Phytoecia geniculata Kolbe m. pusilla Breuning, 1950b: 31. 

?PT: [Côte d'Ivoire, Dimbokro]; ?PT : [Togo]. 
Phytoecia geniculata Kolbe m. subternigra Breuning, 1950c: 44. 

2 ?PT: [Côte d'Ivoire, Dimbokro]. 
Phytoecia haroldii Fâhraeus m. atricornis Breuning, 1951: 453. 

2 ?PT: [Kenya, Ikuta]. 
Phytoecia haroldii Fâhraeus m. bipunctulicollis Breuning, 1951: 454. 

HT: ó\ Tanganyica, Lindi [D. O. Afr.]. 
Phytoecia haroldii Fâhraeus m. holatripes Breuning, 1951: 453. 

HT: 9 , Tanganyica, Lindi [Teir]. 
Phytoecia haroldii Fâhraeus m. ornaticollis Breuning, 1950c: 46. 

?PT: [Kenya, Nairobi, 1660m]. 
Phytoecia haroldii Fâhraeus m. tripuncticollis Breuning, 1950c: 46. 

?PT: [Kenya, Ikuta]; ?PT: [D.O. Afrika, Kigonsera]. 
Phytoecia haroldii Fâhraeus m. unimaculata Breuning, 1951: 453. 

?PT: [N.W. Rhodesia, Lusemska, H.C. Dollman, XII. 1916]; ?PT: [Salisburg, 

ill, Mahon]. 
Phytoecia haroldii Fâhraeus m. verticepunctata Breuning, 1951: 453. 

?PT: [N.W. Rhodesia, Lusemska, H.C. Dollman, XII. 1916]; ?PT: [Nairobi, 

B.E.A]. 
Phytoecia haroldii Fâhraeus m. vitticollis Breuning, 1950c: 46. 

?PT: [Kilimandjaro, Sjöstedt., 1905.6. /Kibonoto, kulturz]; ?PT: [Mozambique, 

Zambezi] . 
Phytoecia kolbei Breuning m. flavoabdominalis Breuning, 1951: 445. 

2 ?PT: [Togoland, L. Conradt, 1892-1893]. 
Phytoecia kolbei Breuning m. rufoantennata Breuning, 1951: 444. 

HT: 9 , Togo, Bismarckburg. 
Phytoecia kolbei Breuning m. togoensis Breuning, 1951: 444. 

HT: 9 , Togo, Bismarckburg. 
Phytoecia kurdistana Ganglbauer m. bitlisensis Breuning, 1947e: 60. 

HT: S, Arménie. Bitlis [1912, Kulzer]; ?PT: [Bitlis, 1912, Kulzer]. 
Phytoecia kurdistana Ganglbauer m. quinquemaculata Breuning, 1947e: 60. 

HT: S, Erivan [Eriwan, 1898, Korb]; ?PT: [Eriwan, 1898, Korb]. 
Phytoecia larvata Pascoe m. fuscoantennata Breuning, 1951: 429. 

?PT: [Shilouvane. H. Junod]. 
Phytoecia maculicollis Peringuey m. anterufa Breuning, 1951: 413. 

HT: S , Transvaal [Grauer]; ?PT: [Transvaal, Grauer]. 
Phytoecia mannerheimi Breuning m. atroapicaloides Breuning, 1951: 369. 

HT: S , Chine, province de Shansi [Licent]. 
Phytoecia millefolii Adams m. obscuripennis Demelt, 1963b: 96. 

PT: Asia Minor, Silifke. 
Phytoecia neavei Aurivillius m. senegalensis Breuning, 1951: 428. 

2 ?PT: [Sénégal]. 
Phytoecia neavei Aurivillius m. unimaculicollis Breuning, 1951: 428. 

HT: 9 , Congo belge. Beni. 



CATALOGUE DES TYPES DE LAMIINAE 197 

Phytoecia nigriventris Kolbe m. anterufipes Breuning, 1951: 447. 

?PT: [Togo, Bismarckburg]. 
Phytoecia nigriventris Kolbe m. dimidiatoides Breuning, 1951: 448. 

?PT: [Belg Kongo]. 
Phytoecia nigriventris Kolbe m. fuscicornis Breuning, 1950c: 45. 

2 ?PT: [Kamerun, Mukonje Farm]. 
Phytoecia nigriventris Kolbe m. nigroantennata Breuning, 1950c: 45. 

?PT: [Belg Kongo, Beni]. 
Phytoecia nigriventris Kolbe m. rufina Breuning, 1950c: 45. 

2 ?PT: [Belg Kongo]. 
Phytoecia nigriventris Kolbe m. uninotata Breuning, 1950c: 45. 

2 ?PT: [Kamerun, Mukonje Farm]. 
Phytoecia orbicollis Reiche et Saulcy m. adanensis Breuning, 1947e: 60. 

HT: Cilicie, Adana [Asia Minor, Adana, coll. Sterba]; ?PT: [Asia Minor, Taurus 

eilic, Kricheldorff]. 
Phytoecia orbicollis Reiche et Saulcy m. ankarensis Breuning, 1964a: 97. 

HT: 8 , Anatolie, Ankara, Dikmen-deresi, 1. VI. 1941, leg. J. Romieux. 
Phytoecia orbicollis Reiche et Saulcy m. maraschensis Breuning, 1951: 54. 

HT: S, Marasch, leg. Sahlberg [Taurus]; ?PT: [Taurus, Marasch, V.28, leg. 

H. Kurzer]. 
Phytoecia pontica Ganglbauer m. discomaculata Breuning, 1947e: 61. 

?HT: Syrie, Damas, coll. Breuning [Kastamani]; ?PT: [Damascus /U. Sahib]. 
Phytoecia pretiosa Falderman m. basiimmaculata Breuning, 1947e: 61. 

HT: 9 , Perse; ?PT: [Armenien, Erze □ um, Reitter]. 
Phytoecia pretiosa Falderman m. nigroconjuncta Breuning, 1947e: 61. 

HT: Perse, Tabris. 
Phytoecia pseudafricana Breuning m. holorufa Breuning, 1951: 449. 

?PT: [Kenya, Ikutha]. 
Phytoecia pseudafricana Breuning m. subteratra Breuning, 1964a: 103. 

HT: o\ Uganda [Afr. c. coll. Plason]; ?PT: [Kenya, Mo □ □ □]. 
Phytoecia pseudolatesuturalis Breuning m. bivittipennis Breuning, 1962b: 15. 

?PT: [Kaimosi]. 
Phytoecia puncticollis Falderman m. nigrifrons Breuning, 1951: 86. 

HT: 9, Luristan, leg. Bodemeyer [Persia]. 
Phytoecia puncticollis Falderman m. nigroscapus Breuning, 1951: 86. 

HT: 2, Luristan, leg. Bodemeyer; ?PT: [Persia, Kereda]. 
Phytoecia puncticollis Falderman m. quadripunctata Breuning, 1951: 87. 

HT: 2, Luristan, leg. Bodemeyer [Persien]. 
Phytoecia puncticollis Falderman m. transitoria Breuning, 1951: 85. 

HT: 9, Luristan, leg. Bodemeyer [Persien]; ?PT: [Persien, Sultarabad, v. 

Bodemeyer]. 
Phytoecia pustulata Schrank m. coeruleomicans Breuning, 1947e: 59. 

HT: Maroc, Meknés. 
Phytoecia pustulata Schrank m. parterufoabdominalis Breuning, 1951: 386. 

HT: S, Anatolie, Ak-Chehir [V.26, Kulzer]; ?PT: [Roumanie, Comana Vlasca, 

A.L. Montandon]. 



198 J- MORATI 

Phytoecia rubropunctata Goeze m. confluentevittata Breuning, 1947e: 60. 

HT: France, Lyon [Kelecsényi]; ?PT: [Macon]. 
Phytoecia ruf a Breuning m. latesuturevittata Breuning, 1965b: 284. 

HT: 9 , Nordrhodesien, Abercorn [Rhodésie du Nord, Abercorn, 4.1 V. 1944, HJ. 

Brédo]. 
Phytoecia ruficollis Aurivillius m. ochreicollis Breuning, 1951: 408. 

HT: S , Oubanghi Chari, Bangui; ?PT: [Sénégal, Dakar]. 
Phytoecia rufiventris Gautier de Cottes m. partenigrecens Breuning, 1947e: 60. 

HT: ó\ Chine, province Foukien; ?PT: [Hondo, Chinzenji]. 
Phytoecia somereni Breuning m. fuscoapicalis Breuning, 1951: 449. 

2 ?PT: [Kenya, Ikutha]. 
Phytoecia sublateralis Breuning m. rufodiscalis Breuning, 1951: 435. 

HT: 9 , Angola, Bailundo. 
Phytoecia sylvatica Hintz m. atroapicalis Breuning, 1950c: 44. 

2 ?PT: [Belg Kongo, Ikutha]. 
Phytoecia vittata Peringuey m. nigrotibialis Breuning, 1951: 423. 

?PT: [Mosambique, Zoub □ □ □]. 
Phytoecia vittipennis Reiche m. ordubadensis Breuning, 1947e: 60. 

?HT: Arménie, Ordubad, coli. Breuning [Caucasus, Araxesthal, Leder. Reitter]; 

?PT: [Ordubad, 13 Klzr.]. 

Prosopocera vitticollis Gahan m. vittipennis Breuning, 1950b: 6. 

HT: 9 , Transvaal, Shilouane, leg. H. Junod, MHNG. {Exemplaire non retrouvé 
dans la collection). 

Saperda alberti Plavilstshikov m. latemaculata Breuning, 1952: 187. 

HT: 9 , Hokkaido, Insel Yesso. 
Saperda fayi Bland m. immaculipennis Breuning, 1952: 167. 

HT: ó\ Ontario, Ridgeway; ?PT: [ill. Ottawa?, 21.V). 
Saperda lateralis Fabricius m. transeuns Breuning, 1952: 165. 

?HT: Pensylvanie, Corwells, coll. Breuning [Kansas]. 
Saperda octopunctata Scopoli m. aurata Breuning, 1952: 183. 

HT: 9 , Lyon. 
Saperda vestita Say m. immaculata Breuning, 1976b: 740. 

HT: 6 , USA, Pennsylvania. 

Scythasis nitida Pascoe m. anticerufa Breuning, 1949: 245. 
?PT: [Burma, ill., Bhurnà]; ?PT: [Malacca, Perak]. 

Stenostola anomala Bates m. ohbayashii Breuning, 1952: 211. 

HT: Shikoku, Omogokei, 24. VII. 1932, S. Miyamoto, meiner Sammlung. 

{Etiqueté S. glenoides ohbayashi dans la collection). 

?HT: [Japan, 13.VIL1948 / Agoumai Prof., North Honshi, Obahata]. {Etiqueté 

S. anomala m. gleneoides dans la collection). 
Stenostola callosicollis Breuning m. incallosa Breuning, 1952: 205. 

HT: o\ Mandschurei, Handaohetzy. [VI. 1938]; ?PT: [idem]. 



CATALOGUE DES TYPES DE LAMIINAE 199 

Stibara tricolor Fabricius m. holorufipennis Breuning, 1961e: 160. 
2 ?PT: [Tonkin, Backan]. 

Thermistis croceocincta Saunders m. reducta Breuning, 1952: 198 (1956d : 696). 
HT: S, Tonkin, Dong- Van, coll. Lepesme; ?PT: [idem]. 

Thyestilla gebleri Faldermann m. subuniformis Breuning, 1952: 195. 

HT: ô , China, Umgebung Peking, leg. Stötzner; ?PT: [Peking, Exp. Stotzner, 

West berge]. 
Thyestilla gebleri Faldermann m. transitiva Breuning, 1952: 195. 

HT: S , China, Provinz Kiang-si, Umgebung Kiukiang. 

Tragocephala freyi Brancsik var. subnubeculosa Breuning, 1980a: 29. 
PT: Anosibe, coll. Dujardin-Delacour, Tananarive [XII. 68]. 

Zosne cachita Heller m. vitticollis Breuning, 1954a: 486. 
HT: 9 , Borneo, Sandakan, leg. Baker. 



REMERCIEMENTS 

Ce travail a bénéficié de l'aide logistique du département d'entomologie du 
Muséum d'histoire naturelle de Genève où il a été réalisé. Je remercie toutes les 
personnes qui m'ont assistée dans la réalisation de ce catalogue et tout particulièrement 
le Dr I. Lobi sous la direction duquel ce travail a été effectué. Sans ses conseils 
précieux, son aide considérable et sa constante disponibilité, il n'aurait pu être mené à 
son terme. Mes remerciements vont également au Dr C. Lienhard pour ses remarques 
pertinentes, au Dr C. Besuchet et à M. Rochas de Rolle pour la communication de la 
liste des «types» de la collection Rochas, à MM. C. Holzschuh de Vienne et P. Téocchi 
d'Avignon pour les références communiquées, ainsi qu'à MM. M. Danilevsky de 
Moscou et M. Sörensson de Lund pour les publications fournies. 



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Breuning, S. 1948. Nouvelles formes de Lamiaires (Première partie). Bulletin du Musée royal 
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du Musée du Congo belge, Tervuren, Sciences Zoologiques 4: 1-46. 

Breuning, S. 1950d. Quelques Lamiaires (Coleoptera, Cerambycidae) du Riksmuseum. Arkiv 
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Museum Journal 5: 373-388. 

Breuning, S. 1951. Révision du genre Phytoecia Muls. (Col. Cerambycidae). Entomologische 
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Breuning, S. 1954b. Nouvelles formes de Lamiaires (Septième partie). Bulletin de l'Institut 
royal des Sciences naturelles de Belgique 30 (41): 1-24. 

Breuning, S. 1954c. Lamiinae (Coleoptera, Cerambycidae). Parc national de l'Upemba, I. 
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Breuning, S. 1955b. Beschreibung einer neuen Rasse des Dorcadion divisum Germ. (Ceramb., 
Coleoptera). Annales Historico-Naturales Musei Nationalis Hungarici (series nova), 6: 
263. 



CATALOGUE DES TYPES DE LAMIINAE 201 



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royale d' Entomologie de Belgique 27: 138-152. 
Breuning, S. 1955e. Neue Lamiinae aus dem South African Museum (Coleoptera, Ceram- 

bycidae). Annals of the South African Museum 41: 345-357. 
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Naturales Musei Nationalis Hungarici, (series nova) 7: 403-404. 
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Humboldt-Universität zu Berlin (Coleoptera, Cerambycidae). Mitteilungen aus dem 

Zoologischen Museum in Berlin 37 : 297-328. 
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(Coleoptera Cerambycidae). Deuxième partie, VII. Description des espèces. Frustula 

Entomologica: 61-140. 
Breuning, S. 1961c. Nouvelles formes de Lamiaires (Treizième partie). Bulletin de l'Institut 

royal des Sciences naturelles de Belgique 37 (20): 1-44. 



202 J- MORATI 



Breuning, S. 1961d. Nouveaux Lamiaires d'Afrique centrale (Coleoptera Cerambycidae 

Lamiinae). Revue de Zoologie et de Botanique Africaines 63: 249-257. 
Breuning, S. 1961e. Neue oder schlecht bekannte Cerambyciden (Col.). Entomologische 

Arbeiten aus dem Museum G. Frey 12: 140-160. 
Breuning, S. 1962a. Révision systématique des espèces du genre Oberea Mulsant du globe 

(Coleoptera Cerambycidae). Frustula Entomologica: 141-232, 1-VI. 
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aus dem Staatlichen Museum für Tierkunde in Dresden 27, 665 pp. 
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Société Entomologique de Mulhouse: 95-103. 
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Onzième partie. Bulletin de la Société royale des Sciences naturelles du Laos 12: 41-62. 
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Reichenbachia Staatliches Museum für Tierkunden in Dresden 5: 283-284. 
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14 e partie. Bulletin de la Société royale des Sciences naturelles du Laos 15: 21-46. 
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Breuning, S. 1966c. Quelques nouvelles formes de Lamiinae de la collection Mussard 

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27-36. 
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Breuning, S. 1968a. Contribution à la connaissance des Lamiens du Laos [Coll. (sic) Céramb.] 

15 e partie et fin. Bulletin de la Société royale des Sciences naturelles du Laos 16: 3-64 

+ addenda: lp. 
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d'Ivoire (J. Decelle, 1961-1964), 21, Coleoptera Cerambycidae Laminae. Annales du 

Musée Royal de l'Afrique Centrale, Tervuren 165: 297-355. 
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42: 34-37. 
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Muséum de Paris. Bulletin du Muséum National d'Histoire naturelle, 2 e série 41: 

187-199. 
Breuning, S. 1970a. Nouveaux Coléoptères Cerambycidae Lamiinae des collections du 

Muséum de Paris (suite). Bulletin du Muséum National d'Histoire naturelle 42: 363-377. 



CATALOGUE DES TYPES DE LAMIINAE 203 



Breuning, S. 1970b. Nouveaux Coléoptères Cerambycidae Lamiinae des collections du 
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Breuning, S. 1971c. Descriptions de Lamiaires nouveaux du Musée Royal de l'Afrique Centrale 
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Breuning, S. 1972b. Quelques rectifications systématiques sur les Lamiaires (Col. Ceramby- 
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Breuning, S. 1972c (1971). Lamiaires nouveaux du nord de l'Inde récoltés par le Dr F. Schmid 
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Breuning, S. 1973a. Lamiaires nouveaux d'Afrique Centrale (Coleoptera Cerambycidae). Revue 
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Breuning, S. 1973b. Lamiaires nouveaux ou peu connus du Muséum de Paris [Col. Ceramby- 
cidae]. Annales de la Société entomologique de France (N. S.) 9: 647-665. 

Breuning, S. 1973c. Beschreibung zweier neuer Arten der Gattung Dorcadion (Col., Ceramby- 
cidae, Lamiinae). Entomologische Zeitschrift 83: 54-55. 

Breuning, S. 1974a. Quelques nouvelles espèces de Lamiinae du Muséum d'histoire naturelle 
de Genève (Coleoptera, Cerambycidae), I. Revue suisse de Zoologie 81: 73-75. 

Breuning, S. 1974b. Quelques nouvelles espèces de Lamiinae du Muséum d'histoire naturelle 
de Genève (Coleoptera, Cerambycidae), II. Revue suisse de Zoologie 81: 371-375. 

Breuning, S. 1974c. Descriptions de nouvelles formes de Lamiinae d'Ethiopie (Coleoptera 
Cerambycidae). Revue de Zoologie et de Botanique Africaines 88: 413-420. 

Breuning, S. 1974d. Descriptions de Lamiaires africains nouveaux (Coleoptera Cerambycidae). 
Revue de Zoologie et de Botanique Africaines 8 : 775-777 '. 

Breuning, S. 1974e. Neue Arten und Gattungen von Lamiinen (Coleoptera, Cerambycidae). 
Mitteilungen aus dem Zoologischen Museum in Berlin 50: 149-165. 

Breuning, S. 1974f. Neue Lamiinae (Coleoptera, Cerambycidae). Mitteilungen aus dem Zoo- 
logischen Museum in Berlin 50: 237-241. 

Breuning, S. 1974g. Nouveaux Mesosini des collections du Muséum national d'Histoire na- 
turelle (Col. Cerambycidae Lamiinae). Bulletin de la Société entomologique de France 
79: 72-79. 

Breuning, S. 1975. Contribution à l'étude des Cerambycidae Lamiinae du massif de l'An- 
dringitra, Madagascar Centre (Col.). Bulletin de la Société entomologique de France 80: 
258-265. 

Breuning, S. 1976a. Nouveaux Apomecynini des collections du Muséum National d'Histoire 
naturelle, Paris (Col. Cerambycidae Lamiinae). Bulletin de la Société entomologique de 
France 81: 207-208. 

Breuning, S. 1976b. Quelques nouvelles espèces de Lamiinae du Muséum d'histoire naturelle 
de Genève, III, (Coleoptera, Cerambycidae). Revue suisse de Zoologie 83: 737-741. 

Breuning, S. 1977a. Nouveaux Lamiaires africains des collections du Muséum de Paris (Col. 
Cerambycidae). Bulletin du Muséum National d'Histoire naturelle, 3 e série, n° 434, 
Zoologie 304: 265-303. 



204 J- MORATI 



Breuning, S. 1977b. Recherches sur le littoral de la Somalie. La plage et la dune de Sar Uanle. 
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dio per la faunistica ed ecologia tropicali del C.N.R.: CXXXIV). Monitore Zoologico 
Italiano [N. S.) 9: 291-298. 

Breuning, S. 1978a. Description de nouvelles espèces de Lamiaires de Nouvelle Calédonie, I. 
Folia Entomologica Hungarica, Rovartani Kozlemények (Series nova) 31: 23-29. 

Breuning, S. 1978b. Description de nouvelles espèces de Lamiaires de Nouvelle Calédonie, II. 
Folia Entomologica Hungarica, Rovartani Kozlemények (Series nova) 31: 67-71. 

Breuning, S. 1978c. Nouvelles espèces de Lamiinae (Cerambycidae) de la coll. Vives. Miscella- 
nea Zoologica 4: 131-136. 

Breuning, S. 1978d. Descriptions de Lamiaires africains nouveaux (Coleoptera Cerambycidae). 
Revue de Zoologie et de Botanique Africaines 92: 93-1 16. 

Breuning, S. 1979a. Beschreibung einer neuen Species der Gattung Dorcadion (Col.: Ceram- 
bycidae). Entomologische Zeitschrift 89: 92-93. 

Breuning, S. 1979b. Nouveaux coléoptères Cerambycidae Lamiinae des collections du Muséum 
de Paris (l ere note). Revue française d 'Entomologie (F.S.) 1: 99-100. 

Breuning, S. 1980a. Diagnoses préliminaires de nouveaux Lamiinae de Madagascar 
[Coleoptera, Cerambycidae]. Revue française d' Entomologie (N.S.) 2: 29-41 

Breuning, S. 1980b. Nouveaux Cerambycidae Lamiinae néotropicaux (Col.). Bulletin de la 
Société entomologique de France 85: 67-71. 

Breuning, S. 1980c. Nouvelles espèces de Lamiinae d'Asie orientale (Col. Cerambycidae). 
Bulletin de la Société Entomologique de Mulhouse: 49-50. 

Breuning, S. 1980d. Nouvelle espèce de Lamiinae des Iles Seychelles. Bulletin de la Société 
Entomologique de Mulhouse: 50. 

Breuning, S. 1980e. Description de nouvelles espèces de Lamiaires des Philippines (Coleoptera, 
Cerambycidae). Mitteilungen aus dem Zoologischen Museum in Berlin 56: 157-182. 

Breuning, S. 1981a. Beschreibungen neuer Lamiinae des Transvaal Museums, Pretoria (Coleo- 
ptera: Cerambycidae). Folia entomologica Hungarica 34(2): 35-54. 

Breuning, S. 1981b. Descriptions supplémentaires de Lamiinae, Sud Africains nouveaux 
(Coleoptera: Cerambycidae). Folia entomologica Hungarica 34(2): 55-59. 

Breuning, S. 1981c. Description d'une nouvelle race du genre Oberea Muls. (Cerambycidae). 
Bulletin de la Société Entomologique de Mulhouse: 26. 

Breuning, S. 1982. Diagnoses préliminaires de nouveaux Lamiinae du Muséum National 
d'Histoire naturelle de Paris [Coleoptera. Cerambycidae]. Annales de la Société ento- 
mologique de France (TV. S.) 18: 9-29. 

Breuning, S. 1984. Stephan von Breuning 1894-1983, Notice autobiographioque & Liste des 
publications. Bulletin de la Société Sciences Nat 41 : 1-17. 

Breuning, S. 1986. Annoted list of Cerambycidae, subfamily Lamiinae collected in Tanzania, 
Zambia and Botswana 1969-80 with description of 86 new taxa. In: Forchhammer, P. & 
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Demelt, C.v. 1963a. Beitrag zur Kenntnis der Cerambycidenfauna Kleinasiens und 13. Beitrag 
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Entomologische Blätter 59: 132-151. 

Demelt, C.v. 1963b. Eine neue Form der Phytoecia (Subgen. Helladia Frm.) millefolii Ad., 
Phytoecia millefolii Ad. m. obscuripennis m. n. Zeitschrift der Arbeitsgemeinschaft 
österreichischer Entomologen 15: 96. 

Dillon, L.S. & Dillon, E.S. 1948. The Tribe Dorcaschematini (Coleoptera: Cerambycidae). 
Transactions of the american entomological Society 73: 173-298. 

Fàhraeus, O. I. 1872. Coleoptera Caffrariae, annis 1838-1845 a J. A. Wahlberg collecta. 
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29-6Ì. 

Fiedler, H. 1938. Systematik und Zoogeographie von Ceroplesis (afrikan. Lamiinae, Col.). 
Deutsche Entomologische Zeitschrift 4: 571-616. 



CATALOGUE DES TYPES DE LAMIINAE 205 



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Cerambycid-fauna of the Loochoo-Archipelago, 2 (Col.). The Entomological Review of 

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Revue suisse de Zoologie 110(1): 207-246; mars 2003 



Small mammal inventory in the Shipstern Nature Reserve (Corozal 
District, Belize, Central America), a preliminary assessment 

Vincent BERSOT 

Muséum d'histoire naturelle, CP 6434, CH-121 1 Genève 6, Switzerland. 



Small mammal inventory in the Shipstern Nature Reserve (Corozal 
District, Belize, Central America), a preliminary assessment. - This 
small mammal inventory was conducted between May and September 2000 
in the Shipstern Nature Reserve located at the northeastern limit of Belize 
in the Corozal District, Central America. In 1 1840 trap nights, 7 species of 
non-volant small mammals were captured, including 5 species of myo- 
morph rodents and 2 species of didelphid marsupials. One species, Oto- 
tylomys phyllotis, accounted for 90.8% of the total captures, and may play a 
major role in the community structure of mammals in the reserve. The 
presence of Peromyscus yucatanicus in Shipstern Nature Reserve consti- 
tutes a first mention for Belize, and suggests that its range is extending more 
southerly to the limit agreed at present. Observations made on Sigmodon 
hispidus lead to formulate the hypothesis of a morphologically smaller 
population in the south of Quintana Roo, Mexico, and northern Belize, and 
further underlines the need for a complete revision of the genus in the 
Yucatan Peninsula. The sympatric diversity concretely obtained in Shipstern 
Nature Reserve is certainly underestimated, but nevertheless gives 
substance to the impression of a rodent community that is clearly affiliated 
to the partly endemic fauna of the Yucatan Peninsula, Mexico. 

Key-words: Belize - Yucatan Peninsula - small mammal inventory - 
Marmosidae - Didelphidae - Heteromyidae - Muridae. 

INTRODUCTION 

The information currently available to assess the magnitude and geographic 
distribution of mammalian diversity in Belize remains incomplete and unevenly repre- 
sentative. Indeed, most collecting has been concentrated in a few historically accessible 
areas in the Maya Mountains, and further north in the Yucatan Peninsula, and today, 
very few areas (if any) are characterized by a long history of field work on mammals. 
Among the most important mammalian surveys, Murie (1935) collected for several 
weeks in 1931 in the vicinity of Belize City and El Cayo (San Ignacio), British 
Honduras (today's Belize), on his way to Uaxactun, Petén, Guatemala. In 1939 and 
1940, after a brief survey in Jamaica and Haiti, LT. Sanderson & A. Sanderson spent 
several months collecting along the coast of British Honduras, from Punta Gorda to the 



Manuscript accepted 09.07.2002 



208 V. BERSOT 

Bahia de Chetumal, prior to continuing their studies in Quintana Roo, Yucatan, and 
Chiapas, Mexico - part of their collection was studied by Hershkovitz (1951). 
Subsequently, additional lists of mammals arose from various research projects. Those 
include Disney (1968), Dieckman (1973), Kirkpatrick & Cartwright (1975), Burton et 
al. (1987), Rabinowitz & Nottingham (1989), and most recently Caro et al. (2001). 
However, few studies ever focused on the small mammal fauna of Belize. Conse- 
quently, methods used to trap marsupials and small rodents often remained adapted to 
common and generalist species, but proved to be unsuitable to catch specialists and/or 
rare species. In this context, recent accounts of mammalian diversity in Belize, such 
as the checklist proposed by McCarthy et al. (1998), still probably underestimate the 
number of small mammal species actually found in the country - this is especially true 
for the northeastern portion of the country (Corozal District) where studies on the 
mammalian fauna remain scarce. 

In Shipstern Nature Reserve (SNR), as far as can be concluded from the 
available sources, non-volant small mammals were only briefly surveyed prior to the 
present study. This was done by Meerman (1993), and Miller et al. (1995), in a first 
attempt to survey the overall mammalian fauna of SNR. Apart from the latter, no other 
studies on mammals were reported from the reserve, with the exception of a bat 
inventory carried out by Bärtschi (1998). Nevertheless, other groups of vertebrates, but 
also invertebrates, were inventoried in SNR. Those include the following checklists: 
birds, reptiles, amphibians, and some miscellaneous invertebrates (Meerman, 1993); 
dragonflies and damselflies (Boomsma, 1993); butterflies (Meerman & Boomsma, 
1993); and freshwater fishes (Bijleveld, 1990). In addition, beside a checklist of the 
flora proposed by Meerman (1993), two research projects focusing on the vegetation 
of SNR were recently carried out by Bijleveld (1998) - structure and floristics - and 
Sayer (2000) - phenology. 

In northern Belize therefore, there is a clear need towards an aggressive pro- 
gramme of short-term mammalian surveys, that are combining the advantage of 
concreteness - listed species are definitely known to have occured together at a par- 
ticular place and time - with the disadvantage of incompleteness. Indeed, although 
indispensable, a long-term inventory project would inevitably be time-consuming and 
expensive, now bulldozers and chain saws work faster than field biologists (Voss & 
Emmons, 1996). In this context, the most important role for this project was to rapidly 
provide crucial diversity data - maximized by focusing on marsupials and rodents, that 
is taxa known to respond to ecological or Zoogeographie gradients - for future research 
and conservation objectives implicating the Shipstern Nature Reserve and its sur- 
rounding areas. In this respect, the primary objective was to obtain fresh material - 
voucher specimens that are well preserved, complete, precisely labeled, and docu- 
mented with exhaustive and informative data - to be subsequently incorporated in the 
collections of the Natural History Museum, Geneva (MHNG), Switzerland. In turn, the 
liver was sampled, and saved for DNA preservation, from each of these voucher spe- 
cimens, so as to constitute a precious tissue collection, completed with ear biopsies 
sampled from the specimens released in the field. These ear and liver samples were 
taken for inclusion into the Collection of Preserved Mammalian Tissues (Catzeflis, 
1991) held at Montpellier, France, where they are at the disposal of scientists. 



SMALL MAMMALS FROM BELIZE 209 

Apart from recording the small mammals and preserving their DNA, the other 
objective of this study was to collect their ectoparasites, and list the endoparasites of 
their digestive tract - giving priority to the classes Trematoda and Cestoidea, and the 
phylum Nemata - to be included in the MHNG museum collection. Indeed, obtaining 
parasites from mammals that are collected during inventories is also time-consuming, 
and such collections have rarely been made. Parasites and other symbionts are im- 
portant components of the biology of the host, however, and must be sampled for a 
complete picture of its ecology and other aspects of its life history. Of course, studies 
of the systematics and ecological characteristics of hosts and parasites require proper 
identification of both groups. Now, as far as the ecto- and endoparasites collected in 
SNR are concerned, this could not be achieved, and deeper analyses will be carried out 
within the framework of a subsequent study. 

STUDY AREA 

Previously owned by a British Limited Company, the area was partly managed 
as a private reserve during the eighties known as the Shipstern Wildlife Preserve. In 
1989, it became the property of the International Tropical Conservation Foundation 
(ITCF), and the name Shipstern Nature Reserve was given. This newly created foun- 
dation followed the opening of the tropical butterfly garden Papiliorama of Marin- 
Neuchâtel in Switzerland, in 1988. Today a major tourist attraction, and further 
developed as the Papiliorama Swiss Tropical Gardens in Kerzers, Fribourg, it is 
actively concerned with the conservation of tropical biodiversity through two sister- 
foundations: the Papiliorama-Nocturama Foundation in Switzerland, and the ITCF in 
Belize. Thus, among other projects involving the local community, two new protected 
areas were opened by the ITCF during the last few years: the Xo-Pol area in 1994; and 
the Sarteneja National Tree Park in 1999. In parallel, the ITCF assisted the Belize 
Coastal Unit in the creation of the Bacalar Chico National Park & Marine Reserve, in 
1996, which is part of the Belizean network of the UNESCO World Heritage Sites. At 
the same time in SNR, scientific work has been and is being further developed by the 
ITCF in conjunction with the universities of Bern, Neuchâtel, and Fribourg, 
Switzerland. 

Located at the northeastern limit of Belize in the Corozal District (Figs 1, 2), the 
Shipstern Nature Reserve covers an area of approximately 9000 hectares (22000 acres). 
It encompasses part of the Shipstern Lagoon, which is in direct contact with the south- 
ern waters of Chetumal Bay, and it also includes the separate surface of Xo-Pol (600 
hectares). By land, the only means of access to the area is an all-weather road starting 
at Orange Walk, and ending at the fishermen village of Sarteneja. 

Apart from a coastal plain stretching all along the Belizean coast, the whole area 
between Belmopan and Corozal (Fig. 1) is covered by lowlands that are further 
extending northward from Chetumal to Cancun on the eastern side of the Yucatan 
Peninsula, Mexico. To the southwest, these northern lowlands give way to the Maya 
Mountain massif, the dominant physical feature in Belize, which is the only area where 
large upfaulted blocks of intrusive and associated metamorphic sedimentary rocks are 
preponderant (Hartshorn et al., 1984). Such a substrata with only a thin soil cover 
means that water is essentially drained on the surface, thus constituing an extensive 



210 



V. BERSOT 




Fig. 1 
Political map of Central America and northern Belize. SNR, Shipstern Nature Reserve. 



system of streams and rivers. By contrast, a small scale karst topography is developed 
in the northern lowlands covered by tertiary limestones, and the area of SNR is thus 
characteristic with numerous sinkholes, caves, and natural wells, virtually no stream 
and river system on the surface, and elevations not exceeding 4-5 m (17 ft). 

Rainfall at the reserve was measured over a period of four years, between 
October 1989 and October 1993 (Meerman & Boomsma, 1993). An average annual 
rainfall of 1260 mm (49.6 inches) was recorded, with a minimum at 1029 mm (40.5 
in.), and a maximum at 1610 mm (63.4 in.) - for comparison, annual rainfall shows a 
considerable increase from north to south in Belize, averaging 1850 mm (73 in.) at 
Belize City (Belize District), and 4526 mm (178 in.) at Barranco (Toledo District). 
Such results suggest that Shipstern Nature Reserve is in one of the dryest areas in 
Belize, with a pronounced dry season - in the year of the present study, the months of 



SMALL MAMMALS FROM BELIZE 21 1 

February and March appeared to be exceptionnally dry, with no rainfall and 10 mm 
(0.4 in.) respectively (Sayer, 2000). As a consequence, the water level of the Shipstern 
Lagoon fluctuates on a yearly cycle, and areas close to the lagoon are temporarily 
flooded during the wet season. 

Indeed, rainfall tends to vary markedly in Belize. It starts with a pronounced dry 
season between the months of January and May. Then, from June to December, rainfall 
increases gradually to reach a maximum in September. And some years, this is pre- 
cisely any time from September to November that the wet season turns to a hurricane 
season. However, if Belize is frequently facing tropical storms, few hurricanes ever 
reach the country. And as far as the Shipstern Nature Reserve is concerned, among 
those that did in the past, only two are known to have caused serious damages. The first 
one, hurricane Janet, hit Belize in 1955, and greatly affected the northern part of the 
country. As a result, the villages of Sarteneja and Shipstern (Fig. 2), as well as Corozal 
Town were completely destroyed, whereas the forests in and around the present reserve 
were almost entirely flattened and burnt by subsequent fires. Because of this, the 
vegetation of Shipstern Nature Reserve can be considered as being 47 years old, and 
most of its vegetation types have not yet reached a structural climax. The second one, 
hurricane Keith, hit the country during the course of the present study, in early October. 
And if the reserve still suffered from localized falls of large trees, damages caused 
directly by the wind remained unextensive in the Corozal District-Keith mostly 
affected San Pedro (Ambergris Cay) and Cay Caulker, both comprised in the range of 
islands spreading all along the coast. However, unusual fluctuations of the water level 
caused by extreme rainfalls and inland accumulation of water carried in by the wind, 
led to serious floods. As a result, the water depth in the Shipstern lagoon was estimated 
at 4-6 feet, and the village of Chunox, as well as Orange Walk remained surrounded by 
floods for weeks. 

Temperatures exceeding 40°C (104°F) have already been measured in the coun- 
try (Walker, 1973). And, it is not uncommon to see the temperatures dropping as low 
as 10°C (50°F) between the months of November and December, when cold air masses 
enter the country from the north-east. More commonly, however, mean monthly 
minima range from 16°C (60.8°F) in winter to 24°C (75.2°F) in summer, and maxima 
from 28°C (82.4°F) to 33°C (91.4°F), with an annual mean temperature of 26°C 
(78.8°F) (King et al, 1992). 

Despite a long-established timber industry, and a history of successive destruc- 
tions due to natural phenomena, Belize remains covered by substantial areas of natural 
vegetation. Nowadays, however, an increased demography in Belize coincides with an 
increased pressure on the forest areas and on the fauna. Of primary concern is the rapid 
expansion of cultivated lands in the surroundings of the Shipstern Nature Reserve. 
Indeed, slash and burn cultivation is more and more common in the vicinity of Xo-Pol, 
whereas a worrying increase in agricultural activities carried on by Mennonites is 
observed from Little Belize towards Shipstern Lagoon. In addition, selective logging 
aimed at species such as Santa Maria {Callophyllum brasiliense), Ciricote (Cor dia 
dodecandra), and Mahogany (Swietenia macrophylla) is expanding throughout the 
eastern forests of Corozal District, and in particular those around Fireburn which are 
virtually empty of their large trees. 



212 



V. BERSOT 





Fig. 2 
Map of Shipstern Nature Reserve (SNR). 1-8, trapping sites; a, New Trail; b, Main Trail; 
c, Main Road; d. Shipstern Old Village. Grid : Universal Transverse Mercator and WGS 84. 

In this context, the Shipstern Nature Reserve plays an important role in the 
country representing one of the very few protected areas in the northern lowlands. 
Indeed, as far as Belize is concerned, the Yucatecan medium-sized semi-evergreen 
forests (Table 6) - still well represented over most of the Yucatan Peninsula, this forest 



SMALL MAMMALS FROM BELIZE 213 

type was probably covering the whole Corozal District before the expansion of sugar 
cane fields - as well as the very rare Yucatecan medium-sized semi-deciduous forests, 
are almost exclusively found in and around the reserve. Furthermore, the latter protects 
the unique patch of Low semi-deciduous Pseudophoenix s.sargentii forest distributed 
on mainland Belize, other patches being localized in the Bacalar Chico National Park 
& Marine Reserve, and in the Yucatan Peninsula mainly. 

With about a third of the surface being occupied by the Shipstern Lagoon, 
habitats in the reserve appear to be diverse and patchily distributed. Indeed, the lagoon 
constitutes an intricate system of shallow waters (1-3 feet) bordered by dwarf man- 
groves and other saline wetlands - listed in the Directory of Neotropical Wetlands 
(IUCN/TWRB, 1986) - and dotted with numerous small mangrove islands. Besides, so 
called "forest islands" are randomly distributed in the transitional areas with true 
forest. And the latter tends to vary considerably in its structural and floristic charac- 
teristics, thus creating a complicated mosaic of various forest types - the patches of 
which are sometimes less than 50 ft wide - that are not always clearly delimited. 
Aiming for a better understanding of its vegetation, Bijleveld (1998) listed seven main 
vegetation types within Shipstern Nature Reserve, to which the Cohune forest 
described by Wright et al. (1959) can be added (Table 6). 

MATERIALS AND METHODS 

Referring to the geographic distribution of the vegetation types listed in table 
6, four areas - named New Trail, Main Trail, Xo-Pol, and Shipstern for more conve- 
nience - were selected in and around Shipstern Nature Reserve. Chosen on the basis 
of their accessibility and the presence of ecotones between characterisitc habitats, two 
trapping sites were attributed to different localities within each of these areas. Thus, 
trapping sites 1/6, 2/5, 3/7, and 4/8, were representative of the New Trail, Main Trail, 
Xo-Pol, and Shipstern areas respectively (Fig. 2). 

Each of the eight trapping sites were identically built by reproducing a stan- 
dardized trap design (Patton et al., 2000). It consisted of two parallel transects 
separated by a distance of 500 ft (150 m). Each trapping line was 940 ft (285 m) in 
length and subdivided into 20 trap stations at 50 ft (15 m) intervals. In turn, four traps 
were positioned at each trap station, for a total number of 160 traps per trapping site. 

One folding Sherman (23 x 7.5 x 7.5 cm) and three folding BTS - Besançon 
Techniques Services, Besançon, France - mesh (28 x 10.5 x 10.5 cm) live traps were 
set at each trap station. Traps were placed in the most likely microhabitat within 7 to 
17 ft (2-5m) of the station marker. Wherever possible, two BTS mesh live traps were 
positioned from a height of 4 to 14 ft (1-4 m), whereas the third one remained terres- 
trial. The Sherman live trap was always set on the ground. For bait, the dried fruit of 
Guaya trees (Talisia olivaeformis) was fixed on the hook of BTS mesh live traps, and 
coated with peanut butter - no baits were used for Sherman live traps. 

All traps were left open for 10 consecutive nights, thus totalling 1600 "trap 
nights" per locality (Wilson et al., 1996). Eight trapping sessions were scheduled over 
a 6 month time interval in the year 2000, and effective trapping periods took place at 
the dates presented in table 3. The last trapping session (site n°8) totalled only 640 trap 



214 V. BERSOT 

nights, having been brutally interrupted under the threat of hurricane Keith. Therefore, 
trapping effort for all 8 trapping sessions together totalled 1 1840 trap nights. 

First captures of newly encountered species were systematically brought back 
to the base camp for a first identification based on the external morphology. At the same 
time, ectoparasites and intestinal endoparasites were sampled along with the liver. 
After collection of several male and female adults within one species, the specimens 
captured successively and satisfactorily identified as belonging to the same species 
were only then released. Standard data - external measurements, weight, sex, age,...- 
along with an ear sample - a triangular incision of 4-5 mm in the ear pinna - were 
collected from each individual to be released. The animal was thus marked and 
recognized if recaptured. 

External measurements are expressed as follows (Nagorsen & Peterson, 1980; 
Catzeflis et al., 1997; Musser et al., 1998): E, ear length (distance from the base of the 
notch of the lower part of the ear to the uppermost margin of the ear); HF, hind foot 
(distance from the end of the heel bone, calcaneum, to the end of the longest toe, 
exclusive of the claw); NEB, nose-ear basis (distance from the tip of the nose to the 
base of the notch of the lower part of the ear); NEC, nose-eye corner (distance from 
the tip of the nose to the anterior corner of the eye); SR, scale rows (number of scale 
rows in a centimeter section of the tail about one-third from the body); TL, total length 
(straight-line distance from the tip of the nose to the end of the last tail vertebra, 
exclusive of hair); TV, tail vertebrae (distance from the base of the tail to the tip of the 
last vertebra, exclusive of hair). 

At MHNG, the voucher specimens were essentially re-identified by referring to 
their corresponding skull measurements and cranial characters. The latter data was 
compared to that found in available literature by first assigning age classes - juvenile, 
immature, subadult, adult, or old adult (Wilson et al., 1996) - based on tooth eruption 
and wear to each specimens. Then, whenever a set of skull measurements was not 
clearly affiliated to one species, distinctive cranial characters were considered as 
reliable. In addition, the data obtained in the field was finally reexamined, along with 
the known distribution of the diagnosed species. As a result, the final identification of 
each specimen was based on the following data: external morphology, external and 
skull measurements, cranial characters, and distribution. 

In murid rodents, skull measurements (Fig. 3) are expressed as follows (Patton 
et al., 2000; Musser et al., 1998; Voss, 1988): BB, braincase breadth; BBP, breadth of 
bony palate; BIF. breadth of incisive foramina: BL. length of auditory bulla; BM1, 
breadth of first upper molar; BOL, basioccipital length; BR, breadth of rostrum; BW, 
breadth of auditory bulla; BZP, breadth of zygomatic plate; CBL, condylobasal length; 
CD, cranial depth; CIL, condyloincisive length; CLM1-3, crown length of maxillary 
toothrow; CLMT, crown length of mandibular toothrow; DI, depth of incisor; HI, 
height of incisor; IB, interorbital breadth; LB, lambdoidal breadth; LBP, length of 
bony palate; LD, length of diastema; LIF, length of incisive foramina; LM, length of 
mandible; MB, mastoid breadth; MPFL, mesopterygoid fossa length; MPFW, 
mesopterygoid fossa width; NL, nasal length; OCB, occipital condyle breadth; OL, 
orbital length; ONL, occipitonasal length; PL, palatal length; PPL, postpalatal length; 
RL, rostral length; ZB, zygomatic breadth. 



SMALL MAMMALS FROM BELIZE 



215 




Fig. 3 
Rodents, limits of 33 cranial and dental dimensions. Abbreviations are defined in text. 



216 V. BERSOT 

In marsupials, skull measurements (Fig. 4) are expressed as follows (Patton et 
al, 2000; Gardner, 1973): BB, braincase breadth; BN, greatest breadth of nasals; BOL, 
basioccipital length; BRF, breadth of rostrum across frontals; BRJ, breadth of rostrum 
across jugals; CBL, condylobasal length; CD, cranial depth; CLMT, molar toothrow 
length (mandible); C-M4, maxillary toothrow length; IB, interorbital breadth; IOC2, 
postorbital constriction; LM, length of mandible; MB, mastoid breadth; M1-M4, mo- 
lar toothrow length (maxillary); NL, nasal length; OCB, occipital condyle breadth; 
ONL, occipitonasal length; PL, palatal length; PPL, postpalatal length; PW, palatal 
width; RL, rostral length; RW, rostral width; ZB, zygomatic breadth. 

A first capture (Table 1) was defined as the first occasion that an individual was 
trapped and marked (Stallings, 1989). The first capture plus subsequent captures of 
each specimen were considered total captures. Recapture indices were calculated by 
dividing total captures by first captures, thus indicating the average number of times an 
individual of each species was captured. The number of trap nights (Table 3) was cal- 
culated by the number of traps multiplied by the number of nights the traps were acti- 
vated per site (Wilson et al., 1996). The trapping success (Table 3) for a site was the 
number of total captures of all species divided by the number of trap nights and ex- 
pressed in percentage. The standard error for each external and cranial measurements 
(Tables 4, 5) was calculated after Fowler et al. (1998). 

RESULTS 

Capture results by species for the eight trapping sites are presented in table 1 . 
Seven small mammal species were sampled: Marmosa mexicana (Merriam, 1897), 
Didelphis virginiana (Kerr, 1792), Heteromys gaumeri (Allen & Chapman, 1897), 
Otonyctomys hatti (Anthony, 1932), Ototylomys phyllotis (Merriam, 1901), Pero- 
myscus yucatanicus (Allen & Chapman, 1897), and Sigmodon hispidus (Say & Ord, 
1825). Overall, O. phyllotis represented 90.8% and 86.6% of the total and first captures 
respectively. On average, individuals of this species were recaptured at a rate of 1.8 
times. Trapping success by trapping site is presented in table 3. A total of 414 captures 
was obtained in 1 1840 trap nights, thus yielding an overall trapping success of 3.5%. 
Table 2 presents for each species the number of total captures per vegetation type 
(Table 6). However, caution should be taken when comparing the latter results, as the 
trapping effort is unequal from one vegetation type to another. External and cranial 
measurements by species presented in tables 4 and 5 combine adult males and females. 
Here again, attention should be given as the single specimens of D .virginiana and 
O. hatti are immature and subadult respectively. 

SPECIES ACCOUNTS 

The following accounts present the essential observations regarding the mor- 
phological description - skull and hindfoot - the local distribution, and the habitat of 
the 2 species of marsupials and 5 species of rodents captured in Shipstern Nature 
Reserve. Additional observations of these species are presented in the referred 
literature. The attribution of a name species to the specimens collected in SNR is also 
discussed, knowing that few genera of Neotropical rainforest mammals are wholly free 
of taxonomic problems. 



SMALL MAMMALS FROM BELIZE 



217 



— BRF— 






Fig. 4 
Marsupials, limits of 9 cranial and dental dimensions. Abbreviations are defined in text. Other 
measurements, see figure 3. 



218 V. BERSOT 

Family Marmosidae 

Marmosa mexicana (Merriam, 1897) Figs 5, 8 

Mexican Mouse Opossum 

Description. The cranium is broad and moderately elongate, with a convex 
dorsal profile (Fig. 5). The rostrum is rather short and broad, concealing the naso- 
lacrimal capsules in dorsal view. The nasals are long and truncate posteriorly, project- 
ing anteriorly beyond the premaxillae to conceal the nasal orifice from dorsal view. 
The zygomatic arches are stout, convex from the dorsal aspect, convergent anteriorly, 
and forming a clear elbow-shaped bend in lateral view. The inferior zygomatic root is 
running on from M4 posteriorly. The interorbital region is broad relative to braincase, 
without pronounced postorbital constriction. The supraorbital ridges are moderately 
grooved dorsally, project laterally only slightly, and transform posteriorly into 
conspicuous parietal ridges on top of braincase. The braincase is narrow, uninflated, 
with slanting lateral, and nearly vertical rear walls. Occipital ridges are present, but 
temporal and lambdoidal ridges are not. The auditory bullae are small and moderately 
rounded, and their anterior margin lies anteriorly to the basisphenoid/basioccipital 
suture. The basisphenoid is narrow and elongate, contrasting with a short and wide 
basioccipital. The bony palate is broad, fenestrated, and extends beyond the molar rows 
posteriorly. The anterior margin of the maxillary /palatine suture lies anteriorly to M2. 
The incisive foramina are short, inconspicuous, and their posterior margin is level with 
the posterior face of the upper canines. The teeth are small and number 50. The 
maxillary toothrows are straight along their labial margins, and convergent anteriorly. 
The canines are rather short and slender, and PM1 is remarkably smaller than PM2. 
The dental formula is : i 5/4. c 1/1, p 3/3. and m 4/4. The dentary has a long and stout 
coronoid process, a well developed condyloid process, a rather delicate angular 
projection, and a moderately deep and elongate ramus. 

Pes has swivelling ankle joints, and opposable hallux (Fig. 8). The plantar sur- 
face of pes is unpigmented, devoid of distinctive scalation, naked from heel to tips of 
the digits, with little fringing hairs along its contour, and with four separate interdigi- 
tal and two separate carpal pads; the thenar and first interdigital, and the hypothenar 
and fourth interdigital pads are nearly fused. Dorsal and lateral surfaces of the meta- 
tarsal region and digits are a pale cinnamon and naked-looking, sparsely covered with 
very short white hairs. Digit I is devoid of claw, and claws on digits II to V are short 
and unpigmented. with inconspicuous tufts of hair at their base. 

Discussion. M .mexicana is easily confused with M.robinsoni. But, whereas 
M.m. is ubiquitous and a typical Central American endemic (McCarthy et al., 1998), 
M.r. was previously known only from Panama, with disjunct populations in southern 
Belize and Roatân Island in Honduras (Reid, 1997). However, M.r. has recently been 
reported in Guatemala and El Salvador, so that it may be present throughout Central 
America, and specimens may have been confused with M.m.. A reappraisal of all 
specimens of the two species in Central America is therefore needed to determine their 
exact distribution. 

On the basis of external characters, M.m. and M.r. are hardly distinguishable, 
although M.r. is usually larger, with a longer and slightly wooly fur (Reid, 1997). Thus. 



SMALL MAMMALS FROM BELIZE 219 

these species are best distinguished by skull characters when adult. In the case of 
VB054 (MHNG n° 18 12.007), external and cranial measurements (Tables 4, 5) tally 
perfectly with those oiM.m. in Alonso-Mejia & Medellin (1992), but are rather at the 
bottom of the ranges given for M.r. in O'Connell (1983). Furthermore, it is clearly 
characterized by its supraorbital ridges that are moderately grooved dorsally, and 
project laterally only slightly (Alonso-Mejia & Medellin, 1992), and its postorbital 
constriction that is not as pronounced as in M.r. (O'Connell, 1983). 

Serious conclusions on habitat and local distribution cannot be drawn from a 
single capture (Table 1). Nevertheless, apart from the strong morphological adaptations 
towards arboreality observed in VB054, the habitat description in Reid (1997) leads to 
infer that M.m. is probably ubiquitous in the forests of Shipstern Nature Reserve. 

Family Didelphidae 

Didelphis virginiana (Kerr, 1792) Figs 5, 8 

Virginia Opossum 

Description. The cranium is elongate and broad, with a convex dorsal profile 
(Fig. 5). The rostrum is long and broad, concealing the nasolacrimal capsules in dorsal 
view. The nasals are long, narrow, and truncate posteriorly, revealing the nasal orifice 
in dorsal view. The lateralmost extent of each nasal, where intercepted by the maxillo- 
frontal suture, is aligned from the point where the same suture meets the lacrimal. The 
lacrimal recedes from the outer margin of the jugal before terminating in an acute point 
on the superior zygomatic root. The zygomatic arches are stout, convex from the dor- 
sal aspect, and convergent anteriorly. The inferior zygomatic root is running on from 
M2 posteriorly; M3 is barely erupting. The dorsal extension of the palatine bones that 
make up part of the inner wall of the orbits is broad. The interorbital region is broad 
relative to braincase, without pronounced postorbital constriction. The supraorbital 
ridges are unextensive, moderately grooved dorsally, project laterally only slightly, and 
transform posteriorly into conspicuous but short and convergent parietal ridges on top 
of braincase. The braincase is narrow, uninflated, with slanting lateral, and nearly 
vertical rear walls. Occipital ridges are present, but temporal and lambdoidal ridges are 
not. The auditory bullae are remarkably small and barely rounded, and their anterior 
margin lies slightly anteriorly to the basisphenoid/basioccipital suture. The basi- 
sphenoid is narrow and rather elongate, contrasting with a short and wide basioccipital. 
The bony palate is broad, fenestrated, and extends beyond the molar rows posteriorly. 
The anterior margin of the maxillary/palatine suture lies slightly posteriorly to the 
margin between Ml and M2. M4 and PM3 are absent, whereas M3 and m4 are 
erupting. The incisive foramina are short, inconspicuous, and their posterior margin is 
level with the anterior face of the upper canines. The teeth are fairly broad and number 
44. The maxillary toothrows are straight along their labial margins, and convergent 
anteriorly. The canines are rather short and stout, and PM1 is remarkably smaller than 
PM2. The dental formula is: i 5/4, c 1/1, p 2/2, and m 3/4. The dentary has a long and 
broad coronoid process, a well developed condyloid process, a small but rather stout 
angular projection, and a deep and elongate ramus. 



220 V. BERSOT 

Pes has swivelling ankle joints, and opposable hallux (Fig. 8). The plantar sur- 
face of pes is dark brown, with distinctive scalation, naked from heel to tips of the 
digits, with little fringing hairs along its contour, and with four separate interdigital and 
one carpal pads; the hypothenar and fourth interdigital pads are fused, and the thenar 
and first interdigital pads are nearly so. Dorsal and lateral surfaces of the metatarsal 
region and digits are blackish and sparsely covered with short black hairs. Digit I is 
devoid of claw, and claws on digits II to V are relatively long and unpigmented, with 
tufts of black hair at their base. 

Discussion. D.virginiana may be difficult to distinguish with certainty from 
D.marsupialis by sight. Also, both species are ubiquitous in Central America - D.m. 
extends from NE Argentina, Bolivia, and Peru in South America to northern Mexico 
(Reid, 1997), whereas D.v., which is thought to have arisen from D.m. (Gardner, 1973), 
is not found any further than NW Costa Rica, but extends northerly to southern Canada 
- and are expected to be sympatric in Shipstern Nature Reserve. 

Perhaps the most distinctive and easily observed external character separating 
D.v. and D.m. is the hair color of the cheek region (Gardner, 1973): it is pure white in 
D.v., and clearly set off from the rest of the head by the darker color of the sides and 
neck behind, and by a dark band extending from the ear through the eye above. Also, 
the extent of the black pigmented basal portion of the tail ranges from half to all black 
in D.v. (Reid, 1997), whereas it tends to be less in D.m., a criterion however considered 
as too variable for broad application by Gardner (1973). Additional characters 
observed in D.v., such as long white mystacial and genal whiskers, and more evenly 
distributed guard hairs over the dorsum, can be useful to separate D.v. from D.m., but 
are still not determining in some cases. Among others, the single specimen captured in 
this study (VB062, MHNG n°l 8 12.008) is assigned to the age class 2 (Gardner, 1973), 
and is probably too young to be distinguished with certainty. For the same reason, the 
external and cranial measurements of VB062 (Tables 4, 5) are not comparable with 
those given for D.v. in McManus (1974), Gardner (1973), and Goodwin (1969). 
Nevertheless, D.v. and D.m. can be separated by skull characters that are independent 
of age classes (Gardner, 1973). Thus, VB062 is clearly characterized by its bone 
configuration in the naso-lacrimal and lacrimo-jugal regions, and on the inner wall of 
the orbit. 

Just as in the case of VB054, the clear morphological adaptations observed in 
VB062 confirm that it is highly scansorial. Furthermore, the habitat description in Reid 
(1997), and McManus (1974), leads to infer that D.v. is also ubiquitous in the forests 
of SNR. 

Family Heteromyidae 

Heteromys gaumeri (Allen & Chapman, 1897) Figs 6, 8 

Gaumer's Spiny Pocket Mouse 

Description. The cranium is elongate and rather slender, with a convex dorsal 
profile (Fig. 6). The rostrum is long and broad, concealing the nasolacrimal capsules 
in dorsal view. The nasals are long with slightly convergent outer margins posteriorly, 
and they project anteriorly beyond the premaxillae to conceal the nasal orifice from 



SMALL MAMMALS FROM BELIZE 



221 





Fig. 5 
Top, Didelphis virginiana (VB062/MHNG n° 18 12.008); bottom, Marmosa mexicana 
(VB054/MHNG n°1812.007). Dorsal, ventral, and lateral view of cranium and lateral view of 
left dentary. Scale bars: 5 (M.m.) and 7 CD. v.) mm. 



dorsal view. The zygomatic arches are slender, only slightly convex from the dorsal 
aspect, and convergent anteriorly. The interorbital region is remarkably broad relative 
to braincase. Supraorbital shelves accentuate the dorsolateral margins of the interorbit 
and transform posteriorly into low temporal ridges extending to inconspicuous lamb- 



222 V. BERSOT 

doidal ridges. The supraorbital ridges are moderately grooved dorsally. Occipital ridges 
are present, and the supraoccipital crest is angular but barely visible in dorsal view. The 
braincase is moderately inflated but not globose, with nearly vertical lateral and rear 
walls. The auditory bullae are moderately large and not clearly rounded; their anterior 
margin lies anteriorly to the basisphenoid/basioccipital suture. The basisphenoid is nar- 
row and short, contrasting with an elongate and moderately wide basioccipital. The 
bony palate is narrow, without fenestration, and extends beyond the molar rows poste- 
riorly. The anterior margin of the maxillary /palatines suture is level with M2. The in- 
cisive foramina are very short and inconspicuous; the premaxillary-maxillary suture 
joins their posterior margin. The teeth number 20. The molar toothrows are straight 
along their labial margins, and convergent anteriorly. Incisors are moderately large 
with smooth orangish enamel faces; in lateral profile, the uppers curve back toward the 
molar rows, an opisthodont configuration. The dental formula is : i 1/1, c 0/0, p 1/1, 
and m 3/3. The dentary has a short coronoid process, a rather long and stout condyloid 
process, a well developed angular projection and a deep but short ramus; the capsular 
process is visible. 

The plantar surface of pes is unpigmented around the interdigital pads and on 
digits, and is otherwise dark brown; it has four separate interdigital and two separate 
carpal pads, with distinctive scalation (Fig. 8); it is thinly covered with short dark 
brown or reddish hairs from heel to thenar and hypothenar pads, with little fringing 
hairs along its contour. Dorsal and lateral surfaces of the metatarsal region and digits 
are whitish, and thinly covered with short white hairs. The three middle digits are 
much longer than the outer two, and interdigital webbing is present between digits II, 
III, and IV. The claw on digit II is enlarged. Claws are otherwise short and unpig- 
mented, with inconspicuous tufts of hair at their base. 

Discussion. H.gaumeri is endemic to the Yucatan Peninsula, and its southern 
range extends no further than N Belize and N Guatemala (Reid, 1997), where it is 
likely to be found in sympatry with H. de smare stianus — although a typical Central 
American endemic (McCarthy et al., 1998) seemingly absent from the state of Yucatan, 
Mexico, H.d. has been reported to the west of Chetumal, Quintana Roo, Mexico (Jones 
et al, 1974). 

However, H.g. differs from all other species of the genus Heteromys by the pre- 
sence of hair on the posterior portion of the sole of the hind foot (Schmidt et al., 1989). 
Also, whereas very few skull characters are known to separate them, H.g. is distin- 
guished from H.d. by a discontinuous pelage coloration between the outer surfaces of 
front legs and dorsum (Reid, 1997), a relatively well haired tail with a conspicuous 
terminal tuft, and by averaging smaller in most external and cranial measurements. 
Thus, VB017 (MHNG n°181 1.098), VB021 (MHNG n°181 1.100), and VB022 (MH- 
NG n° 18 12.001) tally perfectly with the external and cranial measurements (Tables 4, 
5) of H.g. in Schmidt et al. (1989), and Jones et al. (1974), but are below those given 
for H.d. in Rogers & Schmidly (1982), and Goodwin (1969). 

All specimens of H.g. were caught on the ground, and ran away on the ground, 
sometimes directly into a near cavity, when released. Besides, no morphological adap- 
tations towards arboreality were observed from sampled specimens, thus suggesting 
that H.g. is strongly terrestrial. Furthermore, the fact that H.g. was captured in all sites 



SMALL MAMMALS FROM BELIZE 223 

except n°l (Table 1), and in all forest habitats (Table 2), leads to infer that it is ubi- 
quitous in the forests of Shipstern Nature Reserve, and probably absent from exposed 
areas covered by vegetation types 6 and 7. 

Family Muridae 

Otonyctomys hatti (Anthony, 1932) Figs 6, 8 

Yucatan Vesper Mouse 

Description. The cranium is short and rather angular, with a convex dorsal pro- 
file (Fig. 6). The rostrum is short and narrow, not concealing the nasolacrimal capsules 
in dorsal view. The nasals are short, and project anteriorly beyond the premaxillae to 
conceal the nasal orifice from dorsal view. The zygomatic arches are slender, nearly 
parallel from the dorsal aspect, strongly convergent anteriorly, and forming downward 
a pronounced arc-shaped bend in lateral view. The interorbital region is narrow relative 
to braincase. Supraorbital shelves accentuate the dorsolateral margins of the interorbit 
and transform posteriorly into temporal ridges extending to conspicuous lambdoidal 
ridges. The supraorbital ridges are moderately grooved dorsally. Occipital ridges are 
present, and the supraoccipital crest is rounded but barely visible in dorsal view. The 
braincase is well inflated but not globose, with nearly vertical lateral and rear walls. 
The auditory bullae are greatly enlarged and rounded, and their anterior margin lies an- 
teriorly to the basisphenoid/basioccipital suture. Both the basisphenoid and the basioc- 
cipital are narrow and elongate. The bony palate is moderately narrow, without fenes- 
tration, and does not extend beyond the molar rows posteriorly. The anterior margin of 
the maxillary /palatines suture is level with the margin between Ml and M2. The inci- 
sive foramina are rather long and conspicuous, and their posterior margin lies slightly 
posteriorly to the anterior margin of the orbits. The teeth number 16. The molar 
toothrows are straight along their labial margins, and parallel. Incisors are moderately 
large with smooth orangish enamel faces; in lateral profile, the uppers curve back 
toward the molar rows, an opisthodont configuration. The dental formula is: i 1/1, c 0/0, 
p 0/0, and m 3/3. The dentary has a short coronoid process, a very long and stout 
condyloid process, a well developed angular projection, and a rather deep but short 
ramus; the capsular process is barely visible. 

Pes has moderately swivelling ankle joints, but no opposable hallux (Fig. 8). 
The plantar surface of pes is unpigmented on digits and otherwise purplish, devoid of 
distinctive scalation, naked from heel to tips of the digits, with little fringing hairs 
along its contour, and with four separate interdigital and two separate carpal pads. 
Dorsal and lateral surfaces of the metatarsal region and digits are pale brown and 
naked-looking, thinly covered with short white hairs. Claws are short and unpig- 
mented, with a tuft of white hair at their base. 

Discussion. Just as H.gaumeri, O. hatti is endemic to the Yucatan Peninsula, and 
its southern range extends no further than N Belize and N Guatemala (Reid, 1997; 
Peterson, 1965). Consequently, although O.h. may be confused with Nyctomys 
sumichrasti, their respective ranges are not overlapping - N.s. is a typical Central 
American endemic (McCarthy et al., 1998) seemingly absent from the Yucatan Penin- 
sula, Mexico, apart from southern Belize (Jones et al., 1974) - and they have never 
been reported as sympatric as yet. 



224 



V. BERSOT 





Fig. 6 
Top, Heteromys gaumeri (VB017/MHNG n° 181 1.098); bottom, Otonyctomys hatti (VB182/ 
MHNG n° 18 12.009). Dorsal, ventral, and lateral view of cranium and lateral view of left dentary. 
Scale bars: 5 mm. 



The single specimen captured in this study (VB182, MHNG n°l 8 12.009) is 
assigned to the age class II (Genoways & Jones, 1972), and is probably too young to 
be distinguished with certainty, on the basis of external characters. However, even 



SMALL MAMMALS FROM BELIZE 225 

knowing that very few skull characters separate O.h. from N.s., the skull of VB182 is 
easily characterized by its greatly enlarged auditory bullae, which are more than twice 
as large as those of N.s. (Peterson, 1965). 

VB 1 82 was found to have a remarkable agility to climb on all surfaces of its 
mesh wire trap. Also, the morphology of its hindfeet with moderately swivelling ankle 
joints leads to infer that it is scansorial. Besides, by referring to the habitat description 
in Reid (1997), O.h. is likely to be ubiquitous, although rare, in the forests of SNR. 

Ototylomys phyllotis (Merriam, 1901) Figs 7, 8 

Big-eared Climbing Rat 

Description. The cranium is elongate and rather angular, with a flattened dorsal 
profile, gently convex over braincase (Fig. 7). The rostrum is long and narrow, not 
concealing the nasolacrimal capsules in dorsal view. The nasals are rather long and 
terminate in a rounded angle anteriorly, projecting beyond the premaxillae to conceal 
the nasal orifice from dorsal view. The zygomatic arches are stout, fairly angular from 
the dorsal aspect, and strongly convergent anteriorly. The interorbital region is rather 
broad relative to braincase. Supraorbital shelves accentuate the dorsolateral margins of 
the interorbit and transform posteriorly into low temporal ridges extending to pro- 
minent lambdoidal ridges. The supraorbital ridges are moderately grooved dorsally. 
Occipital ridges are present, and the supraoccipital crest is clearly angular in dorsal 
view. The braincase is boxlike, with nearly vertical lateral and rear walls. The auditory 
bullae are large and rounded, and their anterior margin lies anteriorly to the basisphe- 
noid/basioccipital suture. Both the basisphenoid and the basioccipital are moderately 
wide and elongate. The bony palate is moderately narrow, without fenestration, and 
does not extend beyond the molar rows posteriorly. The anterior margin of the maxil- 
lary/palatines suture is level with M2. The incisive foramina are long, broad, conspi- 
cuous, and their posterior margin lies slightly posteriorly to the anterior margin of Ml. 
The teeth number 16. The molar toothrows are straight along their labial margins, and 
parallel. Incisors are moderately large with smooth orangish enamel faces; in lateral 
profile, the uppers curve back toward the molar rows, an opisthodont configuration. 
The dental formula is: i 1/1, c 0/0, p 0/0, and m 3/3. The dentary has a short coronoid 
process, a long and stout condyloid process, a broad and well developed angular 
process, and a moderately deep and short ramus; the capsular process is barely visible. 

The plantar surface of pes is unpigmented on digits and pads, and otherwise 
purplish, devoid of distinctive scalation, naked from heel to tips of the digits, with little 
fringing hairs along its contour, and with four separate interdigital and two separate 
carpal pads (Fig. 8). Dorsal and lateral surfaces of the metatarsal region and digits are 
greyish white, and thinly covered with short white hairs. Claws are short and unpig- 
mented, with a tuft of white hair at their base. 

Discussion. O. phyllotis is a typical Central American endemic (McCarthy et 
ai, 1998), ubiquitous in the Yucatan Peninsula. 

Whereas few skull characters are distinctly separating O.p. from a species such 
as Tylomys nudicaudus - also a Central American endemic, seemingly absent from the 
states of Campeche, Yucatan, and Quintana Roo, Mexico (Reid, 1997; Jones et al., 



226 V. BERSOT 

1974), but likely to be sympatric with O.p. throughout Belize - on the basis of external 
characters, it can hardly be confused with any other species. Indeed, O.p. is character- 
ized by a conspicuous patch of white fur at the ear basis. Also, if both species have a 
shiny, naked-looking, and broadly scaled tail, its distal half is white or yellow 
pigmented in T.n. (Lawlor, 1982). Besides, O.p. averages smaller in most external and 
cranial measurements. In the case of VB001 (MHNG n°18 11.089), VB002 (MHNG 
n°181 1.090), VB004 (MHNG n°181 1.091), VB005 (MHNG n°181 1.092), VB063 
(MHNG n°1811.094), VB105 (MHNG n°181 1.096), VB121 (MHNG n°181 1.097), 
and VB128 (MHNG n° 18 12.061), however, the latter are rather at the bottom, or even 
below (Tables 4, 5), the measurements given for O.p. in Lawlor (1982). Nevertheless, 
the same author (1969) reported that O.p. is characterized by considerable local and 
geographic variation in size and pelage coloration - it varies gradually from small and 
dusky brown in the lowlands of the Yucatan Peninsula, to large and dark greyish-brown 
in Costa Rica, and in the highlands of Chiapas and Guatemala. 

Specimens of O.p. were caught at all levels from the ground to a height 
exceeding three meters. Also, when released, most of them ran away on the ground, 
sometimes directly into a near cavity, whereas a fraction of them climbed on the 
surrounding vegetation before returning to the ground. Such observations were also 
recorded by Lackey (1989), and suggest that O.p. is terrestrial - numerous dens were 
observed in the ground - but also very common in the vegetation composing the forest 
understory. Besides, the fact that O.p. was captured in all sites (Table 1), and in all 
forest habitats (Table 2), leads to infer that it is ubiquitous in the forests of SNR, and 
probably absent from exposed areas covered by vegetation type 7. Furthermore, 
regarding the percentage of total captures recorded for O.p. (Table 1), and the frequent 
observations of predators around the traps, the big-eared climbing rat is certainly 
important in the diets of many sympatric vertebrate carnivores in SNR. 

Peromyscus yucatanicus (Allen & Chapman, 1897) Figs 7, 8 

Yucatan Deer Mouse 

Description. The cranium is elongate and rather slender, with a convex dorsal 
profile (Fig. 7). The rostrum is long and narrow, not concealing the nasolacrimal cap- 
sules in dorsal view. The nasals are long and terminate in a rounded angle anteriorly, 
projecting beyond the premaxillae to conceal the nasal orifice from dorsal view. The 
zygomatic arches are slender, parallel from the dorsal aspect, strongly convergent 
anteriorly, and forming downward a pronounced arc-shaped bend in lateral view. The 
interorbital region is narrow relative to braincase. Supraorbital shelves accentuate the 
dorsolateral margins of the interorbit and transform posteriorly into barely visible low 
temporal ridges extending to inconspicuous lambdoidal ridges. The supraorbital ridges 
are moderately grooved dorsally. Occipital ridges are absent, and the supraoccipital at 
its margin with the interparietal is smooth in dorsal view. The braincase is inflated but 
not globose, with nearly vertical lateral and rear walls. The auditory bullae are 
moderately large and not clearly rounded; their anterior margin lies anteriorly to the 
basisphenoid/basioccipital suture. Both the basisphenoid and the basioccipital are 
moderately wide and elongate. The bony palate is rather broad, without fenestration, 
and does not extend beyond the molar rows posteriorly. The anterior margin of the 



SMALL MAMMALS FROM BELIZE 227 

maxillary /palatines suture is level with the margin between Ml and M2. The incisive 
foramina are long and conspicuous, and their posterior margin is level with the anterior 
margin of the orbits. The teeth number 16. The molar toothrows are only slightly 
convex along their labial margins, and slightly convergent posteriorly. Incisors are 
moderately large with smooth orangish enamel faces; in lateral profile, the uppers 
curve back toward the molar rows, an opisthodont configuration. The dental formula 
is : i 1/1, c 0/0, p 0/0, and m 3/3. The dentary has a falciform coronoid process, a long 
and stout condyloid process, a well developed angular process, and a moderately deep 
and short ramus; the capsular process is barely visible. 

The plantar surface of pes is greyish-brown, devoid of distinctive scalation, thin- 
ly covered with short black hairs on the heel, with little fringing hairs along its contour, 
and with four separate interdigital and two separate carpal pads (Fig. 8). Dorsal and 
lateral surfaces of the metatarsal region and digits are white, thinly covered with short 
white hairs. Claws are short and unpigmented, with a tuft of white hair at their base. . 

Discussion. P.yucatanicus is easily confused with P.mexicanus. Nevertheless, 
P.m. is a typical Central American endemic seemingly absent from the Yucatan Penin- 
sula, Mexico, apart from southern Belize (McCarthy et al., 1998). Whereas P.y. is 
endemic to the Yucatan Peninsula, and its present southern range extends no further 
than in the southern part of the states of Campeche and Quintana Roo, Mexico (Reid, 
1997). Consequently, their respective ranges are not overlapping, and P.y. is considered 
as allopatric from P.m. (Young & Jones, 1983). However, Huckaby (1980) reported that 
the southern limits of the range of P.y. are not clearly known, which is confirmed by 
the present study. Indeed, the presence of P.y. in Shipstern Nature Reserve constitutes 
a first mention for Belize (McCarthy et al., 1998), and suggests that its range is 
extending more southerly to the limit agreed at present. 

On the basis of external characters, P.y. and P.m. are hardly distinguishable; P.y. 
differs in being smaller, both externally and cranially (Young & Jones, 1983). In the 
case of VB225 (MHNG n°1812.010), the external and cranial measurements (Tables 4, 
5) tally perfectly with those of P.y. in Lawlor (1965), Huckaby (1980), and Lackey 
(1976), but are below the ranges given for P.m. in Huckaby (1980), and Goodwin 
(1969). Nevertheless, it is best distinguished from P.m. by the accessory ridges of its 
upper and lower molars that are well-developed and fused with the labial styles 
(Lawlor, 1965; Hooper, 1957). 

Although serious conclusions cannot be drawn from two isolate captures, the 
fact that VB089 and VB225 were captured on the ground and at a height of 2 ft 
respectively, and that no morphological adaptations towards arboreality were observed, 
suggest that it is mainly terrestrial, with a moderate ability to climb on the low vegeta- 
tion composing the understory. Furthermore, the habitat description in Reid (1997) 
leads to infer that P.y. is probably ubiquitous in the forests of Shipstern Nature Reserve, 
although it was only recorded from the Shipstern area (Fig. 2). 



228 



V. BERSOT 






i 1 




SMALL MAMMALS FROM BELIZE 229 

Sigmodon hispidus (Say & Ord, 1825) Figs 7, 8 

Hispid Cotton Rat 

Description. The cranium is short and rather broad, with a moderately convex 
dorsal profile (Fig. 7). The rostrum is short and broad, but not concealing the deeply 
excavated nasolacrimal capsules in dorsal view. The nasals are short and terminate in 
a rounded angle anteriorly, projecting beyond the premaxillae to conceal the nasal 
orifice from dorsal view. The zygomatic arches are stout, convex from the dorsal as- 
pect, and convergent anteriorly. The interorbital region is rather narrow relative to 
braincase. Supraorbital shelves accentuate the dorsolateral margins of the interorbit 
and transform posteriorly into low temporal ridges extending to conspicuous lamb- 
doidal ridges. The supraorbital ridges are grooved dorsally. Occipital ridges are 
present, and in average 3.1 mm distant from temporal ridges. The supraoccipital crest 
is angular in dorsal view, but also when viewed posteriorly. The braincase is only 
slightly inflated, with nearly vertical lateral and rear walls. The auditory bullae are 
moderately large and rounded; their anterior margin lies anteriorly to the basisphe- 
noid/basioccipital suture. The basisphenoid is short, contrasting with a wide and rather 
elongate basioccipital. The diameter of the foramen ovale is less than three-quarters the 
diameter of M3. The bony palate is fairly broad, without fenestration, and with a well- 
developed median crest extending beyond the molar rows posteriorly (spinous 
process). The anterior margin of the maxillary /palatines suture is level with M2. The 
spinous anterodorsal process of the zygomatic plate is blunt, but clearly visible. The 
incisive foramina are long, narrow, conspicuous, and their posterior margin lies slightly 
posteriorly to the anterior margin of the orbits. The teeth number 16. The molar 
toothrows are convex along their labial margins, and slightly convergent anteriorly. 
Incisors are large with ungrooved orangish enamel faces; in lateral profile, the uppers 
curve back toward the molar rows, an opisthodont configuration. The dental formula 
is : i 1/1, c 0/0, p 0/0, and m 3/3. The dentary has a falciform coronoid process, a long 
and stout condyloid process, a broad and well developed angular process, and a deep 
and short ramus; the capsular process is clearly visible. 

The plantar surface of pes is dark grey, with a distinctive scalation, naked from 
heel to tips of the digits, with conspicuous fringing hairs along its contour, and with 
four separate interdigital and two separate carpal pads (Fig. 8). Dorsal and lateral 
surfaces of the metatarsal region and digits are grey, and thinly covered with short 
orangish or silvery hairs. The three middle digits are much longer than the outer two, 
and have enlarged claws. Claws are otherwise greyish, with a conspicuous tuft of 
silvery hair at their base. 

Discussion. The relationships among the seven valid species of the genus 
Sigmodon that are currently recognized in North and Central America have long been 
the subject of controversy, and the genetic integrity of S. hispidus cannot be assumed as 
yet (Elder & Lee, 1985; Voss, 1992; Peppers & Bradley, 2000). Indeed, hispidus as 

Fig. 7 
Top, Ototylomys phyllotis (VB002/MHNG n°l 8 11.090); middle, Peromyscus yucatanicus 
(VB225/MHNG n°l 8 12.010); bottom, Sigmodon hispidus (VB147/MHNG n° 18 12.004). 
Dorsal, ventral, and lateral view of cranium and lateral view of left dentary. Scale bars: 5 mm. 



230 V. BERSOT 

currently recognized may prove to consist of several morphologically cryptic but 
genetically distinct geographical units, and particularly in Central American popu- 
lations that have virtually received no revisionary attention (Voss, 1992). 

Traditionally, the genus Sigmodon is divided into two controversed (Elder & 
Lee, 1985) taxonomic groups : the S.hispidus and the S.fulviventer groups. Apart from 
the members of the S.fulviventer group from which it differs by having a sparsely 
haired tail, with large scales (0.75 mm wide rather than 0.5 mm), S.hispidus could be 
confused with the two other species of the S.hispidus group: S.arizonae and S.mas- 
cotensis. But, whereas S.h. is ubiquitous in Central America - its range extends from 
Northern Venezuela to Central and South-East United States, but it is absent from the 
west coast of Mexico - S.a. and S.m. are confined to the northern and southern part 
respectively of the Mexican west coast (Zimmerman, 1970). As a result, the three 
species are not expected to be sympatric in Belize. 

S.h., S.a., and S.m. are remarkably uniform in external appearance, with vir- 
tually no interspecific variation. S.h. is, nevertheless, distinguishable by having a 
generally shorter hind foot - less than 34 mm, usually averaging 32 mm (Cameron & 
Spencer, 1981; Zimmerman, 1970). With a mean of 26.5 mm, the adult (Voss, 1992) 
specimens examined during the present study are therefore clearly below the average. 
Similarly, the external and cranial measurements of VB146 (MHNG n° 18 12.003), 
VB147 (MHNG n°1812.004), and VB148 (MHNG n°1812.005) are rather at the 
bottom (Tables 4, 5), or below, those given for S.h. in Cameron & Spencer (1981), Voss 
(1992), and Goodwin (1969). However, they better correspond to the measurements 
given in Jones et al. (1974) for specimens that were caught to the west of Chetumal, 
Quintana Roo, Mexico. The latter observations lead to formulate the hypothesis of a 
morphologically smaller S.h. population in the south of Quintana Roo and northern 
Belize, and further underlines the need for a complete revision of the genus in the 
Yucatan Peninsula. 

Several skull characters allowed VB146, VB147, and VB148, to be identified. 
Indeed, the combination of a distance between the temporal and occipital ridges 
inferior to 3.6 mm, a foramen ovale with a diameter inferior to 3/4 the diameter of M3, 
and a well developed crest on the posterior part of the bony palate distinguish them 
clearly from S.a. and S.m. (Zimmerman, 1970). Also, they differ from S.a. by having 
a narrower ventral surface of the presphenoid, a blunt and rather broad spinous 
anterodorsal process of the zygomatic plate, and an angular supraoccipital crest in 
posterior view (Severinghaus & Hoffmeister, 1978). 

S.h. is a habitat specialist, generally found wherever grass is a significant 
component of the local vegetation (Voss, 1992). This is confirmed by the fact that 
specimens of the present study were only captured in vegetation types 6 and 7 of the 
site n°6 (Tables 1, 2); S.h. is therefore expected to be ubiquitous in such habitats, but 
absent from the forests of SNR. Furthermore, all specimens of S.h. were caught on the 
ground, and ran away on the ground when released. Besides, no morphological 
adaptations towards arboreality were observed from sampled specimens, thus 
confirming that 5./?. is strongly terrestrial. 



SMALL MAMMALS FROM BELIZE 



231 







VB054 



VB062 



VB004 



VB182 






VB147 



VB021 



VB225 



Fig. 8 
Marmosa mexicana (VB054), Didelphis virginiana (VB062), Heteromys gaumeri (VB021), 
Otonyctomys hatti (VB182), Ototylomys phyllotis (VB004), Pewmyscus yucatanicus (VB225), 
and Sigmodon hispidus (VB147). Plantar view of left pes. hy, hypothenar pad; th, thenar pad; 
1-4, carpal pads; I-V, digits. Scale bar: 10 mm. 



232 V. BERSOT 



GENERAL DISCUSSION 



Ideally, fieldwork should be continued until asymptotic species lists are 
obtained. This is nearly achieved in the case of several Central American field stations 
with long histories of inventory work, where high diversity - quantified as species 
richness (number of species) - was obtained (Voss & Emmons, 1996); but even in these 
regions, new species were recently added to the overall list. In the vicinity of the La 
Selva Biological Station, nearby the Braulio Carrillo National Park, Costa Rica, up to 
1 14 mammalian species including 5 species of marsupials and 13 species of small ro- 
dents were recorded since the first inventory published by Slud in 1960 (Timm et al., 
1989). Further south in central Panama, up to 113 mammalian species including 6 
species of marsupials and 10 species of small rodents were obtained from Barro Colo- 
rado Island, Gatun Lake, since Enders' surveys from 1929 to 1937 (Eisenberg & 
Thorington, 1973; Voss & Emmons, 1996). In the states of Veracruz and Oaxaca, 
Mexico, two species accounts based on long-term studies were reported by Hall & 
Dalquest (1963), and Goodwin (1969) respectively. In the first case, 156 mammalian 
species including 4 species of marsupials, 48 species of small rodents, and 7 species of 
shrews were recorded since Dalquest 's first account in 1947. And in the state of 
Oaxaca, 204 mammalian species including 6 species of marsupials, 54 species of small 
rodents, and 10 species of shrews were listed since McDougall's surveys from 1943 to 
1967. 

By comparing the present study with the latter inventories, the list of small 
mammals obtained in Shipstern Nature Reserve appears very incomplete with only 2 
species of marsupials, and 5 species of myomorph rodents. However, an expedition of 
six months is considered as very brief, and in such a case adding missing species to the 
inventory is just a matter of time. An essential problem with inventory comparisons is 
highlighted by the latter remark. Indeed, all inventory methods are biased because each 
is suitable for collecting or observing only a fraction of the morphologically and 
behaviorally diverse mammalian fauna that inhabits Neotropical forests, and special 
methods are required to add elusive species to faunal lists (Voss & Emmons, 1996). 
Hence, many methods must be used in combination to census whole communities, and 
virtually all existing inventories are incomplete. Furthermore, the degree of incom- 
pleteness is inversely correlated with inventory duration, so that species lists always 
increase with additional sampling effort up to an asymptote. 

Consequently, results obtained in the present study are better compared to other 
short term inventories. In Belize, Rabinowitz & Nottingham (1989) sampled small 
mammals in the Cockscomb Basin of the Maya Mountains for several months in 1983 
and 1984. Inventory sites such as the latter comprising both riparian and terra firme 
habitats are likely to have more diverse mammalian communities than more arid sites 
such as the one in SNR. This was confirmed by their results, with 4 species of marsu- 
pials, including M.mexicana, and 9 species of myomorph rodents, including O.phyl- 
lotis and S.hispidus. Besides, they recorded that three species - Heteromys desmares- 
tianus, Tylomys nudicaudus, and O.phyllotis - accounted for 67.5% of the total capture. 
With O.phyllotis accounting for 90.8% and H.gaumeri 5.55% of the total capture 
(Table 1), results are even more striking in SNR, and confirm the general trend 



SMALL MAMMALS FROM BELIZE 233 

highlighted by Fleming (1975) that neotropical mammal communities generally 
contain one or two relatively common species and many uncommon species of small 
mammals. Similarly, Disney (1968) reported from his study in central Belize that 
O.phyllotis and H.gaumeri were together the most common species with 76.7% of the 
total capture. Perhaps, these two species are the most agressive of the community in 
northern Belize, and preclude the capture of trap-shy species. 

Recently, another study was completed in the Chiquibul Forest Reserve of the 
Maya mountains (Caro et al., 2001). Their capture results were fairly similar to those 
of Rabinowitz & Nottingham (1989), with 4 species of marsupials including M.mexi- 
cana and D.virginiana, and 7 species of myomorph rodents, including H.gaumeri and 
O.phyllotis. By combining these latter results with those obtained in SNR, it appears 
that at least 4 small mammal species - M.mexicana, D.virginiana, H.gaumeri and 
O.phyllotis - are sympatric throughout Belize. In addition, two species of squirrels - 
Sciurus yucatanensis and S.deppei - were recorded from the Cockscomb Basin, 
Chiquibul Forest Reserve, and SNR by sight (Appendix III), and can also be consid- 
ered as sympatric. However, a main difference in SNR is the presence of species such 
as O.hatti and P.yucatanicus that are endemic to the Yucatan Peninsula (McCarthy et 
al., 1998). In this way, with additional endemics such as H.gaumeri and S.yucata- 
nensis, the small mammal fauna in SNR appears to be closely related to the rodent 
fauna listed by Jones et al. (1974) in Yucatan. 

The overall trapping success (3.5%) realized in this study falls within the range 
of observed success rates in other neotropical small mammal inventories. In the state 
of Minas Gerais, Brazil, Da Fonseca & Kierulff (1989) recorded a trapping success of 
2.4%, and Stallings (1989) 7.5%. In the Maya Mountains, Belize, Rabinowitz & 
Nottingham (1989) recorded a trapping success of 5.23%, and Caro et al. (2001) 0.8%. 
However, the site-to-site trapping success realized in Shipstern Nature Reserve (Table 
3) is fluctuating markedly, ranging from 1.2% in site n°2 to 8.8% in site n°4. Such a 
fluctuation was also reported by Stallings (1989) where the trapping success ranged 
from 2.3% to 18.8% from one site to an other. In the case of SNR, these irregular 
trapping results are unlikely to be due to a temporal fluctuation in density, since both 
sites n°2 and 4 were surveyed during the driest months. Furthermore, the highest re- 
sults (8.8% and 6.6%) were both recorded from sites n°4 and 8, that is in the Shipstern 
area (Fig. 2). The latter observation leads to consider a possible higher density in 
forests situated south to Shipstern Lagoon, at least for O.phyllotis. Indeed, such results 
are biased by the very high predominance of O.phyllotis in sites n°4 and 8 - with a to- 
tal capture of 96.5% and a recapture rate of 2.3, and 95.2% and 1.8, respectively (Table 
1) - and therefore cannot be extrapolated to other species such as H.gaumeri which re- 
mained fairly elusive with a recapture rate of 1.0. 

The results of this small mammal inventory give substance to the impression of 
a diverse rodent community that is clearly affiliated to the partly endemic fauna of the 
Yucatan Peninsula, Mexico. Besides, they demonstrate the important role that O.phyl- 
lotis plays in the community structure of small mammals in SNR. That a single species 
of rodent represents such a high percentage of the total capture, was not recorded in 
any of the studies cited in the text. Thus, the trapping methods used during this inven- 
tory may prove to be particularly effective in trapping O.phyllotis, but this factor alone 



234 V. BERSOT 

is surely not preponderant. Consequently, further study focused on the ecological 
requirements of this species will be of great interest with a view to characterizing the 
overall ecosystem prevailing in SNR. On the other hand, much research at the 
community level will be needed to ascertain the role of O.phyllotis, and other small 
mammals, in their maintenance of predators at higher trophic levels. Finally, by 
re f ering to the very short period in which the collecting took place, and the small mam- 
mal species list predicted from range overlap in appendix II, the sympatric diversity 
concretely obtained in SNR is certainly underestimated. In this way, further sampling 
will be needed to assess the presence of other small mammal species in the reserve, and 
the present inventory is therefore regarded as a preliminary assessment. 

ACKNOWLEDGEMENTS 

This research would not have been possible without the financial support of 
three institutions: Fonds Wiithrich & Mathey-Dupraz, Fondation Hélène & Victor 
Barbour, and Fondation Gertrud Riiegg. Collecting and export permits were provided 
by the Conservation Division of the Forest Department, Ministry of Natural Resources, 
Belmopan, Cayo District, Belize. 

The following people provided unfailing support and advice: Dr Manuel Ruedi, 
Dr Claude Vaucher, Alain De Chambrier, Prof. Peter Vogel, François M. Catzeflis, and 
Prof. Claude Mermod. Special thanks and mention should be given particularly to 
Caspar Bijleveld, Project Coordinator of the ITCF and Director of the Papiliorama 
Swiss Tropical Gardens, Kerzers/Fribourg, Switzerland, for his continuing encoura- 
gement and friendship. At the Shipstern Nature Reserve, I am especially grateful to 
Juan & Argelia Aldana-Mesh, Damian Aldana, Leonardo Balam, Haroldo Mesh & his 
wife, Romei Blanco, and Luis & Theresa Perez for their generous hospitality and 
assistance. Last but not least, thankyou to my friends and colleagues Claude Fischer, 
David Bärtschi, Jean-François Mauffrey, Cynthia Steiner, Alain Merguin, Florence 
Marteau and Bernard Cerroti; Margaret Ducommun who assisted me in the proof- 
reading of the text. My apologies to any other person I have omitted to mention but 
who has nevertheless contributed in any way. 

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238 



V. BERSOT 



Appendix I: Tables 1-6 

Table 1. Capture results. Field numbers in brackets correspond to the specimens captured at 
each site. See text for discussion. 



Siten°l (VB001-VB016) 


18 c 


> 18'493"N/88°11'182' 


•\v 








Species 




Total 
captures 


% Total 




First 
captures 


% Total 


Recapture 
index 


Ototylomys phyllotis 




27 


100.0 




16 


100.0 


1.7 


Total : 




27 


100.0 




16 


100.0 


- 


Site n°2 (VB017-VB031) 


18 c 


>W l\9"N l^n'oW 


'W 








Species 




Total 
captures 


% Total 




First 
captures 


% Total 


Recapture 
index 


Heteromys gaumeri 
Ototylomys phyllotis 




6 

13 


31.6 
68.4 




4 
11 


26.7 
73.3 


1.5 
1.2 


Total : 




19 


100.0 




15 


100.0 


- 


Site n°3 (VB032-VB058) 


18 c 


T5'004"N/88°15'002' 


'W 








Species 




Total 
captures 


% Total 




First 
captures 


% Total 


Recapture 
index 


Marmosa mexicana 
Heteromys gaumeri 
Ototylomys phyllotis 




1 
2 
44 


2.1 
4.2 
93.7 




1 
1 
25 


3.7 
3.7 
92.6 


1.0 
2.0 
1.8 


Total : 




47 


100.0 




27 


100.0 


- 


Site n°4 (VB059-VB123) 


18 c 


Tl'890"N/88°ir209' 


'W 








Species 




Total 
captures 


% Total 




First 
captures 


% Total 


Recapture 
index 


Didelphis virginiana 
Heteromys gaumeri 
Ototylomys phyllotis 
Peromyscus yucatanicus 




1 

3 
136 

1 


0.7 
2.1 
96.5 
0.7 




1 
3 

60 
1 


1.5 
4.6 
92.4 
1.5 


1.0 
1.0 
2.3 
1.0 


Total : 




141 


100.0 




65 


100.0 


- 


Site n°5 (VB124-VB144) 


18 c 


17 , 567 , 'N/88°12'912' 


'W 








Species 




Total 
captures 


% Total 




First 

captures 


% Total 


Recapture 
index 


Heteromys gaumeri 
Ototylomys phyllotis 




7 
29 


19.4 
80.6 




6 

15 


28.6 

71.4 


1.2 
1.9 


Total : 




36 


100.0 




21 


100.0 


- 



SMALL MAMMALS FROM BELIZE 



239 



Site n°6 (VB145-VB1 72) 18°18'159"N / 88°12'046"W 



Species 

Hetewmys gaumeri 
Ototylomys phyllotis 
Sigmodon hispidus 

Total : 



Total 
captures 

3 

36 

10 

49 



% Total 

6.1 

73.5 
20.4 

100.0 



First 
captures 

3 
18 

7 

28 



% Total 

10.7 
64.3 
25.0 

100.0 



Sites n°l-8 
Species 

Marmosa mexicana 
Didelphis virginiana 
Hetewmys gaumeri 
Otonyctomys hatti 
Ototylomys phyllotis 
Pewmyscus yucatanicus 
Sigmodon hispidus 

Total : 



Recapture 
index 

1.0 
2.0 
1.4 



Site n°7 (VB 173 -VB207) 


18 c 


> 15'760' , N/88°15 , 998' 


'W 






Species 




Total 
captures 


% Total 


First 
captures 


% Total 


Recapture 
index 


Hetewmys gaumeri 
Otonyctomys hatti 
Ototylomys phyllotis 




1 
1 

51 


1.9 
1.9 
96.2 


1 
1 

33 


2.9 
2.9 
94.2 


1.0 
1.0 
1.5 


Total : 




53 


100.0 


35 


100.0 


- 


Site n°8 (VB208-VB231) 


18 c 


'13'638"N/88°10'879' 


'W 






Species 




Total 
captures 


% Total 


First 
captures 


% Total 


Recapture 
index 


Hetewmys gaumeri 
Ototylomys phyllotis 
Pewmyscus yucatanicus 




1 

40 

1 


2.4 
95.2 

2.4 


1 

22 
1 


4.2 
91.6 

4.2 


1.0 
1.8 
1.0 


Total : 




42 


100.0 


24 


100.0 


- 



Total 


% Total 


First 


% Total 


Recapture 


captures 




captures 




index 


1 


0.25 


1 


0.45 


1.0 


1 


0.25 


1 


0.45 


1.0 


23 


5.55 


19 


8.15 


1.2 


1 


0.25 


1 


0.45 


1.0 


376 


90.8 


200 


86.6 


1.8 


2 


0.5 


2 


0.9 


1.0 


10 


2.4 


7 


3.0 


1.4 



414 



100.0 



231 



100.0 



240 



V. BERSOT 



Table 2. Capture results per vegetation type. Vegetation types, see table 6. 



Sites n° 1-8 



Species 



Vegetation types 

Total 12 3 4 5 6 7 8 

captures P L Pu Ti Mu Ta 

1 i .......... 

1 ....... i ... . 

23 6 7 3 - - - - 6 - - - 1 

1 i . .......... 

376 233 24 27 7 - - 33 12 6 3 - 31 

2 1 . ......... i 

10 ......... 9 i . 

414 242 31 30 7 - - 33 19 6 12 1 33 



Marmosa mexicana 
Didelphis virginiana 
Heteromys gaumeri 
Otonyctomys hatti 
Ototylomys phyllotis 
Peromyscus yucatanicus 
Sigmodon hispidus 



Total 



Table 3. Trapping success. See text for discussion. 



Site n° 


Trapping periods 


No. trap nights 


No. captures 


% success 


1 


May 1-11 


1600 


27 


1.7 


2 


May 19-29 


1600 


19 


1.2 


3 


June 6-16 


1600 


47 


2.9 


4 


June 21- July 1 


1600 


141 


8.8 


5 


July 11-21 


1600 


36 


2.3 


6 


Aug 28-Sept 7 


1600 


49 


3.1 


7 


Sept 11-21 


1600 


53 


3.3 


8 


Sept 26-30 


640 


42 


6.6 



Total : 



11840 



414 



3.5 



Table 4. External measurements. Body (mm) and mass (g) measurements are presented for 
adults of both sexes, n, sample size. Other abbreviations, mean and standard error, are defined in 
text. 











Mean (n= 


=D 






Variable 


M. 


mexicana 


D .virginiana 





. hatti 


P. yucatanicus 


TL 




323 


389 






199 


191 


TV 




174 


186 






108 


92 


HF 




20 


33 






19 


22 


E 




22 


35 






13 


19 


NEB 




35 


58 






26 


28 


NEC 




16 


26 






13 


15 


SR 




35 


- 






22 


26 


Weight 




66 


170 






24 


24 



SMALL MAMMALS FROM BELIZE 



241 



Heteromys gaumeri 



Ototylomys phyllotis 



Variable 


Mean ± SE 


Range 


n 


Mean 1 SE 


Range 


n 


TL 


266.6 ± 5.58 


252-286 


5 


263.87 1 1.64 


227-295 


91 


TV 


138.413.98 


128-147 


5 


123.761 1.09 


97-150 


94 


HF 


31.13 ±0.9 


25-33 


8 


23.55 10.1 


21-29 


95 


E 


16.88 10.3 


16-18 


8 


22.210.12 


19-26 


92 


NEB 


33.13 10.3 


32-34 


8 


35.910.19 


29-41 


93 


NEC 


18.88 10.77 


17-24 


8 


17.05 10.14 


11-20 


93 


SR 


9.4 1 0.24 


9-10 


5 


7.83 1 0.09 


6-10 


94 


Weight 


60.22 1 3.54 
Sigmodon 


50-75 
hispidus 


9 


65.461 1.15 


43-96 


102 


Variable 


Mean 1 SE 


Range 


n 





TL 

TV 

HF 

E 

NEB 

NEC 

SR 

Weight 



236.2 1 6.3 


220-253 


5 


97.2 1 1.93 


94-104 


5 


26.5 1 0.62 


24-28 


6 


17.810.58 


16-19 


5 


34.01 1.03 


31-37 


6 


16.17 10.7 


14-18 


6 


15.010.0 


15-15 


5 


84.67 1 3.68 


70-92 


6 



Table 5. Cranial measurements (mm), presented for adults of both sexes, n, sample size. Other 
abbreviations, mean and standard error, are defined in text. 







Mean (i 


n=l) 


Variable 


Mean (n=l) 


Variable 


M.mexicana 


D .virginiana 


M.mexicana 


D .virginiana 


ONL 


35.41 




56.99 


MB 


12.68 


18.77 


CBL 


34.42 




55.6 


OCB 


7.11 


12.42 


NL 


15.27 




26.24 


BB 


12.65 


19.21 


BN 


4.71 




8.85 


ZB 


20.02 


29.4 


PL 


18.96 




34.24 


IB 


6.1 


11.33 


PW 


9.53 




17.91 


IOC2 


5.97 


9.39 


PPL 


13.41 




16.77 


CD 


11.43 


16.66 


RL 


12.05 




23.19 


C-M4 


13.38 


20.03 


RW 


5.59 




11.57 


M1-M4 


6.38 


12.96 


BRJ 


10.38 




16.06 


CLMT 


3.29 


14.36 


BRF 


6.65 




9.91 


LM 


24.8 


42.31 


BOL 


4.68 




5.92 












Heteromys gaumeri 




Mean (n 


= D 


Variable 


Mean 1 SE 


Range 


n 


0. hatti 


P. yucatanicus 


ONL 


35.46 1 0.25 


34.97-35.75 


3 


28.67 


28.8 


CBL 


33.27 1 


0.19 


32.95-33.62 


3 


25.77 


26.86 


CIL 


29.4910.1 


29.34-29.67 


3 


24.81 


25.49 


NL 


14.44 1 


0.34 


13.85-15.01 


3 


9.21 


11.09 



242 




V. BERSOT 






PL 


20.28 ± 0.45 


19.44-21.0 


3 


12.4 


14.42 


PPL 


10.51 ±0.23 


10.12-10.93 


3 


11.11 


10.56 


LD 


9.22 ± 0.36 


8.51-9.63 


3 


6.75 


7.98 


RL 


15.56 ±0.18 


15.27-15.88 


3 


9.63 


11.1 


BR 


6.33 ± 0.23 


6.07-6.79 


3 


4.79 


5.05 


LIF 


2.87 ± 0.24 


2.62-3.34 


3 


4.75 


5.86 


BIF 


0.91 ±0.04 


0.84-0.96 


3 


2.38 


2.49 


BZP 


4.36 ± 0.06 


4.29-4.49 


3 


1.46 


2.04 


OL 


8.81 ±0.05 


8.71-8.89 


3 


8.76 


8.92 


LBP 


8.0 ± 0.25 


7.57-8.42 


3 


3.88 


4.09 


BBP 


5.34 ±0.11 


5.2-5.56 


3 


4.95 


5.18 


MPFL 


4.33 ±0.17 


4.06-4.64 


3 


5.03 


4.88 


MPFW 


2.12 ±0.12 


1.98-2.35 


3 


2.11 


1.75 


BOL 


4.95 ±0.1 


4.79-5.14 


3 


4.51 


3.78 


BL 


4.87 ± 0.07 


4.75-5.0 


3 


7.84 


4.19 


BW 


3.54 ± 0.08 


3.45-3.69 


3 


5.54 


3.38 


MB 


14.81 ±0.27 


14.43-15.32 


3 


12.79 


11.55 


OCB 


6.88 ± 0.22 


6.45-7.21 


3 


6.15 


6.53 


LB 


13.52 ±0.33 


13.13-14.17 


3 


10.66 


11.59 


BB 


13.51 ±0.17 


13.3-13.85 


3 


13.1 


11.87 


ZB 


16.41 ±0.15 


16.19-16.7 


3 


14.29 


13.04 


IB 


8.82 ± 0.06 


8.7-8.88 


3 


4.79 


4.67 


CD 


11.56 ± 0.15 


11.32-11.84 


3 


11.58 


9.55 


BM1 


1.35 ±0.05 


1.3-1.45 


3 


1.28 


1.28 


CLM1-3 


4.6 ±0.19 


4.32-4.97 


3 


3.99 


4.22 


DI 


1.45 ±0.05 


1.37-1.55 


3 


1.33 


1.37 


HI 


5.29 ±0.15 


5.06-5.57 


3 


3.57 


3.64 


CLMT 


4.79 ± 0.06 


4.66-4.87 


3 


4.16 


4.29 


LM 


15.01 ±0.14 


14.79-15.28 


3 


13.22 


14.31 



Ototylomys phyllotis 



Sigmodon hispidus 



Variable 


Mean ± SE 


Range 


n 


Mean ± SE 


Range 


n 


ONL 


36.81 ±0.61 


33.86-38.86 


8 


32.82 ± 0.78 


31.59-34.26 


3 


CBL 


34.37 ± 0.67 


30.94-36.84 


8 


30.94 ± 0.74 


29.64-32.19 


3 


CIL 


32.08 ± 0.55 


29.43-34.18 


8 


30.55 ± 0.68 


29.3-31.65 


3 


NL 


12.48 ±0.29 


11.55-13.65 


8 


11.81 ±0.37 


11.07-12.25 


3 


PL 


18.15 ±0.32 


16.44-19.05 


8 


17.34 ±0.43 


16.56-18.05 


3 


PPL 


13.74 ±0.36 


12.27-15.42 


8 


11.39 ±0.34 


10.82-11.98 


3 


LD 


9.02 ±0.14 


8.42-9.55 


8 


9.19 ±0.36 


8.49-9.64 


3 


RL 


12.96 ±0.31 


11.73-13.93 


8 


11.81 ±0.46 


10.9-12.32 


3 


BR 


5.87 ±0.12 


5.36-6.26 


8 


6.53 ± 0.2 


6.21-6.9 


3 


LIF 


7.15 ± 0.12 


6.4-7.55 


8 


7.75 ±0.19 


7.41-8.05 


3 


BIF 


3.28 ± 0.09 


2.97-3.68 


8 


2.44 ±0.14 


2.21-2.69 


3 


BZP 


2.93 ± 0.09 


2.69-3.46 


8 


3.03 ±0.1 


2.86-3.22 


3 


OL 


10.79 ±0.11 


10.33-11.13 


8 


10.44 ± 0.33 


10.1-11.11 


3 


LBP 


4.62 ± 0.07 


4.29-4.82 


8 


5.87 ±0.11 


5.76-6.08 


3 


BBP 


6.2 ±0.11 


5.84-6.67 


8 


6.67 ± 0.29 


6.14-7.15 


3 


MPFL 


6.3 ± 0.2 


5.65-7.37 


8 


4.66 ± 0.26 


4.4-5.18 


3 


MPFW 


2.47 ± 0.04 


2.29-2.58 


8 


2.07 ±0.13 


1.82-2.26 


3 


BOL 


5.16 ±0.14 


4.6-5.89 


8 


5.06 ±0.1 


4.94-5.27 


3 


BL 


6.33 ± 0.09 


5.81-6.64 


8 


5.86 ± 0.07 


5.73-5.95 


3 


BW 


4.47 ±0.12 


4.04-4.98 


8 


4.16 ±0.01 


4.14-4.18 


3 


MB 


13.54 ±0.14 


12.82-14.05 


8 


12.61 ±0.31 


12.1-13.16 


3 


OCB 


7.51 ±0.14 


6.83-7.93 


8 


6.7 ±0.16 


6.44-7.0 


3 



SMALL MAMMALS FROM BELIZE 



243 



LB 


11.93 10.24 


10.76-12.83 


8 


11.89 ±0.44 


11.25-12.73 


3 


BB 


14.33 ±0.18 


13.49-15.13 


8 


13.14 ±0.2 


12.86-13.53 


3 


ZB 


17.82 + 0.28 


16.7-18.78 


8 


18.11 ±0.59 


16.96-18.87 


3 


IB 


5.84 ±0.16 


5.13-6.67 


8 


4.88 ±0.11 


4.72-5.09 


3 


CD 


11.44 ±0.07 


11.22-11.85 


8 


12.16 ±0.25 


11.65-12.43 


3 


BM1 


1.67 ±0.01 


1.61-1.71 


8 


1.74 ±0.04 


1.68-1.81 


3 


CLM1-3 


6.0 ± 0.06 


5.68-6.15 


8 


5.08 ± 0.07 


4.99-5.23 


3 


DI 


1.65 ±0.05 


1.4-1.81 


8 


1.89 ±0.02 


1.86-1.92 


3 


HI 


3.79 ±0.16 


3.11-4.46 


8 


5.73 ±0.14 


5.48-5.95 


3 


CLMT 


5.96 ± 0.05 


5.72-6.08 


7 


5.89 ±0.17 


5.64-6.21 


3 


LM 


17.77 ± 0.3 


16.73-18.77 


7 


17.22 ± 0.65 


16.03-18.26 


3 



Table 6. Vegetation types 1 to 8. The New Trail, Main Trail, Xo-Pol, and Shipstern areas 
correspond to trapping sites 1/6, 2/5, 3/7, and 4/8 respectively (Fig. 2). 

Vegetation type n° Description 

Yucatecan 1 This forest type (30-50 ft) covers most of the land situated to the west 

medium-sized of the reserve, including the Xo-Pol area, as well as the land situated 

semi-evergreen to the south of Shipstern Lagoon, including the Shipstern area. But, 

forest whereas it alternates with extensive Cohune forests in the latter, such 

is not the case in the Xo-Pol area where Cohune trees are virtually ab- 
sent. Dominant species composing its canopy are: Brosimum ali- 
castrum, Hampea trilobata, Coccoloba reflexiflora, Bursera sima- ru- 
ba, Metopium brownei, and Manilkara zapota (emergent tree). 
Dominant understory species are: Pithecellobium stevensonii, Cocco- 
loba schiedeana, Ouratea lucens, Randia aculeata, and Croton 
reflexifolius. Beside Brosimum alicastrum, differential species for this 
vegetation type are: Dendropanax arboreus, Nectandra salicifolia, 
Pouteria campechiana, Protium copal, Quararibea funebris, Sabal 
yapa, Swartzia cubensis, Tetrapteris schiedeana, Talisia olivaeformis, 
and Nectandra coriacea. An additional differential species, Cryo- 
sophila stauracantha, is found in the Xo-Pol area. 

Yucatecan 2 This forest type (30-45 ft) covers part of the land situated north of the 

medium-sized reserve, where it alternates with Low semi-deciduous Pseudophoenix 

semi-deciduous s.sargentii forests. In the Main Trail area, it is transitioning to vege- 

forest tation type 1. Dominant species composing its canopy and its under- 

story are similar to the Yucatecan medium-sized semi-evergreen 
forests, therefore it is very difficult to differentiate between the two. 
Besides, no differential species for it have been clearly brought out 
yet. Hence, it is mainly characterized by the absence of species such 
as Brosimum alicastrum, or Pseudophoenix s.sargentii that are distin- 
guishing vegetation types 1 and 3 respectively. 

Low 3 This forest type (20-40 ft) covers part of the land situated north of the 

semi-deciduous reserve, including the New Trail area, where it alternates with Yuca- 

Pseudophoenix tecan medium-sized semi-deciduous forests. Some patches of it - 

s.sargentii forest coinciding with previously disturbed areas, after the passage of hurri- 

cane Janet in 1955 - are dominated by Lysiloma latisiliquum, and can 
be considered as a structural variant called Lysiloma-dommated 
Pseudophoenix s.sargentii forest. In the latter, the canopy is almost 
exclusively composed of L. latisiliquum, which will be gradually 
replaced by the usual canopy species yet confined to the understory. 
Otherwise, dominant species composing its canopy and its understory 
are fairly the same as in vegetation types 1 and 2, with commonly 



244 



V. BERSOT 



Low 

periodically 
inundated forest 



Bravaisia 
tubiflora- 
dominated 
transitional zone 



Cladium 

jamaicense- 

dominated 

herbaceous 

wetlands 



Discontinuous 
dwarf mangal 
interrupted by 
bare salt fiats 



associated species such as Agave augusti/olia and Diphysa carthag- 
nensis. Differential species from this forest type are: Pseudophoenix 
s.sargentii, Ceiba aesculifolia, Coccothrinax argentata, Erythroxylum 
rotundifolium, Gymnopodium floribundum, Hintonia octomera, Lysi- 
loma latisiliquum, Neomillspaughia emarginata, Plumeria obtusa, 
Eugenia buxifolia, Randia truncata, and Beaucarnea ameliae. 

This heterogeneous type of low forest (10-20 ft) is represented by four 
closely related vegetation types called Pucteal, Tintai, Mucal and 
Tasistal. Often growing in a complicated mosaic, locally called " Bajo 
forest ", these plant communities are patchily scattered all over the 
reserve, and only part of them have yet been localized. Whereas the 
elevation and duration of inundation are gradually decreasing from the 
Mucal to the Pucteal, the Tasistal forms an open habitat (Tasiste 
marsh) close to being permanently inundated. It is dominated and 
differentiated by the palm Accoelorraphe wrightii, which is typically 
found with Erythroxylum areolatum, Eugenia acapulcensis, and 
Jacquinia aurantiaca, along with occasional species such as 
Crescentia cujete, Borrelia verticillata, Eustoma exaltatum, and Cla- 
dium jamaicense. The Mucal forms a shrub forest (Muc marsh) domi- 
nated by Dalbergia glabra, which is typically found with species such 
as Cameraria latifolia and Malpighia lundellii. The Tintai forms a 
semi-open habitat (Tinta marsh) dominated by Haematoxylon cam- 
pechianum and Byrsonima bucidaefolia. The Pucteal forms a low 
forest dominated by Bucida buceras. 

This type of low forest (10-25 ft) is mainly found in the surroundings 
of Shipstern Lagoon, where true forest is transitioning to open wet- 
lands - in most cases, Cladium jamaicense-dominated herbaceous 
wetlands. Such transitional zones are inundated at the peak of the 
rainy season, whereas their soils remain water-saturated for most of 
the year. However, it can also be present in forest areas bordering 
more permanent bodies of water. It is dominated and differentiated by 
Bravaisia tubiflora, which is typically found with Croton reflexifolius, 
Saba! yapa, Metopium brownei, Eugenia rhombea, and Eugenia aca- 
pulcensis, along with occasional species such as Ouratea nitida, 
Cameraria latifolia, and Dalbergia glabra. 

This type of wetland (3-5 ft) is found in the surroundings of Shipstern 
Lagoon, where it forms a transitional belt between true forest and 
mangal flats. Often preceded by a Bravaisia tubiflora-domindXzd tran- 
sitional zone, the belt varies in width from nearly nothing to about 350 
ft (100m). It is dominated and differentiated by Cladium jamaicense, 
which is normally found with species such as Conocarpus erectus and 
Solanum blodgettii. Assimilated to this vegetation type, an omni- 
present variant dominated by Distichlis spicata forms another transi- 
tional belt between the Cladium jamaicense-dominated belt and the 
mangal flats. 

This type of mangal association (3-6 ft) covers most of the periodi- 
cally inundated surface bordering Shipstern Lagoon, and extending to 
areas of transition with true forest. Also found along the coast, it 
represents the dominant vegetation type within the reserve. It is fairly 
intermittent, with large patches of ground being completely bare of 
vegetation, sometimes interrupted by so-called "forest islands" - 
punctual change in ecological conditions allowing various vegetation 
types to establish, including Thrinax /W/ata-dominated low forests 
(not considered as a distinct vegetation type). It is dominated by the 
red mangrove (Rhizophora mangle), which constitutes well-developed 



SMALL MAMMALS FROM BELIZE 245 



communities along the lagoon, but is otherwise found in patches of its 
dwarf counterpart. Typical associated species are the white 
{Laguncularia racemosa) and the black (Avicennia germinans) man- 
groves, only found in dwarf form. Occasionally, species such as 
Distichlis spicata, Fimbristylis spadicea, Batis maritima, and Sali- 
cornia perennis are also found in this vegetation type. 

Cohune forest 8 With the exception of a few small patches localized in the Xo-Pol 

area, this forest type (50-70 ft) is exclusively found in the deep soils 
of the land situated south of the lagoon, including the Shipstern area. 
It alternates with Yucatecan medium-sized semi-evergreen forests, 
and patches of Low periodically inundated forests. The dominant and 
differential species composing the canopy is the cohune palm: 
Orbignya cohune. Associated canopy and understory species are 
otherwise similar to vegetation type 1 . 



Appendix II: Sympatric diversity in SNR 

This appendix presents a diversity estimate for the small mammal fauna of Shipstern 
Nature Reserve. The following list does not include the sampled and sighted species : 

Gray Four-eyed Opossum Philander opossum 

Central American Woolly Opossum Caluwmys derbianus 

Robinson's Mouse Opossum Marmosa robinsoni 

Hispid Pocket Gopher Orthogeomys hispidus 

Forest Spiny Pocket Mouse Heteromys desmarestianus 

Coue's Rice Rat Oryzomys couesi 

Rusty Rice Rat Oryzomys rostratus 

Northern Pygmy Rice Rat Oligoryzomys fulve scens 

Northern Climbing Rat Tylomys nudicaudus 

Slender Harvest Mouse Reithrodontomys gracilis 

Maya Small-eared Shrew Ci-yptotis mayensis 

This small mammal species list was predicted from range overlap - geographic expecta- 
tions (Voss & Emmons, 1996) - and is mainly based on distributional data obtained from Reid 
(1997), Emmons & Feer (1997), and McCarthy et al. (1998). The known ecological requirements 
for each species was also taken into account, so as to minimize inaccuracy in diversity estimate. 
Thus, the Water Opossum (Chironectes minimus) is probably not present in the Shipstern Nature 
Reserve - it is semi-aquatic, favoring fast-flowing, rock- or gravel-bottomed streams in hilly 
country (Reid, 1997) - although its range is seemingly overlapping the area. Inversely, Marmosa 
robinsoni is presently considered as absent from northern Belize, but as already discussed, a 
reappraisal of all specimens of the species in Central America is needed to determine its exact 
distribution, so that it may well be present throughout Central America. Being a habitat general- 
ist, M. robinsoni would therefore be expected in SNR. In the case of the Maya Small-eared Shrew 
{Ci-yptotis mayensis), information on its known geographic and ecological distribution is com- 
plemented by the observation of a non-identified species of shrew in the vicinity of SNR (Walker. 
2000; pers. obs.). 

By combining the present list (1 1 species) with the 7 sampled species of marsupials and 
myomorph rodents, and 3 sighted species of marsupial and squirrels - Didelphis marsupialis, 
Sciurus yucatanensis, and S.deppei - a total of 21 small mammal species is expected in SNR. 
This diversity estimate, however, is subject to several sources of uncertainty, and is to be 
considered as a first approximation. 

Appendix III : Observed mammals in SNR 

This appendix presents an inventory of the mammalian species observed during the field- 
work in Shipstern Nature Reserve. The observations were primarily based on direct sightings 



246 



V. BERSOT 



(D), but also on indirect indices - nests (N), sets of tracks (T), scats (Sc), and sounds (So). A 
total of 19 small, medium-sized, and large mammals was thus obtained from SNR and its direct 
surroundings. In the following list, the numbers are related to the areas - New Trail (1), Main 
Trail (2), Xo-Pol (3), and Shipstern (4) areas (Fig. 2) - from which the corresponding species 
were recorded : 



Common Opossum 


Didelphis marsupiali s 


D/So 


3 


Northern Tamandua 


Tamandua mexicana 


D/So 


4 


Nine-banded Armadillo 


Dasypus novemcinctus 


D 


2 


Yucatan Squirrel 


Sciurus yucatanensis 


D 


3 


Deppe's Squirrel 


Sciurus deppei 


D 


3 


Mexican Porcupine 


Coendou mexicanus 


D 


1 


Central American Agouti 


Dasyprocta punctata 


D 


1,2,4 


Paca 


Agouti paca 


D/So 


1 


Gray Fox 


Urocyon cinereoargenteus 


D/So 


1,2,3 


White-nosed Coati 


Nasua natica 


D/N 


1,3,4 


Kinkajou 


Potos flavus 


D 


4 


Tayra 


Eira barbara 


D 


3 


Puma 


Puma concolor 


T 


3 


Jaguar 


Panthern onca 


T/Sc/So 


2,3 


Ocelot 


Leopardus pardalis 


T/So 


4 


Collared Peccary 


Tayassu tajacu 


D/So 


1 


Baird's Tapir 


Tapirus bairdii 


T 


2,3 


Red Brocket 


Mazama americana 


D/T/So 


3 


White-tailed Deer 


Odocoileus virginianus 


D/T/So 


3 



REVUE SUISSE DE ZOOLOGIE 

Tome 110 — Fascicule 1 



Pages 

Zompro, Oliver & Brock, Paul D. Catalogue of type-material of stick- 
insects housed in the Muséum d'histoire naturelle, Geneva, with 
descriptions of some new taxa (Insecta: Phasmatodea) 3-43 

Chatzaki, Maria, Thaler, Konrad & Mylonas, Moysis. Ground spiders 
(Gnaphosidae; Araneae) from Crete and adjacent areas of Greece. 
Taxonomy and distribution. Ill: Zelotes and allied genera 45-89 

Peng, Xian-Jin, Li, Shu-Qiang & Rollard, Christine. A review of the 
Chinese jumping spiders studied by Dr E. Schenkel (Araneae: 
Salticidae) 91-109 

Schatz, Heinrich. New Sphaewchthonius species from the Neotropical 

region (Acari: Oribatida) 1 1 1-124 

Magnano, Luigi. Una nuova specie di Otiorhynchus Germar, 1822 del 

Pakistan (Coleoptera, Curculionidae, Polydrusinae) 125-128 

Stourac, Petr. Two new species of the genus Quedius from Algeria and 

Pakistan (Coleoptera: Staphylinidae: Staphylinini: Quediina) 129-132 

Haupt, Joachim. Zoogeography in southern Japan as revealed by ground- 
living arachnids 133-139 

Lourenço, Wilson R. New taxonomic considerations on some species of 
the genus Grosphus Simon, with description of a new species (Scor- 
piones, Buthidae) 141-154 

Morati, Jeannine. Catalogue des types de Lamiinae (Coleoptera, Ceram- 

bycidae) conservés au Muséum d'histoire naturelle, Genève 155-206 

Bersot, Vincent. Small mammal inventory in the Shipstern Nature Reserve 

(Corozal District, Belize, Central America), a preliminary assessment 207-246 



REVUE SUISSE DE ZOOLOGIE 
Volume 110 — Number 1 



Pages 

Zompro, Oliver & Brock, Paul D. Catalogue of type-material of stick- 
insects housed in the Muséum d'histoire naturelle, Geneva, with 
descriptions of some new taxa (Insecta: Phasmatodea) 3-43 

Chatzaki, Maria, Thaler, Konrad & Mylonas, Moysis. Ground spiders 
(Gnaphosidae; Araneae) from Crete and adjacent areas of Greece. 
Taxonomy and distribution. Ill: Zelotes and allied genera 45-89 

Peng, Xian-Jin, Li, Shu-Qiang & Rollard, Christine. A review of the 
Chinese jumping spiders studied by Dr E. Schenkel (Araneae: 
Salticidae) 91-109 

Schatz, Heinrich. New Sphaerochthonius species from the Neotropical 

region (Acari: Oribatida) 11 1-124 

Magnano, Luigi. A new species of Otiorhynchus Germar, 1822 from 

Pakistan (Coleoptera, Curculionidae, Polydrusinae) 125-128 

Stourac, Petr. Two new species of the genus Quedius from Algeria and 

Pakistan (Coleoptera: Staphylinidae: Staphylinini: Quediina) 129-132 

Haupt, Joachim. Zoogeography in southern Japan as revealed by ground- 
living arachnids 133-139 

Lourenço, Wilson R. New taxonomic considerations on some species of 
the genus Grosphus Simon, with description of a new species (Scor- 
piones, Buthidae) 141-154 

Morati, Jeannine. A catalogue of the types of Lamiinae (Coleoptera, 

Cerambycidae) housed in the Muséum d'histoire naturelle, Geneva . . 155-206 

Bersot, Vincent. Small mammal inventory in the Shipstern Nature Reserve 

(Corozal District, Belize, Central America), a preliminary assessment 207-246 

Indexed in Current Contents, Science Citation Index 



Volume 1 10 - Number 1 - 2003 

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aim to communicate ideas and information clearly and concisely. Authors not writing in their native language should 
pay particular attention to the linguistic quality of the text. 

Manuscripts must be typed, or printed (high quality printing, if possible by a laser-printer), on one side only 
and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins 
must be at least 25 mm wide. Authors must submit one original and two copies, including tables and figures, in 
final fully corrected form, and are expected to retain another copy. 

We encourage authors to submit the text on a diskette (3,5", Macintosh or IBM compatible, with "Microsoft 
word" or similar programmes). The text should be in roman (standard) type face throughout, including headings, 
except genus and species names which should be formatted in italics (or underlined with pencil); bold, small 
capitals, large capitals and other type faces should not be used. Footnotes and cross-references by page should be 
avoided. 

Papers should conform to the following general layout: 

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, 
name(s) in full and surname(s) of author(s), and full address(es). 

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should 
summarise the contents and conclusions of the paper. The abstract is followed by less than 10 key-words, 
separated by hyphens, which are suitable for indexing. 

Introduction. A short introduction to the background and the reasons for the work. 

Materials and methods. Sufficient experimental details must be given to enable other workers to repeat the 
work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen- 
clature must be strictly followed. Cite the authors of species on their first mention. 

Results. These should be concise and should not include methods or discussion. Text, tables and figures should 
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp. 
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must 
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by 
synonyms, material examined and distribution, description and comments. All material examined should be listed in 
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex 
symbols should be used rather than "male" and "female". 

Discussion. This should not be excessive and should not repeat results nor contain new information, but 
should emphasize the significance and relevance of the results reported. 

References. The Harvard System must be used for the citation of references in the text, e.g. White & Green 
(1995) or (White & Green, 1995). For references with three and more authors the form Brown et al. should be 
used. Authors' names should not be written in capitals. The list of references must include all publications cited in 
the text but only these. References must be listed in alphabetical order of authors, and both the title and name of 
the journal must be given in full in the following style (italics can be formatted by the author): 
Penard, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. 
Penard, E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61. 
Mertens. R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. 
Handley. C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel R. L. & Tipton, V. J. (eds). 

Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. 

References should not be interspaced and, in the case of several papers by the same author, the name has to be 
repeated for each reference. 

Tables. These should be self-explanatory, with the title at the top organised to fit 122 x 180 mm. Each table 
should by typed, double spaced, on a separate page and numbered consecutively and its position indicated in the text. 

Figures. These may be line drawings or half tones and all should be numbered consecutively, and their position 
indicated in the text. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and 
lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors 
should refrain from mixing the drawings and half tones. Original drawings will not be returned automatically. The 
Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If electronically 
scanned figures are submitted on diskettes, this should be clearly indicated on the print-out enclosed with the 
manuscript. 

Legends to figures. These should be typed in numerical order on a separate sheet. 

Proofs. Page proofs only are supplied, and authors may be charged for alterations (other than printer's errors) 
if they are numerous. 

Offprints. The authors receive totally 25 offprints free of charge; more copies may be ordered at current prices 
when proofs are returned. 

Correspondence. All correspondence should be addressed to 
Revue suisse de Zoologie 
Muséum d'histoire naturelle 
CP 6434 

CH-1211 Genève 6 
Switzerland. 
Phone: +41 22 418 63 33 - Fax +41 22 418 63 01 

e-mail: volker.mahnert@mhn,ville-ge.ch 

Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



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