MAN REDET RE a Nahe CIRO rege Na A 16 Au N re tet mg MEAN ea w Kr BASE ORNE un FETE UV CDE PAE and Nota had ela Meehan ed et: MORE PARTE ARAN NI ER EN TEEN N ome da Ai deat Aga He NIENTE DORE O TRPP eni TEMI SNARE va USWA a ss sirena Via SU 6 SRO ERE A AA a RL PCR N SA MN LG BB TA ra HEPAT OA ANIA LAT AU SAR PENARE nen NBA STERN Teta Pr a a Mamata ede Roe eho uve vi SREB edita RS HAAS OF PES OSIO LEN a a Wet ACA ANCES un BSR REEL Tae VANES TEN MoS at Petey aridi ae ge SN aad ae eee, N TINO tact een Aare ne N er AMI ATEI nn NEL ete aon Nennen a Dat det aan, SR WT he AS Ta Ee eisen Weinen BNE RE aN Ney doe! SLPS ATP eet URE Ftd LUNSARSTOTE REN sta DOTE DEEP ave TROIS whee) SS NOT NUS ST LAM prata N IEEE TG} aan ty DÉPOT Me AO LOC Wtveyegy. ICI VOLTI RETTORE Vane yew tay TON ER EP rn Pe ene na Wow alae nn nn Nine un aay RA DECTICIZIA Wet ay OPEN CUITE HET sante Saras nennen un mans at rors eset PTT EE revetses TR ape reno eeeueue Kunert ents yy gen (ETA BA NET pere trio VHD Wd vty Wowie ye etnies COUN EAU ty Dre IRE hen er eye WETTER EO DE thy, Ven porse Vip Mi Dave VAT ENG NENG E AN IE PRIA Là apatia baw DA LICIA è HORE OATES IE CS PILOT en yy CAE DURE CAE My TENTE 1 ee DCE Urt Sun Ab TAS Lena gues CARTE TARDE AVG M A à TAI mont LA Pl ab atib oh Aa Zn DHL] DT DUPONT BLUT TEE ZIONI ies aie runs OCEAN AAV BV BO pe VENT stato CAEN RARE PeD St en ae a bathe HA PN wry Ne whe when Poin BW bat goa mann TZ ACTES CE CETTE EDS PA Ù a NUO: fe ce AIDOTOOZ 10 ASSINS ANAT | | ‘—ADOTOOZ 10 1VNUNOÎ SSIMS IE OOLOG REVUE SUISSE DE ZOOLOGIE TOME 106 — FASCICULE 3 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève FRANCOIS BAUD Conservateur au Muséum d'histoire naturelle de Genève CHARLES LIENHARD Charge de recherche au Muséum d’histoire naturelle de Geneve Comite de lecture President: Ivan LOBL — Muséum de Genéve Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Geneve et de représentants des Instituts de zoologie des universités suisses. Les manuscrits sont soumis a des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, écologie, éthologie, morphologie et anatomie comparee, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www.geneva-city.ch:80/musinfo/mhng/publications/revues.htm PRIX DE L'ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d'abonnement doivent étre adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 4 # A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION Stanley Bowestead Muséum d’histoire naturelle Genéve 1999 Stanley BOWESTEAD A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION The Corylophidae is a family of minute beetles placed in the Cerylonid group of the Cucujoidea. As many allied, they are thought to be mould feeders, being attracted to fine mould that occurs on vegetation as it decays. They are world wide in distribution. The present taxonomic treatment of the Palaearctic taxa is particularly inadequate. Lack of reliable keys makes incertain even identi- fication of common European species. For this revision all available type material and numerous collections totalizing more than 13’000 specimens were examined. The morphological characters are re-appraised. The structures of the head, antennae, prothorax, metathorax, and prolegs relevant for genera, and the habitus, microsculpture, penis with the internal sac, and spermatheca relevant for species are shown on 426 illustrations. The spermathecal characters are used for the first time in the family and they proved to be diagnostic. A cladistic analysis is performed to hypothesize the phylogenetic relationships. All taxa are redefined. Presently, four subfamilies are distinguished: Peltinodinae, Corylophinae, Orthoperinae and Rypobiinae, and 45 species are recognized in the area studied. Latter stretches from the islands of Azores in the west, to the Ural mountains in the east and extends southwards to include Afghanistan and all of North Africa north of Sahara. Five new species and one new tribe are described. Lectotypes are designated for ancient type material. New synonymies of genera and species are indi- cated, and full synonymy is given for all Palaearctic species. Keys to sub- families, tribes, genera and species are provided. Distribution, records and habitat types of species are given. This publication is addressed to systematicians, ecologists, entomologists interested in faunistic research of the individual European countries as well as to scientific librairies. Instrumenta Biodiversitatis n° 3 Ed. Muséum d’histoire naturelle, Ville de Geneve, July 1999, 202 pp., 424 figures ISBN 2-88139-006-5 Price: 60 Swiss francs + handling and postage Orders should be mailed to: Muséum d@’ histoire naturelle, c/o Mme Nathalie Chollet, Case postale 6434, CH-1211 Geneve 6, Switzerland. Fax: +41 22 418 63 Ol, or to e-mail: volker.mahnert @ mhn. ville-ge.ch Werane ordenine.......... copy(ies) of “A Revision of the Corylophidae (Coleoptera) of the West Palaearctic Region”. nee ee ES N 4 \ Er WE # DIE su lang à ek sale va, La a id ER f x t) rent hes der. \ IB BETES: : a un: di | ental AS Nea AL [FREE Bad ivi ANNALES de la SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Geneve tome 106 fascicule 3 1999 E 2 kl GENEVE SEPTEMBRE 1999 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE REVUE SUISSE DE ZOOLOGIE TOME 106 — FASCICULE 3 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève FRANCOIS BAUD Conservateur au Muséum d'histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d'histoire naturelle de Genève Comité de lecture Président: Ivan LOBL — Muséum de Genève 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 a des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www.geneva-city.ch:80/musinfo/mhng/publications/revues.htm PRIX DE L'ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d'abonnement doivent étre 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 106 (3): 539-553; septembre 1999 A revision of the Scopaeus sulcicollis species group, with description of a new species from North-East Anatolia (Coleoptera, Staphylinidae, Paederinae) Johannes FRISCH Institut für Allgemeine und Spezielle Zoologie, Justus-Liebig-Universität Gießen, Stephanstr. 24, D-35390 Gießen, FRG. A revision of the Scopaeus sulcicollis species group, with description of a new species from North-East Anatolia (Coleoptera, Staphylinidae, Paederinae). - The West Palaearctic.Scopaeus sulcicollis species group is defined by external morphological and genital features. It comprises S. am- pliatus Binaghi (North-East Italy, Corsica, Sardinia), S. argonauta Gusarov (Caucasus), S. ponticus sp. n. (North-East Anatolia), S. rubidus Mulsant & Rey (South-West Europe) and S. sulcicollis Stephens (West-, Central-, South Europe), which are illustrated and, except S. argonauta and S. sulcicollis, described using external morphological, genital and meristic characters. A lectotype is designated for S. rubidus. Key-words: Staphylinidae - Paederinae - Scopaeus sulcicollis group - West Palaearctic Region - taxonomy. INTRODUCTION Within Scopaeus (Staphylinidae, Paederinae), both earlier and more recent authors (e. g. BINAGHI 1935; CoIFFAIT 1960, 1968, 1984; FAUVEL 1872; HORION 1965; LOHSE 1964; MULSANT & REY 1854, 1855; REITTER 1909) have frequently employed subgenera which, however, are poorly founded and do not represent monophyletic groups (FRISCH 1996). It was suggested in subsequent revisionary publications on West Palaearctic Scopaeus (e. g. FRISCH 1997, 1998) that these subgenera be replaced by informal groups of related species. Scopaeus sulcicollis Stephens and S. argonauta Gusarov were referred to the S. sulcicollis group (FRISCH 1997) but the complete group has not been revised until now. BINAGHI (1935) noted the close relationship of Scopaeus ampliatus Binaghi, S. rubidus Mulsant & Rey and S. sulcicollis Stephens, 1832 (= S. cognatus Mulsant & Rey sensu BINAGHI 1935), but used inappropriate characters (parameres lacking, apical lobes being fused or nearly fused at apex) to define the species group. A lack of parameres (lateral lobes sensu FRISCH 1994) is a character of several groups of related species, and the apical lobes of the relevant species are, in fact, absolutely separate. Recently, GUSAROV (1992) described S. argonauta from the Caucasus and compared it Manuscript accepted 05.10.1998 540 JOHANNES FRISCH with S. sulcicollis. When designating a lectotype for the latter, FRISCH (1997) referred both species to the S. sulcicollis group, referring to aedeagal characters as the shape of the apical lobes and the apically undivided, ventrally bent dorsal lobe, and to the abdominal sternite 8 having distinct, apico-lateral impressions, which are divided by an elevated middle field. According to the present concept, the Scopaeus sulcicollis group comprises five species in the West Palaearctic realm: $. ampliatus (North-West Italy, Corsica, Sardinia), S. argonauta (Caucasus), S. ponticus sp. n. (North-East Anatolia), S. rubidus (South-West Europe) and S. sulcicollis (West-, Central-, South Europe). These species agree in the discernibly enlarged tempora, in the more or less dilated mesotibiae, in the laterotergites 9 lacking a distinct dorsal tooth, in the aedeagus lacking lateral lobes and in the inner ventral margins of the apical lobes each bearing a row of fine setae. As the first features occur in several groups of related species, only the inner rows of setae of the apical lobes may be an apomorphic feature of the S. sulcicollis group. The species, except S. ampliatus, share furthermore two longitudinal, lateral impressions on abdominal sternit 8 divided by an elevated middle field. As these characters of abdo- minal sternit 8 occur in several species groups but lack in S. ampliatus, they are not usable to define the S. sulcicollis group. Additionally, the aedeagi of S. ampliatus, S. rubidus and S. ponticus sp. n. share somewhat asymmetrical dorsal lobes and apical lobes. The present review gives a more accurate definition of the S. sulcicollis group. It presents redescriptions and the distribution patterns of S. ampliatus Binaghi and S. rubidus Mulsant & Rey and additional notes on S. argonauta, along with illustrations of all species included. Scopaeus argonauta and S. sulcicollis were redescribed in FRISCH (1997). Finally, Scopaeus ponticus sp. n. from North-East Anatolia is included in the S. sulcicollis group. MATERIAL AND METHODS The terminology of the aedeagal features is used as in FRISCH (1994). The sper- matheca and the genital sclerites are named as in FRISCH (1996) and UHLIG (1989). Abdominal sclerites are counted from the first morphological segment. Specimens were measured using a binocular microscope with an eye-piece linear micrometer. Measurements and ratios are based on ten specimens of both sexes, which are including maximum variation range in size and form, and are defined as follows: total length = interval from the apical margin of the mandibles to the end of the abdomen; forebody length = interval from the apical margin of the mandibles to the posterior margin of the elytra at suture; length of head = interval from the apical margin of the clypeus to the posterior margin; HLW = head length : head width; PLW = pronotal length : pronotal width; HPW = width of head : pronotal width; HPL = length of head : pronotal length; PSL = pronotal length : elytral sutural length (excluding scu- tellum); PLL = pronotal length : elytral lateral length; ELW = elytral lateral length : elytral width; ET = eye length : temporal length (both measured laterally); MT = mesotibial length : mesotibial width; A = length (measured without the basal and distal SCOPAEUS SULCICOLLIS SPECIES GROUP 54] tapering): width of the antennal segments 1-11; T = length : width of the central area (between sclerite margins) of the tergite 10; V = length : width of the central area of the female valve. Detailed data are given only for the type specimens. The synonymy of the species presents only primary references. The study material is deposited in the following institutions and private collections: BMNH = The Natural History Museum, London; DEIC = Deutsches Entomologisches Institut, Eberswalde; FCNB = Frey Collection, Naturhistorisches Museum, Basel; HNHM = Hungarian Natural History Museum, Budapest; ISNB = Institut Royal des Sciences Naturelles de Belgique, Brussels; JFCG = J. Frisch Private Collection, Gießen; MCSN = Museo Civico di Storia Naturale “Giacomo Doria”, Genova; MHNG = Muséum d’histoire naturelle, Geneve; MHNL = Musée Guimet d’histoire naturelle, Lyon; MNHN = Muséum National d’Histoire Naturelle, Paris; NHMW = Naturhistorisches Museum, Wien; SMTD = Staatliches Museum fiir Tierkunde, Dresden; TLMF = Tiroler Landesmuseum Ferdinandeum, Innsbruck; VACH = V. Assing Private Collection, Hannover; VGCP = V. Gusavor Private Collection, St. Petersburg; ZMAL = Zoological Museum, Academy of Sciences, St. Petersburg; ZMHB = Museum fiir Naturkunde, Berlin. TAXONOMY Scopaeus argonauta Gusarov Figs 4-6, 17, 22, 25, 29, 34 Scopaeus (s. str.) argonauta Gusarov, 1992: 781. Holotype d, Georgia, Adzharia, Kintrishskiy Nature Reserve, 500 m, 30.07.1990, Gusarov (ZMAL); examined. Scopaeus argonauta; FRISCH 1997: 537. Material examined. Georgia: holotype d, Adzarskaja, Kintrishskiy Nature Reserve, 500 m, 30.07.1990, Gusarov (ZMAL); paratypes 1d, 19, same data as holotype (VGCP, ZMAL); paratype 2, Adzarskaja, Pirveli Maisi, Agdra, 05.08.1990, Gusarov (VGCP); paratypes 29, Abchazskaja, Ganakhleba, 26.07.1990, Gusarov (VGCP); paratype $, Abchazskaja, Gumis- tinskiy Nature Reserve, Tsumuri, Ashamgvara ridge, 900 - 1100 m, 03.07.1981, Gusarov (VGCP); paratype d, Abchazskaja, Gumistinskiy Nature Reserve, Tsumuri, 420 m, 20.07.1990, Gusarov (VGCP). Russia: paratype ¢, Krasnodar Kraj, Lasarewskaja, Aul Kirowa, 29.09.1986, Kirejtshuk (ZMAL). Comments. The similar shape of the aedeagus and the abdominal sternite 8 of males suggests that Scopaeus argonauta (figs 4-6, 17) and S. sulcicollis (figs 1-3, 16) are closely related (FRISCH 1997, GUSAROV 1992). Meanwhile, the holotype d and paratypes (38, 59), which were not traceable then, have been examined, and S. argo- nauta turned out to be a species of quite variable elytral length and not distinguished constantly from S. sulcicollis by the longer elytra. Specimens of S. argonauta with shorter elytra laterally exceeding the pronotal length by just about a seventh (PLL 0,86) cannot be distinguished from S. sulcicollis. Scopaeus argonauta is characterized furthermore by a rather variable shape of the head, which has the tempora rather parallel or notably widened. Scopaeus ponticus sp. n. Figs 79182526 305 35 Material examined. Holotype d, Turkey: Trabzon, Arakli, 100 m, 24.05.1997, collected on the narrow bank of a small stream with damp, fine gravel and sand, Frisch (MHNG). Paratypes. 79, same data as holotype (JFCG, MHNG). 542 JOHANNES FRISCH Fics 1-6 Scopaeus argonauta, 3, Caucasus: aedeagus in 1) lateral, 2) ventral, 3) dorsal view. Scopaeus sulcicollis, 3 lectotype: aedeagus in 4) lateral, 5) ventral, 6) dorsal view. Scale bar = 0.1 mm. SCOPAEUS SULCICOLLIS SPECIES GROUP 543 Description. Length 3.2-3.9 mm; forebody 1.6-1.8 mm. Dark coloured species. Forebody brown, pronotum slightly lighter brown, elytra blackish without distinct sutural or distal lightenings. Abdomen black, segments 8-10 dark brown. Appendages unicolorously light brown. Punctuation distinct, relatively coarse on elytra, but rather fine on pronotum, reticulation rather obsolete, forebody therefore somewhat shining. Head trapezoid with notably widened tempora, distinct hind angles and with a rather concave posterior margin, about a tenth longer than wide and up to almost a fifth wider than pronotum. Eyes rather small, distinctly less than half as long as tempora. Flightless species. Elytra rather short and slender, laterally as long as pronotum or exceeding pronotal length just by about a tenth, at suture up to a fifth shorter than pronotum. Membranous wings strongly reduced, not exceeding double the length of the elytra. Protarsomeres 1-4 in both sexes twice as wide as long. Mesotibia moderately dilated, about five times as long as wide. Antennae rather slender, distal antennomeres almost quadrate. Laterotergite 9 (fig. 23) without a distinct dorsal tooth but slightly dilated apico-dorsally. Male sternite 8 (fig. 18) with a triangular emargination in the apical third and two medio-lateral, lighter coloured, narrow, longitudinal impressions which bear no setae and are divided by a broad, elevated middle field with lateral margins distinct from the lateral impressions. Aedeagus (figs 7-9) with slightly asymmetrical apical lobes and dorsal lobe, with apices somewhat bent to the right in dorsal view. Apical lobes about half as long as phallobase and broadangled bent ventrally in the distal half, their ventral margins in the distal half each extended into a longer, acute distal and a shorter, rounded proximal lobe, which are divided by a semicircular emargination. Proximal lobes bent laterally in ventral view. Outer margins of the apical lobes in dorsal view evenly narrowed in the basal half but somewhat rounded and dilated in the distal half, their inner margins in ventral view each bearing a row of about six short setae. Dorsal lobe conspicuously long and slender, broadangled bent ventrally in the distal half just as the apical lobes, its acute apex strongly projects from the apex of the apical lobes and is extended ventrally to a strong, median tooth. Lateral lobes reduced to lateral groups of few, short setae. Spermatheca (fig. 35) with members slightly bent in lateral view. Chamber strongly triangular, process rather slender. Ratios. HLW 1.06-1.13; PLW 1.15-1.23; HPW 1.1-1.17; HPL 1.0-1.06; PSL 107-1217 PLE 0:87-0:99; ELW 1.16-1.23; ET 0:39-0.48; MT 4:9-5:6; A 22, 1.3; 1:3; SAP OM EL ON Mer DIE VIA. Comments. Externally, S. ponticus sp. n. is difficult to distinguish from S. sul- cicollis and S. argonauta, but may be separated from S. argonauta by the reduced metathoracic wings. Additionally, S. ponticus differs from the specimens examined of S. argonauta, which have elytra exceeding the pronotal length by about a fifth, by their discernibly shorter elytra (see FRISCH 1997). However, bearing in mind the meagre material examined, the length of the membranous wings may be variable in S. ponticus, and the length of the elytra may be as variable as in S. sulcicollis and S. argonauta. Judging from the aedeagal characters, S. ponticus sp. n. is isolated within the S. sulci- collis group. 544 JOHANNES FRISCH Scopaeus rubidus Mulsant & Rey Figs 10-12, 19, 24, 27, 31, 36 Scopaeus rubidus Mulsant & Rey, 1854: 171; 1855: 59. Lectotype d, France, Provence (MHNL); here designated (examined). Scopaeus (Polyodontus) rubidus; FAUVEL 1872: 313. Scopaeus (Heteroscopaeus) rubidus; COIFFAIT 1960: 285. Scopaeus (Anomoscopaeus) rubidus; COIFFAIT 1968: 426. Scopaeus subcylindricus Scriba, 1868: 156; synonymized with S. rubidus by FAUVEL 1871: 136. Material examined (291 specimens). Danmark (MNHN). France: Lectotype d and paralectotype 19 of S. rubidus, Provence (MHNL); paralectotypes 16, 32 of S. rubidus, Lyon (MHNL); Ain (NHMW); Allier (MNHN); Alpes de Haute-Provence (MNHN); Alpes Maritimes (MHNG, MNHN); Aube (ISNB); Aude (FCNB, NHMW, MNHN, SMTD); Bas-Rhin (MNHN); Bouches-du-Rhöne (NHMW); Dröme (ISNB); Gironde (MNHN); Haute-Garonne (ISNB, MNHN); Hautes-Pyrénées (ISNB); Hérault (ISNB, NHMW, MNHN); Isere (MHNG, MNHN); Landes (MNHN); Loiret (NHMW, MNHN); Pyrénées-Atlantiques (ISNB); Rhone (DEIC, NHMW, MNHN); Savoie (ISNB); Var (ISNB, MNHN); Vaucluse (HNHM, MHNG, MNHN). Italy: Abruzzi (NHMW); Emilia-Romagna (JFCG, NHMW, TLMF); Lazio (DEIC, MHNG, NHMW, ZMHB); Liguria (NHMW); Piemonte (BMNH, DEIC, ISNB, MCSN, NHMW, MNHN, ZMHB); Sardinia (HNHM); Toscana (MCSN, SMTD). Spain: Andalucia (ISNB, MHNG, MNHN), Aragon (JFCG, MHNG, NHMW, VACH); Castilla-Leon (MHNG); Catalunya (MHNG); Madrid (MHNG). Switzerland: Genève (MHNG). Description. Length 3.4-3.9 mm; forebody 1.8-2.0 mm. Light coloured species of a rather broad and convex habitus. Forebody unicolorously light reddish brown to orange, pronotum slightly lighter, abdomen brown to dark brown, segments 8-10 somewhat lighter brown. Appendages evenly light brown. Punctuation distinct, rather fine and spacious on pronotum, on elytra with separate punctures and not as granular as in related species. Reticulation omits completely, forebody therefore distinctly shining. Head trapezoid with tempora notably widened, hind angles distinct and with a straight or slightly concave posterior margin, as long as wide or just up to a tenth longer than wide and just about a tenth wider than the rather broad pronotum. Eyes rather large, about half as long as tempora. Elytra rather broad and convex, laterally exceeding pronotal length by about a fifth, at suture about as long as pronotum. Membranous wings entire. Protarsomeres 1-4 in both sexes twice as wide as long. Mesotibia remar- kably dilated, distinctly less than five times as long as wide. Distal antennomeres discernibly transverse. Laterotergite 9 (fig. 24) having a very slight, apico-dorsal dila- tation. Male sternite 8 (fig. 19) with a rather short triangular emargination in just the apical fifth and two lateral, lighter coloured, broad impressions lacking the fine basic setae but studded with only stronger setae, and divided by a broad, slightly elevated middle field, which bears laterally orientated setae. Aedeagus (figs 10-12) with lobes asymmetrical apically. Apical lobes slightly shorter than the phallobase, strongly Fics 7-12 Scopaeus ponticus sp. n., 6 holotype: aedeagus in 7) lateral, 8) ventral, 9) dorsal view. Scopaeus rubidus, 3, France, Isere: aedeagus in 10) lateral, 11) ventral, 12) dorsal view. Scale bar = 0.1 mm. SCOPAEUS SULCICOLLIS SPECIES GROUP 545 546 JOHANNES FRISCH dilated ventrally in the distal half, apically deeply emarginate, each forming a slender, apically truncate and somewhat dilated dorsal projection and a broad, acuminate, distally bent ventral projection, which are notably bent to the right in ventral view. Apical lobes slender and evenly narrowed with asymmetrical apices in dorsal view, ventral inner margins each bearing a long row of about 15-20 short setae. In dorsal view the right lobe is strongly dilated ventro-laterally in the basal half. Dorsal lobe slender at the base but evenly bent ventrally and strongly dilated toward the apex, not projecting from the distal and the ventral margins of the apical lobes but from the deep apical emargination, serrate apico-ventrally and extended apically into an acute spine, which is bent to the right in ventral view. A short, triangular spine projects ventrally from the basis of the apical lobes. Phallobase without lateral setae, but with a remarkably prominent, ventral extension. Spermatheca (fig. 36) with members rather straight in lateral view. Chamber strongly triangular, process slender. Ratios. HLW 1.04-1.11; PLW 1.15-1.25; HPW 1.06-1.13; HPL 0.95-1.01; PSL 0.95-1.07; PLL 0.77-0.85; ELW 1.16-1.28; ET 0.48-0.55; MT 4.2-4.7; A 2.5, 1.4, 1.5, ANS Wall, O24 WS), Oss Wess, Wee Ie 202 WAC) sir. Distribution. Scopaeus rubidus is distributed in Western Europe and in the West Mediterranean region. The species was recorded repeatedly from the north of France (Maine-et-Loire: ABOT 1928; Champagne: CoIFFAIT 1984; Aubé, Allier: FAUVEL 1872; Paris: SAINTE-CLAIRE DEVILLE 1907), and a specimen from Aube was examined. Scopaeus rubidus is confirmed throughout France and Spain southwards to Andalucia, but has not been recorded from the west of the Iberian Peninsula (Galicia, Portugal). Eastwards, the known range reaches from the southern Rhine valley (Strasbourg: CoIFFAIT 1984) and Genève along the South Alpes over the Appenines, where it is confirmed southwards to Lazio and Abruzzi. Scopaeus rubidus occurs on Sardinia but is unknown from Corsica, Sicily, the Balearic Islands and North Africa. The occurance of S. rubidus in Germany is doubtful. Ancient authors recorded the species from South Germany (BERNHAUER & SCHUBERT 1910, GANGLBAUER 1895, Horıon 1965, REITTER 1909, SCHEERPELTZ 1925) and South Bavaria (HORION 1965, KRAATZ 1858), but no specimens from Germany were available from relevant collec- tions. Regarding the confirmed distribution pattern, the record from the upper Rhine valley (South Baden: HORION 1965) is plausible. Records from Austria (REDTENBACHER 1874, SCHEERPELTZ in HORION 1965), the Czech Republic (BoHAc 1985, 1993; KLIMENT 1899), Hungary (LASZLO 1893), Russia (FAUVEL 1872) and the Ukraine (doubted by Horıon 1965) are implausible and based obviously on misidentifications. Alleged specimens from Austria (Lower Austria, Burgenland, Styria, Carinthia), which were given by SCHEERPELTZ (in HORION 1965) and which should be in his collection (NHMW), are absent there. Ancient records from Bohemia and Moravia (KLIMENT 1899) have not been confirmed by recent specimens (BoHAc 1985) and have already been doubted by Horion (1965). Nevertheless, BOHAC (1993) mentions S. rubidus in the Checklist of Czechoslovak Insects. The record of a supposed © of S. rubidus from Crete (SCHEERPELTZ 1964) has to be declined as well. SCOPAEUS SULCICOLLIS SPECIES GROUP 547 Fics 13-15 Scopaeus ampliatus, 3, Italy, Elba: aedeagus in 13) lateral, 14) ventral, 15) dorsal view. Scale bar = 0.1 mm. The record of S. rubidus from Devon in South England (FOWLER 1888), which was adopted by SAINTE-CLAIRE DEVILLE (1907), refers to S. ryei Wollaston (NEWBERRY 1914; EDMONDS 1931, 1932; ALLEN 1968), which was described from this British region. A specimen labelled “Danemark, Venloo” (Jarrige collection, MNHN) is possi- bly mislabelled. The species has not been found in Denmark (M. Hansen, Zoological Museum, Copenhagen, pers. comm.), and its occurrence so far north appears im- plausible. Comments. COIFFAIT (1968, 1984) falsely combined S. rubidus with S. gracilis Sperk and allied species in the ill-based subgenus Anomoscopaeus, which he defined only by asymmetrical apical lobes, an absolute unusable feature in Scopaeus, which occurs in several groups of related species. In fact, S. rubidus does not reveal any relationship to other species of “Anomoscopaeus”. Scopaeus subcylindricus was described from two $ 2 (SCRIBA 1868) collected by Seidlitz. Their origine is not given in the description, and the specimens are not traceable in the Zoologische Staatssammlung, Munich, where the Seidlitz collection 548 JOHANNES FRISCH was given (HORN er al. 1990). The Scriba collection was bought in 1898 by K. Neumann, Bad Homburg, who later published a list of the contained types (NEUMANN 1907), in which he omits Scopaeus subcylindricus. On the basis of the characters given by SCRIBA (1868) for S. subcylindricus, its synonymy with S. rubidus (FAUVEL 1871) appears plausible. Scopaeus ampliatus Binaghi Figs 13-155 201757] Scopaeus (Polyodontus) ampliatus Binaghi, 1935: 95. Holotype 6, Italy, Liguria, Genova, 10.1920, Dodero (MCSN); examined. Scopaeus (Hyposcopaeus) ampliatus; COIFFAIT 1968: 419. Material examined (15 specimens). France: Corsica (BMNH, JFCG, MNHN, NHMG). Italy: holotype & and paratypes 1d, 22, Liguria, Genova (MCSN); paratype d, Sardinia, Laconi (MCSN); Toscana (Elba) (NHMW). Description. Length 3.0-3.2 mm; forebody 1.5-1.7 mm. Forebody unicolorously brown to dark brown, pronotum slightly lighter, abdomen blackish brown, segments 8- 10 somewhat lighter brown. Appendages evenly light brown. Punctuation and reticu- lation as in S. rubidus, but elytra in some specimens with rather granular punctuation and weak reticulation. Head trapezoid with tempora notably widened, hind angles distinct and with a slightly to distinctly concave posterior margin, slightly or up to a tenth longer than wide and up to 0.17 times wider than pronotum. Eyes about half as long as tempora. Elytra relatively short, laterally about as long as pronotum or exceeding pronotal length by just almost a fifth, at suture slightly or up to almost a fifth shorter than pronotum. Membranous wings entire. Protarsomeres 1-4 in both sexes twice as wide as long. Mesotibia slightly dilated, about five to six times as long as wide. Distal antennomeres discernibly transverse. Laterotergite 9 (fig. 21) with a small, apico-dorsal tooth. Male sternite 8 (fig. 20) with a triangular emargination in the apical fourth having the proximal end rather rounded, but without lateral impressions and elevated midline. Lobes of the aedeagus (figs 13-15) slightly asymmetrical and some- what bent to the right in dorsal view. Apical lobes short and stubby, strongly bent ven- trally, each bearing a small, ventro-basal, triangular tooth and a long, dorsally curved apical spine, their ventral margins rather obtuse. Basal two-thirds of the apical lobes asymmetrically convex, afterwards strongly narrowed and parallel toward the apex in dorsal view. Outer margins of the apical lobes strongly convex and evenly narrowed toward the apex in ventral view, their inner ventral margins each bearing a row of about five short setae. Dorsal lobe slender at the base, strongly bent ventrally, claviform dilated distally and deeply divided ventrally into two lateral halves, each bearing an acute, apical tooth which is somewhat protruding from the ventral margins of the apical lobes. Phallobase without lateral setae, somewhat dilated distally and afterwards strongly narrowed toward the apical lobes. Spermatheca (fig. 37) with members some- what bent and strongly triangular. Ratios. HLW 1.05-1.12; PLW 1.19-1.26; HPW 1.09-1.17; HPL 0.97-1.02; PSL 1.05-1.18; PLL 0.83-0.94; ELW 1.06-1.25; ET 0.46-0.57; MT 5.0-5.8; A 2.3, 1.6, 1.3, fe 1821812050:950:9,.0:820:8715512.071,. C)53: Distribution. Scopaeus ampliatus is recorded reliably only from North-West Italy (Liguria, Toscana), Elba, Corsica and Sardinia. 549 SCOPAEUS SULCICOLLIS SPECIES GROUP Fics 16-24 Sternite 8, d: 16) Scopaeus sulcicollis, Bosnia; 17) S. argonauta, Caucasus; 18) S. ponticus sp. - Laterotergite 9, 2: 21) Italy, Elba. S. ampliatus, paratype, Italy, Genova; 22) S. argonauta, Caucasus; 23) S. ponticus sp. n., para- France, Isère, 20) S. ampliatus, type; 24) S. rubidus, Spain, Segovia. Scale bars: a = 0,1 mm (Figs 16-20); b = 0,1 mm (Figs n., holotype; 19) S. rubidus, 21-24). 550 JOHANNES FRISCH SCOPAEUS SULCICOLLIS SPECIES GROUP 551 Comments. Scopaeus ampliatus and S. rubidus share a similar shape of the dorsal lobe of the aedeagus, which is conspicuously dilated toward the apex, and appear related closely. Both species occur sympatrically in Tuscany, Liguria and Sardinia. ACKNOWLEDGEMENTS My sincere thanks are due to the following colleagues for the loan of material from the respective institutes: N. Berti, MNHN; M. J. D. Brendell, BMNH; J. Clary, MHNL; D. Drugmand, ISNB; O. Jäger, SMTD; M. Kahlen, TLMF; A. G. Kirejtshuk, ZMAL; I. Löbl, MHNG; R. Poggi, MCSN; H. Schillhammer, NHMW; E. Sprecher, FCNB; G. Szél, HNHM; M. Uhlig, ZMHB and L. Zerche, DEIC. Specimens from private collections were provided by V. Assing, Hannover, and V. Gusarov, St. Peters- burg. I thank also P. M. Hammond, BMNH, for proof-reading the manuscript. In particular I am indebted to I. Löbl, MHNG, and V. Wolters, University of Gießen, who kindly supported my work in many ways. This paper is part of a Ph. D. thesis at the University of Gießen and was supported by the Studienstiftung des Deutschen Volkes, Bonn-Bad Godesberg. REFERENCES ABOT, G. 1928. Catalogus des Coléoptères observés dans le département de Maine-et-Loire. Paris, 386 pp. ALLEN, A. A. 1968. Notes on some British Staphylinidae (Col. ). 1. The genus Scopaeus Er. with the addition of S. laevigatus Gyll. to our list. Entomologist’s monthly magazine 104: 198-207. BERNHAUER, M. & SCHUBERT, K. 1910: Staphylinidae 1. - In: JUNK W. & SCHENKLING, S. (eds): Coleopterorum Catalogus 19. Berlin, 86 pp. BINAGHI, G. 1935. Studio sul genere Scopaeus Erich. (Coleopt.: Staphylin.). Memorie della Societa Entomologica Italiana 14: 84-115. BoHac, J. 1985. Review of the subfamily Paederinae (Coleoptera, Staphylinidae) in Czechoslo- vakia. Part II. Acta entomologica bohemoslovaca 82: 431-467. BoHac, J. 1993. Staphylinidae. - In: JELINEK, J. 1993: Check-list of Czechoslovak Insects 4 (Coleoptera). Folia Heyrovskyana, Suppl. 1: 39-62. COIFFAIT, H. 1960. Démembrement du genre Scopaeus et description de 4 espèces nouvelles (Coleopt.: Staphylinidae). Revue francaise d’Entomologie 27: 283-290. COIFFAIT, H. 1968. Scopaeus nouveaux ou mal connus de la région paléarctique occidentale. Bulletin de la Société d’Histoire naturelle de Toulouse 104: 405-426. COIFFAIT, H. 1984. Coléoptères Staphylinides de la région paléartique occidentale V: Sous- famille Paederinae Tribu Paederini 2, Sous-famille Euaesthetinae. Publications de la Nouvelle Revue d’Entomologie VIII, 424 pp. Fics 25-37 Valve, 9: 25) Scopaeus argonauta, Caucasus; 26) S. ponticus sp. n., paratype; 27) S. rubidus, Spain, Segovia; 28) S. ampliatus, paratype, Italy, Genova. - Tergite 10, 9: 29) Scopaeus argonauta, Caucasus; 30) S. ponticus sp. n., paratype; 31) S. rubidus, Spain, Segovia; 32) S. ampliatus, paratype, Italy, Genova. - Spermatheca: 33) S. sulcicollis, Bosnia; 34) Scopaeus argonauta, Caucasus; 35) S. ponticus sp. n., paratype; 36) S. rubidus, Spain, Segovia; 37) S. ampliatus, paratype, Italy, Genova. Scale bar = 0,1 mm. 552 JOHANNES FRISCH EDMONDS, T. H. 1931. The British species of the genus Scopaeus Er. (Coleoptera, Staphylinidae). Entomologist’s Monthly Magazine 67: 272-275. EDMONDS, T. H. 1932. Some further notes on the genus Scopaeus Er. (Coleoptera, Staphylinidae) and an addition to the British list. Entomologist’s Monthly Magazine 68: 206-209. FAUVEL, A. 1871. Décades Synonymiques. L’Abeille 7: 136. FAUVEL, A. 1872. Fauna gallo-rhénane 1. Bulletin de la Société Linnéenne de Normandie 2 (7): 8-132. FOWLER, W. W. 1888. The Coleoptera of the British Islands. A Descriptive Account of the Families, Genera and Species Indigenous to Great Britain and Ireland, with Notes as to Localities, Habitats, etc., Vol. II. London, 444 pp., pls 37-70. FRISCH, J. 1994. Neue Arten der Gattung Scopaeus Erichson aus Griechenland, Anatolien und dem Iran (Coleoptera, Staphylinidae, Paederinae). 1. Beitrag zur Kenntnis der Gattung Scopaeus Erichson. Coleoptera 2: 1-46. FRISCH, J. 1996. Revision westmediterraner Scopaeus - Arten (Coleoptera, Staphylinidae: Paederinae) und Beschreibung einer neuen Art aus Südspanien und Marokko. 2. Beitrag zur Kenntnis der Gattung Scopaeus Erichson. Revue suisse de Zoologie 103: 301-318. FRISCH, J. 1997. A revision of some West Palaearctic species of Scopaeus Erichson (Coleoptera, Staphylinidae, Paederinae). Revue suisse de Zoologie 104: 523-557. FRISCH, J. 1998. A revision of some West Palaearctic species of Scopaeus Erichson (Coleoptera, Staphylinidae, Paederinae). Revue suisse de Zoologie 105: 89-124. GANGLBAUER, L. 1895. Die Käfer von Mitteleuropa 2: Familienreihe Staphylinoidea. 1. Theil: Staphylinidae, Pselaphidae. Wien, 880 pp. GUSAROV, V. 1992. New and little known Palaearctic Staphylinids (Coleoptera, Staphylinidae). Entomologicheskoe Obozrenie 71 (4): 775-788, 951. HEYDEN, L. v., Reitter, E. & Weise, J. 1891. Catalogus Coleopterorum Europae, Caucasi et Armeniae rossicae. Berlin, Mödling, Caen, 420 pp. HEYDEN, L. v., REITTER, E. & WEISE, J. 1906. Catalogus Coleopterorum Europae, Caucasi et Armeniae rossicae. Berlin, Paskau, Caen, 774 pp. Horion, A. 1965. Faunistik der mitteleuropäischen Käfer X: Staphylinidae 2. Teil: Paederinae bis Staphylininae: Überlingen, 335 pp. Horn, W., KAHLE, I, FRIESE, G. & GAEDICKE, R. 1990. Collectiones entomologicae. Ein Kompendium über den Verbleib entomologischer Sammlungen der Welt bis 1960. Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik, Berlin, part 1, 220 pp., part 2, 573 pp. KLIMENT, J. 1899. Cesti brouci. Dilo o broucich Cech, Moravy a Slezska. Nem. Brod, 811 pp., 46 +4pl. KRAATZ, G. 1856-1858. Naturgeschichte der Insecten Deutschlands 2, Staphylinii. Berlin, 1080 pp. 1-376: 1856, 377-768: 1857a, 169-1080: 1857b. LASZLO, T. 1983. Coleoptera 2: Staphylinidae 5. Fauna Hungariae 155, 69 pp. Louse, G. A. 1964. Staphylinidae 1 (Micropeplinae bis Tachyporinae). In: FREUDE, H., HARDE, K. W. & LOHSE, G. A. 1964: Die Käfer Mitteleuropas 4. Krefeld, 264 pp. MULSANT, E. & REY, C. 1854. Essai spécifique sur les Scopaeus des environs de Lyon. Annales de la Société Linnéenne de Lyon 2: 161-190. MULSANT, E. & REY, C. 1855. Essai spécifique sur les Scopaeus des environs de Lyon. Opuscules Entomologiques 6: 49-79. NEUMANN, K. 1907. Einiges tiber Staphylinen (Col.). Die Typen und Originale aus der Sammlung des Pfarrers D. Scriba. Deutsche Entomologische Zeitschrift 1907: 327-330. NEWBERRY, E. A. 1914. Scopaeus rubidus of British collections: synonymical note. Entomo- logist’s Monthly Magazine 50: 91-92. REDTENBACHER, L. 1874. Fauna Austriaca. Die Kafer. 3. edition. Wien, 564 pp. SCOPAEUS SULCICOLLIS SPECIES GROUP 553 REITTER, E. 1909. Fauna Germanica. Die Kafer des Deutschen Reiches 2. Stuttgart, 392 pp., 80 plates. SAINTE-CLAIRE DEVILLE, J. 1907. Faune des Coléoptères du bassin de la Seine 2: Staphylinoidea. Paris, 169 pp. SCHEERPELTZ, O. 1925. Staphylinidae. /n: WINKLER, A. (ed.) 1924-1932: Catalogus Coleop- terorum regionis palearcticae. Wien, 1698 pp. SCHEERPELTZ, O. 1964. Staphyliniden von der Insel Kreta (Coleoptera). Annales Historico- naturales Musei Nationalis Hungarici 56, Pars Zoologica: 296-308. SCRIBA, W. 1868. Neue europäische Staphylinen. Berliner Entomologische Zeitschrift 12: 153- 160. 1 STEPHENS, J. F. 1832. Illustrations of British Entomology, a synopsis of indigenous insects containing their generic and specific distinctions. Mandibulata, 5. London, 240 pp. UHLIG, M. 1989. Zur Morphologie der weiblichen Terminalia einiger Staphylinidenarten (Cole- optera), Verhandlungen des 11. SIEEC Gotha 1986: 227-237. REVUE SUISSE DE ZOOLOGIE 106 (3): 555-570; septembre 1999 Abundance, species composition and phenology of Pauropoda (Myriapoda) from a secondary upland forest in Central Amazonia Joachim ADIS!, Ulf SCHELLER?, José Wellington de MORAIS? & José Maria G. RODRIGUES? | Max-Planck-Institute for Limnology, Tropical Ecology Working Group, Postfach 165, D-24302 Plön, Germany. 2 Häggeboholm, Häggesled, S-53194 Järpäs, Sweden. 3 Instituto Nacional de Pesquisas da Amazönia (INPA), C.P. 478, 69.011-970 Manaus/AM, Brazil. Abundance, species composition and phenology of Pauropoda (Myria- poda) from a secondary upland forest in Central Amazonia. - The 6,878 pauropods collected within 12 months in the soil (0-7 cm depth) of a secondary upland forest (1,085.7 + 42.3 ind./m°/month) near Manaus were represented by 41 species of the Pauropodidae (Order Tetramerocerata). The Pauropodinae were represented by 31 species, the Polypauropodinae by 6 species, and the Scleropauropodinae by 4 species. About half of all pauropod specimens obtained inhabited the organic soil layer (0-3.5 cm depth) compared to the mineral subsoil (3.5-7 cm). Abundance of pauropods in the soil was twice as high in comparison to the Symphyla from the same study site. The lack of a distinct reproductive period in eudominant and dominant pauropod species and the presence of juveniles and adults throughout the year indicate a plurivoltine mode of life. Only in one species was the monthly catch of adults positively correlated with maximum temperatures of the soil. Pauropods obtained from the soil of four other upland forests in Central Amazonia (0-14 cm depth) accounted for 1.1-4.4% of the total soil arthropods. A possible parthenogenesis found in three pauropod species 1s discussed. Key-words: abundance - phenology - parthenogenesis - soil fauna - Pauro- poda - Amazon - Neotropics. INTRODUCTION Terrestrial arthropods of Central Amazonian forests have been investigated for several years (cf. ADIS & SCHUBART 1984; Apis 1997; Apis et al. 1997a,b) in a co- operative venture between the National Institute for Amazonian Research (INPA) at Manaus/Brazil and the Tropical Ecology Working Group at the Max-Planck-Institute for Limnology in Plön/Germany (Projeto INPA/Max-Planck). Data on abundance and Manuscript accepted 16.02.1999 556 JOACHIM ADIS ET AL. phenology of Pauropoda sampled during 12 months in 1982/83 in a secondary upland forest are now available, as their taxonomical evaluation has been completed (SCHELLER 1994, 1997). Our data represent the very first contribution on the abundance and phenology of Neotropical pauropod species. Voucher specimens have been deposited at the Systematic Entomology Collections of the Instituto Nacional de Pesquisas da Amazonia (INPA) in Manaus/Brazil and at the Muséum d'histoire naturelle in Geneva/ Switzerland. STUDY AREA, MATERIAL AND METHODS Pauropods were collected between 1981 and 1983 in the course of ecological studies on Central Amazonian arthropods from a previously investigated and fully described secondary upland forest at Rio Taruma Mirim (03°02'S, 60°17'W), a tributary of the Rio Negro near Manaus, where the vegetation was previously cut but unburned (cf. Apis 1992, FRANKLIN et al. 1997). The forest is subject to a rainy season (December-May: average precipitation 1550 mm; 258.8 + 36.8 mm/month) and a "dry" season (June-November: average precipitation 550 mm; 91.8 + 43.8 mm/month, but each month with some rain events; cf. RIBEIRO & Apıs 1984). The yellow latosol (= ferrasol in JORDAN 1984) of the secondary upland forest supported a 2-3 cm thick humus layer, interspersed with fine roots, and a thin surface covering of leaf-litter. One ground photo-eclector (emergence trap with pitfall trap inside) and one arboreal photo- eclector for trunk ascents (funnel trap) were installed in the forest from December 1981 to December 1982 (see ADIs & SCHUBART 1984). The distribution of pauropods in the soil was studied between September 1982 and August 1983 (RODRIGUES 1986). Twelve soil samples were taken once a month every two meter along a randomly selected transect. The split corer composed of a steel cylinder with lateral hinges (diameter 21 cm, length 33 cm), was driven into the soil by a mallet. Each sample of 7 cm depth was then divided into two subsamples of 3.5 cm each. Animals were extracted from subsamples following a modified method of Kempson (Apis 1987). The combined area of 12 samples represented 0.42 m?. Calculated average abundances per m? are given with sample standard deviation. The monthly collection data of pauropods from the two soil layers in relation to changing conditions of precipitation, temperature and humidity of the air near the forest floor as well as moisture content, temperature and pH of the soil were statistically evaluated with a linear, parametric correlation test (CAVALLI- SFORZA 1972) using the original field data (RODRIGUES 1986). Pauropoda (Pauro- podidae) sampled were classified as juveniles (3, 5 and 6 pairs of legs), subadults (8 pairs of legs) and adults (9 pairs of legs). Sex was determined for subadults and adults. RESULTS A total of 6,878 pauropods (body length <1.50 mm without antennae) were collected. Out of these, 90% could be identified to species and developmental stages. All pauropods were obtained by extraction of the forest soil. None were caught on the soil surface (ground photo-eclector), or on the tree trunk (arboreal photo-eclector). PAUROPODA CENTRAL AMAZONIA 3000 2000 Specimens/m’ SP Or ANA D <— 1982 ——— J — Pauropoda + Symphyla Fic. 1 400 Nim? = 14.443 | 300 200 27 N/m = 7 170 100 Ein IA 1983 ° — 557 Precipitation (mm) Distribution of Pauropoda and Symphyla in the soil. Samples taken monthly at 0-7 cm depth between September 1982 and August 1983 in a secondary upland forest near Manaus. (N = total number of specimens). Total precipitation per month given between sampling dates (= in the middle of each month). The low rainfall observed in early 1983 was due to a strong El Nifio- event (cf. ADIS & LATIF 1996). TABLE | Average abundance (N/m?) and dominance (%) of Pauropoda genera (all representing the Pauropodidae) in the soil of a secondary upland forests near Manaus, Brazil. Samples taken monthly at 0-3.5 and 3.5-7.0 cm soil depths between September 1982 and August 1983. PAUROPODIDAE N/m? per month AAN Genus ( Species) 0-3.5 cm 3.5-7.0cm 0-7.0cm 0-7.0cm Allopauropus (28) 486.6 + 24.8 509.1 + 26.3 995.7 + 49.6 91.7 Hemipauropus (2) 84.1 + 6.6 16.4+ 4.7 50.5 + 10.4 4.7 Polypauropus (2) 17.3+ 7.4 12.2+ 6.0 29.5 + 13.3 2.7 Scleropauropus (4) 2.0+ 0.6 1.8+ 0.7 3.8+ 1.2 0.3 Polypauropoides (4) 14+ 0.4 40+ 09 5.4+ 1.1 0.5 Cauvetauropus (1) 0.4+ 0 0.4+ 0 0.8+ 0 0.1 Total 541.8 + 21.1 543.9 + 22.4 1085.7 + 42.3 100.0 558 JOACHIM ADIS ET AL. Pauropoda represented 6.4% of the total arthropods extracted from soil samples within 12 months (Acari & Collembola omitted; cf. RODRIGUES 1986). Their abundance in 0-7 cm soil depth was twice as high than that of the Symphyla (Fig. 1: 14,443 versus 7,170 ind./m?; cf. Apis et al. 1997c). An average abundance of 1,085.7 + 42.3 pauro- pods/m?/month was recorded in 0-7 cm soil depth (Table 1). About half of the pauropods obtained (49.9%) inhabited the organic soil layer (Fig. 2: 0-3.5 cm) compared to the mineral subsoil (3.5-7.0 cm depth). Of the total catch, 68.8% was represented by adults (Fig. 2), 4.6% by subadults and 26.6% by juveniles. The total catch of pauropods collected during the dry season was three times higher compared to the rainy season: 77.5% (1,865.2 + 980.5 ind./m°/month) versus 22.5% (542.0 + 180.2 ind./m?/month). 60 h N/m? = 13 024 6 9 40 (6) 3 © © 20 o 0 0 -3.5 3.5-7.0 Soil Depth (cm) adults Dimmatures J=juveniles S = subadults A = adults FIG. 2 Distribution of Pauropoda in the soil according to depth, and percentage of developmental stages in a secondary upland forest near Manaus. (Total catch = 100%). Samples taken monthly at 0-3.5 and 3.5-7 cm depths between September 1982 and August 1983. (N = total number of specimens). The pauropods collected from the secondary upland forest represented 41 species of the family Pauropodidae (Order Tetramerocerata). Out of these, 38 were new (SCHELLER 1994, 1997) and only 3 known (Table 2). Allopauropus, Hemipauropus and Cauvetauropus represented the subfamily Pauropodinae (31 species), Polypauropus and Polypauropoides the subfamily Polypauropodinae (6 species), and Scleropauropus the subfamily Scleropauropodinae (4 species). Out of the six genera obtained (Tables 1, 2), Allopauropus (28 species) represented 91.7% of the total catch, Hemipauropus (2 species) 4.6%, and Polypauropus (6 species) 2.7%. Representatives of the genus Allopauropus were as abundant in the organic soil layer as in the mineral subsoil PAUROPODA CENTRAL AMAZONIA 559 TABLE 2 Average abundance (N/m) and dominance (%) of Pauropoda species (all representing the Pauropodidae) in the soil of a secondary upland forests near Manaus, Brazil. Samples taken monthly at 0-3.5 and 3.5-7.0 cm soil depths between September 1982 and August 1983. Species Subf. N/m? per month % | No. 0-3.5cm 3.5-7.0cm 0-7.0cm 0-7.0cm 1 |Allopauropus (A.) rodriguesi Sch. Pau. 174.0 + 233.3 184.1 + 223.1 358.1 + 441.2 33.0 2 |Allopauropus (D.) bicornutus Sch. Pau. 78.4+ 94.9 126.9 + 153.4 205.3 + 238.7 18.9 | 3 |Allopauropus (D.) proximus Remy Pau. 48.3+ 76.8 11.64 14.1 59.9+ 88.7 5.5 4 |Allopauropus (A.) dundoensis Remy Pau. 31.3+ 49.5 22.1+ 31.1 53.4+ 79.1 4.9 5 |Allopauropus (A.) tenuis Remy Pau. 27.7+ 35.7 22.6+ 29.0 50.3+ 63.6 4.6 6 |Allopauropus (A.) ovalis Sch. Pau. 22.9+ 18.0 24.8+ 22.3 47.7 + 34.2 4.4 7 |Allopauropus (D.) manausensis Sch. Pau. 23.5 + 17.0 23.1+ 10.9 46.6+ 29.0 4.3 8 |Allopauropus (A.) bicorniculus Sch. Pau. 10.6+ 11.1 33.5 + 24.4 44.1+ 29.2 4.1 9 |Hemipauropus piriformis Sch. Pau. 24.3+ 18.9 EB ea! Vilas) 34.2+ 29.5 3.1 10 |Allopauropus (D.) adisi Sch. Pau. 15.4 + 13.0 13.8+ 9.4 29.2+ 20.7 2.7 11 |Polypauropus tropicus Sch. Pol. 16.2+ 23.1 12.0 + 19.2 28.2+ 41.7 2.6 12 |Allopauropus (D.) anomoios Sch. Pau. 91+ 13.5 12.8+ 14.6 21.9+ 27.0 2.0 13 |Allopauropus (D.) ieenus Sch. Pau. 10.6+ 12.7 72x 14.8 17.8+ 25.7 1.6 14 |Hemipauropus amazonicus Sch. Pau. 99+ 7.9 6.5+ 10.7 164+ 17.1 1.5 15 |Allopauropus (D.) brachypodus Sch. Pau. 6.4+ 10.1 3.4+ 6.0 9.8+ 11.8 0.9 16 |Allopauropus (A.) uncinatus Sch. Pau. 48+ 8.2 44+ 7.2 9.2+ 13.7 0.8 17 |Allopauropus (D.) petiolatus Sch. Pau. 3.6+ 4.8 5.2+ 10.1 8.8+ 12.0 0.8 18 |Allopauropus (D.) irmgardae Sch. Pau. 3.2+ 5.1 44+ 58 76+ 8.2 0.7 19 |Allopauropus (D.) pedicellus Sch. Pau. 56+ 5.8 18+ 2.7 74+ 5.8 0.7 20 |Allopauropus (D.) junki Sch. Pau. 5.6+ 7.9 10+ 1.9 6.6+ 8.9 0.6 21 |Allopauropus (D.) neotropicus Sch. Pau. 12+ 22 26+ 2.9 38+ 4.6 0.4 22 | Allopauropus (D.) tohoius Sch. Pau. 18+ 43 10+ 2.2 28+ 6.0 0.3 23 | Scleropauropus tarumamirimi Sch. Scl. 12+ 0.4 16+ 0.4 28+ 5.1 0.3 | 24 |Allopauropus (D.) sinuosus Sch. Pau. 14+ 2.9 04+ 0.4 18+ 3.6 0.2 | 25 | Polypauropoides biclaviger Sch. Pol. 06+ 1.2 10+ 2.4 16+ 29 0.1 | 26 |Polypauropoides foliolus Sch. Pol. 06+ 1.4 104 2.2 16+ 29 0.1 | 27 Polypauropoides unisetus Sch. Pol. - 16+ 29 16+ 29 0.1 | 28 |Allopauropus (D.) dischides Sch. Pau. 08+ 1.7 06+ 1.4 14+ 24 0.1 29 |Polypauropus latebricolus Sch. Pol. 10+ 2.2 02+ 0.7 12+ 2.2 0.1 30 |Cauvetauropus (P.) biglobulosus Sch. Pau. 04+ 1.4 0.4+ 1.0 0.8+ 1.7 0.1 31 |Allopauropus (D.) acer Sch. Pau. 0.2+ 0.7 0.6+ 1.0 08+ 1.4 0.1 32 |Polypauropoides naous Sch. Pol. 0.2+ 0.7 04+ 1.4 06+ 2.2 0.1 | 33 |Allopauropus (D.) aius Sch. Pau. - 04+ 1.0 04+ 1.0 >0.1 | 34 | Scleropauropus beritae Sch. Scl. 04+ 1.4 - 04+ 1.4 >0.1 | 35 | Scleropauropus rimatus Sch. Scl. 0.2+ 0.7 0.2+ 0.7 04+ 1.4 >0.1 | 36 |Allopauropus (D.) campinaranicus Sch. Pau. 02+ 0.7 - 02+ 0.7 >0.1 | 37 |Allopauropus (D.) korynetes Sch. Pau. - 02+ 0.7 0.2+ 0.7 >0.1 | 38 |Allopauropus (D.) hylaios Sch. Pau. - 0.2+ 0.7 02+ 0.7 >0.1 39 |Allopauropus (D.) mirimus Sch. Pau. - 0.2+ 0.7 02+ 07 >0.1 40 |Allopauropus (D.) tenuilobatus Sch. Pau. - 0.2+ 0.7 0.2+ 0.7 >0.1 41 |Scleropauropus fissus Sch. le 0.2+ 0.7 0.2+ 07 Total 543.9+ 22.4 | 1085.7+ 42.3 | 100.0 Subf. = Subfamily, Pau. = Pauropodinae, Pol. = Polypauropodinae, Scl. = Scleropauropodinae (48.9% versus 51.1%; Fig. 3: top; Table 1). Species representing the genus Hemi- pauropus (Fig. 3) were somewhat more abundant in the organic layer (0-3.5 cm: 67.5% of the total catch; Table 1), whereas representatives of the genus Polypauropoides (Table 1) were more abundant in the mineral subsoil (3.5-7 cm: 74.1%). Out of the total 4] pauropod species, 3 species were only obtained from the organic soil layer and 5 560 JOACHIM ADIS ET AL. solely from the mineral subsoil (Table 3, Fig. 3: bottom). Three species of the genus Allopaurus represented more than half (57.4%) of the total pauropods collected in 0-7 cm soil depth, 27% in the organic soil layer (0-3.5cm) and 30% in the mineralic subsoil (3.5-7cm; Fig. 3: bottom, Table 1). Allpauropus rodriguesi and A. bicornutus were eudominant, representing 33.0% (4,296 ind./m2) and 18.9% (2,460 ind./m?) of the total catch, respectively (Fig. 4, Table 1). Adult specimens predominated in both species (61.9 and 83.3%, respectively), regardless of the soil layer investigated (Fig. 4), whereas subadults were rare (<2%). Juvenile specimens were more abundant in A. rodriguesi (37.3% of the total catch) compared to A. bicornutus (14.9%). A. rodriguesi was likewise abundant in both soil layers (Table 1, Fig. 4), despite its large body size (<1.41mm) and relatively long legs. A. bicornutus was smaller (<0.83mm) and somewhat more abundant in the mineral subsoil (3.5-7cm: 62% of the total catch) compared to the organic soil layer (0-3.5cm). The next two dominant species, A. proximus and A. dundoensis (Fig. 4, Table 1), represented 5.5% (719 ind./m?) and 4.9% (638 ind./m?) of the total catch, respectively. Adults dominated in both soil layers (Fig. 4; total catch: 84.3 and 75.1%, respectively). Subadults were more abundant in A. dundoensis (10.8%) compared to A. proximus (1.7%). A. proximus was more abundant in the organic soil layer (80.6%) compared to the mineral subsoil (Fig. 4: 3.5-7 cm). No males were obtained in A. dundoensis, and only two males each in A. proximus and A. tenuis. Only in Allopauropus proximus was the monthly abundance of adults (250 females, 2 males) in both soil layers positively correlated with maximum soil temperatures (25.1-29.3 °C; average: 25.3 + 0.7 °C), i.e. catch numbers increased with increasing temperatures (total catch: r=+0.85055 (0-3.5cm), r=+0.81230 (3.5-7cm), p<0.01; n=12). For all other species no significant correlations were obtained (p<0.01) between the monthly abundances in the soil and the local abiotic factors. Although the total catches of dominant species obtained during the dry season were generally higher compared to the rainy season, there was no distinct reproductive period because juveniles as well as adults occurred throughout the year (cf. Figs 5-7 for Allopauropus rodriguesi, A. bicornutus, A. ovalis, A. manauensis, A. bicornulus and Hemipauropus piriformis). These results indicate a plurivoltine mode of life. DISCUSSION ABUNDANCE AND VERTICAL DISTRIBUTION Further comparable data on the abundance and vertical distribution of the soil fauna in four different upland forest types of Central Amazonia were obtained by Adis and collaborators (cf. ADIS er al. 1987a,b, 1989a,b; RIBEIRO 1994). Arthropods were collected to a soil depth of 14 cm during rainy and dry seasons and extracted with the Kempson method as described above. Between 75% and 92% of all arthropods were found to inhabit the top 7 cm when Acari and Collembola were included in the total catch numbers and 69%-84% when they were omitted. Data on Pauropoda are now available: PAUROPODA CENTRAL AMAZONIA 561 „am“ 6 496 6 528 13 024 % of total catch 0-3.5 3.5 - 7.0 0-7.0 Soil Depth (cm) FS Allopauropus mw Polypauropus 8 Hemipauropus Dothers (3 gen.) Spp: 38 1 100 > = 80 60 40 LL oO + fa oO Koi + je) — — O > 20 0 - 3.5 35-720: 0-7.0 Soil Depth (cm) = Allopauropus rodriguesi IM Allopauropus E Allopauropus bicornutus bicorniculus m Allopauropus proximus [others Fic. 3 Distribution of dominant genera (top) and species (bottom) of Pauropoda in the soil according to depth in a secondary upland forest near Manaus. (Total catch = 100%). Samples taken monthly at 0-3.5 and 3.5-7 cm depths between September 1982 and August 1983. (N = total number of specimens; Spp. = total number of species). ‘(susundads Jo JaquINU [810] = N) 'E86] IsNsny pur 786] Joquiaydasg usomjoqg Syydap wd /-S’E pur Ç'€-0 12 Ayyuou uoye) sopdurs ‘(22001 = 919 [EIOL) ‘sneuryy wou 159107 puejdn Arepuooes è ur sodes jeyusw -dojaaap Jo asvyusosod pur ‘dop 0) SuIp10998 [IOs oy} ur sısusopunp "y pue snunxoad ‘y ‘sninusooig "y ısond11po4a sndosnpdo]]y Jo uonnqrnsiq i II] x synpe=\y suynpegns=S solueAn[=f soinyewu!O syinpe m synpe=\wy suynpegns=S soelueAn[=f soeinyewwio synpe m (wo) deg |10S (wo) ydeg |I0S OL-G'E ge-0 OL-G'E ge-0 0 L 1 0 02 02 à à (©) = op © a et i a Db = 3 PE È Ors Gi ia] I er n 08 A < 8€9 = „W/N 617 = „W/N s 09 001 FL = sisusopunp snınedojjy snwixoid sndoisnedo||y e) (wo) deg IloS (wo) deg II0S OL-S'E G£-0 OL-GE ge-0 N 0 4 0 0% o x 2 08%, (%8'L) S Ov È le) (©) (%6'vL) f à i (%£'£8) v 09 È Sri 962 + = „W/N a 09h 2 = „W/N 08 i 03 È isenbupos sndoinedol|y sninu1091q Sninedol]|y PAUROPODA CENTRAL AMAZONIA 563 Allopauropus rodriguesi Allopauropus bicornutus OM Adults Ofemales Sa Adults O females males Mi males : 600 E E Pi Pi © © 400 £ E (©) (©) oO oO o a 200 0 SONDJFMAM J JA S ON D J F M A M J J A 20 15 Subadults Ofemales Subadults O females males males a 15 a E E 10 © Pi © 10 © £ £ (©) © à. à 5 D n 5 0 S ON D J F M A M J J A S ON DJ FM AM J JA 800 120 Juveniles m3 Juveniles m3 a 100 ale , 600 i mc E E 80 Pi OD S 400 © 60 = È $ 8 Q © 40 D 200 2 20 0 0 S ON DJJ FMAM J J A S ON D|IJ F M A M J JA <— 1982 >|<——__ 1983 — — 1982 >| 1983 ———__> Fic. 5 Temporal occurrence of developmental stages of Allopauropus rodriguesi and A. bicornutus in the soil (N/m° in 0-7 cm depth) of a secondary upland forest near Manaus. Monthly samples Sag between September 1982 and August 1983. (Classification of juveniles according to pairs of legs). 564 Allopauropus ovalis 190 Adults Ofemales males 80 E =) 60 Cc ® E 2 40 Q D 20 0 SMOMNMDONMIUIMENMPAMMENENU EVA 20 Subadults Ofemales Mi males = 15 E d © 10 E [©] oO LI D 5 0 SRONNMDEJNENMIRMAMMANINU A 60 Juveniles m3 O5 De E d (= oO IE [e] ® Q D SMOMINMID) (STE RMITATMTI J A OOS | 1989 => JOACHIM ADIS ET AL. Allopauropus manauensis 60 Adults O females Emales SE 40 d = © € 3 2 20 0 S$ © N 1D VJ FoMeAeMe nA 10 Subadults Ofemales @ males = d © 5 E (©) ® Qa n 0 so N D J EM A IM eA 60 Juveniles g3 D5 Specimens / m? SOND|IJFMAMJJA PR eg — FIG. 6 Temporal occurrence of developmental stages of Allopauropus ovalis and A. manauensis in the soil (N/m? in 0-7 cm depth) of a secondary upland forest near Manaus. Monthly samples taken between September 1982 and August 1983. (Classification of juveniles according to pairs of legs). PAUROPODA CENTRAL AMAZONIA 565 Allopauropus bicorniculus Hemipauropus piriformis Adults DO females Adults males Ofemales Mmales € E 40 © O Cc (= v oO E E © o 3 ae 0 SON DJFM AM J J A S ON DJF M A M J J A 30 30 Subadults Ofemales Subadults Dfemales males males © a ce Cc oO ® E E È 3 5" È 0 S ON D J F MA M J J A S ON D J F M AM J JA 60 60 Juveniles m3 Juveniles B3 D5 05 E36 B6 E 40 E 40 2 2 fb) E È È È i 20 2 20 0 0 S ON DiJ F M A M J J A S ON DJJ F M A M J JA — 1982 >|<_——_ 1983 — > <— 1982 —>|<—___ 1983. > Fic. 7 Temporal occurrence of developmental stages of Allopauropus bicorniculus and Hemipauropus piriformis in the soil (N/m? in 0-7 cm depth) of a secondary upland forest near Manaus. Monthly samples taken between September 1982 and August 1983. (Classification of juveniles according to pairs of legs). 566 JOACHIM ADIS ET AL. One study was conducted during October 1985 and April 1986 in a secondary upland forest on yellow latosol (Capoeira) at the INPA campus in Manaus (03°08'S, 60°01'W), where the vegetation was previously cut but unburned (ADIs et al. 1987a,b). Pauropods represented 1.1-1.6% of the total arthropods when Acari and Collembola were included (dry season: 50.448 ind./m?, rainy season: 63.850 ind./m?) and 4.6-5.8% when they were omitted from the total catch numbers (dry season: 11.934 ind./m?, rainy season: 17.886 ind./m?). The total abundance of pauropods in 0-14 cm soil depth was higher during the rainy season (1,035 ind/m?) compared to the dry season (549 ind./m?). However, during both rainy and dry seasons percentages of the total catch in the top 7 cm were similar (61.9% and 66.7%, respectively). Another study was made from August 1990 to February 1991 in a secondary upland forest on yellow latosol (Capoeira), about 50 km north of Manaus (03°34'S, 60°60'W), where the vegetation was previously cut and burned (RIBEIRO 1994). Pauro- pods represented 2.5-3.5% of the total arthropods when Acari and Collembola were included (dry season (Aug.-Oct.): 29.064 ind./m?, rainy season (Dec.-Febr.): 19.793 ind./m?) and 9.4-9.8% when they were omitted from the total catch numbers (dry season: 7.720 ind./m2, rainy season: 7.176 ind./m?). The total abundance of pauropods (0-14 cm soil depth) during the rainy and dry seasons was similar (703 versus 727 ind./m2). This was also true for the percentages obtained in the top 7 cm: 42.5% of the total catch during the rainy season and 55.0% during the dry season. A third study was made during March and August 1988 in a primary forest on whitesand soil (Campinarana), about 45 km north of Manaus (02°35'S, 60°01’W; ADIS et al. 1989a,b). Pauropods represented 2.8-4.4% of the total arthropods when Acari and Collembola were included (dry season: 57.703 ind./m?, rainy season: 74.255 ind./m?) and 13.8-18.1% when they were omitted from the total catch numbers (dry season: 14.119 ind./m?, rainy season: 15.023 ind./m?). The total abundance of pauropods (0-14 cm soil depth) during the rainy and dry seasons was similar and high (2,079 versus 2,550 ind./m?). This was also true for the percentages obtained in the top 7 cm: 60.4% of the total catch during the rainy season and 61.9% during the dry season. The fourth study was conducted during April and October 1987 in the primary upland forest on yellow latosol at Reserva Ducke (ADIs er al., unpubl.). Pauropods represented 1.3-1.7% of the total arthropods when Acari and Collembola were included (dry season: 38.727 ind./m?, rainy season: 25.905 ind./m?) and 4.1-4.3% when they were omitted from the total catch numbers (dry season: 11.742 ind./m?, rainy season: 11.005 ind./m?). The total abundance of pauropods (0-14 cm soil depth) during the rainy and dry seasons was similar and low (448 versus 505 ind./m?). No significant difference (Chi-square test) was found between the percentages obtained in the top 7 cm during the rainy season (62.4% of the total catch) and the dry season (84.4%). The lack of pauropods in samples from the ground and arboreal photo-eclector indicates that representatives of this group are rarely active on or above the soil surface in Central Amazonian upland forests. This conclusion is supported by two other studies in a primary upland forest of the Reserva Ducke near Manaus, in which apparently no pauropods were collected in 20 baited pitfall traps and in one or more ground photo- PAUROPODA CENTRAL AMAZONIA 567 eclectors during a sample period of 12 months (Apis et al. 1996, PENNY & ARIAS 1982) or from tree crowns by fogging canopies with pyrethrum (ADIS et al. 1997c). The depth to which pauropods occur in the soil of the Central Amazonian upland forests is unknown. Our studies in various forest types near Manaus (see above) revealed their presence to a soil depth of 14 cm. SERAFINO & MERINO (1978) detected them until 20 cm soil depth in a forest and in plantations in Costa Rica. Pauropoda are, however, found to occur to a depth of at least 75 cm in other parts of the world (SCHELLER 1974). In Allopauropus proximus the vertical distribution was influenced by soil temperatures. In the soil of man-made pastures (0-14 cm depth) adjacent to upland forests in Central Amazonia, pauropods were only found below 7 cm soil depth. One reason might be the low humidity and high temperature of the soil around noon, particularly during the dry season (ADIS & FRANKLIN, unpubl.). The small size of pauropods as well as inefficient sampling methods might explain their "absence" or low abundances in other studies on the Neotropical arthropod fauna in 0-30 cm soil depth (e.g. HARADA & BANDEIRA 1994a,b; HUTHER 1985; MACAMBIRA 1997). PARTHENOGENESIS Two pauropod species of this study had strikingly low sex ratios: Allopauropus proximus (2 males and 250 females; sex ratio 0.008) as well as A. tenuis (2 males and 211 females; sex ratio 0.009). A. dundoensis is only known from 200 females from the secondary upland forest under study. The two first mentioned species have often been reported from tropical and subtropical areas. Although hundreds of specimens were sexed, no males at all were found in A. proximus and a few ones only in A. tenuis (SCHELLER 1970). A. dundoensis has not often been collected outside Amazonia. It was previously known from about the same latitude in tropical West Africa only (REMY 1955, 1956; SCHELLER 1975) from where 29 specimens were sexed, 28 of them females and | male. The latter specimen was subadult and somewhat deviated from the type material. Even if it should belong to A. dundoensis, males seem to be extremely rare. In conclusion, males in A. proximus and A. tenuis seem to be very rare and in A. dundoensis they may not exist at all. As far as known A. proximus is parthenogenetic, A. tenuis probably so, at least in parts of its range, and A. dundoensis might be. In two studies of north African pauropods, REMy (1947) established a striking scarcity of males in two Allopauropus species. In most of the European countries where he had found Allopauropus vulgaris (Hansen) the sex ratio was about 1.0 but in an Algerian material (99 specimens sexed adult and subadults) he could not find males at all (REMY 1947). He found a similar situation in A. productus Silvestri: from Europe, he reported 54 males and 70 females (sex ratio 0.77); in Algerian and Tunisian material with 155 sexed adults and subadults, there were 54 males and 101 females (sex ratio 0.53); from Morocco only 4 specimens among 73 adults and subadults were males (sex ratio 0.05) (REMy 1952). Similar sex ratios have later also been found in these species (material from Corsica: LECLERC 1953; from the Canary Islands: SCHELLER 1979) but also in other species e.g. in A. gracilis (Hansen) from Canada (sex ratio 0.32) 568 JOACHIM ADIS ET AL. (SCHELLER 1984) as well as in A. cuenoti (Remy) from Switzerland (sex ratio 0.07) (SCHELLER 1976) and the Canary Islands with no males at all (SCHELLER 1979). It seems evident that populations with both males and females as well as pure female popu- lations exist in the Pauropoda. Since in millipedes males are much rarer than females and in some species both bisexual and thelytokous forms appear (ENGHOFF 1978), probably due to the presence of bacteria (ADIS et al. 1999), it is plausible that the same types of reproduction have arisen in the Pauropoda too. The sex ratio is generally low in the Pauropoda and there may be a decrease of sexual reproduction in some species (SCHELLER 1970). In many species from all parts of the world males are unknown, but in most cases the numbers of adult and subadult specimens recorded are too low to decide whether they are parthenogenetic or not. Up to now, no laboratory studies have been performed to show parthenogenesis in the Pauropoda. Besides spanandry there is one more factor indicating the occurrence of parthe- nogenetic reproduction in some species. The genus Allopauropus has two subgenera, the nominate subgenus with 3 pairs of setae on the pygidial sternum and 9 pairs of legs in the adult stage, and Decapauropus with 2 pairs of setae on the pygidial sternum and two adult stages, one with 9 pairs of legs and an additional one with 10 pairs of legs. All the specimens of the latter stage, which have been sexed, have been females and this stage seems to be more frequent in areas with an unfavourable environment than in areas with moist and not too cold climates. It might be an adaptation to a partheno- genetic form of reproduction, occurring in areas with in some way harsh climates. Under good environmental conditions in the tropics the additional stage with 10 pairs of legs is very rare, Amazonia included. The development of spanandry or even a thelytokous reproduction in Allopau- ropus seems to have arisen particularly in the subgenus Decapauropus. It 1s note- worthy, however, that in the material from the secondary upland forest near Manaus, a species in Allopauropus s. str., A. (A.) dundoensis, seems to have such a low sex ratio that parthenogenetic reproduction might be conceivable. ACKNOWLEDGEMENTS We wish to acknowledge the valuable support received from PD Dr. W.J. Junk, Head of the Tropical Ecology Working Group at the Max-Planck-Institute (MPI) for Limnology in Plön/Germany. Dr. William Shear, Hampden-Sydney College/USA and Dr. John Lewis, Halse, Taunton/England are thanked for valuable comments on the manuscript. Berit Hansen (MPI Plön) is thanked for making the drawings. Dr. Johann Bauer, MPI for Biochemistry (Martinsried /Germany) assisted us with literature. REFERENCES Apis, J. 1987. Extraction of arthropods from Neotropical soils with a modified Kempson apparatus. Journal of Tropical Ecology 3 (2): 131-138. Apis, J. 1992. Überlebensstrategien terrestrischer Invertebraten in Überschwmmungswäldern Zentralamazoniens. 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SCHELLER, U. 1997. Pauropoda from upland and inundation forests in Central Amazonia, Brazil (Myriapoda, Pauropoda: Millotauropodidae, Pauropodidae). Amazoniana 14 (3/4): 223- 300. SERAFINO, A. & MERINO, J.F. 1978. Poblaciones de microartröpodos en diferentes suelos de Costa Rica. Revista de Biologia Tropical 26 (1): 139-151. REVUE SUISSE DE ZOOLOGIE 106 (3): 571-580; septembre 1999 Freshwater sponges (Porifera, Spongillidae) from the Lake of Geneva, Switzerland Renata MANCONI! & Ruth DESQUEYROUX-FAUNDEZ? 'Dipartimento di Zoologia e Antropologia Biologica dell'Università, Via Muroni 25, I-07100 Sassari, Italy. 2Muséum d'histoire naturelle, Case postale 6434, CH-1211 Geneva 6, Switzerland. Freshwater sponges (Porifera, Spongillidae) from the Lake of Geneva, Switzerland. - A census of the sponge fauna in shallow-waters of the Lake of Geneva, Switzerland, displayed the presence of scattered populations of Spongilla lacustris (Linnaeus, 1758), Eunapius fragilis (Leidy, 1851), Ephydatia fluviatilis (Linnaeus, 1758), and Ephydatia muelleri (Lieberkühn, 1855). Spongilla lacustris was the most frequent species, either alone or associated with other species it was present in numerous sites on a variety of substrata along the entire shoreline. The morphological analysis of speci- mens of S. lacustris of the Lake of Geneva compared with the type material of S. lacustris and Spongilla helvetica, shows a considerable variation of skeletal and gemmular characters. Therefore, the synonymy of S. helvetica with S. lacustris is confirmed. The study also demonstrates the existence of a rich and diversified sponge fauna in the Lake of Geneva. Key-words: Spongilla lacustris - Spongilla helvetica - Lake of Geneva (Lac Léman) - freshwater sponges - distribution - morphology - biodiversity. INTRODUCTION Five species of spongillids are actually known from Switzerland (see map): Spongilla lacustris (Linnaeus, 1758), Eunapius fragilis (Leidy, 1851), Ephydatia fluvia- tilis (Linnaeus, 1758), Ephydatia muelleri (Lieberkhün, 1855) and Trochospongilla horrida Weltner, 1893. Only four of them are presently known to occur in the Lake of Geneva. Trochospongilla horrida has only been reported for the Lake of Neuchatel. ANNANDALE (1909) described Spongilla helvetica, from the Lake of Geneva with no other records from other localities. This species was considered doubtful by some authors (cf. ARNDT 1926; GEE 1931, 1932). JORGENSEN (1946), referring only to the description of thin-walled gemmule, considered S. helvetica conspecific with S. lacus- tris. PENNEY & RACEK (1968) suggested a close affinity between Spongilla helvetica and Spongilla lacustris, but in view of the paucity of the material examined by them, they considered advisable to retain Spongilla helvetica as an available species. After the study of a paratype of the species, EZCURRA DE DRAGO (1972), considered Spongilla helvetica conspecific with Spongilla lacustris. Manuscript accepted 02.03.1999 572 R. MANCONI & R. DESQUEYROUX-FAUNDEZ In this work, we studied type material and different populations of Spongilla lacustris, paying particular attention to the morphology of the gemmules, by making a SEM analysis of the gemmule structure of the neotype of S. lacustris (Figs 5-10) and of the paratype of S. helvetica (Figs 11-16). At the same time, the spicular morphology and micrometry of specimens formerly identified as Spongilla lacustris and Spongilla helvetica from different localities in Switzerland, were analysed (Table 1). In addition, we include here the results of a species census for the Spongillidae represented at different localities of the Lakes of Geneva and Neuchatel (see map). MATERIALS AND METHODS ABBREVIATIONS IZUG = Istituto di Zoologia dell’ Universita di Genova; LEBA = Laboratoire d'Ecologie et de Biologie Aquatique, Université de Genève; MHNG! = Muséum d'histoire naturelle, Genève; NNHML = (Rijksmuseum van Natuurlijke Historie, Leiden), National Natur Historisch Museum, Leiden; SEM = Scanning Electro-Microscopy; US = Universita di Sassari; MURST! = Ministero italiano dell'Universita' e della Ricerca Scientifica e Tecnologica; INTERREG-UE! = European interregional (Sardinia-Corsica) project, European Com- munity. COLLECTIONS Freshwater sponges were collected by C. Vaucher (MHNG) in 1977 and 1993 in the Lake of Neuchatel and by B. Lods-Crozet and D. Cambin (LEBA) from 1989 to 1993 in the Lake of Geneva. Samples were obtained either by hand or by scuba diving at 31 sites along the lake borders (LODS-CROZET 1999: in the press). All specimens were identified to species level and registered for the MHNG collections. A morphological and micrometrical study of spicules and gemmules was made with Spongilla lacustris type material and with the collected samples of the Lakes of Geneva, Neuchatel and River Rhône. Measurements of mega-, micro-and gemmulo-scleres (in um), were obtained with an compound microscope connected to a camera lucida and a Graphtec Digitizer KD 4300 (Table 1). A minimum of 20 spicules were measured. The following additional material was studied for comparison: Spongilla helve- tica Annandale, 1909, holotype and paratype, reg. n° MHNG-18970 INVE and MHNG- 18980 INVE, Morges, Lake of Geneva, Switzerland, Collection Ostroga, respectively, 10.1900 and 07.1902; Spongilla lacustris, neotype designation by PENNEY & RACEK (1968), reg. n° 1053, NNHML, Vlaardingschwaart, the Netherlands, coll. D. v. d. Kuyl, 09. 1941; Spongilla lacustris, det. Ezcurra de Drago, reg. n° FW 22 IZUG and FW 123 First author's research was supported by grants of MHNG, MURS, INTERREG-UE. SPONGILLIDS FROM THE LAKE OF GENEVA 573 IZUG, Bolle de Magadino, Switzerland. Sizes of mega-, micro-and gemmulo-scleres (in um), and presence of gemmular cages and foramina were investigated (Table 2). Spicule slides (prepared as indicated by PRONZATO & MANCONI 1989) and the SEM study were made at the MHNG. A Zeiss Digital Scan Microscope DSM 940 was used, after sputtering the samples with gold-palladium. RESULTS HABITAT AND DISTRIBUTION Several populations (see map) of Spongilla lacustris, Eunapius fragilis, Ephy- datia fluviatilis, and E. muelleri are present in the Lake of Geneva; Trochospongilla horrida was collected only at one locality of the Lake of Neuchatel. Ephydatia fluvia- tilis, and E. muelleri were rare along the northern side of the Lake Neuchatel (two of seven localities studied), but present on both sides of the Lake of Geneva. Eunapius fragilis, was present in five of nine localities of the Lake of Geneva. Spongilla lacustris the most common species, was widespread along the entire shoreline of the lakes of Geneva and Neuchatel (48% of 39 sampled sites), it was very common at depths ranging from 0.2 to 3.5 m, either alone (73%) or associated with Eunapius fragilis (20%) or E. fluviatilis (13%). Spongilla lacustris was present, on a variety of substrata, e.g. wood piles or floating wood (34%), pebbles and boulders (20%), rocks and cliffs (18%), metallic piles (10%), brick walls and piles (8%), shells (8%) and concrete piles (2%). All the represented species are seasonal and active specimens and were found only from spring to autumn. DESCRIPTION Spongilla lacustris is encrusting to massive or cushion-shape to branching. Consistence, fragile, soft. Dimensions, 2 to 70 cm. Colour, white to yellow-orange or light green; light brown or brown after fixation. Surface uneven, hispid, with irregularly scattered small oscula, covered by a fine, fragile ectosomal membrane. Skeleton: Ectosome, short brushes of spicules, perpendicular to the surface, issued from the ends of primary longitudinal fibres. Choanosome, anisotropic network of pauci- to multi-spicular longitudinal primary fibres with a strong sheath of spongin, connected by transversal secondary pauci-spicular fibres. Skeletal meshes irregular in size and form. Spicules (Table 1, Figs 1-4) Megascleres oxeas, bent, fusiform with acerate or blunt, rarely hastate or conical, apices, 90-308 mm length by 2-8 um in diameter. Some of them are slightly spined and associated to gemmules. Microscleres microxeas, bent, fusiform, strongly microspined, 25-178 um length by 2-8 um in diameter; they may be abundant or rare. Gemmules: 98-789 um in diameter, present in active sponges from March to November, in dense clusters or irregularly scattered within the skeletal framework. Gemmular shape ranged from spherical to oval and sometimes a spicular cage composed by megascleres is present. Two gemmular types were observed: either armed (with gemmuloscleres) or naked (without gemmuloscleres). Both types sometimes can 574 R. MANCONI & R. DESQUEYROUX-FAUNDEZ Neuchatel O Yverdon Lausanne Ouchy 00) Lac Léman (Lake of Geneva) Spongillidae species presently collected at different localities in Switzerland, Lakes of Geneva and Neuchatel. 1 = Spongilla lacustris; 2 = Eunapius fragilis; 3 = Ephydatia fluviatilis; 4 = E. muelleri, 5 = Trochospongilla horrida. Fics 1-4. Spongilla lacustris (L., 1759), neotype, NNML 1053, Vlaardingschwaart, the Netherlands and Spongilla helvetica Annandale, 1909, holotype, MHNG 18970 INVE, paratype, MHNG 18980 INVE, Morges, 1909. Fig. 1. Megascleres: smooth to slightly spined oxeas and its apical morphology. Figs la, c, g: S. helvetica, paratype: figs 1b, d, e, f, h, i, j, k: S. lacustris, neotype. Fig. 2. Microscleres: bent oxeas strongly microspined. Figs 2a, b, S. helvetica, paratype; figs 2c, d, S. lacustris, neotype. Fig. 3. Gemmuloscleres, variably bent oxeas, strongly microspined. Figs 3b, c, S. helvetica, holotype; figs 3a, d, S. lacustris, neotype. Fig. 4. S. helve- tica, paratype, cage of spicules on the surface of a gemmule. Scales: Figs la - c, If, g, 2c, 4 = 100 um: Figs Id, e, h - k, 2a, b, d, 3a - d= 10 um. DID SPONGILLIDS FROM THE LAKE OF GENEVA ____{ 576 R. MANCONI & R. DESQUEYROUX-FAUNDEZ Fics 5-10. Spongilla lacustris (L., 1759) neotype NNML 1053, Vlaardingschwaart, the Netherlands. Fig. 5, thick-walled gemmule, foramen with collar. Fig. 6, lateral view of the same type of foramen. Fig. 7, plate-like, enlarged frontal view of the of the same type of foramen. Fig. 8, transversal section of the theca in a thick-walled gemmule. Fig. 9, foramen without collar of a thin-walled gemmule. Fig. 10, transversal section of the theca in a thin-walled gemmule. Scales: Fig. 5 = 100 um, Figs 6, 7, 9 = 20 um., Fig. 8 = 10 um; Fig. 10 = 2 um. SPONGILLIDS FROM THE LAKE OF GENEVA STE) Fics 11-16. Spongilla helvetica Annandale, 1909, paratype. Fig. 11. Gemmule (thick-walled) with a single foramen. Fig. 12, thick gemmular theca with well developed pneumatic layer and inner coat with 5 sub-layers. Fig. 13, gemmule (thin-walled) with two foramina. Fig. 14, thin gemmular theca pneumatic layer and inner coat with 4 sub-layers. Fig. 15, gemmule with two foramina, lateral view of the second foramen, left on top. Fig. 16, gemmular theca of a gemmule with two foramina; well developed pneumatic layer and inner coat with 5 sub-layers. Scales: Figs I, 13; 15 = 100/um, Figs 12,14, 16=5 pm. 578 R. MANCONI & R. DESQUEYROUX-FAUNDEZ TABLE | Mega, micro and gemmuloscleres; gemmule micrometries of Spongilla lacustris (Linnaeus, 1759) specimens presently collected from different localities of Switzerland. Micrometries refer to minimum, mean and maximum range of spicular length and diameter, and gemmular diameter, in ‚um: Locality Megascleres Microscleres Ganmullosclares Gemmilss River Rhöne 163-227-290 49- 69- 90 44-54-64 5 469- 565 3.0-4.8-6.7 1-2.1-4 2-2.4-4 Lake of Geneva 155-187-218 43-62-81 41-63-84 362-473-584 2.5-4.0-5.5 1-1.7-8 1-2.1-4 Lake of Neuchatel 198-207-216 32-46-59 57-70-82 428-502-575 6.2-6.7-7.2 2-2.6-4 2-2.4-3.4 TABLE 2 Mega, micro and gemmuloscleres; gemmule micrometries; presence or not of cage and foramina number in: Spongilla lacustris (Linnaeus, 1758): neotype NNML 1053, Vlaardingschwaart, Netherland; S. lacustris in literature; S. lacustris, FW 22 IZUG, FW 123 IZUG, Bolle de Magadino, coll. Ezcurra de Drago; S. lacustris (= Spongilla helvetica Annandale, 1909), holo- type, MHNG 18970 INVE, paratype, MHNG 18980 INVE, Morges, 1909: (*) From literature or (**) presently remeasured. Micrometries refer to minimum mean ‘and maximum range of spicular length and diameter, and gemmular diameter, in um. N = spicular number, stated in each case. Species name Mega- Micro- Gemmulo- Gemmules Cage Fora- scleres scleres scleres mina Spongilla lacustris | **Neotype 122-273 47-102 35-127 383-636 + I 1053 NNML x 3-16 x 3-7 X 3-7 19510 n=50 n= 100 n=22 *Penney & Racek, 1968 200-350 70-130 80-130 500-800 ? ? x 6-18 x2 = 8 x 3-10 **RW 22 IZUG 160-247 not 44-77 A — I Bolle de Magadino x 6-16 measured x 4-7 n= 22 n=20 **RW 123 [ZUG 160-247 not { 30-77 A — lort Bolle de Magadino x 6-16 measured x 3-6 122 122 *Bolle de Magadino 150-350 50-90 30-150 A — — Ezcurra de Drago, 1972 x 8-30 x 3-8 x 5-10 *S. helvetica not not not not A lor+ Annandale, 1909 measured measured measured measured **Holotype MHNG 18970 INVE 147-241-317 28-49 51-77 578-801 + lor+ Morges x 5-10-18 xo? 3 i = 2510) in n=4 m= INI **Paratype MHNG 18980 INVE 115-294 44-68 35-99 495-810 + lor+ Morges x 4-14 x 3-5 el M=2I0 n=20 n = 30 ni ils SPONGILLIDS FROM THE LAKE OF GENEVA 579 be found in the same specimen. The coat of gemmules ranges from thick-walled with three layers (outer, pneumatic, inner layer) to thin-walled if the coat is composed only of compact spongin in a variable number (3-7) of sub-layers, in total corresponding to the inner layer of the thick walled gemmulae. A distinct horny external layer is present in some gemmules with a thick-walled theca. Thick-walled gemmules bear generally a single foramen and an irregular outer surface due to the presence of gemmuloscleres. Some gemmules with several foramina and a well developed pneumatic layer were observed. Thin-walled gemmules show generally a smooth surface and 1-6 foramina. Slightly elevated foramina without, or with a normal or plate-like collar, are irregularly present. Gemmuloscleres: oxeas to strongyles, straight or variably curved up to a ring- shape and densely microspined, with blunt and sometimes less microspined apices. Abundant, rare or absent, according to the gemmular type concerned, 21 - 130 um, in length by 1-4 m in diameter. Gemmuloscleres are external to or variably embedded in the gemmular theca. DISCUSSION The study of S. lacustris from the Lake of Geneva illustrates the great variability of the taxonomic important characters, both, within the same individual or between individuals (MANCONI & PRONZATO 1991; RICCIARDI & REISWIG 1993). Concerning the spicular and gemmular characters, we observed that thick and thin-walled thecae in gemmules can occur in the same specimen, but some specimen have one gemmular type only. The presence may, depend on the season and hence on the life cycle phase. The cage of megascleres around thin-walled gemmules is a very variable trait. We regularly observed a cage in S. lacustris, but it was rarely recorded by other authors (RETZER 1883; VEJDOVSKY 1887; ANNANDALE 1909; JORGENSEN 1946). The comparative analysis of shape and size of S. lacustris recently collected from the Lake of Geneva, the neotype of S. /acustris and the type material of S. helve- tica, revealed that both diagnostic characters at the spicular level (Figs 1-4), and gemmular morphology (Figs 5-16) fall within the range of S. lacustris Auct. This confirmed the morphological variability of Spongilla lacustris (POTTS 1887; VEIDOVSKY 1887; PENNEY & RACEK 1968; GILBERT & SIMPSON 1976). The morphological analysis by SEM of gemmules (Figs 5-10, 11-16) of S. helvetica type material and S. lacustris neotype, shows that the former species is indistinguishable from the later. The study of intraindividual variability (Table 1) confirms here the extremely high phenotypic plasticity of gemmular morphology. The same remark is valid for several varieties and ‘morphospecies' similar to S. lacustris, that were described from Europe and North America based on gemmular traits (POTTS 1887; VEIDOVSKY 1887; PENNEY & RACEK 1968; EZCURRA DE DRAGO 1972). In addition, our study confirmes that the extreme variability of the gemmular structure in S. lacustris can be attributed to both the position of gemmules within the sponge body and to their life cycle phases as suggested by other authors (JORGENSEN 1946; GILBERT & SIMPSON 1976). 580 R. MANCONI & R. DESQUEYROUX-FAUNDEZ ACKNOWLEDGEMENTS The following persons have greatly contributed to our work and we are grateful: G. Haartog, NNHML; Tiziana Cubeddu, USC; Peter Schuchert, Jean Wiest, Claude Ratton and G. Roth, MHNG. Important collections in the Lake of Geneva, made by B. Lods-Crozet and D. Cambin (LEBA), were at the origin of this work. REFERENCES ANNANDALE, N. 1909. Description d'une nouvelle espece d'éponge d'eau douce du Lac de Genève. Revue suisse de Zoologie 17: 367-369, pl. 9. ARNDT, W. 1926. Die Spongillidenfauna Europas. Archiv fiir Hydrobiologie 17: 337-365. EZCURRA DE Draco, I. 1972. Contributo alla conoscenza delle spugne d'acqua dolce d'Italia (Porifera, Spongillidae). Memorie dell'Istituto Italiano di Idrobiologia Marco de Marchi 29: 109-127. GEE, N.G. 1931. A contribution towards an alphabetical list of the known fresh-water sponges. Peking Natural History Bulletin 5 : 31-52. GEE, N.G. 1932. The known fresh-water sponges. Peking Natural History Bulletin 6: 25-51. GILBERT, J.J. & Simpson, T.L. 1976. Gemmule Polymorphism in the Freshwater Sponge Spongilla lacustris. Archiv fiir Hydrobiologie 78(2): 268-277. JORGENSEN, C.B. 1946. On the gemmules of Spongilla lacustris Auct. together with some remarks on the taxonomy of the species. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjobenhavn 109: 69-79. Lops-CROzET, B. 1999. (In the press). Invertébrés benthiques de la zone littorale du Léman. 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Die deutschen Süsswasserschwämme. /naugural Dissertation, Universität Tiibingen, 1-30. RICCIARDI A. & REISWIG, H.M. 1993. Freshwater sponges (Porifera, Spongillidae) of eastern Canada: taxonomy, distribution, and ecology. Canadian Journal of Zoology 71: 665-682. SCHELLENBERG, U. & BURLA, H. 1972. Schwämme (Porifera) im Zürichsee. Revue suisse de Zoologie 79 (suppl.): 319-332. VEJDOVSKY, F. 1887. Diagnosis of the European Spongillidae, pp. 172-180. In: Ports, E. (ed.). Contributions towards a synopsis of the American forms of freshwater Sponges with descriptions of those named by other authors and from all parts of the world. Proceedings of the Academy of Natural Sciences of Philadelphia. REVUE SUISSE DE ZOOLOGIE, 106 (3): 581-590; septembre 1999 Endoparasitic helminths of fishes in three Alpine lakes in France and Switzerland Vladimira HANZELOVA!, Tomas SCHOLZ?, Daniel GERDEAUX? & Alain de CHAMBRIER* i ! Parasitological Institute, Slovak Academy of Sciences, Hlinkova 3, 040 01 Koëice, Slovak Republic. ? Institute of Parasitology, Academy of Sciences of the Czech Republic, Brani$ovskä 31, 370 05 Ceské Budéjovice, Czech Republic. 3 Institute of Hydrobiology, INRA, BP 511, F-74 203 Thonon-les-Bains, France. 4 Natural History Museum, P.O. Box 6434, CH-1211 Geneva 6, Switzerland. Endoparasitic helminths of fishes in three Alpine lakes in France and Switzerland. — A systematic survey of endoparasitic helminths of 9 species of fishes from three Alpine lakes, Annecy, Bourget (both France) and Geneva Lake (Switzerland and France), is presented. Eleven species of intestinal helminths (2 Digenea, 7 Cestoda, 1 Nematoda and 1 Acantho- cephala) were found in 272 of 372 (i.e. 73%) fish species examined. The dominant parasites were pseudophyllidean tapeworms Eubothrium crassum (Bloch, 1779) and E. salvelini (Schrank, 1790) in lake trout (Salmo trutta m. lacustris) and Arctic char (Salvelinus alpinus) respectively, and the proteocephalidean Proteocephalus longicollis (Zeder, 1800) in whitefish (Coregonus lavaretus). Key-words: Endohelminths — fishes — Alpine lakes — France — Switzerland. INTRODUCTION Little attention has been paid to studies on the parasite fauna of Alpine lakes despite their importance for fishery and sport fishing (GERDEAUX et al. 1995). This is also valid for three large lakes in the western part of the Alps, namely Annecy and Bourget lakes in France and Geneva Lake situated between Switzerland and France. Up to now, only a few data on the helminths parasitizing fishes in these lakes have been published, with most records related to Geneva Lake. ZSCHOKKE (1884) reported several taxa of /chthyotaenia (= Proteocephalus) from fishes in Geneva Lake but most are now considered invalid (see SCHOLZ & HANZELOVA 1998). JOYEUX & BAER (1936) listed several helminth species from this lake, including tapeworms Cya- thocephalus truncatus, Eubothrium crassum, E. salvelini, Triaenophorus crassus, Pro- teocephalus dubius (= P. percae), P. longicollis, P. fallax, P. neglectus, P. salmoni- sumblae (three latter species synonymized with P. longicollis — SCHOLZ & HANZELOVA Manuscript accepted 02.03.1999 582 V. HANZELOVA, T. SCHOLZ, D. GERDEAUX & A. DE CHAMBRIER 1998), and P. torulosus. DOBY & JARECKA (1964), JARECKA & Dosy (1965), and MORANDI & PONTON (1989) studied the biology of Proteocephalus tapeworms (most probably P. longicollis) in whitefish (Coregonus lavaretus) from Geneva Lake and in experimentally infected copepods. However, except for a study by GERDEAUX ef al. (1995), who reported the occurrence of the pseudophyllidean cestode Eubothrium salvelini in Arctic char (Salve- linus alpinus), almost no information exists about species composition and richness of the helminth fauna of fishes in Annecy and Bourget lakes. During studies on the ecology of fishes in Annecy and Bourget lakes, heavy infections of salmonids, in particular of Arctic char with E. salvelini, were recorded since December 1992 (D. Gerdeaux — unpublished data; GERDEAUX et al. 1995). In order to provide more detailed information about helminth parasites, samples of fishes from the above mentioned lakes and Geneva Lake were taken between 1992 and 1998. In this paper, a survey of endoparasitic helminths found is presented. MATERIALS AND METHODS Samples were taken from three peri-alpine lakes in the western part of the Alps, namely Annecy Lake (Lac d’Annecy), Bourget Lake (Lac du Bourget; both Haute Savoie, France) and Geneva Lake (Lac Léman; Switzerland and France). All lakes are of glacial origin and they have steep banks and a relatively flat bottom. Annecy Lake is an oligotrophic lake with a surface of 2,800 hectares and maximum depth 65 metres. It is highly productive in fish, with highest natural production of coregone (whitefish) and stocking production of Arctic char. Bourget Lake is an eutrophic lake with a surface of 4,500 hectares and maximum depth 145 m. It is rich mainly in perch (Perca fluviatilis), lake trout (Salmo trutta m. lacustris) and cyprinid fishes; sustained by stocking yields of whitefish and Arctic char are rather low. Geneva Lake is a mesotrophic lake situated between Switzerland and France; it has a TABLE | Number of fish examined Species Annecy Bourget Geneva Total Esox lucius 2 4 6 Salmo trutta m. lacustris 8 4 16 Salvelinus alpinus 34 16 2 62 Coregonus lavaretus 40 14 21 75 Lota lota 18 7 22 Alburnus alburnus I 1 Rutilus rutilus 10 40 50 Tinca tinca 3 3 Perca fluviatilis 48 4 76 128 Total 160 45 158 363 ENDOPARASITIC HELMINTHS OF FISHES IN THREE ALPINE LAKES 583 surface area of 58,000 hectares and its maximum depth reaches 310 m. The fish community is halfway between the two others; perch, salmonids and cyprinids are of equal importance (GERDEAUX 1990). A total of 363 fishes of 9 species was examined in September 1997 and June 1998 (Table 1). Fish were caught by local fishermen using gill nets or by angling. Digestive tracts were removed, placed into boxes with ice, transported to the laboratory and examined the same or next day. The worms found were fixed by hot 4% formalin and nematodes by hot 4% solution of formalin and saline, identified and counted. Voucher specimens have been deposited in the helminthological collections of the Natural History Museums in Paris and Geneva; and at the Institute of Parasitology, Ceské Budèjovice. Some additional records from the collection (INVE) of the Natural History Museum of Geneva are included. RESULTS A total of 11 species of endoparasitic helminths was found: 2 species of trematodes (Digenea), 7 species of tapeworms (Cestoda), | nematode (Nematoda) and | spiny-headed worm (Acanthocephala). DIGENEA 1. Asymphylodora tincae (Modeer, 1790) Host: tench (Tinca tinca). Site: intestine. Locality: Bourget, June 1998 (infected 2 of 3 fish examined; intensity of infection 3-296). Comments: This is a frequent parasite specific to tench, occurring in the Palearctic Region (BYKHOVSKAYA-PAVLOVSKAYA & KULAKOVA 1987). The life cycle of this trematode includes only one intermediate host, a lymnaeid snail, in which tailless cercariae (cercariaeum) develops. The fish acquire infection after consuming the snail harbouring cercariaea (NASINCOVA & SCHOLZ 1994). 2. Bunodera luciopercae (Miiller, 1776) Host: perch (Perca fluviatilis). Site: intestine. Locality: Annecy, September 1997, June 1998 (12/48; 41.8; 3-100). Comments: B. luciopercae is a typical intestinal parasite of percid fishes in Europe and the ex-USSR, with perch representing the most suitable host (BYKHOVS- KAYA-PAVLOVSKAYA & KULAKOVA 1987). In some regions, the parasite exhibits marked seasonal patterns in the occurrence and maturation (see CHUBB 1979 for review). The life cycle includes 2 intermediate hosts (YAMAGUTI 1975). 584 V. HANZELOVA, T. SCHOLZ, D. GERDEAUX & A. DE CHAMBRIER CESTODA 3. Caryophyllaeus laticeps (Pallas, 1781) Host: roach (Rutilus rutilus). Site: intestine. Locality: Annecy, September 1997, (one of 10 fish examined infected with 1 juvenile cestode). Comments: It seems that this tapeworm, which is rather common parasite of cyprinid fishes in Europe and Eurasia (DUBININA 1987), is quite rare in the lakes studied. In Geneva Lake, it was not present in any of 40 roach examined in June 1995 and September 1997. Oligochaetes (Tubificidae) serve as intermediate hosts of this parasite (SEKUTOWITCZ 1932). 4. Eubothrium crassum (Bloch, 1779) Hosts: lake trout (Salmo trutta m. lacustris); perch as paratenic host. Site: intestine. Localities: lake trout: Annecy, September 1997, June 1998 (8/8; 47.8; 4-172; of these, 358 adult specimens and 24 juvenile ones); Bourget, September 1997, June 1998 (4/4; 59.8; 35-89; 239 adults and 48 juvenile tapeworms); perch: Annecy, September 1997, June 1998 (4/48; 1.5; 1-3 juvenile specimens); Bourget, September 1997 (1/4; more than 50 juveniles). Comments: E. crassum is a common parasite of salmonoid fishes, widely distributed in the Holarctic with two biological races, one freshwater and one marine (KENNEDY 1978a,b). In Europe, E. crassum occurs from the Arctic part of Norway to southern France and Switzerland (VIK 1963: KENNEDY 1978b, 1996). JOYEUX & BAER (1936) reported E. crassum from trout in Geneva Lake. Trout (Salmo trutta) is consi- dered to be preferred host of the cestode, with prevalence often reaching 70-100% (WOOTTEN 1972). In the present study, E. crassum has not been recorded in the lake trout from Geneva lake (four hosts examinated). However, this parasite is common in the lake trout as indicated by recent samples from the Geneva Museum collection: Chens, 16.01.1992, 24.04.1998; Genthod 22-23.01.1992; Prangins, 08.03.1992; Her- mance, 31.03.1993; Tougues, 14.01.1996; Coppet, 18.01.1998. Prevalence: 8/9. In Annecy and Bourget lakes adult tapeworms were found exclusively in lake trout, whereas juveniles with the beginning of segmentation occurred in trout and perch. On the basis of the presence of juvenile tapeworms in prey fish as perch, ruff (Gymnocephalus cernuus) or three-spined stickleback (Gasterosteus aculeatus), it was supposed that two intermediate hosts are involved in the life cycle of this cestode (ROSEN 1918; JOYEUX & BAER 1936; AKHMEROV 1962). However, field data (VIK 1963; WOOTTEN 1972; REIMER 1984) indicate that there is only one obligatory intermediate host, a copepod, and prey fish serve as paratenic hosts (KENNEDY 1978a,b). JoyEUX & BAER (1936) reported common occurrence of juvenile Eubothrium tapeworms from several fish species in Geneva Lake. Although these juvenile cestodes could not be identified, it can be assumed that they might belong to E. crassum. ENDOPARASITIC HELMINTHS OF FISHES IN THREE ALPINE LAKES 585 5. Eubothrium salvelini Schrank, 1790 Host: Arctic char (Salvelinus alpinus). Site: intestine. Localities: Annecy, September 1997, June 1998 (34/34; 19.6; 1-60; 458 adult specimens and 209 juveniles); Bourget, September 1997, June 1998 (16/16; 19.4; 2-39; 283 adults and 27 juveniles). Comments: E. salvelini is considered a specific parasite of Arctic char in Europe and Eurasia, with its distribution largely coincident with that of its host (KENNEDY 1978a,b). It is a fairly common parasite of Arctic char and it has been reported from this fish species in Switzerland and France (JOYEUX & BAER 1936; GERDEAUX er al. 1995), Austria (RYDLO 1970, 1985; KRITSCHER 1991), Germany (REIMER 1984; HOFFMANN er al. 1986a; ENGELHARDT & MIRLE 1993) as well as in northwestern Europe (NYBELIN 1922; WooTTEN 1972; KENNEDY 1978a,b; KRISTOFFERSEN 1995; KNUDSEN et al. 1997). Similar to the present study, high prevalence (70-100%) and intensity of infection (up to 60 worms) were reported in other localities (SOBECKA & PIASECKI 1993), including Alpine lakes (REIMER 1984). In North America it occurs also in other salmonid fishes as Oncorhynchus (SMITH 1973; KENNEDY 1978a,b; Boyce 1979; MuzzaL 1995). It has been demonstrated experimentally (BOYCE 1974) that copepods serve as the only intermediate hosts. Field data suggest that the infection level and its variation among individual localities depend largely upon proportion of planktonic copepods in the food of Arctic char (REIMER 1984). In contrast to E. crassum, no juvenile tape- worms were found in other fish than Arctic char during the present study. 6. Proteocephalus longicollis (Zeder, 1800) Host: whitefish (Coregonus lavaretus). Site: intestine. Localities: Annecy, September 1997, June 1998 (36/40; 113 adults and a great number of juvenile cestodes, with intensity of infection ranging from 17 to more than 1000 (precise number not counted); Bourget, September 1997, June 1998 (11/14; 7.8; 1-30; 60 adults and 26 juveniles); Geneva Lake, September 1997 (8/21; 14,9; 2-58; 36 adults and 233 juveniles). Comments: It has been demonstrated that P. longicollis (syns. P. exiguus, P. fallax and P. neglectus) is the only Proteocephalus species occurring in salmonid and coregonid fishes in Europe (SCHOLZ & HANZELOVA 1998). It is a highly variable species occurrung in a wide range of fishes including brown, rainbow and brook trouts and whitefish (HANZELOVA er al. 1995, 1996; SCHOLZ & HANZELOVA 1998). In the lakes investigated, adult worms were found exclusively in the intestine of whitefish whereas Arctic char and lake trout were free of P. longicollis infection. However, a fry of Arctic char and lake trout from a hatchery on Annecy lake were heavily infected with P. longi- collis (D. Gerdeaux - unpublished data). JOYEUX & BAER (1936) also reported P. longicollis from lake trout, Arctic char and whitefish in Geneva Lake. Proteocephalus tapeworms, most probably conspecific with P. longicollis, were found in whitefish from Geneva Lake by DoBy & JARECKA (1964), and MORANDI & PONTON (1989). 586 V. HANZELOVA, T. SCHOLZ, D. GERDEAUX & A. DE CHAMBRIER Intermediate hosts of P. longicollis are planktonic copepods (see SCHOLZ 1999 for review). In Geneva Lake, Cyclops strenuus and C. abyssorum have been reported as intermediate hosts of Proteocephalus sp. (most probably P. longicollis) by JARECKA & DoBy (1965). In Annecy lake, Cyclops prealpinus serves as dominant intermediate host of this parasite (V. Hanzelova - unpublished data). A high number of metacestodes was found in whitefish in all three lakes. Much lower number of adult worms in these fish indicates that only a very small proportion of juvenile worms reaches maturity (about 0.5-1% estimated for P. filicollis — HOPKINS 1959). 7. Proteocephalus percae (Miiller, 1780) Hosts: perch (Perca fluviatilis); burbot (Lota lota) as alternative host. Site: intestine. Localities: perch: Annecy, September 1997, June 1998 (27/48; 6.4; 1-53); Bourget, September 1997, June 1998 (2/4; 4 and more than 100 juveniles); Geneva, June 1996 (20/76; 2.2; 1-14); burbot: Annecy, June 1998 (2/18; 2.5; 1-4). Comments: P. percae 1s a common parasite of perch, reported also from other percid fish in the Palearctic (SCHOLZ & HANZELOVA 1998). In lakes under consi- deration, it occurs fairly frequently in perch. Burbot represents only an alternative, probably postcyclic host of P. percae. It is known that predatory fishes as pike, trout, eel or burbot may harbour P. percae tapeworms and serve as alternative (paradefinitive, postcyclic or accidental) hosts (SCHOLZ 1998). Life cycle of this cestode involves a copepod intermediate hosts (WOOTTEN 1974). 8. Proteocephalus torulosus (Batsch, 1786) Host: bleak (Alburnus alburnus). Site: intestine. Locality: Geneva, June 1995 (1/1; 1). Comments: This is a typical parasite of cypriniform fishes (Cyprinidae and Cobitidae), occurring in Europe, Asia and western part of North America (SCHOLZ & HANZELOVA 1998). It occurs preferably in riverine ecosystems rather than in lakes. Only one bleak was examined in this study and it is possible that P. torulosus occurs more frequently in the lakes studied. JOYEUX & BAER (1936) also reported bleak as the definitive host of this tapeworm from Geneva Lake. Intermediate hosts of P. torulosus are planktonic copepods, in particular Cyclops spp. (WAGNER 1917; SCHOLZ 1993). 9. Triaenophorus nodulosus (Pallas, 1781) Hosts: pike (Esox lucius); perch as the second intermediate host. Sites: intestine (adult tapeworm); liver (plerocercoids in perch). Localities: pike: Annecy, June 1998 (1/2; 12); perch: Annecy, September 1997, June 1998 (20/48; 3.5: 1-10); Geneva, June 1995 (number not counted). Comments: T. nodulosus is a specific parasite of pike in the Palearctic Region (KUPERMAN 1973). It frequently occurs in Alpine lakes in Austria and Germany (RYDLO ENDOPARASITIC HELMINTHS OF FISHES IN THREE ALPINE LAKES 587 1985; HOFFMANN et al. 1986b). The life cycle of this cestode includes two intermediate hosts, a copepod in which a procercoid develops, and a fish where a plerocercoid is encysted in the liver. Infection level with adults and plerocercoids in fishes from the lakes studied is fairly low compared to other localities (see CHUBB 1982 for review). NEMATODA 10. Cammallanus lacustris (Zoega, 1776) Hosts: perch (Perca fluviatilis), burbot (Lota lota). Site: intestine. Localities: perch: Annecy, September 1997, June 1998 (36/48; 8.1; 1-44); Bourget, September 1997, June 1998 (4/4; 11.3; 4-21); Geneva, June 1995 (number not counted); burbot: Annecy, June 1998 (5/18; 3.2; 1-6). Comments: C. lacustris is a quite common parasite of predatory fish as pike, perch or burbot, widely distributed in the Palearctic Region (MORAVEC 1994). In Annecy and Bourget lakes, perch is apparently the principal host of this nematode, which develops through planktonic copepods (MORAVEC 1994). The absence of nema- todes in pike from the lakes concerned may be related to the low number of fish examined. ACANTHOCEPHALA 11. Acanthocephalus lucii (Miiller, 1780) Hosts: perch (Perca fluviatilis), pike (Esox lucius), burbot (Lota lota). Site: Intestine. Localities: perch: Annecy, September 1997, June 1998 (31/48; 9.6; 1-65); Bour- get, September 1997, June 1998 (3/4; 2.0; 1-4); Geneva, June 1995 (number not coun- ted); pike: Annecy, June 1998 (1/2; 6); burbot: Annecy, June 1998 (5/18; 3.2; 1-6). Comments: A. lucii 1s a very frequent parasite occurring in a wide spectrum of fishes of different families in the Palearctic Region (BAUER 1987). In the lakes studied, it forms an important part the helminth fauna of perch and occurs commonly also in burbot and pike. Intermediate host of A. /ucii is the isopod Asellus aquaticus (KOMAROVA 1950). DISCUSSION A total of 11 species of endohelminths was found in fishes from the three Alpine lakes studied. The most characteristic feature of these helminth communities is a high number (7) of cestode species, which represents the dominant part of the helminth fauna in all localities. Prevalence of E. salvelini, E. crassum and P. longicollis in Arctic char, lake trout and whitefish, respectively, reached 90-100%, and intensity of infection was also considerably high. Common occurrence of Eubothrium tapeworms and P. longicollis have also been reported from other lakes in the Alps and from northern Europe (KENNEDY 1978a,b; REIMER 1984; BRISTOW & BERLAND 1991; KRISTOFFERSEN 1995). 588 V. HANZELOVA, T. SCHOLZ, D. GERDEAUX & A. DE CHAMBRIER Considering existing literature data on the helminth fauna of fishes from the lakes studied, it is worth of mention that two other tapeworms, reported by JOYEUX & BAER (1936) from Geneva Lake, have not been found in this study. Instead of Triaeno- phorus crassus, T. nodulosus has been found. It is possible that former species is rather rare in pike population. The absence of Cyathocephalus truncatus, in our samples, a parasite which was reported from trout, whitefish and burbot by JOYEUX & BAER (1936), is difficult to explain. 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Occurence and distribution of Eubothrium crassum (Bloch 1779) and E. salvelini (Schrank 1790) (Cestoda) in Norway, with notes on their life cycles. Nytt Magazin for Zoologi, Oslo 11: 47-73. WAGNER, O. 1917. Uber den Entwicklungsgang und Bau einer Fischtaenie (Ichthyotaenia torulosus Batsch). Jenaische Zeitschrift fiir Naturwissenschaft 55: 1-66. WOOTTEN, R. 1972. Occurrence of Eubothrium crassum (Bloch, 1779) (Cestoda: Pseudo- phyllidea) in brown trout Salmo trutta L., and rainbow trout S. gairdneri Richardson, 1836, from Hanningfield Reservoir, Essex. Journal of Helminthology 46: 327-339. Wootten, R. 1974. Studies on the life history and development of Proteocephalus percae (Miiller) (Cestoda: Proteocephalidae). Journal of Helminthology 48: 269-281. YAMAGUTI, S. 1975. A Synoptical Review of Life Histories of Digenetic Trematodes of Vertebrates. Keigaku Publishing Co., Tokyo, 590 pp. ZSCHOKKE, F. 1884. Recherches sur l’organisation et la distribution zoologique des vers parasites des poissons d’eau douce. Archives de Biologie, Paris 5: 153-241. REVUE SUISSE DE ZOOLOGIE 106 (3): 591-598; septembre 1999 Un nouveau genre et une nouvelle espèce de scorpion d'Egypte, appartenant a la famille des Buthidae Simon Wilson R. LOURENCO Laboratoire de Zoologie (Arthropodes), Muséum national d’ Histoire naturelle, 61, rue de Buffon, F-75005 Paris, France; e-mail: arachne @ mnhn.fr New genus and new species of scorpion from Egypt, belonging to the family Buthidae Simon. - The new genus Egyptobuthus and the new species Egyptobuthus vaissadei belonging to the Buthidae family are described from Egypt. This new genus is probably an enclaved endemic element, isolated from the typical saharian and perisaharian faunas. Its dis- covery represents an interesting contribution to the knowledge of the distri- butional patterns and affinities of some elements of the African and Northern African faunas. Key-words: Scorpion - Buthidae - new genus - new species - Egypt - endemism. INTRODUCTION Dans l’ensemble de l’ordre Scorpiones, des 15 familles reconnues à l’heure actuelle (LOURENÇO 1998a, b), celle des Buthidae Simon est de loin la plus importante, tant pour le nombre d’especes, que par le nombre de genres reconnus. SissoM (1990) sıgnale un total de 50 genres valables au sein de la famille des Buthidae. Ce nombre cependant ne cesse de croitre, et depuis la synthese présentée par Sissom, plus d’une dizaine de nouveaux genres de Buthidae ont été créés (e.g. KOVARIK 1995, 1996, 1997a, b, 1998; LoURENCO 1995a, 1996a, 1997a, b, 1998c; LOURENCO & VACHON 1995, 1997). Il est vrai qu’une relative majorité des nouveaux taxa génériques créés corres- pondent a des scorpions pouvant étre classés comme des micro-buthidés. Ceci est le cas des Microcharmus Lourengo et Pseudouroplectes Lourenco de Madagascar ou encore Himalayotityobuthus Lourenço de l'Himalaya. Dans ces cas précis, leur petite taille peut expliquer leur découverte tardive, qui n’a pu être possible qu’à travers l’utilisation des techniques de collecte et d’extraction plus sophistiquées telles celles de Winkler, ou de lavage de sols, méthodes rarement utilisées pour la collecte des scorpions. Par contre, la découverte de certains nouveaux genres de taille standard (3 à 6 cm), pourrait s'expliquer soit par: (1) l’extr&me rareté des individus d’une population donnée, soit (ii) une meilleure résolution des caractères génériques valables en parallèle avec une connaissance plus précise de l’ensemble des groupes génériques de la famille donnée. Manuscrit accepté le 09.02.1999 592 WILSON R. LOURENCO Bical Habitus d’ Egyptobuthus vaissadei, holotype femelle. NOUVEAU GENRE DE SCORPION D’EGYPTE 593 L’étude d’une petite collection comprenant entre autres quelques scorpions d’Egypte nous a permis de déceler un curieux scorpion appartenant a un nouveau genre et a une espece nouvelle. Ceci est d’autant plus interessant si on tient compte du fait que la faune scorpionique des pays d’ Afrique du Nord a été intensivement étudiée par divers auteurs (POCOCK 1895; SIMON 1910), et en particulier par VACHON (1952). Egyptobuthus n. gen. Espece type: Egyptobuthus vaissadei n. sp. Diagnose Scorpions de petite taille; longueur totale des adultes d’environ 30 mm. La morphologie générale rappelle en partie celle des genres Butheoloides Hirst et Uro- plectes Peters. Granulation générale faiblement marquée. Carènes des Pédipalpes, corps et metasoma moyennement ou faiblement marquées; les dorsales des anneaux II a IV du metasoma avec un granule postérieur bien développé en forme de mamelon. Peignes avec fulcres; dent interne normale. Sternum pentagonal. Eperons tibiaux présents sur les pattes III et IV; éperons basitarsaux présents à la quatrième patte, mais réduits sur la troisième. Chélicères avec la dentition des Buthidae (VACHON 1963); dents basales du doigt mobile tres discretes; dent distale externe plus petite que la distale interne. Tranchant des doigts mobiles des mains des pédipalpes avec 12 séries de granules; les doigts sont tres courbés comme ceux observés dans le genre Darchenia Vachon. Trichobothriotaxie du type A avec la disposition « pour les trichobothries de la face dorsale du fémur des pédipalpes (VACHON 1973, 1975). Position du nouveau genre au sein de la famille des Buthidae Le nouveau genre Egyptobuthus présente quelques caracteres proches de ceux retrouvés chez d’autres genres africains tels Uroplectes et Darchenia, mais globalement l’ensemble de ses caractères l’isolent au sein des Buthidae. Son apparence générale présente quelques similitudes avec le genre Butheoloides, cependant ce dernier est classé parmi les «micro-buthidés». Leurs points communs sont une anatomie générale tres voisine, et la présence d’un sternum a forme pentagonale. Ce dernier caractere les place parmi les exceptions au sein des Buthidae, qui globalement présentent un sternum triangulaire. Egyptobuthus ne peut pas cependant étre considéré comme un genre véritablement associé a Butheoloides. Ce dernier, faisant partie intégrale des «micro- buthidés» est en réalité associé à d’autres genres africains ou asiatiques tels Orthochirus Karsch, Butheolus Simon ou Karasbergia Hewitt. Ils ont tous en commun une structure cuticulaire non-granulée mais ponctuée sur le metasoma; ces ponctuations présentant en effet des structures sensorielles spécifiques (Fet in litt. Lourenco obs. pers.). Le genre Egyptobuthus ne présente pas ce type de structure sur la cuticule ds anneaux du meta- soma, mais plutöt une structure plus classique, avec une granulation peu importante. Les affinités exactes du nouveau genre restent à préciser comme cela est le cas pour d’autres genres rares de Buthidae tel Birulatus Vachon, également connu uniquement de la zone de transition entre |’ Afrique et le Moyen Orient (VACHON 1973; LOURENÇO sous presse). 594 WILSON R. LOURENCO Fics 2 à 7. Egyptobuthus vaissadei, holotype femelle. 2. Opercule génital et peignes. 3. Tranchant du doigt mobile des pédipalpes. 4. Chelicere avec la dentition caractéristique. 5. Sternum. 6. Fémur avec la disposition a des trichobothries dorsales. 7. Extrémité de la patte IV, avec les éperons tibial et basitarsaux. NOUVEAU GENRE DE SCORPION D’ EGYPTE 595 En ce qui concerne la répartition géographique respective des deux genres, Butheoloides apparait comme un élément typiquement péri-saharien. Sans reprendre ici la totalité des considérations biogéographiques sur le genre Butheoloides (LOURENCO 1995b, 1996b), il me parait utile de les résumer brievement. Lors de sa création par Hirst le genre Butheoloides n était connu que par la seule espèce B. maroccanus Hirst, habitant les flancs des montagnes de |’ Atlas dans la région sud de Marrakech au Maroc, donc a l’extrémité Nord-occidentale de |’ Afrique. Par la suite, avec la découverte de deux nouvelles especes (Butheoloides milloti Vachon, des falaises de Bandiagara au Mali et Butheoloides monodi Vachon, décrite de l’ouest du Sénégal) la répartition du genre s’est élargie vers le Sud et l'Est du continent africain, affichant déja une répartition du type péri-saharienne. Bien plus tard avec la description de Butheoloides annieae Lourengo de la Côte d'Ivoire, de Butheoloides wilsoni Lourenço du Burkina Faso, de Butheoloides polisi Lourengo d’Ethiopie et de Butheo- loides hirsti Lourenco du Soudan, ce modele de répartition s’est de plus en plus confirmé. Egyptobuthus, au contraire, semble étre un élément isolé dans une des zones refuges présentes sur les régions arides du Sahara et du Moyen Orient, comme c’est le cas du genre Birulatus, dont la distribution est limitée a des régions refuges en Jordanie et Israel (LOURENCO sous presse). Egyptobuthus vaissadei n. sp. Holotype femelle: Egypte, région nord du Sinaï, proche ds côtes de la Mer Rouge, II- 1997 (H.-W. Herrmann) collecté avec un exemplaire de Buthus occitanus Amoreux). Déposé au Museum d'histoire naturelle de Genève. Etymologie: Le nom spécifique est créé en hommage a M. Alain Vaissade, Conseiller administratif de la Ville de Geneve. Description Coloration générale jaunatre avec des taches brunatres. Plaque prosomienne jaunatre avec des taches brunatres, surtout dans la région antérieure. Mesosoma jaunatre avec deux bandes longitudinales brunätres qui commencent dans la partie distale de la plaque prosomienne et se prolongent sur le mesosoma, s’élargissant vers l’arriere. Anneaux metasomaux I a III jaunätres; IV et V plus foncés, jaune-rougeatre; présence de taches brunatres longitudinales sur tous les anneaux. Vésicule jaunatre avec des taches brunatres très estompées; aiguillon avec la base jaunatre et à l’extrémité rou- geätre. Peignes, opercule génital, sternum, hanches et processus maxillaire ocre-jaune. Pattes jaunatres avec des taches brunätres tres estompées; pédipalpes jaunätres, avec des nuances de taches brunätres. Cheliceres jaunätres avec la présence d’un trame brun- rougeatre faiblement marqué. Morphologie. Prosoma: front de la plaque prosomienne avec une concavité très peu profonde, presque droit; tubercule oculaire très antérieur par rapport au centre de la plaque prosomienne; yeux médians séparés par plus d’un diamètre oculaire; trois paires d’yeux latéraux; la première paire composée d’yeux plus petits. Carènes pratiquement absentes; granulations réduites à un minimum; tégument presque lisse. Mesosoma: 596 WILSON R. LOURENCO FIG. 8 Carte avec la distribution géographique péri-saharienne du genre Butheoloides (cercles noirs) et la localité-type d’ Egyptobuthus vaissadei (étoile). tergites pratiquement lisses, comme pour la plaque prosomienne. Carene axiale présente sur les tergites I à VI, mais faiblement marquée; pratiquement absente sur le VIIème ou quatre autres carenes sont faiblement représentées. Metasoma: anneaux arrondis avec des granulations et des carenes peu marquées, surtout sur les trois premiers; carenes dorsales des anneaux II à IV avec une grosse épine postérieure, en forme de mamelon. Vésicule pratiquement lisse; aiguillon plus court que la vésicule, très incurvé, pourvu d’une épine sous-aiguillonnaire peu développée, également en forme de mamelon. Sternum pentagonal. Sternites à stigmates petits et aplatis, linéaires. Peignes avec 17-16 dents; fulcres présents; lame basilaire intermédiaire non-dilatée. Pédipalpes: fémur a 5 carenes peu marquees; tibia et pince tres lisses, avec des carenes presque imper- ceptibles. Douze series de granules sur le tranchant des doigts mobiles. Cheliceres avec la dentition des Buthidae (VACHON 1963); les deux dents basales du doigt mobile presque fusionnées et peu perceptibles; dent distale externe plus petite que la dent distale interne. Trichobothriotaxie du type A-a (VACHON 1973, 1975), orthobothrio- taxique. Eperons tibiaux presents sur les pattes III et IV; éperons basitarsaux présents sur la patte IV et réduits sur la III. NOUVEAU GENRE DE SCORPION D’EGYPTE 597 TABLEAU I Mensurations (en mm) de l’holotype d’Egyptobuthus vaissadei Prosoma - Longueur 32 - Largeur antérieure 23 - Largeur posterieure 3,4 Anneau caudal I - Longueur 1,8 - Largeur 22 Anneau caudal V - Longueur 3,4 - Largeur DO - Hauteur 2,0 Vésicule - Largeur 2,0 - Hauteur 167 Pedipalpe - Fémur longueur 2,6 - Fémur largeur 1,0 - Tibia longueur 3,2 - Tibia largeur IS) - Pince longueur 52 - Pince largeur 1,4 - Pince hauteur 1,2 Doigt mobile - Longueur 3,3 REMERCIEMENTS Je suis très reconnaissant à M. Philippe Bouchard, Muséum national d’ Histoire naturelle, Paris pour sa contribution à la réalisation de plusieurs dessins illustrant le présent travail, au Dr Sabine Jourdan (Paris) pour la lecture du manuscrit et à M. T. Ziegler (Bonn) pour m'avoir confié à l’étude le spécimen décrit. BIBLIOGRAPHIE KOVARIK, F. 1995. Review of Scorpionida from Thailand with descriptions of Thaicharmus mahunkai gen. et sp. n. and Lychas krali sp. n. (Buthidae). Acta Societatis zoologicae Bohemicae 59: 187-207. KOVARIK, F. 1996. Baloorthochirus becvari gen. et sp. n. from Pakistan, and taxonomic position of Orthochirus luteipes (Scorpiones: buthidae). Acta Societatis zoologicae Bohemicae 60: 177-181. KOVARIK, F. 1997a. Afroisometrus gen. n. from Zimbabwe (Scorpiones: Buthidae). Acta Societatis zoologicae Bohemicae 61: 35-37. KOVARIK, F. 1997b. Results of teh Czech Biological Expedition to Iran. Part 2. Arachnida: Scorpiones, with descriptions of /ranobuthus krali gen. n. et sp. n. and Hottentotta zagrosensis sp. n. (Buthidae). Acta Societatis zoologicae Bohemicae 61: 39-52. Kovarik, F. 1998. Three new genera and species of Scorpiones (Buthidae) from Somalia. Acta Societatis zoologicae Bohemicae 62: 115-124. 598 WILSON R. LOURENCO LOURENCO, W. R. 1995a. Description de trois nouveaux genres et de quatre nouvelles especes de scorpions Buthidae de Madagascar. Bulletin du Muséum national d'Histoire naturelle, Paris, 4e sér. 17 (A-1-2): 95-106. LOURENÇO, W. R. 1995b. Considérations sur la répartition géographique du genre Butheoloides Hirst avec la description de Butheoloides wilsoni n. sp. (Scorpiones, Buthidae). Bulletin du Muséum national d’ Histoire naturelle, Paris, 4e ser. 16 (A-2-4): 475-480. LOURENÇO, W. R. 1996a. A new genus and a new species of scorpion (Buthidae) from Iran. Zoology in the Middle East 12: 93-98. LOURENÇO, W. R. 1996b. A propos de deux espèces nouvelles appartenant au genre Butheoloides Hirst (Scorpiones, Buthidae). Revue Arachnologique 11 (9): 87-94. LOURENÇO, W. R. 1997a. A new genus and new species of Buthidae scorpion from Pakistan. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 12 (155): 153- 1578 LOURENÇO, W. R. 1997b. Description of a new genus and new species of Buthidae scorpion from the Himalayas of India and Nepal with some biogeographic implications. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 12 (156): 183-188. LOURENÇO, W. R. 1998a. Panbiogéographie, les distributions disjointes et le concept de famille relictuelle chez les scorpions. Biogeographica 74 (3): 133-144. LOURENÇO, W. R. 1998b. Une nouvelle famille est nécessaire pour des microscorpions humicoles de Madagascar et d’Afrique. Comptes Rendus de l'Académie des Sciences, Paris, Sciences de la Vie 321: 845-848. LOURENCO, W. R. 1998c. Uroplectoides abyssinicus new genus and new species of scorpion (Scorpiones, Buthidae) from Ethiopia. Entomologische Mitteilungen aus dem Zoo- logischen Museum Hamburg 12 (158): 309-316. LOURENCO, W. R. (sous presse). Complementary notes on the phylogenetic position of the genus Birulatus Vachon, 1973 (Scorpiones, Buthidae) and redescription of Birulatus haasi. Zoology in the Middle East. LOURENÇO, W. R. & VACHON, M. 1995. Un nouveau genre et deux nouvelles espèces de scor- pions Buthidae d’Iran. Bulletin du Muséum national d'Histoire naturelle Paris, 4e ser. 17 (A3/4): 297-305. LOURENCO, W. R. & VACHON, M. 1997. Un nouveau genre et quatre nouvelles especes de scorpions Buthidae pour le Moyen-Orient. Zoosystema 19 (2-3): 327-336. Pocock, R. I. 1895. On the Arachnida and Myriopoda obtained by Dr. Anderson’s Collector during Mr. T. Bent’s Expedition to the Hadramaut, South Arabia; with a supplement upon the Scorpions obtained by Dr. Anderson in Egypt and Eastern Soudan. The Linnean Society's Journal-Zoology 25: 292-316. SIMON, E. 1910. Révision des Scorpions d’Egypte. Bulletin de la Société entomologique d’Egypte, Cairo: 57-87. SıssoM, W. D. 1990. Systematics, Biogeography, and Paleontology (pp. 64-160). In: PoLIS, G. A. (ed.). The Biology of Scorpions. Stanford University Press. VACHON, M. 1952. Etudes sur les scorpions. Publications de l’Institut Pasteur d’Algerie, Alger: 482 pp. VACHON, M. 1963. De l’utilité, en systematique, d’une nomenclature des dents des chélicères chez les Scorpions. Bulletin du Muséum d’Histoire naturelle, Paris, 2e ser. 35(2): 161- 166. VACHON, M. 1973. 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 Museum national d Histoire naturelle, Paris, 3e 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 des Séances de l’Académie des Sciences, Paris, ser. D, 281: 1597-1599. REVUE SUISSE DE ZOOLOGIE, 106 (3): 599-610; septembre 1999 Nuovi Anillina (Coleoptera, Carabidae: Bembidiini) del Kenya Roberta GARETTO! & Pier Mauro GIACHINO? I Via Milano 13, I-10122 Torino, Italia. 2 Museo Regionale di Scienze Naturali, Via Giolitti 36, I-10123 Torino, Italia. New species of Anillina (Coleoptera, Carabidae: Bembidiini) from Kenya. - Three new species of Anillina are described from Kenya: Micro- dipnites mahnerti n. sp. from Kirimiri forest (Embu district), Microdipnites perreti n. sp. from Taita Hills (near Kwale) and Selenodipnus embuanus n. sp. from Irangi forest (Embu district). The zoogeographical interest of these new taxa is stressed. Key-words: Coleoptera - Carabidae - Bembidiini - Anillina - new species - Kenya. Lo studio di una piccola serie di Carabidae Anillina provenienti dalle raccolte effettuate dal personale del Muséum d'Histoire Naturelle de Genève nell’ Africa orien- tale (Kenya) e gentilmente concessa in studio ad uno degli Autori (P.M.G.) dal Dr. Ivan Lobl, Conservatore del Dipartimento di Entomologia dello stesso Museo, ha permesso l’identificazione di tre specie nuove per la scienza. Le nuove specie, appartenenti ai generi Microdipnites Jeannel, 1957 e Seleno- dipnus Jeannel, 1963, costituiscono oggetto di descrizione nella presente nota. Microdipnites mahnerti n. sp. Figg. 1-5 Holotipo d Kenya, Embu district, Kirimiri forest, east of Runyenje, 1550 m, thifting of forest litter, 13.X.77, V. Mahnert et J.-L. Perret leg. (Muséum d'histoire naturelle, Genève). Paratipi: 2 d, 9 2, idem (Muséum d’histoire naturelle, Genève; Museo Regionale di Scienze Naturali, Torino; Coll. Giachino, Torino; Coll. Casale, Torino). DIAGNOSI. Un Microdipnites con caratteri perfettamente conformi alla defi- nizione del genere fornita da JEANNEL (1937, 1963): dente labiale assente, un solo arti- colo dei protarsi dilatato e profondamente asimmetrico nel d, elitre profondamente smarginate all’apice, serie ombelicata di tipo C (sensu JEANNEL 1963); edeago non arcuato, privo di restringimento basale, parameri armati di due setole apicali. M. mahnerti n. sp. differisce invece dalle due specie note del genere M. kahu- zianus (Basilewsky, 1957) e M. minutissimus (Basilewsky, 1957), oltre che per la conformazione edeagica, anche per la conformazione della smarginatura dell’apice elitrale, che si presenta molto più accentuata, e per la posizione del 7° poro della serie Manoscritto accettato il 15.02.1999 600 R. GARETTO & P. M. GIACHINO Fic. 1 Microdipnites mahnerti n. sp. Holotipo d. Habitus. Scala: 0.1 mm. NUOVI ANILLINA DEL KENYA 601 Fico. 2-5 Microdipnites mahnerti n. sp.: Holotipo d edeago in visione dorsale (2); Paratipo d zampa protoracica destra in visione dorsale (3); Paratipo d labium in visione ventrale (4); Paratipo d palpo labiale sinistro in visione ventrale (5). Scala: 0.05 mm. ombelicata. Da M. perreti n. sp., descritto oltre nella presente nota e il cui maschio è sconosciuto, differisce per la smarginatura apicale delle elitre nettamente più accen- tuata. DESCRIZIONE. Lunghezza totale (dall’apice delle mandibole all’apice delle elitre) mm 0.83 - 0.92 (dall’apice delle mandibole all’apice dell'ultimo urotergite mm 1.01- 1.14). Corpo poco allungato e relativamente tozzo, depigmentato, fulvo-testaceo con elitre e addome più chiari, giallo-testacei; tegumenti lucidi, a microscultura quasi total- mente svanita, cosparsi di pubescenza rada e breve (fig. 1). Capo robusto, quasi ipertrofico, più stretto del pronoto, anoftalmo. Antenne ispessite, nettamente moniliformi a partire dal terzo antennomero, raggiungenti, distese all'indietro, la base delle elitre. Solco clipeo-frontale indistinto; margine anteriore dell’epistoma subrettilineo. Due setole sopraorbitali per lato, abbastanza distanziate fraloro e poste su linee parallele, non convergenti all’indietro. Mandibole brevi, semplici, senza creste dorsali; labbro provvisto di 6 setole marginali anteriori; labium stretto, articolato, inerme sul margine anteriore, con epilobi molto sviluppati (fig. 4). Palpi mascellari (fig. 5) con penultimo articolo grande, ovoidale allungato, e ultimo piccolissimo, poco differenziato. Pronoto trasverso (rapporto larghezza max./ lunghezza max. = 1.38 - 1.42), allargato in avanti, ristretto alla base, con lati lungamente arcuati, non sinuati avanti la 602 R. GARETTO & P. M. GIACHINO base; non denticolati, né smarginati anteriormente agli angoli posteriori. Angoli ante- riori arrotondati, ma prominenti; posteriori ottusi e smussati al vertice. Disco appena convesso, con pubescenza corta e rada; doccia mediana molto superficiale, appena accennata. Doccia marginale larga e spianata, bruscamente ristretta presso la base; setole marginali anteriori inserite all’interno della doccia marginale, circa all'altezza del quarto anteriore; setole basali nettamente avanti agli angoli posteriori. Elitre brevi, tozze, profondamente smarginate nella zona preapicale, senza traccia di strie (esclusa la stria suturale). Disco poco convesso; tegumenti lucidi, con microscultura quasi indistinta e con pubescenza corta, molto rada ed eretta. Omeri poco indicati, arrotondati; margine post-omerale denticolato, con crenellatura molto fine ma distinta fino all’altezza del 3° poro omerale; smarginatura preapicale profonda, angolosa; apici subtriangolari, separatamente arrotondati. Doccia marginale larga ed evidente nella meta basale. Chetotassi: poro ombelicato basale grande, foveato, munito di una breve setola. Serie ombelicata di tipo C [i grandi pori setigeri della serie ombelicata corrispondono al 2°, al 6° e al 7° (sensu JEANNEL 1963)]; primi tre pori del gruppo omerale circa equidistanti fra loro, 4° poro leggermente più distanziato e inserito anteriormente alla metà dell’elitra; 5° poro posto circa al terzo apicale dell’elitra, 6° posto presso l'angolo della smarginatura apicale, 7° nettamente preapicale, posto circa alla base del triangolo apicale. Pori discali assenti; piccoli pori setigeri, in numero di due o tre, sono presenti nell’area apicale presso il 6° e 7° poro della serie ombelicata e nell’area immedia- tamente preapicale. Zampe (fig. 3) relativamente allungate, non ispessite; tarsi anteriori nei maschi con un solo tarsomero asimmetricamente dilatato. Edeago (fig. 2) piccolo, poco arcuato; base del lobo mediano profondamente incisa, biloba in visione laterale; margine ventrale del lobo mediano subrettilineo. Apice ingrossato, largamente membranoso, con lama apicale poco evidente e irrego- larmente curvata. Sacco interno munito di una lamella copulatrice piccola, poco chiti- nizzata, fortemente arcuata, quasi a ferro di cavallo, con lato concavo rivolto verso l’alto. Parameri muniti ciascuno di due setole apicali. DERIVATIO NOMINIS. Dedichiamo con piacere questa nuova specie al Dr. V. Mahnert, Direttore del Muséum d’histoire naturelle de Genève, che per primo la raccolse insieme al Dr. J.-L. Perret. DISTRIBUZIONE, ECOLOGIA. Microdipnites mahnerti n. sp. è noto al momento solo della località tipica: la foresta di Kirimiri a 1550 m di altitudine in Kenya, sito a SE del più noto Monte Kenya. Gli esemplari sono stati raccolti al vaglio in foresta mista. Microdipnites perreti n. sp. Figg. 6-10 Holotipo 9 Kenya, Taita district, Taita Hills, near Wundanyi, 1350 m, thifting in mixed forest, 2.XII.1974, V. Mahnert et J.-L. Perret leg. (Muséum d’histoire naturelle, Genève) DiaGNOSI. Un Microdipnites con caratteri perfettamente conformi alla defi- nizione del genere fornita da JEANNEL (1937, 1963): dente labiale assente, elitre smar- ginate posteriormente, serie ombelicata di tipo C (sensu JEANNEL 1963). NUOVI ANILLINA DEL KENYA FIG. 6 Microdipnites perreti n. sp.: Holotipo 2. Habitus. Scala: 0.1 mm. 604 R. GARETTO & P. M. GIACHINO M. perreti n. sp. differisce invece da M. minutissimus (Basilewsky, 1957) per la smarginatura dell’apice elitrale, che si presenta più accentuata, e per la posizione del 7° poro della serie ombelicata; da M. mahnerti n. sp. per la conformazione della smargi- natura apicale delle elitre, che si presenta molto meno accentuata e da M. kahuzianus (Basilewsky, 1957) per la diversa conformazione dell’apice elitrale e per la differente localizzazione del 6° poro della serie ombelicata. DESCRIZIONE. Lunghezza totale dell’olotipo @ (dall’apice delle mandibole all’apice delle elitre) mm 0.86 (dall’apice delle mandibole all’apice dell’ultimo uro- tergite mm 1.03). Corpo tozzo, depigmentato, fulvo-testaceo con elitre e addome piu chiari, giallo-testacei; tegumenti lucidi, a microscultura quasi totalmente svanita, cosparsi di pubescenza rada e breve (fig. 6). Capo robusto, appena piu stretto del pronoto, anoftalmo. Antenne ispessite, nettamente moniliformi a partire dal terzo antennomero, raggiungenti appena, distese all’indietro, la base del pronoto. Solco clipeo-frontale indistinto; margine anteriore dell’epistoma subrettilineo. Due setole sopraorbitali perlato, abbastanza distanziate fra loro e poste su linee parallele, non convergenti all’indietro. Mandibole brevi, semplici, senza creste dorsali; labbro provvisto di 6 setole marginali anteriori; labium stretto, articolato, privo di dente sul margine anteriore, con epilobi poco sviluppati (fig. 10). Palpi mascellari (fig. 9) con penultimo articolo grande, ovoidale allungato, e ultimo piccolissimo, poco differenziato. _ Pronoto trasverso (rapporto larghezza max./ lunghezza max. = 1.31), allargato in avanti, ristretto alla base, con lati poco e lungamente arcuati, non sinuati avanti la base, non denticolati né smarginati anteriormente agli angoli posteriori. Angoli anteriori arrotondati, non prominenti; posteriori ottusi e smussati al vertice. Disco appena convesso, con pubescenza corta e rada; doccia mediana molto superficiale, appena accennata. Doccia marginale larga e spianata, non ristretta posteriormente presso la base; setole marginali anteriori inserite all’interno della doccia marginale, circa all'altezza del quarto anteriore; setole basali appena avanti agli angoli posteriori. Elitre brevi, tozze, smarginate nella zona preapicale, senza traccia di strie (esclusa la stria suturale). Disco poco convesso; tegumenti lucidi, con microscultura quasi indistinta e con pubescenza corta, molto rada ed eretta. Omeri sfuggenti, largamente arrotondati; margine post-omerale impercettibilmente denticolato, con crenellatura estremamente fine e poco distinta fino all'altezza del 3° poro omerale; smarginatura preapicale presente ma poco accentuata; apici elitrali separatamente arrotondati. Doccia marginale larga ed evidente fino al terzo apicale. Chetotassi: poro ombelicato basale grande, foveato. Serie ombelicata di tipo C (i grandi pori setigeri della serie ombelicata corrispondono al 2°, al 6° e al 7° (sensu JEANNEL 1963)); primi tre pori del gruppo omerale circa equidistanti fra loro, 4° poro leggermente più distanziato e inserito anteriormente alla metà dell’elitra; 5° poro posto circa al terzo apicale dell’elitra, 6° posto prima della smarginatura apicale, 7° posto presso la smarginatura apicale. Pori discali assenti; un piccolo poro setigero è presente nell’area preapicale fra il 6° e il 7° poro della serie ombelicata. Zampe (fig. 8) relativamente allungate, non ispessite. NUOVI ANILLINA DEL KENYA 605 (ALL ‘4 hy Wn, ane a 7X = 10 Fico. 7-10 Microdipnites perreti n. sp., Holotipo 9: stilomeri 1 - 2 sinistri in visione dorsale (7); zampa protoracica destra in visione dorsale (8); palpo labiale sinistro in visione ventrale (9); labium in visione ventrale (10). Scala: 0.1 mm. 606 R. GARETTO & P. M. GIACHINO Stili, nell’olotipo 2 (fig. 7), gracili, con fossetta sensoria munita di una setola. Maschio sconosciuto. DERIVATIO NOMINIS. Dedichiamo con piacere questa nuova specie al Dr. J.-L. Perret che per primo la raccolse insieme al Dr. V. Mahnert. DISTRIBUZIONE, ECOLOGIA. Microdipnites perreti n. sp. è noto al momento solo della località tipica: Taita Hills, presso Kwale a SW di Mombasa (Kenya). L’unico esemplare noto è stato raccolto al vaglio in foresta mista ad una quota di 1350 m. Selenodipnus embuanus n. sp. Figg. 11-15 Holotipo ? Kenya, Embu district, Irangi Forest Station, 2100 m, thifting of forest litter, 12.X.1977, V. Mahnert et J.-L.Perret leg. (Muséum d’histoire naturelle, Genève) DriaGnosI. Un Selenodipnus con caratteri perfettamente conformi alla defi- nizione del genere fornita da JEANNEL (1937, 1963): dente labiale assente, setole pro- notali presenti in numero di due, elitre non smarginate posteriormente, serie ombelicata di tipo B (sensu JEANNEL 1963), setole discali presenti, bordo omerale denticolato. S. embuanus n. sp. differisce invece da S. parallelus (Jeannel, 1957) per le elitre di forma ovoidale, a lati non paralleli, e per la posizione del 7° poro della serie ombeli- cata; da S. humerosus (Jeannel, 1957) per l’apice elitrale non allargato, ma regolar- mente arrotondato e per la differente localizzazione del 7° e 8° poro della serie ombelicata. Descrizione. Lunghezza totale dell’olotipo © (dall’apice delle mandibole all'apice delle elitre) mm 1.23 (dall’apice delle mandibole all’apice dell’ultimo uro- tergite mm 1.27). Corpo tozzo, depigmentato, fulvo-testaceo con elitre e addome più chiari, giallo-testacei; tegumenti lucidi, a microscultura quasi totalmente svanita, cosparsi di pubescenza rada e breve (fig. 11). Capo robusto, più stretto del pronoto, anoftalmo. Antenne ispessite, nettamente moniliformi a partire dal terzo antennomero, raggiungenti appena, distese all'indietro, la base del pronoto. Solco clipeo-frontale indistinto; margine anteriore dell’epistoma subrettilineo. Due setole sopraorbitali per lato, abbastanza distanziate fra loro e poste su linee convergenti posteriormente, due setole occipitali presenti. Mandibole brevi, semplici, senza creste dorsali; labbro grande, provvisto di 6 setole marginali anteriori; labium stretto, trasverso, articolato, privo di dente sul margine anteriore, con epilobi grandi ma moderatamente sporgenti (fig. 15). Palpi mascellari (fig. 14) con penultimo articolo poco rigonfio, ovoidale molto allungato, e ultimo lungo circa un terzo del penultimo. Pronoto subquadrato (rapporto larghezza max./ lunghezza max. = 1.17), poco allargato in avanti, moderatamente ristretto alla base, con lati poco arcuati, quasi rettilinei e convergenti nella metà basale; non sinuati ma distintamente denticolati avanti la base. Angoli anteriori poco prominenti, posteriori retti al vertice. Disco appena convesso, con pubescenza corta e rada; doccia mediana molto superficiale, appena accennata. Doccia marginale larga e spianata, non raggiungente la base e non ristretta posteriormente; setole marginali anteriori inserite all'interno della doccia marginale, circa all'altezza del quarto anteriore; setole basali appena avanti agli angoli posteriori. NUOVI ANILLINA DEL KENYA 607 Fic. 11 Selenodipnus embuanus n. sp.: Holotipo ©. Habitus. Scala: 0.1 mm. 608 R. GARETTO & P. M. GIACHINO 13 8 FIGG. 12-15 Selenodipnus embuanus n. sp., Holotipo 2: stilomeri | - 2 destri in visione dorsale (12); zampa protoracica destra in visione dorsale (13); palpo labiale sinistro in visione ventrale (14); labium in visione ventrale (15). Scala: 0.05 mm. NUOVI ANILLINA DEL KENYA 609 Elitre brevi, tozze e largamente ovoidali; senza traccia di strie. Disco poco convesso; tegumenti lucidi, con microscultura quasi indistinta e con pubescenza corta, molto rada ed eretta. Omeri appena accennati, largamente arrotondati; margine post- omerale sensibilmente denticolato, con crenellatura fine ma distinta fino all'altezza del 4° poro omerale; apici elitrali separatamente e largamente arrotondati, privi di smargi- natura preapicale. Doccia marginale larga ed evidente fino al livello del 9° poro della serie ombelicata. '@ Kisangani Fic. 16 Cartina di distribuzione dei generi e specie di Anillina nell’ Africa centro-orientale. 9 Micro- dipnites Jeannel, 1957: M. kauzianus (Basilewsky, 1951) (1); M. minutissimus (Basilewsky, 1954) (2); M. vanschuytbroecki Bruneau De Miré, 1990 (3); M. perreti n. sp. (4); M. mahnerti n. sp. (5). A Selenodipnus Jeannel, 1963: S. humerosus Jeannel, 1957 (6); S. parallelus Jeannel, 1957 (7); S. embuanus n. sp. (8). V Pelonomites Jeannel, 1963: P. leleupi (Basilewsky, 1953) (9); P. celisi (Basilewsky, 1954) (10); P. coiffaiti Bruneau De Miré, 1990 (11); P. vignai Zaballos & Casale, 1997 (12). Elgonotyphlus Sciaky & Zaballos, 1993: E. zoiai Sciaky & Zaballos, 1993 (13). Chetotassi: poro ombelicato basale grande, foveato. Serie ombelicata di tipo B [1 grandi pori setigeri della serie ombelicata corrispondono al 2°, al 6° e al 9° (sensu JEANNEL 1963)]; primi tre pori del gruppo omerale circa equidistanti fra loro, 4° poro distanziato di circa il doppio e inserito anteriormente alla metà dell’elitra; 5° poro posto prima del terzo apicale dell’elitra, 6° posto al terzo apicale dell’elitra, 7° circa equidistante e posto all’interno sul disco; 8° e 9° estremamente ravvicinati fra loro. Pori 610 R. GARETTO & P. M. GIACHINO discali presenti in numero di tre; un piccolo poro setigero € pure presente nell’area preapicale dopo il 9° poro della serie ombelicata. Zampe (fig. 13) relativamente allungate, non ispessite. Stili, nell’olotipo © (fig. 12), robusti, muniti di due setole spiniformi e di una fossetta sensoria con due setole. Maschio sconosciuto. DERIVATIO NOMINIS. La nuova specie prende nome dal distretto dell’ Embu, posto a Sud del Monte Kenya, dove è sita la località tipica. DISTRIBUZIONE, ECOLOGIA. Selenodipnus embuanus n. sp. è noto al momento solo della località tipica: Irangi Forest, sita a 2100 m di quota sulle propaggini meridionali del Monte Kenya (distretto di Embu). L’unico esemplare conosciuto è stato raccolto vagliando il terriccio posto sotto alberi morti in foresta mista. CONSIDERAZIONI ZOOGEOGRAFICHE Le nuove specie descritte nella presente nota rivestono un particolare significato zoogeografico. Del Kenya erano finora note due specie di Anillina: Pelonomites vignai Zaballos & Casale, 1997 e Elgonotyphlus zoiai Sciaky & Zaballos, 1993, ambedue del Monte Elgon. La scoperta di specie appartenenti ai generi Microdipnites Jeannel, 1957 e Selenodipnus Jeannel, 1963 in Kenya, in zone relativamente lontane dagli areali finora conosciuti per questi due generi (essenzialmente la catena montuosa del Kiwu in Congo) appare quindi di grande interesse, e conferma uno scenario di popolamento (fig. 16), per alcuni generi della linea filetica di Microtyphlus (Microdipnites, Pelonomites, Selenodipnus, Elgonotyphlus), ampiamente illustrato e discusso da JEANNEL (1961), LELEUP (1965) e ZABALLOS & CASALE (1997). RINGRAZIAMENTI Siamo molto grati al Dr. Ivan Löbl del Muséum d’histoire naturelle di Ginevra per averci concesso in studio 1 preziosi materiali oggetto della presente nota. Un ringraziamento particolare va al Prof. Achille Casale del Dipartimento di Zoologia e Antropologia Biologica dell’Università di Sassari per la revisione critica del manoscritto. BIBLIOGRAFIA JEANNEL, R. 1937. Les Bembidiides endogés (Col. Carabidae). Revue francaise d’Entomologie 3: 241-339. JEANNEL, R. 1961. La Gondwanie et le peuplement de |’ Afrique. Annales du Musée royal du Congo Belge 102: 5-161. JEANNEL, R. 1963. Monographie des “Anillini”, Bembidiides endogés (Coleoptera Trechidae). Mémoires du Muséum national d’Histoire naturelle, Paris (A) 28: 33-204. LELEUP, N. 1965. La faune entomologique cryptique de |’ Afrique intertropicale. Annales du Musée royal de l’Afrique Centrale, Tervuren 8, 141: 1-186. SCIAKY, R. ZABALLOS, J. P. 1993. Elgonotyphlus zoiai: new genus and new species of phane- rodont Anillini from Kenya (Coleoptera Carabidae). Journal of African Zoology 107: 321-327. ZABALLOS, J. P., CASALE, A. 1997. Un nuevo Pelonomites Jeannel, 1963 del Monte Elgon (Kenya) (Coleoptera: Carabidae: Bembidiinae: Anillini). Elytron 11: 105-113. REVUE SUISSE DE ZOOLOGIE 106 (3): 611-620; septembre 1999 Histiostrongylus spineus n. sp. (Nematoda: Trichostrongylina), parasite de Phyllostomus discolor (Chiroptera: Phyllostomidae) et nouvelles données sur les genres Histiostrongylus Molin, 1861 et Parahistiostrongylus Perez Vigueras, 1941 Claude VAUCHER! & Marie-Claude DURETTE-DESSET? ! Muséum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Suisse. 2 Muséum national d'Histoire naturelle, Laboratoire de Biologie parasitaire: Protozoologie, Helminthologie, associé au CNRS, 61, rue de Buffon, F-75231 Paris cedex 05, France. Histiostrongylus spineus n. sp. (Nematoda: Trichostrongylina), parasite of Phyllostomus discolor (Chiroptera: Phyllostomidae), and comments on the genera Histiostrongylus Molin, 1861 and Parahistiostrongylus Perez Vigueras, 1941. - Histiostrongylus spineus n. sp. is described from the bat Phyllostomus discolor caught in Peru and in Nicaragua. Parahistio- strongylus octacanthus Lent & Freitas, 1940 is redescribed. H. spineus is distinguished from the other species of the genus, H. coronatus Molin, 1861 mainly by the disposition of the bursal rays 6 and 8, the dorsal ray and the oesophagus. The validity of both genera is confirmed. Key-words: Nematoda - Molineoidea - Molineidae - Anoplostrongylinae - Histiostrongylus - Parahistiostrongylus - neotropical bats. INTRODUCTION Parmi les Anoplostrongylinae (Molineidae) parasites de Chiropteres, plusieurs genres sont caractérisés par des têtes armées d’épines. Nous terminons dans ce travail la revision des especes du Nouveau Monde commencée précédemment (VAUCHER & DURETTE-DESSET 1986). MATERIEL Le matériel a été collecté au Nicaragua par A. de Chambrier et T. Jaccoud (Muséum de Genéve), au Pérou par C. Vaucher, au Paraguay par C. Dlouhy (Asuncion) ou lors des missions du Muséum de Genève (EMGP). D’autres exemplaires résultent de la dissection de Chiropteres provenant du Surinam et de Colombie, conservés au Département de mammalogie du Muséum de Genéve. Les parasites sont déposés dans les collections du Muséum de Genéve (INVE) et du Muséum de Paris (MNHN, KP). Manuscrit accepté le 01.02.1999 612 C. VAUCHER & M.-C. DURETTE-DESSET Histiostrongylus spineus n. sp. MATERIEL-TYPE Chez Phyllostomus discolor (intestin gréle): Pérou, prov. Loreto, Samiria, male holotype (25150 INVE), femelle allotype (25151 INVE) et 2 paratypes mâles et 1 paratype femelle (25152 INVE), 25.10.1980, leg. C. Vaucher et F. Bona. AUTRE MATERIEL Chez Phyllostomus discolor: Pérou. prov. Loreto, Rio Yanayacu, | femelle (MNHN, 844 KP), 20.10.1980; Pérou, prov. Loreto, Cocha Yuracyacu, | fragment (25153 INVE), 20.10.1980, leg. C. Vaucher et F. Bona; Nicaragua, Prov. Leon, rives du lac Momotombo, 1 male (25154 INVE), 20.01.1983, leg. A. de Chambrier et T. Jaccoud. Nematodes de taille moyenne, non enroulés. Synlophe constitué d’environ 89 (male) a 105 crétes (femelle) au milieu du corps, orientées perpendiculairement a sa paroi, a peine plus serrées sur les faces latérales que sur les faces médianes (Fig. 3). Musculature présentant une portion non contractile tres développée. Téte portant un capuchon céphalique arme de 8 grosses épines réunies a leur base, a lame acérée (Figs 1, 4). Vue apicale montrant une bouche triangulaire avec une minuscule dent dorsale incisée. Cycle de papilles labiales internes et cycle de papilles labiales externes réduits aux deux latérales, réunies en une papille double voisine des amphides. Quatre papilles céphaliques (Fig. 4). Anneau nerveux, pore excréteur et deirides très antérieurs et situés au méme niveau (Figs 1, 2). Glandes excrétrices de grande taille. Oesophage pourvu d’une valvule, précédant une constriction délimitant un bulbe distal distinct. Présence d’un court canal oesophago-intestinal (Fig. 1). Male: male holotype long de 8.1 mm sur 175 um de large. Téte haute de 155 um et de 240 um de diametre, épines incluses. Anneau nerveux, pore excréteur et deirides situés respectivement a 235, 280 et 290 um de l’apex. Oesophage long de 570 um. Bourse caudale de type 2-3 mesurant 295 x 175 um, relativement petite, aux côtes 4 plus courtes que les côtes 3 et 5. Papilles des côtes 6 situées environ a mi-distance des côtes 5 et 8. Côtes 8 naissant au 1/3 de la côte dorsale, longues, mais n’atteignant pas le bord de la membrane bursale. Cote dorsale divisée en 3 branches a son extrémité distale, les deux branches externes elles-mémes subdivisées (Figs 5-7). Spicules tres longs, subégaux, de 1.5 mm (19% de la longueur du corps) pourvus d’une double aile, élargie surtout dans sa portion proximale; extrémité distale divisée en trois branches, l’externo-latérale se séparant nettement de l’interno-dorsale et de |’ interno-ventrale; branches internes renflées a leur extrémité, l’interno-ventrale étant caractérisée par sa forme en pince (Fig. 8). A l’extrémité de chaque branche, présence de membranes, collapsées par la fixation. Fics 1-8 Histiostrongylus spineus n. sp. chez Phyllostomus discolor. 1 - femelle, partie antérieure, vue laterale droite (la flèche indique l’emplacement de la deiride); 2 - paratype, deiride; 3 - paratype male, coupe transversale au milieu du corps; 4 - vue apicale d’un fragment antérieur; 5 - male, bourse caudale, vue latérale droite; 6 - premier paratype mâle, detail de la côte dorsale, vue ventrale; 7 - second paratype male, bourse caudale, vue ventrale; 8 - premier paratype male, détail de l’extrémité du spicule droit disséqué; el = branche externo-latérale; id = branche interno- dorsale; iv = branche interno-ventrale. (1: MNHN, 844 KP; 2, 3, 6-8: 25152 INVE; 4: 25153 INVE; 5: 25154 INVE). 613 HISTIOSTRONGYLUS ET PARAHISTIOSTRONGYLUS I ET DLE RAN za 614 C. VAUCHER & M.-C. DURETTE-DESSET Cone génital avec lèvre dorsale longue, lèvre ventrale élargie et peu proémi- nente. Papilles 0 non observées, papilles 7 filiformes. Gubernaculum très visible, d’aspect spongieux, long de 150 um (Fig. 7). Femelle: allotype longue de 9.5 mm sur 170 um, adulte mais non ovigère. Téte haute de 160 um et de 245 um de diamètre, épines incluses. Oesophage long de 695 um. Anneau nerveux, pore excréteur et deirides situés respectivement à 190, 183 et 202 um de l’apex. Vulve à 2.8 mm de l’extrémité postérieure (70% de la longueur totale). Ovéjecteur amphidelphe (les deux femelles du matériel-type) ou opisthodelphe (1 spécimen), présentant toujours une très forte asymétrie du vestibule (Fig. 18); branche antérieure: vestibule 810 um, sphincter 60 um et trompe 175 um; branche postérieure: vestibule 80 um, sphincter 60 um et trompe 180 um. Utérus ne contenant pas d’oeufs (de méme que chez les deux autres femelles). Queue longue de 90 um, avec deux pointes latéro-ventrales et une dorsale dédoublée entourant une pointe médiane acérée (Figs 19, 22). REMARQUES L’espece que nous avons décrite ci-dessus possede les caractéristiques du genre Histiostrongylus Molin, 1861 decrit chez Phyllostomus discolor au Brésil et particu- lièrement bien illustré par PEREZ VIGUERAS (1941) et BARUS & VALLE (1967). Ces caractéristiques sont les suivantes: ovéjecteur fortement asymétrique, spicules très longs (plus de 1 mm), se divisant en trois branches a leur extrémité distale et présence d’un gubernaculum. De plus, elle possede comme H. coronatus Molin, 1861, une valvule oesophago-intestinale. Cependant, plusieurs caracteres la distinguent de H. coronatus, espece-type du genre: alors que chez celle-ci, les cötes 6 et 8 sont paralleles et que leurs extrémités sont très rapprochées, chez A. spineus n. sp. au contraire, ces côtes sont divergentes et leurs extrémités écartées d’une distance environ égale a celle qui sépare les extrémités des côtes 5 et 6. L’oesophage est différent également, l’espece-type ne possédant pas de bulbe proximal comparable. D’autre part, dans notre matériel, la côte dorsale ne se termine pas par une pointe dépassant très nettement les côtes 9 et 10, comme c’est le cas chez H. coronatus (Perez Vigueras 1941; Barus & Valle 1967). Enfin, les épines céphaliques sont plus robustes, d’aspect triangulaire, épaisses a la base et présentent une petite indentation distale qui n’existe pas chez l’espece-type. Etymologie: spineus, adjectif latin signifiant épineux. Fics 9-17 Parahistiostrongylus octacanthus Lent & Freitas, 1940 chez Phyllostomus elongatus. 9 - femelle, extrémité antérieure (la fleche indique l’emplacement de la deiride); 10 - femelle, coupe transversale au milieu du corps; 11 - femelle, vue apicale; 12 - 14: variations de la bourse caudale selon la provenance: 12, matériel colombien; 13 - matériel paraguayen; 14 - vue latérale droite, matériel péruvien; 15 - variation de l’extrémité de la côte dorsale; 16 - mâle, detail de l’extrémité d’un spicule droit disséqué, vues ventrale, dorsale, laterale; 17 - deiride. (9 - 11, 14, 17: 25174 INVE; 12: 25160 INVE; 13, 15, 16: 25156 INVE). HISTIOSTRONGYLUS ET PARAHISTIOSTRONGYLUS 615 9 15 10 12-14 11,16 100 um GIG C. VAUCHER & M.-C. DURETTE-DESSET Bien que le nombre de spécimens examinés soit réduit, les prévalences semblent élevées: 3 P. discolor parasités sur 5 examinés provenant du Pérou; | parasité sur 1 examiné provenant du Nicaragua. Parahistiostrongylus octacanthus Lent & Freitas, 1940 MATERIEL ETUDIE Chez Phyllostomus hastatus: Pérou, prov. Loreto, Rio Yanayacu, 1 mâle, 1 partie antérieure (25155 INVE), 18.10.1980, leg. C. Vaucher et F. Bona; Paraguay, prov. Concepcion, 20 km au sud de l’estancia “Estrellas”, 6 mâles, 11 femelles, 3 fragments antérieurs (25156 INVE: MNHN, 202 KP), 16.10.1979, leg. EMGP; Brésil, prov. Rio de Janeiro, estancia “Belfa”, Vale do Saò Joa6, 2 mâles, 1 femelle (25157 INVE), 10.01.1986, leg. C. Dlouhy; Colombie, 24 mâles et 23 femelles chez 14 hôtes (25158-25171 INVE: MNHN, 842, 843 KP), 1961-1965, leg. B. Mechler; Surinam, 3 males et 7 femelles, chez 2 hötes (25172, 25173 INVE). Chez P. elongatus: Pérou, prov. Loreto, Samiria, 4 males, 8 femelles (25174 INVE), 25.10.80; 2 fragments antérieurs, 1 femelle (25175 INVE), 08.11.1980; 2 femelles (25176 INVE), 08.10.1980, leg. C. Vaucher et F. Bona. Chez Sturnira lilium: Paraguay, prov. Itapua, Caicisa, 1 male, 1 femelle (25177 INVE), 06.11.1982; 1 femelle (25178 INVE), 07.11.1982; 1 femelle, 1 partie antérieure (25179 INVE), 07.11.1982, leg. EMGP. Nématodes de taille moyenne, non enroulés. Synlophe constitué de petites crétes a peine plus serrées sur les faces latérales que sur les faces médianes, orientées perpen- diculairement a la paroi du corps (Fig. 10). Chez un male (25156 INVE), il y a 81 crétes au milieu du corps; chez deux femelles (25158 INVE), 81 a 83 crétes au niveau de la fin de l’oesophage, 86 à 91 au quart antérieur du corps, 88 a 91 au tiers antérieur, 88 et 87 au milieu, 74 et 71 au quart postérieur; chez une autre (25174 INVE), 96 crétes au milieu du corps. Musculature circulaire présentant une nette portion contractile. Téte avec une vésicule céphalique armée de 8 grosses épines en forme de lame mince et acérée, réunies a leur base (Fig. 9). Vue apicale montrant une bouche triangulaire pourvue d’une dent dorsale. Cycle de papilles labiales internes réduit aux deux latérales; cycle complet de papilles labiales externes. Papilles labiales interno- et externo-latérales formant deux papilles doubles associées aux amphides; autres papilles labiales externes formant quatre papilles doubles avec les quatre papilles céphaliques (Fig. 11). Anneau nerveux, pore excréteur et deirides très antérieurs, situés au méme niveau. Oesophage présentant un élargissement médian et un bulbe terminal, pourvu d’une discrete valvule au niveau de la constriction posterieure. Présence d’un tres court canal oesophago-intestinal (Fig. 9). Male: un male de 5.6 mm a 220 um de diamètre. Téte haute de 112 um et de 218 um de diamètre, épines incluses. Anneau nerveux, pore excréteur et derides situés respectivement a 230, 250 et 250 um de l’apex. Oesophage long de 610 um. Bourse caudale de type 2-3, mesurant 385 x 155 um (Figs 12-14). Cötes 4 plus courtes que les côtes adjacentes. Côtes 6 écartées des côtes 5. Espace entre les côtes 6 et 8 un peu plus court que l’espace entre les côtes 5 et 6. Cône génital avec des lèvres dorsale et ventrale élargies. Cötes 8 naissant au tiers proximal de la hauteur de la cöte dorsale, plus courtes que celle-ci et n’atteignant pas le bord de la membrane bursale. Côte dorsale divisée de façon variable a son extérmité distale: côtes 9 et 10 naissant au tiers distal de la côte dorsale, parfois séparées à la base, parfois portées par un court tronc commun (Fig. 15). HISTIOSTRONGYLUS ET PARAHISTIOSTRONGYLUS 617 Cötes 10 portant un appendice interne plus ou moins développé. Cotes 11 fusionnées en une pointe acérée dépassant souvent le lobe dorsal de la membrane. Spicules courts, de 255 um (230-285 um chez 6 autres individus). Extrémité distale des spicules recourbée dorsalement et terminée par une boule arrondie; du côté ventral au contraire, le spicule est aplati et forme une palette élargie (Fig. 16). Pas de papilles observables au niveau du cone génital. Gubernaculum absent. Femelle: une femelle longue de 7.3 mm a un diametre de 215 um au milieu du corps. Téte haute de 110 um et de 160 um de diametre, épines incluses. Oesophage long de 610 um. Anneau nerveux, pore excréteur et deirides situés respectivement a 202, 250 et 265 um de l’apex. Vulve s’ouvrant a 2.67 mm de l’extrémité postérieure (61% de la longueur du corps). Ovéjecteur amphidelphe, vestibule symétrique ou presentant une légère asymetrie (Fig. 21). Branche antérieure: vestibule 270 um, sphincter 155 um et trompe 350 um; branche postérieure: vestibule 350 um, sphincter 155 um et trompe 570 um. Les dimensions des différentes parties de l’ovéjecteur présentent une variabilité marquée; chez la femelle dont l’ovéjecteur est illustré (Fig. 21), les dimensions suivantes ont été relevées: Branche antérieure: vestibule 135 um, sphincter 100 um et trompe 290 um; branche posterieure: vestibule 130 um, sphincter 100 um et trompe 190 um. Uterus antérieur long de 1.9 mm avec 35 œufs sur 1-2 rangs; utérus postérieur long de 1.7 mm avec 15 œufs sur 1-2 rangs. Oeufs de 78-96 x 46-57 um. Queue longue de 85 um, terminée par une pointe médiane recourbee ventralement, une pointe dorsale dédoublée et deux pointes latéro-ventrales (Figs 20, 23). REMARQUES Cette espece a été décrite par LENT & FREITAS (1940) chez P. hastatus au Brésil. Il n’y a pas de differences notables entre nos observations et la description originale. La bourse caudale a été observée en détail et dessinée par ces auteurs, qui relevent une forte variation dans la disposition des côtes constituant la côte dorsale. Nous avons observé également une certaine variation (Fig. 15), mais pas de disposition comparable aux figures 12 et 17 de LENT & FREITAS (1940), qui nous paraissent par ailleurs sortir d’une variation intraspécifique et relever plutot de la tératologie. Les prévalences observées sont élevées chez les hötes du genre Phyllostomus: 1 P. hastatus parasité sur 1 examiné au Paraguay, au Brésil ainsi qu’au Pérou: 2 parasités sur 2 examinés provenant du Surinam; 14 parasités sur 17 examinés provenant de Colombie; 2 P. elongatus parasités sur 8 examinés au Pérou. Au contraire, 3 Sturnira lilium parasités sur 121 examinés au Paraguay, ce qui pourrait indiquer que cet höte est accidentel. DISCUSSION Les Trichostrongles parasites de Chiropteres possédant une tête armée sont classés dans 7 genres. Biacantha Wolfgang, 1954 (2 especes) est caractérisé par 2 épines ventrales et Websternema Vaucher & Durette-Desset, 1986 (1 espece) par 2 épines latérales. Trois genres, Histiostrongylus Molin, 1861 (2 especes), Parahistiostrongylus Perez Vigueras, 1941 (1 espèce) et Neohistiostrongylus Barus & RySavy, 1971 618 C. VAUCHER & M.-C. DURETTE-DESSET a 20 um 18.217 1920 100 um HISTIOSTRONGYLUS ET PARAHISTIOSTRONGYLUS 619 (1 espèce) possèdent une tête armée de 8 épines disposées régulièrement. Les deux premiers genres sont proches puisque différents auteurs (YAMAGUTI 1961; BARUS & RySAvy 1971; DURETTE-DESSET & CHABAUD 1975) les ont considérés comme syno- nymes. Cependant DURETTE-DESSET (1983) revalide Parahistiostrongylus en s’appuyant sur les caractères du vestibule, fortement asymétrique chez Histiostron- gylus, Sur la morphologie des spicules et sur absence de gubernaculum chez Para- histiostrongylus. Barus & RySAvy (1971) ont proposé le genre Neohistiostrongylus pour classer Parahistiostrongylus viguerasi Lopez-Neyra, 1946, parasite de Myotis myotis (Vesper- tilionidae) en Espagne. Ce genre est caractérisé par deux ailes latérales, présentes tout le long du corps, par des spicules courts divisés profondément en 3 branches, par un revétement épineux de la face interne de la bourse caudale et par la présence d’un gubernaculum. Ce genre nous parait donc valide. Linustrongylus Vaucher & Durette-Desset, 1986, monotypique, possede aussi 8 épines, mais elles sont réparties en deux groupes symetriques sur les faces médianes; de plus, la bouche est entourée d’un cadre buccal (VAUCHER & DURETTE-DESSET 1986). Le dernier genre, Spinostrongylus Travassos, 1935 (2 especes) présente 14-16 épines simples ou dédoublées, suivies de rangées de très petites épines en arrière de la téte; il est donc très différent des précédents. Spinostrongylus est connu d’ Egypte et de Chine. Avec Neohistiostrongylus, décrit d'Espagne, il se distingue, par sa répartition zoogéographique, des autres taxa a téte armée, qui sont connus seulement du Nouveau Monde. DURETTE-DESSET & CHABAUD (1975) considéraient les espèces possédant un petit nombre de crochets comme primitives et estimaient que les espèces pourvues d’une armature plus complexe tendaient a perdre leur synlophe. Avec les nouvelles données que nous apportons, cette corrélation paraît moins nette, puisque Histiostron- gylus et Parahistiostrongylus, qui possèdent des structures céphaliques déjà complexes, ont un synlophe formé de nombreuses crétes, peu développées il est vrai. De toute manière, tous les synlophes présents dans ce groupe a téte armée sont caractérisés par de petites crétes, orientées perpendiculairement a la paroi du corps. Seuls Spino- strongylus et Linustrongylus sont totalement dépourvus de synlophe. Les mémes auteurs se demandaient si les formes armées formaient un groupe monophylétique. Il ne nous a pas été possible de mettre en évidence une ou plusieurs synapomorphies communes à ces quelques genres. Au contraire, elles forment un ensemble hétérogène: les bourses caudales présentent des dispositions de cötes diverses. Par exemple, les côtes 4 sont nettement plus courtes que les côtes adjacentes chez Histiostrongylus, Fics 18-23 Figs 18, 19, 22: Histiostrongylus spineus n. sp. chez Phyllostomus discolor. 18 - paratype femelle, ovéjecteur; 19 - queue d’une femelle; 20 - femelle, extrémité postérieure. (18, 22: 25152 INVE; 19: MNHN, 844 KP). — Figs 20, 21, 23: Parahistiostrongylus octacanthus Lent & Freitas, 1940. 20 - femelle, ovéjecteur; 21 - queue d’une femelle; 23 - femelle, extrémité postérieure. (20, 23: 25174 INVE, Phyllostomus elongatus; 21: 25156 INVE, Phyllostomus hastatus). 620 C. VAUCHER & M.-C. DURETTE-DESSET Parahistiostrongylus, Websternema et Neohistiostrongylus. Chez Linustrongylus, toutes les côtes sont courtes et trapues, elles rappellent par conséquent la disposition rencontrée chez Cheiropteronema Sandground, 1929, lobe dorsal excepté (DURETTE- DESSET & VAUCHER 1988). Des spicules longs caractérisent Websternema, Histio- strongylus et Linustrongylus. L’extrémité des spicules est divisée en trois branches chez Neohistiostrongylus, Histiostrongylus et Biacantha. La queue des femelles est pourvue d’une seule pointe chez Spinostrongylus, dans tous les autres genres il y a plusieurs pointes. Par conséquent, nous estimons qu’il n’existe pour l’instant aucun argument décisif pour considérer les especes a tète armée comme un groupe naturel. REMERCIEMENTS Aux participants de l’expédition du Muséum de Genève au Paraguay en 1979 et en 1982 et particulièrement a François J. Baud (Muséum de Genève) pour I identi- fication des hötes; a Alain de Chambrier, Thierri Jaccoud (Muséum de Geneve) et Carlo Dlouhy (Asuncion) pour leurs récoltes au Nicaragua et au Brésil; au ministére de l'Agriculture et des Elevages du Paraguay, au ministère de l’Agriculture du Pérou a Lima et Iquitos ainsi qu’au personnel de la station biologique “Pithecia” de Samiria (prov. Loreto, Pérou) pour les facilités accordées lors du travail de terrain. BIBLIOGRAPHIE Barus, V. & RySAvy, B. 1971. An analysis of the biogeography of Nematodes of the family Trichostrongylidae parasitizing Bats of the suborder Microchiroptera. Folia Parasito- logica 18: 1-14. BARUS, V. & DEL VALLE, M. T. 1967. Systematic Survey of Nematodes Parasitizing Bats (Chiroptera) in Cuba. Folia Parasitologica 14: 121-140. DURETTE-DESSET, M. C. 1983. Keys to genera of the superfamily Trichostrongyloidea. In: ANDERSON, R. C. & CHABAUD, A. G. (eds). CIH Keys to the nematode parasites of vertebrates. Commonwealth Agricultural Bureaux, Slough, 86 pp. DURETTE-DESSET, M. C. & CHABAUD, A. G. 1975. Nématodes Trichostrongyloidea parasites de Microchiropteres. Annales de Parasitologie humaine et comparée 50: 303-337. DURETTE-DESSET, M. C. & VAUCHER, C. 1988. Trichostrongyloidea (Nematoda) parasites de Chiropteres néotropicaux. II. Nouvelles données sur le genre Cheiropteronema Sandground, 1929. Revue suisse de Zoologie 95: 889-899. LENT, H. & TEIXEIRA DE FREITAS, J. F. 1940. Histiostrongylus octacanthus n. sp. (Nematoda: Strongyloidea). Arquivos do Instituto Benjamin Baptista 6: 91-96. LOPEZ-NEYRA, C. R. 1946. Parahistiostrongylus viguerasi sp. n. Trichostrongylidae, nuevo de Quiropteros en Espana. Revista ibérica de Parasitologia 6: 245-256. MOLIN, R. 1861. Il sottordine degli acrofalli ordinato scientificamente secondo i risultamenti delle indagini anatomiche ed embriogeniche. Memorie del Reale Istituto Veneto di Scienze, Lettere ed Arti, Venezia 9: 427-633. PEREZ VIGUERAS, I. 1941. Nota sobre el genero Histiostrongylus Molin, 1861. Revista de Medicina Tropical y Parasitologia 7: 67-72. VAUCHER, C. & DURETTE-DESSET, M.-C. 1986. Trichostrongyloidea (Nematoda) parasites de Chiroptères néotropicaux I. Websternema parnelli (Webster, 1971) n. gen. n. comb. et Linustrongylus pteronoti n. gen. n. sp., parasites de Preronotus au Nicaragua. Revue suisse de Zoologie 93: 237-246. YAMAGUTI, S. 1961. Systema Helminthum vol. 3: The Nematodes of Vertebrates. Interscience Publ., New York, 1261 pp. REVUE SUISSE DE ZOOLOGIE, 106 (3): 621-626; septembre 1999 A new apterous and microphthalmic species of Anemadus (Coleoptera: Leiodidae: Cholevinae) from China Jan RUZICKA Department of Ecology, Faculty of Forestry, Czech Agricultural University CZ-165 21 Praha 6, Czech Republic; e-mail: ruzicka@If.czu.cz A new apterous and microphthalmic species of Anemadus (Coleoptera: Leiodidae: Cholevinae) from China. - Anemadus smetanai sp. n. is descri- bed from China, Yunnan province. Its diagnostic characters including male and female genitalia are illustrated. The species differs from all other species of Anemadus Reitter, 1885 by microphthalmy, absence of metathoracic wings, elytra coalescent and coined with scutellum, ventrites with a pair of antero-lateral processes and female sternum VIII with concave spiculum ventrale. Key-words: Taxonomy - Coleoptera - Leiodidae - Cholevinae - Anemadus - China - Yunnan province. INTRODUCTION The genus Anemadus Reitter, 1885 is the largest genus of the tribe Anemadini Hatch, 1928 (sensu NEWTON & THAYER 1992). The genus was reviewed by GIACHINO & VAILATI (1993) with 28 valid species recognized, distributed from central and southern Europe, Turkey, Middle East, Iran to Pakistan and northern India. Later, a further species was described from China, Sichuan province (PERREAU 1996a), and two species were described from Taiwan and Japan (PERREAU 1996b). In this paper, a new species of this genus (collected recently by AleS Smetana in China) is described. It has several remarkable characters absent from other Anemadini, but known from endogenous and cavernicolous taxa. The following abbreviations are used throughout the text: JRUC - collection of Jan Ruzicka, Praha; MHNG - Muséum d’histoire naturelle, Genéve. Exact label data are cited for type material, separate lines are indicated by “/”, separate labels by “//”. The code in brackets, attached to locality data, refers to the habitat notes in the field book of the collector. Anemadus smetanai sp. n. Figs 1-13 MATERIAL EXAMINED. Holotype (4), allotype (2) and 3 paratypes (1 4,2 2 ©), labelled: “CHINA N Yunnan [prov.], Xue / Shan [mts] nr. Zhongdian / 3900m 25.V1.1996 / 27°49N 99°34E C41 // collected by / A. Smetana, J. Farkaë / and P. Kabätek”; 1 paratype (9 ): ditto, but “4050m, 24.V1.1996, C39”. Holotype is deposited in MHNG, allotype and paratypes in JRUC. Manuscript accepted 16.04.1999 JAN RUZICKA ETYMOLOGY. The new species is named after its collector, Ales Smetana from Ottawa. 22 En ae <8 ae SU on ERE zi. WM Bes — -2: 8 © 1 INN \ 237 Mi .& - Ou III) AA 5 DE 2 - Ah ; a a a4 N N + VON NS | EE SER 2: OS & HS = Sos 3: habitus setation and microsculpture of elytra dorso- VENDE : 4: setation of pronotum dorso-laterall laterally. J Anemadus smetanai Sp. n., female paratype. 1: head laterally; 2: left antenna dorsally dorsally ANEMADUS FROM CHINA 623 Head 1.05 and 1.07 times as long as wide. Surface smooth, without distinct microsculpture (under 50x magnification), covered by fine, regularly distributed punc- ° tures, separated by 2-4 times of their diameter. Eye reduced to cluster of about 10 facets (Fig. 1). Clypeo-frontal suture distinctly developed. Clypeus transverse, trapezoidal. Antenna (Fig. 2) elongated, 1.17 times as long as maximum pronotal width. Proportions of antennomeres in holotype (I to XI, length x maximum width): 24 x 9, PREIS NET NAT NOX 713 x Sal Fx 1ON8 x93 13 kT, 12 x 12 and 23x13. Pronotum 1.60 times as wide as long, widest behind the mid-length, 1.95 times as wide as head. Lateral margin regularly rounded, posterior angles acute, produced posteriorly. Posterior margin regularly concave. Pronotal surface regularly convex, with a pair of weak, postero-lateral impressions (Fig. 3). Pronotal surface covered by distinct transverse microsculpture; intermixed granulate punctures arranged as dense as on head, but more superficially (Fig. 4). Elytra oval, with slender, elongate apex. Elytra 1.45 and 1.49 times as long as maximum width, 2.50 and 2.52 times as long as pronotum and 1.07 times as wide as pronotum. Elytra coalescent, coined also with scutellum (Fig. 13). Each elytron apically rounded. Elytral surface with distinct transverse microsculpture. Punctures granulate, arranged into transverse, irregular rows (elytral type “a” in GIACHINO & VAILATI 1993; Fig. 5). Sutural stria absent. Large intermixed punctures, arranged into 8 irregular, longitudinal rows visible in elytron immersed to glycerine (not perceptible in dry mounted specimens). Metathoracic wings absent. Protibia slender, gradually expanded toward apex. Protarsus slightly expanded, basiprotarsomere as wide as maximum protibial width. Mesotibia slightly curved, two basal mesotarsomeres expanded. Metatibia almost straight, 1.35 times as long as metatarsus, metatibia 0.77 and 0.78 times as long as maximum combined elytral width. Mesosternal carina low, mesopraesternum posteriorly with sharp angle in lateral view. Metatergal apparatus reduced (Fig. 13). Metendosternite reduced, anterior part widely emarginate, posterior part shortened (Fig. 10). Ventrites IV to VII with sub- lateral part isolated by distinctly developed apodeme, antero-laterally present pro- minent process, tucked under posterior part of previous ventrite (as on Fig. 12). Genital segment as on Fig. 8. Aedeagus regularly narrowed toward tip with elongate, subtruncate apex (Fig. 6). Internal sac with a pair of lateral rows, composed by sclerotized spines of different size and shape, medial part covered only by very minute toothlets, arranged in regular pattern (Fig. 6). Paramere constricted subapically, apex incurvate, bearing three large and one minute seta on inner margin (Fig. 7). Female. Body length 2.75-3.05 mm (mean 2.92 mm in 4 specimens; for the same number of specimens given all means below), maximum body width 1.15-1.35 mm (mean 1.25 mm). Similar to the male, but body more robust, appendages less prolonged (Fig. 3). Head 1.01-1.10 times as long as wide (mean 1.05). Antenna 1.06-1.12 times as long as maximum pronotal width. Proportions of antennomeres in one paratype (I to XI, length x maximum width): 26 x 9, 22 x 8, 18x 4, 15 x 8, 16x 7, 12x 8, 16x 11,8 x 10, 132 el and 25! x13) 624 JAN RUZICKA Pronotum 1.61-1.73 times as wide as long (mean 1.65), 1.87-2.00 times as wide as head (mean 1.94). Elytra 1.44-1.51 times as long as maximum width (mean 1.46), 2.60-2.70 times as long as pronotum (mean 2.65) and 1.08-1.12 times as wide as pronotum (mean 1.10). Pro- and mesotarsomeres not expanded. Metatibia 1.25-1.29 times as long as metatarsus, 0.67-0.71 times as long as maximum combined elytral width (mean 0.69). Ventrites IV to VII with similar sublateral apodeme and antero-lateral process as in males. Ventrite VIII with elongate spiculum ventrale with anterior margin only weakly sclerotized, distinctly concave (Fig. 9). Tergum IX with lateral sclerotized line; apically with a single, minute seta; anterior and medial margin only weakly sclerotized. Tergum X ovoid, only weakly sclerotized anteriorly; posterior margin with two pairs of larger setae and several minute ones. Coxite small, subtriangular, bearing a single seta. Stylus elongate, with one basal, three subapical and one large, apical seta (Fig. 11). AFFINITIES. Anemadus smetanai sp. n. belongs to the genus Anemadus Reitter, 1885 (sensu GIACHINO & VAILATI 1993) having the internal sac of aedeagus with two longitudinal rows of spines and lacking the medial dent. The new species clearly differs from all other species of this genus by the following set of characters: (a) micropthalmy (Fig. 1; eyes fully developed in other species of Anemadus); (b) absence of metathoracic wings, combined with elytra coalescent and coined also with scutellum (Fig. 13; metathoracic wings fully deve- loped, elytra not coalescent in other species of Anemadus); (c) ventrites with a pair of antero-lateral processes (Fig. 12; such processes are lacking in other species of Ane- madus); (d) ventrite VIII with spiculum ventrale concave in females (Fig. 9; ventrite VII with spiculum ventrale convex in other species of Anemadus). GIACHINO & VAILATI (1993) proposed several species groups in Anemadus, based mainly on the shape of the aedeagus, the spine pattern of the internal sac and the type of elytral microsculpture. Presently, it is impossible to placed A. smetanai sp. n. to any of these species groups, the because the characters of the aedeagus differ from those in other groups of Anemadus. The new species lacks basal enlarged spines and possesses two lateral, longitudinal rows of larger spines of the internal sac, a similar shape of the apex of the aedeagus and elytra with microsculpture of the type “a” (according to GIACHINO & VAILATI 1993) similarly as in members of the A. strigosus (Kraatz, 1852) species group (A. strigosus, A. bianchii Reitter, 1906 and A. arcadius Reitter, 1885, from Europe and Asia Minor). However, in A. smetanai sp. n., the spines of the internal sac are generally much more larger (minute and more homogenous in species of the A. strigosus group). Furthermore, A. smetanai sp. n. differs in the shape of apex of the parameres which are distinctly incurvate (always distinctly turned outwards in members of the A. strigosus species group). Two of the three other recently described species of Anemadus from China, Taiwan and Japan were not placed in any species group (PERREAU 1996a, b). Further- more, other species of this genus are currently described from China (M. Perreau, pers. comm.). Consequently, I presently refrain from defining a new species group for A. smetanai sp. n. ANEMADUS FROM CHINA 625 Fics 6-13 Anemadus smetanai sp. n. 6: aedeagus dorsally; 7: apex of left paramera dorsally; 8: male genital segment ventrally; 9: female ventrite VIII ventrally; 10: female metendosternite postero-dorsally; 11: female genitalia dorsally; 12: lateral part of female ventrite V ventro-laterally, a - apodeme, p - antero-lateral process; 13: female metatergal apparatus, basal part of left elytron and scutellum ventrally. The isolated position of A. smetanai sp. n. may be strongly influenced by the life of the species in the montane deep litter conditions. Similar changes in external mor- phology (reduction of eyes, coalescent elytra and/or reduction to loss of metathoracic wings) are known also in other edaphophile species of Cholevinae, e.g. Choleva leuco- pthalma Fiori, 1899 (SOKOLOWSKI 1941), several species of Ptomaphagus (Adelops Tellkampf, 1844) (PECK 1973, 1977, 1978a, PECK & GNASPINI 1997) and all species of Ptomaphagus (Appadelopsis Gnaspini, 1996) (PECK 1978b). However, A. smetanai sp. n. 1s the first such known case in Anemadini (sensu NEWTON & THAYER 1992). BIONOMICS. Most of the specimens was sifted from deep layers of rotten leaves and detritus in montane primary forest (with dominant Abies, Betula, Carpinus and Rhododendron spp.), a single specimen was sifted from layers of moss, rotting bark and humus under it on a huge fallen fir (Abies sp.) in an primary high montane forest with dominant Abies and tree-like Rhododendron spp. (A. Smetana, pers. comm.). 626 JAN RUZICKA ACKNOWLEDGEMENTS I am grateful to Ales Smetana (Ottawa) for providing Chinese material of Cholevinae for study. Many thanks are due to Jana Nebesärovä and Antonin Polak (Laboratory of Electron Microscopy, ASCR, Ceské Budéjovice) for help with the SEM photographs, Jakub Roléfk (Bi-MAC Graphics Inc., Praha) for assistance with moun- ting the SEM photographs and David Kral for comments on the previous version of the manuscript. REFERENCES NEWTON, A.F. Jr. & THAYER, M.K. 1992. Current classification and family-group names in Staphyliniformia (Coleoptera). Fieldiana: Zoology, New Series 67: I-IV,1-92 pp. GIACHINO, P.M. & VAILATI, D. 1993. Revisione degli Anemadinae Hatch, 1928 (Coleoptera Cholevidae), Monografie di Natura Bresciana No. 18, Museo Civico di Scienze naturali di Brescia, Brescia, 314 pp. Peck, S.B. 1973. A systematic revision and the evolutionary biology of the Ptomaphagus (Adelops) beetles of North America (Coleoptera; Leiodidae; Catopinae), with emphasis on cave-inhabiting species. Bulletin of the Museum of Comparative Zoology 145: 29-162. Peck, S.B. 1977. The subterranean and epigean Catopinae of Mexico (Coleoptera: Leiodidae). Bulletin of the Association for Mexican Cave Studies 6: 185-213. Peck, S.B. 1978a. New montane Promaphagus beetles from New Mexico and zoogeography of southwestern caves (Coleoptera; Leiodidae; Catopinae). The Southwestern Naturalist 23: 227-238. Peck, S.B. 1978b. Systematics and evolution of forest litter Adelopsis in the southern Appa- lachians (Coleoptera: Leiodidae; Catopinae). Psyche 85: 355-382. PECK, S.B. & GNASPINI, P. 1997. Ptomaphagus inyoensis n.sp., a new microphthalmic montane beetle from California (Coleoptera; Leiodidae; Cholevinae; Ptomaphagini). The Cana- dian Entomologist 129: 769-776. PERREAU, M. 1996a. Contribution a la connaissance des Cholevidae du Japon et de Taiwan (Coleoptera). Revue suisse de Zoologie 103: 283-297. PERREAU, M. 1996b. Nouveaux Cholevinae d’Asie (Coleoptera Leiodidae). Revue suisse de Zoologie 103: 939-949. SOKOLOWSKI, K. 1941. Choleva leucopthalma Fiori (Col., Catopidae) (Catopiden-Studien III). Mitteilungen der Miinchner Entomologischen Gesellschaft 31: 943-948. REVUE SUISSE DE ZOOLOGIE 106 (3): 627-641; septembre 1999 The genus Proscopia Klug, 1820 (Orthoptera, Caelifera, Eumastacoidea, Proscopiidae) in Central America, with description of a new species Alba BENTOS-PEREIRA! & C. Hugh F. ROWELL? | Facultad de Ciencias - Entomologia, Universidad de la Repüblica, Casilla 490, 11200 Montevideo, Uruguay. ? Zoologisches Institut der Universität Basel, Rheinsprung 9, CH-4051 Basel, Switzerland. The genus Proscopia Klug, 1820 (Orthoptera, Caelifera, Eumastacoidea, Proscopiidae) in Central America, with description of a new species. - Corynorhynchus septentrionalis (Bruner) from Panama and Costa Rica is the only proscopiid known to occur in Central America. Here we show that the specimens previously allocated to this taxon actually comprise at least two and probably three species of the genus Proscopia. We describe P. pana- mensis n. sp. and redescribe P. septentrionalis (Bruner) (reinstated comb.). We are unable to describe the third taxon due to lack of adult material. Key-words: Orthoptera - Caelifera - Proscopiidae - taxonomy - Central America. INTRODUCTION In the most recent revision of the family Proscopiidae, JAGO (1989) noted that the genus Corynorhynchus was in need of revision, as many of its species were in his opinion badly identified. One of the most difficult taxa to place has been the species septentrionalis Bruner, 1905, the most northerly proscopiid known and the only one recorded from Central America. This species was originally described in the poorly defined genus Taxiarchus, later transferred to Proscopia by HEBARD (1924), and most recently to Corynorhynchus by JAGO (1989). The confused situation has been exacer- bated by the paucity of specimens. Bruner's description was based on two specimens from Pozo Azul de Pirris, Costa Rica, which he described as being a d and @; in fact both are 2 2, one adult and one larva, as originally noted by HEBARD (1924). HEBARD also cited two further specimens collected by D.E. Harrower in Gatün, Canal Zone, Panama, one adult d and one ® larva. JAGO (1989) cited a further ¢ collected by Schrader from Barro Colorado Island, Canal Zone, Panama, belonging to the collection of the Academy of Natural Sciences, Philadelphia (ANSP); he also mistakenly recorded the lectotype 9 as coming from Panama, instead of Costa Rica. We recently located two more adult © © and one adult ¢ in the collections of the Instituto Nacional de Biodiversidad (INBio), Costa Rica, and an adult d and a £ Manuscript accepted 02.03.1999 628 A. BENTOS-PEREIRA & C.H.F. ROWELL larva in those of the Smithsonian Tropical Research Institute in Panama (STR). Additionally we collected two further specimens (an adult 2 and a @ larva) on Cerro Copé, Panama, in September 1997. We have used all these specimens, including the type series from the ANSP, to review the Central American proscopiids, as part of an on-going revision of the entire family (Bentos-Pereira, in progress)'. We show here that there are in fact at least two Central American species, one from Costa Rica (septentrionalis Bruner), which we redescribe, and another species (panamensis n. sp.) from Panama. There is probably a second new species among the specimens, but there is to date insufficient material for a description. All three must be attributed to the genus Proscopia as it is currently defined. METHODS For the description of the d internal genitalia we use the terminology proposed by JAGo (1989); his diagram is reproduced in Figure 1. Numbers in the text following an anatomical term refer to this diagram. We also use the structure of the 2 sperma- theca as a systematic character, based on the description and classification of this structure in the Proscopiidae given originally by DESCAMPS (1973). All measurements were made under the microscope with a digital read-out micrometer stage and an eyepiece graticule. The measured dimensions are defined in Figures 2, 3, & 4. DESCRIPTIONS 1. Proscopia septentrionalis (Bruner, 1905), reinstated combination SYNONOMY Taxiarchus septentrionalis Bruner, 1905: 313-315, pl. 1; 1908: 342. KirBy 1910: 87 (mistakenly assigns the species to Rehn). CAUDELL 1911: 159 (mentions Kirby's error). OTTE 1978: 34 (location of types). Proscopia septentrionalis (Bruner); HEBARD 1924:93 (lectotype selected). MELLO LEITAO 1939: 417. CARBONELL 1977: 24. Corynorhynchus septentrionalis (Bruner); JAGO, 1989: 273 (mistakenly assigns the species to Brunner). SPECIMENS EXAMINED COSTA RICA: Prov. San José: Pozo Azul de Pirris, June 1905 (M. A. Carriker), 1 adult 3d, specimen nos. H233, CSC2670, ABP224. Lectotype of Taxiarchus septentrionalis Bruner, 1905 (ANSP). COSTA RICA: Prov. Puntarenas: Osa Peninsula, Corcovado National Park, 18-23 March 1978 (D.H. Jansen), 1 adult 4, in copulation, specimen nos. CRIO01 687843, ABP229 (INBio). This is the first authentic record of a 4 of this species. Same data as previous specimen, 1 adult ©, in copulation, specimen nos. CRI001 687844, ABP 228 (INBio). COSTA RICA: Prov. Puntarenas: Osa Peninsula, Agua Buena, Fila Casa Loma, 500 mts, LS N296300 E514300, 28.2.1993 (M.A. Zumbado), 1 adult 2, specimen nos. 146 MAZ.93, CRI002 510601, ABP 227 (INBio). ! The exceptions are the specimens collected by Harrower, which now appear to be missing from the ANSP collections. According to HEBARD (1924) their data were as follows: Panama: Prov. Panama: Gatün, 25-31 July 1916 (D.E. Harrower), one adult d and one @ larva. CENTRAL AMERICAN PROSCOPIIDS 629 2 4 3 Fics 1-4 Fic. 1. Diagram and nomenclature of the d genital sclerites of the family Proscopiidae, after JAGO, 1989. 1: transverse plate of ectophallic membrane; 2: pair of hook-like lophi; 3: median dorsal slit or genital opening; 4: pair of valvular plates lateral to genital opening; 5: sublophal pair of accessory plates; 6: pair of anterior supplementary lateral plates; 7: distal ejaculatory sac; 8: proximal part of endophallic duct; 9: pair of semicircular lateral struts giving flexible attach- ment for lophi; 10: pair of plates overlying the anterior supplementary lateral plates (6). — Fic. 2. Dimensions of the head. a: rostrum; b: eye; c: total length of head; d: width of head. — Fic. 3. Dimensions of the thorax. a: pronotum; b: mesonotum; c: metanotum. — Fic. 4. Measure of length of the subgenital plate. 630 A. BENTOS-PEREIRA & C.H.F. ROWELL 7 Fics 5-8 5. Head of 3 of Proscopia septentrionalis. Lateral view. Scale bar 3 mm. — 6. Head of d of Proscopia panamensis. Lateral view. Scale bar 3 mm. — 7. Head of ? of Proscopia septen- trionalis. Lateral view. Scale bar 5 mm. — 8. Head of 2 of Proscopia panamensis. Lateral view. Scale bar 5 mm. REDESCRIPTION Male: Genitalia (Figures 9, 11). Anterior transverse plate (1) united with the lophi (2), which terminate in strong hooks, curving upwards and converging towards the midline. This unitary plate is equivalent to an epiphallus. It is located in a delicate membrane which extends towards the anterior part of the aedeagus. Laterally this membrane unites the transverse plate with lateral plates (10) in the same plane. The same membrane extends posteriorly, covering the median slit (3), which extends almost to the posterior extremity of the aedeagus. In this species (but not in panamensis) the slit 1s closed posteriorly and below by the fused lateral plates (4), which form a very obtuse angle extending from in front of plate (1) almost to below plate (10). The fused lateral plates (4) articulate with the anterior complementary plates (6), which are not straight but twisted. The median slit communicates with a pleated membranous chamber, in the floor of which opens a strongly sclerotized duct. This duct is tubular and opens proximally by means of a sclerotized valve. It is probable that during copulation the duct is not everted via the median slit, but that the pleated chamber acts as the intromittent organ, as in the closely related genus Astromascopia (Bentos, unpubl.), and that the sclerotized duct acts as an ejaculatory duct. CENTRAL AMERICAN PROSCOPIIDS 631 13 14 Fics 9-14 9. Aedeagus of Proscopia septentrionalis. Dorsal view. Scale bar 1.5 mm. — 10: Aedeagus of Proscopia panamensis. Dorsal view. Scale bar 1.5 mm. — 11. Aedeagus of Proscopia septen- trionalis. Lateral view. Scale bar 1.5 mm. — 12. Aedeagus of Proscopia panamensis. Lateral view. Scale bar 3 mm. — 13. Spermatheca of Proscopia septentrionalis. Scale bar 1 mm. — 14. Spermatheca of Proscopia panamensis. Scale bar | mm. 632 A. BENTOS-PEREIRA & C.H.F. ROWELL External morphology. Head delicate and elongate, markedly constricted behind the large globular eyes. Rostrum short, slightly inclined downwards, dorso-ventrally compressed, pointed tip somewhat rounded, the four edges weakly carinate (Figure 5). Coronary suture present. A small weak carina runs from the gena to the base of the eye. Antenna with 7 segments, antennal organs present on the sixth and seventh segments. Thorax. Prothorax tubular, anterior and posterior edges of the pronotum smooth, the latter forming a raised annulus. Anteriorly and laterally there are two slight depressions lighter in colour than their surround, and medially two others matching the surround. Integument ornamented with relatively large tubercles, particularly noticeable above the well-developed pleural suture. Apterous. Mesonotum and metanotum not inflated. Midline with tegumentary ornamentation, but finer and less marked than in the prothorax. Mesothoracic pleural suture smooth, pleura without ornament. Metathoracic pleural suture carinate, pleura smooth. A single mid-line tubercle somewhat posteriorly on the metanotum. Prothoracic legs inserted near the midline of the prosternum, not laterally. Prothoracic femora with fine denticles, tibiae smooth, square in cross-section and with minutely serrate carinae, the external and internal ventral carinae bearing additionally 15 and 13 small spines respectively (15/13). Mesothoracic legs very similar, with 12/12 femoral spines. Metathoracic femur slightly inflated at the base, two dorsal carinae, integument finely denticulate. Metathoracic knees with two short dorsal spines. Metathoracic tibae with 17/11 small dorsal spines, those of the inner row being longer; two small ventral and two large dorsal tibial spurs. Abdomen. First abdominal notum similar to the metathoracic notum. The rest of the abdomen smooth, cuticle sparsely punctate, medial carina well-marked. Supraanal plate (Figure 17) with slightly convex margins terminating in a rounded process. Cerci simple, slightly inward curving, two thirds as long as supraanal plate; subgenital plate short and rounded (Figures 15, 19). Coloration. Specimen CRIOO1 687843 was photographed alive in colour by the collector, P. De Vriess: the transparency shows a dark brown insect devoid of other markings. Female: Genitalia. Spermatheca of type 4 (DESCAMPS 1973), having a terminal ampulla with a small preapical diverticulum and one large and three small vermiform diverticula (Figure 13). TABLE |. Dimensions of ¢ of P. septentrionalis (mm). Specimen number INBio CRIOO1 687843 Length of head 9.9 Length of rostrum 1.38 Largest diameter of eye 2.88 Width of head (dorsally, at the level of the pronotum) 3.22 Length of pronotum 26.22 Length of mesonotum 259 Length of metanotum 2.36 Length of subgenital plate 2.67 CENTRAL AMERICAN PROSCOPIIDS 63 (US) 15 16 18 19 20 Fics 15-20 15. Tip of abdomen of d of Proscopia septentrionalis. Lateral view. Scale bar | mm. — 16. Tip of abdomen of 3 of Proscopia panamensis. Lateral view. Scale bar 1 mm. — 17. Supraanal plate of 3 of Proscopia septentrionalis. Scale bar 1 mm. — 18. Supraanal plate of & of Proscopia pana- mensis. Scale bar 1 mm. — 19. Subgenital plate of d of Proscopia septentrionalis. Scale bar I mm. — 20. Subgenital plate of d of Proscopia panamensis. Scale bar 1 mm. 634 A. BENTOS-PEREIRA & C.H.F. ROWELL External morphology. Larger and more robust than the d, but generally very similar. Head. Rostrum as in d but slightly inclined upwards (Figure 7). The general shape of the head is somewhat less elegant than in the d; the same carinulae are present on the rostrum and on the genae. Coronal suture obsolete. Antennae broken. Thorax. Prothorax similar to that of d. The ornamention is proportionally somewhat coarser. There are no definite granules above the pleural suture, and the posterior margin of the pronotum is less marked. Meso- and metathorax very similar to d, not inflated and without wing rudiments. Pleura granulated, not smooth. A well- marked line of short spines on both episterna, similar to that of Corynorhynchus spi- nosus. Prothoracic legs as in ¢, bearing 12/13 small spines. Mesothoracic femur similar to prothoracic but with weaker carinae, tibia with 11/9 spines. Metathoracic femur slightly inflated basally, covered with denticles and bearing two prominent dorsal carinae. Metathoracic knees with two short dorsal spines. Metathoracic tibiae with 9/16 dorsal spines, smaller than expected in an insect of this size. Abdomen. First segment similar to the thoracic segments, but flatter and with fewer granules. Pleural suture almost obsolete, without the carinae present in the à. The remaining abdominal segments are completely smooth, with a weak medial carina. Supraanal plate with slightly thickened margin, ending in an acute process (Figure 21). Cerci small, conical, about one fifth the length of the supraanal plate. Subgenital plate rounded, with a spatulate projection medially, which is slightly narrower basally than apically (Figure 23). Ovipositor valves large and strong, smooth. Coloration. The specimen CRI002 510601 was photographed alive in colour by its collector, M. Zumbado. The photograph shows a uniformly dark reddish-brown insect, with a slightly paler abdomen, lacking all other markings. Tibial spines black. The specimen CRIOO1 687844 was similarly photographed by the collector, P. De Vriess. In this © the ventral edges of the first 4 abdominal tergites are conspicuously edged with white, forming a tapering lateral stripe. The two collecting localities are only a few kilometres apart. In the holotype © the pale markings are asymmetrical, being better developed on the left side than on the right. TABLE 2. Dimensions of 2 2 of P. septentrionalis (mm) Specimen number INBio INBio Holotype CRI002 CRIOO1 ABP 224 510601 687844 Length of head 14.71 13.85 15.3 Length of rostrum 3.06 2978 3.0 Largest diameter of eye 3.74 3.34 3.4 Width of head 6.76 6.69 62 (dorsally, at the level of the pronotum) Length of pronotum 35.66 31.94 Bow Length of mesonotum 6.31 4.47 5 Length of metanotum 4.23 4.84 Sal CENTRAL AMERICAN PROSCOPIDS 635 2. Proscopia panamensis sp. n. SPECIMENS EXAMINED Holotype. PANAMA: Prov. Panama: Km 7.5, El Llano-Carti Road (87°55'W, 9°17'N), 9.5.96, (Hermögenes Fernandez Marin), adult d, specimen nos. 98047, ABP 225 (STRI). Paratypes. PANAMA: Prov. Coclé: Cerro Copé, 830 m, 20.9.1997 (C.H.F. Rowell & A. Bentos-Pereira), adult 2, specimen nos 97532, ABP 81, alcohol specimen (ANSP). PANAMA: Same data as previous specimen (specimen no. 97533). @ larva, alcohol specimen (Museo Fairchild, University of Panama). PANAMA: Proy. Panama: Barro Colorado Island, 27.8.77 (R. Silberglied & A. Aiello), © larva, specimen no. 98048 (STRI). DESCRIPTION Male: Similar to the 6 of P. septentrionalis, from which it differs as follows: Genitalia (Figures 10, 12). The genital complex is larger, robuster and (at least in our specimens) more strongly sclerotized, especially the lophi and the lateral plates. The medial slit is not closed below by fused lateral plates; in this species these remain separate, and extend laterally and posteriorly as far as the anterior complementary lateral plates (6). These latter are of a similar twisted shape to those of P. septen- trionalis but are additionally folded back on themselves at the anterior end and are provided with sensory hairs. External morphology. In general less gracile than P. septentrionalis. Head (Figure 6). Rostrum short with truncate tip. Rostral edges weakly carinate, the carinae not continuing beyond the eyes. Medial carina of occiput short, not continuing poste- riorly beyond the midddle of the eyes. Cuticle smooth. Thorax. Pronotum with straight parallel lateral margins. Anterior margin smooth, posterior margin somewhat thickened and with a very slight medial embay- ment. Pleural sutures marked only by a line, not carinate. Cuticle of meso- and meta- notum with numerous granules and transverse striae. Prothoracic femur square in cross- section, with serrate carinae and small tubercles, tibia with 11/12 spines. Mesothoracic femur practically smooth, tibia with 11/12 spines. Metathoracic femur strongly thickened in its basal half, with two weak dorsal carinae, denticles sparser and smaller than in septentrionalis. Metathoracic tibia with 10/16 relatively large spines. Abdomen. Ist abdominal notum transversely striate, this condition continuing on succeeding abdominal segments in the midline only, the remaining areas smooth. Medial carina absent. Subgenital plate (Figure 16) normally short and rounded, longer and more pointed when the aedeagus is protruded (Figure 20). Supraanal plate shield- shaped with a sharply pointed medial terminal process (Figure 18). Coloration. General coloration blackish-brown, tibial spines black. Female: The 2 2 of the species septentrionalis and panamensis are easier to distinguish than are the d d, especially by the structure of their spermathecae and subgenital plates. They are of the same length but P. panamensis is more gracile than P. septentrionalis. Genitalia: Two spermathecae present, both with long, fine and convoluted ducts with numerous diverticula. The two distal ampullae differ. One is large, entire and 636 A. BENTOS-PEREIRA & C.H.F. ROWELL Fics 21-24 21. Supraanal plate and ovipositor valves of Proscopia septentrionalis. Scale bar 5 mm. — 22. Supraanal plate and ovipositor valves of Proscopia panamensis. Scale bar 5 mm. — 23. Subgenital plate of ® of Proscopia septentrionalis. Scale bar 5 mm. — 24. Subgenital plate of 2 of Pro- scopia panamensis. Scale bar 5 mm. CENTRAL AMERICAN PROSCOPIIDS 637 TABLE 3. Dimensions of 4 of P. panamensis (mm) Specimen number 98047, ABP 225 (Holotype) Length of head Sal Length of rostrum 1.66 Largest diameter of eye 2.63 Width of head (dorsally, at the level of the pronotum) 4.2 Length of pronotum 23:69 Length of mesonotum 4.8 Length of metanotum 2252 Length of subgenital plate 1.39 smooth; the other is smaller and divided into two diverticula, apical and preapical, the latter with a smaller digitiform prolongation (Figure 14). External morphology. Head. Rostrum as in d (Figure 8). Seen from the side the dorsal margin is decidedly more sinuous than in septentrionalis. Thorax. Pronotum with straight margins and spiny cuticle. Some spines are large and conspicuous, particularly above the pleural suture. Margins as in P. septentrionalis. Meso and metanotum with a wide median band of granular cuticle. The extreme posterior of the metanotum is elevated and pleated, ending in a raised wavy margin. Pleural suture smooth in the mesonotum and marked by a carina in the metanotum. Pleura granular. The metathoracic epimeron has a medial carina. Apterous. Prothoracic leg. Femur rounded and smooth; tibia quadrangular in cross section, 13/12 spines. Mesothoracic leg similar, 11/15 tibial spines. Metathoracic femur basally inflated and bearing 2 medial carinae, each with a line of tubercles; ventral surface with a single weak carina. Metathoracic knees with two short dorsal spines. Femoral cuticle is granular and similar to that of the pronotum. 16/11 tibial spines. The two more ventral tibial spurs are smaller, the two dorsal spurs somewhat larger and stronger. Abdomen. The Ist abdominal segment bears a small anterior tubercle and trans- verse striations. The remaining abdominal segments smooth and minutely punctate, with a weak medial dorsal carina. Subgenital plate rounded and smooth, supranal plate pointed, cerci small and pointed. Ovipositor large and strong. Edges of the valves minutely toothed (Figures 22, 24). Coloration. The living 2 is olive green-brown on the head and thorax. The abdominal tergites are dark blackish brown with light yellow ventral margins, forming a conspicuous pale stripe along the sides of the abdomen from segments 2-8. Supraanal plate light brown, contrasting with the other abdominal tergites. Eyes reddish brown. Legs, dark brown, shading to black distally. 638 A. BENTOS-PEREIRA & C.H.F. ROWELL TABLE 4: Dimensions of 2 2 of P. panamensis (mm). Specimen number Adult, Larva, 98048 Larva 91532, 97338, ABP 81 alcohol specimen Length of head 12.6 7.6 11.4 Length of rostrum 129 1.66 15S Largest diameter of eye 33 2.63 2.1 Width of head 6.2 5.41 52 (dorsally, at the level of the pronotum) Length of pronotum 52% 24.68 26.7 Length of mesonotum 3:3 3a 4.0 Length of metanotum 4.8 3.88 4.1 3. Proscopia sp. SPECIMENS EXAMINED PANAMA: Prov. Panama: Barro Colorado Island, 24 May 1957 (Schrader) adult à, specimen no. 98049 (ANSP). Specimen bears labels "A86 BCI. Panama 24.5.57 Schrader coll.", "Compared with type, ANSP has type. Proscopia septentrionalis è Bruner. Det. Rehn 1958" in the handwriting of J.A.G. Rehn. "Male. Genitalia already removed (NDJ)" in handwriting of N.D. Jago. COSTA RICA: Prov. San José: Pozo Azul de Pirris, June 1905 (M. A. Carriker), juvenile 9, specimen no. 98050. Originally described by Bruner as the d type of T. septentrionalis (ANSP). As indicated on the label cited above, the genitalia were dissected from the only d specimen (by Rehn?) prior to Jago's examination in the 1980s, but are no longer present and must be assumed lost. The external morphology differs considerably from that of the previous species, and appears to indicate a different taxon. In the absence of an adult 2 specimen and of the d internal genitalia, however, the possibility cannot be excluded that either P. panamensis or P. septentrionalis is morphologically very variable or polymorphic and actually includes these specimens. The differences are as follows: Male: Head. Fastigium shorter than in septentrionalis, and somewhat more truncated than in panamensis. A medial carina runs from the tip of the fastigium to the occiput. Thorax: A prominent medial carina runs along the entire thorax. Thorax without ornament, either dorsally or on the pleura, and lacking a metathoracic median tubercle. The pronotum is divided into two clearly marked regions, the anterior with a granular cuticle, the posterior smooth. The legs are damaged but seem different from those of the those other two species. Abdomen. Well marked medial carina. Supraanal plate with a sharply pointed terminal process. Subgenital plate rounded and obtuse as in other Proscopia species. Female (larva): Rostrum identical to that of d in shape and ornamentation, but proportionately longer. The adult © rostrum is probably longer that that of septentrionalis. CENTRAL AMERICAN PROSCOPIIDS 639 Thorax. Meso- and metanotum have one lateral additional carina relative to the other two species. Legs: these are in better condition than in the d specimen. Pro- thoracic femur square in cross-section with serrate edges, tibia similar with 16 external and 12 internal spines. Mesothoracic femur (only segment preserved) similar to that of prothorax. Metathoracic femur scarcely at all inflated at base, two well-marked dorsal carinae, knees with two large dorsal spines. Metathoracic tibia square in cross-section with serrate edges, 8 internal and 17 external spines, which are large and expanded at the base. Spurs identical to those of other species. Abdomen. Terga similar to meso- and metanotum. Tip of supraanal plate broken, probably as in d. Subgenital plate straight. TABLE 5. Dimensions of d of P. sp. (mm) Specimen number 98049 Length of head 9.0 Length of rostrum 2.0 Largest diameter of eye 22 Width of head (dorsally, at the level of the pronotum) 5.8 Length of pronotum 18.25 Length of mesonotum (broken) Length of metanotum 2.80 Length of subgenital plate (broken) DISCUSSION GENERIC ATTRIBUTION JAGO (1989: 273) defined the genus Corynorhynchus on the structure of the d genitalia. He clearly states that the phallic complex of this genus lacks the transverse plate (1), that the lophi are greatly reduced and that the lateral plates (4) are reduced to a pair of angular spicules. The genus also lacks supplementary plates (6) and the genital duct is not sclerotized. The two species which we describe above do not fit this description at all. The presence of a transverse plate (1), strongly fused to prominent and well sclerotized lophi and supported by lateral complementary plates (6), and the presence of well- developed lateral plates (4), are by contrast in accordance with JAGo's definition (1989: 282) of Proscopia. Additionally, his description of the manner in which the sclerotized 3 genital duct joins the membrane in Proscopia corresponds exactly to the structure we see in our specimens. Our specimens conflict in some characters with Jago's definition: especially, in panamensis the lateral plates (4) are not fused posteriorly (though they are in septentrionalis) and they form a pod-like structure similar to that of Cephalocoema. There are also differences in the supraanal plate. However, Jago examined only two species of Proscopia (aberrans Hebard and gigantea Klug) and his definition may be unduly restrictive. No other genus fits our material better. In this article we also present data on the © spermatheca, which has previously been shown (BENTOS-PEREIRA 1997) to be useful within this group, particularly to 640 A. BENTOS-PEREIRA & C.H.F. ROWELL distinguish between closely related species. DESCAMPS (1973) did not describe the spermatheca of Corynorhynchus, but in other investigations (Bentos-Pereira, unpu- blished) it has proven to be of his Type 1, a simple spermatheca consisting of an ampulla at the end of a long duct. As described above, the spermathecae of the taxa described here are in contrast extremely complex; P. panamensis has a Type 4 sper- matheca with two independent terminal ampullae and numerous diverticula, while that of P. septentrionalis is somewhat less complex, with one terminal ampulla and a single tube but with various diverticula. We have also found the subgenital plate to be useful, especially in the case of P. septentrionalis. This character was first used within the Proscopiidae by LIANA (1972, 1980). Although it is not as definitive as the aedeagus or the spermatheca, it supplies useful information. All the described species of Corynorhynchus are markedly sexually dimorphic in the structure of the rostrum: the d rostrum is roughly conical, but that of the $ has larger or smaller apical expansions. The rostra of Proscopia, on the other hand, like those of the closely-related (vide JAGO 1983) Cephalocoema, are not more dimorphic than one would expect from the difference in size between the sexes. In this character too the taxa treated here agree with other species of Proscopia, but not with those of Corynorhynchus. All in all we have no doubt that the former is the correct genus to which to assign the Central American proscopiid species. In the absence of the internal genitalia we cannot be sure as to the generic placing of the third taxon described above, but to judge from the external morphology it too probably belongs to Proscopia. DISTRIBUTION P. septentrionalis has long been known as the most northerly proscopiid, and Pozo Azul de Pirris near the south western coast of Costa Rica is often quoted as the northern limit of the family. The great rarity of the Central American species, however, and their very cryptic form and coloration, makes us doubt the necessity of this conclusion. It seems quite possible that these or similar species have a wider distri- bution in Central America than is currently known, at least on the Caribbean side, which has much wet forest. This speculation is supported by the probable presence of a third, as yet undescribed, species in both Panama and Costa Rica, which has previously evaded detection. ACKNOWLEDGEMENTS We thank A. Solis (InBio, Costa Rica), D. Quintero (University of Panama), A. Aiello (STRI, Panama) and D. Azuma (Academy of Natural Sciences, Philadelphia) for permission to examine specimens from their collections, D. Quintero, A. Aiello and N. Scott Pezet for logistic help in Panama, C. S. Carbonell for help and useful discussion and P. De Vriess and M. Zumbado for photographs of living specimens. Our Panamanian field work was performed during a visit to the Smithsonian Tropical Research Institute in Gamboa, Panama. CENTRAL AMERICAN PROSCOPIIDS 641 REFERENCES BENTOS-PEREIRA, A. 1997. El genero Astromascopia Jago 1989 (Orthoptera - Proscopiidae). Tesis, PEDECIBA, Universidad de la Republica, Montevideo. BRUNER, L. 1905. Two remarkable new Costa Rican locusts. Entomological News 16: 313-315, pl 1. BRUNER, L. 1908. Acrididae. /n: Frederick DU CANE GODMAN (ed.). Biologia Centrali Americana. Insecta, Orthoptera, 2: 1-342, plates 1-4 (1900-1909). London, published for the Editor by R.H. Porter, 1893-1909. CARBONELL, C.S. 1977. Orthopterorum Catalogus 17. The Hague: Dr. W. Junk. 29 pp. CAUDELL, A.N. 1911. Some remarks on Kirby's Synonymic Catalogue of Orthoptera, Vol. 3, with additional notes on Vols. 1 and 2. Entomological News 22: 158-167. Descamps, M. 1973. Notes préliminaires sur les genitalia de Proscopioidea (Orthoptera, Acrido- morpha). Acrida 2: 77-95. HEBARD, M. 1924. Studies in the Acrididae of Panama (Orthoptera). Transactions of the Ameri- can Entomological Society 50: 75-140, plates VI-VII. Jago, N.D. 1989. The genera of the Central and South American grasshopper family Pro- scopiidae (Orthoptera: Acridomorpha). EOS (Madrid) 65: 249-307. KIRBY, W.F. 1910. A synonymic catalogue of Orthoptera. Vol. 3, Orthoptera Saltatoria. Part 2 (Locustidae vel Acrididae). British Museum, London. 674 pp. KLUG, F. 1820. Proscopia, novum Insectorum Orthopterorum genus. Horae Physicae Bero- linensis, Bonnae, 15-26, plates 3, 4. Liana, A. 1972. Etudes sur les Proscopiidae (Orthoptera). Polska Akademia Nauk, Institut Zoolo- giczny, Annales Zoologici 29: 381-459. LIANA, A. 1980. Materiaux pour la connaissance des Proscopiidae (Orthoptera). Mitteilungen des Hamburger zoologischen Museums und Instituts 77: 229-260. MELLO LEITAO, C. 1939. Estudio monogräfico de los proscöpiidos. Revista del Museo de La Plata (NS) 1: 280-449, pl XII. OTTE, D. 1978. The primary types of Orthoptera (Saltatoria, Mantodea, Phasmatodea & Blattodea) at the Academy of Natural Sciences of Philadelphia. Proceedings of the Academy of Natural Sciences, Philadelphia 130: 26-87. AT nn nni ES REVUE SUISSE DE ZOOLOGIE 106 (3): 643-661; septembre 1999 Glomeris undulata Koch and G. conspersa Koch are conspecific. - Enzyme electrophoretic evidence and taxonomical consequences (Diplopoda: Glomeridae) René HOESS & Adolf SCHOLL University of Berne, Institute of Zoology, Division of Population Biology, Baltzerstr. 3, CH-3012 Berne, Switzerland. Glomeris undulata Koch and G. conspersa Koch are conspecific. - Enzyme electrophoretic evidence and taxonomical consequences (Diplopoda: Glomeridae. - Syntopic and allotopic populations of Glomeris undulata and G. conspersa have been investigated with enzyme electropho- resis. There was no evidence for separate gene pools of these taxa. However, G. romana, a species supposed to be closely related to G. undulata and G. conspersa, prooved to be well differentiated based on allozyme data. We therefore conclude that the taxa undulata and conspersa are conspecific. Consequently, G. conspersa C. L. Koch, 1847, is a junior subjective synonym of G. undulata C. L. Koch, 1844 (nov. syn.). An analysis of the colour pattern provides arguments for the discussion why no intermediate forms exist. Key-words: Diplopoda - Glomeris - revision - local alleles - morphology - distribution - new synonymy. INTRODUCTION Glomeris undulata C.L.Koch, 1844, and G. conspersa C.L.Koch, 1847, are traditionally treated as two closely related but distinct species (cf. VERHOEFF 1911). They may be separated with reliability only by the form of the dorsal dark spots on the segments 2 to 12. These spots are parallel-sided in G. undulata and convergent posteriorly in G. conspersa (seen in walking position). Transitional forms that could indicate interbreeding of these taxa are very rare and have been named G. undulata var. pseudoconspersa (but see also VERHOEFF (1928b) for a case of assumed hybridisation). G. undulata and G. conspersa are often found syntopic, and large parts of their ranges overlap. Within the scope of an analysis of the genetic differentiation of the Central European Glomeris species by means of enzyme electrophoretic methods, we also examined syntopic and allotopic populations of G. undulata and G. conspersa. For comparison we included G. romana Verhoeff, 1900, a species which is supposed to be closely related to G. undulata and G. conspersa (cf. VERHOEFF 1911) but with a mainly allopatric distribution. Manuscript accepted 12.01.1999 644 RENE HOESS & ADOLF SCHOLL MATERIAL AND METHODS Population samples of Glomeris undulata, G. conspersa, and G. romana were analyzed genetically by using routine enzyme electrophoretic methods of our labo- ratory (SCHOLL et al. 1978). Vertical starch gel electrophoresis was conducted using the same buffer systems as in previous studies on Glomeris (HOESS et al. 1997). 18 enzyme loci were analyzed. The enzymes investigated and the loci scored (in brackets) are: aspartate aminotransferase (Aat-1, Aat-2), glyceraldehyd-3-phosphate dehydrogenase (Gapdh), glucose-6-phosphate isomerase (Gpi), hexokinase (Hk), leucine aminopeptidase (Lap), L-lactate dehydrogenase (Ldh-1, Ldh-2), malate dehydrogenase (Mdh-1, Mdh-2), malic enzyme (Me), mannose-6-phosphate isome- rase (Mpi), peptidase (Pep), 6-phosphogluconate dehydrogenase (Pgd6), phospho- glucomutase (Pgm), superoxide dismutase (Sod-1, Sod-2) and sorbitol dehydrogenase (Sodh). The zymograms were photographed on Polaroid for reference. We refer to observed electromorphs as alleles which are identified by their electrophoretic mobility (in mm) relative to electromorphs of previously studied species (HOESS et al. 1997). Mendelian inheritance of the alleles at a given locus was not tested by cross- breeding experiments but was concluded by analogy to results of previous studies of our laboratory (e.g. ZIMMERMANN & SCHOLL 1993). Allele frequencies and genetic distances were calculated with the BIOSYS-1 programme package (SWOFFORD & SELANDER 1989). Nei-distance D (NEI 1978) was used for the construction of an UPGMA dendrogram (SNEATH & SOKAL 1973). Bootstrap analysis was conducted with the PHYLIP programme package (FELSENSTEIN 1986-95). The bootstrap values are estimates from 100 replicates (bootstrap values levelled out at 100 replicates) with UPGMA as cluster algorithm and an unrooted consensus tree. The following population samples (Fig. 1) were examined (number of speci- mens in brackets): Glomeris undulata: Switzerland: Bösingen (13), Brunnen (9), Ingenbohler Berg (15), Loucherhorn (6), Malans (6), Meride (7), Merishausen (8), Niderhorn (6), St-Gingolph (15); Germany: Ehingen (17); G. conspersa: Switzerland: Biel (10), Bösingen (20), Castagnola (3), Chatollion (14), Densbiiren (10), Ingen- bohler Berg (11), Loucherhorn (8), Malans (12), Meride (8), Merishausen (13), St-Gingolph (11), Valangin (10); Germany: Ehingen (20); France: Cruseilles (13); G. romana: Italy: Camaiore (18), Lucca (4). Several other population samples have been collected (often with low sample sizes) and have been used in addition to literature data to produce a distribution map of the taxa undulata and conspersa. Tergites of two selected specimens of G. undulata were drawn using a camera lucida. The specimens were turned around their axes in order to draw every part of the tergite in vertical view. RESULTS Allele frequencies of the 18 loci of all samples are given in Table 1. For unknown reasons, several specimens in most samples failed to show activity at some REVISION OF GLOMERIS CONSPERSA Ehingen Merishausen An I RS @ Densbüren i (\ CH iy #3 @ Malans Ingenbohler Berg Bösingen Pan | ° Niderhorn i St-Gingolph © 4 >> Biel Chätollion DoS ® Loucherhorn Cruseilles Fic. 1 Collecting sites in Switzerland and adjacent countries. Camaiore 645 & ADOLF SCHOLL RENE HOESS 646 Ovo EEO - 600 TEO 670 900 - 3 = © 900 £TO - 900 €0'0 800 STO i j i IO EGO = SEQ = 970 SITO i 110 £S0 SS0 900 05°0 = 3 = - _£0'0 SIO S0'0 j i 900 070 600 90'0 €0° OKO), De 2% z = = SCO 9]9]]V SNO] = ie] Gai =; we un ies! i SEI ee > Elsa 0 Be A 2 za log 8 SR. © © NE a @ wes == to = E: FTT cD dd © Q SIC NT LS D 3 = © > sr oO 5 © 5 © È = = = sr je OS = D © o ho 4 5 = 2 = 2 ol aS) È 3 I Da 5 3 lai =P 5 œ œ © ig’) aa aa DUDUIOA on psiadsuod “Dp DIDINpun *H ‘sn90] 2A1H99dS91 ou} Je paloos suaundads JO Joquinu = N ‘DUDIUOI 1) pue vssadsuod “D ‘pipynpun S142W0]9 UL paıpnys 190] aUIAZUS QT AU} je sorauanbaay 9[9A][V - | FIAVL 647 REVISION OF GLOMERIS CONSPERSA me en RR 2.7002 = È = a = 3 3 7 È 2 È 3 3 È = {VII MH TO a OLO 80°0 €90 L90 TIO 090 LL0 6€0 060 SS’O 0S°0 SEO 050 IL'0 MO 2 = = 5 & ADOLF SCHOLL RENE HOESS 648 ee 1¢0 LIO 170 610 050 950 PEO 050 PHO 8€ 0 LEO PrO cr0 EEO [TO LI°0 £E0 LIO 050 6£0 SO 8£ 0 £90 LE'O 96 rr0 ECO 9770 Pro LrO £10 VEO 070 LIO 800 Fio 870 670 6901620 85°0 050 290 LV0 190 €770 95°0 I€0 £9'0 [001 STO 050 EEO I£O £00 1EO 670 O10 6€0 SO S70 870 670 £00 £0°0 900 900 90°0 610 Z7°0 SL'0 ZIO ÿl'O OTO 60°0 LIO SL'O 8L°0 83 0 980 080 160 STO 990 980 90'0 SLO LIO LIO OT €I 9 CC SOS E) v60 SL'OÏI80 LEO 880 +60 890 SL'0 890 LTO OL'0 050 S90 0£ 0 180 SSOÏE6O 760 £80 760 ELO E80 £90 TEO LEO 050 649 REVISION OF GLOMERIS CONSPERSA 81 LIO L0°0 — S80 E610 = 1 74610 23221900, 25036: Selig ge ee = 2 CUO 126380 ZI Cis 227 2050 ENTE = È ? Z 5 = = = 5 = = SOON CLONE ze ee nn Me Se sa OURS O0 RE I nr COUR AE LEE di ele dre in et ESTONE RO 8 a eG PIO i eek ase sete OFS TALS [WOLIGOES SONO ren a= E HOMO OO AO 820 a1 Zion al Ais SO I ee ieee me COO ea i? | POS ea OGM Von ARE £0°0 TL'O 8L'O|SL'O STO 050 SLO 050 CLO €60 LIO 800 L0°0 9 ON SÌ LIO 800 £00 & ADOLF SCHOLL RENE HOESS 650 69°0 900 LI°0 LIO 760 TE'O 670 910 S60 ITO LEO S00 SL'O 980 610 600 FIO 050 09°0 OT'0 070 800 070 LO'0 £0°0 060 60 [TO LL'O 05°0 STO TTO €70 070 E10 09°0 1840 98°0 Le 651 LOMERIS CONSPERSA J REVISION OF € 90°0 LI°0 160 €800 STO SL'O €80 SL'O 9€0 LEO L90 980 870 €0°0 0°0 OTO FIO 080 980 S60 61 6£°0 €EO €0°0 L90 85’0 €8 0 180 L480 980 090 S80 €1'0 $00 IEO 050 LVO tt'O LIO ITO 180 70 9 eg! E = = €£€0 E00 0800 Z60 TO #90 STO 010 SLO 690 CL'O T0 LO'0 80°0 ££'O O10 c60 090 #90 L90 [80 Lh'0|00I 1€0 €10 0E°0 OS'0 £70 [86 & ADOLF SCHOLL RENE HOESS 652 ee Men AEH OE 160 Sco ey pro =. iro, ore Beagle = RS a -< er are coo Spd sro Ii 2000) 800 860 IPL SER Red Et: ee sy ee 0 Ne STO BO TION e |<: eee 00K le 000 A (cs = ie SO === S00 (NO PHOTO 000 LEO. 110 O10 720 6:0 | = 601000 Syn SINGH HG 5 = 5 2 = = > = > i z 7 a z A x = = - = È 5 > = [96 SD PS NE NC UE ROUES 050 = = 200 SCOR US te 760 €80 S60 690 #80 ZLO FEO OO ZLO 6LO 190 090 890 SL'O|GL'O 650 OSO ESO 080 #60 £90 110 850 890 JOO! D NON SD TO 11:0. LIO = 060 = > = 060 50:0, = 1/2010 £60 ALTO O10 = 01:0) No) pono) coven TE ey ee oe le fi CT ao om Tr II cnr 653 REVISION OF GLOMERIS CONSPERSA 90° IGOR SOS ss oO 1 ' ' 61°0 STO IT i À evo pl'O i i i i 70 870 ETO STO £€0 Str0 À : ; 670 E i i i i 0 €£0 €50 880161 \ \ i 0 050 YIO LOO ErO OF \ i i À i i 61°0 S80 £0 670 90°0 |” À j j ‘0 €TO €770 050 LS0 05 i ; ‘0 6£0 ETO ECO 770 LIO OVO OL : S£O STO Coe eles Ol 9 9 c60 €800 S80 060 SL’ i i ‘0 €80 +80 #8°0 IL'O 0S°0 ILO 190 800 LIO 110 O10 ST i i à 800 800 STO 0S°0 670 870 LIO 800 Sav: 8 5 Ol 8 ol ww "9 Of) G Gh” © OF VN ol & ADOLF SCHOLL RENE HOESS 654 €10 90°0 €10 £00 760 €60 960 900 L0°0 REVISION OF GLOMERIS CONSPERSA 655 loci or their zymograms were not scorable. These specimens were not included for calculations of allele frequencies at the respective loci. Therefore, the number of specimens is not always identical in Table | for all loci scored in a particular popu- lation. Most enzyme loci are highly polymorphic in all three taxa, viz. Glomeris undu- lata, G. conspersa and G. romana. Although the samples of G. undulata and G. conspersa cover large parts of the ranges of these taxa, only minor variation in allele frequencies of the respective populations was observed. While G. romana differs from G. undulata and G. conspersa at several loci by allele substitution (Sod-1) or highly different allele frequencies (Aat- 1, Gapdh, Ldh-2, Mdh-1, Mpi, Pgm, Sodh), G. undulata and G. conspersa, in contrast, usually show the same alleles and very similar frequencies. The analysis of the distribution of rare alleles reveals that some local alleles which were each found at one locality only, are present in both G. undulata and G. conspersa from the particular locality. These local alleles are: Aat-2°9 in the Merishausen sample, allele Me?? at Meride, allele Ldh-2% at Ehingen, allele Ldh-2°0 at Malans and allele Sodh®® at Meride. The UPGMA dendrogram that resulted from a cluster analysis of the popu- lations according to their genetic distances is shown in Fig. 2. G. romana is clearly separated from the other two taxa with an average genetic distance (Nei-D) of 0.65 and a bootstrap value of 100. G. undulata and G. conspersa, in contrast, do not form separate clusters. The syntopic samples are of particular interest because in seven from eight such situations studied, G. undulata and G. conspersa from the same locality cluster with lowest genetic distances, and four of these clusters are supported by bootstrap values > 70 (Fig. 2). Genetic distances among the undulata/conspersa popu- lations are rather low. Thus, separations of the samples according to geographical regions or differentiations according to geographical distances are not evident. DISCUSSION Several species of the genus Glomeris are known to be highly variable in their colour pattern, and about 400 varieties, including the nominotypic varieties, have been described which belong to about 70 nominal species (many names, however, are not available according ICZN Art. 16 + 45f, g). If the biological species concept (e.g. Mayr & ASHLOCK 1991) is applied that defines species as genetic units of naturally interbreeding populations which are reproductively isolated from other such groups, allozyme data provide a very powerful tool to test whether or not two taxa or forms share a common gene pool and thus belong to the same species. Species boundaries and specific separation within the genus Glomeris have recently been successfully analysed with allozyme data (HoEss et al. 1997) where G. intermedia Latzel, 1884, was shown to be a separate species, contrary to traditional taxonomy that treated G. intermedia as a subspecies of G. hexasticha Brandt, 1833. G. undulata and G. conspersa have always been treated as two distinct species which are assumed to be closely related. Intermediate forms that might indicate hybrids have not been observed even though both taxa are often found syntopic and 656 RENE HOESS & ADOLF SCHOLL 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 e e er Nei-distance D 100 Ehingen (c) Ehingen (u) Biel (c) St-Gingolph (c) 78 St-Gingolph (u) Densbüren (c) - Chatollion (c) Merishausen (u) Merishausen (c) Ingenbohler Berg (c) 92 Ingenbohler Berg (u) Brunnen (u) Meride (c) Meride (u) 78 Malans (c) Malans (u) Bosingen (c) — Bösingen (u) Loucherhorn (c) Loucherhorn (u) Valangin (c) G. undulata/conspersa 100 Niderhorn (u) Cruseilles (c) Castagnola (c) Camaiore G. romana Lucca FIG. 2 UPGMA dendrogram based on Nei-distance D values in pairwise comparisons of populations | of Glomeris undulata (u), G. conspersa (c) and G. romana. Bootstrap values > 70 are indicated (percentage over 100 replicates). REVISION OF GLOMERIS CONSPERSA 657 large parts of their ranges overlap. Only VERHOEFF (1901) described a light form of G. undulata as var. pseudoconspersa indicating to him a mixture of the colour pattern of both taxa, and, later, postulated some hybrids (VERHOEFF 1928b, p. 320-321). Thus, as judged from the colour patterns, specific separation of G. undulata and G. conspersa seems to be completed despite some rare possible hybrids. The genetic basis of va- riation in colour patterns, however, has never been analysed in Glomeris. The difference in the colour patterns of G. undulata and G. conspersa might result from two alleles in a dominant/recessive relation at a single locus. In this case, we would not expect to find intermediate forms. ALLOZYME DATA Based on the allozyme data, there is no evidence for separate gene pools of G. undulata and G. conspersa. The alleles observed and their frequencies are rather similar in all populations of both taxa. Consequently, the degree of genetic differen- tiation of all populations studied is very low. Furthermore, the analysis of the distri- bution of rare alleles is very informative because it revealed that local alleles (cf. BusAck 1986) are repeatedly found in syntopic populations of both taxa. The Aat-2 locus is of particular interest because this locus is highly conservative in Glomeris (Hoess et al. unpublished), most Glomeris species are fixed for the allele Aat-2100, and allele Aat-2° is only found at the locality Merishausen where it was recorded in both G. undulata and G. conspersa. In the dendrogram, the populations of G. undulata and G. conspersa do not form separate clusters, but syntopic populations of both taxa usually cluster at lowest levels of genetic differentiation, and most of these clusters are supported by bootstrap values > 70. Consequently, we regard the taxa G. undulata and G. conspersa as a single species (see "Conclusion" below) and treat conspersa in the following as a form of G. undulata. In comparison to G. undulatalconspersa, G. romana clearly has a separate gene pool. Genetic differentiation, however, is rather low. This agrees with current classification where G. romana is assumed to be closely related to G. undu- lata/conspersa (VERHOEFF 1911). COLOUR PATTERN Although both forms of G. undulata (nominate form and f. conspersa) are often found syntopically, no transition in colour pattern is known. In fact, G. undulata var. pseudoconspersa and the "hybrids" of VERHOEFF (1928b) do not represent transition forms at the dorsal spots, that means from the parallel spots of undulata to the conver- gent spots of conspersa. Instead, VERHOEFF (1928b) used the colour pattern of the remaining parts of the tergites to qualify these specimens as transitional. Consequently, the question remains: why are the two forms of G. undulata separated so strictly? A specially patterned G. undulata specimen (from near Mendrisio in Ticino, Switzerland) may help to understand the real nature of the colour pattern of G. undu- lata. The fifth tergite of this specimen is illustrated in Fig. 3a. Unlike most individuals of this species where the speckles seem to be scattered irregularly next to the central spot, the speckles of this specimen are arranged in a sinuous line that strikingly 658 RENE HOESS & ADOLF SCHOLL FIG. 3 Vertical view of the fifth tergite of Glomeris undulata: a) a specimen from Mendrisio (Switzerland), b) a specimen from Ingenbohler Berg (Switzerland), c) schematic diagram of the basic pattern of G. undulata (sinuous line) with the nomenclature of the dark spots sensu VERHOEFF (1928a), d) “normally coloured” specimen of nominate form, e) "normally coloured" specimen of f. conspersa. REVISION OF GLOMERIS CONSPERSA 659 resembles the dark spots of many other Glomeris species. Possibly, this specimen reflects the original pattern of G. undulata. Varieties with a light spot IV (sensu VERHOEFF 1928a) between the dark spots IV (as in the illustrated specimen) are already known. But this pattern cannot explain the difference in the central dark spot between the nominate form and f. conspersa. However, other varieties are known - e.g. the specimen shown in Fig. 3b (from Ingenbohler Berg) - that show an additional dark spot in the light spot IV. From these two specimens we can conclude a basic pattern for the species which is illustrated in Fig. 3c. The sinuous line (with an extra line in the region of the muscular insertions on the prozonite) may be traced from one side to the other. In real specimens, certain parts of this sinuous line may be invisible due to non-expression of the dark pigment or they may be connected by dark pigment filling up the space between the dark spots. Thus, we can conclude that in the nominate form of G. undulata the dark spots IV are well expressed and normally connected, and that in f. conspersa only the dark spot within the light spot IV - in Verhoeffs terminology this dark spot was treated as part of the dark spots IV - is well expressed and the normal dark spots IV are missing. With this argumentation no transition between the nominate form of G. undulata (Fig. 3d) and the form conspersa (Fig. 3e) can be imagined. A single gene or a group of coupled genes may be responsible for the expression or absence of the dark spots IV. DISTRIBUTION Fig. 4 shows the distribution of G. undulata (nominate form and f. conspersa) based on a compilation of literature data and on own collections. In large parts of its range, G. undulata is present only in one form. The reasons for this sitation are not known. But if the above hypothesis of the genetic basis of the two forms would be confirmed experimentally, the discordant distribution of both forms could be explained by local differences in allele frequencies, either due to selection or stochastic processes. CONCLUSION As we have shown, there is no evidence for separate gene pools in syntopic populations of Glomeris undulata and G. conspersa. Thus, we conclude that these taxa are conspecific. Therefore, we consider G. conspersa C.L.Koch, 1847, as a junior subjective synonym of G. undulata C.L.Koch, 1844 (nov. syn.). There is no need to preserve the better known name G. conspersa because G. undulata has been used many times in the literature (at least 37 entries for G. undulata and at least 69 entries for G. conspersa). ACKNOWLEDGEMENTS We thank L. Beer, L. Frauchiger and V. Siegfried for assisting in the laboratory work. P. Benninger, M. Braunwalder, B. Gantenbein, R. Köpfli, M. Lörtscher, S. Minder, A. Pedroli-Christen, A. Schuppisser and P. Sonderegger helped us gather the glomerids used in this study. E. Obrecht made valuable comments on an earlier draft of the manuscript. 660 RENE HOESS & ADOLF SCHOLL G. undulata (nominate form) G. undulata f. conspersa both tons Fic. 4 Distribution of Glomeris undulata (nominate form and f. conspersa). The section covered in Fig. | is framed. REFERENCES Busack, S. D. 1986. Biogeographic analysis of the herpetofauna separated by the formation of the Strait of Gibraltar. National Geographic Research 2: 17-36. FELSENSTEIN, J. 1986-95. PHYLIP (Phylogeny Inference Package), version 3.5c, native-mode executables for PowerMac. Univ. Washington. Hoess, R., SCHOLL, A. & LORTSCHER, M. 1997. The Glomeris-taxa hexasticha Brandt and intermedia Latzel: species or subspecies? - allozyme data (Diplopoda, Glomerida, Glomeridae). Entomologica Scandinavica, Supplement 51: 133-138. REVISION OF GLOMERIS CONSPERSA 661 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 1985. International code of zoological nomenclature - 3rd edition. Charlesworth, Huddersfield, XX + 338 pp. KocH, C. L. 1844. Deutschlands Crustaceen, Myriapoden und Arachniden. /n: Deutschlands Insecten ... fortgesetzt von G. A. W. HERRICH-SCHÄFFER, Heft 40 (= G. W. F. PANZER, Fauna Insectorum Germanicae initia, Heft 190). Pustet, Regensburg. KocH, C. L. 1847. System der Myriapoden, mit den Verzeichnissen und Berichtigungen zu Deutschlands Crustaceen, Myriapoden und Arachniden, Heft 1-40. F. Pustet, Regens- burg, V + 272 pp. Mayr, E. & ASHLOCK, P. D. 1991. Principles of systematic zoology - 2nd edition. Mc Graw Hill, New York, XX + 475 pp. NEI, M. 1978. Estimation of avarage heterozygosity and genetic distance from a small number of individuals. Genetics 89: 583-590. SCHOLL, A., CORZILLIUS B. & VILLWOCK, W. 1978. Beitrag zur Verwandtschaftsanalyse altweltlicher Zahnkarpfen der Tribus Aphaniini (Pisces, Cyprinodontidae) mit Hilfe elektrophoretischer Untersuchungsmethoden. Zeitschrift fiir zoologische Systematik und Evolutionsforschung 16: 116-132. SNEATH, P. H. A. & SOKAL, R. R. 1973. Numerical Taxonomy. Freeman, San Francisco, XV + 573 pp. SWOFFORD, D. L. & SELANDER, R. B. 1989. BIOSYS-1. A computer program for the analysis of allelic variation in population genetics and biochemical systematics. Release 1.7., D. L. Swofford, Illinois Natural History Survey. VERHOEFF, K. W. 1901. Beiträge zur Kenntnis palaearktischer Myriopoden. XVIII. Aufsatz: Uber Diplopoden aus Süddeutschland und Tirol. Jahresheft des Vereins für vater- ländische Naturkunde in Württemberg 57: 81-111. VERHOEFF, K. W. 1911. Über Diplopoden. 20. (40.) Aufsatz: Neuer Beitrag zur Kenntnis der Gattung Glomeris. Jahresheft des Vereins für vaterländische Naturkunde in Württem- berg 67: 78-147. VERHOEFF, K. W. 1928a. Klasse Diplopoda. /n: Bronns Klassen und Ordnungen des Tier- Reichs, 5, II. Abtl., 1. Teil, 1.-6. Lieferung. Akademische Verlagsgesellschaft, Leipzig: XI + 1071 pp. VERHOEFF, K. W. 1928b. Neue und besonders ostalpine Chilognathen-Beiträge. (108. Diplo- poden-Aufsatz). Zoologisches Jahrbuch für Systematik 55: 253-328. ZIMMERMANN, M. & SCHOLL, A. 1993. Specific status of Aquarius cinereus (Puton) and A. najas (De Geer) (Hemiptera: Gerridae) and the extent of hybridisation in the Medi- terranean region. Entomologica Scandinavica 24: 197-210. pi ecm 214 a f 3 REPAS pi rae ats)! FA a rl EEE 1-4 ta ge na 4 ER è. | . DENT à | hi i li + 40 au POR al > pi. My (180 vie a! re La REVUE SUISSE DE ZOOLOGIE, 106 (3): 663-689; septembre 1999 Aleocharinae di Hong Kong (Coleoptera, Staphylinidae)* Roberto PACE Via Vittorio Veneto 13; I-37032 Monteforte d'Alpone (Verona), Italia. Aleocharinae from Hong Kong (Coleoptera, Staphylinidae). - In this paper, based on the study of new material collected by means of modern sampling methods 22 species are described as new. These new species belong to following tribes: Deinopsini (1 n. sp.), Pronomaeini (2), Gyrophaenini (2), Placusini (1), Homalotini (1), Deremini (1), Athetini (10), Pygostenini (1), Termitopaediini (1) and Myrmedoniini (2). Two new synonymies are proposed. The main diagnostic characters are illustrated. Key-words: Coleoptera - Staphylinidae - Aleocharinae - taxonomy - new species - Hong Kong. INTRODUZIONE Fino ad ora le specie della sottofamiglia Aleocharinae segnalate per Hong Kong città e suo territorio circostante, sono state raccolte occasionalmente e in modo sporadico. Al fine di una loro conoscenza il più possibile completa, erano necessarie ricerche mirate e l'adozione di tecniche di raccolta nuove. Così è stato effettuato dal collega Guillaume de Rougemont, noto studioso di Staphylinidae di Londra, che grazie ai suoi lunghissimi soggiorni a Hong Kong al fine di perfezionare il suo apprendimento della lingua cinese, ha curato anche la ricerca di Aleocharinae su tutto il territorio di Hong Kong. Per questo fine ha adottato varie tecniche di ricerca e ha scelto tra l’altro, come campo delle sue indagini, un’oasi naturalistica di Hong Kong, quale il “Kadoorie Agricultural Research Centre”, legato all’ Università di Hong Kong e proprietà dei fratelli Kadoorie, noti benefattori della città. L'intera serie di Aleocharinae raccolte da G. de Rougemont durante il suo lungo soggiorno a Hong Kong, sono state a me affidate in studio, insieme a piccole raccolte effettuate da altri entomologi inglesi, quali S.J. Reels e G. Ades. I risultati di questo esame sono il contenuto del presente lavoro. Gli holotypi delle nuove specie sono conservati al “Museum d’histoire naturelle” di Ginevra (MHNG). * (151° contributo alla conoscenza delle Aleocharinae) Manoscritto accettato il 13.02.1999 664 ROBERTO PACE ELENCO SISTEMATICO DELLE TRIBU E DELLE SPECIE DEINOPSINI Sharp, 1883 Adinopsis chinensis sp. n. (figg. 1-4) PRONOMAEINI Ganglbauer, 1894 (=Myllaenini) Myllaena hongkongiphila sp. n. (figg. 5-6) Myllaena reelsi sp. n. (figg. 7-8) GYROPHAENINI Kraatz, 1856 Gyrophaena (s. str.) anguli sp. n. (figg. 9-12) Pseudobrachida hongkongensis sp. n. (figg. 13-14) PLACUSINI Mulsant & Rey, 1871 Placusa kadooriorum sp. n. (figg. 15-19) HOMALOTINI Heer, 1839 Coenonica javana Bernhauer, 1914 Coenonica javana Bernhauer, 1914: 106 1 d, Hong Kong, Shek Kong, 20.11.1991, G. Ades leg.; 1 2, Hong Kong, Kadoorie Agricultural Research Centre, flight interception trap, X-XII.1996, G. de Rougemont leg.; | 4, Hong Kong, Lantau Is., 8.11.1997, G. de Rougemont leg. Distribuzione. Giava, Cina e Hong Kong. Anomognathus (s. str.) bohaci Pace, 1992 Anomognathus (s. str.) bohaci Pace, 1992; 126 1 ©, Hong Kong NT. sifted litter, 1.11.1997, G. de Rougemont leg. Distribuzione. Thailandia. Nuova per Hong Kong. Linoglossa (s. str.) hongkongensis sp. n. (figg. 20-22) DEREMINI Seevers, 1965 Demerinda rougemonti sp. n. (figg. 23-24) ATHETINI Casey, 1910 Hydrosmecta subalgarum sp. n. (figg. 25-29) Atheta (Acrotona) iperanomala sp. n. (figg. 30-32) Atheta (Acrotona) shekkongensis sp. n. (figg. 33-36) Atheta (Acrotona) tricholutea sp. n. (figg. 37-38) Atheta (Coprothassa) roridanotha sp. n. (figg. 39-42) Atheta (Microdota) vagans Bernhauer, 1907 Atheta (Microdota) vagans Bernhauer, 1907: 404; PACE 1998: 150. Atheta (Microdota) tronquetiella Pace, 1988: 328, syn. n. ALEOCHARINAE DI HONG KONG 665 1 d e2 ©, Hong Kong, Tsinfai Tong, 25.1V.1997, G.T. Reels leg. Distribuzione. Sri Lanka, Taiwan, Giappone, Cina. - Atheta (Microdota) ocularis Cameron, 1939 (figg. 43-46) Atheta (Microdota) ocularis Cameron, 1939: 327 Atheta (Microdota) triflexa Pace, 1986: 196, syn. n. 1 3, Hong Kong, C.U.R.K., flight interception trap, III-IV.1997, G. de Rouge- mont leg. Distribuzione. India, Giava, Hong Kong. Atheta (Microdota) pseudovagans sp. n. (figg. 47-50) Atheta (Microdota) persimplex sp. n. (figg. 51-54) Atheta (Poromicrodota) hoihaensis sp. n. (figg. 55-56) Atheta (Sipalatheta) algarum sp. n. (figg. 59-62) Tomoglossa fuliginosa sp. n. (figg. 63-64) Pelioptera opaca Kraatz, 1857 Pelioptera opaca Kraatz, 1857: 56; PACE 1998: 152 1 ©, Hong Kong, Kao Tam Tso, yellow pan, 18.V.1986, G.T. Reels leg. Distribuzione. Sri Lanka, Nepal, India, Birmania, Cina, Hong Kong, Sabah, Giappone. PYGOSTENINI Fauvel, 1899 Odontoxenus reelsi sp. n. (figg. 65-66) TERMITOPAEDIINI Seevers, 1957 Dioxeuta rougemonti sp. n. (figg. 67-69) MYRMEDONIINI Thomson, 1867 Zyras (Sinozyras subgen. n.) pygmaeus sp. n. (figg. 70-73) Zyras (Zyras) hongkongensis sp. n. (figg. 74-77) Zyras (Glossacantha) yangi Bernhauer, 1938 Zyras (Glossacantha) yangi Bernhauer, 1938: 147 1 2, Hong Kong, Shek Kong, at light, 19.111.1992, G. Ades leg. Distribuzione. Cina. Nuova per Hong Kong. Nota. Il bulbo distale della spermateca dell’esemplare di Hong Kong è netta- mente più lungo di quello della spermateca dei tipi di yangi. 666 ROBERTO PACE DESCRIZIONI DEINOPSINI Sharp, 1883 Adinopsis chinensis sp. n. Figg. 1-4 Holotypus ©, Hong Kong, X-XII.1996, G. de Rougemont leg. (MHNG). DESCRIZIONE. Lungh. 2,0 mm. Corpo debolmente opaco e giallo-bruno con capo bruno e margine anteriore et posteriore del pronoto gialli; antenne gialle, zampe rossicce. L’intera superficie del corpo è coperta di tubercoletti estremamente super- ficiali e di pubescenza sericea fittissima; spermateca fig. 2, urotergite X della femmina con due valvule fig. 3, apice dello stesso fig. 4. COMPARAZIONI. La nuova specie è affine ad A. rufobrunnea Cameron, 1919, di Singapore e ad A. nepalensis Pace, 1987, a motivo della presenza della papilla molto stretta all’apice dell’undicesimo antennomero. Se ne distingue per i caratteri dati nella seguente chiave: | Lunghezza 1,8 mm; rapporto larghezza/lunghezza del pronoto pari a 1,45 o1.60Xfemmina sconosciuta ....2.2.2.5/. + OR eee 2 - Lunghezza 2,0 mm; rapporto larghezza/lunghezza del pronoto pari a 1-3 04maschio:sconosciutosHons Kong... 2%)... rear ar chinensis sp. n. 2 Rapporto larghezza/lunghezza del pronoto pari a 1,60; apice dell’ edeago appuntito e armatura genitale interna dell’edeago semplice. Singapore EEE Eye RR TREE SA 03s ILS rufobrunnea Cameron - Rapporto larghezza/lunghezza del pronoto pari a 1,45; apice dell’ edeago non appuntito e armatura genitale interna dell’edeago ben sviluppata. Nepalis. ete eee III lA AR RR TTI nepalensis Pace PRONOMAEINI Ganglbauer, 1895 (=Myllaenini) Myllaena hongkongiphila sp. n. Figg. 5-6 Holotypus ©, Hong Kong, Kadoorie Agricultural Research Centre, flight interception trap, X-XII.1996, G. de Rougemont leg. (MHNG). DESCRIZIONE. Lungh. 2,2 mm. Corpo debolmente lucido e giallo sporco (imma- turo); antenne gialle con i tre antennomeri basali giallo sporco; zampe gialle. L’intero corpo è coperto di pubescenza sericea fittissima; spermateca fig. 6. COMPARAZIONI. La nuova specie presenta parte prossimale della spermateca non avvolta a spirale. Per questo carattere la specie tassonomicamente e geograficamente piu vicina è M. himalayca Cameron, 1939, dell’ India settentrionale. Ma la spermateca di quest’ultima specie ha bulbo distale assai stretto, con introflessione apicale minus- cola, e non larghissima come quella della nuova specie. Inoltre la nuova specie mostra occhi più lunghi delle tempie, mentre himalayca li ha più corti delle tempie. ALEOCHARINAE DI HONG KONG 667 FIGG. 1-4 Adinopsis chinensis sp. n. Habitus: 1; spermateca: 2; urotergo della femmina con due valve: 3; apice dello stesso: 4. Myllaena reelsi sp. n. Figg. 7-8 Holotypus 9, Hong Kong, Ma On Shan, at light, 17.X.1996, G.T. Reels leg. (MHNG). DESCRIZIONE. Lungh. 3,7 mm. Corpo debolmente lucido e giallo-bruno; antenne brune con antennomero basale giallo sporco e undicesimo giallo paglierino; zampe giallo sporco. L’intera superficie del corpo è coperta di pubescenza sericea fittissima; spermateca fig. 8. COMPARAZIONI. La nuova specie presenta la parte prossimale della spermateca che descrive due spire. Per questo carattere va comparata con M. chinoculata Pace, 1998, della Cina. Ma questa specie presenta tali spire relativamente più ampie, taglia corporea nettamente inferiore (1,9 mm), occhi molto più corti delle tempie ed elitre più corte del pronoto. ETIMOLOGIA. La nuova specie è dedicata al suo raccoglitore Graham Reels, entomologo inglese. 668 ROBERTO PACE GYROPHAENINI Kraatz, 1856 Gyrophaena (Gyrophaena) anguli sp. n. Figg. 9-12 Holotypus 6, Hong Kong, sifted litter, 1.11.1997, G. de Rougemont leg. (MHNG). Paratypus: 1 2, Hong Kong, Kadoorie Agricultural Research Centre, X-XII.1996, G. de Rougemont leg. DESCRIZIONE. Lungh. 1,7 mm. Corpo lucido e bruno con addome giallo avente gli uroterghi liberi IV e V nero-bruni; antenne e zampe gialle. La punteggiatura del capo e del pronoto è netta e distribuita come da fig. 11. Le elitre sono coperte di tuber- coletti superficiali. Solo il quinto urotergo libero presenta reticolazione, ma super- ficiale: sul resto del corpo la reticolazione è assente. Edeago figg. 9-10, sesto urotergo libero del maschio fig. 12. COMPARAZIONI. La nuova specie presenta due caratteri che permettono di avvici- narla tassonomicamente a G. gratella Cameron, 1939, dell’ India: il margine posteriore del sesto urotergo libero del maschio con quattro denti, di cui i due interni più esili degli esterni e apice dell’edeago flesso al lato ventrale e senza appendice preapicale. Tuttavia 1 denti laterali del sesto urotergo libero del maschio di gratella sono stretti alla base e lunghi il doppio degli interni, mentre gli interni della nuova specie sono lunghi meno del doppio degli esterni. Inoltre l’apice dell’edeago della nuova specie, in visione laterale, è stretto, mentre è assai largo in gratella. il pronoto di gratella ha doppia punteggiatura, cioè fine e sparsa tra i quattro punti discali in quadrato, mentre il pronoto della nuova specie ha solo punti isolati robusti. Pseudobrachida hongkongensis sp. n. Figg. 13-14 Holotypus ©, Hong Kong, Kadoorie Agricultural Research Centre, 31.V.1996, G. de Rougemont leg. (MHNG). DESCRIZIONE. Lungh. 1,6 mm. Corpo lucidissimo e bruno con margine posteriore degli uroterghi rossiccio; antenne giallo sporco con antennomeri 8 a 10 giallo-bruni e undicesimo bruno; zampe gialle. Solo le elitre sono coperte di reticolazione distinta, il resto del corpo è senza reticolazione. La punteggiatura del capo e del pronoto è evidente e distribuita come da fig. 13. Gli uroterghi sono coperti di tubercoli allungati; sper- mateca fig. 14. COMPARAZIONI. Dato che presenta il penultimo antennomero fortemente tras- verso, la nuova specie sembra simile a P. siwalikensis Cameron, 1939, dell’ India. Ma questa specie presenta addome bruno-rossiccio, con fascia nericcia davanti al pigidio, il quinto urotergo libero senza tubercoli e taglia corporea di 2,3 mm. PLACUSINI Mulsant & Rey, 1871 Placusa kadooriorum sp. n. Figg. 15-19 Holotypus 6, Hong Kong, Kadoorie Agricultural Research Centre, flight interception trap, X-X11.1996, G. de Rougemont (MHNG). Paratypus: 1 2, Hong Kong N.T.., sifted litter, 1.11.1997, G. de Rougemont leg. ALEOCHARINAE DI HONG KONG 669 FIGG. 5-12 Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. 5-6: Myllaena hongkongiphila sp. n.; 7-8: Myllaena reelsi sp. n.; 9-12: Gyrophaena (Gyro- phaena) anguli sp. n. 670 ROBERTO PACE DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e nero-bruno con meta posteriore delle elitre di un giallo sporco e con il margine posteriore degli uroterghi rossiccio; antenne nere con i quattro antennomeri basali giallo paglierino; zampe giallo-rossicce con femori bruni. La reticolazione del capo è distinta, quella del pronoto è estrema- mente superficiale e quella delle elitre e dell’addome è svanita. La punteggiatura del capo è distinta. Il pronoto, le elitre e l'addome sono coperti di tubercoletti superficiali. Il pronoto ha quattro punti isolati profondi; edeago figg. 16-17, spermateca fig. 18, sesto urotergo libero del maschio fig. 19. COMPARAZIONI. In base alla forma dell’edeago, la nuova specie mostra d’essere tassonomicamente affine a P. pygmaea Kraatz, 1859, dello Sri Lanka, ma i denti del margine posteriore del sesto urotergo libero del maschio, sono assai differenti, cioè in pygmaea vi è un largo dente mediano ad apice arrotondato tra due spine laterali ricurve. Anche l’edeago delle due specie presenta differenze sensibili. Il dente preapicale ventrale dell’apcie della lama sternale dell’edeago di pygmaea, è molto vicino all’apice, e piuttosto distante è quello della nuova specie. ETIMOLOGIA. La nuova specie è dedicata ai fratelli Kadoorie, noti benefattori di Hong Kong. Essi hanno concesso a G. de Rougemont di effettuare ricerche sulle Aleocharinae nel loro Centro di Ricerca sull’ Agricoltura. HOMALOTINI Heer, 1839 Linoglossa (s. str.) hongkongensis sp. n. Figg. 20-22 Holotypus ©, Hong Kong, Lantau Tai O, in Auricularia fungus. 15.11.1997, G. de Rouge- mont leg. (MHNG). DESCRIZIONE. Lungh. 3,0 mm. Corpo lucidissimo e rossiccio con 1 tre uriti basali giallo-rossicci; antenne bruno-rossicce con i due antennomeri basali e l'undicesimo giallo-rossicci; zampe gialle. Sul corpo non vi è traccia di reticolazione. La punteggiatura del capo e del pronoto è fitta e molto superficiale. Sulle elitre sono sparsi alcuni punti netti su una superficie indistintamente punteggiata. Tubercoletti fini coprono la superficie degli uroterghi. La fronte è impressa trasversalmente. Il pronoto ha una profonda concavità medianea posteriore con superficie rugosa sulla linea mediana; spermateca fig. 20); sesto urotergo libero della femmina fig. 21. COMPARAZIONI. Recente è la descrizione di una nuova specie del genere Linoglossa Kraatz, 1859, della Cina: L. chinensis Pace, 1998, ma la nuova specie presenta caratteri più simili a quelli osservabili su L. angustata (Motoschulsky, 1858), dell’ India meridionale, pertanto geograficamente più lontana, tanto che non appartiene neanche al medisimo sottogenere (sottogenere Axinocolya per L. chinensis). La nuova specie, che presenta una fossetta del pronoto simile, è distinta da L. angustata per i caratteri dati nella seguente chiave: - Decimo antennomero appena trasverso; disco del capo profondamente solcato; pronoto nettamente trasverso; bulbo distale della spermateca tronco-conico. Lungh. 3,5 mm. India meridionale. . . . . angustata (Motschulsky) ALEOCHARINAE DI HONG KONG 671 01 mm E E 18 20 ON se E E 5 | E E o Fico. 13-21 Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del maschio (19) e della femmina (21). 13-14: Pseudobrachida hongkongensis sp. n.; 15-19: Placusa kadooriorum sp. n.; 20-21: Linoglossa hongkongensis sp. n. Decimo antennomero nettamente trasverso; disco del capo senza solco; pronoto quasi lungo quanto largo; bulbo distale della spermateca subel- litiico Sun hys:0imm*> Hone Kona hongkongensis sp. n. 672 ROBERTO PACE DEREMINI Seevers, 1965 Demerinda rougemonti sp. n. Figg. 23-24 Holotypus ©, Hong Kong N.T., sifted litter, 1.11.1997, G. de Rougemont leg. (MHNG). DESCRIZIONE. Lungh. 1,7 mm. Corpo lucido e giallo-rossiccio con capo bruno- rossiccio e pronoto rossiccio; antenne bruno-rossicce con i tre antennomeri basali rossicci; zampe gialle. Sul corpo non vi è reticolazione. La punteggiatura del capo e del pronoto è fitta e distinta. Tubercoletti salienti coprono la superficie delle elitre e dell’addome. Il solco laterale del pronoto è profondo e largo, quello mediano è poco profondo. Le elitre presentano un debole spigolo laterale, dall’omero fino presso l’angolo posteriore esterno; spermateca assai minuscola, fig. 24. COMPARAZIONI. In base alla forma della spermateca, la nuova specie si situa tassonomicamente lontano sia da D. termophila Cameron, 1924, dell’India, che da D. hongkongensis Pace, 1998a, di Hong Kong. Infatti queste due specie hanno sper- mateca ben sviluppata in lunghezza, memtre la nuova specie ha spermateca assai corta. Inoltre l’addome della nuova specie ha pubescenza rada, mentre è fitta quella dell’addome delle due specie citate a paragone. ATHETINI Casey, 1910 Hydrosmecta subalgarum sp. n. Figg. 25-29 Holotypus 4, Hong Kong, Tai Long, under seaweed on sand, 30.III.1997, G. de Rouge- mont leg. (MHNG). Paratypi: 3 ©, stessa provenienza. DESCRIZIONE. Lungh. 2,2 mm. Avancorpo debolmente opaco, addome lucido. Corpo bruno con pronoto ed elitre giallo-bruni; antenne brune con i due antennomeri basali giallo-rossicci; zampe gialle. La reticolazione del capo e del pronoto è netta, quella delle elitre è quasi vigorosa e quella dell’addome è distinta. La punteggiatura del capo e delle elitre è estremamente superficiale, quella del pronoto è indistinta; edeago figg. 25-26, spermateca fig. 27, sesto urotergo libero del maschio fig. 29. COMPARAZIONI. La nuova specie ha habitus simile a quello di H. cooteri Pace, 1998a, della Cina, ma gli antennomeri 4 a 10 sono nettamente trasversi e non più lunghi che larghi come in cooteri. La spermateca della nuova specie ha bulbo prossimale non così nettamente dilatato come quello di cooteri. Atheta (Acrotona) iperanomala sp. n. Figg. 30-32 Holotypus ©, Hong Kong, Cheung Sheung, yellow pan, 27.1V.1997, G.T. Reels leg. (MHNG). DESCRIZIONE. Lungh. 2,1 mm. Corpo lucido e giallo-bruno (immaturo); antenne brune con i due antennomeri basali e la base del terzo di un giallo sporco; zampe gialle. La reticolazione della superficie del capo è estremamente superficiale, quella delle elitre è svanita e quella del pronoto e dell'addome è assente. La punteggiatura del capo è superficiale. I tubercoletti che coprono la superficie del pronoto sono fini e poco ALEOCHARINAE DI HONG KONG 673 mm FIGG. 22-28 Habitus, spermateca ed edeago in visione laterale e ventrale. 22: Linoglossa hongkongensis sp. n.; 23-24: Demerinda rougemonti sp. n.; 25-28: Hydrosmecta subalgarum sp. n. 674 ROBERTO PACE FicG. 29-32 Sesto urotergo libero del maschio, habitus e spermateca da due differenti punti di vista. 29: Hydrosmecta subalgarum sp. n.; 30-32: Atheta (Acrotona) iperanomala sp. n. ALEOCHARINAE DI HONG KONG 675 salienti, quelli delle elitre sono distinti e quelli dell’addome sono salienti; spermateca figg. 31-32. COMPARAZIONI. L’habitus della nuova specie è pressoché identico a quello di A. suspiciosa (Motschulsky, 1859) dell'India, Sri Lanka, Singapore, Cina meridionale e Filippine, cioè l'addome è appena ristretto all’indietro e la reticolazione del capo è estremamente superficiale. Ma la spermateca è nettamente differente nelle due specie. La parte prossimale della spermateca di suspiciosa descrive una sola spira, mentre nella nuova specie ne descrive due e il bulbo distale è senza introflessione apicale in suspi- ciosa, mentre è molto sviluppato in profondità nella nuova specie. Atheta (Acrotona) shekkongensis sp. n. Figg. 33-36 Holotypus d, Hong Kong, Shek Kong, at light, 24.VIII.1991, G. Ades leg. (MHNG). Paratypus: | 2, stessa provenienza. DESCRIZIONE. Lungh. 2,2 mm. Corpo lucido e bruno-rossiccio con quarto urite libero bruno; antenne brune con antennomero basale bruno-rossiccio; zampe gialle. La superficie del capo è coperta di reticolazione estremamente superficiale, il resto della superficie del corpo è senza reticolazione. La superficie del capo e del pronoto è coperta di tubercoletti fini e distinti; le elitre presentano tubercoletti superficiali; edeago figg. 34-35, spermateca fig. 36. COMPARAZIONI. Un numero di caratteri dell’edeago e della spermateca permette di avvicinare tassonomicamente la nuova specie ad A. vicaria (Kraatz, 1859), diffusa dallo Sri Lanka all'India e alle Filippine. Le differenze sono le seguenti: i penultimi antennomeri di vicaria sono nettamente trasversi, mentre nella nuova specie sono poco trasversi; gli occhi in vicaria sono lunghi quanto le tempie, mentre nella nuova specie gli occhi sono molto più lunghi delle tempie; l’addome di vicaria è appena ristretto all’indietro, mentre nella nuova specie è fortemente ristretto; il dente preapicale dorsale dell’edeago di vicaria è vicinissimo all’apice, mentre nella nuova specie è piuttosto lontano; i due pezzi falciformi interni dell’edeago di vicaria sono nettamente più lunghi di quelli corrispondenti nella nuova specie. Atheta (Acrotona) tricholutea sp. n. Figg. 37-38 Holotypus 9, Hong Kong N.T., Kadoorie Agricultural Research Centre, flight inter- ception trap, X-XII.1996, G. de Rougemont leg. (MHNG). DESCRIZIONE. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio con capo rossicci, con elitre brune orlate di giallo alla sutura e al margine posteriore e con quarto urite libero bruno; antenne brune con i due antennomeri basali e la base del terzo gialli; zampe gialle. Solo le elitre mostrano una reticolazione distinta e fine: sul resto della superficie del corpo non vi è traccia di reticolazione. La punteggiatura del capo e del pronoto è fitta e superficiale. I tubercoletti che coprono la superficie delle elitre sono fitti e superficiali, quelli degli uroterghi sono salienti; spermateca fig. 38. COMPARAZIONI. Il colore prevalentemente giallo-rossiccio del corpo e il tipo di spermateca, permettono di avvicinare tassonomicamente la nuova specie ad A. mi- mannuliventris Pace, 1998b, pure di Hong Kong. Il corpo della nuova specie però è più 676 ROBERTO PACE Fico. 33-38 Habitus, edeago in visione laterale e ventrale e spermateca. 33-36: Atheta (Acrotona) shekkon- gensis sp. n.; 37-38: Atheta (Acrotona) tricholutea sp. n. ALEOCHARINAE DI HONG KONG 677 robusto di quello di mimannuliventris, gli occhi sono piu lunghi delle tempie, mentre in mimannuliventris sono più corti delle tempie. La spermateca della nuova specie ha parte prossimale più prolungata e più sottile della parte corrispondente della spermateca di mimannuliventris e l’introflessione apicale del bulbo distale è nettamente meno pro- fonda nella nuova specie. Atheta (Coprothassa) roridanotha sp. n. Figg. 39-46 Holotypus d, Hong Kong, Kadoorie Agricultural Research Centre, flight interception trap, X-XII.1996, De Rougemont leg. (MHNG). Paratypus: | 2, Hong Kong, Ma On Shan, at light, 16.X.1996, G.T. Reels. DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e bruno con elitre giallo-brune; antenne brune con i due antennomeri basali giallo-bruni; zampe gialle. Sul capo e sul pronoto la reticolazione è assente. La reticolazione delle elitre è estremamente super- ficiale e quella dell’addome è distinta. L’avancorpo è coperto di tubercoletti salienti, fitti e regolari; edeago figg. 40-41, spermateca fig. 42. COMPARAZIONI. La nuova specie è chiaramente distinta sia per i caratteri dell’edeago, che della spermateca, da A. iucunda Pace, 1998b della Cina. L’edeago della nuova specie è appena ricurvo al lato ventrale, mentre è profondamente arcuato in iucunda e il bulbo distale della spermateca è meno dilatato nella nuova specie che in iucunda. Atheta (Microdota) pseudovagans sp. n. Figg. 47-50 Holotypus 4, Hong Kong, C.U.H.K., flight interception trap, HI-IV.1997, G. de Rouge- mont leg. (MHNG). DESCRIZIONE. Lungh. 1,8 mm. Avancorpo debolmente opaco, addome lucido. Corpo bruno con elitre giallo-brune e addome nero-bruno; antenne brune con i due antennomeri basali bruno-rossicci; zampe gialle. La reticolazione del capo e del pronoto è netta, quella delle elitre è indistinta e quella dell'addome è superficiale. L'intero corpo è coperto di tubercoletti distinti, tranne sul disco del capo dove mancano; edeago figg. 48-49, sesto urotergo libero del maschio fig. 50. COMPARAZIONI. La nuova specie ha habitus simile a quello di A. vagans Bernhauer, 1907, diffusa dallo Sri Lanka a Taiwan, Cina e Giappone. Solo gli occhi sono più sviluppati e gli antennomeri 8 a 10 meno fortemente trasversi. Il margine posteriore del sesto urotergo libero del maschio della nuova specie ha forma come da fig. 50, mentre in vagans è plurinciso a metà. Inoltre l'apice dell’edeago della nuova specie è stretto, mentre è molto largo in vagans. Atheta (Microdota) persimplex sp. n. Figg. 51-54 Holotypus 4, Hong Kong, C.U.H.K., flight interception trap, III-IV.1997, G. de Rouge- mont leg. (MHNG). Descrizione. Lungh. 1,4 mm. Corpo lucido e bruno (immaturo); antenne nero- brune; zampe di un giallo sporco. La reticolazione del capo è molto svanita, quella del 678 ROBERTO PACE 01 mm Oi mm FIGG. 39-46 Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 39- 42: Atheta (Coprothassa) roridanotha sp. n.; 43-46: Atheta (Microdota) ocularis Cameron, es. di Hong Kong. ALEOCHARINAE DI HONG KONG 679 01 mm Imm 01mm N / Ae UE ET Eur, Fico. 47-54 Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. 47-50: Atheta (Microdota) pseudovagans sp. n., 51-54: Atheta (Microdota) persimplex sp. n. 680 ROBERTO PACE pronoto è assente, quella delle elitre è superficiale e quella dell’addome è distinta. I tubercoletti che coprono la superficie del capo sono superficiali, quelli del pronoto sono salienti, quelli delle elitre distinti e quelli dell'addome sono fini; edeago figg. 52-53, sesto urotergo libero del maschio fig. 54. COMPARAZIONI. La nuova specie ha edeago simile a quello di A. pauxilla Cameron, 1944, dell’India, ma l’apice di quello della nuova specie è nettamente più stretto di quello di pauxilla. Gli occhi della nuova specie sono più lunghi delle tempie, mentre quelli di pauxilla sono molto più corti delle tempie e il quarto antennomero è lungo quanto largo nella nuova specie, mentre è nettamente trasverso in pauxilla. Atheta (Poromicrodota) hoihaensis sp. n. Figg. 55-56 Holotypus 9, Hong Kong, Hoi Ha, flight interception trap, 14.11.1996, G.T. Reels leg. (MHNG). DESCRIZIONE. Lungh. 2,4 mm. Corpo lucido e nero pece, con elitre e base dell’addome brune; antenne nere con i due antennomeri basali nero-bruni; zampe gialle con femori giallo-bruni. La reticolazione della superficie dell’avancorpo è molto superficiale, quella degli uroterghi è distinta e composta di maglie molto trasverse. I tubercoletti che coprono la superficie del capo e delle elitre sono fini e poco salienti, quelli del pronoto sono distinti; spermateca fig. 56. COMPARAZIONI. La nuova specie ha spermateca simile a quella di A. sororcula Cameron, 1939, dell’India e Nepal, ma la nuova specie ha occhi più lunghi delle tempie, mentre sororcula li ha molto più corti e l’introflessione apicale del bulbo distale della spermateca della nuova specie è larghissima, mentre è stretta in sororcula. Atheta (Sipalatheta) algarum sp. n. Figg. 59-62 Holotypus d, Hong Kong, Ting Kok, seaweed, 8.11.1997, G. de Rougemont leg. (MHNG). Paratypi: 2 d e 2 2, stessa provenienza; 1 9, Hong Kong, Sha Lang, seaweed on beach, 8.1IL.1997, G. de Rougemont leg. Descrizione. Lungh. 2,3 mm. Corpo lucido e bruno; antenne brune con 1 due antennomeri basali rossicci; zampe giallo-rossicce. La reticolazione del capo e del pro- noto è netta, quella delle elitre è distinta e quella dell’addome è superficiale. La pun- teggiatura del capo e del pronoto è estremamente superficiale. Tubercoletti molto svaniti coprono la superficie delle elitre; spermateca fig. 59, edeago figg. 60-61. COMPARAZIONI. La struttura dell’edeago della nuova specie, priva di apofisi laterali, richiama la forma dell’edeago di A. ciu Pace, 1993, della Cina. Tuttavia l’ede- ago della nuova specie presenta un dente preapicale dorsale, assente in civ e il bulbo basale dello stesso organo è più sviluppato in algarum sp. n. che in ciu. Inoltre il bulbo distale della spermateca della nuova specie è poco sviluppato, senza distinta intro- flessione apicale, mentre in ciu il bulbo distale è molto sviluppato, con profonda intro- flessione apicale. In più gli occhi della nuova specie sono più corti delle tempie, mentre in ciu sono lunghi quanto le tempie. 681 ALEOCHARINAE DI HONG KONG Imm 01 mm 01 mm 58 Fico. 55-61 Habitus, spermateca ed edeago in visione laterale e ventrale. 55-56: Atheta (Poromicrodota) hoihaensis sp. n.; 57-58: Gnypeta lucidula Pace di Hong Kong; 59-61: Atheta (Sipalatheta) algarum sp. n. 682 ROBERTO PACE Tomoglossa fuliginosa sp. n. Figg. 63-64 Holotypus ©, Hong Kong N. T., sifted litter, 1.11.1997, G. de Rougemont leg. (MHNG). Paratypus: | es. (privo di pigidio), stessa provenienza. DESCRIZIONE. Lungh. 2,3 mm. Corpo lucido e bruno con capo nero-bruno ed elitre giallo-brune; antenne brune con i due antennomeri basali bruno-rossicci; zampe gialle. Tutto il corpo è coperto di reticolazione distinta, sull’addome è a maglie molto trasverse. Tubercoletti salienti sono diffusi fittamente su tutta la superficie del corpo; spermateca fig. 63. COMPARAZIONI. La nuova specie è ben distinta da 7. subcorticalis (Cameron, 1939) dell’India, per il pronoto nettamente trasverso (pronoto appena trasverso in subcorticalis) e per l’introflessione apicale del bulbo distale assai largo nella nuova specie e strettissimo in subcorticalis. PYGOSTENINI Fauvel, 1899 Odontoxenus reelsi sp. n. Figg. 65-66 Holotypus 2, Hong Kong, Kau Tam Tso, yellow pan, 18.V.1996, S.J. Reels leg. (MHNG). DESCRIZIONE. Lungh. 2,7 mm. Corpo lucidissimo e giallo-rossiccio con capo bruno e con margine posteriore del pronoto, metà posteriore delle elitre e pigidio giallo- rossicci chiari; antenne bruno-rossicce con antennomero basale giallo-rossiccio; zampe rossicce. Il corpo è senza reticolazione. La punteggiatura dell’avancorpo è rada e superficiale. Solo gli uroterghi liberi IV, V e VI sono coperti di pubescenza; spermateca fig. 66. COMPARAZIONI. A Hong Kong è presente un’altra specie: O. rougemonti Pace, 1998c. La nuova specie si distingue da essa per i caratteri dati nella seguente chiave: - Antenne lunghe: volte all'indietro raggiungono il margine posteriore delle elitre; pronoto fortemente trasverso; capo con doppia punteggiatura; uroterghi liberi IV e V coperti di setoline allineate trasversalmente; femminaksconoseiufa. Euneh. 1-9-mm... non... E rougemonti Pace - Antenne corte: volte all’indietro raggiungono la base delle elitre; pronoto debolmente trasverso; capo con punteggiatura rada e superficiale; uro- terghi liberi IV et V diffusamente coperti di pubescenza; maschio sco- nosciutos/kungh. 27mm... ie le eee reelsi sp. n. TERMITOPAEDIINI Seevers, 1957 Dioxeuta rougemonti sp. n. Figg. 67-69 Holotypus d, Hong Kong, Pak Ngao Shek, leaf litter, 26.X.1996, J. Fellowes leg. (MHNG). Paratypi: 2 d, Hong Kong, Lin Au, flight interception trap, 26.X.1997, G.T. Reels leg. DESCRIZIONE. Lungh. 2,0 mm. Corpo lucido e giallo con elitre giallo-brune e addome oscurato di bruno agli uriti liberi III e IV; antenne e zampe gialle (immaturo). La reticolazione sulla superficie del capo e del pronoto è assente, quella delle elitre è molto svanita e quella dell’addome è superficiale. Non è presente una visibile 683 ALEOCHARINAE DI HONG KONG mes) Ar hier ters! fai | È 4 01 mm 01mm 67 FIGG. 62-68 Habitus, spermateca ed edeago in visione laterale e ventrale. 62: Atheta (Sipalatheta) algarum sp. n.; 63-64: Tomoglossa fuliginosa sp. n.; 65-66: Odontoxenus reelsi sp. n.; 67-68: Dioxeuta rouge- monti sp. n. 684 ROBERTO PACE punteggiatura su tutto il corpo. Rade setole sono inserite sul corpo come da fig. 69; edeago figg. 67-68. COMPARAZIONI. La nuova specie é affine a D. flavescens (Cameron, 1939) della Birmania. Se ne distingue per le antenne ancor più lunghe, cioè lunghe quanto quattro volte e mezzo la lunghezza del pronoto (e non solo quattro volte come in flavescens). Inoltre il pronoto della nuova specie è quasi lungo quanto largo, mentre in flavescens è nettamente trasverso (rapporto larghezza/lunghezza pari a 0,08 nella nuova specie e pari a 0,78 in flavescens). Di flavescens non è noto l’edeago. MYRMEDONIINI Thomson, 1867 Zyras (Sinozyras subgen. n.) pygmaeus sp. n. Figg. 70-73 Holotypus ©, Hong Kong, Lung Kwu Chao, at light, 18.X.1997, S.J. Reels leg. (MHNG). DESCRIZIONE. Lungh. 2,8 mm. Corpo lucidissimo e giallo-rossiccio con capo e uriti liberi IV, V e VI bruno-rossicci; antenne bruno-rossicce con i due antennomeri basali e la base del terzo gialli; zampe gialle. L'intera superficie corporea è senza reticolazione. La punteggiatura del capo è molto superficiale e assente sulla fascia longitudinale mediana, quella del pronoto è distinta e assente sulla fascia mediana. Le elitre sono coperte di tubercoletti fini e superficiali. L’addome presenta tubercoletti distinti. Il pronoto ha una profonda fossetta mediana posteriore; spermateca fig. 73. COMPARAZIONI. La taglia corporea ridotta, gli occhi molto più corti delle tempie e gli antennomeri IV a X notevolmente trasversi, sono caratteri distintivi che per- mettono di attribuire la nuova specie a un sottogenere nuovo, Sinozyras, tassonomi- camente vicino a Zyras (Zyras) da cui si distingue anche per i caratteri dati nella seguente chiave: - Ligula divisa in due larghı lembi, con setole apicali; mento poco tras- verso; mesocoxe largamente separate tra loro dato che il processo meta- sternale è larghissimo, tronco all’estremità e insinuato profondamente tra JESMESOCOXE EE N IO CEL Subgen. Zyras (Zyras) - Ligula divisa in due lembi stretti, senza setole apicali; mento estrema- mente trasverso; mesocoxe appena separate tra loro dato che il processo metasternale è appuntito e insinuato per breve tratto tra le mesocoxe RR PRETI URL Ao eI pe as Spe ats “ol Ses A Sr Subgen. Zyras (Sinozyras n.) Typus subgeneris: Zyras (Sinozyras) pygmaeus sp. n. Zyras (Zyras) hongkongensis sp. n. Figg. 74-77 Holotypus 6, Hong Kong, Tai Long, at light, 11.11.1997, G.T. Reels leg. (MHNG). Paratypi: 2 9, stessa provenienza. DescRIZIONE. Lungh. 5,0 mm. Corpo lucidissimo e nero con pronoto e base dell’addome giallo-rossicci; antenne giallo-rossicce con i tre antennomeri basali bruni e l’undicesimo giallo; femori gialli con estremità distale bruna, tibie anteriori giallo- brune, medie e posteriori gialle; tarsi gialli. La superficie del corpo è senza reticola- ALEOCHARINAE DI HONG KONG 685 FIGG. 69-72 Habitus, labio con palpo labiale e mento. 69: Dioxeuta rougemonti sp. n.; 70-72: Zyras (Sino- zyras subgen. n.) pygmaeus sp. n. ROBERTO PACE 686 FicG. 73-77 Spermateca, habitus ed edeagö in visione laterale e ventrale. 73: Zyras (Sinozyras subgen. n.) pygmaeus sp. n.; 74-77: Zyras (Zyras) hongkongensis sp. n. ALEOCHARINAE DI HONG KONG 687 79 FIGG. 78-79 Zyras (Glossacantha) yangi Bernhauer, es. di Hong Kong. Habitus della femmina: 78; sperma- tecan 79: zione. La punteggiatura del capo e del pronoto è finissima, assente sulla linea mediana del capo. Tubercoletti fini coprono la superficie delle elitre. La punteggiatura dell’ad- dome è netta e distribuita come da fig. 74; edeago figg. 75-76, spermateca fig. 77. COMPARAZIONI. Poiché la nuova specie presenta capo e pronoto non fortemente punteggiati, è comparabile solo con Z. chinkiangensis Bernhauer, 1939, della Cina, che ha il medesimo carattere. Se ne distingue per i caratteri dati nella seguente chiave: - Pronoto molto trasverso, con fossetta mediana posteriore profonda e circolare; decimo antennomero appena trasverso; edeago di un terzo più lungo e bruscamente arcuato al lato ventrale, con apice appuntito, in vi- sione ventrale. Lungh. 7,5 mm. Cina nord-occidentale. DR Rf ia tea fn ci Biba ape senile RE chinkiangensis Bernhauer = Pronoto appena trasverso, con impressione mediana posteriore trasversa; edeago di un terzo meno lungo, arcuato al lato ventrale in modo debole, con apice a larga ogiva, in visione ventrale. Lungh. 5,0 mm. Hong Kong. SATA AO TI NA ae ae ar RR ents tee hongkongensis sp. n. 688 ROBERTO PACE RINGRAZIAMENTI Ringrazio vivamente il collega Guillaume de Rougemont, noto studioso di Staphylinidae di Londra, per avermi affidato in studio il materiale oggetto del presente lavoro, frutto di sue ricerche e di G.T. Reels, G. Ades e J. Fellowes che pure senti- tamente ringrazio insieme ai conservatori del Natural History Museum di Londra e Field Museum of Natural History di Chicago per il prestito di tipi e di materiale di confronto. BIBLIOGRAFIA BERNHAUER, M. 1907. Zur Styphyliniden-Fauna von Japan. Verhandlungen der zoologisch- botanischen Gesellschaft in Wien 57: 371-414. BERNHAUER, M. 1938. Zur Staphyliniden-Fauna von China und Japan (XI. Beitrag). Entomo- logisches Nachrichtenblatt Troppau 12: 145-158. CAMERON, M. 1919. New species of Staphylinidae from Singapore. Part II. Transactions of the Entomological Society of London 1918: 231-246. CAMERON, M. 1927. Descriptions of two new genera of termitophilous Staphylinidae from India. Entomologist’s Monthly Magazine 63: 222-224. CAMERON, M. 1939. The Fauna of British India, including Ceylon and Burma. Coleoptera. Staphylinidae 4: 410 pp., London. CAMERON, M. 1944. Descriptions of new Staphylinidae (Coleoptera). Proceedings of the Royal Entomological Society of London 13: 49-52. Casey, T.L. 1910. New species of the Staphylinid tribe Myrmedoniini. In: Memoirs on the Coleoptera I: 183 pp., Lancaster. FAUVEL, A. 1899. Sur une tribu nouvelle de Staphylinides (Pygostenini). Jn: RAFFRAY, A. & FAUVEL, A.: Genres et especes de Staphylinides nouveaux d’Afrique. Revue d’Ento- mologie 18: 1-44. GANGLBAUER, L. 1895. Die Käfer von Mitteleuropa. Vol. 2. Familienrehie Staphylinoidea. I. Teil. Staphylinidae, Pselaphidae: 881 pp., Vienna. KRAATZ, G. 1856. Naturgeschichte der Insekten Deutschlands. Abteilung I, Coleoptera. Vol. 2, Staphylinii: 376 pp.. Berlin. KRAATZ, G. 1859. Die Staphyliniden-Fauna von Ostindien, insbesonders der Insel Ceylan. Archiv für Naturgeschichte 25: 1-196. MOTSCHULSKY, V. DE. 1858. Enumération des nouvelles espèces de Coléoptères rapportées de ses voyages. Bulletin de la Societe imperiale des Naturalistes de Moscou 3: 204-246. MOTSCHULSKY, V. DE. 1859. Insectes des Indes orientales et des contrées analogues. Etudes Entomologiques 8: 25-118. MULSANT, E. & Rey, C. 1871. Histoire Naturelle des Coléopteres de France. Brévipennes. Aléochariens: 321 pp., Paris. PACE, R. 1986. Aleocharinae dell’ Asia Sudorientale raccolte da G. de Rougemont. Bollettino del Museo civico di Storia naturale di Verona 13: 139-237. PACE, R. 1987. Staphylinidae dell’ Himalaya Nepalese. Aleocharinae raccolte dal Prof. Dr. J. Martens (Insecta: Coleoptera). Courier Forschungsinstitut Senckenberg 93: 386-441. PACE, R. 1988 (1997). Aleocharinae dello Sri Lanka raccolte da Marc Tronquet. Bollettino del Museo civico di Storia naturale di Verona 14: 315-338. Pace, R. 1992. Aleocharinae del Vietnam (Coleoptera, Staphylinidae). Nouvelle Revue d’Entomologie (N.S.) 9: 119-129. ALEOCHARINAE DI HONG KONG 689 Pace, R. 1993. Aleocharinae della Cina. Bollettino del Museo civico di Storia naturale di Verona 17: 127-180. Pace, R. 1998. Aleocharinae della Cina: Parte I (Coleoptera, Staphylinidae). Revue suisse de Zoologie 105: 139-220. PACE, R. 1998a. Aleocharinae della Cina: Parte II (Coleoptera, Staphylinidae). Revue suisse de Zoologie 105: 395-463. PACE, R. 1998b. Aleocharinae della Cina: Parte III (Coleoptera, Staphylinidae). Revue suisse de Zoologie 105: 665-732. Pace, R. 1998c. Aleocharinae della Cina: Parte IV (Coleoptera, Staphylinidae). Revue suisse de Zoologie 105: 000-000. SEEVERS, C.H. 1957. A Monograph on the Termitophilous Staphylinidae (Coleoptera). Fieldiana: Zoology 40: 344 pp. SEEVERS, C.H. 1965. The systematics, evolution and zoogeography of Staphylinid Beetles associated with Army Ants (Coleoptera, Staphylinidae). Fieldiana: Zoology 47: 137-351. SHARP, D. 1883. Fam. Staphylinidae. /n: Biologia Centrali-Americana. Insecta. Coleoptera, Vol. I: 145-312. T u an | | i MO Aare wt IRA x aS | Ve atest en - È RER Wy Stars Far #i rt ‘. x ti AL È “jh! | cri REVUE SUISSE DE ZOOLOGIE 106 (3): 691-744; septembre 1999 A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People’s Republic of China, I Ivan LOBL Muséum d'histoire naturelle, Case postale 6434, CH-1211 Geneva 6, Switzerland. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People’s Republic of China, I. - The present paper is the first part of a review of the Scaphidiinae of People’s Republic of China. It treates 78 species in 12 genera. New species are: Cyparium sichuanum sp. n., Ascaphium irregulare sp. n., A. alticola sp. n., Episcaphium catenatum sp. n., E. strenuum sp. n., E. haematoides sp. n., Scaphidium inexspectatum sp. n., S. jizuense sp.n., S. kubani sp. n., S. becvari sp. n., S. kurbatovi sp. n., S. lunare sp. n., S. schuel- kei sp. n., S. stigmatinotum sp. n., S. sichuanum sp. n., Baeocera xichangana sp. n., B. huashana sp. n., B. cooteri sp. n., B. kubani sp. n., B. yunnanensis sp. n., Scaphobaeocera simplex sp. n., S. molesta sp. n., S. pseudovalida sp.n., and Toxidium villosum sp. n. Scaphidium tsushimense Shirözu & Mori- moto, 1963 is placed in synonymy of Scaphidium amurense Solsky, 1871. Lectotypes are designated for Cyparium yunnanum (Achard); Ascaphium minus Pic, A. sinense Pic, Scaphidium delatouchei Achard, S. fukiense Pic, S. klapperichi Pic, S. vernicatum (Pic), S. yunnanum Fairmaire, Pseudobiro- nium sinicum Pic. Keys to species are provided. Key-words: Coleoptera - Staphylinidae - Scaphidiinae - taxonomy - China. INTRODUCTION The scaphidiines with almost 1400 species currently recognized is one of the more speciose groups of mycophagous beetles. While large collections from the Indian subcontinent, including Himalaya, Thailand and Japan were studied previously by me and the faunas of these areas appear reasonably well documented, very little is known about the scaphidiines of China. The gaps in the knowledge obscure reco- gnition of eventual centres of endemism and understanding distributional patterns of the taxa recorded from neighbouring countries. Thus, the collections of scaphidiines of the People’s Republic of China which became available since the last decade provide significant new information. Based on this material, a review of the scaphidiines of China is given. The present paper deals with all genera but Scaphisoma Leach which will be treated in a separate study. In respect to the size and habitat diversity of China, the scaphidiines appear under- represented compared to other parts of Asia (e.g. LOBL 1979, 1990b, 1992). Obviously, the available information is still anecdotal, and strongly depends on the Manuscript accepted 07.01.1999 692 IVAN LOBL accesibility of the sites and methods used by the collectors. In particular, the field work in high altitudes yield significant material from several mountains of Yunnan and Sichuan, while the material from low altitude sites is inadequate, with the possible exception of that from Hong Kong. Since 1886, when FAIRMAIRE described the first species from China, Scaphidium yunnanum, little information was published afterwords. ACHARD (1920a, 1920b, 1920c), Pic (1916, 1920, 1954), LôBL (1965, 1972, 1984b), and LI (1992) provided isolated descriptions of 18 species, two of which were synonymized. LOBL (1966, 1968, 1970, 1984b) and Li & CHEN (1993) recorded a few additional species, and Rougemont (1996) published a list of species of Hong Kong, based on identifications made by me. Presently, 31 identified and three additional unidentified species have been reported from the People’s Republic of China (referred below to as China). The classification of the higher taxa follows LESCHEN & LOBL (1995) and methods follow LößL (1992). A key to the genera is given in the latter paper but for distinguish characters of Episcaphium and Ascaphium see also LESCHEN & LOBL (1995). For references to previous descriptions and synonymy see the catalogue by opr (1997): Acronyms of the collections examined: MHNG Muséum d histoire naturelle, Genève MMUE The Manchester Museum, Manchester MNHN Museum National d’Histoire Naturelle, Paris NHMB Naturhistorisches Museum, Basel NHRM Naturhistoriska Riksmuseet, Stockholm NMPC Narodni Muzeum, Praha PCAP Private collection A. Pütz, Eisenhüttenstadt PCMS Private collection M. Schülke, Berlin SMNS — Staatliches Museum für Naturkunde, Stuttgart SYSTEMATICS Cyparium Erichson This genus comprises 47 valıd species occuring in all major biogeographical areas, except New Guinea and Australia. A single species, Cyparium yunnanum (Achard), was known previously from China. Five additional species are mentioned in this study, one of which is new. KEY TO THE SPECIES OF CYPARIUM OF CHINA | Elytra dark, with yellow humeral spot and yellow apical fascia So cast aa cae rR ae er RA cE RO I Pelee in C. montanum Achard = Elytratunicoloured’™*. 2! bott. MI, BUSS. So ee ee Z 2 Pronotum and elytra microsculptured and iridescent. Large species 4.5- OlOhmmylOn Se. i RR EE C. mikado Achard - Pronotum and elytra without microsculpture and not iridescent ............. 3 SCAPHIDINAE OF CHINA, I 693 (D) Prohypomera with microsculpture, lateral portion of metasternum coar- Selyzpünetater. ne, RME el EE RUE sce ee = - Prohypomera without microsculpture, lateral portions of metasternum SHTOO UI. REIN AURA SWANS FAN EEE ROLE OD ETE EN a SUAS N 5 - Mesepisternaswithimieroseulpturene Ser C. sibiricum Solsky - Mesepisterna without microsculpture................ C. yunnanum (Achard) 5 Elytra with distinct puncture rows. Metacoxal process with evenly Concavexapicalied ge tt Fin... en. ANSE ER C. siamense Löbl - Elytra without distinct puncture rows. Metacoxal process with tridentate ApICAled Ser meh WY SERRE DEA ISO BIBI I AA C. sichuanum sp. n. Cyparium mikado Achard Material examined. China, Beijing, Badaling, 13.XII.1993, G. de Rougemont, 1 (MHNG); Shaanxi, Qin Ling Shan, 110°06°E, 34°27’N, Hua Shan, north Valley, 1200-1400 m, 118 km E Xian, 18-20.VIIL.1995, M. Schülke, 1; same data but leg. D. W. Wrase, 1; same but leg. A. Piitz, 3 (PCAP, PCMS, MHNG). Distribution. Japan; China: Beijing and Shaanxi. Comments. This species is locally common in Japan and very active (personal observation). It may be easily distinguished from its Asian congeners by the pronotum and elytra with microsculpture, in combination with the uniformly dark and large body. The shape of the parameres and the structures of the internal sac of the aedeagus (Figs 1 to 3) are also diagnostic. Cyparium montanum Achard Material examined. China, Yunnan, Jizu Shan, 25°50’N 100°21’E, 2500-2700m, 6- 10.VII.1994, V. Kuban, 9 (NHMB, MHNG). Distribution. North India; Bhutan; China: Yunnan. Comments. This species may be easily distinguished by the elytral colour pattern. Cyparium siamense Löbl a Material examined. China, South Yunnan, Mengyang Nat. Res. ca 500m, 14.IX.1994, S. A. Kurbatov, litter, 3 (MHNG). Distribution. China: Yunnan. Comments. The specimens of China are slightly smaller than those from Thailand. They differ also in having the elytral margin keels entirely exposed, and the aedeagi 0.90-0.92 mm long. Cyparium sibiricum Solsky Figs 1-3 Material examined. China, Yunnan, Heishui, 35km north Lijiang, 27°13’N, 100°19°E, 18.VI.-4.VII.1993, S. Beévar, 10 (MHNG); North Yunnan, Lijiang, 2600m, 30.VI.-2.VII.1990, L. & M. Bocäk, 1 (NHMB); Sichuan, Ganzi Pref., Daxue Shan, 101°57’E 30°03’N, north Kanding, 2600-270m, 22-24.V.1997, A. Piitz, 1 (PCAP); South Sichuan, 20km S Muli (Bowa), 101°13’E, 27°45’N, 3500m, 29.VI.1998, M. Bocäk, 1 (BMNS); Shaanxi, Qin Lin Shan, 100°06°E 34°27’N, Hua Shan, north valley, 1200-1400m, 118km E Xian, 18.-20.VIIL.1995, M. Schülke, 1 (MHNG). 694 IVAN LOBL Distribution. Russia: Siberia, Transbaikal, Far East Russia; China: Shaanxi, Sichuan, Yunnan. Comments. The specimens of China possess finer and shorter elytral puncture rows than the specimens from Russia. Cyparium tenenbaumi Pic from Siberia is very similar to, and likely just an infrasubspecific form of C. sibiricum. More material is needed to examine the validity of C. tenenbaumi. Cyparium yunnanum (Achard) Type material examined. Lectotype d and paralectotype 2, labelled Ht. Yunnan, Tali / Type (red) / Cyparium yunnanum n. sp. J. Achard det Type (handwritten by J. Achard) / Mus. Nat. Pragae Inv. 18714 (lectotype) and Inv. 18713 (paralectotype) (NMPC) - by present designation. Additional material. China, Yunnan, | 2 (MHNG). Distribution. China: Yunnan. Comments. This species possesses following diagnostic characters: Length 1.8- 1.9 mm. Body black, not iridescent. Apex of abdomen, femora, tibiae and antennal club brown. Antennal segments | - 6 and tarsi ochraceous. Prohypomera dull, with conspi- cuous microsculpture. Mesepisterna smooth. Metasternum without microsculpture. Most of middle portion of metasternum smooth, fine punctures near intercoxal process. Lateral portions of metasternum distinctly punctate, puncture diameters smaller than puncture intervals. Abdominal segments with microsculpture consisting of punctures. First exposed sternite punctate as lateral portions of metasternum, and with laterobasal row of elongate pits. Aedeagus very similar to that in C. sibiricum. Cyparium sichuanum sp.n. Figs 4-6 Holotype d: China, Central Sichuan, Wolong Nat. Res., 100m, 24.V.1994, in bamboo litter, S. A. Kurbatov (MHNG). Description. Length 3.7 mm, width 2.3 mm. Body black, frons and abdominal apex dark brown, legs reddish-brown, palpi and antennal segments | to 6 yellowish, segments 7 to 11 light brown. Pronotum and elytra without microsculpture, not iri- descent. Head with frons finely punctate, at narrowest point 0.17 mm. Antennal segments 3 and 5 evenly long, segment 4 distinctly shorter, segment 6 1.4 times as long as wide, segment 7 about as long as wide, segments 8 to 10 gradually wider, each wider than long. Pronotum with lateral edges barely sinuate near base, lateral keels not exposed in dorsal view, punctation irregular, partly very fine, diameters of largest punctures usually much smaller than puncture intervals. Exposed portion of scutellum wider than long. Elytra with adsutural areas impressed and flat; lateral margin keels concealed at level of maximal width and near apex (dorsal view); discal punctation very irregular, four irregular, fairly coarse puncture rows hardly distinguishable; punctation Fics 1-7 Aedeagi in Cyparium; 1 to 3. C. sibiricum Solsky, internal sac extruded, parameres (2) and internal sac (3) in detail; 4 to 6. C. sichuanum sp. n., paramere (5) and internal sac (6) in detail; 7. C. siamense Löbl, internal sac. Scale bars = 0.1 mm in figs 2, 3, 5, 6, 7; = 0.2 mm in figs 1 and 4. 695 SCAPHIDIINAE OF CHINA, I ues Ò Gi > = SSA FE SS 696 IVAN LOBL extremely fine between puncture rows, along lateral edges, and at base; punctation coarse and dense on apical third of elytra and on area posterior humeral protuberance. Coarse elytral punctures larger than largest pronotal punctures. Prohypomera smooth. Mesepisterna and metasternum extremely finely punctate. Median portion of metaster- num slightly raised in middle, with two wide and shallow admesal impressions. Metacoxal process raised in middle, with apical edge tridentate, more prominent in middle than at angles. Exposed abdominal segments with mesh-like microsculpture, in addition to brickwall-like microsculpture of intersegmental membranes. Propygidium and pygidium almost evenly finely punctate. First exposed sternite with fairly coarse and sparse punctation, following sternites very finely punctate. Tibiae slighty curved. Male sexual characters. Segments | to 3 of protarsi and mesotarsi almost evenly wide, distinctly wider than segments 4 and 5. Aedeagus (Figs 4 to 6) 1.22 mm long. Apical portion of median lobe perpendicular to basal bulb. Parameres sinuate, widest subapically (lateral view), with apical lobe. Internal sac with membranes forming scale- like and denticulate structures, robust sclerites absent. Comments. This species would key to the couplet «5» in the key of the Asian species of Cyparium (LOBL 1990a), together with C. laevisternale Nakane. It may be distinguished from C. laevisternale by the elytral punctation which is mostly coarse, and by the shape of the metacoxal process which is concave apically and not raised medially in C. laevisternale. The internal sac of the aedeagus resembles that of C. siamense (Fig. 7), although it is more complex. The parameres are not lobed in C. siamense. Ascaphium Lewis This genus comprises seven described species which are keyed in LOBL (1992). Three of them occur in Japan, two are described from China (one of them described originaly as a variety), one from Vietnam, and one from Nepal. Three unidentified species from Taiwan and Burma are represented in collection of the MHNG. Two additional, new species from China are described below. KEY TO THE SPECIES OF ASCAPHIUM OF CHINA I Large species 6.5-7 mm long. Antennal segment 7 about 2 times as long as wide. Elytra with puncture rows almost reaching posterior tenth of TR TO ni a Te du A. sinense Pic - Smaller species 5-6 mm long. Antennal segment 7 about 1.6 times as long as wide. Elytra with puncture rows ending far anterior apical tenth ON CN ATEN ESSE RO ciro did OE EE 2 Antennae uniformly brown, or segments 7 to 10 slightly darkened. Elytra with four almost equal, long, discal puncture rows and one short lateral puncture row starting about at mid-length of elytral disc......... A. minus Pic - Antennae with segments 7 to 10 much darker than segments | to 6 and II Elytra wath discal puncture rows. different "NES RC EC EEE 3 D SCAPHIDIINAE OF CHINA, I 697 3 Elytra with four discal puncture rows not or hardly impressed, without additionalishontilateralinower ares eis esa Re Ort A. alticola sp. n. - Elytra with four discal puncture rows distinctly impressed and two lateral, very short additional puncture row............... A. irregulare sp. n. Ascaphium minus Pic Type material examined. Lectotype d labelled Kuatun Fukien China 18.11.46 (Tchung Sen.) / war. minor mihi (handritten by Pic) (MHNG); paralectotypes 14 and 19 labelled as the lectotype but from 25.4. and 10.5.46, respectively (MHNG) - by present designation. Additional material. Same data but from 14.9.46 and without the original identification labels, 4 (MHNG). Distribution. China: Fujian. Comments. Specimens without Pic’s original identification labels are likely syntypes but in doubt they are not considered as such. Additional specimens from Kuatun are in MNHN. The previous spelling “minor” combined with a neuter genus name is incorrect. Ascaphium sinense Pic Type material examined. Lectotype ¢ labelled Kuatun Fukien China 6.12.46 (Tschung Sen.) / 9739 E91/ Type Ascaphium sinense n. sp. (red handwritten) (NHRS); paralectotypes 3d, 22 with same label data as the lectotype but from 6.3., 2.4., 22.10., 2.11. and 18.11.46, respectively. Each bearing Pic’s handwritten label Ascaphium sinense (MHNG) - by present designation. Additional material. From the same locality but from various dates in 1946, 29 (MHNG). Distribution. China: Fujian. Comments. As under A. minus. Ascaphium irregulare sp.n. Figs 8-10 Holotype d : China, Yunnan, 25°58’N, 100°21’E Jizu Shan, 2500-2700m, 6-10. VII.1994, V. Kuban (MHNG). Paratypes: same data as holotype, 1 d,4 9 (MHNG). Description. Length 5.0-5.5 mm. Head, body, femora and tibiae uniformly black. Palpi, antennal segments | to 6 and 11, tarsi and apical abdominal segments light brown to ochraceous. Antennal segments 7 to 10 blackish. Frons at narrowest point 0.43-0.46 mm, with punctation irregular, fine and sparse anteriorly, gradually denser and coarser toward neck. Neck smooth, impunctate, not microsculptured dorsally. Antennae moderately long; segment 5 about 3 times as long as wide; segment 6 2.5 times as long as wide, club stout; segments 7, 8, 9 and 11 each 1.5 to 1.6 times as long as wide; segment 10 about 1.2 times as long as wide. Pronotum at base 2.0-2.2 mm wide; lateral edges oblique in basal half, arcuate in apical half; antebasal puncture row slightly impressed, contiguous in middle, consisting of coarse punctures; discal punc- tation sparse and very fine. Elytra uneven, somewhat flattened posterior basal fourth, impressed laterally and apically. Each elytron with four uneven discal puncture rows situated in relatively wide impressions, and with two additional lateral puncture rows consisting each of two to six coarse punctures. Two inner puncture rows straight, IVAN LOBL 698 SCAPHIDIINAE OF CHINA, I 699 starting distant from base and ending usually anterior mid-length of elytron; third puncture row sometimes interrupted, starting usually at base and ending posterior mid- length of elytron; fourth puncture row starting at base and curved, or starting posterior base, straight, ending near mid-length of elytron. Remaining elytral punctation very fine. Metasternum with two shallow, elongate medio-apical impressions. First exposed abdominal sternite with mediobasal protuberance; microsculpture absent, except near apical edge. Microsculpture obsolete on following three sternites, consisting of extre- mely fine striae on apical abdominal segments. Protibiae straight, mesotibiae and meta- tibiae slightly curved. Male sexual characters. Segments | to 3 of protarsi distinctly widen. Aedeagus (Figs 8 to 10) 1.25-1.35 mm long. Median lobe with apical portion curved, widen toward tip; basal process large, prominent. Parameres diverging apically, curved and slightly widen at tip. Internal sac with asymmetrical complex of subapical sclerites joined to two strongly sclerotized rods. Comments. This species is characterized by the pattern of the elytral puncture rows which are short, irregular, and impressed. Additional diagnostic characters are: antennal segments 7 to 10 strongly darkened and metasternum with two shallow medio- apical impressions. See also comments under A. alticola. Ascaphium alticola sp. n. Holotype 2: China West Sichuan, 29°35’N 102°00°E 2900-3200m, Gonggashan- Hailuogou, 3-6. VII.1994, J. Farkaë & D. Kral (MHNG). Paratype 2: same data as holotype (MHNG). Description. Length 6.3 mm. Colour as in A. irregulare. Frons at narrowest point 0.48 mm, with punctation very irregular, consisting of more or less large punc- tures mostly smaller than intervals between them. Largest punctures situated near neck. Neck entirely microsculptured, with few fine punctures in middle of dorsal surface. Antennae moderately long, with segment 5 about 3 times as long as wide; segment 6 twice as long as wide; club stout; segment 7 1.6 times as long as wide; segment 8 as wide as and slightly shorter than 7, about 1.4 times as long as wide; segments 9 and 10 evenly large, each 1.6 times as long as wide, slightly wider than segments 7 and 8; segment 11 1.6 times as long as wide. Pronotum at base 2.2 mm wide; lateral edges oblique in basal half, arcuate in apical half; antebasal puncture row slightly impressed laterally, contiguous in middle, consisting of fairly coarse punctures; discal punctation very fine. Elytra slightly flattened or impressed between basal fourth and mid-length, with uneven, inflexed sides. Each elytron with four discal puncture rows: first and second rows short, straight, not or slightly impressed, starting just posterior basal fifth and ending about at level of mid-length of elytron; third puncture row distinctly Fics 8-13 Aedeagi in Ascaphium and Episcaphium; 8 to 10. A. irregulare sp. n., internal sac (10) in detail; 11'to 13. E. catenatum sp. n., internal sac (13) in detail. Scale bars = 0.1 mm in figs 10 and 13, = 0.2 mm in figs 8, 9, 11 and 12. 700 IVAN LOBL impressed, slightly oblique, approximate to base, ending just posterior middle third of elytral length; fourth puncture row deeply impressed, curved, starting posterior humeral protuberance, ending slightly anterior apical third of elytron. Punctation very fine between and posterior puncture rows. Metasternum with two median striae shallow, approximate, diverging posteriorly. First exposed abdominal sternite with mediobasal protuberance; microsculpture consisting partly of extremely fine punctures, partly forming more distinct mesh-like pattern and transverse striae. Following sternites with microsculpture consisting of meshes and/or transverse striae. Protibiae straight, meso- tibiae and metatibiae slightly sinuate. Comments. This species is very similar to A. irregulare from which it may be distinguished by the pattern of the elytral puncture rows and by the metasternal striae diverging posteriorly. Episcaphium Lewis This genus comprises four species, from Japan, Nepal, East Malaysia, and Sri Lanka. The collections coming from China include three new species which may be easily distinguished by their colour pattern. KEY TO THE SPECIES OF EPISCAPHIUM I Flead:pronotumiandielytraumfonmniyiblack .. ........ ROSS SS NSA 2 - Colour pattern different... - . Anni ca bacon ACCO TONE 4 2 Abdomen uniformly black . . . 729422 ee ae ah is Se ee 3 - Abdomen ochraceous, elytra without puncture rows........ E. strenuum Sp. n. 3 Elytra with fine puncture rows, without large basal protuberance. Meta- SEEMUMMMOHMIELOSEUlpturede were cia ede eee E. catenatum sp. n. - Elytra without puncture rows, with large basal protuberance. Metaster- num with microsculpture consisting of transverse striae. a es. sec engaged A RAIL Pelo centra Ur E. callosipenne (Achard) 4 Elytra uniformly reddishior ochraceous 21... 42.1: 2.222 Sr ee 5 - Elytra,with one:or two: dark transverse fasciae.. "PP 20.02 os eee 6 5 Head, orhead and pronotum, black "eme PC er Leer E. semirufum Lewis - Heads pronotumvandielytraneddish Sf beh: 222.222 gro E. uniforme Löbl 6 Elytra with dark central and apical fasciae; puncture rows absent. Pro- notumareddishie ya ya Amerie ire sai ia cone E. saucineum (Motschulsky) - Elytra with dark apical transverse spot and four puncture rows. Pronotum reddish, often with pair of dark basal spots ............. E. haematoides sp. n. Episcaphium catenatum sp. n. Figs 11-13 Holotype d: China, Central Sichuan, Wolong Nat. Res., 1500m, 21.V.1994, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 2 4,3 © (MHNG). Description. Length 5 mm. Body and legs black, antennal segments | to 6, tip of antennal segment 11, palpi and apex of abdomen dark brown. Frons at narrowest point SCAPHIDIINAE OF CHINA, I 701 0.42 mm, with punctation sparse and fine anteriorly, becoming more dense and coarse posteriorly, puncture diameters larger than puncture intervals near neck. Punctation fine and dense on tempora, fine and sparse on neck. Neck microsculptured, except on smooth centre. Antennae short, club segments stout. Antennal segment 5 distinctly longer than segment 6 (index 28:22), almost 2 times as long as wide; segment 6 about 1.4 times as long as wide; segments 7 and 8 each about 1.3 times as long as wide; segments 9 and 10 as long as wide or slightly longer than wide; segment 11 about 1.5 times as long as wide. Pronotum at base 1.9 mm wide, with lateral margin sinuate, concave near base, arcuate in apical half; antebasal puncture row irregular, sparse, not impressed laterally, interrupted at middle; discal punctation sparse and fine. Elytra without impressions, lacking discal grooves or striae, with four discal puncture rows (in addition to adsural, basal, and marginal puncture rows): first to third rows short, ending near mid-length of elytron, fourth row reaching apical fourth of elytron; first and second rows starting distant basal row, third and fourth rows joined to basal row. Punctures forming rows about as large as or slightly larger than punctures forming antebasal pronotal row; remaining punctures very fine. Humeral protuberance hardly distinct. Mesal groove of mesosternal process narrow or absent. Metasternum lacking microsculpture, extremely finely punctate, with median impression shallow, narrowed anteriorly, and with two small oval impressions near metacoxae. Abdominal tergit 7 with distinct lateral striae. Abdominal sternites microsculptured; microsculpture on first exposed sternite consisting of points and transverse striae, that on following sternites mesh-like and striate. First exposed sternite very finely and sparsely punctate. Tibiae slightly curved. Male sexual characters. Segments 1 to 3 of protarsi distinctly enlarged. Aedeagus (Figs 11-13) 1.35-1.43 mm long. Median lobe with curved, cylindrical apical portion, and robust, prominent ventral process. Compression plate small. Parameres straight in lateral view, slightly arcuate in dorsal view. Internal sac with robust, apically bifid sclerite joined to pair of vertical laminae and to subapical lobe formed by flat denticles. Apical portion of internal sac covered by very fine spine-like structures oriented proximally. Comments. This species may be distinguished from its congeners, E. callosi- penne excepted, by the uniformly black body. It differs notably from E. callosipenne by the absence of the metasternal microsculpture, shape of the elytra and size and shape of the segments of the antennal club. Episcaphium strenuum sp. n. Figs 14-17 Holotype d : China, Yunnan, 27°06’N 115°15’E, Yulonshan, 3000-3500m, Camaizi pass, 18-25. VII.1990, V. Kuban (MHNG). Paratypes: Sichuan, Gongga Shan, Hailuogou, 29°35’N 102°00° E, in front of Glacier 1, 2800m, 9.VII.1996, A. Smetana, 1 d (MHNG); Sichuan, Daxue Shan, Gongga Shan, Hailuogou Glacier Park, above Camp III, 30.V.1997, A. Pütz, 1 4 (PCAP); Yunnan, Heishui, 35 km N Lijiang, 27°13’N, 100°19’E, 18.VI.-4.VII.1993, S. Beëväï, 3 2 (MHNG); same but 1-19.VII. 1992, 1 2; Yunnan, Jizu Shan, 2500-2700m, 25°58’N 100°21’E, 6-10. VII.1994, 1 2 (MHNG). Description. Length 6.5-7.5 mm. Head, thorax, elytra and legs, tarsi excepted, black. Abdomen and tarsi reddish-brown or ochraceous. Antennae very dark brown to 702 IVAN LOBL SCAPHIDIINAE OF CHINA, I 703 blackish. Frons at narrowest point 0.50-0.60 mm wide, with punctation irregular, mostly coarse, fine anteriorly, dense along eye edges, at centre and near neck, sparse on admesal and anterior areas. Frontal punctures mostly much larger than intervals, as large as or smaller than intervals on surface sparsely punctate. Tempora finely punctate. Vertex sparsely and mostly finely punctate, with distinct microsculpture consisting of striae, smooth on small medio-anterior area. Antennae long, club with stout segments. Antennal segment 5 almost 4 times as long as wide; segment 6 distinctly shorter and hardly wider than segment 5, about 2.5 to 3 times as long as wide; segments 7 and 8 evenly wide, 8 slightly shorter than 7, 9 and 10; segment 7 about 1.8-1.9 times as long as wide; segments 8 and 9 each about 1.5-1.6 times as long as wide; segments 9 to 11 becoming gradually, slightly wider, 9 about 1.5-1.6 times as long as wide, 10 1.4 times as long as wide, 11 about 1.6-1.7 times as long as wide. Pronotum at base 2.2-2.4 mm wide, with lateral margin hardly sinuate, antebasal puncture row almost even, not or hardly interrupted at middle, slightly impressed laterally; punctation on disc rather dense and fine. Elytra with small, shallow, mediolateral impression and larger apical impression, lacking discal grooves, striae, without trace of puncture rows; punctation very fine, finer than that on pronotum. Humeral protuberance low. Mesosternal process without mesal groove. Metasternum extremely finely punctate, lacking microsculpture, with medio-apical impression shallow, narrowed anteriorly, and two small impressions near metacoxae; apical edge of metacoxal process slightly concave. Abdominal tergite 7 with distinct lateral striae. Abdominal sternites very finely punctate. First exposed sternite with punctation very fine and sparse, becoming denser mediobasally, and with microsculpture consisting of punctures. Following sternites with distinct microsculpture consisting of striae. Tibiae slightly curved. Male sexual characters. Segments 1 to 3 of protarsi widen. Aedeagus (Figs 14- 17) 2.05-2.10 mm long. Median lobe very weakly widen toward mid-length, almost evenly wide, sinuate and curved in lateral view. Compression plate large. Parameres slender, slightly curved in dorsal view, straight in lateral view. Internal sac with complex sclerites forming rods and plates, membranes bearing spine-like structures visible only while extruded. Comments. This species may be easily distinguished from its congeners by the colour pattern. Episcaphium haematoides sp. n. Figs 18-20 Holotype d: China, Yunnan, Heishui 35km west Lijiang, 27°13’N 100°19’E, 1- 19.V11.1992, S. Beévat (MHNG). Fics 14-20 Aedeagi in Episcaphium; 14 to 17. E. strenuum sp. n., internal sac (16 and 17) in detail, and extruded (17); 18 to 20. E. haematoides sp. n., internal sac (20) in detail. Scale bars = 0.1 mm in figs 16, 17 and 20, = 0.2 mm in figs 14, 15, 18 and 19. 704 IVAN LOBL Paratypes: as holotype, 4 4,4 ©, 64 (MHNG, NHMB); same but 18.VI.-4.VII.1993, 127 (MHNG); Yunnan, Yulong Mts, 3900m, 27°10’N 100°13’E, 16-19.VI.1993, Bolm, 1 6 (NHMB); Yunnan, Jizu Shan, 2500-2700m, 25°58’N 100°21E, 6-7.VII.1994, V. Kuban, 1 6, 1 © (MHNG); Habashan, southeastern slope, 27°20°N 100°11’E, 3-6.VI.1995, S. Beévar, 20 (MHNG), NHMB); same but 2000-3000m, 10-13.VII.1992, V. Kuban, 1 6 (MHNG); Yunnan, Habashan, eastern slope, 3800-4600m, 27°19’N 100°08’E, 15.V11.1992, D. Kral, 1 (MHNG); Sichuan, Sabde, ca 3000m, 1-3. VII.1992, R. Dunda, 1 (MHNG). Description. Length 4.8-5.5 mm. Head black. Pronotum ochraceous or reddish, usually with two small, well delimited black spots situated between antebasal puncture row and basal edge, near lateral edges. Elytra as pronotum ochraceous or reddish, but each with an apical transverse black spot not touching lateral and sutural edges. Abdomen ochraceous or reddish with apical segments slightly lighter. Most of ventral surface ochraceous or reddish. Prosternum, prohypomera along posterior edge, meso- sternum, mesepisterna, coxae, femora and tibiae black. Tarsi dark brown to blackish. Antennae with segments | to 6 very dark, almost black, club black. Frons at narrowest point 0.45-0.48 mm wide, with punctation irregular, mostly sparse and fine, denser and more coarse on centre and near neck. Punctures mostly smaller than intervals between them, some coarse punctures larger than intervals. Tempora impunctate. Vertex finely and sparsely punctate, lacking microsculpture. Antennae moderately long, with club stout. Antennal segment 5 2.3-2.4 times as long as wide; segment 6 shorter and stouter than segment 5, about 1.3 times as long as wide; segments 7 to 10 becoming gradually, slightly wider; segments 7, 9 and 10 each as long as segment 5; segment 7 slightly longer than wide; segment 8 slightly shorter than segment 7 and slightly wider than long; segments 9 and 10 each slightly wider than long; segment 11 about 1.3-1.4 times as long as wide. Pronotum as base 1.9-2.3 mm wide, with lateral margin somewhat rounded, antebasal puncture row interrupted at middle, slightly impressed laterally, discal punctures fairly sparse and very fine. Elytra lacking lateral and apical im- pressions, with four discal puncture rows consisting of evenly fine punctures, or punc- tures becoming finer porteriorly. Puncture rows starting posterior basal fifth or fourth of elytron and ending far anterior apical edge. Inner puncture row parallel to adsutural stria, outer puncture rows slightly oblique. Mesosternal process with raised, ridge-like edges, flattened in middle. Metasternum extremely finely punctate, lacking micro- sculpture, median impressions extremely shallow or absent. Abdominal tergit 7 with distinct lateral striae. Abdominal sternites very finely and sparsely punctate. Lateral portion of first exposed sternite with microsculpture consisting of punctures, median portion without microsculpture. Following three sternites with microsculpture consis- ting of transverse striae. Tibiae curved. Male sexual characters. Segments | to 3 of protarsi distinctly widen. Aedeagus (Figs 18-20) 1.45-1.55 mm long. Median lobe narrowed apically, inflexed and slightly sinuate in lateral view. Basal process small, slightly prominent. Compression plate large. Parameres slightly arcuate in dorsal view, almost straight posterior curved base in lateral view. Internal sac with two median pairs of sclerotized rods, inner rods much smaller than outer rods. Comments. This species may be easily distinguished by the colour pattern. SCAPHIDIINAE OF CHINA, I 705 4 ba 4 3 3 2 Atimessasllonerasiwiderenn. sn 2 ee ae B. breveapicalis (Pic) - Antennal segment 6 elongate and narrow, slightly shorter than segment 7 and 3-5 times as longas widé .. i.e: I ee 7 7 Parameres of aedeagus expanded dorso-apically to form very large, stronelyiselerotizedllobei 17. tate Sor soor oe bee e B. kubani sp. n. - Parameres of aedeagus without lobe or with small, slightly sclerotized, subapical lobes +10 2) eile ns LEONE ee 8 SCAPHIDIINAE OF CHINA, | 723 Apical portion of parameres strongly widen .............. B. huashana sp. n. Apicalsportion of parameresinotor. hardly widens...) sey. eee 9 Antennal segment 11 about twice as long as wide. Lateral contours of pronotum and elytra continuously arcuate in dorsal view. Elytral punc- tauonmostlyiobsoleter ee eae B. xilingana sp. n. Antennal segment 11 about 3 times as long as wide. Lateral contours of pronotum and elytra separately arcuate in dorsal view. Elytral punctation wsually shiner DUE DISC ter twee ck. tye ech RO ce SER HERE RAT 10 Internal sac of aedeagus without strongly sclerotized guide-sclerite EN D get Mee I A II B. xichangana sp. n. Internal sac of aedeagus with strongly sclerotized guide-sclerite........... Il Guide-sclerite of internal sac narrow, angulate basally ........ B. callida Löbl Guide-sclerite of internal sac robust, not angulate basally................ 12 Guide-sclerite of internal sac gradually tapering apically ......... B. freyi Löbl Guidesselerite\ofinternalisac noftapening apically; ). inno Mir 18 Guide-sclerite of internal sac straight, stouter apically........ B. cooteri sp. n. Guide-sclerite of internal sac abruptly curved in middle, widen subapi- Cally@gpomied'atipo Dee ee es le ccs Sy a Re Ne B. hammondi Löbl Metatarsi short, as long as or slightly longer than half of metatibiae. Elytra with sutural stria not extendend along basal edge. Body length Se IRE el DES DEE een B. mussardi Löbl Metatarsi long, as long as or longer than 4/5 of metatibiae. Elytra with sutural stria extended along basal edge. Body length 1.2 - 1.5 mm......... 15 Elytra each with basal stria not joined to lateral stria. Parameres of aedeasusmolcheda per Se ee B. franzi (Löbl) Elytra each with basal stria joined to lateral stria. Parameres of aedeagus MOCMOLCN EG ie sche is Se Me aia a Nese 16 Elytralgpunctation/obsoletes eo «oe eae ae B. sordidoides Löbl Elytral,punetation.distinetpe.. te tt yes a Sane PER Re 17 Punctation distinct only on basal half of elytral disc ...... B. pseudinculta Löbl Runctation distinct on mostofelytralldise m un nn an 18 Metepisternum distinctly separated from metasternum by deep suture. Antennal segment 3 short, about as long as 2/3 of segment 4 BERNER Où Po de Ces LAS A ea rie I tA ae eer B. serendibensis (Löbl) Metepisternum indistinctly separated from metasternum by outer punc- ture row. Antennal segments 3 and 4 almost equally long ............... 19 Aedeagus with parameres straight, conspicuously narrow....... B. vidua Löbl Aedeagus with parameres slightly sinuate or arcuate, not particularly IAGO WARE NE ev N ence Beamline Nek Mn ty A eh I IT 20 Aedeagus with parameres diverging apically; internal sac without row of dENTICIE ST ER AA te Se Re I UNS En B. longicornis (Löbl) Aedeagus with parameres converging at apex; internal sac with one or two row of denticles situated posterior sclerite complex ....... B. pigra (Löbl) 724 IVAN LOBL Baeocera serendibensis (Löbl) Material examined. China, Hong Kong, V. 1996, G. de Rougemont, 2 (MHNG). Distribution. India; Nepal; Sri Lanka; Thailand; China: Hong Kong. Baeocera sordidoides Löbl Material examined. China, Sichuan, Xiling Mts, 1600-2400m, litter, 30. VII.-4. VIII.1996, S. A. Kurbatov, 1 (MHNG). Distribution. Nepal; China: Sichuan. Comments. This specimen is tentatively assigned to B. sordidoides. It differs conspicuously from the type series by the metasternal punctation which is highly reduced, fine and sparse. Baeocera mussardi roberti Lob! Material examined. China, Hong Kong, III, V, and IX. 1996, G. de Rougemont, 19 (MHNG); Zhejiang, Lin’an County, ca 200m, W Tienmu Shan Nat. Res., 22.V.1996, J. Cooter, 1 (MHNG); Yunnan, Gaoligong Mts, 1500-2500m, 25°22’N 98°49’E, 1.-24.V.1995, O. Semela, 1 (NHMB); Yunnan, Shi Lin, 8.X.1985, G. de Rougemont, 1 (MHNG). Distribution. Bhutan; India; Nepal; China: Hong Kong, Zhejiang, Yunnan. Baeocera longicornis (Löbl) Material examined. China, Hong Kong, Ill. 1996, G. de Rougemont, 2 (MHNG); South Yunnan, Mengyang Nat. Res., ca 500m, 11. and 14.IX.1994, S. A. Kurbatov, 2 (MHNG). Distribution. India; Nepal; Sri Lanka; Thailand; China: Hong Kong, Yunnan. Baeocera pigra (Löbl) Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 9. and 11.IX. 1994, S. A. Kurbatov, 2 (MHNG). Distribution. India; Nepal; Sri Lanka; Thailand; China: Yunnan. Baeocera vidua Löbl Material examined. China, East Hubei, near Macheng, ca 500m, 25.V.1995, S. Kurbatov, Il (MHNG); northeast Guangxi, 10km S Longsheng, ca 1000m, 15.VI.1995, S. A. Kurbatov, 6 (MHNG); Sichuan, Mt. Emei, 1500m, 21.IX.1994, S. A. Kurbatov, 3 (MHNG); Yunnan, Gaoligong Mts, 1500-2500m, 25°22° N 98°49’E, 17.-24.V.1995, O. Semela, 2 (NHMB). Distribution. Thailand; China: Hubai, Guangxi, Sichuan, Yunnan. Baeocera pseudinculta Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 10.1X.1994, S. A. Kurbatov, 1 (MHNG). Distribution. Thailand; China: Yunnan. Comments. The parameres of the aedeagi are narrower in the Chinese specimens than those in the specimens from Thailand. SCAPHIDIINAE OF CHINA, I 725 Baeocera franzi (Löbl) Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 9-14.[X.1994, S. A. Kurbatov, 55 (MHNG); Yunnan, Yipinglang, 1800-2000m, 17-20.VI.1994, V. Kuban, I (NHMB); Sichuan, Mt. Emei, 1700 and 1800m, 22. and 24.IX.1994, S. A. Kurbatov, 2 (MHNG); Sichuan, Qingshen Shan, 65km NW Chengdu, 8km W Taiping, 103°33’E 30°53’N, 18.V.- 4.V1.1997, 800-1000m, A. Pütz, 3 (PCAP); Sichuan, east Xian, Mt. Huashan, 9-11.V.1994, S. A. Kurbatov, 2 (MHNG); South Sichuan, near Xichang, 1600m, 28.VII.1996, S. A. Kurbatov (MHNG); East Hubei, near Macheng, ca 500m, 25.V.1995, S. A. Kurbatov, 5 (MHNG); Hubei, Wuhan, park, 200m, 28.V.1995, S. A. Kurbatov, 4 (MHNG); Jiangsu, Nanjing, Zijinshan, 8.V.1996, J. Cooter, 3 (MHNG); Shaanxi, Qin Lin Shan, 110°06’E, 34°27’N, Hua Shan, north valley, 1200-1400m, 118km east Xian 18-20. VIII.1995, M. Schülke, 1 (PCMS). Distribution. Thailand; China: Yunnan, Sichuan, Hubei, Jiangsu, Shaanxi. Baeocera callida Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 14.IX.1994, S. A. Kurbatov, 1 (MHNG); Yunnan, Xishan, 2300m, 24°57’N, 102°38’E, 27.V11.1983, Bolm, 3 and V. Kuban, | (NHMB). Distribution. Pakistan; North India; Nepal; China: Yunnan. Baeocera freyi Löbl Material examined. China, Shaanxi, Qin Lin Shan, 108°49’E, 33°55’N, River Valley 40 km south Xian, autoroute km 50, River bank, 1200m, 31.VIII.1995, M. Schiilke, 1 and A. Pütz, 6; Jiangsu, Nanjing Zijinshan, 8.V.1996, J. Cooter, 1 (MHNG, PCAP). Distribution. Far East Russia; North Korea; China: Shaanxi. Comments. This species may be easily distinguished from its congeners by the shape of the sclerites of the internal sac (Figs 58, 59). Baeocera hammondi Löbl Material examined. China, Shaanxi, East Xian, Mt. Huashan, 1700m, 9-11.V.1994, litter, S. A. Kurbatov, 1 (MHNG). Distribution. China: Shaanxi. Baeocera gilloghyi (Löbl) Material examined. China, South Yunnan, Mengyang Nat. Res. ca 500m, 9 and 11.IX. 1994, S. A. Kurbatov, 5 (MHNG). Distribution. Northeast India; Vietnam; China: Yunnan. Comments. This species is unique in having the apical portion of the left para- mrere strongly widen to form a large dorsal lobe. Baeocera pubiventris Löbl Material elxamined. China, South Yunnan, Mengyang Nat. Res. ca 500m, 9 and 11.1X.1994, S. A. Kurbatov, 4 (MHNG). Distribution. North India; Nepal; Thailand; China: Yunnan. Comments. This species may be easily distinguished from its congeners by the shape of the parameres, as figured in LOBL (1990b). 726 IVAN LÖBL Baeocera breveapicalis Pic Material examined. China, Northeast Guangxi, 10 km S Longsheng, ca 1000m, 15.VI. 1995, S. A. Kurbatov, 2 (MHNG); S. Yunnan, Mengyang Nat. Park, ca 500m, 12.IX.1994, S. A. Kurbatov, 3 (MHNG). Distribution. Vietnam; China: Guangxi, Yunnan. Comments. This species may be distinguished from the members of the B. mons- trosa group by the aedeagal characters only (Fig. 63). Baeocera satana Nakane Material examined. China, Northeast Guangxi, 10km south Longsheng, ca 1000m, litter, 15.V1.1995, S. A. Kurbatov, 1 (MHNG). Distribution. Japan; China: Guangxi. Comments. This species is member of a small group of species (B. dentipes Löbl, B. errabunda Löbl, B. monstrosetibialis Löbl, B. thoracica Löbl and B. tuber- culata Löbl) which have modified metatibiae in males. All but B. satana are known from India and Nepal. Baeocera satana may be easily distinguished from its Chinese congeners by the shape of the aedeagus (see LOBL 1994a) and male metatibiae. Baeocera xichangana sp. n. Figs Sl; 52 Holotype d: China, South Sichuan, near Xichang, 1600m, 28.VII.1996, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 2 d,2 2 (MHNG). Description. Length 2.0 - 2.15 mm. Body reddish-brown to blackish. Apex of abdomen, antennae and tarsi lighter. Relative length of antennal segments 3 to 11 as: 15: 18: 20: 19: 20: 15: 19: 17: 23 (holotype); segments 3 to 6 evenly slender; segments 7 and 8 evenly wide, slightly wider than segment 6; segment 7 about 5 times as long as wide; segment 8 about 4 times as long as wide; club segments much wider than segment 8; segment 11 about 3 times as long as wide and twice as wide as segment 8. Lateral contours of pronotum an elytra separately arcuate in dorsal view. Lateral pronotal keel sinuate posteriorly (lateral view). Pronotum and elytra lacking micro- sculpture, very finely punctate. Punctation on basal portion of elytra as that on pronotal disc, punctation on posterior half of elytra slightly more distinct. Scutellum completely covered by pronotal lobe. Elytra with entire basal stria joined to lateral stria, and middle portion of lateral edge straight in dorsal view. Prohypomera and mesepisterna im- punctate. Lateral portions of metasternum extremely finely punctate. Mesocoxal lines parallel to coxa, bordered by fairly small, not elongate pits. Metepisternum flat, wide, slightly narrowed anteriorly or parallel-sided, with deep, straight, impunctate suture. Exposed abdominal sternites very finely punctate, without microsculpture. Protibiae straight, mesotibiae and metatibiae barely curved. Fics 51-55 Aedeagi in Baeocera; 51 and 52. B. xichangana sp. n.; 53. B. huachana sp. n.; 54 and 55. B. cooteri sp. n. Scale bars = 0.2 mm. VAY SCAPHIDIINAE OF CHINA, I 728 IVAN LOBL Male sexual characters. Segments 1 and 2 of protarsi strongly widen, slightly narrower than protibia; segment 3 of protarsi distinctly widen. Aedeagus (Figs 51, 52) 0.97-0.98 mm long. Median lobe slightly asymmetrical, with basal buld moderately sclerotized, about as long as apical portion. Apical portion gradually narrowed in dorsal view, inflexed, arcuate, and almost evenly wide (tip excepted) in lateral view. Orifice of dorsal valve long, situated along right edge. Parameres conspicuously long, symme- trical, with inner and outer edge sinuate, almost evenly wide between expanded basis and apex. Internal sac with U-shaped basal sclerite, extended by long flagellum sclero- tized along edges. Long apical portion of flagellum weakly sclerotized, extruded. Fla- gellar guide-sclerite absent. Comments. Baeocera xichangana 1s unique by its aedeagal characters and cannot be placed in any known species group (LOBL 1992, LOBL & STEPHAN 1993). It may be easily distinguished from its congeners by the dorsal valve of the median lobe open along the right edge, the long parameres and the internal sac with very long flagellum sclerotized along both edges. Baeocera huashana sp. n. Fig. 53 Holotype d: China, East Xian, Mt. Huashan, 1700m, 9-11.V.1994, S. A. Kurbatov (MHNG). Description. Length 2.10 mm. Body black, apices of elytra and abdomen dark brown. Legs brown, tarsi lighter than tibiae and femora. Antennae light brown. Relative length of antennal segments 3 to 11 as: 16: 18: 20: 18: 20: 17: 19: 17: 23. Segments 3 to 6 almost evenly wide; segments 7 and 8 evenly wide, slightly wider than 6, segment 7 about 5 times as long as wide, segment 8 about 4 times as long as wide; clubs segments much wider than segments 7 or 8, segment 11 about 2.5 times as long as wide and 2 times as wide as segment 7. Lateral contours of pronotum and elytra separately arcuate in dorsal view. Pronotum and elytra without microsculpture. Pronotal punctation very fine. Scutellum completely covered by pronotal lobe. Elytra with basal stria entire, joined to lateral stria; lateral edge straight in middle portion (dorsal view). Elytral punctation fine, punctures notably larger than those on pronotal disc. Prohypomera, mesepisterna and lateral portions of metasternum extremely finely punctate. Mesocoxal lines convexly arcuate, bordered by coarse, slightly elongate pits. Metepisternum wide, flat, slightly widen posteriorly, with almost straight, impunctate suture. Sternite I with coarse, partly elongate basal pits, extremely finely punctate, not microsculptured. Following sternites with microsculpture consisting of punctures. Protibiae and mesotibiae straight, metatibiae slightly curved. Male sexual characters. Segments | of protarsi and mesotarsi strongly widen, narrower than tibiae. Segment 2 of protarsi strongly widen, narrower than segment 1. Segment 3 of protarsi and segment 2 of mesotarsi slightly widen. Aedeagus (Figs 53) 0.95 mm long. Basal bulb fairly strongly sclerotized. Apical portion of median lobe slightly asymmetrical, shorter than basal bulb, gradually narrowed in dorsal view, with tip blunt, strongly inflexed ventrally. Ventral process large, prominent, with tubercle in middle. Parameres arcuate, with long basal process, gradually narrowed toward apical SCAPHIDIINAE OF CHINA, I 729 portion, apex widen. Internal sac with wide, bisinuate, strongly sclerotized flagellum, extended by narrow, weakly sclerotized, extruded portion. Flagellar guide-sclerite long, slender, widen at base and at apex. Comments. This species is a member of the B. curtula group (LOBL 1992). It may be distinguished from most members of that group (B. kapfereri Reitter, B. freyi Löbl, B. hammondi Löbl, B. hamifer Löbl, B. mustangensis Löbl, and B. cooteri and B. kubani described below) by the shape of the parameres. It resembles B. curtula Achard by the shape of the apically widen parameres, althought they are narrower in the latter species. It differs conspicuously from B. curtula by the elytral punctation, which is in B. curtula as fine as the pronotal punctation, by the larger aedeagus with bisinuate flagellum of the internal sac, and by the tuberculate basal process of the median lobe. Baeocera cooteri sp. n. Figs 54, 55 Holotype d: China, Zhejiang, Lin’an county, ca 500m, W Tianmu Shan N. R., 16.V.1996, J. Cooter (MHNG). Paratypes: same data as holotype, 3 d, 2 9; same data but 300m, 22.V., 1 d ; same data but 350, 16-22.V., 1 9; Zhejiang, Lin’an County, Lin Long Shan, 250m, 22.V.1996, J. Cooter, 5 3,1 ©; Hong Kong, III.1996, R. de Rougemont, 1 d (all MHNG). Description. Length 1.80-2.15 mm. Body dark brown to black, apical abdominal segments and appendages lighter. Most external characters as in B. huashana but pits bordering mesocoxal lines and base of sternite I distinctly elongate. Relative length of antennal segments 3 to 11 as: 15: 20: 20: 19: 20: 16: 19: 17: 25 (holotype). Elytra with distinct discal punctation. Male sexual characters. Segments | to 3 of protarsi strongly widen, segment 1 almost as wide as protibiae, following segments gradually narrowed. Mesotarsi with segment | strongly widen, segments 2 and 3 slightly widen. Aedeagus (Figs 54, 55) 0.80-0.90 mm long. Basal bulb fairly sclerotized. Apical portion of median lobe shorter than basal bulb, slightly asymmetrical and with apex blunt in dorsal view, inflexed, almost evenly concave and narrowed toward tip in lateral view. Ventral process mode- rately projecting. Parameres with long basal process, sinuate and almost evenly wide in dorsal view, moderately curved in lateral view, with small, dorso-apical lobe. Internal sac with flagellum joined to incurved rods; flagellar guide-sclerite almost straight, narrow, except strongly expanded apical portion in dorsal view, robust and hook-like in lateral view. Comments. This species is characterised by the shape of the flagellar-guide sclerite and parameres. It may be readily distinguished from other members of the B. curtula group known from China (B. callida, B. hammondi, B. freyi, B. huashana, B. kubani) by the distinct elytral puncation. Baeocera kubani sp. n. Figs 56, 57 Holotype d: China, Yunnan, Jizu Mts, 2800m, 26°58’N 100°21’E, 30.V.-3.V1.1993, S. Kuban (MHNG). Description. Length 2.47 mm. Body black, apical abdominal segments and tibiae dark brown, tarsi and antennae ochraceous. Antennae apparently long (segments IVAN LOBL 730 SCAPHIDIINAE OF CHINA, I 73] 8 to 11 of both antennae brocken off and lost). Relative length of segments 3 to 7 as: 21: 19: 20: 20: 20. Segments 3 to 6 almost equally wide, segment 7 wider than 6, about 5 times as long as wide. Lateral contours of pronotum and elytra separately arcuate in dorsal view. Pronotum and elytra without microsculpture. Pronotal punctation dense, extremely fine. Scutellum completely covered by pronotal lobe. Lateral edges of elytra slightly rounded in dorsal view, basal stria entire. Elytral punctation dense, consisting of punctures extremely shallow, mostly about as small as pronotal punctures, some punctures distinctly larger. Prohypomera, mesepisterna and lateral portion of meta- sternum extremely finely punctate. Mesocoxal lines parallel to coxa, bordered by coarse, not elongate marginal pits. Metepisterna large, almost flat, with suture deep, impunctate, almost straight. Exposed abdominal tergites very finely punctate. Sternites I to V without microsculpture, very finely punctate; basal pits of sternite I not elongate. Protibiae and mesotibae hardly curved, metatibiae straight. Male sexual characters. Segments | to 3 of protarsi strongly widen, narrower than protibiae; segment 2 slightly narrower than segment 1, segment 3 distinctly narrower than segment 2. Segment | of mesotarsi strongly widen, segments 2 und 3 moderately widen. Median portion of exposed abdominal sternites bearing very sparse pubescence. Aedeagus (Figs 55, 56) 1.27 mm long. Median lobe with moderately sclerotized basal bulb. Apical portion of median lobe gradually narrowed in dorsal view, strongly inflexed, slightly asymmetrical, shorter than basal bulb. Ventral process small, not prominent. Parameres with short basal processes, almost evenly narrow between widen basis and mid-length, strongly expanded dorsally between mid-length and apex, forming conspicuous, large, relatively strongly sclerotized lobe. Internal sac with complex sclerotized basal structures. Flagellar guide-sclerite robust, partly extruded, flattened vertically, raised subapically, with apical hook. Comments. This species may be easily distinguished from its congeners by the shape of the parameres expanded by a dorsal lobe conspicuously large, and by the shape of the flagellar guide-sclerite. See comments under B. cooteri. Baeocera yunnanensis sp. n. Figs 60-62 Holotype dé: China, South Yunnan, Mengyang Nat. Res., ca 500m, 9.IX.1994 S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 2 d, and same data but from 11.IX., 1 d (MHNG). Description. Length 1.7 - 1.8 mm. Body, femora and tibiae uniformly ochra- ceous, or prothorax and abdomen lighter than elytra and metasternum. Antennae and tarsi lighter than prothorax and abdomen. Antennae long, relative length of segments 3 to 11 as: 12: 16: 18: 15: 20: 18: 22: 20: 30 (holotype). Segments 3 to 6 equally narrow, segment 5 about 4 times as long as wide; segments 7 and 8 equally wide, slightly wider than segment 6, segment 7 about 4 times as long as wide; segments 9 and 10 each FiGs 56-59 Aedeagi in Baeocera; 56 and 57. B. kubani sp. n.; 58 and 59. B. freyi Löbl. Scale bars = 0.2 mm. IVAN LOBL 62 63 SCAPHIDIINAE OF CHINA, I 733 slightly wider than 8; segment 11 as wide as or wider than 10, about 4 times as long as wide. Lateral contours of pronotum and elytra separately arcuate. Pronotum and elytra lacking microsculpture. Pronotum with extremely fine punctation. Exposed portion of scutellum large. Lateral edge of elytra almost straight in middle third. Elytra with basal stria interrupted at humeral area. Humeral area slightly raised. Elytral punctation fairly fine, consisting of well delimited punctures, intervals mostly 2 to 5 times as large as puncture diameters; basal portion of elytra extremely finely punctate. Prohypomera and mesepisterna impunctate. Lateral portions of metasternum extremely finely punctate. Mesocoxal lines parallel, with marginal pits extended laterally, not or slightly elongate. Metepisternum large, flat, narrowed anteriorly, with impressed, slightly rounded suture. Abdominal tergites with microsculpture consisting of punctures; punctation fairly fine, about as that of apical portion of elytra, but more dense. Abdominal sternites with extremely fine punctation, lacking microsculpture; basal pits of sternite I not or hardly elongate. Tibiae slightly curved. Male sexual characters. Protarsi and mesotarsi with segments | strongly widen, about as wide as tibiae; segments 2 slightly widen, segments 3 hardly widen. Aedeagus (Figs 60-62) 0.83-0.87 mm long. Basal bulb large, weakly sclerotized. Apical portion of median lobe not inflexed, short, asymmetrical, with tip blunt in dorsal view, pointed in lateral view. Ventral process small, not prominent. Parameres strongly asymmetrical, relatively narrow. Left paramere moderately narrowed apically, with large mediodorsal lobe, and abruptly inflexed apex. Right paramere abruptly narrowed in posterior third, and with ventral lobe. Internal sac very complex. Comments. This new species is a member of the B. monstrosa group. Is it very similar to B. pubiventris and B. gylloghyi with which it was found. It may be easily distinguished from these two species by the right paramere abruptly narrowed and lobed ventrally, and the left paramere lobed dorsally. Bironium Csiki The genus comprises 27 species, all known from subtropical or tropical ares of Asia and in New Guinea. Members of Bironium were not yet reported from China. One species is present within the collections examined. Bironium bidens Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 9-14.1X.1994, S. A. Kurbatov, 14 (MHNG). Distribution. Thailand; China: Yunnan. Mysthrix Champion Tis genus comprises two likely termitophilous species, one of which occurs also in China. The second species, M. kistneri Lobl, is known from Sumatra. Fics 60-63 Aedeagi in Baeocera; 60 to 62. B. yunnanensis sp. n., internal sac (62) in detail; 63. B. breveapicalis (Pic). Scale bars = 0.2 mm in figs 60, 61; = 0.1 mm in figs 62, 63. 734 IVAN LOBL Mysthrix termitophilum Champion Material examined. China, Hong Kong, III, V, and X-XII.1996, G. de Rougemont, 31 (MHNG). Distribution. North India; Nepal; China: Hong Kong. Comments. The specimens from China are slightly larger than the few ones which I have seen from the Himalayan region. Scaphicoma Motschulsky This genus comprises eleven species currently recognized as valid, distributed in South East Asia, New Ireland, and tropical Africa. Although some species of Scaphi- coma may be readily recognized (e.g., S. nigrovittata (Achard), S. flavovittata Motschulsky) by the colour pattern and punctation, reliable diagnostic characters are unknown from most of them. For this reason I could not identify one of the two species found in China. Scaphicoma arcuata (Champion, 1927) Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 11.[X.1994, S. A. Kurbatov, | (MHNG). Distribution. North India; Nepal; Thailand; China: Yunnan. Comments. This species differs from the similar S. apicalis (Pic) by the abdominal sternite I with dense basolateral punctation. Scaphicoma sp. Material examined. China, Northeast Guangxi, 10km S Longsheng, ca 1000m, 15.VI. 1995, S. A. Kurbatov, 2 2 (MHNG). Comments. These specimens represent possibly a separate species. They differ from S. apicalis and S. arcuata, and resemble S. cincta (Pic) by their colour pattern. The body is ochraceous but the bases of the pronotum and elytra, the elytral apices and lateral edges, large portions of the prohypomera, mesepisterna, metasternum, and the entire metepisterna are strongly darkened. Scaphobaeocera Csiki This genus is widely distributed and found in Asia, New Guinea, Australia, Micronesia, Mascarene archipelago and tropical Africa. It comprises 60 described species. Seven are known to occur in Japan (LOBL 1981), one from Far Eastern Russia (LOBL 1993), and one unidentified species was recorded from China (ROUGEMONT 1996). The new material raises the number of Chinese species to eleven, three of which are new. SCAPHIDIINAE OF CHINA, I 735 KEY TO THE SCAPHOBAEOCERA OF CHINA I Antennal segment 11 shorter than, or as long as, segment 10. . S. spinigera Löbl - Antennalkseoment i Jonserithanssesment- [Or ee: . 2.2 er ESSE 2 2 Antennal segment 11 about as long as segments 9 and 10 combined. Bodyalenethl1=1:3:mm N rar. In san. HR Re. ee S. dorsalis Löbl - Antennal segment 11 shorter than segments 9 and 10 combined ............ 3 3 Prohypomera with longitudinal stria. Antennal segment 8 about 4 times aslone asswide. Body, lensth 6 mm... ann S. pseudovalida sp. n. - Prohypomera without stria. Antennal segment 8 about as long as wide, or upto 2 times as long as wide. Body length 1.1-1.5mm.................... 4 4 Aedeagus with parameres notched subapically................ S. incisa Löbl - Nedeasusswithlparameresmotnotched yrı.2....0. 02000 0 en 5 5 Aedeagus with median lobe forming ventral plate prominent posterior AU GUIA PFOCESSEE TO RN er O S. lamellifera Löbl - Aedeacus with median lobe lacking ventraliplate.) eee 6 Aedeagus with flagellum simple, evenly, slightly arcuate, narrowed gra- dualllygapicalllye fox e o S. simplex sp. n. - Medeacusawithyiiacellumuditteremt varie tee atk ee a aie en 7 7 Aedeacusiwith flagellum spiral'and simplen 27. Mn ane Ee 8 - Aedeagus with flagellum not spiral, or spiral and complex basally.......... 9 8 Aedeagus with flagellum forming two or three complete circles S. difficilis Löbl - Aedeagus with flagellum forming six or seven complete circles . . . S. spira Löbl 9 Aedeagus with basal bulb prominent apicoventrally ........... S. nobilis Löbl - Aedeagus with basal bulb not prominent apicoventrally ................. 10 10 Apical third of parameres conspicuously widen in lateral view; flagellum Spiral, with basevlanserand complex an na ee er S. molesta sp. n. - Apical two thirds of parameres moderate widen in lateral view; flagellum irregularly incurved, not spiral, with base fairly small, hook-like Ba AIRS FERIE UMN PIR CR UV 4 ep € pc allel pearl preg S. cognata Löbl Scaphobaeocera lamellifera Löbl Material examined. China, Yunnan, Gaoligong Mts, 25°22’N 98°49’E, 1500-2500m, 17- 24.V.1995, O. Semela, 1 (MHNG). Distribution. Northeast India; China: Yunnan. Comments. This species was known from two males taken in the Garo Hills, Meghalaya. The single specimen from Yunnan exhibits exactly the same conspicuous aedeagal characters as the types. Scaphobaeocera incisa Löbl Material examined. China, South Yunnan, Mengyang Nat. Res. ca 500m, 9-13.IX.1994, S. A. Kurbatov, 6 (MHNG). Distribution. Thailand; China: Yunnan. Comments. This is the only species of Scaphobaeocera which possesses para- meres notched subapically. 736 IVAN LOBL Scaphobaeocera cognata Lob! Material examined. China, Shaanxi, Qin Lin Shan, 107°56’E 33°45’E autoroute km 93 S of Zhouzhi, 108 km southwest Xian, 1650m, mountain forest, 1-2.IX.1995, A. Pütz, 1 (PCAP); Central Sichuan, Wolong Nat. Res., 1500m, 22. and 24.V.1994, S. A. Kurbatov, 11 (MHNG); same data but 1700m, 19.V., 1 (MHNG); Yunnan, Mengyang Nat. Res. ca 500m, 10.1X.1994, S. A. Kurbatov, 1 (MHNG); Yunnan, Jızu Shan, 25°58°N, 100°21’E, 2500-2700m, 6-10. VII.1994, V. Kuban, 1 (NHMB); same data but 2800m, 20.V.-3.VI.1993, Bolm, 1 (NHMB); Yunnan, Lijiang, 26°53’N, 100°18’E, 1800m, 23.VI.-21.VII.1992, S. Beëväï, 2 (MHNG); Yunnan, Heishui, 35km north Lijiang, 27°13’N 100°19’E, 18.VI.-4.VII.1993, S. Beévar, 1 (MHNG). Distribution. North India; Nepal; China: Shaanxi, Sichuan, Yunna. Comments. Females from the Wolong Nat. Res. are tentatively associated with the males. All specimens recorded from other localities are males and were dissected. Scaphobaeocera nobilis Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 8-12.1X.1994, S. A. Kurbatov, 9 (MHNG). Distribution. Bhutan; Thailand; China: Yunnan. Scaphobaeocera spinigera Löbl Material examined. China, Central Sichuan, Wolong Nat. Res., 1500m, 21.V.1994, S. A. Kurbatov, 1 (MHNG); Hong Kong, V. and IX-X.1996, G. de Rougemont, 3 (MHNG). Distribution. Pakistan; India; Nepal; Thailand; China: Sichuan, Hong Kong. Scaphobaeocera difficilis Löbl Material examined. China, West Hubei, Shennongia Nat. Res., 2000-2200m, 3-8.VI. 1995, S. A. Kurbatov, 6 (MHNG). Distribution. Pakistan; India; Nepal; Thailand; China: Hubei. Scaphobaeocera spira Löbl Material examined. China, Yunnan, Gaoligong Mts, 2200-2500m, 24°57’N 98°45’E, 8- 16.V.1995, O. Semela, 11 (NHMB, MHNG). Distribution. Nepal; Thailand; China: Yunnan. Scaphobaeocera dorsalis Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 8-14.[X.1994, S. A. Kurbatov, 28 (MHNG); Central Sichuan, Wolong Nat. Res., 1700, 19.V.1994, S. A. Kur- batov, 2 (MHNG). Distribution. India; Nepal; Thailand, Taiwan; China: Sichuan, Yunnan. Fics 64-69 Aedeagi in Scaphobaeocera; 64 and 65. S. simplex sp. n.; 66 and 67. S. molesta sp. n.; 68 and 69. B. pseudovalida sp. n. Scale bar = 0.1 mm. 780 SCAPHIDIINAE OF CHINA, I 69 67 65 738 IVAN LOBL Scaphobaeocera simplex sp. n. Figs 64, 65 Holotype d: China, Central Sichuan, Wolong Nat. Res., 1500m, 22.V.1994, litter, S. A. Kurbatov (MHNG). Description. Length 1.4 mm, dorsoventral diameter 0.78 mm. Body very dark, almost black, with reddish lustre. Abdominal ventrites 2 to 5, coxae, femora and tibiae reddish-brown, abdominal apex and tarsi yellowish. Pronotum, elytra, lateral portions of metasternum and abdominal sternites distinctly microsculptured and iridescent. Punctation very fine on dorsal and ventral surfaces of body. Relative length of antennal segments 3 to 11 as: 8: 10: 12: 15: 9: 13: 14: 20. Segments 3 to 6 almost equally narrow, slightly widen apically, segment 3 about 2.5 times as long as wide; segments 7 and 8 each about 2 times as long as wide; segment 11 twice as long as wide. Prohypomera without longitudinal stria. Elytra with distinct parasutural striae. Median portion of metasternum shallowly impressed, with anteriomedian stria. Metepisterna flat, about 0.10 mm wide, slightly narrowed anteriorly. Mesocoxal areas 0.02 mm long, with minute marginal pits. Basal pits of sternite I minute. Tibiae straight. Male sexual characters. Segments | to 3 of protarsi strongly widen, almost as wide as apex of protibia. Aedeagus (Figs 64, 65) 0.43 mm long. Median lobe with minute ventral process, apical portion inflexed, concave ventrally, tip truncate in dorsal view, acute in lateral view. Internal sac with simple, arcuate, gradually narrowed flagellum. Parameres straight, converging apically, widen in apical two thirds. Comments. This species may be easily distinguished from its congeners by the shape of the simple flagellum. Scaphobaeocera australiensis Löbl possesses also a flagellum very simple, but it is bent irregularly, and much narrower than in the new species. In addition, the colour patter and the shape of the median lobe in S. australiensis are clearly different. Scaphobaeocera molesta sp. n. Figs 66, 67 Holotype d: China, South Yunnan, Mengyang Nat. Res., ca 500m, 9.IX.1994, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 1 8,5 2; same data as holotype but 11.IX., 1 d; same data but 14.IX., 1 4 (all MHNG). Description. Length 1.4-1.5 mm, dorsoventral diameter 0.80-0.86 mm. Body reddish-brown, elytra darkened toward apex. Apex of abdomen, legs and antennae lighter. Body microsculptured and iridescent. Punctation extremely fine and indistinct on pronotum, lateral portions of metasternum and abdomen, distinct on elytra. Relative length of antennal segments 3 to 11 as: 12: 20: 22: 19: 22: 17: 24: 25: 30 (holotype). Segments 3 to 5 almost equally narrow, not widen apically, segment 3 about 3 times as long as wide; segment 6 distinctly wider than segment 5, 4 times as long as wide; segment 7 about 3 times as long as wide; segment 8 about twice as long as wide; segment 11 3 times as long as wide (holotype). Apex of scutellum exposed. Elytra with distinct parasutural striae. Prohypomera without longitudinal stria. Middle portion of metasternum flat, lacking stria or impression, distinctly, very densely punctate and pubescent. Metasternal sides microsculptured. Mesocoxal areas 0.02-0.03 mm long, with marginal pits obsolete. Metepisternum 0.0.4-0.05 mm wide, flat, parallel-sided, with straight suture. Basal pits of sternite I obsolete. Tibiae straight. SCAPHIDIINAE OF CHINA, I 739 Male sexual characters. Segments | to 3 of protarsi strongly widen, segment | wider than base of protibiae, segment 2 about as wide as base of protibiae, segment 3 distinctly narrower than segment 2. Aedeagus (Figs 66, 67) 0.43-0.45 mm long. Median lobe with apical portion strongly inflexed, ventral process distinct, tip acute. Internal sac with flagellum narrow, forming two circles, widen basally, lacking obvious additional sclerotized pieces. Parameres straight in dorsal view, slightly curved in lateral view, rather abruptly widen apically, extending behind level of tip of median lob. Comments. This species is similar and likely closely related to S. dispar Löbl, with which it shares most characters. It may be distinguished by the aedeagus with parameres more abruptly widen apically and the spiral flagellum of the internal sac. Scaphobaeocera pseudovalida sp. n. Figs 68, 69 Holotype d: China, South Yunnan, Mengyang Nat. Res., ca 500m, 11.1X.1994, litter, S. A. Kurbatov (MHNG). Paratypes: 1 d same data as holotype, 1 d same data but 10.IX. (MHNG). Description. Length 1.55-1.60 mm, dorsoventral diameter 0.85 mm. Body, femora and tibiae uniformly reddish-brown. Tarsi and antennae lighter than body. Body microsculptured and iridescent. Punctation extremely fine and indistinct on pronotum, lateral portions of metasternum and abdomen, distinct on elytra. Relative length of antennal segments 3 to 11 as: 9: 16: 20: 20: 21: 17: 21: 24: 30 (holotype). Segments 3 and 4 almost equally wide, not or hardly widen apically; segment 3 about 2.5 times as long as wide; segments 5 and 6 slightly wider than segment 4; segments 4 to 6 each 4 times as long as wide; segment 7 about 3 times as long as wide, segments 8 and 11 each about 4 times as long as wide, parallel-sided. Scutellum concealed. Elytra with distinct parasutural striae. Prohypomera with deep longitudinal stria. Middle portion of meta- sternal flat, lacking stria, distinctly punctured and pubescent. Metasternal sides micro- scultured. Mesocoxal areas 0.02-0.03 mm long, with marginal pits minute. Metepister- num 0.04-0.05 mm wide, flat, with slightly concave suture. Basal pits of sternite I obsolete. Protibiae and mesotibiae straight, metatibiae slightly curved. Male sexual characters. Segments | and 3 of protarsi strongly widen, segments | and 2 each about as wide as base of protibiae, segment 3 slightly narrower. Aedeagus (Figs 68, 69) 0.39-0.42 mm long. Median lobe with apical portion hardly inflexed, lacking ventral process, tip blunt. Internal sac with flagellum narrow, forming one complete circle, widen basally, and with additional subbasal sclerites. Parameres almost straight, gradually widen apically, extending slightly posterior level of tip of median lobe. Comments. This species is similar to S. aberrans Löbl, S. pecki Löbl, S. robus- tula Löbl and S. valida Löbl by the external and aedeagal characters. It may be distinguished from these four species by the long antennal segment 8. In addition, it differs from these species, except S. valida, by the parameres which become gradually wider in lateral view and by the flagellum of the internal sac curved in middle to form one complete circle. The new species may be distinguished from S. valida, in addition to the antennal characters, by the smaller size of the body and by the tip of the para- meres not curved mesially. 740 IVAN LOBL Scaphoxium Lob! This genus comprises 26 species distributed in Asia, Australia, Melanesia, and New Zealand. It was not yet recorded from China. Two species of Scaphoxium were found in the collections examined. Scaphoxium intermedium Löbl Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 11.1X.1994, S. A. Kurbatov, 2 (MHNG). Distribution. North India; Thailand; China: Yunnan. Comments. The single Chinese specimen is a male. It possesses exactly the same aedeagal characters as the numerous specimens from India and Thailand I have examined. Scaphoxium ?taiwanum Löbl Material examined. China, Central Sichuan, Wolong Nat. Res., 1000m, 24.V.1994, S. A. Kurbatov, 3; same data but ca 500m, 15-16.V., 1; Sichuan, Mt. Emei, 1700m, 22.IX.1994, S. A. Kurbatov, 1; same data but 1000m, 4-20.V.1989, S. Kuban, 2 (all MHNG). Comments. Scaphoxium taiwanum is known from India, Nepal, Thailand, and Taiwan. The species 1s well defined by the internal sac of the aedeagus which bears two pairs of conspicuous rods. The basal rods are straight, the more distal rods are apically curved, hook-like. Only two males were found in the material from China. One, from the Wolong Nat. Res., has the internal sac extruded, the second, from Mt. Emei, has the internal sac apparently deformed. Both have the hook-like sclerites more robust than those ın males examined from other localities, and the pair of parallel rods untraceable. Toxidium LeConte This genus is probably a paraphyletic assemblage (LOBL 1992). The ten Asian species of Toxidium which are placed in the T. aberrans group, together with Scaphi- schema poupilieri (Reiche) from the western Mediterranean area, are similar and likely related, in respect to the remaining species assigned to Toxidium. An additional species comes from China and is described below. The key to the Asian species of Toxidium in LÒBL (1992) gives an incorrect statement in the couplet «8», concerning the basal striae of the elytra. The three involved species (T. diffidens, T. pubistylis, and T. robustum), possess elytra with basal striae not joined to the lateral striae. A new key is provided to replace the inconvenient couplet, and to accomodate the new species. KEY TO THE ASIAN SPECIES OF TOXIDIUM l Elytra bright reddish, with blackish adsutural and apical areas 7. spectabile Löbl - Blytra unicolour or with light transverse fasciae . . 5). PE EEE 2 2 Metasternum short, mesocoxal areas about as long as shortest intervals hetweensmesocoxallmes andmetacoxat ha. ee T. vagans Löbl - Metasternum long, mesocoxal area much shorter than interval between mesocoxal lines and'metacoxa.... .... LL STE 3 SCAPHIDIINAE OF CHINA, I 74] 3 Elytraswithoutibasaksnaee te. 1 me ee ee PRE et. he ARRE 4 - Elytrawithbasalstnlaeree fits) sf. atei Apes Eee RE RE 5 4 Elytral punctation fine and irregular. Parameres of aedeagus smooth D PA ys rear hr. I LISSONE ite ad AN SPO. Ne T. incompletum Lob! - Elytral punctation coarse and fairly regular. Parameres of aedeagus DUDESCENt HE en see Pe EN mer cui E T. villosum sp. n. 5 Mesepimeral ridge present. Lateral portions of metasternum fairly coar- SELNADUNC IAE Apa ies 2 o ood Ge eae T. indicum Achard - Mesepimeral ridge absent. Lateral portions of metasternum very finely (LUVIN CLEATS ic Sepals ty Na ee akc td braves te armen tas EEE ES 6 6 Pronotal punctation coarse, distinct at 12x magnification . . . 7. aberrans Achard - Pronotal punctation very fine, hardly visible at 24x magnification.......... 7 7 Elytra with sutural striae long, starting neat elytral base .................. 8 - Elytra with sutural striae very short, present only near elytral apices . ....... 9 8 Ely tratentinely. coarselypunetate an 2m. N. T. curtilineatum Champion i Apicaleportioniomelytrastinely punctate). "Fe een. EURE T. styligerum Löbl 9 Internal sac of aedeagus with long, apically bifid rod and short, V-shaped subapiealaselerite 2 3) rg AMONT SIINO ee T. pubistylis Löbl - Internal sac of aedeagus with sclerotized rod not bifid, and without Additional Sele nites: u. me ee ase esas Cranky Lut BE REVUE SUISSE DE ZOOLOGIE TOME 106 — FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève FRANCOIS BAUD Conservateur au Muséum d'histoire naturelle de Genève CHARLES LIENHARD Charge de recherche au Muséum d’histoire naturelle de Geneve Comite de lecture Président: Ivan LOBL — Muséum de Genéve Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Geneve et de représentants des Instituts de zoologie des universités suisses. Les manuscrits sont soumis a des experts d’institutions suisses ou étrangères selon le sujet étudié. La preference sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L'ABONNEMENT: (en francs suisses) Les demandes d'abonnement doivent étre adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse SUISSE Fr. 225.— UNION POSTALE Fr. 230.— REVUE SUISSE DE ZOOLOGIE 106 (4): 745-787; décembre 1999 ZOOLOGIA ET BOTANICA ‘99 Zurich, 17-19 February 1999 (Joint meeting of the Swiss Society of Zoology and the Swiss Society of Botany) 17 February 1999: Satellite Symposium: Evolution, Ecology and Behaviour 18-19 February 1999: Main Symposium: Genes, Populations and Politicians: Issues in Conservation Biology Author Index ANHOLT Bradley R., see THURNHEER Sylvie et al. ANTOINE Céline, MULLER Jessica & CASTELLA Emmanuel Assemblages of freshwater molluscs in a fringing wetland (Lake Neuchatel, Switzerland) see also OERTLI Beat et al. ARNOLD Claire, GILLET Francois & GOBAT Jean-Michel Ecological knowledge and species conservation: the example of the European wild grapevine Vitis vinifera subsp. sylvestris (Gmel.) Hegi AUDERSET-JOYE Dominique, see OERTLI Beat ef al. BAER Boris, MAILE Roland, SCHMID-HEMPEL Paul, MORGAN E. David & JONES Graeme R. Chemistry and possible function of a mating plug in bumblebees BAER Boris & SCHMID-HEMPEL Paul Multiple mating: an advantage against parasites BAKKER Theo C. M., see KUENZLER Reto et al. BALLOUX Francois, see LUGON-MOULIN Nicolas et al. & PERRIN Nicolas et al. BAZIN Alain, see GOVERDE Marcel er al. BERSIER Louis-Félix, see HOFER Ulrich er al. BOLLENS Ursula & RYSER Peter Nutrients and water regime: Effects on 16 wetland species BORCARD Daniel, see HOFER Ulrich ef al. BRASCHLER Brigitte Influences of small-scale habitat fragmentation on ants BRAUER Ingo, see MATTHIES Diethart et al. BREM Dominik & LEUCHTMANN Adrian Transmission mode of a grass endophyte with mixed strategy of reproduction BRUNNER Harald, see LUGON-MOULIN Nicolas et al. BUCHLER Urs, see GUGERLI Felix et al. CAMBIN Diana, see CASTELLA Emmanuel et al., OERTLI Beat et al. & PULFER Corinne et al. CASTELLA Emmanuel, Lops-CROZET Brigitte, CAMBIN Diana, ILG Christiane, KNISPEL Sandra, MARION Thomas, MARMONIER Pierre & SIMANT Helene 746 ZOOLOGIA ET BOTANICA ‘99 Physical habitat and zoobenthic diversity of a glacial stream (the Mutt, Swiss Alps) CASTELLA Emmanuel, SPEIGHT Martin C. D. & GOELDLIN DE TIEFENAU Pierre Using species pool and digitised biological data for biodiversity assessment and conservation: a test case with Syrphidae (Diptera) see also ANTOINE Céline et al., ILG Christiane et al., OERTLI Beat et al. & PULFER Corinne et al. CASTELLA Vincent, RUEDI Manuel, EXCOFFIER Laurent & IBANEZ Carlos Do males and females mouse-eared bats cross the Gibraltar Strait? Molecular markers say no! CHAUTEMS Alain, see PERRET Mathieu ef al. CONEDERA Marco, see MORETTI Marco et al. DEGEN Thomas, see GOUINGUENE Sandrine et al. Di GiuLIo Manuela, EDWARDs Peter J. & MEISTER Erhard Enhancing insect diversity in agricultural grasslands: the role of management and landscape structure DIETRICH Barbara & WEHNER Rüdiger Coexistence of parapatric species: how do desert ants solve this contradiction ? DUELLI Peter, see MORETTI Marco et al. EDWARDS Peter J., see Di GruLio Manuela et al., KELLER Michael et al., MORETTI Marco et al., STUDER Sibylle et al. & ULLRICH Karin S. et al. EEK Liina, see RYSER Peter er al. ERHARDT Andreas, see GOVERDE Marcel ef al. & RUSTERHOLZ Hans-Peter ef al. EXCOFFIER Laurent, see CASTELLA Vincent ef al. FINKELDEY Reiner Molecular markers and the conservation of forest resources - 1s there an impact ? FISCHER Markus, VAN KLEUNEN Mark & SCHMID Bernhard Relations among population size, genetic variation, and plant performance: the example of Ranunculus reptans FORSTNER Michael R. J., see PASTORINI Jennifer et al. FREIMANN Barbara & WARD Paul I. Male seminal products influence female sperm storage muscles in Scatho- phaga stercoraria FREYMOND Hervé & GALLAND Nicole Genetic diversity and conservation of Saxifraga hirculus FUMAGALLI Luca & TABERLET Pierre Genetic affiliation of historical wolves from Switzerland GAECHTER Fani, see Roy Bitty et al. GALEUCHET David J., RUTISHAUSER Rolf & SCHNELLER Johann Jakob Typha minima - formerly common, now endangered. Genetic variability and relationship between fragmented and ex situ populations GALLAND Nicole, see FREYMOND Hervé et al. GANTENBEIN Benjamin, LARGIADER Carlo R. & SCHOLL Adolf Nuclear and mitochondrial gene variation among populations of Buthus occi- tanus (Amoreux, 1789) across the Strait of Gibraltar ZOOLOGIA ET BOTANICA ‘99 747 GAUTSCHI Barbara & TENZER Isabel Development of microsatellite markers for the Bearded Vulture (Gypaetus barbatus) and their application to the reintroduction project GEBHARDT-HENRICH Sabine Reproductive success was predicted by body mass loss in female great tits (Parus major) GERLOFF Christine U., OTTMER Birgit K. & SCHMID-HEMPEL Paul Inbreeding in bumble bees GILLET Francois, see ARNOLD Claire er al. GOBAT Jean-Michel, see ARNOLD Claire ef al. GOELDLIN DE TIEFENAU Pierre, see CASTELLA Emmanuel et al.» GOUDET Jérome A population genetics view of multivariate statistical methods see also LUGON-MOULIN Nicolas et al. & PERRIN Nicolas et al. GOUINGUENE Sandrine, DEGEN Thomas & TURLINGS Ted How parasitoids of caterpillars find suitable hosts ? GOVERDE Marcel, BAZIN Alain, SHYKOFF Jacqui & ERHARDT Andreas Influence of leaf chemistry of Lotus corniculatus (Fabaceae) on larval deve- lopment of Polyommatus icarus (Lepidoptera, Lycaenidae): effects of elevated CO, and plant genotype GUGERLI Felix, SPERISEN Christoph, BUCHLER Urs & SENN Josef Genetic diversity in tree species of subalpine forests: a case study of Norway spruce (Picea abies) GUSEWELL Sabine Does the public support restoration measures in fen meadows invaded by common reed ? HAGGSTROM Hakan, RAHIER Martine & HARTMANN Thomas Deterrent effect of pyrrolizidine alkaloids and detoxification by a generalist predator (Mitopus morio) HARTMANN Thomas, see HAGGSTROM Hakan et al. HAUSSER Jacques, see LUGON-MOULIN Nicolas et al. HEEB Philipp Within-pair copulation frequency predicts female reproductive performance in tree sparrows, Passer montanus HELLRIEGEL Barbara & REYER Heinz-Ulrich Which factors influence the composition of mixed populations of a hemiclonal hybrid (Rana esculenta) and its sexual host ? HERGER Franziska, LARGIADER Carlo R., LURTSCHER Mathias & SCHOLL Adolf The Austropotamobius pallipes species complex in the Alpine region - phylo- geographic studies using nuclear and mitochondrial markers HEYLAND Andreas Behaviour and mutual influencing of two polychaete species with different life-cycles during settlement: Poecilochaetus serpens E. Claparede and Sphae- rosyllis sp. (Polychaeta: Syllidae: Exogoninae) 748 ZOOLOGIA ET BOTANICA ‘99 HOFER Ulrich, BERSIER Louis-Felix & BORCARD Daniel Spatial organization of a tropical forest herpetofauna on an elevational gradient revealed by null model tests HÖRDEGEN Philipp, see MORETTI Marco et al. IBANEZ Carlos, see CASTELLA Vincent et al. ILG Christiane, CASTELLA Emmanuel & Lops-CROZET Brigitte Macroinvertebrate biodiversity in glacial catchment: the rule of water sources and tributaries see also CASTELLA Emmanuel er al. JENSEN Deborah B. Conservation by design: the theory and practice of conservation priority setting JOKELA Jukka, see RIGBY Mark C. et al. JONES Graeme R., see BAER Boris et al. JUGE Raphaelle, see OERTLI Beat er al. KALBERER Nicole & RAHIER Martine The cost of dispersal by flight in an alpine leaf beetle Oreina cacaliae (Cole- optera: Chrysomelidae) KELLER Laurent, see REUTER Max et al. KELLER Michael, KOLLMANN Johannes & EDWARDS Peter J. Genetic introgression from distant provenances reduces fitness in local weed populations - a case study with species for ecological compensation areas KERY Marc Inferring the absence or extinction of a species from a site: a case study with three snake species KNADEN Markus Enemy recognition in the desert ant Cataglyphis fortis KNISPEL Sandra, see CASTELLA Emmanuel et al. KOLLIKER Mathias Heritability of offspring begging and its co-evolutionary potential with paren- tal feeding behaviour see also ROULIN Alexandre er al. KOLLMANN Johannes, see KELLER Michael er al. & PFLUGSHAUPT Kaspar et al. KOLNAAR Rogier Effect of Puccinia lagenophorae on competition between Senecio vulgaris L. and Capsella bursa-pastoris KUENZLER Reto & BAKKER Theo C.M. A study of multiple female preferences in sticklebacks using computer ani- mations KUSKE Stefan Does the mass release of the exotic egg parasitoid Trichogramma brassicae pose a risk to populations of endemic natural enemies ? LACHAVANNE Jean-Bernard, see OERTLI Beat er al. LANDOLT Martin, see Roy Bitty et al. ZOOLOGIA ET BOTANICA ‘99 749 LARGIADER Carlo R., see GANTENBEIN Benjamin et al. & HERGER Franziska et al. LAWTON John H. Biodiversity and ecosystem function LEUCHTMANN Adrian, see BREM Dominik et al. Lops-CrOZET Brigitte, see CASTELLA Emmanuel et al. & ILG Christiane et al. LuGON-MOULIN Nicolas, BRUNNER Harald, BALLOUX Francois, GOUDET Jerome & HAUSSER Jacques Do riverine barriers, history or introgression shape the genetic structuring of a common shrew (Sorex araneus) population ? LURTSCHER Mathias, see HERGER Franziska et al. MACDONALD David Alien invaders MAIBOM Wiebke, see MATTHIES Diethart ef al. MAILE Roland, see BAER Boris et al. MARION Thomas, see CASTELLA Emmanuel et al. MARMONIER Pierre, see CASTELLA Emmanuel et al. MARTIN Robert D., see PASTORINI Jennifer er al. MATTHIES Diethart, BRÄUER Ingo, MAIBOM Wiebke & TSCHARNTKE Teja Population size and risk of extinction in rare plants MEISTER Erhard, see Di GruLio Manuela et al. & STUDER Sibylle er al. MEYER Andrea, see STEPHAN Andre et al. MICKASCH Tatjana M. Is there a relationship between expanding Orconectes limosus and decreasing Austropotamobius pallipes populations in Switzerland ? Mikos Z. Maja & WARD Paul I. Cryptic female choice of male genotype in the yellow dung fly (Scathophaga stercoraria) MORETTI Marco, HORDEGEN Philipp, CONEDERA Marco, DUELLI Peter & ED- WARDS Peter J. Response of invertebrates to wildfires. First results about spiders (Araneae) and ground beetles (Carabidae) in deciduous forests on the south part of the Alp (Ticino, Switzerland) MORGAN E. David, see BAER Boris et al. MÜLLER Jessica, see ANTOINE Céline et al. & OERTLI Beat et al. MULLER Jiirg P. The reintroduction of the Bearded Vulture, Gypaetus barbatus, in the Alps — an example for the practical project realisation OERTLI Beat, AUDERSET- JOYE Dominique, JUGE Raphaelle, CASTELLA Emmanuel, MULLER Jessica, ANTOINE Céline, SAAM Mirko, CAMBIN Diana & LACHA- VANNE Jean-Bernard Biodiversity in Swiss ponds: how biotic communities respond to altitude see also PULFER Corinne et al. 750 ZOOLOGIA ET BOTANICA ‘99 OLIVIERI Isabelle Contribution of metapopulation-level studies to the understanding of extinc- tion, adaptation, and speciation OTTMER Birgit K., see GERLOFF Christine U. et al. PASCHKE Melanie Does genetic variation matter ? PASINELLI Gilberto Effects of habitat structure on home range size and breeding success of the middle spotted woodpecker Dendrocopos medius PASTORINI Jennifer, FORSTNER Michael R. J. & MARTIN Robert D. Phylogenetic relationships among Lemuridae (Primates) inferred from DNA sequences PERRET Mathieu, SAVOLAINEN Vincent, CHAUTEMS Alain & SPICHIGER Rodolphe Pollination systems and speciation in Sinningieae (Gesneriaceae, Angiosperm) PERRIN Nicolas, BALLOUX Francois & GOUDET Jerome Breeding systems inferred from genetic structures PFLUGSHAUPT Kaspar & KOLLMAN Johannes Reproductive traits in small and isolated populations of Prunus mahaleb PFUNDER Monika, see SCHURCH Stéphanie et al. PULFER Corinne, OERTLI Beat, CASTELLA Emmanuel & CAMBIN Diana Biodiversity of Trichoptera in Swiss ponds RAHIER Martine, see HAGGSTRÖM Hakan et al. & KALBERER Nicole et al. REUTER Max & KELLER Laurent Conflict over caste determination in eusocial hymenoptera REYER Heinz-Ulrich, see HELLRIEGEL Barbara ef al. RIBI Georg Habitat alterations facilitate genetic introgression between European Viviparus species (Mollusca, Prosobranchia) see also VORBURGER Christoph et al. RICHNER Heinz, see ROULIN Alexandre er al. RIGBY Mark C. & JOKELA Jukka High costs of predator avoidance compromise immune defences in snails (Lymnaea stagnalis) ROMER Jann Astacus leptodactylus - a threat to native species in Lake Aegeri, Switzerland ? ROULIN Alexandre, KOLLIKER Mathias & RICHNER Heinz Hawk-dove games among offspring of Tyto alba Roy Barbara A., see SCHURCH Stéphanie et al. & UTELLI Anna-Barbara et al. Roy Bitty, LANDOLT Martin, GAECHTER Fani & SCHMID Bernhard Relationships between biodiversity and enemy attack rates RUEDI Manuel, see CASTELLA Vincent ef al. & SALAMIN Nicolas et al. RUSTERHOLZ Hans-Peter & ERHARDT Andreas Influence of human land-use change on the genetic population structure of the Adonis Blue butterfly (Lysandra bellargus Rott) ZOOLOGIA ET BOTANICA ‘99 751 RUTISHAUSER Rolf, see GALEUCHET David J. et al. RYSER Peter, ZOBEL Kristjan, EEK Liina, URBAS Pille & WAHL Stefan Plant characteristics which drive succession see also BOLLENS Ursula et al. SAAM Mirko, see OERTLI Beat et al. SALAMIN Nicolas, RUEDI Manuel & SAVOLAINEN Vincent Inferring complex phylogenies: which strategy to adopt ? Saucy Francis, see VIETTE Michele er al. SAVOLAINEN Vincent, see PERRET Mathieu er al. & SALAMIN Nicolas et al. SCHIEGG Karin - Patch size and patch connectivity: habitat relationships of saproxylic Diptera and Coleoptera - Small but beautiful: influence of dead wood dimensions on species richness and diversity of saproxylic insects SCHMID Bernhard, see FISCHER Markus ef al. & Roy Bitty er al. & STEPHAN André et al. SCHMID-HEMPEL Paul, see BAER Boris et al. & GERLOFF Christine U. et al. SCHNELLER Johann Jakob, see GALEUCHET David J. et al. SCHOLL Adolf, see GANTENBEIN Benjamin er al. & HERGER Franziska et al. SCHURCH Stéphanie, PFUNDER Monika & Roy Barbara A. Interactions between ants, Euphorbia cyparissias, and its pathogenic rust fungi SENN Josef, see GUGERLI Felix et al. SIMANT Hélène, see CASTELLA Emmanuel ef al. SHYKOFF Jacqui, see GOVERBE Marcel et al. Soliva Marco & WIDMER Alex Phylogenetic relationships within the orchid genus Ophrys based on multiple DNA sequences SPEIGHT Martin C. D., see CASTELLA Emmanuel et al. SPERISEN Christoph, see GUGERLI Felix et al. SPICHIGER Rodolphe, see PERRET Mathieu et al. STEPHAN André, MEYER Andrea & SCHMID Bernhard Plant diversity positively affects soil microbial diversity STUDER Sibylle, EDWARDS Peter J. & MEISTER Erhard Enhancing biodiversity in agricultural grasslands: plant community structure under different managements SUTER Werner A biodiversity strategy as a national nature conservation concept TABERLET Pierre, see FUMAGALLI Luca et al. TENZER Isabel, see GAUTSCHI Barbara et al. TETTAMANTI Chiara, see VIETTE Michéle et al. THURNHEER Sylvie & ANHOLT Bradley R. Higher order interactions: The influence of predator-induced indirect effects on competitive ability in waterfrog larvae (Rana esculenta and Rana lessonae) TSCHARNTKE Teja, see MATTHIES Diethart et al. 752 ZOOLOGIA ET BOTANICA ‘99 TURLINGS Ted, see GOUINGUENE Sandrine et al. ULLRICH Karin S. & EDWARDS Peter J. Enhancing biodiversity in arable land: the role of management and landscape structure URBAS Pille, see RYSER Peter ef al. UTELLI Anna-Barbara & Roy Barbara A. Altitudinal variation in nectar robbery in Aconitum lycoctonum VAN KLEUNEN Mark, see FISCHER Markus et al. VIETTE Michele, TETTAMANTI Chiara & SAUCY Francis Preference for acyanogenic white clover (Trifolium repens) in the vole, Arvi- cola terrestris in simple and repeated choice tests VORBURGER Christoph & RIBI Georg Mechanisms of impact of the introduced crayfish Pacifastacus leniusculus on the native crayfish Austropotamobius torrentium in Switzerland VOUILLAMOZ José The genus Onosma (Boraginaceae) in Switzerland: chromosome numbers, allozymic polymorphism and evolutionary history of a group of rare taxa WAHL Stefan, see RYSER Peter et al. WALTHER Gian Reto Why palms in Switzerland ? WARD Paul I., see FREIMANN Barbara ef al. & Mikos Z. Maja et al. WEHNER Rüdiger, see DIETRICH Barbara et al. WEIBEL Urs Nest site selection and breeding success of skylarks (Alauda arvensis) in an arable landscape with wild flower strips WEISSER Wolfgang W. Predator-induced morphological shift in the pea aphid WIDMER Alex, see SOLIVA Marco ef al. & ZUBER Doris et al. ZOBEL Kristjan, see RYSER Peter er al. ZUBER Doris & WIDMER Alex Intraspecific phylogenetics of the European mistletoe (Viscum album) based on nuclear and chloroplast DNA sequences ZOOLOGIA ET BOTANICA ‘99 753 ‘ Abstracts (alphabetically by first author) ANTOINE Céline, MULLER Jessica & CASTELLA Emmanuel Laboratoire d'Ecologie et de Biologie Aquatique; Université de Geneve; chemin des Clochettes 18; CH-1206 Genève; Switzerland Assemblages of freshwater molluscs in a fringing wetland (Lake Neuchatel, Switzer- land) Freshwater gastropod assemblages were surveyed in 9 contrasted ponds of the largest Swiss wetland (fringe of Lake Neuchatel). The survey was carried out as a preliminary assessment of molluscan biodiversity in ponds selected along gradients of age and hydrological connections. It is intended as a basis for 1) selection of sites for monitoring of the dynamics of key species, and 2) prioritising management options aiming at biodiversity preservation/enhancement. Over 22 taxa were recorded among which Valvata macrostoma and Bathyomphalus contortus (L.) are of prominent interest. Relationships between habitat types, water/sediment characteristics, molluscan diversity and species composition are discussed. ARNOLD Claire, GILLET Francois & GOBAT Jean-Michel University of Neuchatel; Institute of Botany; Laboratoire d'écologie végétale; rue Emile- Argand 11; CH-2007 Neuchatel; Switzerland Ecological knowledge and species conservation: the example of the European wild grapevine Vitis vinifera subsp. sylvestris (Gmel.) Hegi Since the beginning of the century there has been a very marked decrease of wild grapevine in Europe, resulting in the fragmentation of its distribution area. This reduction of the populations is directly or indirectly linked to human activities and to a lack of knowledge of this subspecies. From an historical point of view, the discovery of typical wild grapevine seeds in Neolithic sites shows that this subspecies is anterior to subspontaneous cultivars on the European territory. New genetical molecular technologies based on microsatellites show the presence of alleles specific to wild grapevine, which prove the originality of subsp. sylvestris. The ecolo- gical study of wild grapevine highlights both the necessary natural conditions for maintaining population dynamics, and the dangers which threaten it. Seedlings and young plants were observed in only 11% of the 163 studied stations. This established fact leads us to suppose that present conditions are no longer propitious to regeneration in a majority of sites whose per- manence is endangered. All attempts at reintroduction in the Rhine floodplain forests have so far been unsuccessful. Precise studies of active regeneration stations should make it possible to define the optimum conditions for site revitalisation or for the successful reintroduction of wild grapevine in stations where it existed in the past. BAER Boris!, MAILE Roland, SCHMID-HEMPEL Paul!, MORGAN E. David? & Jones Graeme R.? 4 Experimental Ecology; ETH Zürich; Switzerland “ Keele University; Chemistry Department; Keele Staffordshire; United Kingdom Chemistry and possible function of a mating plug in bumblebees Previous work showed that queens of the bumblebee Bombus terrestris would benefit from multiple mating due to lower parasite load and higher fitness. Nevertheless, B. terrestris is mostly singly mated in Central Europe. One possible solution to this phenomena is that the male bumblebee does not want to share its fitness with other males and therefore prevents mul- tiple mating by transferring a sticky plug into the queen's sexual tract. Chemical compounds in 754 ZOOLOGIA ET BOTANICA ‘99 the plug may influence the queen's behaviour, i.e. perhaps preventing her from mating again ? Using gas chromatography we identified cyclo-prolylproline and 4 different fatty acids present in the plug. We then mixed the same concentration of these compounds to create a "synthetic plug". This will enable us to test effects of these chemicals on male and queen mating be- haviour. BAER Boris & SCHMID-HEMPEL Paul Experimentelle Ökologie; ETH Zentrum NW; CH-8092 Zürich; Switzerland Multiple mating: an advantage against parasites Multiple mating by males (polyandry) and females (polygyny) is common in animal species. Whereas the advantages for males are obvious, the reasons for a female to mate several times are more difficult to understand. In social insects multiple mating by males and females reduces relatedness within colonies and thus is problematic for a kin selection explanation for the maintenance of eusociality. Nevertheless, multiple mating is common in social insects and several hypothesis have been proposed to explain this fact. One of them states that multiple mating increases genetic variability providing an advantage against parasites. We tested this hypothesis using the bumblebee Bombus terrestris (L.). Using artificial insemination, we were able to produce bumblebee colonies of high and low genetic diversity. These colonies were placed in the field and worker samples were collected and checked for the presence of parasites. For the two most common parasites, Crithidia bombi and Nosema bombi, we found that inten- sities and prevalence were lower in colonies of high genetic diversity than in colonies of low genetic diversity. Additionally, for all parasite species found, parasite load and parasite richness were significantly lower in colonies of high genetic diversity than in colonies of low genetic diversity. Finally, the fitness of colonies of low genetic diversity was lower. These results show that queens benefit from multiple mating with respect to parasitism and fitness. The fact that B. terrestris ıs singly mated in central Europe may be explained by the mating behaviour of the male, which includes mate guarding and the production of a sticky mating plug injected into the female after sperm transfer. BOLLENS Ursula & RYSER Peter Geobotanisches Institut ETH; Zürichbergstrasse 38; CH-8044 Zürich; Switzerland Nutrients and water regime: Effects on 16 wetland species Although protected by law, Swiss wetlands suffer damage from nutrient inflow and interference to the water regime. Because species diversity is highest at a rather low level of biomass pro- duction, it is an objective of wetland management to lower site productivity. Should we focus mainly on keeping nutrients away from the surroundings or is flooding also an appropriate means to promote typical species composition ? To test the response of various fen species to different nutrient availabilities and water levels, a factorial growth experiment was established in spring 1998. The treatments are: base-level fertilisation (0.9 mg N/individual/week, 0.06 mg P/individual/week), threefold nitrogen supply, threefold phosphorus supply: wet conditions, flood periods, and drought periods. Biomass and nutrient content were recorded in September 1998. The first season’s results show a distinct rise of biomass due to additional nitrogen fertilisation and only little effect of the phosphorus treatment. These results apply to all species whether they originate from nutrient poor or nutrient rich sites. Even under enhanced nitrogen supply there is little evidence of phosphorus limitation. All species are affected by flood. Periodic drought causes negative or positive effects on biomass production. Conclusions: (1) Nitrogen was the growth limiting element in the experiment. The growth reducing effect of flooding was probably due to its reducing effect on N-uptake. (2) The results suggest that flooding is an appropriate means of reducing site productivity. However, the water ZOOLOGIA ET BOTANICA ‘99 755 used for flooding should not be nitrogen rich. (3) One growing season is too short for the assessment of species-specific effects of nutrients and flooding. The experiment will be conti- nued for another year. BRASCHLER Brigitte Institut für Natur-, Landschafts-und Umweltschutz; Universität Basel; Johanns-Vorstadt 10; CH-4056 Basel; Switzerland Influences of small-scale habitat fragmentation on ants In many parts of the world, human activities have greatly altered natural landscapes, converting them in mosaics of agricultural, industrial and urban patches. Habitat fragmentation is one of the most apparent consequences of human use of nature. Research has often focused on large- scale habitat fragmentation and its effects on conspicuous species groups like mammals and birds. The influence of small-scale habitat fragmentation on less conspicuous species groups, including many small invertebrates, is less studied. In my work I concentrate on the ant populations in calcareous grasslands in the Swiss Jura Mountains. I especially study influences of the fragmentation on species composition, nest dispersion and changes in interactions between colonies and between the ants and other organisms. Several aspects of foraging behaviour of ants seem to be changed in fragments compared to control plots. For example ants were more numerous at baits in fragments than at baits in control plots. Also ants preferred to forage on baits in the fragment rather than in the surrounding isolation area. Additionally, aphids, an important sugar resource of many ant species, were more abundant in fragments than in control plots. Effects on nest dispersion and species composition are less pronounced. BREM Dominik & LEUCHTMANN Adrian Geobotanisches Institut ETH; Zollikerstrasse 107; CH-8008 Zürich; Switzerland Transmission mode of a grass endophyte with mixed strategy of reproduction Epichloe sylvatica (Ascomycota, Clavicipitaceae) is a host specific endophyte of the woodland grass Brachypodium sylvaticum with two alternative modes of reproduction. The predominant asexual forms are seed transmitted (vertical), whereas the sexual forms are capable of contagious transmission (horizontal) mediated by ascospores. However, transmission rate and mechanisms of infections are undetermined in this system. A transplant experiment and phenological observations were designed to estimate the rate of horizontal transmission and to explore possible routes of new infections. We observed clear coincidence of time of host flowering and availability of ascospores indicating that infection through stigmata is a possible mechanism of horizontal transmission. At sites where sexual forms of E. sylvatica were present 34% or 17% respectively, of the uninfected transplants became infected after two years, whereas at sites with only asexual forms all transplants remained uninfected. This suggests that contagious spread of the endophyte to established plants can occur frequently and that ascospores probably serve as inoculum. Besides the stigmata, stem or leaf tissues may provide an alternative route of entry, which can lead to new infections. CASTELLA Emmanuel!, Lops-CROZET Brigitte’, CAMBIN Diana!, ILG Christiane!, KNISPEL Sandra”, MARION Thomas, MARMONIER Pierre? & SIMANT Hélène | Laboratoire d'Ecologie et de Biologie Aquatique; Université de Geneve; chemin des Clo- „ee 18; CH-1206 Genève; Switzerland „ Musée de Zoologie; Place Riponne 6; CH-1005 Lausanne; Switzerland > GRETI; Université de Savoie; Le Bourget du Lac; France Physical habitat and zoobenthic diversity of a glacial stream (the Mutt, Swiss Alps) 756 ZOOLOGIA ET BOTANICA ‘99 Within the framework of a European research programme for "Arctic and Alpine Stream Ecosystem Research", studies of the geomorphology, water physico-chemistry and macro- benthic communities were carried out in 1996 and 1997 on the Mutt, a glacier-fed tributary of the Rhone River (Switzerland). The objective of the programme is to test and refine Milner and Petts' model (1994) suggesting water temperature and channel stability as the main variables determining the occurrence and longitudinal succession of invertebrate taxa in glacial streams. Diversity and community composition patterns recorded on the Mutt, with sites ranging from 2600 to 1800 m a.s.l., are confronted with the model. Results evidencing high taxonomic richness for groups like Diptera Chironomidae and Trichoptera Limnephilidae are discussed, together with the current relative low level of taxonomic expertise regarding these groups in alpine ecosystems. CASTELLA Emmanuel!, SPEIGHT Martin C. D.2 & GOELDLIN DE TIEFENAU Pierre? | Laboratoire d'Ecologie et de Biologie Aquatique; Université de Genève; chemin des Clo- chettes 18; CH-1206 Genève; Switzerland ? Research Branch; National Parks and Wildlife Service; 51 St Stepens Green; Dublin 2; Ireland 3 Musée de Zoologie; Place Riponne 6; CH-1005 Lausanne; Switzerland Using species pool and digitised biological data for biodiversity assessment and con- servation: a test case with Syrphidae (Diptera) An overview is given of the use of digitised information about the European species of one group of invertebrates, the Syrphidae (Diptera), in addressing biodiversity maintenance issues. They probably represent no more than 1-2% of Europe's invertebrate biodiversity, so it cannot be claimed that they may be used as proxies for the entire European invertebrate fauna. However, they provide insights into what might be achieved if invertebrates were effectively incorporated into biodiversity maintenance work, rather than being almost universally ignored as at present - although invertebrates do represent approximately 75% of the biodiversity sup- ported by the European continent. The species pool concept is essentially that the biota of part of a region is a sub-set of the biota of that region and so can be predicted from the regional species list, given sets of assembly or deletion rules. In the present case, examples are shown of regional and local syrphid lists, coupled with digitised information on the listed species, invoking the predictive capacity of the regional species pool. The predicted species lists gene- rated are used for assessment of the performance of the biodiversity maintenance function at site level, and for identification of biodiversity maintenance priorities at various spatial levels. The information about Syrphid species is extracted from the "Syrph the Net" database covering some 400 syrphid species known from Atlantic parts of Europe and including spreadsheet files providing information on the habitats, microhabitats, traits, range and status of the included species. CASTELLA Vincent, RUEDI Manuel, ExCOFFIER Laurent & IBANEZ Carlos Institut de Zoologie et d'Ecologie Animale: Université de Lausanne; BB; CH-1015 Lausanne; Switzerland Do males and females mouse-eared bats cross the Gibraltar Strait? Molecular markers say no! Because of their role in limiting gene flow, geographic barriers like mountains or seas often coincide with large genetic discontinuities. Although the Strait of Gibraltar represents such a potential barrier for terrestrial organisms, no studies have been conducted on its impact on gene flow. Here we test this effect on a flying mammal (Myotis myotis) which is distributed on both sides of this Strait. Based on a sample of four populations, mtDNA haplotypes reveal a complete and ancient isolation between Northern and Southern bats. Results based on seven ZOOLOGIA ET BOTANICA ‘99 757 microsatellite loci confirm the presence of this strong barrier to gene flow, suggesting that neither males nor females do actually cross the 15 km of the Gibraltar Strait. This conclusion is surprising since Mouse-eared bats are able to cover daily up to 25 km between roosts and foraging territories. Several alternative explanations to this surprising pattern of genetic divergence are discussed. Di GIULIO Manuela, EDWARDS Peter J. & MEISTER Erhard Swiss Federal Research Station for Agroecology and Agriculture; Reckenholzstrasse 191; CH-8046 Zürich; Switzerland Enhancing insect diversity in agricultural grasslands: the role of management and landscape structure During the last two decades great efforts have been made in Switzerland to preserve and en- hance biodiversity. In this context we are interested in the effects of different management systems and landscape structure on the insect diversity of agricultural grasslands. The research area is located in the Schaffhauser Randen (Jura formation). It is an extensive forested area with a number of more or less isolated enclaves of agricultural land. Four such enclaves have been selected in order to provide areas with varying mixtures of arable and grassland habitats. In each enclave (area), meadows of two grassland management types (medium intensive and 'extensive') have been investigated. As an indicator group for insect diversity true bugs (Heteroptera) were chosen. Previous studies have shown that the richness of the bug fauna correlates strongly with total insect diversity. The bug communities of extensive sites demonstrate clearly a more even rank abundance distribution compared to those of medium intensive sites. Extensive meadows have both more individuals and more species. We used ordination techniques to assess the factors affecting the structure of bug communities. A statistical method based on Canonical Correspondence Analysis (Borcard 1992) was used to partition the variance of the species data into spatial (area) and management components. Our model explains 72.3% of the variance (p=0.01). Management accounts for 29.7% (p=0.01), area for 35.4% (p=0.01) and the interaction ‘management x area’ for 7.2% of the species variation. DIETRICH Barbara & WEHNER Riidiger Institute of Zoology; University of Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Coexistence of parapatric species: how do desert ants solve this contradiction ? The parapatric Cataglyphis-species, bicolor and savignyi inhabit different parts of the lowland/ highland steppes of central Tunisia and the presaharan semidesert of southern Tunisia, respecti- vely. Due to their morphological similarity these two species have been mistaken as one single species since the beginning of this century. Within the study area both species overlap in their distributional ranges. The temporal and spatial foraging patterns of the two Cataglyphis species were investigated particularly in the steppe area around the oasis of El Guettar in central Tunisia. By recording the ants foraging paths we showed that C. bicolor exhibits significantly shorter foraging runs than C. savignyi with respect to both length and time: 90% of the foraging time is spent within an area of radius 14.5 m and 21.0 m, respectively (Mann-Whitney U-test: p = 0.014 (length), p = 0.038 (time)). This result is in accord with the outcome of geobotanical studies of the nesting sites. We were able to demonstrate that they exhibit clear-cut micro- habitat differences. C. bicolor colonies usually occupy the more plentiful micro-habitats than C. savignyi. Furthermore experiments with artificial food baits showed that C. bicolor dominates C. savignyi in direct encounters. These results suggest that coexistence is maintained by the interrelationship of the two species: the dominant species drives the subordinate one out of the higher quality microhabitats. 758 ZOOLOGIA ET BOTANICA ‘99 FINKELDEY Reiner Swiss Federal Institute for Forest, Snow and Landscape Research (WSL); Ziircherstrasse 111; CH-8903 Birmensdorf; Switzerland Molecular markers and the conservation of forest resources - is there an impact ? During the last decade a rapidly increasing number of molecular genetic markers was success- fully applied to forest trees. The urgent need to conserve forest genetic resources has often motivated this research. However, the practical impact of most molecular genetic studies is negligible. At least three main reasons account for this situation: first, research has concentrated on a few widespread, common tree taxa of the temperate and boreal zone (in Europe mainly Picea, Pinus and Quercus species), although the need for conservation measures is most urgent for endemic, rare species of the tropics. Second, research focused on an investigation of spatial patterns of genetic variation at marker loci without straightforward implications for patterns of adaptive variation. Third, new molecular techniques have shifted research priorities from problem-orientation towards method-orientation. Researchers tend to concentrate on the development and validation of new marker types rather than on the application of existing tools to urgent conservation problems. Biochemical and molecular markers have the potential to contribute significantly to the conservation of forest resources if they are used to investigate key processes in endangered forest ecosystems. This implies their application in a population genetic context, their combination with other investigations including ecological field obser- vations and quantitative genetic experiments, and a focus on both spatial and temporal dyna- mics of genetic structures. A recently published study on the reproduction and gene flow of tropical figs (Ficus spp.) based on allozyme markers illustrates the usefulness of gene markers in this regard. FISCHER Markus, VAN KLEUNEN Mark & SCHMID Bernhard Institut fiir Umweltwissenschaften; Universitit Ziirich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Relations among population size, genetic variation, and plant performance: the example of Ranunculus reptans Firstly, we studied selectively neutral RAPD-variation in samples representing comparable areas of 17 populations (3 at Lake Como, 14 at Lake Constance) of the clonal Ranunculus reptans, a rare lake-shore plant. Variation among populations (p < 0.001) corresponds to a low gene flow of Nem = 0.84. Molecular variance was smaller in small populations covering < 100 m2 area than in larger ones (p < 0.03), indicating genetic drift. Secondly, we grew vegetative offspring of 104 genotypes from the 14 Lake Constance populations with and without competition by the grass Agrostis stolonifera in a plant room and measured the number of nodes (rooted, flowering, and total) produced per planted rosette. Plants from larger populations produced more nodes than plants from smaller populations (total p < 0.03; rooted p < 0.1; flowering p < 0.01), whereas positive correlations with molecular variance were not statistically significant. Plasticity (mean without competition minus mean with competition) of node numbers was larger for plants from larger populations (significant for total number of nodes) and for plants from populations with higher molecular variance (significant for number of rooted nodes). Non-significant correlations between broad-sense heritabilities and population size respective molecular variance were positive for the numbers of nodes and negative for plasticities therein. Numbers of nodes and plasticity therein were negatively correlated with the corresponding heritabilities. Because we grew plants in controlled environments our study indicates a genetic basis of observed patterns. Because differences among populations were more closely related to population size than to molecular variance they rather indicate effects of selection than of genetic erosion. Directed selection may also explain observed negative correlations between performance measures and corresponding heritabilities. Our study supports the notion that genetics matter for plant conservation. ; ZOOLOGIA ET BOTANICA ‘99 759 FREIMANN Barbara & WARD Paul I. Zoological Museum of the University of Ziirich; Winterthurerstrasse 190; CH-8057 Ziirich; Switzerland Male seminal products influence female sperm storage muscles in Scathophaga stercoraria The Yellow dungfly Scathophaga stercoraria (L.) is frequently used as a model organism to examine questions about reproductive strategies. In this study we examine the influence of male seminal products on the muscles of the female sperm storage organs, the spermathecae. There are normally three spermathecae, arranged as a singlet and a doublet, which are connec- ted to the bursa copulatrix by long ducts. Isolated female reproductive tracts were treated with different male extracts: from testes, ductus seminalis or as a control, Ringer. It was shown that male testes extracts increase the contraction rate of female spermathecal muscles. There may also be differences between the reactions of singlet and doublets. It is suggested that this mechanism helps to transport sperm into the female spermathecae, as sperm cannot move extensively on their own. The control of muscles responsible for sperm uptake is therefore a "hot spot" in the conflict of interests between male and female in this polygamous fly. FREYMOND Hervé & GALLAND Nicole Institute of Ecology; University of Lausanne; CH-1015 Lausanne; Switzerland Genetic diversity and conservation of Saxifraga hirculus Saxifraga hirculus L. is a tetraploid arctic-boreal plant species; it has a wide circumpolar distribution, whereas its distribution outside Polar regions is highly fragmented. In Switzerland, Saxifraga hirculus was known to occur in at least 25 localities but now only occurs in one locality (Col du Marchairuz, VD, the southernmost European population of this species). As a consequence of such fragmented distribution and of restricted gene flow, genetic drift and inbreeding may occur in the relic populations. Thus, in a conservation perspective, we have addressed three major questions: 1) What is the level of genetic diversity within and among populations ? 2) Is there a spatial genetic structure within populations, and if so, what is the size of the breeding units ? 3) How critical is the vegetative reproduction versus sexual one ? For this purpose, a survey of allozymes/AFLPs diversity of six populations (from Switzerland, Denmark and Iceland) is being undertaken. Methodology and sampling scheme are presented, with the advantages and limits of allozymes and AFLPs. The results of this study will help in designing conservation plans in the future. FUMAGALLI Luca & TABERLET Pierre Laboratoire de Biologie de la Conservation; Institut d'Ecologie; Batiment de Biologie; Univer- sité de Lausanne; CH-1015 Lausanne; Switzerland & Laboratoire de Biologie des Populations d'Altitude; CNRS UMR 5553; Université Joseph Fourier; BP 53; F-38041 Grenoble Cedex 9; France Genetic affiliation of historical wolves from Switzerland Human persecution and forest clearance has led to the eradication of wolf Canis lupus popu- lations in the Alps in general and in Switzerland in particular before the end of the 19th century, even if few isolated individuals of unknown origin have been killed during the 20th century. The species started to recolonize this region from the Italian population in the early 1990s and this process is still going on. We have sequenced a portion of the mitochondrial DNA control region from three historical samples: one individual from a Middle Age site, and two individuals killed in 1947 and 1954, respectively. The mitochondrial data can be useful to characterise whether the population presently colonising the Alps corresponds to the historical lineage disappeared from Switzerland. The analysis reveals that: (1) the DNA sequence from 760 ZOOLOGIA ET BOTANICA ‘99 the Middle Age sample is closely related to a wolf mitochondrial lineage found in Central Europe; (2) the two individuals killed in Switzerland during this century show a mtDNA haplo- type typical of North American wolf populations. Conservation implications of these results are discussed. GALEUCHET David J., RUTISHAUSER Rolf & SCHNELLER Johann Jakob Institute of Systematic Botany; University of Zurich; Zollikerstrasse 107; CH-8008 Ziirich; Switzerland Typha minima - formerly common, now endangered. Genetic variability and relation- ship between fragmented and ex situ populations In Europe Typha minima, a pioneer plant in alluvial zones of alpine rivers, has become endangered. During the last decades the number of populations strongly decreased due to loss of natural habitats. We asked the following questions: Which consequences has fragmentation on the genetic variability of Typha minima ? What are the consequences for the conservation ? Our results showed that in spite of strong fragmentation no genetic drift was detected along the river Rhine. This might be due to the fact that fragmentation of populations took place only recently and/or to clonal growth of the species. Gene flow may be still possible by water transport (seeds) downstream. However, the distance between populations seems to be too far for effective wind dispersal (pollen, seed). To preserve 7. minima from extinction and to rebuild a functional metapopulation along the river Rhine it is necessary to reintroduce plants from ex situ populations. A reintroduction into the Rhone valley is questionable, because the ex situ populations in the botanical gardens of Fribourg and Geneva are not known. GANTENBEIN Benjamin, LARGIADER Carlo R. & SCHOLL Adolf Institute of Zoology; Division of Population Biology; University of Bern; Baltzerstrasse 3; CH- 3012 Bern; Switzerland Nuclear and mitochondrial gene variation among populations of Buthus occitanus (Amoreux, 1789) across the Strait of Gibraltar We analyzed population samples of Buthus occitanus (Amoreux, 1789) (Scorpiones: Buthidae) across the Strait of Gibraltar using allozyme electrophoresis (15 loci scored) and sequencing a 460 bp fragment of the 16S rRNA mitochondrial gene. The samples were collected from three sites in Spain and France (ssp. occitanus), eleven sites in Morocco (ssp. occitanus, ssp. mardochei and ssp. paris) and one site in Tunisia (ssp. tunetanus). Also, B. atlantis from five sites along the Atlantic coast of Morocco were included, this taxon was alternatively treated as a subspecies of B. occitanus. A sample of Androctonus mauretanicus (Pocock, 1902) from Morocco and a sample of Androctonus crassicauda (Olivier, 1807) from Turkey were included as outgroups in the allozyme analysis. Centruroides exilicauda (Wood, 1863) from Baja California was the outgroup for DNA sequence analysis. Generally, the observed genetic variability of allozymes was low within populations. The UPGMA phenogram, based on Neifs D (1972), shows a low genetic differentiation among all Buthus samples from Morocco (inclu- ding B. atlantis). B. o. occitanus from Europe and B. o. tunetanus, however, clearly branch off at a high genetic distance. In contrast to the allozymes, the mtDNA data revealed more differentiation among the Moroccan samples, showing several lineages. The divergence among these lineages is about the same as between populations separated by the Strait of Gibraltar (approximately 12% sequence divergence). GAUTSCHI Barbara & TENZER Isabel Institut fiir Umweltwissenschaften; Universitat Zirich-Irchel; Winterthurerstrasse 190; CH- 8057 Ziirich; Switzerland & Institute of Plant Sciences; Pathology Group; Universitàtstrasse 2; ETH Zentrum; CH-8092 Ziirich; Switzerland ZOOLOGIA ET BOTANICA ‘99 761 Development of microsatellite markers for the Bearded Vulture (Gypaetus barbatus) and their application to the reintroduction project We describe the construction of a genomic library of G. barbatus enriched for the simple sequence repeats (CA)n and (GA)n, respectively. We found 29 different microsatellite repeats in 62 positive clones sequenced. So far, unique primer pairs for 16 of these 29 loci could be designed. Eleven primer pairs amplified in a locus specific manner. We describe the characteristics of these microsatellite markers and present preliminary results of polymorphism. The use of these newly developed microsatellite markers for the reintroduction project will be discussed. GEBHARDT-HENRICH SABINE Ethologische Station Hasli; Universität Bern; Wohlenstrasse 50a; CH-3032 Hinterkappelen; Switzerland Reproductive success was predicted by body mass loss in female great tits (Parus major) Body mass changes between incubation and when nestlings were 14 days old were measured in a great tit population breeding in artificial nestboxes. Half of the pairs were experimentally fed from nestbuilding until the hatching day of their young. Females lost 1.82 g (STD: 1.15, N = 67) between incubation and when their nestlings were 14 days old. Mass loss was not affected by experimental feeding. In control (unfed) females fledgling mass was significantly negatively correlated with body mass loss of the females. Under additional feeding this relationship tended to disappear. Fledgling mass is a predictor of fledgling survival in tit species and thus a measure of reproductive success. Fledgling number was not correlated with loss of body mass. Therefore, females losing less body mass during breeding had a higher current reproductive success then females with a greater loss of body mass. GERLOFF Christine U., OTTMER Birgit K. & SCHMID-HEMPEL Paul Experimental Ecology; ETH-Zentrum NW; CH-8092 Ziirich; Switzerland Inbreeding in bumble bees In small and isolated populations low levels of genetic variation may increase the likelihood of inbreeding, leading to inbreeding depression. In addition to more general negative effects, inbreeding in social hymenoptera leads to the production of diploid males. These males develop from eggs that should have produced workers or young queens but instead develop into diploid males because they are homozygous at the sex determining locus. The production of diploid males is very costly for the colony because it decreases the work force by 50%, and furthermore, the diploid males do not sire viable colonies. We studied the effect of inbreeding on the performance of colonies of the bumble bee Bombus terrestris (L.). We compared colonies of queens mated to their brother to those mated to an unrelated male (33 inbred and 44 outbred colonies out of 16 maternal families). We scored the number of workers and sexuals produced and calculated the caloric investment into sexuals to estimate fitness at the colony level. At the level of the individual we measured the efficiency of immune defences. The results show that inbreeding has no consistent negative effect but that the effect depends on the genetic background of the family and the colony. Diploid males had significantly lower immune defences than haploid males. Thus, when assessing the effects of inbreeding in a population, its genetic structure has to be considered. GOUDET Jérome Institut de Zoologie et d'Ecologie Animale; Batiment de Biologie; Université de Lausanne; CH-1015 Lausanne; Switzerland 762 ZOOLOGIA ET BOTANICA ‘99 A population genetics view of multivariate statistical methods While multivariate statistics are appealing to the population geneticist, they suffer from two drawbacks. The first is the lack of correspondence between inertia and a genetic parameter. The second is the absence of good testing procedure. Here I identify the relation between inertia and Fst, and suggest a test for both total inertia and the inertia of the different axes. The analyses are applied to both simulated and real data sets. Advantages and limits of multivariate methods are discussed. GOUINGUENE Sandrine, DEGEN Thomas & TURLINGS Ted Université de Neuchatel; Institut de Zoologie; Laboratoire d'Ecologie Animale et d'Entomo- logie; rue Emile-Argand 11; CH-2007 Neuchatel; Switzerland How parasitoids of caterpillars find suitable hosts ? Many parasitoids that attack phytophageous insects make use of plant odours to locate the habitat of their hosts. These odours are specifically emitted by a plant after it has been damaged by a herbivore, and not after mechanical damage. The odour emissions occur systemically throughout the plant. Factors in the oral secretion of the herbivores are the main elicitors of the plants’ reaction. One such elicitor, volicitin, was recently isolated and identified. The induced plant odours are useful cues for the parasitoids and indicate the presence of a potential host. Studies on these volatile plant signals have been conducted most extensively with maize plants and have led to the exact identification of the blend of compounds (mainly terpenoids) that is emitted after caterpillar attack. Our recent studies show that plants of a single maize variety consistently emit similar odour blends, but there can be considerable variation among different maize genotypes. This variation is also observed in the emissions of wild relatives of maize. We discuss these results in the context of the reliability of plant-induced signals as cues that allow parasitic wasps to find suitable hosts. GOVERDE Marcel, BAZIN Alain, SHYKOFF Jacqui & ERHARDT Andreas Institut für Natur-, Landschafts- und Umweltschutz; St. Johanns-Vorstadt 10; CH-4056 Basel: Switzerland Influence of leaf chemistry of Lotus corniculatus (Fabaceae) on larval development of Polyommatus icarus (Lepidoptera, Lycaenidae): effects of elevated CO, and plant genotype 1) Four Lotus corniculatus genotypes differing in their cyanoglycoside and condensed tannin concentrations were grown in either low (350 ppm) or high (700 ppm) atmospheric CO, environments. Larval performance, consumption and conversion efficiency of Polyommatus icarus feeding on this plant material was measured. 2) Plants grown under elevated CO, contained less cyanoglycosides, more condensed tannins and more starch than control plants. However, water content, nitrogen and protein as well as the nitrogen content in relation to carbon concentration did not differ between CO, treatments. 3) The four genotypes differed significantly in condensed tannins, cyanoglucoside, leaf water and leaf nitrogen but no genotype x CO, interaction was detected, except for total phenolics and condensed tannins in which two plant genotypes showed stronger increases under elevated CO, than the other two. 4) Larvae of P. icarus consumed more plant material and used and converted it more efficiently from plants grown at high atmospheric CO,. 5) Larvae developed significantly faster and were significantly heavier when fed plant material grown under elevated CO,. The observed difference in weight disappeared in the pupal and adult stages. However, lipid concentration of adults from the elevated CO, treatment was marginally significantly higher than of controls. 6) We conclude that the higher carbohydrate content of L. corniculatus plants grown at elevated CO, renders leaves more suitable and better digestible to P. icarus. Furthermore, differences in allelochemicals might influence the palatability of L. corniculatus leaves for this specialist on Fabaceae. ZOOLOGIA ET BOTANICA ‘99 763 GUGERLI Felix, SPERISEN Christoph, BUCHLER Urs & SENN Josef Swiss Federal Institute for Forest, Snow and Landscape Research; Ziircherstrasse 111; CH- 8903 Birmensdorf; Switzerland Genetic diversity in tree species of subalpine forests: a case study of Norway spruce (Picea abies) From a human perspective, one of the predominant tasks of forests in mountainous areas is to protect against natural hazards. Stability of mountain forest stands, however, is increasingly threatened by air pollution, cease of management, lack of regeneration due to overabundant ungulates, and novel environmental conditions because of climate change. The project ‘Biodiversity in Alpine Forest Ecosystems: Analysis, Protection and Management‘, funded by the European Union, addresses the genetic diversity of carrier species in Alpine forest ecosystems. The overall objective of the project is to create a common data pool which will help to better understand the function and dynamics of forest ecosystems and to transfer results into practice. At 14 locations across the Alps, each consisting of three elevational levels, populations of adult and juvenile forest tree species are sampled. The target species are Abies alba, Larix decidua, Picea abies, Pinus cembra, and P. mugo. Nuclear (isoenzymes, micro- satellites) as well as plastid markers (chloroplast microsatellites, mitochondrial tandem repeats) are investigated by respective expert groups. Our group so far screened 25 populations of Norway spruce (Picea abies (L.) Karst.) for a length polymorphism in the mitochondrial fragment of a nad/ intron. We found a decrease of within-population variation from eastern to western populations, which we relate to a serious bottleneck during re-colonization of the Alpine range by Norway spruce. GUSEWELL Sabine Geobotanisches Institut ETH; Gladbachstrasse 114; CH-8044 Ziirich; Switzerland Does the public support restoration measures in fen meadows invaded by common reed ? Public support is indispensable to the long-term success of measures for nature conservation and restoration. Such measures are more likely to be approved if their aims are understandable and if they are considered useful by the general public; aesthetical aspects might be particularly important in strongly frequented areas. This study investigated whether the invasion of fen meadows by common reed (Phragmites australis) is perceived as a degradation by the general public and whether measures to restore lower, more open vegetation types are supported. Quali- tative interviews were carried out in the field with visitors of a wetland nature reserve (Roben- hauser Ried/ZH); moreover, students of various disciplines answered a questionnaire based on colour photographs. The field survey revealed no general preference for sites with either a high or a low abundance of Phragmites. In the questionnaire survey, the tendency to prefer fen meadows with a low abundance of P. australis differed among disciplines and increased with the (presumable) knowledge of students about nature conservation. Likewise, the acceptance of restoration measures appeared to mainly depend on ecological knowledge. Both surveys suggested that people reject restoration measures (regardless of their aesthetical evaluation) if they can not conceive (a) that fen meadows are semi-natural areas needing management, (b) that the spread of P. australis might be a consequence of human impacts and (c) that changes in management can be necessary to maintain a status quo. More information about these points would probably enhance public support to nature conservation and restoration. HAGGSTRÖM Hakan!, RAHIER Martine! & HARTMANN Thomas? ! Laboratoire d'Ecologie Animale et Entomologie; Institut de Zoologie; Université de Neu- chatel; rue Emile-Argand 11; CH-2007 Neuchatel, Switzerland * Institut für Pharmazeutische Biologie der Technischen Universität Braunschweig; D-38106 Braunschweig; Germany 764 ZOOLOGIA ET BOTANICA ‘99 Deterrent effect of pyrrolizidine alkaloids and detoxification by a generalist predator (Mitopus morio) Several leaf beetles in the genus Oreina sequester pyrrolizidine alkaloid N-oxides (PAs) from their host plants (e.g. Adenostyles spp., Asteraceae). We studied the efficiency of these compounds as defence against a harvestman (Mitopus morio, Phalangidae) which is common in habitats where Oreina spp. occur. M. morio was observed to feed on larvae of O. speciosissima and O. cacaliae both in the field and in a laboratory experiment. Two populations of M. morio were compared with respect to the deterrent effect of PAs and the efficiency of detoxification. One of the populations comes from a site dominated by Adenostyles alliariae. These indivi- duals are likely to have come into contact with prey containing PAs. The other population comes from a site where Adenostyles spp. are absent and is therefore likely to have little experience of PAs. For the test of deterrence, the individuals were offered artificial food (minced meat) containing the PA senecionine N-oxide in different concentrations. The effi- ciency of PA detoxification was studied in a tracer feeding experiment. The two populations differed in both behaviour and detoxification ability. Individuals from the population lacking Adenostyles plants were more deterred by the alkaloid compared to those from the other site. The metabolic treatment of PAs differed between the populations, but both populations had efficient ways to eliminate PAs and did not sequester these compounds for their own defence. We suggest that the efficiency of PAs as defence for sequestering herbivores may differ between populations. HEEB Philipp Centre for Behaviour and Evolution; Hasli Ethol. Station; University of Bern; CH-3032 Hinter- kappelen; Switzerland Within-pair copulation frequency predicts female reproductive performance in tree sparrows, Passer montanus Female birds can solicit and accept copulations from males and are thus able to control their copulation frequency. Engaging in copulations is expected to be costly since copulations take up time and can increase the risks of predation and pathogen transmission. Given these costs, the high frequency of within-pair copulations shown by many monogamous birds remains puzzling. A number of hypotheses have been proposed to explain the benefits that female birds obtain from frequent copulations with their male. These non-exclusive hypotheses predict that high within-pair copulation frequencies should be positively associated with components of female reproductive performance. In agreement with this prediction results from a tree sparrows population show that within-pair copulation frequency was positively correlated with the number of eggs laid and the number of young raised. The results suggest that, in tree sparrows, within-pair copulation frequency can be considered as an early expression of the pairs’ reproductive ability. The possibility that within-pair copulation frequency could function as a signal for assessment of phenotypic quality by both males and females is discussed. HELLRIEGEL Barbara! & REYER Heinz-Ulrich? ! Zoological Museum; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland 2 Zoological Institute; University of Zurich: Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Which factors influence the composition of mixed populations of a hemiclonal hybrid (Rana esculenta) and its sexual host ? Hemiclonal hybrids combine advantages of sexual and asexual reproduction, but their need for backcrossing tightly couples their population dynamics with that of their sexual hosts. Two discrete-time models are constructed to identify important factors influencing the composition and dynamics of the resulting mixed populations. Hybridogenetic waterfrogs serve as an ZOOLOGIA ET BOTANICA ‘99 765 example. The results can explain the remarkable stability over time of observed species ratios within hybridogenetic populations, even when population sizes fluctuate. They also make suggestions why species ratios vary so widely between populations: the precondition is that either one or several of the four factors mate choice, relative fecundities, (larval) competitive abilities, dispersal and their interactions differ among localities. While differences in fecundity influence species coexistence, larval superiority seems to be more important for the hybrids spread in mixed populations; but differences between host and hybrid are not necessary for their coexistence. Dispersal affects the species composition mainly in the habitat where host and hybrid are competitively equal, but ecological and reproductive dispersal have opposing effects. These results suggest interesting consequences for other tightly coupled systems. HERGER Franziska, LARGIADER Carlo R., LURTSCHER Mathias & SCHOLL Adolf Division of Population Biology; Institute of Zoology; University of Bern; Baltzerstrasse 3; CH-3012 Bern; Switzerland The Austropotamobius pallipes species complex in the Alpine region - phylogeo- graphic studies using nuclear and mitochondrial markers In a joint analysis of nuclear (allozyme) and mitochondrial markers (sequence and RFLP analysis) five genetically distinct groups of the Austropotamobius pallipes (Lereboullet 1858) species complex were detected in the Alpine region. The geographic distribution of these evolutionary lineages coincided largely with several taxa (A. p. pallipes, A. berndhauseri, A. p. italicus) defined on the basis of morphological characters. A low level of genetic variability was found within these lineages. In some cases, the combination of the two marker classes allowed to assess the natural or artificial origin of populations. The interlocking distribution of the five evolutionary lineages indicated that conservation efforts must aim at the level of local populations. HEYLAND Andreas Zoological Museum; University of Zurich; Winterthurerstrasse 190; CH-8057 Ziirich; Switzer- land Behaviour and mutual influencing of two polychaete species with different life-cycles during settlement: Poecilochaetus serpens E. Claparede and Sphaerosyllis sp. (Poly- chaeta: Syllidae: Exogoninae) The influence of a small interstitial polychaete (Sphaerosyllis sp.) on the settlement behaviour of a polychaete with a benthoplanktonic life-cycle (P. serpens) was examined in a series of laboratory experiments. Sphaerosyllis sp. was found exclusively in the first cm of the sediment and was among the most abundant species at this site, with an average of around 11,000 animals per m2. From direct observations of the settlement process of P. serpens with a digital camera-system it was found that Sphaerosyllis sp. disturbed the settlement behaviour of P. serpens planktonic larvae that were in an advanced developmental stage. However this in- fluence was not strong enough to prevent P. serpens larvae from settling. P. serpens settled immediately on the sediment but a decision to leave the substratum could be made after the animal was partially burrowed. Possible reasons and consequences of this behaviour are discussed. The nearest neighbour distances between settling planktonic P. serpens larvae were measured under laboratory conditions. No avoidance behaviour was observed between these larvae, which might explain the patchy distribution of P. serpens in the field. Horer Ulrich!, BERSIER Louis-Felix? & BORCARD Daniel? ! Department of Vertebrates; Natural History Museum; Bernastrasse 15; CH-3005 Bern; Switzerland 166 ZOOLOGIA ET BOTANICA ‘99 ? Zoological Institute; University of Neuchatel; rue Emile-Argand 11; CH-2007 Neuchâtel: Switzerland 3 Département de sciences biologiques; Université de Montréal; C.P. 6128; Montréal, Québec H3C 3J7; Canada Spatial organization of a tropical forest herpetofauna on an elevational gradient revealed by null model tests Five null model tests were applied to the herpetofaunal assemblage on the western slope of Mount Kupe, Cameroon. Based on the pattern of species range boundaries and abundances along the primary forest elevational gradient, ranging from 900 to 2,000 m, the relative impor- tance of interspecific competition and ecotones in structuring the assemblage was assessed. Tests were run for 1) all species, 2) amphibians, 3) reptiles, 4) amphibians dependent on streams for reproduction, and 5) amphibians that don't use streams for reproduction. For three null models, the observed patterns do not differ from random expectations. The results indicate that there are very few species whose gradient distributions may be limited by interspecific competition between congeners. Significant discontinuities in abundance patterns and range boundary dispersion revealed zonations in all subsets analyzed, but neither indicate distinct species groups with sharp exclusion boundaries nor a strong response to vegetational ecotones. Physical factors varying in parallel with the gradient and specific habitat components, particu- larly waterbodies suitable as amphibian breeding sites, are suggested to be the dominant factors in limiting gradient distributions of amphibians and reptiles on Mount Kupe. The zonations revealed suggest a pattern of three spatially non-exclusive species groups: physical factors separate distinct lowland and montane species limited by physiological constraints and produce the faunal discontinuities in the lower submontane forest around 1,300 m; this boundary 1s encompassed by the range of a group of anuran species, whose distributions on the gradient are centred at intermediate elevation, and appear to be limited by specific habitat requirements. The response to predominantly abiotic factors suggests a basic difference to endotherms, where biotic factors seem to be of major importance in limiting elevational distributions. ILG Christiane, CASTELLA Emmanuel & Lops-CROZET Brigitte Laboratoire d'Ecologie et de Biologie Aquatique; Université de Genève: chemin des Clochettes 18; CH-1206 Geneve; Switzerland Macroinvertebrate biodiversity in glacial catchment: the rule of water sources and tributaries Alpine streams remain relatively understudied, regarded as sheltering poor zoobenthic commu- nities, mainly made up of Diamesa (Diptera, Chironomidae). If this is true for glacial rivers, at the snout of the glacier, it is not valid for rhithral or krenal segments, snowmelt- or ground- water-fed, that allow relatively diverse communities to develop. The study presented here, illustrates the case of the Mutt (Valais, Swiss Alps). The main channel is typically kryal, fed by glacial meltwater. Its tributaries are glacial outlets or krenal stream ecosystems. These ground- water fed streams show a broad scale of physico-chemical characteristics depending on altitude, geological substratum or season. There is a very strong correlation between environmental variables (temperature, conductivity, turbidity and discharge) and the composition of the ma- croinvertebrate communities. In the glacial brooks harsh conditions prevail: low water tempe- rature, high turbidity, large diel discharge fluctuations. Never more than 5 taxa, essentially chironomids, were found in each station. On the other hand, the krenal tributaries are inhabited by highly diversified macroinvertebrate communities (up to 20 taxa at one station). These species, most of them belonging to various families of Plecoptera, Ephemeroptera, Trichoptera, Diptera, prevail in rheocrene habitats, characterised by high physico-chemical stability. The habitat heterogeneity, generated by the different water origins, favours the coexistence in a rather small catchment area of diversified macroinvertebrate communities, and plays therefore an important part in the biodiversity of the Mutt basin. ZOOLOGIA ET BOTANICA ‘99 767 JENSEN Deborah B. Conservation Science Division; The Nature Conservancy; 1815 N. Lynn Street; Arlington, VA 22209; USA Conservation by design: the theory and practice of conservation priority setting One key challenge in biodiversity conservation is deciding what actions must be taken today, and which can be put off until tomorrow. Not all species nor all places are equally at risk. Over the past two decades conservation biologists have developed practical but rigorous methods for setting priorities for species, sites and even countries or regions. While most agree that bio- diversity value and threat are important criteria for priority setting, disagreements on methods of selecting conservation sites abound. This lecture will review the main approaches for conservation priority setting and argue for the systematic design of reserve networks. Current examples of efforts to design reserve networks will be taken from The Nature Conservancy's work in North and South America. KALBERER Nicole & RAHIER Martine Institut de Zoologie; Université de Neuchatel; rue Emile-Argand 11; CH-2007 Neuchatel; Switzerland i The cost of dispersal by flight in an alpine leaf beetle Oreina cacaliae (Coleoptera: Chrysomelidae) One part of the population of Oreina cacaliae leaves the host plant patches in autumn and flies to overwintering places away from the host plant. The other part of the population overwinters next to the host plant and does not fly. Flying females emerge earlier in the season and start reproducing earlier than their nonflying conspecifics. In 1997 the flyers took advantage of an early start and the fecundity was not significantly reduced compared to the nonflyers although the flyers died earlier. Due to the extremely high mortality in the flyer group in 199$ fecundity was significantly lower compared to the nonflyers. The cost of dispersal by flight seems to be the higher risk of mortality compared to beetles that overwinter next to the host plant and do not engage in flight. KELLER Michael, KOLLMANN Johannes & EDWARDS Peter J. Geobotanisches Institut; ETH; Zürichbergstrasse 38; CH-8044 Zürich; Switzerland Genetic introgression from distant provenances reduces fitness in local weed popu- lations - a case study with species for ecological compensation areas Various seed mixtures have been suggested for ecological compensation areas in intensively used farmland. Seed mixtures which contain foreign seed provenances may reduce fitness in local populations due to genetic introgression. 2) This hypothesis was tested with the weed species Agrostemma githago, Papaver rhoeas and Silene alba. Hybrids (F2 backcrosses) between local Swiss plants and English, French, German and Hungarian provenances (plus one French Papaver and one US source in Silene) were tested in the field for effects on fecundity. Aboveground biomass after one growing season was taken as a measure for fitness under natural environmental conditions. In some cases, survivorship in the field assay and seed mass were determined. The results were compared with the parents, with the Fl and with crossings among other Swiss provenances. 3) Negative outbreeding effects on fecundity were found in all four hybrids of Papaver and in the F2 of Agrostemma with introgressions by the German provenance, but not in Silene. Mortality was slightly higher in the Fl of Papaver, and even more in the F2, whereas it was unaffected in the Silene hybrids. Seed mass of Agrostemma and Silene decreased in the F2 compared with the F1. 4) The results are discussed with respect to the use of foreign provenances on agricultural compensation areas and in the context of re- enforcement of endangered plant populations. The relevance of the measured fitness compo- nents, and long-term effects of genetic introgressions are addressed as well. 768 ZOOLOGIA ET BOTANICA ‘99 KERY Marc Institut für Umweltnaturwissenschaften; Uni Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Inferring the absence or extinction of a species from a site: a case study with three snake species Many animal species are difficult to see and even during visits to known occupied sites no specimen may be detected. For such species local extinction or absence from a site may be difficult to ascertain. However, if we know with what probability a species is detected at any one visit to an occupied site, the number of times that a site has to be visited unsuccessfully, before it can be presumed unoccupied, can be calculated. The probability of detection can be estimated from visits to sites known to be occupied. From 1994 to 1997 I studied the probability of detection in three snake species, Vipera aspis, Coronella austriaca and Natrix natrix at 88 sites in the Jura mountains. In all species, probability of detection was higher in larger populations. There was no effect of the type of habitat nor of the size of a site. Overall, there was a 38%, 13%, and 21% chance per visit to see at least one specimen of V. aspis, C. austriaca and N. natrix, respectively. This translates into 7, 23 and 13 unsuccessful visits, respectively, that are necessary before a site can be assumed unoccupied. Sites whose occupancy status is unknown are most likely occupied by small populations. For them, a minimum of 15, 28, and 35 visits are necessary in V. aspis, C. austriaca and N. natrix, respectively. Hence, to infer the absence of these species, a much larger search effort than usually applied may be necessary. Furthermore, some of the species may be more widespread than hitherto thought. KNADEN Markus Zoologisches Institut; Universität; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Enemy recognition in the desert ant Cataglyphis fortis Twenty ants of colony A were given into individual encounters with each 10 ants of colony B and C. A mean aggression level was calculated. After these fights 10 ants of colony C were put into colony A where they were killed immediately. In the next day the individual encounters between ants of colony A and B as well as C were repeated. There was an increased aggression level of colony A against C, but not against B. Cataglyphis ants seem to be able to recognize ants which have been intruders in the past. KOLLIKER Mathias Zoology Department; University of Bern; CH-3032 Hinterkappelen; Switzerland Heritability of offspring begging and its co-evolutionary potential with parental feeding behaviour Models of parent-offspring conflict predict that the differing evolutionary interests of genes expressed in parents and their dependent offspring lead to an arms race between offspring begging and parental feeding behaviour. In theory, the evolutionary stable resolution varies depending on 1) mating system, 2) offspring relatedness, 3) fitness cost of begging to offspring and 4) fitness cost of parental effort to parents. In the great tit (Parus major), male and female parents feed their young to variable relative amounts, offspring relatedness varies among families due to extra-pair paternity and parents seem to make individual trade-off decisions between current and future reproduction. Such within-population variation may lead to the maintenance of a range of conflict resolutions, rather than one single outcome. We investigated this hypothesis using a cross-fostering experiment where sibships were split and transferred in different nests of unrelated parents. The vocal begging performance of individual nestlings was measured under controlled laboratory conditions. The similarity in vocal begging intensity of ZOOLOGIA ET BOTANICA ‘99 769 siblings growing in different nests was used as an estimate for its heritability. We tested parental response to begging calls using begging-playback experiments. The broad-sense heri- tability of vocal begging intensity, and the covariation between the biological parent's respon- siveness and the offspring's begging behaviour were estimated. Information about heritable variances and covariances in parent-offspring communication may be necessary to understand its evolutionary potential. KOLNAAR Rogier Institut de Biologie végétale; Université de Fribourg; rue Albert Gockel 3; CH-1700 Fribourg; Switzerland Effect of Puccinia lagenophorae on competition between Senecio vulgaris and Cap- sella bursa-pastoris The annual plants Senecio vulgaris L. and Capsella bursa-pastoris (L.) Medic. belong to similar communities occupying ruderal habitats. The host-specific rust fungus Puccinia lagenophorae Cooke infects S. vulgaris and infection alters the outcome of competition between S. vulgaris and C. bursa-pastoris. Depending on time of infection, competition is more or less affected. Epidemics of P. lagenophorae develop and may infect S. vulgaris plants within a population at various times. Can the development of P. lagenophorae epidemics be predicted, and thus, the effect of P. lagenophorae on plant competition at the population level ? In the study presented the effect of temperature on epidemiological parameters of S. vulgaris on P. lagenophorae was determined. Plants were grown at different temperatures in two experiments. Incidence, latent period and severity were assessed. In one of the experiments, infectious period and spore production were assessed. Velocity of epidemic spread was estimated for plants grown at 10, 16 and 22°C. A temperature effect on latent period was detected in both experi- ments, whereas a temperature effect on incidence and severity was detected in only one experiment. Latent period increased by decreasing temperature. Spore production curves were fitted using the gamma function. A temperature effect was detected on infectious period and spore production. Data obtained in the study presented were used to estimate velocity of epidemic spread. A temperature effect on velocity was detected. Velocity of epidemic spread was highest on plants grown at 22°C and lowest on plants grown at 10°C. KUENZLER Reto & BAKKER Theo C.M. Abt. Verhaltensoekologie; Zoologisches Institut; University of Bern; Wohlenstrasse 50a; CH-3032 Hinterkappelen; Switzerland A study of multiple female preferences in sticklebacks using computer animations The study of multiple mate preferences is handicapped by the impossibility of experimentally manipulating particular traits let alone combinations of traits. Previously used methods such as direct or indirect tests with live animals and video playback techniques can often not get round confounding variables and suffer from non-standardised variance within and between repli- cates. Choice tests using reality-based computer animations offer a solution because they feature fully controllable virtual animals in a defined surrounding. A 3D-model of a male stickleback was built based on the dimensions and proportions of an average male and its courtship movements were animated. Females were allowed to choose between two virtual males which only differed in the desired ornament(s) and that were presented simultaneously on a high-quality computer display. The female's attention towards animated males correlated positively with their attention towards a real courting male. First results of preference tests will be presented. KUSKE Stefan Federal Research Station for Agroecology and Agriculture; CH-8046 Ziirich; Switzerland 770 ZOOLOGIA ET BOTANICA ‘99 Does the mass release of the exotic egg parasitoid Trichogramma brassicae pose a risk to populations of endemic natural enemies ? Lydella thompsoni Hert. (Diptera, Tachinidae) is a native larval parasitoid of the European Corn Borer and shows parasitism rates up to 50% in Ticino. The first generation of the tachinid develops on hosts in pristine habitats, such as Archanara spp. (Lepidoptera, Noctuidae), while the European Corn Borer is the main host for the subsequent generations. Maize seed producers carry out mass releases of the exotic egg parasitoid Trichogramma brassicae Bedz. (Hyme- noptera, Chalcidoidae) to control the European Corn Borer, which is the key pest of maize in Ticino. These releases coincides with the oviposition period of known spring hosts of L. thompsoni. Preliminary studies have shown that a high percentage of T. brassicae can disperse from maize fields into pristine habitats in the surroundings where they search and might para- sitize alternative hosts, such as Archanara spp. The present study investigates the importance of the exotic egg parasitoid 7. brassicae in displacement processes of the endemic competitor L. thompson. LAWTON John H. NERC Centre for Population Biology; Imperial College; Silwood Park; Ascot, Berks SL5 7PY; United Kingdom Biodiversity and ecosystem function Field and laboratory experiments with model plant communities show clear evidence of a reduction in ecosystem processes (for instance productivity and nutrient cycling) with declining plant species richness within any one site. Comparisons made across different sites, however, yield more complex relationships, so that mixing within- and between-site effects is confusing. A robust body of emerging theory underpins and explains these empirical results. This talk will start by reviewing the theory, before briefly summarising the empirical data. I will then go on to present preliminary results from the European-wide BIODEPTH experiment to illustrate the key points. BIODEPTH has simulated the loss of plant species from model, grass/herb commu- nities using identical experimental protocols at eight sites across Europe. The implications of these data and theory for conservation, in the face of the rapid loss of species from many surviving natural and semi-natural habitats, and for more intensely managed ecosystems and landscapes will be discussed. The remaining scientific challenges are: (1) To understand better the mechanism(s) underpinning ecosystem responses to loss of biodiversity. (11) To distinguish between the role of biodiversity in maintaining ecosystem processes under constant, or benign environmental conditions, and under extreme environmental perturbations. The "insurance hypothesis" predicts that many apparently "redundant species" are essential for maintaining ecosystems under extreme events. (111) To start to consider the role of animal and microbial diversity in ecosystem processes. LUGON-MOULIN Nicolas, BRUNNER Harald, BALLOUX Francois, GOUDET Jerome & HAUSSER Jacques Institut d'Ecologie; Laboratoire de Zoologie et Ecologie Animale; Bätiment de Biologie; Uni- versité de Lausanne; CH-1015 Lausanne-Dorigny; Switzerland Do riverine barriers, history or introgression shape the genetic structuring of a common shrew (Sorex araneus) population ? The common shrew (Sorex araneus) is subdivided into numerous chromosome races. The Valais and Cordon chromosome races meet and hybridize at a mountain river in Les Houches (French Alps). A significant genetic structuring was recently reported among populations found on the Valais side of this hybrid zone. In this paper, a phylogenetic analysis and partial Mantel tests are used to investigate the patterns and causes of this structuring. A total of 185 shrews ZOOLOGIA ET BOTANICA ‘99 771 were trapped at 12 localities. All individuals were typed for nine microsatellite loci. Although various mountain rivers are found in the study area, riverine barriers are not found to have a significant influence on gene flow. Partial Mantel tests show that our result is due to the influence of the hybrid zone with the Cordon race. The geographic patterns of this structuring are discussed in the context of the contact zone, which appears to extend up to a group of two rivers. The glacier they originate from is known to have cut the Arve valley as recently as 1818. The recent history of this glacier, its moraine and possibly rivers, may therefore be linked to the history of this hybrid zone. MACDONALD David Zoology Department; South Parks Road; Oxford; Oxon OX1 3PS; United Kingdom Alien invaders Introduced alien species are a major factor in conservation. In particular, the American mink has been widely introduced around the world. This has revealed not only major problems for conservation, but also theoretically interesting insights in ecological theory. In Britain, for example, the American mink is largely responsible for a radical decline in the population of water voles. In Eastern Europe the American mink is responsible for the collapse in populations of European mink. These results, and their wider relevance, will be discussed. MATTHIES Diethart!, BRAUER Ingo”, MAIBOM Wiebke? & TSCHARNTKE Teja” ! Institut für Umweltwissenschaften; Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland ? Fachgebiet Agrarökologie; Universität Göttingen; Waldweg 26; D-37073 Gottingen; Ger- many Population size and risk of extinction in rare plants In the mid 1980s the distribution and size of populations of endangered plant species in Lower Saxony (N-Germany) were recorded by the conservation authorities. We used these data as the basis for a study of the effects of population size on the probability of survival of local popu- lations in eight rare plant species (Gentianella germanica, G. ciliata, Lepidium campestre, Melampyrum arvense, M. nemorosum, Rhinanthus minor, R. serotinus and Thlaspi perfolia- tum). In 1996 we visited 359 known sites of these plants and recorded whether they still had extant populations. The risk of extinction of local populations was very high. Of the studied populations 28% had become extinct in the 10 years since the last census. Logistic regressions revealed significant negative relationships between population size and risk of extinction in seven of the eight studied species. The form of the relationship and the population size required for 90% probability of survival over 10 years varied strongly among species, suggesting that minimum population sizes required for persistence will vary among plant species. MICKASCH Tatjana M. Zoological Museum of the University of Zurich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Is there a relationship between expanding Orconectes mosus and decreasing Austro- potamobius pallipes populations in Switzerland ? The white-clawed crayfish (Austropotamobius pallipes) is considered an endangered species in Switzerland. It is native to the waterbodies of the canton of Berne, Waadt and Wallis, where the American species Orconectes limosus occurs too. The first recording of O. limosus was 1979 in Lake Biel. Eversince, it expanded its range continuously, and early in 1998 it was found in the rıver Aare 90 km downstream of Lake Biel. Because of its resistance to crayfish plague (Aphanomyces astaci) it is a potential threat to the white-clawed crayfish. The aim of this TAD ZOOLOGIA ET BOTANICA ‘99 poster is to assess the actual and potential threat of A. pallipes by O. limosus. In summer and fall of 1997, 63 streams were checked and O. limosus was found in 17 out of 43 streams draining either into Lake Biel or the river Aare, but not in the remaining streams. A. pallipes was found only in 5 out of 20 streams where the species had been recorded since 1990 by local authorities. O. limosus was restricted to the immediate surroundings of the rivermouths. The probability of his occurrence was the higher, the wider the stream was and the denser it was in the river or the lake. There is no obvious connection between the decline of native white- clawed crayfish populations and the expansion range of O. limosus. Alternative explanations of the decline of the native species are either chemically or organically polluted water entering the streams more or less continuously, or short intervals of high phosphate and nitrate concentrations after heavy rainfall. Mikos Z. Maja & WARD Paul I. Zoologisches Museum der Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich, Switzerland Cryptic female choice of male genotype in the yellow dung fly (Scathophaga ster- coraria) Sperm competition in the yellow dung fly, (Scathophaga stercoraria), has been examined in great detail. -Strong male-biased size dimorphism allows male behavioural control over copulation and fertilisation. Recent investigations point to internal cryptic control over fertilisation by the female. Females were mated to two males, with either a few hours or just over a day between the matings. All flies came from two different lines (Pgm-2, Pgm-3) selected for different alleles of the Pgm locus. I tested for effects of the male's genotype on copulation duration, latency and fertilisation success. The proportion of offspring fathered by the second male (P2) was scored using the Pgm marker. Results indicate that both the male's genotype and his genetic similarity to the female's genotype affected paternity. Paternity of males from Pgm-2 line was overall higher but females at the same time preferred males of a genotype different from her own. Males homozygous for allele 2 (Pgm-2) copulated longer and their latency to copulation was shorter. The experiment showed that female S. stercoraria uses cryptic choice to control paternity to overcome her lack of behavioural control. Moretti Marco!, HORDEGEN Philipp’, CONEDERA Marco!, DUELLI Peter? & Epwarops Peter J.° ! Swiss Federal Institute for Forest, Snow and Landscape Research; Station South of the Alps; _ CH-6504 Bellinzona; Switzerland 2 Swiss Federal Institute for Forest, Snow and Landscape Research; CH-8903 Birmensdorf/ZH; Switzerland 3 Geobotanical Institute; Swiss Federal Institute of Technology: Zürichbergstrasse 38; CH-8044 Zürich; Switzerland Response of invertebrates to wildfires. First results about spiders (Araneae) and ground beetles (Carabidae) in deciduous forests on the south part of the Alps (Ticino, Switzerland) Current knowledge on the consequences of wildfire for faunistic biodiversity is fragmentary, particularly in deciduous forests south of the Alps, which is the region of Switzerland most affected by this phenomenon. After the second year of a doctoral thesis about "The influence of sporadic and regular fires on invertebrate biodiversity” planned for three years, we present some preliminary results. The aim of the project is to explore the effect of single and regular fires on the invertebrate fauna. The investigation is taking place in a hilly region in the chestnut belt (Castanea sativa forest). The study area is a south facing slope. We have sampled several invertebrate groups in 23 test areas: 7 areas affected by a single fire, 10 areas with repeated ZOOLOGIA ET BOTANICA ‘99 773 fires and 6 control areas of intact forest. The soil fauna have been assessed using several stan- dardised sampling methods: pitfall traps for epigeic arthropods, litter-soil samples and soil eclectors for soil dwelling groups, and yellow window traps for flying insects. Sampling period between January and October 1997. The first results show that among spiders and carabid beetles species react differently to fire. Some species of spider are already present a few days after the fire; however, other species seem to avoid burned areas. The highest number of species of spiders was recorded in 2 years old burned areas. The number of species of carabid beetles appears to be higher in forest areas which have burned frequently. Some first hypo- theses based on these preliminary results are presented. MULLER Jürg P. Bündner Natur-Museum; Masanerstrasse 31; CH-7000 Chur; Switzerland The reintroduction of the Bearded Vulture, Gypaetus barbatus, in the Alps — an example for the practical project realisation Due to a large-scale reintroduction project the first Bearded Vulture, Gypaetus barbatus, rele- ased into the wild are breeding again in the Alps, 100 years after their extinction. The project, which has been successful up to now, has the following milestones: the development of a bree- ding program, the choice of a suitable means of release, the choice of four release sites in the Alps as well as the establishment of a monitoring program. Of great importance was the setting up of the breeding network and the decision not to take any animals from the endangered populations. Therefore, only a breeding program with about 50 birds, already in captivity and belonging to the Eurasian sub-species Gypaetus barbatus bar- batus could be considered. Breeding stations were established in Vienna, Haute-Savoie and Goldau. For the breeding program additional pairs are kept in different zoos. Today the breeding program functions very well. About 150 bird were brought up in captivity. 50% were set free, 50% were taken into the breeding stock. A further decisive step was the choice of the method for release. Using the so-called "Hacking method", the birds are brought at an age of about 100 days without parents into an artificial nesting site. At that time, they are not able to fly, but start flying usually 20 days later. During this period the birds get a strong connection to the release site: a behaviour also called philopatry. The choice of 4 release sites, distributed at regular intervals across the Alps, has a strong bea- ring on the release method. It is expected that the birds, because of their philopatric behaviour will, in the majority of cases, stay in the range of their release area and preferably hatch there. In this way, nuclei of populations are created from which a whole population can build up in the Alps. The next step was to drawn up a monitoring project: Monitoring is understood as the compari- son of the target situation, i.e. a self-sustaining population in the Alps, with the present situation at a certain point in time. As long as these situations differ from each other, the project should be continued. In detail, monitoring also serves to discover wrong developments and therefore wrong decisions. So the monitoring program encompasses a whole range of scientific projects. At the present time following scientific programs are being carried out in the Swiss project: The spread of the birds over the Alps and their survival is monitored with the help of amateur field observers such as ornithologists and game guards, reporting their sightings to the project data bank. The development of the young at the release sites and the formation of pairs is followed up by zoologists, charged by the reintroduction project. The fact of small populations both in the breeding network and in the wild, require a genetic management plan. The basic study is carried out by Barbara Gautschi from Zurich University. I would be wrong to believe that the monitoring program allows us a very precise regulation of the project. If the breeding methods were unsuitable and the birds would not hatch in the wild then one would have to rethink the whole project entirely. Monitoring also has an important 774 ZOOLOGIA ET BOTANICA ‘99 documentary and scientific value. In this way one can follow the establishment of a population in a previously uninhabited area. Up to now 80 birds have been released, of which about 50 survived. Breeding of released birds took place in Haute-Savoie (1997 and 1998) and in the Stelvio National Park (1998). There, the birds obviously have started breeding again; so that there is much hope they will reproduce also this year. In a reintroduction project spanning the whole region of the Alps and crossing country borders, great observance had to be given to the internal flow of information. An own bulletin, annual meetings and workshops serve to train the people involved. The public relation work proves an extremely important feature. It is prerequisite for the acceptance of the project by the popu- lation and the authorities (permission) and for successful funding. Contributions to television, radio and printed media are decisive to the project becoming well known. Books, brochures and exhibitions have a long term educational effect while lectures are suited to winning over local interest groups (local population, hunters, authorities, ornithologists) and to discussing criticism of the project. A difficult exercise in managing the project besides the specific expertise problems is the lea- dership of the many different persons involved. Many take part for very different reasons and therefore have very different expectations of the project. In order to carry out this sort of project and to make the necessary decisions, information of various types are needed: practical knowledge about keeping animals, results of specially ini- tiated basic research, studies with the characteristics of environmental impact assessments, extensive and intensive monitoring, project management and public relations. Zoologists who want to take part in such projects need more and better-organised opportunities for practical training and for further education in addition to their purely scientific field. OERTLI Beat, AUDERSET- JOYE Dominique, JUGE Raphaelle, CASTELLA Emmanuel, MULLER Jessica, ANTOINE Céline, SAAM Mirko, CAMBIN Diana & LACHAVANNE Jean-Bernard Laboratoire d'Ecologie et de Biologie aquatique; University of Geneva; chemin des Clochettes 18; CH-1206 Geneve; Switzerland Biodiversity in Swiss ponds: how biotic communities respond to altitude This large-scale project supported by the Swiss Federal Office for Environment, Forest and Landscape is an investigation of environmental conditions and biodiversity over a wide range of small waterbodies. It aims at: (1) categorising waterbodies and their associated plant and invertebrate diversity, (2) deriving predictive relationships between a set of environmental characteristics of the ponds (structure of the environment, water quality ...) and their biotic rich- ness and composition, (3) setting up a standardised method for the assessment of the conser- vation value of ponds. The first results demonstrate the preponderance of altitude in governing diversity of Swiss ponds and a quantification of this tendency is proposed. Altitude also plays a key rule in the typology of the ponds. OLIVIERI Isabelle Institut des Sciences de |'Evolution; Université Montpellier 2; place Eugène Bataillon; F-34095 Montpellier cedex 05; France Contribution of metapopulation-level studies to the understanding of extinction, adap- tation, and speciation The evolutionary potential of a species is of primary concern in conservation biology. Such evolutionary potential may be described by the genetic and phenotypic diversity of a species, its colonising ability, as well as by its propensity to locally adapt and eventually speciate. Clearly, studies above the population-level are a necessary step towards an understanding of what ZOOLOGIA ET BOTANICA ‘99 775 determines a species niche breadth. In this talk, I will describe how the evolution of dispersal might affect the evolutionary potential of a species, by taking examples from widespread and endemic plant species. I will then describe how the evolution of dispersal and habitat selection through host choice can lead to local adaptation and eventually speciation in herbivorous insects. Methods used range from mathematical models to demographic surveys of natural populations, through population and quantitative genetics studies. PASCHKE Melanie Institut für Umweltwissenschaften; Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Does genetic variation matter ? The theoretical background of loss of genetic variation in small populations of plant species is well discussed. Reduced genetic variation is thought to decrease the future capability of a popu- lation to adapt to environmental changes. However, theoretical predictions so far have not been critically evaluated in the field. It is still unclear if under stressful environmental conditions genetic variation in itself is more important for the survival of a population than the phenotypic plasticity of the individuals. The hexaploid Cochlearia bavarica (Brassicaceae, Vogt 1985) is endemic to Bavaria and highly endangered. It is a perennial, self-incompatible plant. In a previous study a reduced genetic variation and reproduction was observed in small populations of the species. In the present study I now ask the following questions: 1) Is high genetic varia- tion within populations important for plant performance in the field and is this relationship independent of population size ? 2) Is high genetic variation within populations important for the coping with environmental stress ? I carried out two experiments to address these questions: transplantation experiment: performance of plants in own/foreign environments. PASINELLI Gilberto Zoological Museum and Institute; University of Zurich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Effects of habitat structure on home range size and breeding success of the middle spotted woodpecker Dendrocopos medius The endangered middle spotted woodpecker is restricted to old oak forests, which are rapidly disappearing due to inappropriate forest management. The relationships between habitat factors, home range size and breeding success of this species was investigated during a five year study based on 33 radio-tracked birds. Multivariate analyses revealed that home range size was inversely related to a) the density of potential cavity trees and b) three variables potentially linked to food supply, namely the densities of large oaks (>= 36 cm dbh) and rough-barked trees as well as the amount of dead limbs in crowns. None of these four variables was related to breeding success, suggesting that the middle spotted woodpecker adjusts home range size to habitat quality. Management decisions in favour of this woodpecker should not only focus on the conservation and promotion of large oaks, but also on the supply of trees suited for cavity excavation. PASTORINI Jennifer!, FORSTNER Michael R. J.2 & MARTIN Robert D.! ! Anthropologisches Institut; Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland * Florida Atlantic University; Division of Science; Dept. of Biology; 2912 College Avenue: Davie FL 33314; USA Phylogenetic relationships among Lemuridae (Primates) inferred from DNA se- quences 776 ZOOLOGIA ET BOTANICA ‘99 The family Lemuridae includes 4 genera: Eulemur, Hapalemur, Lemur, Varecia. Phylogenetic trees based on morphology generally place Hapalemur as the most basal offshoot. By contrast, genetic and behavioural studies have consistently placed Hapalemur as the sister-group to Lemur, with Varecia as a basal offshoot. Nodal relationships among Eulemur species also remain contentious. A mitochondrial DNA sequence data set from the ND3, ND4L, ND4 genes and 5tRNAs (Gly, Arg, His, Ser, Leu) was generated to try to clarify phylogenetic relationships within the Lemuridae. We extracted DNA from hair, blood or tissue samples from all species of all four lemurid genera and various outgroup taxa were also included. The ~2400 bp sequences were analysed using maximum parsimony, neighbour-joining and maximum likelihood. The results support a basal position for Varecia, a sister-group relationship for Lemur/Hapalemur and monophyly of Eulemur, but relationships among Eulemur species were not clearly resol- ved. One clade contains mongoz, rubriventer, and fulvus; macaco and coronatus form the second clade. H. griseus and H. aureus form a clade with strong support, but the sequence data do not yield resolution of the trichotomy involving A. simus, H. aureus/H. griseus and L. catta. Pairwise distances among all 4 taxa are approximately equal; there is no increase in genetic distance between Lemur and Hapalemur. Although our preliminary results do not definitively show whether Hapalemur is paraphyletic, they do at least strongly support a sister-group rela- tionship for Lemur and Hapalemur. PERRET Mathieu, SAVOLAINEN Vincent, CHAUTEMS Alain & SPICHIGER Rodolphe Conservatoire et Jardin botaniques de la Ville de Genève; chemin de l'Impératrice 1; CH-1292 Chambésy; Switzerland Pollination systems and speciation in Sinningieae (Gesneriaceae, Angiosperm) In angiosperms, plant-pollinator interactions have been suggested to play a major role in speciation, but few studies have addressed how such ecological relationships originate and evolve. In the present study, we have chosen a phyletic approach in the plant tribe Sinningieae. These herbaceous/shruby plants from the Neotropics are pollinated by bees, butterflies, moths, hummingbirds or bats. The major question that we address is how speciation occurs at the plant-animal interface, viz. how pollination systems versus allo/sympatry took part in the biodiversification process of these gesneriads. For this purpose, we inferred the phylogeny of all plant species (75 sp.) using DNA data from three noncoding regions of the plastid genome (trnL-trnF, rbcL-atpB and rpl16). In addition, since sugar ratios have been correlated with pollinator identity, nectar has been collected and analysed. These rewards, together with the morphological syndromes, have been mapped onto the phylogeny to trace the evolutionary events that should have given rise to the modern distribution of ecological features. In the case of species with a wide geographical distribution, we show how sympatric speciation could have occurred with pollinator switches. By contrast, in the case of clades containing species with the same pollinator, we show that speciation has relied on geographical isolation or minute mor- phological specializations to specific pollinators. Finally we discuss complementary approaches (population/field experiments, developmental genetics) that will be developed in the near future. PERRIN Nicolas, BALLOUX Francois & GOUDET Jérome Institute of Ecology; Biology Building; University of Lausanne; CH-1015 Lausanne; Switzer- land Breeding systems inferred from genetic structures We investigate the possibility to infer the parameter values of breeding systems (number and size of breeding groups within subpopulations, sex-specific dispersal, polygyny level) from field estimates of the amount and apportionment of genetic variance (hierarchical Fst). The approach is illustrated for the case of Crocidura russula, a monogamous shrew with female- biased dispersal. ZOOLOGIA ET BOTANICA ‘99 aga) PFLUGSHAUPT Kaspar & KOLLMAN Johannes Geobotanisches Institut ETH; Ziirichbergstrasse 38; CH-8044 Ziirich; Switzerland Reproductive traits in small and isolated populations of Prunus mahaleb At the margin of its distribution in central Europe Prunus mahaleb is confined to steep, south- facing cliffs and scree slopes which have a particular sunny and warm microclimate. The fleshy-fruited shrub is widespread in the Mediterranean basin and extended its area of distribution towards central Europe during a warmer period after the last glaciation. Thus, the relictic populations in central Europe have been isolated for about 5,000-6,000 years. To investigate the effects of isolation and population size on reproduction of P. mahaleb several flower and fruit traits were sampled in eleven small and isolated populations in Switzerland. Mean fruit pulp mass and seed mass were tightly correlated across all populations, and positively correlated with flower size. Flower size, pulp and seed mass correlated neither with population size nor with the degree of isolation. However, the three variables showed a significant negative correlation with the altitude of the study sites. In a Principle Component Analysis (PCA) seven of the populations were clearly differentiated by their reproductive traits. The first axis of the PCA was mainly determined by flower size, pulp and seed mass; this axis ran parallel to a gradient in precipitation. The second and third axes correlated with the number of flowers and fruits per raceme, and percentage abortion of immature fruits within the racemes. These results suggest that climatic factors are more important to explain the observed variation in flower and fruit traits than population size or the degree of isolation. PULFER Corinne, OERTLI Beat, CASTELLA Emmanuel & CAMBIN Diana Laboratoire d'Ecologie et de Biologie Aquatique; University of Geneva; chemin des Clochettes 18; CH-1206 Geneve; Switzerland Biodiversity of Trichoptera in Swiss ponds Within the framework of a larger project aiming at the classification of ponds in Switzerland and the set up of an assessment method for the conservation value of these ecosystems, larval Trichoptera assemblages were surveyed in 56 ponds varying in altitudinal range, morphometry and nutrient status. Sampling was carried out on a habitat basis, accounting for all the major habitat types occurring in the ponds. Relationships between the occurrence of the 35 taxa found and a set of environmental variables (structure of the pond surrounding, water quality) are presented. They stress altitude as a prominent constraint for the prediction of trichopteran diversity. REUTER Max & KELLER Laurent Institute of Zoology and Animal Ecology; University of Lausanne; Batiment de Biologie; CH-1015 Lausanne; Switzerland Conflict over caste determination in eusocial hymenoptera Haplodiploidy makes that in the eusocial hymenoptera workers favour a female-biased sex ratio whereas the queen prefers an equal investment in male and female sexuals. Although both workers and gynes develop from female eggs (caste determination occurring during the larval Stage), we show in a simple model, that the evolutionary stable (ES) investment in workers for the queen and the workers do not differ, if the number of female eggs available is not limited. In this case, workers are not produced at the expense of gynes. Introducing a constraint on the adult sex ratio (gynes and workers to males) in our model, we find that the workers’ ES investment in worker production is lower than that of the queen. At the workers optimum, overall sexual production (a function of investment in workers) is smaller than at the queen's ES strategy (ESS), but the sex ratio is more female biased. If worker-gyne determination is under control of the developing larvae themselves, the ES investment in workers is slightly 778 ZOOLOGIA ET BOTANICA ‘99 lower than at the workers' ESS. Our results indicate that considerable conflict over caste determination arises between the queen on one hand and the workers and female larvae on the other. Worker-larvae conflict, in contrast, is almost insignificant. RIBI Georg Zoologisches Museum der Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Habitat alterations facilitate genetic introgression between European Viviparus spe- cies (Mollusca, Prosobranchia) Viviparus ater lives in lakes and rivers, whereas V. contectus originally lived in semi-perma- nent swamps, ponds and small streams. However, swamps have been drained throughout Europe and V. contectus has lost most of its preferred habitats. On the other hand, new habitats such as canals and harbours have been created which are suitable to both species, and where they now live sympatrically. Since V. ater and V. contectus have been found to coexist only in disturbed or man-made habitats, it can be assumed that the two species occupied different habitats during most of their evolutionary history, and that it is only now that coexistence between them is widespread. This may also be an explanation for the fact that the two species, despite pronounced differences in many characters, can still hybridise. Hybrids are fertile in backcrosses with both parental species. Natural hybrids and backcrosses were found in Lake Garda, Italy, and electrophoretic data suggest that genetic introgression may occur between V. ater and V. contectus. Although introgression may have only just started, the high frequency of interspecific copulations in nature, the high survival rate and the high fecundity of hybrids and backcrosses suggests that there is a high potential for introgression. This is an example of genetic changes of natural populations probably brought about by man made habitat alterations. RIGBY Mark C. & JOKELA Jukka ETH Zürich; Experimental Ecology; ETH Zentrum NW; CH-8092 Zürich; Switzerland High costs of predator avoidance compromise immune defences in snails (Lymnaea stagnalis ) Organisms are often attacked simultaneously by several kinds of enemies, requiring different defences. The optimal level of defence is a balance between the costs and benefits of defence. However, with multiple enemies, the allocation to one defence may constrain the necessary allocation to another defence. Here, we show that two different defence strategies have sub- stantial interacting fitness costs. We manipulated the allocation to predator avoidance and immune defence in the freshwater pond snail Lymnaea stagnalis. Increasing predator avoidance correspondingly reduced energetic reserves, the proportion of snails reproducing, and repro- ductive output. Immune challenge imposed high costs when predator avoidance was infrequent, but lower costs when predator avoidance was frequent. This suggest that snails trade immune defences for predator avoidance, weakening their ability to defend against parasites when predators are abundant. ROMER Jann Zoologisches Museum der Universitat Ziirich; Winterthurerstrasse 190; CH-8057 Ziirich; Switzerland Astacus leptodactylus - a threat to native species in Lake Aegeri, Switzerland ? Lake Ageri hosts three crayfish species: Austropotamobius torrentium, Astacus astacus and Astacus leptodactylus. The last species, originally from Eastern and South-eastern Europe, was caught for the first time in Switzerland in 1979. It has a higher reproductive rate and grows ZOOLOGIA ET BOTANICA ‘99 779 faster than the two indigenous species (Au. torrentium & As. astacus). To estimate the potential of As. leptodactylus to affect indigenous species in Lake Ageri, I examined the distribution and relative densities of the three species and their association with different substrate types. collecting plot samples by SCUBA-diving. I found crayfish at 29 of 33 sites between April and September 1997. I found only one species at 8 sites, two species at 15 sites and all three species at 6 sites. Species composition differed significantly between sites according to substrate composition. Au. torrentium prefers stony substrate, while the two Astacus species were more frequent at sites with muddy substrate and reed vegetation. In all three species, I found gravid females and recently hatched juveniles and all size classes were present. Animals missing appendages were more often As. astacus than either of the two other species. Limb loss was significantly more frequent in As. astacus at sites where the introduced species (As. lepto- dactylus) was present at high densities. This suggests that As. leptodactylus is dominant in competitive interactions with As. astacus and the potential for competitive exclusion exists. ROULIN Alexandre, KOLLIKER Mathias & RICHNER Heinz Department of Zoology; University of Bern; CH-3032 Hinterkappelen; Switzerland Hawk-dove games among offspring of 7yto alba Conflicting parent-offspring interactions are among the most common behaviours in both humans and animals. Models of offspring begging propose two possible outcomes of the conflict where either scramble competition among siblings leads to escalated begging level to attract the parents’ attention, or where begging is a honest indicator of food need directed at the parents. However, nestlings may also use each other's vocalisation behaviour to decide whether to escalate or retreat in a contest over food, as in a game theoretical model with conditional Hawk-Dove strategies. Following game theory, a nestling should play Dove when a nestmate vocalises more intensely and may therefore be willing to escalate in the contest over the next food item delivered, and he should then increase vocalisation once this nestmate has consumed the prey. Such a change in vocalisation is not predicted by both scramble competition and honest signalling models. All three experiments in wild barn owls Tyto alba suggest that nest- lings are sensitive to each others vocalisation level in a manner consistent with the conditional Hawk-Dove strategies. The findings suggest that the understanding of parent-offspring conflict may require the incorporation of a game theoretical approach of asymmetric contest into cur- rent begging models. Roy Bitty, LANDOLT Martin, GAECHTER Fani & SCHMID Bernhard Geobotanisches Inst. ETH: Zürich; und Institut für Umweltwissenschaften; UNI Zürich; Switzerland Relationships between biodiversity and enemy attack rates Theory suggests that disease and herbivory should be more prevalent where plant biodiversity is low, because effective host density is correspondingly high, and high host density should facilitate transmission. To assess this idea experimentally, we quantified disease levels and herbivore attack in a the BIODEPTH experimental plots in Switzerland. Plant diversities per plot were 1, 4, 8, or 32 plant species per plot. Our 1998 data show that of the nine natural ene- mies studied on three different hosts, 5 were more prevalent in the lowest-diversity plots (in agreement with theory), but 9 were more prevalent in higher-diversity plots (violating the predictions of theory). These results suggest that the individual ecology of each disease or insect must be considered before host diversity levels can be used to predict attack rates. RUSTERHOLZ Hans-Peter & ERHARDT Andreas Department of Integrative Biology; Section of Conservation Biology (NLU); St. Johanns- Vorstadt 10; University of Basel; CH-4056 Basel; Switzerland 780 ZOOLOGIA ET BOTANICA ‘99 Influence of human land-use change on the genetic population structure of the Adonis Blue butterfly (Lysandra bellargus Rott) Today, habitat fragmentation is one of the most important processes causing losses of biodiversity and increasing species extinction rates. Consequently, assessment of population genetic structure and diversity is increasingly relevant for the management for threatened species. In this study, RAPD-PCR analyses are used to investigate the effects of habitat fragmentation on the genetic structure of Adonis Blue butterfly populations in the northwestern Jura mountains. Twelve populations ranging from small (estimated population size: 10-30 individuals) to medium (35-70 individuals) to large (200-500 individuals) were investigated. Our study shows that: (1) genetic diversity and hence molecular variance are reduced in the small populations, and (2) these two measurements do not differ between medium and large populations, (3) molecular variance between populations are high (35%), (4) there was neither a correlation between genetic distances and geographical distances, nor did clustering of the genetic phenotypes correspond to the geographic locations of the study sites, and (5) all investigated populations are highly isolated (range of calculated gene flow between populations: 1 x 10-4 to 8 x 10-5). This example shows that man-made changes of landscape can reduce and fragment butterfly populations with the associated genetic consequences. These findings are of particular relevance for conservation, since they suggest that fragmentation of formerly large butterfly populations causes disruption of gene flow and genetic erosion in small relic butterfly populations. RYSER Peter, ZOBEL Kristjan, EEK Liina, URBAS Pille & WAHL Stefan Geobotanisches Institut ETH Zürich: Gladbachstrasse 114; CH-8044 Zürich; Switzerland Department of Botany and Ecology; Tartu University; Lai 40; EE-2400 Tartu; Estonia Plant characteristics which drive succession Artificial grasslands were grown in pots for two years at two levels of irradiance and nutrients. Species mixtures consisted of three combinations of ten ecologically contrasting grass species each. Although the treatments provoked a clear response in growth and plant morphology, there were no differences in species composition of the swards after the first growing season. The proportion of a species in the sward after one growing season correlated negatively with its leaf tissue density and its root tissue density. A low tissue density is known to be associated with a high inherent relative growth rate of a species. Initially dominant species had shorter leaf life- spans and faster root turnover rates. In full daylight the average leaf life-span of the sward increased and the average root turnover rate decreased in the second year due to shifts in species composition. In shade, however, a strong increase of two species with high turnover rates, Poa trivialis and Dactylis glomerata, counteracted the general increase of species with low turnover rates. We conclude that the initial course of succession is largely independent of environmental conditions. Among the species present at a site, fast growing species with a low tissue density initially dominate the vegetation regardless of irradiance or nutrient supply. These species have shorter leaf life spans and faster root turnover rates, which indicate that their relative abundance would diminish in the course of a succession. However, this was only partly confirmed by the data of the second growing season. SALAMIN Nicolas, RUEDI Manuel & SAVOLAINEN Vincent Institute of Ecology, University of Lausanne, CH-1015 Lausanne; Switzerland Inferring complex phylogenies: which strategy to adopt ? During the evolution of species, speciation and extinction events were far from uniform. In particular, many groups of organisms experienced rapid diversification known as adaptive radiation. These radiations occurred during short intervals of time, making phylogenetic recons- truction difficult. Moreover, these radiations may happen at different times within the same ZOOLOGIA ET BOTANICA ‘99 78] organismal history, further complicating reconstruction. Among several possible strategies to overcome such complexities, it is often proposed to combine a rapidly evolving gene (i.e. expected to best resolve recent radiations) with data from a slowly evolving gene (i.e. expected to best resolve ancient radiations). We have investigated this hypothesis by a series of simu- lations. Sequences evolving at different rates were simulated along a model tree involving a recent and an ancient radiation. Performances of phylogenetic reconstruction were compared for rapidly and slowly evolving genes, analysed separately, or in combination. Even with extensive data sets, neither strategy was completely successful. We then argue for a more refi- ned strategy and test these recommendations with real DNA data sets. SCHIEGG Karin Chair of Nature and Landscape Protection; Swiss Federal Institute of Technology; c/o WSL; CH-8903 Birmensdorf; Switzerland Patch size and patch connectivity: habitat relationships of saproxylic Diptera and Coleoptera The intensive forest exploitation during the past centuries almost eradicated dead wood from Central European forests. Species confined to dead wood do not only suffer from habitat destruction but also from the effects of fragmentation. I studied the influence of volume and spatial arrangement of dead wood on saproxylic Diptera and Coleoptera at different scales. Dead wood connectivity was a better predictor of species richness and diversity than dead wood volume. The abundances of saproxylic Diptera and Coleoptera were influenced by the presence of dead wood within 150 m around the traps. Thus, a network of 'dead wood islands' each of about 7 ha in size would be optimal for the effective conservation of saproxylic insects in Central European forests. SCHIEGG Karin Chair of Nature and Landscape Protection; Swiss Federal Institute of Technology; c/o WSL; CH-8903 Birmensdorf; Switzerland Small but beautiful: influence of dead wood dimensions on species richness and diversity of saproxylic insects Dead wood is a major component of forest ecosystems providing habitat for a variety of species. The intensive forestry during the past centuries nearly eradicated all dead wood, at least in Central European forests and, as a consequence, the saproxylic fauna depauperated. Efforts made to promote dead wood mostly focus on logs and snags as these are thought to be ecologically more important than branches or twigs. I studied species richness, diversity and species abundance distributions of saproxylic Diptera and Coleoptera reared from logs and branches of beech (Fagus sylvatica) using eclectors (emergence traps). In both insect groups, branches hosted more species, more rare and exclusive species and higher diversity than logs. Thus, activities for maintaining and enhancing forest biodiversity must also include the preservation of dead branches. SCHÜRCH Stéphanie, PFUNDER Monika & Roy Barbara A. Geobotanisches Institut ETH; Ziirichbergstrasse 38; CH-8044 Ziirich; Switzerland Interactions between ants, Euphorbia cyparissias, and its pathogenic rust fungi Ants form a mutualism with Euphorbia cyparissias L. since they disperse its seeds which are specially provisioned with a food body (elaiosome). Ants also make up more than 35% of all visitors to the flowers of E. cyparissias and to its parasitic rust fungi (which require insects for reproduction). In this study, we asked whether the mutualism with ants also extends to 782 ZOOLOGIA ET BOTANICA ‘99 pollination of the flowers. In addition, we wondered whether ants fertilised its pathogens as well. A mating system experiment established that E. cyparissias can self, but it set more seeds when outcrossed. Insect exclusion experiments showed that ants do pollinate the flowers, but do not fertilise the fungi. Thus ants have positive interactions with the plants and no effect on its pathogens. SoLIVA Marco & WIDMER Alex Geobotanisches Institut ETH; Zollikerstrasse 107; CH-8008 Zürich; Switzerland Phylogenetic relationships within the orchid genus Ophrys based on multiple DNA sequences The genus Ophrys shows a highly specialised pollination system. Flowers imitate female hymenopterans and provoke males to copulate with the flowers. If a male tries to copulate with flowers from different plants, pollination can occur. This type of pollinator exploitation is known as pollination by sexual deceit and is often highly specific. Due to this specificity, strong isolation among plant gene pools should be given. On the other hand, hybrids among species have been reported and morphological boundaries are often blurred. To assess phylo- genetic relationships among Ophrys taxa, we have chosen DNA sequences from the nuclear and the chloroplast genome. The data set from the nuclear genome (ITS sequences) comprises 706 characters, 34 of which were parsimony-informative for the ingroup. Maximum parsimony (MP) analyses produced 6 shortest trees. The strict consensus tree contains 3 polytomies. The cpDNA data set comprises 978 characters of which 28 were parsimony-informative for the ingroup. Cladistic analyses produced 32.677 most-parsimonious trees. Two polytomies exist in the strict consensus tree. Strict consensus trees based on nuclear and chloroplast DNA sequence variation are largely congruent. One apparent difference between the two trees is the position of the O. fuciflora complex. According to the ITS sequences they form a separate lineage. In contrast, in the cpDNA tree the complexes O. argolica, O. fuciflora and O. sphegodes form a large polytomy. This difference may be due to different levels of variability of the two consi- dered regions. Over all, the large polytomies suggest that most Ophrys species have diverged relatively recently. STEPHAN André, MEYER Andrea & SCHMID Bernhard Institut für Umweltwissenschaft: Universität: Winterthurerstrasse 190; CH-8057 Zürich; Switzerland Plant diversity positively affects soil microbial diversity Soil microbial diversity is neither easily measurable nor precisely defined. Measurements of potential C-source utilisation by microbial communities can be done with BIOLOG plates. The resulting patterns of C-source utilisation represent an informative projection of the soil micro- bial community structure. We applied this technique (BIOLOG EcoPlates with 31 C-sources) to soil samples taken from an experiment designed to assess the effects of plant diversity on ecosystem functioning. In this experiment special emphasis was given to true replication of the plant diversity levels and to the influence of the functional groups grasses, herbs and legumes. We found that increasing plant species richness (log-scale) led to linear increases in (1) total soil bacterial activity (p<0.01) and (2) soil bacterial diversity measured by the C-source utili- sation patterns (richness p<0.05, Simpson-Index p<0.01, Shannon-Wiener index p<0.05). More precisely, (3) the microbial activity increased with plant species richness on more than half of the 31 investigated C-sources (p<0.001 for 3, p<0.01 for 7, p<0.05 for 2, p<0.1 for 4 C- sources), whereas no decline in activity was found for any C-source. Furthermore, (4) soil mi- crobial activity linearly increased with increasing number of plant functional groups (p<0.01). Finally, (5) the three plant functional groups differentially stimulated the soil microbial activity in the order legumes>herbs>grasses (p<0.05 for 3, p<0.01 for 2, p<0.001 for 1 of the 31 C- sources). ZOOLOGIA ET BOTANICA ‘99 783 STUDER Sibylle!, EbWARDS Peter J.2 & MEISTER Erhard! | Swiss Federal Research Station for Agroecology and Agriculture; Reckenholzstrasse 191; CH-8046 Ziirich; Switzerland 2 Geobotanical Institute ETH Zurich; Ziirichbergstrasse 38; CH-8044 Zürich; Switzerland Enhancing biodiversity in agricultural grasslands: plant community structure under different managements We are studying the effects of different management systems and of landscape structure on insect (true bugs) and plant diversity in grasslands. This poster focuses on plant community structure of differently managed grasslands. The study site is an area of some 3.5 km? in the Schaffhauser Randen in Northern Switzerland. The three administratively recognised mana- gement types are: extensive (no fertiliser, one cut), little intensive (little manure addition, two cuts) and medium intensive (moderate fertilisation (mineral or manure/slurry)). For each management type plant species composition has been assessed in twelve locations (replicates). As expected, species richness was highest in extensively used meadows (42 species per 6 m?) and lowest in the medium intensive type (30 species per 6 m2). The little intensive type was intermediate (35 sp. per 6 m7). However, in the ordination analysis (CA) the little intensive type clearly forms a separate group. The results concerning community structure (discussed below) also emphasise the special position of this grassland type. The dominance structure in the little intensive type differs considerably from the other two. In contrast to the extensive and the medium intensive types, the rank abundance distributions of the little intensive type show no clearly dominant species. Furthermore, rank consistency of the 10 most abundant species is lowest in the little intensive type. This means that a relatively large number of species can reach intermediate to high cover values and that a broad range of species can survive in these habi- tats. We conclude that the little intensive grassland type is of importance and that further work is required to assess it's value for conservation of biodiversity in grasslands. SUTER Werner WSL; Birmensdorferstrasse 111; CH-8903 Birmensdorf; Switzerland A biodiversity strategy as a national nature conservation concept Nature conservation today is in transition from the period, when action meant mainly reaction, to a new era of planned conservation as an integral part of land-use activities. Since there is little difference between nature and biodiversity conservation, the mandate from the convention on biological diversity (Rio de Janeiro 1992) to prepare and implement national biodiversity strategies should be taken by the member countries as an opportunity to develop comprehensive plans for nature conservation. Using Switzerland as an example of the many countries which have not yet embarked on preparing such a strategy, it is argued that monitoring schemes, no matter how good they are, cannot substitute for genuine conservation strategies, but should be seen as instruments for measuring their success. Conservation biology should play a stronger role in the process leading towards a comprehensive national conservation plan. There will be a number of elements central to such a strategy: 1) To define the national role in conserving bio- diversity. 2) To state the main goal of the strategy (e.g. conservation of net species numbers). 3) To define target entities of the strategy at different levels (genetic, species, and ecosystems) and the criteria for their measurement (e.g. size of minimum viable populations, minimum size and distribution patterns of biotopes, etc.). 4) To set priorities according to the country's respon- sibility for species and biotopes. 5) To quantify goals separately for each biogeographical region (habitats ?). 6) To develop ideas of how not only populations and biotopes, but also ecological processes can be conserved, and evaluate where alternative management options (managing the cultural landscape versus letting wilderness develop) are appropriate. 784 ZOOLOGIA ET BOTANICA ‘99 THURNHEER SYLVIE! & ANHOLT BRADLEY R.? ! Institute of Zoology; University of Zurich; Winterthurerstrasse 190; CH-8057 Ziirich; Swit- zerland ? Department of Biology; University of Victoria; PO Box 3200; Victoria BC; V8W 3N5; Canada Higher order interactions: The influence of predator-induced indirect effects on competitive ability in waterfrog larvae (Rana esculenta and Rana lessonae) Predators do not only reduce prey density, but also induce behavioural responses in prey species. In waterfrogs, tadpoles of the hybrid Rana esculenta and the parental species R. lesso- nae, reduce their activity in the presence of predators. Since the amount of reduction differs between the genotypes, this non-lethal effect of predators can potentially change the compe- titive ability of their larvae. We investigated this possibility by measuring feeding rates and development for tadpoles of R. lessonae and 3 hemiclones of R. esculenta in the absence and presence of an odonate predator (Aeshna cyanea). At the beginning of the larval period (first month), R. esculenta fed and grew more in the presence than in the absence of Aeshna, contrary to our expectation. R. lessonae fed and grew equally in both treatments. Thereafter, R. escu- lenta reversed the pattern, feeding equally but growing less when Aeshna was present. For R. lessonae time to and size at metamorphosis did not differ between treatments. In contrast, R. esculenta hemiclone GUT3 metamorphosed earlier or hemiclone GUT2 slightly heavier when Aeshna was present. This surprising outcome may result from differential reduction of the swimming activity and, hence, energy expenditure of the genotypes. But whatever the mecha- nism, Aeshna predators seem to favour the competitive ability of R. esculenta. ULLRICH Karin S. & EDWARDs Peter J. Geobotanical Institute; ETHZ; Zürichbergstrasse 38; CH-8044 Zürich; Switzerland Enhancing biodiversity in arable land: the role of management and landscape struc- ture The intensification of agriculture in Europe within the last decades has led to changes in landscape structure and considerable reduction of biological diversity in agricultural systems. There is now great interest in finding ways of managing such systems to preserve and enhance biological diversity, partly for conservation reasons and partly to achieve greater sustainability. To this purpose the Swiss policy strongly promotes the introduction of ecological compensation areas, such as wild flower strips, set-asides and hedges, in intensively-used arable land. The objective of this study is: 1. To compare the effectiveness of different management schemes in enhancing floristic and faunistic diversity within some of the ecological compensation areas, especially wild flower strips. 2. To examine how ecological compensation areas contribute to biological diversity at a landscape level. The abundance and diversity of plants and insects (Heteroptera) in wild flower strips is being compared and interpreted in terms of the management and the landscape structure. In particular interest focuses on the contribution of spontaneously appearing species. An experimental approach has been used to investigate the importance of dispersal processes and the data will be modelled using GIS. UTELLI Anna-Barbara & Roy Barbara A. Geobotanisches Institut ETH; Zürichbergstrasse 38; CH-8032 Zürich; Switzerland Altitudinal variation in nectar robbery in Aconitum lycoctonum Studies of nectar robbery have shown that positive, neutral and negative effects on reproduction of the plant can be found. Only few studies also considered the possible geographic variation in the plant-nectar robber interaction. We examined the causes and consequences of nectar robbery in Aconitum lycoctonum in high (2000 m) and low (500 m) elevation populations in ZOOLOGIA ET BOTANICA ‘99 785 Switzerland. At high elevation, nectar robbery was more frequent and was caused by short-ton- gued bumblebees. A nectar robber exclusion experiment showed that nectar robbery by short- tongued bumblebees had no effect on plant female reproductive success. This result can be explained by the fact that robbing bees rarely damaged the nectaries, and piercing does not decrease flower longevity. In addition, we found that about 10% of the visits these bees made were pollen collecting. Thus, these bees may act as low efficiency pollinators and may at times be mutualistic associates, not just robbers. At low elevation, the holes in the flowers were probably caused by beetles (Meligethes viridescens). These beetles are mostly pollen eaters, and as such reduce plant male fitness. In addition, they damage ovaries and nectaries when feeding. Overall, our results show that there can be strong geographic variation in nectar robbe- ry, in the nectar robber community, and the consequences of robbery for host plant fitness. VIETTE Michele, TETTAMANTI Chiara & SAUCY Francis Department of Biology, Ecology & Biology of Organisms; University of Fribourg; Perolles, CH-1700 Fribourg; Switzerland Preference for acyanogenic white clover (Trifolium repens) in the vole, Arvicola terrestris in simple and repeated choice tests In this paper we report investigations on the defensive role of plant secondary compounds in the reciprocal interactions between the vole, Arvicola terrestris, and its preferred food item Trifolium repens. In white clover, the production of cyanogenic glycosides (liberating HCN when the plant is attacked) is controlled by a simple genetic system involving two diallelic genes. The maintenance of a polymorphism for cyanogenesis in natural populations of white clover is poorly understood and could be explained by a selective pressure of herbivores on different cyanotypes. In this study, we conducted two series of cafeteria tests during which the voles were offered two bunches of white clover. In simple choice tests, 181 animals were tested in 13 combinations using six varieties of 7. repens (3 acyanogenic and 3 cyanogenic). When offered opposite cyanotypes, they showed a strong preference for the acyanogenic plants, while no significant preference could be detected in tests with similar cyanotypes. However, in the latter case, they ate significantly more in tests with acyanogenic varieties. In repeated choice tests, 40 voles were tested 8 times at one or two weeks intervals for a particular choice in order to investigate the constancy of their behaviour. Again, voles showed a significant preference for acyanogenic plants and most of them showed constant choices. The consequences of these results for the maintenance of polymorphism for cyanogenesis in T. repens will be discussed. VORBURGER Christoph & RIBI Georg Zoological Museum; University of Ziirich; Winterthurerstrasse 190; CH-8057 Ziirich; Switzer- land Mechanisms of impact of the introduced crayfish Pacifastacus leniusculus on the native crayfish Austropotamobius torrentium in Switzerland Interspecific behavioural interactions by which the introduced North American crayfish Paci- fastacus leniusculus could affect and possibly displace the native crayfish Austropotamobius torrentium in Europe were studied. We conducted a series of laboratory experiments on sub- strate selection, aggressive interactions and competition for shelter, and we compared size and growth between species. In choice experiments the two species differed in substrate preference with A. torrentium mainly selecting gravel and P. leniusculus mainly mud. Neither species was inherently dominant in antagonistic interactions, but dominance was size-dependent, favouring the larger and faster growing species P. leniusculus. Shelter possession was generally deter- mined by aggressive dominance, although A. torrentium showed a stronger preference for shel- ters and sometimes defended them against larger P. leniusculus. Experimental P. leniusculus were infected with the crayfish plague fungus Aphanomyces astaci, to which they are resistant. 786 ZOOLOGIA ET BOTANICA ‘99 During the experiments, the disease was transmitted to non-resistant A. torrentium, which died 10 to 20 days after contact with P. leniusculus. We conclude that antagonistic interactions could play an important role in species replacements among crayfish by determining access to limited resources. However in the present case, a crucial factor which may promote an eventual species replacement is the crayfish plague. VOUILLAMOZ José Institute of Ecology, Systematic Botany and Geobotany; University of Lausanne; CH-1015 Lausanne; Switzerland The genus Onosma (Boraginaceae) in Switzerland: chromosome numbers, allozymic polymorphism and evolutionary history of a group of rare taxa The genus Onosma is represented in Switzerland by two rare species distinguished by the numbers of rays (0-20) arising from the base of the main setae, showing distinct chromosome numbers: O. pseudoarenaria Schur s.l., with stellate setae (5-20 rays), and O. helvetica (A. DC.) Boiss. with simple setae (0-5 rays). The populations of O. pseudoarenaria Schur s.l. are split up and located only in Valais, between Martigny and Sierre, and show 2n=26 or 2n=28, depending on the presence of B chromosomes. They reveal bimodal chromosomal comple- ments with 12 large (L) and 14 or 16 small (S) chromosomes. The populations of O. helvetica (A. DC.) Boiss. form two poles on both sides of those of O. pseudoarenaria Schur s.l., one in Ollon (VD) and the other in Haut-Valais, between Sierre and Brig. They both show 2n=20 presenting bimodal chromosomal complements (12 L + 8 S) as well. The caryological compa- rison of these two species with those from Italy (O. echioides agg. 2n=14 S) and France (O. fastigiata s.1., 2n=12 L) suggests the hypotheses of an allotetraploid origin for O. pseudo- arenaria Schur s.l. and of a hybrid origin for O. helvetica (A. DC.) Boiss. through back- crossing. These hypotheses are under investigation using starch gel electrophoresis to analyse the allozymic variability and try to confirm or invalidate the putative parent species. Along with the clarification of their exact karyograms, these data may allow to put forward both the origins and the postglacial recolonization routes of the genus Onosma in Switzerland. WALTHER Gian Reto Geobotanisches Institut ETH; Zürichbergstrasse 38; CH-8044 Zürich; Switzerland Why palms in Switzerland ? The present vegetation in a given site is coupled with the local climatic and edaphic conditions. Changes in climate may also cause changes in the species composition of the vegetation. The presented study analyses possible causes of vegetation changes. Especially the spreading of exotic evergreen broad-leaved shrub and tree species points to a possible link with the changing climate in recent times. WEIBEL Urs Geobotanisches Institut ETH & Schweiz. Vogelwarte Sempach; Ziirichbergstrasse 38; CH- 8044 Ziirich; Switzerland Nest site selection and breeding success of skylarks (Alauda arvensis) in an arable landscape with wild flower strips Modern intensive farming regimes in Central Europe have led to severe decreases in skylark populations in the last few decades. The lack of sufficient nesting habitats throughout the breeding season due to the more frequent applications of fertilisers and a restricted crop rotation led to a reduction of the breeding attempts. Additionally, brood losses caused by far- ming operations, mainly mowing, led to a not sustainable breeding success. In a four year ZOOLOGIA ET BOTANICA ‘99 787 survey of a skylark population in the Klettgau region (Canton Schaffhausen, northern Switzer- land) nest site selection and breeding success was investigated. The study site (5.3 km?) is mainly used for intensive winter cereal farming, but the fields are small and wild flower strips ("Buntbrachen") are established on 6 ha. The breeding success was calculated with the May- field method. Throughout the breeding period skylarks showed a shift from winter cereals towards root vegetables as nesting sites, according to the vegetation structure. Wild flower strips, set-asides and cultivated grassland had the highest relative use as nesting habitat. Nests were more often found near to the border of the fields. The daily survival rate of eggs was significantly higher than that of nestlings. The estimated breeding success probability was about 20% between 1995 and 1997, but 38% in 1998. Broods in cereals reached the highest success probabilities, and that of nests in wild flower strips was mediocre. Nests near to the field border had a lower daily survival probability. Predation caused about 70% of all nest losses. WEISSER Wolfgang W. Zoology Institute; University of Basel; Rheinsprung 9; CH-4051 Basel; Switzerland Predator-induced morphological shift in the pea aphid Aphids exhibit a polymorphism whereby individual aphids are either winged or unwinged. The winged dispersal morph is mainly responsible for the colonisation of new plants and, in many species, 1s produced in response to adverse environmental conditions. Aphids are attacked by a wide range of specialised predators and predation has been shown to strongly influence the growth and persistence of aphid colonies. We reared colonies of pea aphids in the presence and the absence of an adult ladybird and noted the types of morphs that were produced. Winged phenotypes first appeared in the presence of a ladybird more than one aphid generation before they were found in the control. This indicates that aphids responded to the presence of the predator by producing the dispersal morph which can escape by flight to colonise other plants. In contrast to previous examples of predator-induced defence this shift in prey morphology does not lead to better protection against predator attack, but enables aphids to leave plants when mortality risks are high. ZUBER Doris & WIDMER Alex Geobotanisches Institut; ETH Zürich; Zollikerstrasse 107; CH-8008 Zürich; Switzerland Intraspecific phylogenetics of the European mistletoe (Viscum album) based on nuclear and chloroplast DNA sequences The European mistletoe (Viscum album) is a widely distributed, hemiparasitic plant species. Currently, three subspecies are recognized which differ in their host preferences. In central Europe, V. album ssp. austriacum occurs on pine (Pinus spp.) and rarely on Norway spruce (Picea excelsa); V. album ssp. abietis is restricted to silver fir (Abies alba), while V. album ssp. album grows exclusively on a wide variety of deciduous trees. Crossinfection experiments reported in the literature suggest that the host-parasite interaction is highly specific, although few successful crossinfections have been observed. The aim of this project was to assess phylogenetic relationships among Visum album isolates from different host species to test whether molecular evidence supports the distinction of three host races and to assess their phylogenetic relationships. is Au ve I Be XII { A } yy tati ike >, RR; ‘fey = F1 LEA Age 96 im! a aa ee te Le ni BY nina Sn tik bat Er a Rh DEI aa n REVUE SUISSE DE ZOOLOGIE 106 (4): 789-811; décembre 1999 Psélaphides paléarctiques nouveaux ou méconnus (Coleoptera: Staphylinidae: Pselaphinae) Claude BESUCHET Muséum d'histoire naturelle, case postale 6434, CH-1211 Geneve 6, Suisse. New or poorly known Palaearctic pselaphids (Coleoptera: Staphylini- dae: Pselaphinae). - One genus (Thelotia gen. n.) and ten new species are described: Octomicrus dentifrons sp. n. from Iraq, Faronus distinctus sp. n. from Turkey, Thelotia cebennica sp. n. from France, Amauronyx caudatus sp. n. from Yugoslavia, Namunia terricola sp. n. from Greece, Bryaxis cor- sus sp. n. from Corsica, Trissemus bellax sp. n. from Spain, 7. sulcifrons sp. n. from Morocco, 7. trilobatus sp. n. and 7. holzschuhi sp. n. from Pakistan. Trichonychini, Dimerus and six species are placed in synonymy. A neotype of Bryaxis opuntiae Schmidt-Göbel and lectotypes of Rybaxis marquardti Reitter and Bythinus banaticus Reitter are designated. Trans- fers of three species are proposed and new distributional records of Octo- micrus staphylinoides (Fiori) are provided. Key-words: Coleoptera — Staphylinidae — Pselaphinae — taxonomy — Palaearctic. INTRODUCTION La collection de Psélaphides du Muséum d'histoire naturelle de Genève compte près de 180000 exemplaires préparés, dont environ 21000 pour la Suisse et 45000 pour la région paléarctique. C'est le résultat de nombreuses recherches sur le terrain, d'achats de collections, de dons importants et d'échanges. Les identifications sont plus ou moins nombreuses et complètes suivant les régions ou les tribus. Des centaines d'espèces nouvelles sont encore a décrire, particulierement pour l'Asie où mes collegues et moi- méme avons beaucoup travaillé. Je me suis limité dans cette publication aux espèces paléarctiques, principale- ment pour "The Catalogue of the order Coleoptera of the Palaearctic region" où j'avais encore quelques synonymes et transferts à proposer, mais aussi des espèces nouvelles à décrire. J'ai également abordé des problèmes d'un intérét plus général, en relation en particulier avec la valeur taxonomique attribuée a certains caractères. Je tenais a discuter depuis longtemps, exemples a l'appui, de l'écartement des hanches postérieures des Euplectini-Trichonychini et de celui des carénules du premier tergite apparent des Trissemus. Manuscrit accepté le 22.07.1999 790 CLAUDE BESUCHET ABREVIATIONS UTILISEES POUR LES MUSEES HNHM = Hungarian Natural History Museum, Budapest MHNG = Muséum d'histoire naturelle, Genève DESCRIPTIONS Octomicrus dentifrons sp. n. Holotype è. IRAQ. Misan, Azair-Qalat Salih, 14.12.1977, tamisage de detritus (Topäl & Zilahy) (HNHM). Paratypes. Même provenance, 16 59 (HNHM), 26 29 (MHNG). Long. 1,3 a 1,5 mm. Coloration entierement d’un brun rougeätre clair. Forme grêle, subparallèle, un peu déprimée. Tête presque carrée, un peu plus large que longue, le lobe frontal tres large. Yeux bien développés, saillants, plus longs que les tempes; celles-ci arrondies. Antennes legerement moins longues que la tete et le pronotum réunis, relativement gréles; scape cylindrique, une fois et demie plus long que large; pédicelle un peu plus long que large; articles 5 et 7 un peu plus larges que les autres articles du funicule; 3 aussi long que large, 4 a 8 transverses; massue formée de trois articles, le premier de taille intermédiaire entre les articles 8 et 10, bien séparé de ce dernier, transverse; articles 10 et 11 tout à fait contigus, de même largeur; 10 trans- verse, 11 deux fois plus long, légèrement plus long que large. Pronotum aussi long que large, a peine moins large que la téte; dépression basale transverse, tres profonde, prolongée en avant par un sillon médian profond n’atteignant pas le bord antérieur du pronotum; côtés de celui-ci ornés chacun, en arrière, de deux petites fossettes pro- fondes, l’une située presque sur les angles postérieurs, l’autre étant la fossette basale externe; une petite dent latérale saillante entre ces deux fossettes. Elytres réunis a peine plus longs que larges, nettement plus longs et plus larges que le pronotum; plus grande largeur située au tiers postérieur; élytres ornés chacun de deux fossettes basales profondes, d’une strie suturale entiere, d’un sillon dorsal court et d’une carénule laterale tres fine, entiere; fossette laterale indistincte. Ailes fonctionnelles. Métasternum tres convexe. Abdomen un peu moins long que toute la partie antérieure du corps, parallele, les quatre premiers tergites apparents de méme longueur, avec la méme ornementation; base déclive, ornée de chaque côté d’une petite fossette profonde, prolongée sur le bord interne par une strie oblique rejoignant la suture tergite-pleurite. Caractères sexuels du male. Lobe frontal prolongé en avant par une dent trapé- zoidale saillante, légèrement relevée à son sommet; la base de cette dent occupe un tiers de la largeur du lobe frontal. Trochanters HI armés sur le bord postérieur d’une petite dent; fémurs III un peu renflés: tibias III prolongés à l’apex par un éperon relativement robuste (fig. 1); le tiers apical du tibia porte de longues soies sur le bord interne comme sur le bord externe, supprimées sur le dessin. Les sternites apparents II, III et IV portent chacun une dépression arrondie dans la moitié postérieure du segment; sternite V orné sur son bord postérieur de deux petites dents saillantes tres rapprochées, séparées par une échancrure étroite profonde; sternite VI avec une grande échancrure arrondie pro- fonde accompagnee de part et d’autre d’une rangée de grandes soies plus ou moins regulierement disposées; dernier sternite formé de deux hémisternites triangulaires libres (fig. 2). PSELAPHIDES PALEARCTIQUES 79] Edeage (long. 0,12mm) tres complexe, formé de deux parties bien distinctes (fig. 3). Partie basale (fig. 4) en position ventrale, avec deux petits parameres arrondis portant chacun trois soies et, a son sommet, deux longues épines courbées. Partie api- cale (fig. 5) insérée sur le milieu de la face dorsale de la partie basale, coudée presque en angle droit par rapport a celle-ci. Octomicrus dentifrons sp. n. de Mésopotamie présente des affinités évidentes avec les Octomicrus indo-malais, en particulier avec O. longulus Schaufuss, 1877, espece-type du genre Octomicrus Schaufuss, 1877. Les caracteres sexuels des males relatifs aux sternites abdominaux sont quasiment identiques. Octomicrus dentifrons differe des especes indo-malaises par les antennes plus gréles, par le lobe frontal plus large, orné chez le mâle d’une dent saillante très caractéristique et par |’ édéage. Octomicrus staphylinoides (Fiori) Dimerus staphylinoides Fiori, 1899 Nouvelles provenances. FRANCE, ALPES-MARITIMES: St. Jean-la-Riviere, au bord de la Vésubie, 30.05.1941 et 20.06.1958, 22 (Ochs). HAUTE-SAVOIE: Vongy-Thonon, au bord de la Dranse dans des débris de crue, 19.05.1964, 1 (Besuchet) et 12.07.1980, 14 32 (Besuchet). SUISSE, Genève: Malval, au bord de l’Allondon dans des débris de crue, 1.04.1965, 1? (Besuchet). YEMEN, Wadi Sharez, 03.1985, 1d (de Rougemont) (tous MHNG). à Caractères sexuels du mâle. Trochanters inermes. Avant-dernier sternite avec une grande échancrure triangulaire accompagnée de part et d’autre de soies nombreuses assez longues; dernier sternite formé de deux hémisternites triangulaires libres. Edéage (fig. 6). Long. 0,17 à 0,19mm. Base assez longuement pédonculée. Paramères atrophiés, soudés sur la face ventrale de ce pédoncule où ils sont reconnais- sables par leurs six soies. Pédoncule prolongé du côté ventral par une grande lame atténuée en pointe: cette lame porte, au milieu de sa face dorsale, une pièce complexe repliée sur elle-même, c’est-à-dire rabattue en avant. L’édéage est donc aussi formé de deux parties bien distinctes. JEANNEL a donné deux représentations de l’édéage du Dimerus staphylinoides Fiori, très différentes l’une de l’autre; la première (1950: 55, fig. 21d) résulte de l’étude d’un mâle monté in toto en préparation microscopique à la gélatine glycérinée; la seconde (1952a: 11, fig. 4), dessinée d’après un édéage extrait de l’abdomen, a été reprise dans une autre publication du même auteur (1956b: 93, fig. 12). J’ai pu monter en préparation microscopique au baume du Canada quadre édéages (4 4 de France, d'Italie et du Yémen) de cette espèce, ce qui me permet d’en donner un dessin correct et une bonne interpretation, mais aussi une description précise des caracteres sexuels du male. Octomicrus staphylinoides (Fiori), 1899 (espece-type du genre Dimerus Fiori, 1899) ne differe des Octomicrus longulus Schaufuss, 1877 (espece-type du genre Octo- micrus Schaufuss, 1877) et dentifrons sp. n. que par quelques caractéres: corps nette- ment déprimé; pronotum régulièrement atténué en arriére jusqu’aux angles postérieurs, sans dent latérale saillante; dépression basale du pronotum peu profonde, presque carrée, progressivement relevée en avant; pas de caractéres sexuels sur les trochanters 792 CLAUDE BESUCHET III et les fémurs HI, ni sur les sternites abdominaux II a V; édéage assez longuement pédonculé. Par contre de nombreux caractéres sont tout a fait identiques: structures de la tête, du pronotum, des élytres et des tergites abdominaux; même division du dernier sternite des males (fig. 2); tarses de deux articles seulement (fig. 1); antennes avec les articles 5 et 7 un peu élargis; massue triarticulée, les articles 10 et 11 complètement contigus; palpes maxillaires tout a fait semblables; méme réticulation du mésosternum; édéage également formé de deux parties bien distinctes. JEANNEL (1956: 84) avait déja remarqué cette affinité étroite, mais il avait cependant maintenu Dimerus comme sous- genre d’Octomicrus. En réalité Dimerus Fiori, 1899 n’est qu’un synonyme d’ Octo- micrus Schaufuss, 1877 (syn. nov.). Faronus distinctus sp. n. Holotype 3. TURQUIE, Antalya: 18km au sud-est de Gazipasa, 27.04.1978 (Besuchet & Löbl) (MHNG). Paratypes. Même provenance, 18 19 (MHNG); entre Antalya et Kemer, à 25km d’Antalya, 4.05.1975, 19 (Besuchet & Löbl) (MHNG); Geris près de Akseki, 23.- 29.03.1997, 14 19 (Brachat) (Coll. Brachat); env. Antalya, 1d (Franz) (MHNG). Mugla: Bayir, à 25km au nord-est de Kemer, 950m, 3.05.1975, 24 (Besuchet & Löbl) (MHNG); Gökova, 30.04.1975, 26 19 (Besuchet & Löbl) (MHNG); Cetibeli, a 15km au nord de Marmaris, 1.05.1975, 16 (Besuchet & Löbl) (MHNG); 10km au nord-ouest de Mugla, 30.04.1975, 19 (Besuchet & Löbl) (MHNG). Izmir: Bornova, 20.05.1974, 19 (Jaccoud) (MHNG); Agamemnon près d’Izmir, 28.04.1975, 15 29 (Besuchet & Löbl) (MHNG); env. Camlik, à 14km à l’est de Selcuk, 8.05.1975, 16 (Besuchet & Löbl) (MHNG). GRECE, Rhodes: Mont Profitis Ilias, 600m, 24.04.1973, 1d (Besuchet) (MHNG). Tous dans des tamisages de mousses et de feuilles mortes. Long. 1,6 à 2,0mm. Ne diffère exterieurement de Faronus parallelus Besuchet que par l’absence complete et constante des ailes, ce qui provoque quelques modifi- cations des élytres, et par abdomen toujours nettement élargi de la base jusqu’au bord postérieur du 3° tergite apparent. Elytres toujours plus courts et moins larges que ceux de F. parallelus, élargis d’avant en arriere, le calus huméral effacé; proportion lon- gueur/largeur chez sept exemplaires de tailles différentes: 0,37/0,45mm (2); 0,39/ 047mm (9); 0,40/0,48mm (3); 0,42/0,50mm (4); 0,43/0,49mm (2); 0,43/0,51mm (3); 0,44/0,53mm (2). Caracteres sexuels. Tibias III du male armés sur le bord interne apical d’un petit éperon (inermes chez parallelus). Sclérite génital de la femelle assez robuste, en arc de cercle épais (gréle chez parallelus). Edéage (long. 0,45 à 0,50mm) bien plus grand que celui de parallelus (0,32 a 0,36mm), mais cependant assez semblable, que ce soit au niveau des parameres ou du grand lobe falciforme; mais le second lobe de la partie médiane de |’ édéage (fig. 8) diffère totalement de celui de parallelus (fig. 9). Ce type d’édéage, formé dans la partie médiane de deux grands lobes bien sclérifiés, à savoir un lobe falciforme toujours semblable et un lobe portant les carac- teres spécifiques, existe chez trois espèces de Faronus. Faronus lafertei Aubé, largement répandu dans l’Europe méridionale occiden- tale (Portugal, Espagne, France, Corse, Italie et Suisse: Genève), est normalement ailé; mais les exemplaires aptères ne sont pas rares. Les édéages sont rigoureusement sem- PSELAPHIDES PALEARCTIQUES 793 blables chez les uns et les autres; c’est toujours le lobe gauche qui porte les caracteres spécifiques; il n’y a donc jamais d’inversion. Faronus pararellus Besuchet, largement répandu dans la région méditerra- néenne orientale (Dalmatie, Grece, Rhodes, Turquie, Chypre, Liban et Israél), est presque toujours ailé; je n’ai vu qu'un seul mâle aptere de Dalmatie. Les édéages sont toujours inversés par rapport a l’espece précédente; c’est donc toujours le lobe droit (fig. 9) qui porte les caracteres spécifiques. Faronus sahlbergi Besuchet (1960: 11. Holotype d: Mus. Helsinki. Loc. typ.: Liban, fleuve Lycus) n’est qu’un synonyme de Faronus parallelus Besuchet (1958: 897. Holotype 9 : Mus. Bâle, Coll. Frey. Loc. typ.: Turquie, Lyciae Taurus) (syn. nov.), car les deux femelles décrites sous ce dernier nom sont ailées et présentent un sclérite génital gréle. Faronus distinctus n. sp., localisé dans le sud-ouest de la Turquie jusqu’a Rhodes, est toujours aptere. Les inversions de l’édéage sont fréquentes; sur 12 édéages étudiés, 4 portent les caracteres spécifiques sur le lobe gauche (fig. 8) et 8 sur le lobe droit; ces inversions peuvent se manifester dans la méme localité. Thelotia gen. n. Espece-type: Thelotia cebennica sp. n. Fig. 7. Euplectini robuste, aplati, subparallèle, microphtalme et aptere. Téte avec les deux fossettes du vertex profondes, tomenteuses, largement séparées; dépression frontale bien marquée, un peu plus longue que large, prolongée de chaque còté jusqu'aux fossettes du vertex par un sillon profond. Toute la face dorsale de la tête avec de nombreux petits tubercules saillants, y compris la région oculaire et les tempes; celles-ci arrondies. Yeux réduits a quelques ommatidies. Palpes maxillaires petits. Mandibules gréles, armées d’une rangée de petites dents sur le bord interne, la partie apicale en pointe acérée. Antennes relativement courtes avec une massue de trois articles distincts. Pronotum peu convexe, plus large que long, les côtés ornés de petits tubercules saillants un peu plus gros que ceux de la téte; trois fossettes basales égales, profondes, tomenteuses, presque alignées, reliées par un sillon transversal bien marqué mais fin; pronotum orné sur toute sa longueur d’un sillon médian bien marqué, un peu moins étroit que le sillon transversal; fossettes basales latérales prolongées en arriere chacune par un sillon distinct jusqu’au bord postérieur du pronotum; encore deux fossettes tangentes a ce bord postérieur, soit une de chaque côté entre le sillon médian et les sillons latéraux. Elytres aplatis, larges et courts (de méme longueur sur la suture que le pronotum); bord postérieur concave. Chaque élytre avec trois fossettes basales tomenteuses; fossette basale interne plus grande et plus profonde, formée en réalité de deux fossettes accolées a peine distinctes, prolongée par une strie suturale entiere, courbée de façon à joindre la suture sur le bord postérieur de l’élytre; fossette basale médiane simple, plus petite, profonde, prolongée par une strie dorsale trés courte; fossette basale latérale un peu plus grande, profonde, prolongée sur son bord externe par une carene laterale saillante sur toute la longueur de l’élytre et sur son bord postérieur par un sillon parallele a la carène et accentuant ainsi son relief; il y a encore 794 CLAUDE BESUCHET Fics 1-6 Octomicrus. 1: O. dentifrons sp. n., tibia et tarse postérieurs du mâle: 2: Id., hémisternite du mâle, face ventrale; 3: Id., édéage, face laterale; 4 et 5: Id., partie basale et partie apicale de l’édéage au méme grossissement, face ventrale; ...= points d’insertion sur la figure 4 de la partie apicale; 6: O. staphylinoides (Fiori), édéage, face ventrale. une petite dent humérale saillante à côté de la fossette latérale. Abdomen plus long que le pronotum et les élytres réunis, subparallele; premier tergite apparent nettement un peu plus long, orné a la base d’une dépression tomenteuse profonde, trés courte, un peu moins large que la moitié de la largeur basale de ce segment; pas de carénules pour limiter cette dépression; tergites apparents II, III et IV de méme longueur, simples, tres largement rebordés sur les côtés sauf le 4° avec un rebord atténué d’avant en arrière. Pattes relativement courtes, sans caractéres particuliers; tarses terminés par un ongle PSELAPHIDES PALEARCTIQUES 795 gréle assez long, simple, sans épine paronguéale. Face ventrale. Téte avec la région gulaire non carénée, suivie en arriere d’une grande fossette transverse profonde, tomenteuse. Prosternum entièrement caréné, orné de chaque côté, pres du bord antérieur, d’une fossette tomenteuse profonde. Trois petites fossettes tomenteuses sur le bord antérieur du mésosternum, deux de part et d’autre de l’apophyse mésosternale, la dernière sur le bord antérieur de celle-ci. Métasternum court, orné de chaque còté de deux fossettes tomenteuses profondes, l’une sur le bord postérieur des cavités coxales II, l’autre sur le bord externe de ces mêmes cavités. Hanches I et II contigués, III très legerement séparées (0,04mm, soit la largeur apicale des tibias III). Premier sternite apparent avec le processus intercoxal étroit, atténué en pointe en avant; sternites apparents II, II et IV de même longueur, V de moitié plus court, le dernier un peu plus long, arrondi a l’apex; base du 2° sternite occupée toute entière par une bande tomenteuse très courte. Caractères sexuels du male encore inconnus. Thelotia cebennica sp. n. Holotype 2. FRANCE, Garp: Roquedur près du Vigan dans les Cévennes, 4.04.1982, dans un lavage de terre au pied d’un noisetier, dans un ravin boisé (Thelot) (MHNG). Fig. 7. Long. 2,05mm. Coloration entierement d’un brun rougeätre peu foncé. Pubescence couchée, relativement courte. Face ventrale avec les téguments lisses, brillants et non ponctués. Téte un peu plus large que longue (0,33/0,38mm), avec une saillie arrondie relativement peu élevée entre les fossettes du vertex; tubercules antennaires arrondis, peu saillants, accompagnés sur leur bord postérieur d’une fossette tomenteuse, nettement plus petite et moins profonde que les fossettes du vertex. Yeux réduits a deux ou trois ommatidies dépigmentées. Longueur des antennes: 0,70mm; scape cylindrique, presque deux fois plus long que large; pédicelle legerement moins large que le scape, un peu plus long que large; article 3 un peu plus petit, un peu plus large que long, 4 et 5 légèrement plus larges que longs, 6 et 7 très nettement plus larges que longs, l’article 7 légèrement plus large que les autres articles du funicule; 8 transverse, 9 bien plus large que les articles précédents, transverse; 10 encore plus large, transverse; dernier article (0,15/0,11mm) orné d’une grande fossette tomenteuse sur la moitié apicale de la face ventrale. Dernier article des palpes maxillaires (0,09/0,045mm) en ovale allongé. Pronotum (0,38/0,45mm) plus large que la tête, nettement plus large que long, sa plus grande largeur située au tiers antérieur, ses côtés atténués en arriere en ligne droite jusqu’aux angles postérieurs, ceux-ci obtus; disque du pronotum lisse et brillant. Elytres réunis nettement plus larges que le pronotum, nettement plus larges que longs (0,42/0,58mm avec la longueur prise sur le bord externe, 0,37/0,58mm le long de la suture). Abdomen (0,90/0,60mm au niveau du bord postérieur du 2° tergite apparent) plus long que le pronotum et les élytres réunis; longueur respective des cinq tergites apparents: 0,27mm; 0,20mm; 0,20mm; 0,20mm et 0,17mm. Longueur des tibias III: 0,40mm. Male encore inconnu. Ce genre nouveau est dédié a M. Jean-Pierre Thelot de Forcalqueiret-La Roque- brussanne, qui m’a généreusement donné le seul exemplaire connu. MM J.-P. Thelot, I. Löbl, S. Vit et moi-méme avons fait de gros lavages de terre dans cette région des 796 CLAUDE BESUCHET Cévennes méridionales pour retrouver ce Psélaphide extraordinaire. En vain malheureu- sement. Mais nous avons tous pris le rare Tychobythinus lavagnei (Deville) déja present dans le lavage de terre du 4.04.1982. Chez de nombreux Psélaphides, la partie de l’Elytre courbée sur le còté présente une petite fossette laterale prolongée en arriere par une carénule laterale. Chez Thelotia, cette fossette laterale est en position dorsale et sa carénule, particulierement bien développée, saillante sur toute la longueur de l’élytre, forme la carène latérale. C’est l’aboutissement d’une Evolution qui se manifeste déja chez les genres Trichonyx Chaudoir et Amauronyx Reitter, où la fossette latérale est encore sur le côté de l’élytre, mais avec une carene laterale bien développée, très distincte en vue dorsale. Thelotia a exactement la méme structure du pronotum que les Amauronyx, dont certaines especes portent aussi de petites granulations sur les cOtés, mais peu nombreuses. Les structures générales de la face ventrale et de l’abdomen sont aussi tres semblables; mais les Amauronyx présentent deux carénules sur le premier tergite apparent et l’écartement des hanches III est chez eux très prononcé; leurs élytres sont toujours plus longs que le pronotum, même chez les espèces microphtalmes. A mon avis les genres Thelotia et Amauronyx sont malgré tout apparentés; mais faut-il les placer dans la tribu des Euplectini LeConte, 1861 ou dans celle des Trichonychini Reitter, 1882? Les Trichonychini different des Euplectini, d’apres RAFFRAY (1903: 489; 1908: 12, 113) et JEANNEL (1950: 72, 155; 1956a: 51) par l’écartement plus ou moins grand des hanches postérieures (contigués chez les Euplectini) et par la structure des tarses, formés de deux ongles inégaux (un seul ongle chez les Euplectini). PARK (1942: 63) et quelques auteurs américains avant lui avaient remarqué que cette séparation n’était pas justifiée. Je m’étais moi aussi (BESUCHET 1956: 371) opposé à cette distinction. Avec le genre Thelotia, le probleme se pose a nouveau, avec encore plus d’acuité, car les tribus des Euplectini et des Trichonychini ont été maintenues dans le catalogue de NEWTON & CHANDLER (1989: 12, 52). L’écartement ou la contiguité des cavités coxales postérieures est un caractére d’une “importance majeure” selon JEANNEL (1959: 21), a tel point que les Euplectini (p. 95) et les Trichonychini (p. 460), jadis étroitement apparentés, se trouvent maintenant séparés dans deux sous-familles de JEANNEL différentes: Euplectitae et Bythinitae! L’écartement plus ou moins grand des hanches postérieures est un caractere difficile a évaluer. Par contre le processus intercoxal du premier sternite apparent, qui s’intercale dans l’espace laissé libre par les hanches postérieures, permet des observations précises. Ce processus intercoxal est toujours atténué en pointe étroite chez les Euplectus Leach, Bibloplectus Reitter, Meliceria Raffray, Plectophloeus Reitter, Pseudoplectus Reitter, Pseudozibus Jeannel, Trimium Aubé et Thelotia. Il est étroit, allongé, subparallele et arrondi a son sommet chez les Saulcyella Reitter (Trichonychini pour JEANNEL 1950) et les Zibus Saulcy. Ce processus est moins étroit, trapézoidal et un peu plus large que long chez Aphiliops Reitter (Trichonychini pour JEANNEL 1950). Enfin il est nettement plus large, transverse et plus ou moins arrondi a son sommet chez les Amauronyx Reitter, Namunia Reitter et Trichonyx Chaudoir (ces trois genres généralement placés dans les Trichonychini). Les Thelotia et Amauronyx sont complètement séparés dans ce schéma! 7977 PSELAPHIDES PALEARCTIQUES FIG. 7 Thelotia cebennica gen. n., sp. n., face dorsale de la femelle. Del. S. Vit. 798 CLAUDE BESUCHET Les Amauronyx, Namunia et Trichonyx présentent toujours deux ongles inégaux bien distincts. Ce deuxieme ongle plus petit, inséré obliquement par rapport au premier (épine paronguéale de JEANNEL), existe également chez différents Euplectini où il a été bien observé par CASEY (1893) (in PARK 1942: 64) et par moi (BESUCHET 1956: 372). Il n’existe pas chez Thelotia. JEANNEL (1950, 1956, 1959) a ajouté à la définition des Trichonychini: “Elytres visiblement rebordés” ou “élytres a gouttière marginale bien visible de haut”. C’est bien le cas des Amauronyx et des Trichonyx, de Thelotia aussi, mais pas des Namunia. Cette carène latérale est également bien visible chez les Plectophloeus! Il n'y a ainsi plus de caractères distinctifs entre les Trichonychini et les Euplectini puisque |’ observation annule completement la validité de ceux-ci. Les Trichonychini doivent donc étre ratta- ches aux Euplectini; ils tombent dans la synonymie de ceux-ci (Syn. nov.). Amauronyx caudatus sp. n. Holotype 6. YOUGOSLAVIE, Serbie, Tara Planina, Mitrovac (route Mitrovac-lac d’accumulation), 900m, 23.05.1984, tamisage de feuilles mortes dans une forét de hétres, érables et sapins (Besuchet) (MHNG). Paratypes. Même provenance, 1d 49 (Besuchet) (MHNG); Tara Planina, Mitrovac (route Perucac-Mitrovac), 950m, 23.05.1984, tamisage d’une veille souche de hêtre, 16 (Besuchet) (MHNG); Tara Planina, 1000m, 11.06.1984, 38 49 (Nonveiller) (Coll. Nonveiller) et 18 19 (MHNG). Long. 2,2 a 2,5 mm. Coloration entierement d’un brun rougeätre. Pubescence couchée, formée de soies de longueur moyenne. Téte un plus large que longue; bosse du vertex arrondie, ornée en arrière d’une carene médiane bien marquée jusque sur la constriction collaire. Yeux petits, formés chacun de 6 a 9 ommatidies; tempes grandes, arrondies. Antennes robustes, légèrement moins longues que la téte et le pronotum réunis; scape deux fois plus long que large, distinctement rétréci à la base; pédicelle nettement plus long que large; articles du funicule de méme largeur; 3 a peine ou légèrement plus long que large; 4, 6 et 7 nettement plus larges que longs ou transverses; 5 legerement plus large ou aussi large que long; 8 transverse; massue formée de trois articles, les deux premiers transverses, le dernier deux fois plus long que les deux articles précédents réunis. Pronotum légèrement plus large que long, un peu plus large et plus long que la tête: côtés très légèrement crénelés et rebordés en arrière de la plus grande largeur, celle-ci située un peu en avant du milieu; sillon médian entier, bien marqué du bord antérieur du pronotum jusqu’a son bord postérieur; celui-ci orné de sept fossettes contigués. Elytres réunis nettement plus larges que longs, bien plus larges et plus longs que le pronotum; calus huméral effacé; carène latérale bien visible en vue dorsale, particulierement dans sa partie antérieure; sillon dorsal bien marqué au moins jusqu’au milieu de l’élytre. Abdomen nettement plus long que les élytres, de même largeur, terminé dans les deux sexes par une dent saillante, robuste et arrondie, enticrement localisée sur le pygidium; premier tergite apparent un peu plus grand (long. 0,25 a 0,25mm), distinctement élargi d’avant en arrière; carénules basales très courtes (0,05mm), subparallèles, largement séparées (0,20 à 0,23mm) par un espace glabre à peine approfondi. Premier sternite apparent avec un processus intercoxal transverse (largeur: 0,10mm), arrondi à son sommet. PSELAPHIDES PALEARCTIQUES 799 Caractères sexuels du male. Tous les trochanters inermes. Tibias II prolongés sur le bord interne par un petit éperon. Métasternum a peine déprimé dans sa partie postérieure. Avant-dernier sternite avec une grande échancrure arrondie sur le bord postérieur, sans soies ou épines particulières. Opercule légèrement plus long que large, arrondi (0,22 à 0,24/0,18 à 0,19mm). Edéage (fig. 10). Long. 0,44 a 0,45mm. Parameres avec chacun quatre soies, deux en position dorsale et deux en position ventrale; paramere droit bien développé, élargi dans sa partie apicale; paramère gauche petit, gréle, subparallèle. Lobe interne bifide dans sa moitié apicale. Amauronyx caudatus sp. n. differe de toutes les autres especes du genre par la dent saillante de son pygidium; son édéage présente une structure assez semblable a celle de l Amauronyx cobosi Besuchet de I’ Andalousie, mais inversée. Namunia terricola sp. n. Holotype 4. GRECE, Eubée, 5km au sud-est de Kariotos, 30.03.1983, tamisage au pied d’un platane creux (Vit) (MHNG). Paratype. Méme provenance mais dans un tamisage de feuilles mortes, 1 2 (Vit) (MHNG). Long. 2,40mm. Proche de Namunia cavernicola Besuchet, 1978, dont il differe par sa taille bien plus faible, par ses antennes plus courtes (tous les articles moins allongés, spécialement les articles 9 et 10, réciproquement légèrement plus long que large et aussi long que large), par les élytres un peu plus courts, par la pubescence du métasternum plus longue et par les trochanters II du male inermes mais ornés sur le bord postérieur de soies assez longues et nombreuses. C’est la partie de l’Elytre courbée sur le côté qui porte la fossette latérale et la carénule latérale, cette dernière entière mais fine, totalement indistincte en vue dorsale. Les deux especes présentent de grosses fossettes profondes et tomenteuses sur le bord postérieur des hanches II et une tomen- tosité abondante a la base du 2° sternite apparent, caractères que je n’avais pas relevés dans ma révision (BESUCHET, 1978: 127) et qui sont nettement moins développés chez les autres especes du genre. Edéage (fig. 11). Long. 0,37mm (soies non comprises). Paramère droit un peu plus long que chez N. cavernicola; sac interne semblable, avec des centaines d’épines minuscules serrées. Quelques mensurations de holotype de Namunia terricola sp. n. Tête, lon- gueur/largeur: 0,49/0,45mm; lobe frontal, largeur: 0,28mm; scape: 0,20/0,09mm; pro- notum: 0,48/0,50mm; élytres: 0,63/0,82mm (longueur prise sur la suture); soies des élytres, longueur: 0,17-0,19mm; premier tergite apparent: 0,37/0,82mm; processus intercoxal, largeur apicale: 0,16mm; tibias III, longueur: 0,68mm. Bryaxis corsus sp. n. Holotype è homéomorphe. FRANCE, Corse: Haut Asco, 3.06.1971 (Senglet) (MHNG). Paratypes. Même provenance, 3d hom. 89 (Senglet) (MHNG); Haut Asco, 1500m, 10.07.1974, 1dhom. (Löbl) (MHNG); Haut Asco, 1450m, 17.08.1981, 4d hom. (Brachat) (Coll. Brachat): 1500m, 31.05.1986, 1 Shom. 52 (Brachat) (Coll. Brachat); Forêt de Bonifato, 700m, 04.1976, 1dhom. 12 (Deharveng) (MHNG): Forêt de Tartagine, 04.1976, 1 Soed. 16hom. 19 (Dehar- veng) (MHNG). 800 CLAUDE BESUCHET Long. 1,35 a 1,65mm. Coloration d’un brun rougeatre clair. Téte légèrement plus large que longue, le tégument lisse et brillant; lobe frontal subparallele, transverse, tronqué en avant, orné d’une dépression bien marquée; carène du vertex entiere, normalement développée, égale sur toute la longueur. Yeux plutöt petits, formés de 5 a 7 ommatidies chez la femelle, de 7 a 9 chez le male. Tempes arrondies en vue dorsale, distinctement anguleuses sous les yeux, cet angle prolongé par quelques soies. Palpes maxillaires normalement développés, avec quelques granules écrasés dans la partie apicale du 2° article; dernier article deux fois et demie plus long que large, la plus grande largeur située au tiers basal. Antennes a peine moins longues que la téte et le pronotum réunis; article 3 très nettement plus long que large, 4 et 5 égaux, un peu plus longs que larges, 6 et 7 sphériques, 8 un peu plus large que long; massue formée de trois articles progressivement élargis, les deux premiers transverses, le dernier aussi long que les quatre articles précédents réunis. Pronotum aussi long que large, le tégu- ment lisse, brillant, non ponctué; plus grande largeur située un peu en avant du milieu. Elytres réunis un peu plus larges que longs, la ponctuation légère, éparse; calus huméral effacé. Caractères sexuels de la femelle. Scape simple, cylindrique, deux fois plus long que large; pédicelle simple, une fois et demie plus long que large. Tibias I et III simples, gréles. Caractères sexuels des males. Scape (fig. 12) fortement renflé, orné au milieu de son bord interne d’un gros tubercule arrondi mais aplati dorso-ventralement; pédicelle (fig. 12) un peu plus long que large, assez largement rebordé sur son bord interne, l’angle apical saillant. Male homéomorphe avec les fémurs non renflés, les tibias I gréles mais nettement échancrés au tiers apical, les tibias III gréles, simples et légère- ment courbés près de l’apex. Male oedimère plus robuste, le pronotum légèrement plus large, les fémurs fortement renflés, les tibias I un peu plus robustes, aussi nettement échancrés au tiers apical; tibias III robustes, armés d’une petite dent saillante un peu en arriére du milieu du bord interne; partie apicale du tibia moins large, le bord interne légèrement concave, prrolongé a l’apex par un éperon bien distinct. Edéage (fig. 13) parfaitement symétrique. Long. 0,33 à 0,34mm chez les mâles homéomorphes, 0,37mm chez le male oedimere. Paramères parallèles dans la partie apicale chez les males homéomorphes, assez régulièrement atténués chez le mâle oedimére. Lobe interne gréle, subparallele, simple dans sa partie basale, bilobé dans sa partie apicale. Il y a des Bryaxis endémiques dans presque tous les massifs montagneux de la région métiterranéenne; ce genre n’était pourtant pas encore connu des montagnes de la Corse. Ce Bryaxis corsus sp. n. est apparenté au Bryaxis pedator (Reitter) de l’Italie moyenne par le scape antennaire du mâle, par les tibias postérieurs du mâle oedimère et par la structure générale de |’ édéage. Mais il en diffère par les yeux distinctement plus petits et par les tempes anguleuses dans les deux sexes, par le pédicelle antennaire du male fortement dilaté et assez largement rebordé sur le bord interne (fig. 12) (chez B. pedator, pédicelle petit, simple, semblable à celui de la femelle), enfin par l’édéage dont les paraméres sont inermes et le lobe interne moins franchement bilobé. PSELAPHIDES PALEARCTIQUES 801 Trissemus bellax sp. n. Holotype 6. ESPAGNE, Prov. CIUDAD REAL: Venta de Cardenas, 16.05.1960, dans le gravier au bord du Rio Magaña (Besuchet) (MHNG). Paratypes. Même provenance, 1d (MHNG). Prov. CIUDAD REAL: Ojos del Guadiana, 14.08.1969, 35 (Senglet) (MHNG). Prov. JAEN: Hinojarez, 19.07.1971, 1d (Senglet) (MHNG). Prov. ZARAGOZA: Caspe, 28.09.1966, dans un marais (Comellini) (MHNG). Long. 1,6 à 1,8mm. Coloration d’un brun rougeätre plus ou moins foncé. Tégu- ments de la face dorsale non ponctués, lisses et brillants, avec une pubescence couchée, formée de soies courtes. Tête un plus large que longue; face dorsale légèrement convexe, ornée de trois fossettes profondes, tomenteuses, la fossette frontale étant légè- rement plus grande; lobe frontal court, transverse, atténué d’arrière en avant; tempes subparallèles en arrière des yeux, puis atténuées et arrondies, un peu plus courtes que ceux-ci. Antennes de longueur moyenne; scape et pediceile nettement plus longs que larges; article 3 à 6 de même largeur, un peu moins larges que le pédicelle; 3, 5 et 6 nettement plus longs que larges, 4 légèrement ou un peu plus long que large. Pronotum un peu plus large que long, un peu plus large que la tête, la plus grande largeur située un peu en avant du milieu; côtés fortement arrondis, subparallèles près des angles postérieurs; fossette basale médiane petite et profonde; base du pronotum ornée sur toute la largeur de petits points serrés. Elytres réunis nettement plus larges que longs, deux fois plus larges que la tête, la plus grande largeur située un peu en avant de l’apex; callosité humérale assez bien marquée, arrondie; strie dorsale interrompue un peu avant le bord postérieur des élytres. Ailes bien développées. Premier tergite apparent (0,28- 0,31/0,70-0,75mm) légèrement atténué d’avant en arrière; carénules basales légèrement divergentes, courtes, marquées tout au plus jusqu’au tiers antérieur du tergite, séparées à la base par un espace (0,21-0,22mm) un peu inférieur au tiers de la largeur basale du segment. Caractères sexuels de la femelle. Articles 7 et 8 des antennes de même largeur que les articles précédents, 7 nettement un peu plus large que long, 8 transverse; massue triarticulée, progressivement élargie; article 9 transverse, 10 un peu plus large que long; dernier article relativement petit (0,15/0,10mm), la base obliquement tronquée. Caractères sexuels du mâle. Massue antennaire formée de cinq articles; articles 7 et 8 un peu plus larges que les articles précédents, 7 transverse, 8 très transverse; article 9 nettement plus large, très transverse, un peu déprimé sur la face ventrale; article 10 très grand (0,16-0,17/0,14mm), obliquement tronqué à l’apex, convexe sur la face dorsale, déprimé sur la face ventrale de la base au quart apical; cette dépression ornée de deux sillons longitudinaux profonds, contigus; quart apical ventral avec deux petites dents saillantes; dernier article petit (0,15-0,16/0,10-0,11mm) comme chez la femelle, la base aussi obliquement tronquée. Trochanters simples; hanches II armées chacune, a la base, d’une dent aiguë robuste; tibias II armés sur le bord interne, juste avant I’ apex, d’un éperon gréle pointu. Depression du métasternum peu profonde. Edéage (fig. 14). Long. 0,33 à 0,36mm. Paramères convergents fortement d’avant en arrière, prolongés chacun par une petite lame apicale subparallèle, ces deux lames contigués. Armature du sac interne formée d’une grande épine et d’une douzaine d’épines gréles séparées en deux faisceaux. 802 CLAUDE BESUCHET Fics 8-13 8: Faronus distinctus sp. n.. lobe de l'édéage portant les caractères spécifiques. face dorsale: 9: Faronus parallelus Besuchet, id.. face dorsale au même grossissement: 10: Amauronyx caudatus sp. n.. édéage. face dorsale: 11: Namunia terricola sp. n.. édéage. face dorsale: 12 et 13: Brvaxis corsus Sp. n.. base de l’antenne du mâle. face dorsale et édéage du male homéomorphe. face dorsale. PSELAPHIDES PALEARCTIQUES 803 Les Trissemus antennatus (Aubé), heterocerus (Aubé) et bellax sp. n. sont nettement apparentés par la grande massue antennaire des males mais cependant bien distincts les uns des autres par la structure de cette massue. Trissemus sulcifrons sp. n. Holotype 8. MAROC, environs de Rabat, 03.1963, dans des débris accumulés par une crue (Mussard) (MHNG). Long. 1,8mm. Coloration d’un brun noirätre, les élytres rougeatres. Espece tres proche et très semblable a Trissemus quedenfeldti (Reitter) dont elle diffère par les antennes plus courtes, par les caractéres sexuels du front et des antennes plus marqués et par l’édéage. Tête aussi longue que large, le lobe frontal très large (0,33mm), forte- ment transverse; tubercules antennaires aplatis; épistome rugueux; fossettes du vertex petites, profondes, tomenteuses, séparées par un espace convexe (0,19mm) presque double de celui qui se trouve entre le bord supérieur de |’ceil et la fossette voisine. Antennes un peu plus longues que la téte et le pronotum réunis; scape court, a peine plus long que large; pédicelle petit, légèrement plus long que large; article 3 long, modifié chez le male; articles 4 a 8 de méme largeur, un peu moins larges que le pédi- celle; 4 un peu plus long que large, 5 une fois et demie plus long que large, 6 légère- ment plus long que 4 mais moins long que 5; article 7 a peine plus long que large, 8 transverse; massue triarticulée, progressivement élargie; article 9 nettement plus large que long, 10 un peu plus large que long; dernier article (0,18/0,11mm) un peu asymé- trique, la face dorsale nettement plus convexe, la base obliquement tronquée. Premier tergite apparent (0,33/0,73mm) atténué d’avant en arrière; carénules basales nettement divergentes, marquées presque jusqu’au milieu du tergite, séparées a la base par un espace (0,10mm) égal à 1/7 de la largeur basale du segment. Caractères sexuels du male. Base du lobe frontal limitée par un sillon profond en arc d’accolade, la pointe dirigée en arrière, les extrémités atteignant presque le bord latéral; lobe frontal légèrement surélevé au milieu, orné près du sillon d’une très petite carène médiane partiellement masquée par une touffe de soies serrées. Antennes avec l’article 3 particulièrement grand (0,125/0,07mm), aussi long que les articles 4 et 5 réunis, un peu plus large que le pédicelle, distinctement élargi de la base a l’apex, la face externe nettement convexe, la face interne distinctement concave avec de très petits granules serrés. Tous les trochanters simples; tibias II prolongés a l’apex du bord interne par un éperon robuste. Dépression du métasternum peu profonde. Chez T. quedenfeldti, le sillon frontal est moins profond, simplement un peu courbé; l’article 3 des antennes est aussi très allongé mais à peine modifié. Edéage (fig. 15). Long. 0,36mm. Diffère de celui de 7. quedenfeldti (fig. 16) (0,43 a 0,48mm) par les paramères de largeur plus égale et les dents du sac interne plus courtes. Trissemus trilobatus sp. n. Holotype 5. PAKISTAN, Prov. Swat: Madyan, 1400m, 19.06.-4.07.1971, à la lumière (Holzschuh) (MHNG). Paratypes. Même provenance, 476 19 (MHNG), 504 (Coll. Holz- schuh). 804 CLAUDE BESUCHET Long. 1,55 a 1,7mm. Coloration entièrement d’un brun rougeätre peu foncé. Téguments de la face dorsale non ponctués, lisses et brillants, avec une pubescence couchée, formée de soies courtes. Téte légèrement plus large que longue; face dorsale un peu convexe, ornée de trois fossettes profondes, tomenteuses, égales; lobe frontal court, transverse, un peu atténué d’arriere en avant; tempes atténuées des le bord postérieur des yeux, arrondies, un peu plus courtes que ceux-ci. Antennes légèrement plus longues que la téte et le pronotum réunis chez le male, un peu plus courtes chez la femelle; scape une fois et demie plus long que large; pédicelle nettement plus long que large; articles 3 a 7 de méme largeur, un peu moins larges que le pédicelle; 3, 4 et 5 nettement plus longs que larges, 4 et 5 souvent très légèrement plus courts; article 6 légérement ou a peine plus long que large, 7 transverse. Pronotum un peu plus large que long, légèrement plus large que la tête, la plus grande largeur située au milieu; côtés fortement arrondis et atténués jusqu’a la base; fossette basale médiane petite, profonde; base du pronotum ornée sur toute la largeur de petits points serrés. Elytres réunis nettement plus larges que longs, presque deux fois plus larges que la tête, la plus grande largeur située un peu en avant de l’apex; callosité humérale assez bien marquée, arron- die; strie dorsale interrompue un peu avant le bord postérieur des élytres. Ailes bien développées. Premier tergite apparent (0,32-0,35/0,69-0,72mm) légérement atténué d’avant en arrière; carénules basales nettement divergentes, assez longues, dépassant légèrement le milieu du tergite, séparées a la base par un espace (0,17-0,19mm) Egal environ au quart de la largeur basale du segment. Caractéres sexuels de la femelle. Article 8 des antennes de méme largeur que les articles précédents, très transverse; massue triarticulée, progressivement élargie; articles 9 et 10 transverses; dernier article relativement petit (0,18/0,11mm), la base oblique- ment tronquée. Caractères sexuels du male. Article 8 des antennes légèrement plus large que les articles précédents, un peu prolongé du côté ventral; massue triarticulée (fig. 17); article 9 transverse, échancré au milieu de sa face ventrale; article 10 transverse, prolongé sur sa face ventrale par trois lobes allongés (fig. 18); dernier article assez grand (0,24- 0,26/0,15mm), la base formée, du còté ventral, par une lame portant un petit lobe arrondi de chaque còté et une lamelle trapézoidale au milieu; cette lame est immédia- tement suivie d’une cavité transverse très profonde qui se prolonge, sur la face ventrale de cet article 11, par une dépression progressivement atténuée en profondeur et en largeur. Trochanters II ornés, sur le bord postérieur, d’une épine très fine; tibias II armés sur le bord interne, un peu avant l’apex, d’un éperon très long (fig. 19), sub- parallèle et tronqué à son extrémité. Dépression du métasternum peu profonde. Edéage (fig. 20). Long. 0,40 à 0,43mm. Paramères élargis dans la partie apicale. Armature du sac interne formée de 12 à 14 épines assez grandes. Trissemus trilobatus sp. n. est proche du 7. akinini (Reitter) de |’ Asie centrale tant par la massue antennaire du mâle que par l’édéage, presque semblable; les tibias II portent aussi un éperon très long, tronqué chez trilobatus, atténué en pointe chez akinini. Ce dernier présente chez le male un net renflement des articles antennaires 4, 5 et 6 tandis que la massue du mâle de trilobatus est d’une complexité bien plus grande (fig. 17). PSELAPHIDES PALEARCTIQUES 805 Trissemus holzschuhi sp. n. Holotype 3. PAKISTAN, Prov. Swat: Madyan, 1400m, 19.06.-4.07.1971, à la lumière (Holzschuh) (MHNG). Long. 1,6mm. Coloration entierement d’un brun rougeätre peu foncé. Tégu- ments de la face dorsale non ponctués, lisses et brillants, avec une pubescence couchée formée de soies courtes. Tête à peine plus large que longue; face dorsale légèrement convexe, ornée de trois fossettes profondes, tomenteuses, la fossette frontale étant un peu plus grande; lobe frontal court, transverse, ses côtés presque parallèles; tempes atténuées dès le bord postérieur des yeux, arrondies, un peu plus courtes que ceux-ci. Antennes aussi longues que la tête et le pronotum réunis; scape et pédicelle nettement plus longs que larges; articles du funicule de même largeur, nettement moins larges que le pédicelle; article 3 presque deux fois plus long que large, 4 et 6 nettement plus longs que larges, 5 une fois et demie plus long que large, 7 légèrement plus long que large, 8 distinctement plus large que long; massue triarticulée, progressivement élargie; article 9 un peu plus large seulement que les articles précédents, un peu plus large que long, 10 nettement plus large que long; dernier article relativement petit (0,15/0,09mm), un peu plus convexe sur la face dorsale. Pronotum un peu plus large que long, un peu plus large que la tête, la plus grande largeur située un peu en avant du milieu; côtés fortement arrondis, subparallèles près des angles postérieurs; fossette basale médiane petite, profonde; base du pronotum ornée sur toute la largeur de petits points serrés. Elytres réunis nettement plus larges que longs, presque deux fois plus larges que la tête, la plus grande largeur située un peu en avant de l’apex; callosité humérale assez bien marquée, arrondie; strie dorsale distincte presque jusqu’au bord postérieur des élytres. Ailes bien développées. Premier tergite apparent (0,28/0,66mm) légèrement atténué d’avant en arrière; carénules basales à peine divergentes, assez longues (0,16mm), marquées jusqu’au tiers postérieur du tergite, largement séparées (0,26mm) par un espace égal aux 2/5 de la largeur basale du segment. Caractères sexuels du mâle. Antennes non modifiées. Trochanters II légèrement anguleux sur la base du bord postérieur; tibias II prolongés sur le bord interne par un éperon robuste, tibias III par un petit éperon gréle. Disque du métasternum occupé par une dépression arrondie très profonde. Edéage (fig. 21). Long. 0,34mm. Paramères complètement soudés dans la partie médiane, courts mais bien séparés dans la partie apicale. Armature du sac interne complexe, avec en position dorsale un faisceau de nombreuses soies très longues et en position ventrale deux slérites de chaque côté. Trissemus holzschuhi sp. n. n’a pas d’affinités directes avec les autres espèces paléarctiques du genre. Il est bien caractérisé par les carénules du premier tergite apparent largement séparées et par l’édéage dont les paramères sont complètement soudés dans la partie médiane et l’armature du sac interne formée de soies nombreuses. D’après JEANNEL (1959:529), les Trissemus paléarctiques se répartissent dans trois sous-genres: Trissemus Jeannel s. str. pour les T. antennatus (Aubé) et heterocerus (Aubé); Trissemellus Jeannel pour les T. maroccanus (Raffray), arabicus (Raffray), brittoni Jeannel; Trissemosus Jeannel pour les T. niloticus (Motschulsky) et tetuanicus 806 CLAUDE BESUCHET Fics 14-21 Trissemus. 14: T. bellax sp. n., édéage, face dorsale; 15: T. sulcifrons sp. n., édéage, face dorsale; 16: T. quedenfeldti (Reitter), édéage, face dorsale; 17: T. trilobatus sp. n., extrémité de l’antenne du mâle, face latérale; 18: Id., article 10 de l’antenne du mâle, vue de bout; 19: Id., extrémité du tibia intermédiaire du male; 20: Id., édéage, face dorsale; 21: T. holzschuhi sp. n., édéage, face dorsale. PSELAPHIDES PALEARCTIQUES 807 Jeannel, auxquels il faut encore ajouter Trissemites Jeannel pour T. militaris (Saulcy) d’après son tableau d’identification. Celui-ci est basé principalement sur l’écartement plus ou moins grand des carénules basales du premier tergite apparent, la localisation des caractères sexuels sur les antennes et quelques structures de I’ édéage. Le sous-genre Trissemus s. str. est bien défini, selon JEANNEL, par la grande massue antennaire du mâle et par l’édéage avec les parameres soudés. Trissemus bellax sp. n. présente une massue antennaire assez semblable a celle de 7. antennatus (Aubé), espèce-type du genre 7rissemus; mais ses parameres (fig. 14) sont indépendants, contigus seulement dans la partie tout a fait apicale. Toujours d’après JEANNEL, le sous- genre Trissemites présente des carénules distantes du quart ou du cinquieme de la largeur du tergite, avec des antennes semblables dans les deux sexes. C’est bien le cas pour 7. militaris (Saulcy). Mais que penser du 7. trilobatus sp. n., Trissemites par ses carénules du premier tergite apparent, mais totalement différent par la massue anten- naire du male (fig. 17)? Enfin 7. holzschuhi sp. n., d’apres ce méme tableau d’iden- tification, serait un Trissemellus, mais dont les paramères de l’édéage sont soudés (fig. 21). La valeur taxonomique des caractères retenus par JEANNEL pour les Trissemus me paraît bien discutable. L’écartement des carénules de ce premier tergite apparent est sujet à une certaine variation intraspécifique; il y a d’autre part, à l’intérieur du genre, tous les intermédiaires entre les écartements minimum (1/7) et maximum (2/5). Quant aux caractères sexuels des antennes, ils peuvent concerner, suivant les espèces, tous les articles (sauf le scape), un ou plusieurs a la fois. L’édéage présente normalement deux parameres bien séparés, indépendants; mais ceux-ci peuvent étre exceptionnellement contigus ou soudés, sur des longueurs plus ou moins grandes. Les caracteres appliqués pour les sous-genres des Trissemus sont par trop variables et arbitraires; ils sont sans intérét pour les espèces paléarctiques et ne peuvent pas étre retenus. Par conséquent je transfère toutes les espèces paléarctiques, y compris celles du Japon, dans le sous-genre Trissemus s. str. NOUVEAUX SYNONYMES ET TRANSFERTS Rybaxis marquardti Reitter (1900: 50. Syntype d ici désigné comme lectotype: Mus. Paris. Loc. typ.: Ouzbékistan: Buchara) n’est qu’un synonyme de Rybaxis longi- cornis (Leach, 1817) (syn. nov.). Ce lectotype appartient à la forme d longicornis S. str. Pselaphus ruber Beck (1817: 10, pl. II, fig. 7. Types perdus. Loc. typ.: Allemagne, Baviere) est d’apres la description un synonyme de Rybaxis longicornis (Leach, 1817) (syn. nov.). Pselaphus castaneus Beck (1817: 11, pl. II, fig. 9. Types perdus. Loc. typ.: Allemagne, Baviere) est d’aprés la description un synonyme de Brachygluta fossulata (Reichenbach, 1816) (syn. nov.). Trissemus sahlbergi Karaman (1963: 79, fig. 1, 2 et 3. Holotype $: Mus. Budapest. Loc. typ.: Palestine, Jourdain) n’est qu’un synonyme de Trissemus dentipes (Baudi, 1869) (syn. nov.). Je connais cette espece de Chypre, du sud-est de la Turquie, d'Israël et de Jordanie. 808 CLAUDE BESUCHET Le genre Kunzea Leach (1826: 448) est souvent cité dans les catalogues et les bibliographies comme synonyme de Bryaxis Kugelann, 1794 = Arcopagus Leach, 1817 = Bythinus auct.; mais l’espece-type nigriceps Leach (1826: 449) a toujours été oubliée par les auteurs. Les diagnoses données en latin et en anglais sont pourtant assez précises, en particulier pour les antennes; d’autre part l’espece a été découverte dans une forét de pins des Alpes-Maritimes frangaises. Seul Bryaxis latebrosus (Reitter), pas rare dans le sud-est de la France et en Ligurie occidentale, correspond aux descriptions de Leach: “Antennae with their first and second joints thicker than the others; the first elongato-cylindric, internally abruptly dilated; the second minutely globose, narrower than the first one”... “Corpore toto ferruginea”. D’autre part on peut éliminer les espèces décrites précédemment par LEACH (1817: 83), a savoir “Bythinus securiger”, “Bythinus Curtis”, “Arcopagus glabri- collis”, “Arcopagus clavicornis” et “Arcopagus bulbifer”, de même “Arcopagus rugi- collis”, décrit en 1826. Kunzea nigriceps Leach (1826: 449. Types perdus semble-t-il. Loc. typ.: France, Alpes-Maritimes) et Bythinus latebrosus Reitter (1884: 72. Syntypes d : Mus. Paris. Loc. Typ.: France, Alpes-Maritimes: Nice) sont synonymes (syn. nov.) dans le genre Bryaxis Kugelann. Linderia (s. str.) leleupi Jeannel (1952b: 292). Holotype 9: Inst. Bruxelles. Loc. typ.: France, Ariege: Saint-Girons) appartient en réalité au genre Bryaxis Kugelann (comb. nov.) et plus précisément au groupe de B. fauconneti (Fauvel). Coiffait a retrouvé cette espèce dans différentes localités de l’Ariège: les femelles sont tout à fait semblables a holotype de B. leleupi; les males sont fortement oedimères, avec des yeux formes d’une dizaine d’ommatidies et un scape subcylindrique deux fois et demie plus long que large, simple; pédicelle un peu plus long que large, simple. Macrobythus klimeschi W. & C. Blattny (1914: 175. Lectotype d : Mus. Bâle, Coll. Frey. Loc. typ.: Croatie, Dalmatie centrale, Svilaja Planina) appartient indis- cutablement au genre Bryaxis Kugelann (comb. nov.). Il est bien difficile de préciser les affinités de cette espece endogée, car ce lectotype, étiqueté en 1956 par un M. Kamp., n’a pas d’édéage. Je juge utile de compléter la description des frères Blattny par quelques mensurations. Long. 1,70mm. Yeux formés chacun de 3 a 4 ommatidies dépigmentées, saillantes. Téte: 0,33/0,35mm, yeux compris; largeur du lobe frontal: 0,20mm. Pro- notum: 0,41/0,43mm. Elytres réunis: 0,66/0,73mm. Antennes, long. 0,94mm; scape: 0,25/0,08mm, légèrement élargi de la base a l’apex; pédicelle: 0,08/0,06mm; article 11: 0,19/0,10mm. Dernier article des palpes maxillaires: 0,30/0,10mm. Tibias III gréles: 0,62/0,05mm. Carène du vertex entière, bien marquée sur toute sa longueur, légèrement surélevée en avant et en arrière. Bythinus banaticus Reitter (1884: 74. Syntype d ici désigné comme lectotype: Mus. Paris. Loc. typ.: Roumanie, Mehadia) est effectivement un synonyme de Bythinus hopfgarteni Reitter, 1881, comme l’a montré KARAMAN (1948: 11, 16). Mais la varia- bilité du 2° article antennaire des males est mal interprétée. B. hopfgarteni et banaticus ont quasiment le même pédicelle, correspondant à la fig. li de KARAMAN (“B. hopf. ab. rectangulus 3”); la fig. 1h (“B. hopf. ab. banaticus 3”) et la fig. 1g (“B. hopfgarteni PSELAPHIDES PALEARCTIQUES 809 3”) représentent d’autres stades de cette variabilité, toujours fréquente chez les mâles des Bythinus balkaniques. Tychus fournieri Saulcy (1864: 259. Syntypes: Mus. Paris. Loc. typ.: France, Var: Draguignan) a été déplacé par JEANNEL (1960: 344) dans le genre Tychomorphus Jeannel. Il appartient en réalité au genre Tychus Leach (comb. nov.) où il est apparenté à Tychus aretinus Dodero. Bryaxis opuntiae Schmidt-Göbel (1836: 31, pl. I fig. 17. Type d perdu. Loc. typ.: Italie, Sicile). MACHULKA (1930: 120) a fait remarquer le premier que ce Bryaxis opuntiae de Sicile est un Tychini et que le Bryaxis opuntiae sensu AUBE (1844: 115) est identique au Reichenbachia opuntiae auct., aujourd’hui Trissemus olivieri (Raffray). SCHMIDT-GOBEL (1836) a décrit deux Tychini de Sicile, l’un au front étroit (Tychus dichrous, p. 18, fig. 16), l’autre au front large (Bryaxis opuntiae, p. 31, fig. 17); les types sont perdus pour les deux espèces. Les Psélaphides de la collection Schmidt- Gobel ne se trouvent pas au musée de Prague; B. opuntiae, découvert en Sicile par Helfer, n’existe pas dans la collection de celui-ci, conservée au musée de Prague (communication du Dr J. Jelinek), mi d’ailleurs dans la collection Motschulsky a Moscou, ou se trouvent aussi des Psélaphides de Helfer. La faune des Psélaphides de la Sicile est maintenant bien connue grace a SABELLA (1998); celui-ci fait d’ailleurs correctement remarquer que les Tychomorphus qui peuplent cette île, 7. jacquelini (Boieldieu, 1859) et integer (Reitter, 1881), pourraient correspondre a Bryaxis opuntiae Schmidt-Göbel (SABELLA, 1998: 337); mais il n’a pris aucune decision a ce sujet. Toujours d’apres SABELLA (1998: 209), T. integer est plus abondant (Fig. 121), et compte tenu des grands ports ou arrivaient jadis les entomologistes, a une probabilité plus grande d’étre le Bryaxis opuntiae Schmidt- Göbel. D’apres la figure originale 17f, la massue antennaire est relativement grande, caractere qui milite en faveur de 7. integer. Pour la stabilité de la nomenclature et pour résoudre ce probleme d’identite, je designe pour néotype de Bryaxis opuntiae Schmidt- Göbel un male de Tychomorphus integer (Reitter) de Palerme, Sicile (MHNG) (syn. nov.). Ce néotype porte une petite étiquette manuscrite “Palermo Si.” de 7 sur 3mm. REMERCIEMENTS Mes remerciements vont à MM. Ivan Löbl pour ses remarques et suggestions judicieuses, Stanislav Vit pour la tres belle figure 7, Carolus Holzschuh de Vienne et Jean-Pierre Thelot de Forcalqueiret-La Roquebrussanne pour le don de trois belles especes nouvelles. Je tiens aussi a remercier les collegues qui m’ont communiqué pour étude des Psélaphides des collections entomologiques dont ils sont responsables: Mme Eva Sprecher du musée de Bâle (Coll. Frey), MM. Georges Coulon de l’Institut royal des Sciences naturelles de Belgique et Otto Merkl du musée de Budapest; ou de leur collection personnelle: MM. Volker Brachat de Geretsried et Guido Nonveiller de Zemun. 810 CLAUDE BESUCHET BIBLIOGRAPHIE AUBE, C. 1844. Revision de la famille des Psélaphiens. Annales de la Société Entomologique de France, 2° série, II: 73-160, pl. 3. BECK, L. von 1817. Beiträge zur baierischen Insektenfaune, oder Beschreibung und Abbildung neuentdeckter Käfer. Augsburg, 47 pp., 7 pl. BESUCHET, C. 1956. Revision des genres Zibus, Saulcyella, Aphiliops et description d’un genre nouveau (Col. Pselaphidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 29: 363-372. BESUCHET, C. 1958. Coleoptera Pselaphidae et Scydmaenidae. Revue suisse de Zoologie 65: 891- 919: BESUCHET, C. 1960. Coléopteres Psélaphides de la collection J. Sahlberg. Annales Entomologici Fennici 26: 11-31. BESUCHET, C. 1978. Le genre Namunia Reitt. (Coleoptera, Pselaphidae). Revue suisse de Zoo- logie 85: 127-133. BLATTNY, W. et C. 1914. Beitrag zur Kenntnis der Pselaphiden und Scydmaeniden. Cole- opterologische Rundschau 3: 175-178. JEANNEL, R. 1950. Coléoptères Psélaphides. Faune de France 53: II + 421 pp. JEANNEL, R. 1952a. Exploration du Parc National de l’Upemba. Pselaphidae. Institut des Parcs Nationaux du Congo Belge, fasc. 13: 3-61. JEANNEL, R. 1952b. Sur quelques Pselaphides de France. Bulletin et Annales de la Societe Entomologique de Belgique 88: 291-294. JEANNEL, R. 1956a. Les Psélaphides de l’Afrique du Nord. Mémoires du Museum National d’Histoire Naturelle, Paris, n.s. (Série A: Zoologie) 14: 1-233. JEANNEL, R. 1956b. Sur les genres Dimerus Fiori et Octomicrus Schaufuss (Coleoptera Psela- phidae). Revue Francaise d’Entomologie 23: 84-100. JEANNEL, R. 1959. Révision des Psélaphides de I’ Afrique intertropicale. Annales du Musée Royal du Congo Belge, Tervuren (Série in 8°, Sciences Zoologiques) 75: 1-742. KARAMAN, Z. 1948. Revizija Pselaphida (Kol.). Zagreb: Jugoslavenska Akademija Znanosti i Umjetnosti, 19 pp. KARAMAN, Z. 1963. Nouvelles espèces de Trissemus et de Reichenbachia (Col. Psel.). Bulletin de la Société Entomologique de Mulhouse 1963: 79-81. LEACH, W. E. 1817. On the stirpes and genera composing the family Pselaphidea; with the names of the British species. Zoological Miscellany 3: 80-87. LEACH, W. E. 1826. On the stirpes and genera composing the family Pselaphidae; with descrip- tions of some new species. Zoological Journal 2: 445-453. MACHULKA, V. 1930 (1929). Beitrag zur Synonymie der palaearktischen Pselaphiden (Coleop- tera). Clasopis Cleskoslovenské Spoleclnosti Entomologické 26: 119-121. NEWTON, A. F. Jr. & CHANDLER, D. S. 1989. World Catalog of the Genera of Pselaphidae (Coleoptera). Fieldiana, Zoology, New Series 53: II + 93 pp. Park, O. 1942. A Study in neotropical Pselaphidae. Northwestern University Studies in the Biological Sciences and Medicine 1: 1-403, 21 pl. RAFFRAY, A. 1903-1904. Genera et Catalogue des Psélaphides. Annales de la Société Entomo- logique de France 72: 484-604; 73: 1-476, 635-678, 3 pl. RAFFRAY, A. 1908. Coleoptera. Fam. Pselaphidae. In: WYTSMAN, P. (ed.): Genera Insectorum, fasc. 64: 487 pp., 9 pl. REITTER, E. 1884. Bestimmungs-Tabellen der europäischen Coleopteren. X. Nachtrag zu dem V. Theile, enthaltend: Clavigeridae, Pselaphidae und Scydmaenidae. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 34: 59-94. REITTER, E. 1900. Beitrag zur Coleopteren-Fauna des russischen Reiches. Deutsche Entomo- PSELAPHIDES PALEARCTIQUES 811 logische Zeitschrift 1900: 49-59. SABELLA, G. 1998. Pselafidi di Sicilia. Museo Regionale di Scienze Naturali Torino; Monografie 25:415 pp. SAULCY, F. H. C. DE 1864. Faune française et européenne. Descriptions et remarques. Annales de la Société Entomologique de France, 4° série, 4: 253-260. SCHMIDT-GOBEL, H. M. 1836. Dissertatio inauguralis zoologica de Pselaphis faunae pragensis cum anatomia Clavigeri. Pragae: Filiuorum Theophili Haase, IV et 50 pp., 2 pl. ait GA rl 1 ; REVUE SUISSE DE ZOOLOGIE 106 (4): 813-904; décembre 1999 Androdeloscia gen. n., a new genus of South American terrestrial isopods with description of 13 new species (Crustacea: Oniscidea: ''Philosciidae"') Andreas LEISTIKOW Universitat Bielefeld, Abteilung fiir Zoomorphologie und Systematik, Morgenbreede 45, D-33615 Bielefeld / Ruhr-Universität Bochum, Lehrstuhl für spezielle Zoologie, Universitätsstraße 150, D-44780 Bochum, Germany; e-mail: Leiste@Biologie.Uni-Bielefeld.de Androdeloscia gen. n., a new genus of South American terrestrial isopods with description of 13 new species (Crustacea: Oniscidea: ‘“Philosciidae”). - A new genus of philosciid Oniscidea is described tor the Venezuelan species Prosekia hamigera (Vandel, 1952) and its Brazilian congener P. silvatica Lemos de Castro & Souza, 1986. They differ remar- kably from Prosekia rutilans (Vandel, 1952) and are linked by several synapomorphies with other genera of small philosciids from South America, particularly with Erophiloscia Vandel, 1972 and Andenoniscus Verhoeff, 1941. Several new species of this genus have been found in the collections of Dr W. Hanagarth in Peru, which are now courtesy of the Staatliches Museum fiir Naturkunde, Stuttgart (SMNS) and in the collec- tions undertaken by Dr C. Schmidt (Ruhr-Universität) in Venezuela in spring 1998. A diagnosis of the new genus is given, all the species included are described in detail, the inter- and infrageneric phylogenetic relationships are discussed. A key to the species is given. Key-words: Isopoda - Oniscidea - South America - biogeography - phylogeny - taxonomy. INTRODUCTION There are some species of Oniscidea from South America, which had been described as belonging to the genus Chaetophiloscia Verhoeff, 1908, a genus with linea frontalis reduced and long noduli laterales (VERHOEFF 1908), the generic diagnosis was completed by SCHMALFUSS (1990). These species were separated in the genus Prosekia Vandel, 1968 mainly due to some biogeographic considerations; VANDEL (1968) stated, that Chaetophiloscia is a genus of eastern mediterranean distri- bution widely absent from southern Spain and Portugal. A wide gap between the two distributional areas, circum-mediterranean and northern South America would result even when continents were transformed to their position in early Cretaceous. Addi- Manuscript accepted 06.04.1999 814 ANDREAS LEISTIKOW tionaly, he gave some characters for differentiation (VANDEL 1968, p. 118): the shape of the antennula and the length of the noduli laterales. Whereas the biogeographical argument is difficult to follow, the remarkable differences in morphology justify the separation of Prosekia as a genus on its own. Unfortunately, the members of Prosekia showed up to be a polyphyletic assembly. To reduce this genus to a monophyletic taxon, a re-examination of all its members is attempted. The first results lead to the separation of Prosekia hamigera (Vandel, 1952) and P. silvatica Lemos de Castro & Souza, 1986 into a new genus Androdeloscia gen. n.. They differ in many characters from P. rutilans (Vandel, 1952) which was the first species described of those belonging to this genus (VANDEL 1952) and should considered as type of this genus. The examination of a collection of Oniscidea performed in spring 1998 in Venezuela by Dr C. Schmidt (Ruhr-Universität-Bochum) revealed the presence of additional three species of Androdeloscia gen. n. in this region; another important collection of Oniscidea was made by Dr W. Hanagarth in Peru. The material was first examined by Dr H. Schmalfuss from the Staatliches Museum fiir Naturkunde, Stuttgart. He described three species of the philosciid genus /schioscia Verhoeff, 1928 (SCHMALFUSS 1980) and gave a summary of the material. The other material remains undescribed until now. In this collection, members of Androdeloscia gen. n. were plentiful, so that 10 new species can be distinguished. Within this work, the systematic section is followed by a paragraph on the phylogeny and biogeography of this interesting new genus. The following acronyms are used: SMNS Staatliches Museum für Naturkunde, Stuttgart USNM United States National Museum, Smithonian Institution, Washington MNHG Muséum d'histoire naturelle, Genève MUMV Museo de la Universidad de Maracay, Venezuela SYSTEMATIC SECTION Androdeloscia gen. n. Diagnosis: Cephalothorax with linea supra-antennalis, linea frontalis in most species reduced, small lateral lobes, compoud eyes consisting of about 6 ommatidia. Antennula with divergent setal tufts on distal article (fig. 64), antenna with three- articulated flagellum bearing a long apical organ. Mandible with molar penicil composed of 3 to 6 branches, maxilla with lateral endite bearing 4+6 or 4+5 teeth, hyaline lobe rarely present, maxilla with lateral lobe twice as broad as medial lobe, maxillipedal endite with small knob-like penicil. Pereopods slender, with small antenna-grooming brush on carpus 1, orna- mental sensory spine double-fringed serrate, carpus with mediodistal setal tuft, coxal plates bearing flagelliform nodulus lateralis (fig. 64), sulcus marginalis present to rudimentary, nodulus lateralis IV more dorsally inserted than others. Pleopods without respiratory area, exopodites rhomboid but exopodite 5, which is triangular, male exopodite 5 with straight medial margin, bearing caudally A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 815 guide slot for flagelliform endopodite 2 (fig. 64). Male exopodite 1 rounded, endopodite bearing tubercles, in some species hyaline lamellae. Genital papilla stout, with elliptic ventral shield slightly surpassed by ductus ejaculatori, apically obtuse. Uropod with triangular protopodite laterally grooved, endopodite inserting proximally of exopodite. Type species: Chaetophiloscia hamigera Vandel, 1952. Comments: As VANDEL (1968) stated, the genus Prosekia is distinguished by the shape of the antennula and the complexity of the male pleopod endopodites. A thoroughly revision of the type material of Andenoniscus silvaticus Verhoeff, 1941 revealed the presence of this type of antennula also in this genus (LEISTIKOW 1998). Thus, it may be a character of a larger monophylum - provisorically called the Pre- sekia-group. The new genus is similar to Andenoniscus Verhoeff, 1941 and Erophi- loscia Vandel, 1972 in the overall habitus, tiny species of maximal 5mm with slender pleon and rounded telson. In contrast to the above mentioned species, a linea frontalis is reduced in Androdeloscia gen. n.; another autapomorphy is the shape of the male pleopod 5 exopodite, which is apically drawn out to some extend. An even more complex character is a furrow parallel to the medial margin of the caudal surface, covered with pectinate scales. This furrow works as a guide slot for the flagelliform pleopod 2 endopodite, fitting so tight that it is sometimes impossible to remove the endopodite from the furrow without damaging it. A furrow of similar structure and function has been evolved several times within the Crinocheta, it can be found in Chaetophiloscia and Philoscia Latreille, 1804 as well (pers. obs.). The details of this structure and the lack in its closest relatives may warrant the autapomorphic status. A medially drawn out pleopod 5 exopodite in the male can be found in Erophiloscia, too (VANDEL 1972). In contrast to the new genus, there is no trace of a guide slot for the pleopod 2 endopodite. Most probably, Erophiloscia is the adelphotaxon of Andro- deloscia gen. n. This will be dealt with in another contribution to these genera. Fur- thermore, several species of Prosekia described from the Amazon region (LEMOS DE CASTRO & SOUZA 1986; LIMA 1997) have to be transferred to Androdeloscia. This is the result of a re-examination of the type material from the Museo Nacional, Rio de Janeiro. Etymology: The genus name refers to the complex male copulatory devices of the pleopods: "andros" greek for "male", "del-" is abbreviated from greek "delos" for obvious/visible, whereas "oscia" is a suffix commonly used for Crinocheta with philosciid facies. KEY TO THE SPECIES I Malesiwath) percopod /*beanneobes On IMEUS wer ee: 10 = Males with pereopod 7 without meral differentiation.................... 2 2 Fields of cuticular scales ventrally on male pereopod 4 to 6 . . . . A. taitii sp. n. - INOFSUCHEMELASTOMPETCOPOASME RA RE I RIA EN STAN CIS 5 3 Male pleopod 1 exopodite comparatively big, more or less circular SIC. D RER SNe RMN Ne LE © A URSS LE PER ET EVA À. opercularis Sp. n. 816 ANDREAS LEISTIKOW Male pleopod | exopodite rounded but smaller than half the length of endopodite A354 54!laatk oa «LL eevee eis el ae e 4 Male pleopod | endopodite slender, apex directed laterally, with small knobs and saddle-shaped lobe, without prominent protrusions, distal sensory spine of propus | with double-serrate fringe ......... A. dalensi sp.n. Pleopod 1 endopodite of different shape, distal sensory spine of propus lswathvoneito)three subapical points PA kt Jee saan sa 5 Male pleopod 1 endopodite with prominent protrusions ................. 6 Male pleopod | endopodite without protrusions, apex with small knobs, lookines firrcone-like? .... u... no ee oan) ne 8 Protrusion on endopodite directed laterally, sickle-shaped...... A. hamigera Male pleopod 1 endopodite with two small protrusions .................. 7 Apex of endopodite finger-shaped, protrusions on halflength A. poeppigi sp. n. Apex stout, hammer-shaped due to distal protrusions ....... A. malleus sp. n. Male pleopod 1 endopodite stout with broad base, pleopod 5 exopodite PALCHI Ary FA WI UE as. See aia I RN A. conipus sp. n. Male pleopod | endopodite with almost equally-sided triangular base . ..... 9 Male endopodite | distinctly bent laterally in last third . . . . A. plicatipus sp. n. Malefendopoditeziàmore Omless:stralohtenmias 1. 6. 0er A. feistae sp. n. Merusiwithione lobe or. protrusion: PAPE RO IRR 11 Merusiwithimorethanone lobe On protrusion. 2.22... . 2 er ee EE 14 Protiusionion menus mediodistally. ee 2. 2 ne 13 Protrusion on merus medially, more or less on halflength ............... 12 Pleopod 1 endopodite apically bulbous with flagelliform tip; short inner CLAW AOL MENCODOGS ois seat Masala mue VII A. ferrarai sp. n. Pleopod 1 endopodite apically bulbous, obtuse, appearing spiny; inner claw almost as long as interungual seta............... A. longiunguis sp.n. Protrusion finger-shaped, directed distally, with sensory spine apically . RENNER Wd In re Guat aan este elle Claes ATEN PS A. digitata sp. n. Protrusion small lobe, pleopod | endopodite with flagelliform tip SN doe een EE RN RER REES Re A. merolobata sp. n. Male pleopod 1 endopodite with distally directed protrusion of mus- CUAT PALI 2 MESURE I MISERI UAT get ER ee eer RIONE ET A. silvatica Male pleopod 1 without such a protrusion .......... A. pseudosilvatica sp. n. SPECIES ACCOUNT Androdeloscia hamigera (Vandel, 1952) comb. n. Chaetophiloscia hamigera Vandel, 1952 Prosekia hamigera (Vandel): VANDEL 1968 Material: Venezuela: 23 (max. 3.5mm) 22 (ovigerous) Caripe, 10°10.51'N 63°30. 21'W, garden with palms, orchids, Poaceae, ferns, under flower pots, leg. 07.04.1998 C. Schmidt, MHNG coll.; 18 22 (ovigerous) same data, MUMV coll.; 98 72 (ovigerous) 99 6imm, same data, author's coll.; 25 39 (ovigerous) 62 2imm, Caripe, surroundings of Cueva del Guacharo, 10°35.94'N 63°11.81'W, moist forest, under rotting logs and leaf litter near A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 817 brook, leg. 07.04.1998 C. Schmidt, author's coll.; 98 1d 119% 9imm, Peninsula de Paria, Puy Puy, 10°42.00'N 62°58.05'W, bay with sandy beach, coconut palms, surrounded by dry mountains, swamp near beach, under logs and leaf litter, leg. 30.03.1998 C. Schmidt, author's coll. Colour: Colour pattern similar to A. dalensi sp. n. with light spots in medial line of pereon, pleon unmarked. Cephalothorax: Linea supra-antennalis straight, lateral lobes small, lamina frontalis present, linea frontalis lacking, compound eyes consisting of about 8 ommatidia (pl.1, Ctf). Pereon: Tegument smooth and shiny, bearing scattered tricorn-like setae, coxal plates without gland pores, sulcus marginalis poorly individualized, noduli laterales flagelliform, on coxal plate IV more dorsally inserted (fig. 1, Cx3/Cxp). Pleon: Set apart from pereon, neopleurae of pleonites 3 to 5 rather short, pleotelson with rounded apex, laterally straight, bearing few tricorn-like setae. Appendages: Antennula: Three-articulate, rather slender with distal article bearing prominent tuft of aesthetascs medially and 2 aesthetascs apically (fig. 1, Anl). Antenna: Comparatively slender, length ratio of peduncular articles 1 to 5 1: 2: 2: 4: 5, flagellum three-articulate with articles subequal in length, distal article slightly longer, apical organ half as long as flagellum (fig. 1, An2). Mandible: Molar penicil dichotomized, composed of about seven branches, pars intermedia with two penicils on left and one on right mandible, additional plumose seta more proximally (fig. 2, Mdl/r). Maxillula: Medial endite with two pointed penicils, apical tip medially, lateral endite with apically 3+5 teeth, four of inner set cleft, laterally fringed (fig. 2, Mx1). Maxilla: Lateral lobe almost twice as broad as medial one, bearing pectinate scales, medial endite bearing some cusps apically (fig. 2, Mx2). Maxilliped: Basipodite with sulcus lateralis, palp with one seta on proximal article, two setal tufts on distal articles, endite with two teeth caudally, knob-like penicil smooth (pl.2, Mxp). Pereopods: Rather slender, especially pereopod 1, carpus with antenna- grooming brush and serrate ornamental sensory spine (fig. 3, Sc1), dactylus with short inner claw and simple dactylar seta (fig. 3, Dac). Sexual differentiation: Pereopods 2 to 4 with fields of proximally directed’ cuticular setae medially on carpus and merus (fig. 3, PE4). Pleopods: Similar to other members of the genus (fig. 3, PL1-5). Sexual differentiation: Male pleopod exopodite circular, endopodite with falciform protrusion laterally on halflength, apex obtuse, with hyaline lamellae and hyaline lobe, caudal row of sensory spine only on distal third of pleopod. Pleopod 2 similar to next species, pleopod 5 exopodite with medial slot guide, distally pro- truding. Uropod and genital papilla: As in generic diagnosis. Comments: As in most of the species of Androdeloscia gen. n., the species A. hamigera (Vandel, 1952) is best distinguished by the shape of the male pleopod 818 ANDREAS LEISTIKOW 1 endopodite. The lateral sickle-shaped protrusion is found in no other species, thus being the best character to separate the species. First described as belonging to the genus Chaetophiloscia Verhoeff, 1908, the species was grouped with some other South American species of Chaetophiloscia in a new genus Prosekia Vandel, 1968 mainly for biogeographic reasons (VANDEL 1968). Since it differs remarkably from Prosekia rutilans (Vandel, 1952) it is now part of the monophyletic group Andro- deloscia from tropical South America. Prosekia rutilans has big compound eyes, a faint linea frontalis and structurally different male pleopods. E. g., pleopod 2 endopodite is short with a club-like apex, while in all members of Androdeloscia the endopodite is slender with a more or less flagelliform apex. A discussion of the characters of P. rutilans will be given elsewhere. Androdeloscia conipus sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE d 4mm "A2", 1.X11.1975 leg. W. Hanagarth, SMNS coll. T446; PARATYPES 46 59, same data as holotype, SMNS coll. T447. Colour: Dorsally brown with several white markings on the tergites, medially forming a white band which is accompanied by two paramedian rows of prominent white spots on pereonites | to 4, pleon with some light spots, cephalothorax dorsally densely marked white. Cephalothorax: No linea and lamina frontalis, linea supra-antennalis and small lateral lobes present, profrons between compound eyes slightly bulbous (fig. 4, Ctf). Pereon: Smooth and shiny tegument, coxal plates with sulcus marginalis along lateral border, no gland pores, insertion of nodulus lateralis bordered by concentric structure of the cuticle, noduli long and flagelliform, maximum of d/c-coordinates on coxal plate IV (fig. 4, Cx4/Cxp). Pleon: Retracted from pereon, neopleurae of pleonites 3 to 5 conspicuous, pleotelson with straight margins, bearing some prominent tricorn-like setae. Appendages: Antennula: Three-articulate, medial tuft consisiting of about 10 aesthetascs (fig. 4, Anl). Antenna: Rather stout, length ratio of peduncular articles 1 to 5 1: 2: 2: 3: 4, flagellum three-articulate, joints subequal in length, apical organ longer than distal article (fig. 4, An2). Mandible: Pars molaris consisting of a four-branched molar penicil, pars intermedia bearing two penicils on left and one on right mandible, intermedial penicil slender (fig. 5, Mdl/r). Maxillula: Medial endite bearing two slender penicils and apical tip, lateral article with 4+5 teeth, four of inner set cleft (fig. 5, Mx1). Maxilla: Lateral lobe two times broader than medial one, fine hair-like setae arranged in parallel lines, medial lobe with 7 cusps apically (fig. 5, Mx2). Maxilliped: Basipodite with sulcus lateralis, palp with proximal article bearing one seta, endite with knob-like penicil rostrally and tooth caudally (fig. 5, Mxp). A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 819 Pereopods: Slender, dactylus with short inner claw and simple dactylar seta, carpus of pereopod | with antenna-grooming brush reaching far laterally, ornamental sensory spine with one serrate fringe reaching more proximally than other (fig. 6, Sel)! Sexual differentiation: No apparent sexual differentiation (fig. 6, PEI-7). Pleopods: Exopodites of pleopod 3 and 4 prominent, bearing laterally 4 sensory spines, no respiratory areas discernible, endopodites bilobate (fig. 7, PL1-5). Sexual differentiation: Male pleopod | exopodite almost circular, endopodite stout and coniform, apically pointed, with some distinct dentations laterally and caudally, caudal row of small spines reduced to area near apex (fig. 7, PL1). Pleopod 2 exopodite triangular with two lateral sensory spines, endopodite with long, flagelli- form apex (fig. 7, PL2), pleopod 5 exopodite with guide slot for pleopod 2 endo- podite, apically extensively drawn out (fig. 7, PLS). Uropod: Similar to generic diagnosis. Genital papilla: As in generic diagnosis. Comments: This species belongs to a group which is characterized by the reduction of the hyaline lamellae on the male pleopod endopodite, which is typical for most of the members of the Prosekia-group. Together with the following two species, A. feistae sp.n. and A. plicatipus sp. n., is united by the apomorphic structure of the male pleopod | endopodite, which is pointed and bears a variety of small bosses and knobs. As in the other species of this group, the pereopods of the males are un- modified, but males can be at once separated from other species of Androdeloscia gen. n. by the stout pleopod | endopodite. Etymology: The species name is derived from latin "conus" and "pus", meaning cone and foot, referring to the coniform endopodite of pleopod 1. Androdeloscia feistae sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE d 3mm, forest, IV-V.1975 leg. W. Hanagarth, SMNS coll. T478; PARATYPES 1d 102 4 juveniles max. 4mm, same data as holotype, SMNS coll. T448; 15 3mm, matorral at river side, clay soil, leg. W. Hanagarth, SMNS coll. T449; 35 4 juveniles, max 3mm, matorral at river side, 4.-27.XII. 1975 leg. W. Hanagarth, SMNS coll. T450; 25 129 max. 3.5mm, forest, II.-1II.1975, leg. W. Hanagarth, SMNS coll. T451. Colour: Most specimens were faded to yellowish brown, some light spottings on the dorsum. Cephalothorax: Linea frontalis missing, linea supra-antennalis prominent, only slightly bent between antennal sockets, small lateral lobes, compound eyes consisting of about 7 ommatidia (fig. 8, Ctf). Pereon: Tegument smooth and shiny, with scattered tricorn-like setae, sulcus marginalis reduced, noduli laterales flagelliform, more dorsally on coxal plate IV (fig. 8 €x3/Cxp). Pleon: Set apart from pereon, neopleurae of pleonites 3 to 5 adpressed, pleo- telson with rounded lateral margins, bearing scattered tricorn-like setae. 820 ANDREAS LEISTIKOW Appendages: Antennula: Distal article apically with small tip and two aesthetascs, medially with tuft of about 10 aesthetascs, medial article slightly shorter than proximal one (fig. 8, Anl). Antenna: Peduncle rather stout, article 4 only slightly longer than article 3, flagellum with distal article half as long as all three joints together, apical organ of two thirds the length of flagellum (fig. 8, An2). Mandible: Pars intermedia with two penicils left and one right, molar penicil consisting of about 4 branches, additional penicil distally (fig. 9, Mdl/r). Maxillula: Medial endite with two penicils and small tip apically, lateral endite with 4+6 teeth apically, inner set consisting of 5 cleft and one vestigial tooth, small spine subapically, cuticular lobe sublaterally, both on rostral surface (fig. 9, Mx1). Maxilla: Lateral lobe more than twice as broad than medial lobe, bearing scattered pectinate setae, medial lobe with 9 cusps apically (fig. 9, Mx2). Maxilliped: Basipodite with short sulcus lateralis, endite bearing two teeth caudally, small knob-like penicil rostrally, palp with single seta on proximal article, medial and distal article with setal tufts (fig. 9, Mxp). Pereopods: Slender with short setal tufts on carpus laterodistally, antenna- grooming brush of carpus | very small, tricorn-like setae slender, ornamental sensory spine serrate (fig. 10, Scl), dactylus with short inner claw, dactylar seta simple (fig. 10, Dac). Sexual differentiation: Apparently no prominent sexual differentiation, male ischium 6 and 7 with only one sensory spine laterally (fig. 10, PE1-7). Pleopods: Exopodites of pleopods 2 to 4 rhomboid, of pleopod 5 almost triangular, laterally with about | to 3 sensory spines, no respiratory areas discernible in light microscope, endopodites bilobate (Fig. 10, PLx; fig. 11, PL1-5). Sexual differentiation: Male pleopod 1 exopodite rounded, endopodite straight, muscular basal part short, apex acute, slightly twisted, i. e. caudal row of spines running on rostral side, some cuticular hooks directed proximally, hyaline serrate lobe fairly motile (fig. 11, PL1). Pleopod 2 as in other species of this genus (pl.11, PL2), exopodite of pleopod 5 with sulcus medially for pleopod 2 endopodite, distally only pointed (fig. 11, PLS). Uropod: As in other species of genus (fig. 10, UR). Genital papilla: Similar to generic diagnosis. Comments: Androdeloscia feistae sp. n. together with A. plicatipus sp. n. are the sister species of the preceding species, they share as an apomorphy the shape of the male pleopod 1 endopodite which is rather slender with a basal part of triangular shape containing the muscle M49 (cf. ERHARD 1997). Different to its sister species, the endopodite is straight and not laterally bent. Further apomorphies are the reduction of sensory spines on ischium 6 to one in contrast to two. Etymology: This species is named for Mrs R. Feist, University of Bielefeld in honour for her help in SEM preparations and photograpy. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 821 Androdeloscia plicatipus sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE d 3mm, field with maniok 12.- 27.XII.1975 leg. W. Hanagarth, SMNS coll. T479; PARATYPES 26 92 2 juveniles max. 3.5mm, same data as holotype, SMNS coll. 452; 5d 19, max. 2.5mm, matorral at river side, 4.-27.XII.1975, leg. W. Hanagarth, SMNS coll. T453; 38 189 2 juveniles, max. 3.5mm, area with kudzu tropical, III.-V.1975, leg. W. Hanagarth, SMNS coll. T454; 64 559 2 juveniles, max.3.5mm, forest, XII.75-1.1976, leg. W. Hanagarth, SMNS coll. T455; 14 29 same data, USNM coll. Colour: Reddish brown, slightly faded, dorsum with light patches at least on coxal plates, cephalothorax heavily spotted yellowish. Cephalothorax: Linea frontalis absent, area slightly bulbous, linea supra- antennalis prominent, almost straight between antennal sockets, small lateral lobes present, compound exes consisting of about 6 ommatidia (fig. 12, Ctf). Pereon: Tegument shiny, dorsum bearing scattered tricorn-like setae, coxal plates with reduced sulcus marginalis and flagelliform noduli laterales, insertion area with small concentrical grooves, more dorsally on coxal plate IV (fig. 12, Cx4/Cxp). Pleon: Retracted from pleon, neopleurae of pleonites 3 to 5 conspicuous, telson with straight margins, bearing few slender tricorn-like setae. Appendages: Antennula: Three-articulate, medial article shortest, distal article apically with small tip and two aesthetascs, medially with about 10 aesthetascs (fig. 12, Anl). Antenna: Peduncle rather stout, article 5 only slightly longer than flagellum and peduncular article 4, three-articulate flagellum with long apical organ, half as long as flagellum (fig. 12, An2). Mandible: Molar penicil consisting of 4 branches, left pars intermendia with two penicils and several coniform setae, right bearing few setae and one penicil (fig. 13, Md//r). Maxillula: Medial endite with two penicils apically, lateral endite with stepped setal fringe laterally, apically with 4+6 teeth, five of inner set cleft, innermost simple tooth vestigial (fig. 13, Mx1). Maxilla: Lateral lobe two times broader than medial, bearing several trichifrom setae and pectinate scales, apically about 8 clusps (fig. 13, Mx2). Maxilliped: Basipodite with short sulcus lateralis, endite with two teeth caudally and knob-like penicil rostrally, palp with two unequal setae on proxinal article, medial and distal article each bearing setal tuft, lateral setae rather stout (fig. 13, Mxp). Pereopods: Pereopods rather slender, with setal tuft laterodistally on carpus, carpus | with antenna-grooming brush, ornamental sensory spine serrate (fig. 14, Scl), dactylus with short inner claw and simple dactylar seta, interungual seta curved (fig. 14, Dac). Medial sensory spines of merus 6 closer together than in preceding species (fig. 14, Sm6). Sexual differentiation: Only slight differentiation, male pereopod 7 ischium medially lacking sensory spines (fig. 14, PE1-7). Pleopods: Similar to the preceding species, medial protrusion of pleopod 3 protopodite rather long (fig. 15, PL1-5). 822 ANDREAS LEISTIKOW Sexual differentiation: Male pleopod 1 exopodite rounded, endopodite with short basal part containing muscle M49, distal part slender, one third before apex strongly grooved and bent laterally, twisted, short row of spines, apex with several decurved cuticular hooks (fig. 15, PL1). Pleopod 2 exopodite pyriform, without sensory spines, endopodite slender, not as flagelliform as in preceding species (fig. 15, PL2), pleopod 5 exopodite with guide slot for endopodite 2, apically only slightly drawn out (fig. 15, PLS). Uropod: Similar to other species. Genital papilla: As in other members of genus. Comments: This species is close to the preceding, A. feistae sp. n. from which it is immediately distinguished by the shape of male pleopod 1 endopodite as described above. Therefore, its structure is more complex in comparison to the other two species of the "cone pleopod"-group which also includes A. conipus sp. n. A synapomophy of A. plicatipus sp. n. and A. feistae sp. n. 1s that there is only one medial sensory spine on the male ischium 7 instead of two as in other species. Etymology: The species name is derived from latin "plicatus" meaning "folded" and "pus" for "foot" due to the shape of pleopod 1 endopodite. Androdeloscia taitii sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°7'S 74°6'W, altitude 250m: HOLOTYPE 4 4mm, matorral at iver side, sandy soil, 20.VII.1975 leg. W. Hanagarth, SMNS coll. T456; PARATYPES 36 32 1 juvenile, max. 4mm, same data as holotype, SMNS coll. T457; 16 4.5mm, secondary forest of 10 years in flood area, 23.1V.1975, leg. W. Hanagarth, SMNS coll. T458. Colour: Dorsum faded to reddish brown, few light spots visible, cephalothorax heavily spotted, pleon uniformly coloured. Cephalothorax: Linea supra-antennalis strongly decurved between antennal sockets, small lateral lobes, linea frontalis lacking, compound eyes consiting of about 7 ommatidia (fig. 16, Ctf). Pereon: Pereonites with few tricorn-like setae, tegument smooth, coxal plates with proximally reduced sulcus marginalis, noduli laterales long, flagelliform, inser- tion on coxal plate IV considerably dorsal (fig. 16, Cx3/Cxp). Pleon: Retracted from pereon, rather short, with small neopleurae on pleonites 3 to 5, pleotelson with straight lateral margins. Appendages: Antennula: Three-articulate, proximal article rather slender, distal article with two aesthetascs apically, broken in dissected antennule, medial set of about 10 aesthetascs (fig. 16, Anl). Antenna: More slender than in preceding species, especially flagellum, medial article shortest, apical organ as long as distal article (fig. 16, An2). Mandible: Pars intermedia only sparsely setose, molar penicil composed of 4 rather short branches, right lacina mobilis deeply sinuous apically (fig. 17, Mdl/r). Maxillula: Medial endite with small tip and two penicils apically, medial endite with hyaline cuticular lobe and 4+6 teeth, four of innermost set cleft (fig. 17, Mxl). A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 823 Maxilla: Lateral lobe two times broader than medial one, bearing trichiform setae, medial lobe bearing stronger setae, apically with about six cusps (fig. 17, Mx2). Maxilliped: Basipodite with sulcus lateralis almost reduced, palp with two setae on proximal article, setal tufts on distal articles, innermost consisting of 3 setae, endite bearing knob-like penicil rostrally and two teeth caudally (fig. 17, Mxp). Pereopods: Rather slender, especially pereopod 1, carpus with antenna- grooming brush and apically serrate ornamental sensory spine, carpus of pereopods 3 and 4 with hyaline cuticular fringe mediodistally (fig. 18, PE3/4), laterodistal tufts small, dactylus with simple dactylar seta and short inner claw (fig. 18, Dac). Sexual differentiation: Male merus of pereopod 2 and 3 with quadrangular, of pereopod 4 to 6 with acute, proximally directed cuticular scales medially (fig. 18 PEI- 4; fig. 19, PE 6-7). Pleopods: Exopodites of pleopod 3 and 4 rhomboid, laterally with 3 to 4 sensory spines, exopodite 5 more triangular, lacking respiratory areas, endopodites bilobate (fig. 20, PL1-5). Sexual differentiation: Male pleopod | exopodite pointed, shape characteristic: almost straight from lateral insertion on protopodite to apex, medially much more rounded at the base, endopodite with short basal area containing muscle M49, distally straight up to the bulbous apex, bearing two horns, medially with groove, and twisted, row of spines beginning caudally, ending more apically on rostral side, lateral area of bulb rostrally with hyaline lamellae, caudally granulate (fig. 20; PL1). Pleopod 2 exopodite similar to exopodite 1, even more drawn out medioproximally, no sensory spines, endopodite slender, less tapered than A. feistae sp. n. (fig. 20, PL2), exopodite 5 triangular with lateral furrow, pointed (fig. 20, PLS). Uropod and genital papilla: As in generic diagnosis. Comments: This and the following four species form a monophyletic group within Androdeloscia gen. n. characterized by the shape of the pleopod | exopodite in the male which appears to be inverted, i.e. the lateral margin turned medially. A synapomorphy of A. taitii sp. n., A. merolobata sp. n. and A. longiunguis sp. n. is the distorted pleopod 1 endopodite in the male: the spermatic channel is turning from the caudal to the rostral side, thus, the caudal row of spines is ending on the rostral side. Beside the shape of the male pleopod | endopodite, A. faitii sp. n. is at once distin- guished by the lack of lobes on the merus 7, it only shows areas of cuticular scales on most of the pereopods. Etymology: The species is dedicated to Dr S. Taiti, Florence, one of the leading specialists of Oniscidea. Androdeloscia merolobata sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE dé 5mm, forest, X.-XI.1975 leg. W. Hanagarth, SMNS coll. T459; PARATYPES 2¢ 10% 1 juvenile, max. 5.5mm, same data as holotype, SMNS coll. T460; 54 several © and juveniles, max. 5mm, cattle meadow with kudzu, 1975-1976 leg. W. Hanagarth, SMNS coll. T461; 35 J42 1 juvenile, max.5.5mm, forest, VI.-VII.1975 leg. W. Hanagarth, SMNS coll. T462; 98 99 1 juv., max. 5.5mm, forest, 824 ANDREAS LEISTIKOW XI.1975-1.1976 leg. W. Hanagarth, SMNS coll. T463; 18 29 same data, USNM coll.; 38 29 max. 5.5mm, area with kudzu tropical, 12.VI.1975 leg. W. Hanagarth, SMNS coll. T464; 34 22 max. 5.5mm, area with kudzu tropical, 27.1V.1975 leg. W. Hanagarth, SMNS coll. T465; 33 max. 4.5mm, forest, II.1975 leg. W. Hanagarth, SMNS coll. T466. Colour: Dorsally reddish brown with yellowish spottings, medial line dark brown, on pereonites 5 to 7 bifurcated, pleonites uniformly reddish brown, cephalo- thorax with light spotting. Cephalothorax: Linea supra-antennalis straight, lateral lobes small, linea fron- talis lacking, compound eyes consisting of about 7 ommatidia (pl.21, Ctf). Pereon: Tegument smooth and shiny, bearing scatterd tricorn-like setae, coxal plates without gland pores, sulcus marginalis poorly individualized, noduli laterales flagelliform, on coxal plate IV more dorsally inserted (pl.21, Cx3/Cxp). Pleon: Set apart from pereon, neopleurae of pleonites 3 to 5 rather short, pleotelson with rounded apex, laterally straight, bearing few tricorn-like setae. Appendages: : Antennula: As in preceding species (pl.21, Anl). Antenna: Fairly slender, peduncular article 2 stout, flagellum three-articulate, joints subequal in length, apical organ half as long as flagellum (pl.21, An2). Mandible: Pars intermedia with two penicils on left, one on right side, some coniform setae, molar penicil consisting of four branches, additional penicil slender (fig. 22, Mdl/r). Maxillula: Medial endite with two penicils and lateral tip apically, lateral endite with fringe of stepped setae, more distally stouter and individualized, apical teeth 4+6, four of innermost cleft, simple teeth short, subapical vestigial tooth on rostral surface (fig. 22, Mx1). Maxilla: Lateral lobe twice as broad as medial one, covered with rows of trichiform setae, medial lobe apically cuspidate, bearing trichiae (fig. 22, Mx2). Maxilliped: Basipodite with conspicuous sulcus lateralis, endite apically setose, with two teeth on caudal, small knob-like penicil on rostral surface, proximal article of palp bearing two setae, distal lobes with setal tufts (fig. 22, Mxp). Pereopods: Pereopods slender, small setal tuft laterodistally on carpus, carpus 1 with small antenna-grooming brush and apically serrate ornamental sensory spine (fig. 23, Scl), dactylus with short inner claw and simple dactylar organ (fig. 23, Dac). Sexual differentiation: Male pereopod 2 to 3 merus with quadrangular cuti- cular scales, merus 4 to 6 with acute cuticular scales, pereopod 7 merus with semicircular lobe mediodistally on caudal side, bearing cuticular scales, covering a third of length of merus (fig. 23, PE1-4; pl.24, PES-7). Pleopods: Pleopod exopodites 3 to 4 rhomboid with 3 to 4 sensory spines laterally, without respiratory area, endopodites bilobate, pleopod 5 more pointed (pli25 sR 1-5): Sexual differentiation: Male pleopod | exopodite similar to A. taitii sp. n., endopodite at its base slightly smaller than protopodite, basis containing M49 trian- gular, distal part twisted, row of spines turned to rostral side, apex bulbous with transverse groove on caudal side, sides of spermatic furrow serrate, rostral side with A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 825 some cuticular granules and hyaline lamellae, bulb terminated by laterally directed flagellar extension (fig. 25, PL1). Pleopod 2 exopodite with broad rounded apex, bearing 3 sensory spines laterally, endopodite slender, distal half flagelliform (fig. 25, PL2), pleopod 5 exopodite triangular with slightly drawn out apex, caudally with medial guide slot for endopodite 2 (fig. 25, PLS). Uropod and genital papilla: As in other members of genus (fig. 24, UR). Comments: As the species name infers, A. merolobata sp. n. bears a medio- distal lobe on merus 7 in the male. This and the complex shape of the male pleopod 1 endopodite are autapomorphies of this species which forms the adelphotaxon of A. longiunguis sp. n., the apomorphy is the shape of pleopod 2 exopodite with two sen- sory spines laterally. It is united with A. taitii and its sister species by the distortion of the male pleopod 1 endopodite. A. merolobata sp. n. was the by far commonest of all its congeners in the Panagua area, it was collected at many sites in big numbers. It occurred in both forest and open areas as cleared places for agriculture. Etymology: The species name refers to the lobe on the male pereopod 7 merus. Androdeloscia longiunguis sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE 4 4mm, area with kudzu tropical S.IX.1975 leg. W. Hanagarth, SMNS coll. T467. Colour: The only male of this species rather faded, so that spotting invisible, dark median line dorsally. Cephalothorax: Linea frontalis lacking, linea supra-antennalis present, small lateral lobes and slight lamina frontalis, compound eyes consisting of about 8 omma- tidia (fig. 26, Ctf). Pereon: Pereonites covered with small tricorn-like setae, tegument smooth, coxal plates without sulcus marginalis and gland pores, noduli laterales long, flagelli- form, nodulus of coxal plate IV the most dorsal one (fig. 26, Cx3/Cxp). Pleon: Set apart from pereon, neopleurae of pleonites 3 to 5 small but clearly visible, pleotelson with rounded lateral margin, bearing several tricorn-like setae. Appendages: Antennula: Similar to other species of the genus, apical tuft with 3 aesthetascs (fig. 26, Anl). Antenna: Rather stout, length ratio of peduncular articles 1 to 5 is 1: 2: 2: 3: 4, flagellum three-articulate, distal article longest, apical organ slightly longer (fig. 26, An2). Mandible: Molar penicil consisting of 3 branches, pars intermedia with 2 penicils and coniform setae on left, one penicil and few setae right, additional seta long (fig. 27, Mdl/r). Maxillula: Medial endite with small tip and 2 penicils apically, lateral endite with lateral fringe similar to A. merolobata sp. n., tooth formula 4+6, with four teeth of inner set cleft, innermost simple tooth short, hyaline cuticular lobe present, rostrally with subapical vestigial tooth (fig. 27, Mx1). 826 ANDREAS LEISTIKOW Maxilla: Lateral lobe more than two times broader than medial, heavily covered with trichiform setae, medial lobe bearing about 6 cusps (fig. 27, Mx2). Maxilliped: Basipodite with sulcus lateralis, palp with two setae on proximal article, two setal tufts on distal articles, endite with two teeth caudally, knob-like penicil rostrally (fig. 27, Mxp). Pereopods: Rather slender, especially pereopod 1, carpus with antenna- grooming brush and serrate ornamental sensory spine (fig. 28, Scl), dactylus with long inner claw and simple dactylar seta (fig. 28, Dac). Sexual differentiation: Male pereopod 2 to 4 with cuticular scales on merus, or cuticular plaques in pereopod 5 and 6, merus 7 with sinuous lobe bearing sensory spine proximally of halflength (fig. 28, PE1-7). Pleopods: As in the preceding species (fig. 29, PL1-5). Sexual differentiation: Male pleopod | exopodite similar to A. merolobata, endopodite structurally similar, twisted with row of small spines rostrally, two caudal grooves vertically, apex less bulbous, rather square, long cuticular spines directed medially, hyaline lamellae on rostral surface (fig. 29, PL1). Pleopod 2 and 5 similar to preceding species (fig. 29, PL2/5). Uropod and genital papilla: As in generic diagnosis (fig. 26, UR). Comments: The name-giving long inner claw, the location of the meral lobe and the shape of the male pleopod | endopodite are the autapomorphies of this species. A. longiunguis sp. n. is linked with the preceding species by the shape and setation of the male pleopod 2 exopodite. Etymology: The species name is derived from latin for stressing the long inner claw of the dactylus. Androdeloscia ferrarai sp.n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE 6 5mm, "VT 57a RC22-21", 15.V.1976 leg. W. Hanagarth, SMNS coll. T496; PARATYPES 36 6% 5 juveniles, max. 5.5mm, same data as holotype, SMNS coll. T470; 68 389 6 juveniles, max. 3mm, field with annual maniok, 12.-27.XI1.1975 leg. W. Hanagarth, SMNS coll. T471; 16 22 same data, USNM coll. Colour: Reddish to purplish brown, pereon with pale dorsal spotting, light patches on coxal plates, pleon uniformly coloured. Cephalothorax: No linea frontalis, linea supra-antennalis present, small lateral lobes and slight lamina frontalis, compound eyes consisting of about 10 ommatidia (fig. 30, Ctf). Pereon: Smooth and shiny tegument, coxal plates with sulcus marginalis, no gland pores, noduli laterales long and flagelliform, most dorsally on coxal plate IV (fig. 30, Cx3/Cxp). Pleon: Retracted from pereon, neopleurae of pleonites 3 to 5 conspicuous, pleotelon with straight margins, bearing several prominent tricorn-like setae. Appendages: Antennula: Three-articulate, rather slender with distal article bearing pro- minent tuft of aesthetascs medially and 2 aesthetascs apically (fig. 30, An1). A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 827 Antenna: Comparatively slender, length ratio of peduncular articles I to 5 1: 2: 2: 4: 5, flagellum three-articulate with articles subequal in length, distal article slightly longer, apical organ even longer (fig. 30, An2). Mandible: Molar penicil consisting of 4 branches, left pars intermendia with two penicils and several coniform setae, right bearing few setae and one penicil (pl.31, Mdl/r). Maxillula: Medial endite with small tip and two penicils apically, medial endite with hyaline cuticular lobe slender, 4+5 teeth, four of innermost set cleft (fig. SIM): Maxilla: Lateral lobe slightly broader than medial, almost without setation, medial lobe setose with about 10 apical cusps (fig. 31, Mx2). Maxilliped: Basipodite with conspicuous sulcus lateralis, endite apically setose, with two teeth on caudal surface, small knob-like penicil lacking, proximal article of palp bearing two setae, distal lobes with setal tufts, proximal one consisting of two setae (fig. 31, Mxp). Pereopods: Pereopods rather slender, with setal tuft laterodistally on carpus, carpus | with antenna-grooming brush, ornamental sensory spine serrate (fig. 32, Scl), dactylus with short inner claw and simple dactylar seta, interungual seta curved (fig. 32, Dac), merus 3 to 4 with hyaline cuticular fringe (fig. 32, PE3/4). Sexual differentiation: Male pereopod merus 5 with hyaline cuticular fringe, merus 6 with cuticular scales medially, pereopod 7 merus with small setose hump distally at halflength (pl.33, PES-7). Pleopods: Similar to preceding species, endopodites with straight to concave distal margin (fig. 34, PL1-5). Sexual differentiation: Male pleopod 1 exopodite as in preceding species, endopodite with rather broad basal part containing M49, of half-length of endopodite, apical part bulbous, terminally drawn out, laterally with bulbous lobes, each separated by groove, distally some cuticular hooks, medially some granules, caudally with subapical row of spines, apically some hyline lamellae (fig. 34, PL1). Pleopod 2 endopodite with two lateral sensory spines, endopodite with flagelliform distal part (fig. 34, PL2), pleopod 5 exopodite with medial guide slot caudally, slightly pointed (fig. 34, PLS). Uropod: Protopodite only slightly grooved between insertion of endo- and exopodite. Genital papilla: Margins of ventral shield almost parallel, slightly surpassed by ductus ejaculatorii (fig. 34, Gen). Comments: The male pleopod | endopodite may be referred to as somehow near to the ground plan of the "inverted exopodite"-group. The presence of a lamina frontalis is somewhat surprising and must be interpreted as a silent gene expression. The maxilliped endite lacks the penicil on the rostral surface, this reductive character is an autapomorphy of this species. As in the preceding species, the shape of both male merus 7 and pleopod | endopodite at once distinguish A. ferrarai sp. n. from its congeners. 828 ANDREAS LEISTIKOW Etymology: This species is dedicated to Dr F. Ferrara, Florence, one of the leading scientists of Oniscidea systematics. Androdeloscia poeppigi sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE ¢ 2.5mm, "A3.4", 16.X1.1975, leg. W. Hanagarth, SMNS coll. 472; PARATYPES 26 49 1 juvenile, max.3.5mm, same data as holotype, SMNS coll. T473. Colour: Yellowish brown, rather faded due to alcohol preservation, thus spotting badly discriminable. Cephalothorax: Linea frontalis absent, area slightly bulbous, linea supra- antennalis prominent, almost straight between antennal sockets, small lateral lobes present, compound exes consisting of about 7 ommatidia (fig. 35, Ctf). Pereon: Tegument smooth and shiny, coxal plates with sulcus marginalis rostrally, long flagelliform nodulus lateralis, nodulus lateralis of coxal plate IV inserting more medially than others (fig. 35, Cx2/Cxp). Pleon: Set apart from pereon, neopleurae of pleonites 3 to 5 adpressed, pleotelson with straight lateral margins, bearing few scattered tricorn-like setae. Appendages: Antennula: Three-articulate, proximal article rather slender, distal article with two aesthetascs apically, broken in dissected antennule, medial set of about 10 aesthe- tascs (fig. 35, Anl). Antenna: Peduncle rather stout, article 4 only slightly longer than article 3, flagellum with distal article half as long as all three joints together, apical organ of two thirds the length of flagellum (fig. 35, An2). Mandible: Molar penicil consisting of 4 branches, left pars intermendia with two slender penicils and several coniform setae, right bearing few setae and one penicil, additional penicil present (fig. 36, Mdl/r). Maxillula: Medial endite with two penicils and lateral tip apically, lateral endite with fringe of stepped setae, more distally stouter and individualized, apical teeth 4+5, four of inner set cleft, second lateral tooth rather short, decurved (fig. 36, Mxl). Maxilla: Lateral lobe more than two times broader than medial, heavily covered with pectinate scales, medial lobe bearing about 7 cusps (fig. 36, Mx2). Maxilliped: Basipodite with sulcus lateralis, endite caudally with two teeth, rostrally bearing knob-like penicil, palp with one seta on proximal article, medial article with proximal tuft of 2 short setae, distal tuft of 2 setae (fig. 36, Mxp). Pereopods: Slender with short setal tufts on carpus laterodistally, antenna- grooming brush of carpus | dense, lateroproximally with 2 tricorn-like setae, orna- mental sensory spine serrate (fig. 37, Scl), dactylus with short inner claw, dactylar seta simple (fig. 37, Dac). Sexual differentiation: Appartently no sexual dimorphism (fig. 37 PE1-7). Pleopods: Expodites of pleopods 3 to 4 rhomboid, of pleopod 5 almost triangular, laterally with about 2 to 4 sensory spines, no respiratory areas discernible in light microscope, endopodites bilobate (fig. 38, PL1-5). A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 829 Sexual differentiation: Male pleopod 1 exopodite rounded with inconspicuous point, basal part of endopodite containing M49 slightly individualized, distal part pointed, no caudal row of spines, apex obtuse, two protrusions on lateral and medio- rostral side at about halflength, the latter better discernible when endopodite separated from protopodite (fig. 38, PL1). Pleopod 2 exopodite triangular, bearing one lateral sensory spine subapically, endopodite more than twice as long as exopodite, not as flagelliform as in preceding species (fig. 38, PL2). Pleopod 5 exopodite with medial guide slot and drawn out apex (fig. 38, PLS). Uropod: Similar to other species of the genus (fig. 35, UR). Genital papilla: Rather stout, ventral shield slightly ovoid (fig. 38, Gen). Comments: Together with A. malleus sp. n., A. poeppigi sp. n. forms the adel- photaxon of the "fir-cone"-species group, the small finger-shaped humps on the male endopodite of pleopod 1 and the reduction of the caudal row of spines are their synapomorphies. Different to A. malleus sp. n., this species has a narrow male ple- opod 1 endopodite with the humps arranged more proximally in contrast to the former. Etymology: This species is named after E. F. Poeppig, one of the first explorers of the Amazon basin. Androdeloscia malleus sp. n. Material: Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station "Panguana" 9°37'S 74°56'W, altitude 250m: HOLOTYPE ¢ 3mm, "A2" 1.X11.1975, leg. W. Hanagarth, SMNS coll. T474; PARATYPES 35 39 max. 3.5mm, same data as holotype, SMNS coll. 475, 38 282, max. 3.5mm, forest, VIIL-IX.1975, leg. W. Hanagarth SMNS coll. T476, 58 239 2 juveniles, max. 3.5mm, forest, VI.-VII.1975 leg. W. Hanagarth, SMNS coll. T477. Colour: As in other species of the genus, dorsally reddish brown with light markings of muscle insertions, cephalothorax heavily spotted. Cephalothorax: Linea frontalis missing, area slightly bulbous, linea supra- antennalis prominent, only slightly bent between antennal sockets, small lateral lobes, compound eyes consisting of about 7 ommatidia (fig. 39, Ctf). Pereon: Tegument shiny, dorsum bearing scattered tricorn-like setae, coxal plates with reduced sulcus marginalis and flagelliform noduli laterales, insertion area with small concentrical grooves, maximum of d/c-coordinates on coxal plate IV (fig. a9 €x3/ Exp). Pleon: Retracted from pereon, neopleurae visible, pleotelson with rounded distal margin, laterally straight, bearing few, short tricorn-like setae. Appendages: Antennula: Distal article apically with small tip and two aesthetascs, medially with tuft of about 9 aesthetascs, medial article slightly shorter than proximal one (fig. 39, Anl). Antenna: More slender than in preceding species, especially flagellum, distal article longest, proximal articles subequal in length, apical organ longer than distal article (fig. 39, An2). 830 ANDREAS LEISTIKOW Mandible: Pars molarıs consisting of a four-branched molar penicil, pars intermedia bearing two penicils on left and one on right mandible, intermedial penicil slender (fig. 40, Mdl/r). Maxillula: Similar to preceding species (fig. 40, Mx1). Maxilla: Lateral lobe only slightly broader than medial lobe, bearing pectinate scales, medial lobe sparsely coverd with trichiae, apically cuspidate (fig. 40, Mx2). Maxilliped: As in A. poeppigi sp. n., but distal setal tuft of media article of palp comprising about 4 setae (fig. 40, Mxp). Pereopods: Rather slender, especially pereopod 1, carpus with antenna-groo- ming brush and serrate ornamental sensory spine (fig. 41, Scl), propus 1 with 2 distal sensory spines serrate on one side, dactylus with long inner claw and simple dactylar seta (fig. 41, Dac). Sexual differentiation: Pereopods in both sexes similar (fig. 41, PE1-7). Pleopods: Pleopod exopodites 3 and 4 elongate rhomboid with 3 to 4 sensory spines laterally, exopodite 5 more or less triangular, endopodites bilobate, no respiratory areas discernible in light microscope (fig. 42, PL1-5). Sexual differentiation: Male pleopod 1 exopodite rounded, endopodite rather stout, apex hammer-shaped, some minute spines caudally present, spermatic furrow laterally open, on lateral side of apex few granules (fig. 42/43, PL1). Pleopod 2 and 5 similar to preceding species (fig. 42, PL2/5). Uropod: As in other species of genus. Genital papilla: Ventral shield stout, ovoid, slightly surpassed by ductus ejaculatorii (fig. 42, Gen). Comments: This is the sister species of A. poeppigi sp.n., differing mainly in the shape of male pleopod | endopodite. A. malleus has the two lobe more apically thus forming the "hammer"-like apical region. The endopodite is rather stout and as in the preceding species, it lacks the hyaline lamellae which are so characteristic for this genus although they do not represent an autapomorphy of Androdeloscia gen. n.. Etymology: The latin term "malleus" means "hammer" and refers to the shape of the male pleopod | endopodite. Androdeloscia opercularis sp. n. Material: Venezuela: HOLOTYPE d 3mm, Falcon, Parque Morrocoy, Peninsula de Morrocoy, northern shore, Cueva del Indio (carstic cave with crushed ceiling, within detritus in small niches and edges of the rock, under stones on bottom leg. 18.03.1998 C. Schmidt MHNG coll., PARATYPES 16 32, same data as holotype, MHNG coll.; 96 59, about 80 imm., same dat as holotype, author's coll. Colour: Purplish brown with pereon and pleon dorsally bearing light spots, medial line of pereon dark brown. Cephalothorax: Linea frontalis absent, area slightly bulbous, linea supra- antennalis prominent, almost straight between antennal sockets, small lateral lobes present, compound eyes consisting of about 7 ommatidia (fig. 44, Ctf) Pereon: Smooth and shiny tegument, coxal plates with sulcus marginalis along lateral border, no gland pores, insertion of nodulus lateralis bordered by concentric A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 831 structure of the cuticle, noduli long and flagelliform, on coxal plate IV more dorsally (fig. 44, Cx1/Cxp) Pleon: Retracted from pereon, neopleurae of pleonites 3 to 5 adpressed, pleotelson with straight margins, bearing some prominent tricorn-like setae. Appendages: Antennula: Distal article apically with small tip and two aesthetascs, medially with tuft of about 10 aesthetascs, medial article much shorter than proximal one (fig. 44, Anl). Antenna: Peduncle rather slender, article 4 two times longer than article 3, flagellum with distal article as long as other joints, apical organ of two thirds the length of flagellum (fig. 44, An2). Mandible: Pars molaris consisting of a five-branched molar penicil, pars intermedia bearing two penicils on left and one on right mandible, intermedial penicil slender (fig. 45, Mdl/r). Maxillula: Medial endite with two penicils and apical tip, lateral endite bearing 4+5 teeth, inner set cleft, one short subapical tooth caudally, lateral fringe of trichi- form setae stepped (fig. 45, Mx1). Maxilla: Lateral lobe slightly broader than medial one, fine hair-like setae arranged in parallel lines, medial lobe with 7 cusps apically (fig. 45, Mx2). Maxilliped: Basipodite with sulcus lateralis, palp with proximal article bearing two setae, endite with knob-like penicil rostrally and tooth caudally (fig. 45, Mxp). Pereopods: Slender with many tricorn-like setae, pereopod | to 6 with hyaline fringes on medial border, pereopod | with antenna-grooming brush on carpus, ornamental seta double-fringed serrate (fig. 46, Sc1), dactylus with short inner claw and simple dactylar seta, interungual seta apically swollen (fig. 46, Dac). Sexual differentiation: Medial margin of pereopod 2 to 4 merus with fields of proximally directed cuticular setae (fig. 46, PEI-7). Pleopods: Exopodite of pleopods 3 rhomboid, pleopod 5 almost triangular, pleopod 4 intermediate, laterally with 2 sensory spines, no respiratory areas discer- nible in light microscope, endopodites bilobate (fig. 47, PL1-5). Sexual differentiation: Male pleopod 1 exopodite circular with transverse fol- ding medially of insertion in protopodite, comparatively large, endopodite proximally stout, muscle insertion area of M49 slightly protruding distally, apical part with lobe rostrally, apex protruding, rostrally with two rows of knobs, laterally serrate, caudally with hyaline lamellae, caudal row of spines from halflength of interlocking area with genital papilla not reaching apex (fig. 47, PL1). Pleopod 2 exopodite with broad rounded apex, bearing 3 sensory spines laterally, endopodite slender, distal half flagelliform (fig. 47, PL2), pleopod 5 exopodite triangular with slightly drawn out apex, caudally with medial guide slot for endopodite 2 (fig. 47, PLS). Uropod: As in other species of the genus. Genital papilla: Ventral shield rather slender with parallel margins, ductus ejaculatorii slightly surpassing ventral shield (fig. 47, Gen). Comments: This is one of the species with rather simple apical part of the 832 ANDREAS LEISTIKOW pleopod 1 endopodite, from its congeners A. opercularis sp. n. is at once separable by the huge rounded exopodites of pleopod 1, which almost covers the pleoventral area. Etymology: The species name "opercularis" refers to the big pleopod 1 exo- podites of the male. Androdeloscia silvatica (Lemos de Castro & Souza, 1986) comb. n. Prosekia silvatica Lemos de Castro & Souza, 1986 Material: Venezuela: 25 (max. 3mm) 32 (ovigerous), Peninsula de Paria, eastern part of south coast, 10°35,94'N 63°11,81'E, along brook with coacoa plantations and moist forest, upper part with water, concrete pond with moist bottom under leaf litter, leg. 09.04.1998 C. Schmidt, MNHG coll.; 34 119 same data, author's coll.; 24, 39 (ovigerous) Peninsula de Paria, Puy Puy 10°42.00'N 62°58.05'W, bay with sandy beach, coconut palms, surrounded by dry mountains, banana plantation at the eastern part of the bay, in leaf litter, leg. 31.03.1998 C. Schmidt, MUMV coll.; 174 129 (ovigerous) 6imm same data, author's coll. Colour: Dorsally purplish brown with pale spots on pereon, medial line dark brown, white central stripe on pereonites I to IV, double line on pereonites V to VII, pleon unmarked. Cephalothorax: Linea frontalis lacking, linea supra-antennalis and lamina fron- talis present, small lateral lobes, compound eyes consisiting of seven ommatidia (fig. 48, Ctf). Pereon: Tegument smooth and shiny, coxal plates lacking gland pores, sulcus marginalis reduced, noduli laterales present, long and flagelliform, with maximum of d/c-coordinates on coxal plate IV (fig. 48, Cx3/Cxp). Pleon: Retracted from pereon, neopleurae of pleonites 3 to 5 adpressed, pleo- telson with straight distal margin, bearing some tricorn-like setae. Appendages: Antennula: Three-articulate with prominent proximal article, distal joint bulbous, bearing two distinct sets of aesthetascs (fig. 48, Anl). Antenna: Antennal peduncle composed of five articles with length ratio 1: 2: 2: 3: 4, densely covered with tricorn-like setae, flagellum composed of three articles, distal one bearing prominent apical organ, as long as flagellar articles 1 and 2 together (fig. 48, An2). Mandible: Pars intermedia only sparcely setose, molar penicil composed of 3 rather short branches, right lacina mobilis deeply sinuous apically (fig. 49, Mdl/r). Maxillula: Medial endite with small tip and two penicils apically, medial endite with subapical tooth and 4+5 teeth, four of innermost set cleft (fig. 49, Mx1). Maxilla: Lateral lobe two times broader than medial one, bearing trichiform setae and pectinate scales, medial lobe bearing stronger setae medially, apically with about six cusps (fig. 49, Mx2). Maxilliped: Basipodite with conspicuous sulcus lateralis, endite with two teeth on caudal surface, small knob-like penicil rostrally, proximal article of palp bearing two setae, distal lobes with setal tufts, proximal one consisting of two setae (fig. 49, Mxp). Pereopods: Pereopods rather slender, with setal tuft laterodistally on carpus, A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 833 carpus | with antennal-grooming brush, ornamental sensory spine serrate (fig. 50, Scl), dactylus with medium-sized inner claw and simple dactylar seta, interungual seta straight (fig. 50, Dac), merus 2 to 4 with hyaline cuticular scales (fig. 50, PE2). Sexual differentiation: Male pereopod 7 with two small lobes mediodistally, caudally of distal sensory spine, proximally directed lobe on medioproximal edge, covered with small scales (fig. 50, PEI-7). Pleopods: Similar to preceding species, endopodites more triangular (fig. 51, PL1-5). Sexual differentiation: Male pleopod | exopodite with slight point, endopodite with broad basis, followed by broad part without musculature with distally directed lateral protrusion, apical part decurved laterally, apex bearing three rows of knobs, caudally a trace of hyaline lamellae, row of spines short (fig. 51, PL1). Pleopod 2 exopodite trianular with one sensory spine laterally, endopodite with flagelliform distal half (fig. 51, PL2). Pleopod 5 exopodite with guide slot for pleopod 2 exopodite (igo PIES): Uropod and genital appendage : See generic diagnosis (fig. 51, Gen). Comments: A. silvatica shares the character, that the pereopod 7 merus bears three lobes medially with the following species. This character was not depicted by Lemos de Castro & Souza in the original description (1986), possibly the examined specimen was a young male or the pereopod 7 was substitutional and, therefore, does not show all the characters of a mature male. Its autapomorphies are the shape and position of lobes on merus 7 and the shape of pleopod 1 endopodite, which looks like a tin-opener of a Swiss Officer's knife. Androdeloscia pseudosilvatica sp. n. Material: Venezuela: HOLOTYPE 16 3mm, Quebrada de la Virgen, 8°58.20'N 69°47.57'W, dry plain, agriculture, near a well under flower pots beneath a tree in puddle, leg. 25.03.1998 C. Schmidt; PARATYPES 1d 12 (marsupium), same data as holotype, MHNG coll.; 45 19 (marsupium) limm, same data as holotype, author's coll. Colour: Dorsally reddish to purplish brown, pereon spotted with light mar- kings, pereonites V to VII with light medial spot, pleon uniformly coloured. Cephalothorax: Vertex strongly arched, linea frontalis reduced, linea supra- antennalis and slight lateral lobes present, no lamina frontalis, compound eyes composed of about seven ommatidia (fig. 52, Ctf). Pereon: Tegument rather smooth with scattered tricorn-like setae, coxal plates with sulcus marginalis and nodulus lateralis (fig. 52, Cx3/Cxp), no gland pores visible at 400x magnification. Pleon: Set apart from pereon, small neopleurae on pleonites 3 to 5, laterally adpressed, pleotelson with straight lateral margins, bearing few tricorn-like setae. Appendages: Antennula: As in other species of the genus. (fig. 52, Anl) Antenna: Peduncle rather stout, length ratio from proximal to distal 1: 2: 2: 3: 4, flagellum three-articulate, joints subequal in length, medial and distal article with pair of aesthetascs, apical organ long and slender, half of length of flagellum, short 834 ANDREAS LEISTIKOW free sensilla (fig. 52, An2). Mandible: Molar penicil consisting of about 4 branches, pars intermedia bearing two on left and one penicil on right mandible, additional plumose seta proximally (fig. 53, MdVr). Maxillula: Medial endite with two penicils and inconspicuous subapical tip, lateral endite apically bearing 4+5 teeth, four of inner set cleft, laterally with trichiae, medial area with trichiae fused and forming hyaline plaques (fig. 53, Mx1). Maxilla: Lateral lobe broader than medial one, slightly setose, medial lobe bearing about 5 cusps (fig. 53, Mx2). Maxilliped: Basipodite with sulcus lateralis, endite with small knob-like penicil on rostral side, palp with three setal tufts medially (fig. 53, Mxp). Pereopods: Pereopod 1 with antenna-grooming brush, ornamental sensory spine double-fringed serrate (fig. 54, Sc1), hyaline cuticular scales on merus 1 and 2, dactylus with short inner claw and simple dactylar seta (fig. 54, Dac). Sexual differentiation: Male pereopod 7 merus with two crenulate lobes mediodistally, another lobe more proximally at insertion of basal sensory spine (fig. 54, PE1-7). Pleopods: Pleopod endopodites slightly bilobate, exopodites 3 and 4 rhomboid, with 3 lateral sensory spines, no respiratory areas (fig. 55, PLI-S). Sexual differentiation: Male pleopod | exopodite rounded with small distal point, endopodite basis strong, a small furrow separating muscle insertion M49 from muscle-free part, apical third much more slender, slightly directed laterally, bearing small knobs rostrally, caudal row of spines short, lateral margin of spermatic channel not reaching proximally to interlocking area with genital papilla, trace of hyaline lamellae apically (fig. 55, PL1). Pleopod 2 and 5 similar to preceding species (fig. 55, PE2/5); Uropod and genital papilla: As in other species (fig. 55, Gen). Comments: Similar to A. silvatica, A. pseudosilvatica sp. n. bears three lobes on merus 7 in the male, the position is a little bit different. The pleopod 1 endopodite is not as drawn out as in the preceding species, only a small furrow is present laterally. Different to A. silvatica, this species has a complete sulcus marginalis on the coxal plates and lacks the lamina frontalis. Androdeloscia dalensi sp. n. Material: Venezuela: HOLOTYPE 4 4mm, Andes, road from Timotes to Mérida, 8°53.72' N 70°47.99'W 3400 +- 500m, very steep northern slope, covered with Bryophyta, Pteridophyta, Poaceae, Ericaceae, between Bryophyta, leg. 23.03.1998 C. Schmidt, MNHG coll.; PARATYPES 16 2? (ovigerous), same data as holotype, MHNG coll. 23 3% (ovigerous) 1 imm., same data as holotype, author's coll. Colour: Pereon and pleon dorsally purplish brown, both with prominent pale patches. Cephalothorax: Linea frontalis lacking, linea supra-antennalis and lamina frontalis present, small lateral lobes, compound eyes consisiting of eight ommatidia (fig. 56, Ctf). Pereon: Tegument smooth and shiny, coxal plates lacking gland pores and A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 835 sulcus marginalis, noduli laterales present, long and flagelliform, more dosally inserted on coxal plate IV (fig. Cx3/Cxp). Pleon: Retracted from pereon, neopleurae visible, adpressed; pleotelson with straight distal margin, bearing some slender tricorn-like setae. Appendages: Antennula: Three-articulate, medial article shortest, distal article apically with small tip and two aesthetascs, medially with about 10 aesthetascs (fig. 56, Anl). Antenna: Peduncle rather stout, article 5 only slightly longer than flagellum and peduncular article 4, three-articulate flagellum with long apical organ, half as long as flagellum (fig. 56, An2). Mandible: Molar penicil consisting of 5 branches, left pars intermedia with two penicils and several coniform setae, right bearing few setae and one penicil (fig. 57, Mdl/r). Maxillula: Medial endite with two penicils apically, lateral endite with stepped setal fringe laterally, apically with 4+5 slender teeth, five of inner set cleft, subapical tooth caudally (fig. 57, Mx1). Maxilla: Lateral lobe two times broader than medial, bearing several trichifrom setae and pectinate scales, apically about 6 cusps (fig. 57, Mx2). Maxilliped: Basipodite with short sulcus lateralis, endite with two teeth caudally and prominent knob-like penicil rostrally, palp with two unequal setae on proximal article, medial and distal article each bearing setal tuft, lateral setae rather stout (fig. 57, Mxp). Pereopods: Slender with short setal tufts on carpus laterodistally, antenna- grooming brush of carpus | dense, lateroproximally with 3 tricorn-like setae, orna- mental sensory spine serrate (fig. 58, Scl), sensory spine of propus | pointed, sub- apically serrate (fig. 58, Spl), dactylus with short inner claw, dactylar seta simple (fig. 58, Dac). Sexual differentiation: Male pereopod 7 carpus with proximal sensory spine very strong, acute and decurved, standing on protrusion (fig. 58, PE1-7). Pleopods: Pleopod exopodites 3 rhomboid, pleopod 4 subquadrangular, pleopod 5 triangular, with | to 3 sensory spines laterally, without respiratory area, endopodites bilobate (fig. 59, PL1-5). Sexual differentiation: Male pleopod | exopodite rounded, endopodite slender, transverse groove one third before apex, distal part decurved laterally, bearing a saddle-shaped lobe and several small knobs, caudal row of spines only on last third subapically (fig. 59, PL1). Pleopod 2 with triangular exopodite, bearing two sensory spines laterally, endopodite with distal half flagelliform (fig. 59, PL2), pleopod 5 exopodite with guide slot for apex of pleopod 2 endopodite (fig. 59, PLS). Uropod: As in the other species. Genital appendage: Ventral shield slightly surpassed by ductus ejaculatorii, rather slender (fig. 59, Gen). Comments: A. dalensi sp. n. is a somewhat isolated species within the genus. The male has a strong sensory spine on the carpus of pereopod 7, the pleopod is bent laterally, some knobs are present apically, but no lamellae. The sulcus marginalis of 836 ANDREAS LEISTIKOW the coxal plates is completely reduced. Within this genus these characters can be found only in this species. The outline of the body is more slender than in other congeners. Etymology: This species is dedicated to Dr H. Dalens (Toulouse) in honour of his life’s work on Oniscidea. Androdeloscia digitata sp. n. Material: Brazil: HOLOTYPE d 3.5mm, Amazon rain forest near Manaus, EMBRAGA field project, MNHG coll.; PARATYPES 1% (ovigerous) MNHG coll.; 54 19 limm same data as holotype, author's coll. Colour: Purplish brown with many light spots dorsally on pereon, medial line marked by white spots on pereonite III to VII and pleonites. Cephalothorax: Linea frontails lacking, linea supra-antennalis present, small lateral lobes and slight lamina frontalis, compound eyes consisting of about 8 ommatidia (fig. 60, Ctf). Pereon: Pereonites covered with small tricorn-like setae, tegument smooth, coxal plates without sulcus marginalis and gland pores, noduli laterales long, flagelliform, nodulus of coxal plate IV the most dorsal one (fig. 60, Cx3/Cxp). Pleon: Retracted from pereon, neopleurae of pleonites 3 to 5 conspicuous, pleotelson with straight margins, bearing some prominent tricorn-like setae. Appendages: Antennula: Three-articulate, medial tuft consisiting of about 10 aesthetascs (fig. 61, Anl). Antenna: Fairly slender, peduncular article 2 stout, flagellum three-articulate, joints subequal in length, apical organ half as long as flagellum (fig. 60, An2). Mandible: Pars molaris consisting of a four-branched molar penicil, pars intermedia bearing two penicils on left and one on right mandible, intermedial penicil slender (fig. 61, Mdr). Maxillula: Medial endite bearing two penicils apically, lateral tip, lateral endite with 4+4 teeth, inner set cleft, subapical tooth caudally (fig. 61, Mx1). Maxilla: Lateral lobe slightly broader than medial one, bearing scattered pecti- nate scales, medial endite apically with 5 cusps (fig. 61, Mx2). Maxilliped: Basipodite with sulcus lateralis, palp with one small seta on proximal article, endite with knob-like penicil rostrally and two strong teeth caudally (fig. 61, Mxp). Pereopods: As in the preceding species, dactylar seta rather short (fig. 62, Scl/Dac). Sexual differentiation: Male pereopod 7 with rostral protrusion on mediodistal margin, half as long as carpus with distally inserted sensory spine (fig. 62, PE1-7). Pleopods: Pleopod exopodites subtriangular, laterally bearing 2 to 4 sensory spines, endopodites slightly bilobate, no respiratory areas (fig. 63, PL1-5). Sexual differentiation: Male pleopod 1 with pointed exopodite, medially widely rounded, endopodite with small distal protrusion of basal part, distal third A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 837 slender, apex slightly bulbous, crenulate, with hyaline lamellae caudally and rostrally directed lobe (fig. 63, PL1). Pleopod 2 exopodite triangular with two lateral sensory spines, exopodite with flagelliform distal half (fig. 63, PL2), pleopod 5 with medial guide slot and distinctly pointed apex (fig. 63, PLS). Uropod and genital appendage: As in other species of the genus (fig. 63, Gen). Comments: Another member of Androdeloscia gen.n. bearing a lobe on the male merus 7, but different to the preceding species, in A. digitata sp.n. the lobe is very long, directed distally and bears a sensory spine at its tip. Together with its characteristically shaped male pleopod 1 endopodite, the species cannot be confused with other species. The lack of a sulcus marginalis is most probably explained as covergence to other species of Androdeloscia gen. n. Etymology: The species name is derived from the latin "digitus" which means "finger" and refers to the protrusion on the male pereopod 7 merus. PHYLOGENY AND BIOGEOGRAPHY Within the genus, some distinct groups can be traced. The phylogeny of the whole genus shall be elucidated in the following, the relationships are figured in fig. 65. For the outgroup comparison the species Prosekia rutilans (Vandel, 1952), Andenoniscus silvaticus Verhoeff, 1941 and Ischioscia martinae Leistikow, 1997 were used. As stated above, Androdeloscia gen. n. belongs to a group of South American "philosciids" which are characterized by the peculiar shape of the antennula. The autapomorphies of the genus are the reduction of the linea frontalis and the shape of the pleopod 5 exopodite in the male with its guide slot structure (character set 1). The former character is not strongly supporting the monophyly since it is a reductive character. Thus, it can easily occur in evolution by a single substitutional event in the genome. The latter character is rather complex and may thus warrant monophyly. The evolution of several characters are of importance within this genus: sexual differen- tiations of the pereopods and the structure of the male pleopod 1. The primitive character state of the male pleopod | endopodite is characterized by a set of hyaline lamellae on the medial margin of the apex. The most basal species of this genus is A. dalensi which shows the autapomorphies of the genus but has a serrate distal sensory spine of the propus | and a mandibular penicil composed of more than 5 branches, both plesiomorphic characters even found in P. rutilans; autapomorphies are the reduction of both lamina frontalis and hyaline lamellae on the male pleopod 1 endo- podite. A. hamigera is the next derived species in the shape of the sensory spine (character 2) but has still a prominent molar penicil. All the other species are united by the molar penicil consisting of a long and three short branches, a reductive character but may be used for the phylogenetic reconstruction because of the correspondence of the number of branches and their length (character 3). Half of the species, the members of group A, are equipped with meral lobes and humps in the male, the pleopod | exopodite is enlarged, bearing a small point apically (character 838 ANDREAS LEISTIKOW set 4). The former character can also be found in the genus Andenoniscus Verhoeff, 1941. Since the latter genus 1s excluded from the monophyletic taxon Androdeloscia by the plesiomorphic character state of the male pleopod 5 as far as it is known and the slightly different shape of the cephalothorax, an autapomorphy of Andenoniscus, there is evidence for either an analogy or the expression of a silent gene in both genera (cf. STURM 1994). Additionaly the lobes are different in their position in Andenoniscus and Androdeloscia. Within group A, four species can be separated in group B by having the male pleopod | endopodite with slight lateral humps (character 5). All the species save A. digitata have this slight hump more prominent and somewhat enlarged distally (character 6). The species A. silvatica (Lemos de Castro & Souza, 1986) and A. pseudosilvatica have the synapomorphy of three lobes on the merus, one subproximally and two distally on the medial margin (character 7). The following four species, group C, the adelphotaxon of the preceding ones, are charac- terized by these apomorphies (character set 8): Distal sensory spine of propus | with one tip, pleopod 1 endopodite apically bulbous, pleopod | exopodite with distal point laterally (not medially as in pleopod 2, the exopodite looking "inverted"). In A. ferrarai the lobes of the merus are slight, the pleopod 1 endopodite is straight, the plesiomorphic character state, while A. faitii, A. merolobata, A. longiunguis the endo- podite is twisted, the spermatic channel ending on the rostral side, males have fields of cuticular scales on the merus of pleopod 2 to 6 (character set 9). In this group, A. merolobata and A. longiunguis most probably are sister species, with having strong meral lobes and a pleopod 2 exopodite which is distally obtusely rounded (character set 10). The latter character is less evolved in A. faitii and the meral lobe is missing, this is most parsimonously explained by reduction of this structure. Another phyletic lineage heads to a species group of Caribbean and Amazo- nian distribution, the group D: The last five species are united by the lack of hyaline lamellae on the male pleopod | endopodite, all the species share a rather slender maxilliped with the distolateral edge of the basipodite almost rectangular and not rounded as in the other species (character set 11). In the next evolutionary step the knobs and bumps are concentrated apically on both sides of the endopodite, the apex looking like a fir cone (group E, character 12). Within this group, A. feistae and A. plicatipus are sister species. They are united by the shape of the basal part of pleopod 1 endopodite, it is broad but then narrowed laterally. Thus, the part containing M49 is short compared to the other species (character 13). The adelphotaxon of group E are A. poeppigi and A. malleus with the male pleopod 1 endopodite bearing two strong lobes which are twisted out of the sagittal plane, the spermatic channel is medially open (character set 14). From the biogeographic point of view, the different groups are found in the Amazonian and Caribbean subregion of the Neotropical realm. The basal species, A. dalensi, A. hamigera and A. opercularis are distributed in Venezuela, which is part of the Caribbean subregion. The group of species with meral lobes can be found in most of Amazonia and even in Venezuela, they may have been split off in the two groups with the species of the group C are exclusively found in the foothills of the Andes, part of the Amazonian subregion, as are the members of the group D, while their A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 839 sister group (group B) is distributed in Venezuela and the central Amazonian sub- region. Interestingly, A. si/vatica can be found in both subregions; in Venezuela it was found in the east, on Peninsula de Paria, while its sister species was found in the western part of Venezuela in the foothills of the Cordillera de Mérida. They may vicariate each other. The species from Amazonia are more basal and may be close to A. dalensi save A. tarumae (Lemos de Castro, 1984) which has two lobes on the male merus 7, and is a link between A. digitata and A. silvatica. The other species are described in LEMOS DE CASTRO & SOUZA (1986) and Lima (1997). Until now, the collecting of Oniscidea in South America was very scarce. Therefore, our knowledge on the distribution of genera is rather patchy. As one can deduce from the above mentioned, the genus Androdeloscia gen. n. is widely distri- buted in the Amazonian and Caribbean subregion of the Neotropis. From the point of our knowledge, it has a radiation centre in the eastern slopes of the Andes, where many species can be found occurring sympatrically. They may be found in different habitat, but there is more research needed to elucidate this question. The relatively high diversity in the Andean foothills was explained by the successive settlement by forms from the Amazon valley via dispersal along rivers and colonization by forms from the higher Andean regions. Possibly, a mix of species from both altitudinal regions can be found in this area. Such a faunal composition was postulated by SIMPSON & HAFFER (1978), since these areas have strongly been affected by both climatic changes and oreographic transformantions from the Tertiary on. The species in this area have the most diverse pleopodal structures in the males. This diversity must be explained by evolving in isolated habitats for a considerable period. Once the species barriers were established, there was a selective pressure to evolve unequivocal copulatory devices, a process of character displacement, selecting the extremes within the intra-specific variation. Hence, the some most complex male pleopods can be found in the eastern slopes of the Andes with their high number of sympatric species while in Venezuela the male pleopod | endopodites are structurally simple. ACKNOWLEDGEMENTS For the loan of the material, the author thanks Dr H. Schmafuss, Staatliches Museum fiir Naturkunde, Stuttgart and Dr P. S. Young, Museo Nacional, Rio de Janeiro. He is indepted to Prof. Dr J. W. Wägele and Dr C. Schmidt, Ruhr-Universität Bielefeld both for giving the opportunity to examine the collections made by them in Venezuela and for fruitious discussion on the manuscript. 840 ANDREAS LEISTIKOW + Tr 200um Cxp FIG. 1 Androdeloscia hamigera (Vandel, 1952). Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 84] Fic. 2 Androdeloscia hamigera (Vandel, 1952). Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx! maxillula with detail of apex of lateral endite; Mx2 maxilla. 842 ANDREAS LEISTIKOW Fic. 3 Androdeloscia hamigera (Vandel, 1952). Dac dactylus 1 in rostral view; PE1/4 pereopods 1 (rostral view) and 4 (caudal view); PL1-5 pleopods 1 to 5, with details of pleopod 1 endopodite; Scl ornamental sensory spine of carpus 1; Sp! distal sensory spine of propus 1. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 843 100um * _ AS NS È à 100um NX i —y Fic. 4 Androdeloscia conipus sp. n. holotype d 4mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx4 coxal plate 4; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 844 ANDREAS LEISTIKOW Mx2 Fic. 5 Androdeloscia conipus sp. n. holotype ¢ 4mm. Mdl/r left and right nvandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 845 m 100um FIG. 6 Androdeloscia conipus sp. n. holotype d 4mm. Dac dactylus 1 in rostral view; PE1-7 pere- opods 1 (rostral view), 2, 6, 7 (caudal view); Sci ornamental sensory spine of carpus 1; Spl distal sensory spine of propus 1; Sp2 sensory spine of propus 2. 846 ANDREAS LEISTIKOW 100um m 100um Fic. 7 Androdeloscia conipus sp. n. holotype d 4mm. PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 847 200um Fic. 8 Androdeloscia feistae sp. n. holotype d 3mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales: Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 848 ANDREAS LEISTIKOW Fic. 9 Androdeloscia feistae sp. n. holotype d 3mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx! maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 849 Androdeloscia feistae sp. n. holotype ¢ 3mm. Dac dactylus | in rostral view; PE1-7 pereopods 1, 6, 7 (caudal view) with carpus 1 in rostral view; PLx pleopod 5 exopodite of female (Paratype 3.5mm); Scl ornamental and medial sensory spine of carpus 1; Sc6/7 sensory spines of carpus 6/7; UR uropod. 850 ANDREAS LEISTIKOW 100um PL4 Fic. 11 Androdeloscia feistae sp. n. holotype d 3mm. PL1-5 pleopods 1-5, rostral view with detail of endopodite | in caudal view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 85] 300um Fic. 12 Androdeloscia plicatipus sp. n. holotype d 3mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx4 coxal plate 4; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 852 ANDREAS LEISTIKOW Fic. 13 Androdeloscia plicatipus sp. n. holotype d 3mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 100um Androdeloscia plicatipus sp. n. holotype d 3mm. Dac dactylus 1 in rostral view; PE1-7 pereopods 1, 3, 6, 7 (caudal view), with detail of carpus | in rostral view; Scl ornamental sensory spine of carpus 1; Sc6 sensory spine of carpus 6; Sm6 sensory spine of merus 6; Spl distal sensory spine of propus 1. 853 854 ANDREAS LEISTIKOW 100um 100um PL3 PL4 Fic. 15 Androdeloscia plicatipus sp. n. holotype d 3mm. PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 855 200um 200um Fic. 16 Androdeloscia taitii sp. n. holotype ¢ 4mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales: Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; UR uropod. 856 ANDREAS LEISTIKOW FIG. 17 Androdeloscia taitii sp. n. holotype d 4mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 857 Fic. 18 Androdeloscia taitii sp. n. holotype d 3.5mm. Dac dactylus 4 in rostral view; PE1-4 pereopods 1-4 (caudal view), with detail of carpus 1 in rostral view; Scl ornamental and longest sensory spine of carpus 1; Sc2 sensory spine of carpus 2; Sd4 dactylar seta of dactylus 4. 858 ANDREAS LEISTIKOW Fic. 19 Androdeloscia taitii sp. n. holotype d 4mm. PES-7 pereopods 5-7 (caudal view), with details of merus 6 in rostral and caudal view; Si7 lateral sensory spine of ischium 7; Sm7 sensory spine of merus 7; Sb5 tricorn-like seta of basis 5. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 859 Fic. 20 Androdeloscia taitii sp. n. holotype ¢ 4mm. PL1-5 pleopods 1-5, rostral view, with details of endopodite | in caudal and rostral view. 860 ANDREAS LEISTIKOW 200um Fic. 21 Androdeloscia merolobata sp. n. holotype d 5mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 861 Fic. 22 Androdeloscia merolobata sp. n. holotype d Smm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite in caudal and rostral view; Mx2 maxilla. 862 ANDREAS LEISTIKOW Fic. 23 Androdeloscia merolobata sp. n. holotype 4 5mm. Dac dactylus 4 in rostral view; PE1-4 pereopods 1-4 (caudal view), with detail of carpus 1 in rostral view; Scl ornamental and longest sensory spine of carpus 1; Sc7 sensory spine of carpus 7. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 863 Fic. 24 Androdeloscia merolobata sp. n. holotype & Smm. PES-7 pereopods 5-7 (caudal view), with details of lobes on merus 7 in rostral and caudal view; UR uropod. 864 ANDREAS LEISTIKOW JES \ 200um FIG. 25 Androdeloscia merolobata sp. n. holotype ¢ 5mm. PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 865 Fic. 26 Androdeloscia longiunguis sp. n. holotype d 4mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view. 866 ANDREAS LEISTIKOW Fic. 27 Androdeloscia longiunguis sp. n. holotype ¢ 4mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx! maxillula with detail of apex of lateral endite in caudal and rostral view; Mx2 maxillula. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 867 en —4 ] 00um Fic. 28 Androdeloscia longiunguis sp. n. holotype d 4mm. Dac dactylus 4 in rostral view; PE1-7 pereopods I(rostral view), 4, 5, 7 (caudal view), with detail of merus 7 in rostral view; Scl ornamental and longest sensory spine of carpus 1; Sp! distal sensory spine of propus 1. 868 ANDREAS LEISTIKOW Fic. 29 Androdeloscia longiunguis sp. n. holotype ¢ 4mm. PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 869 500um 100um 4 el 50um VII NI Cx3 Fic. 30 Androdeloscia ferrarai sp. n holotype d Smm. Anl antennula; An? antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 870 ANDREAS LEISTIKOW Fic. 31 Androdeloscia ferrarai sp. n. holotype d 5mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 871 Fic. 32 Androdeloscia ferrarai sp. n. holotype d 5mm. Dac dactylus 3 in rostral view; PEI-4 pereo- pods 1-4 (caudal view), with detail of carpus | in rostral view, and detail of merus 3; Scl ornamental and longest sensory spine of carpus 1; Sc2 sensory spine of carpus 2; Sd3 dactylar seta of dactylus 3; Spl distal sensory spine of propus 1. 872 ANDREAS LEISTIKOW Fic. 33 Androdeloscia ferrarai sp. n. holotype d 5mm. PES-7 pereopods 5-7 (caudal view), wit details of merus 6 in rostral and caudal view; Sb5 tricorn-like seta of basis 5; Sc6 lateral sensory spine of carpus 6; Sm7 sensory spine of merus 7; UR uropod. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 873 100um 4 e 9 4 H D 2 z 4 dg a [4 Fic. 34 Androdeloscia ferrarai sp. n. holotype d Smm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. 874 ANDREAS LEISTIKOW Fic. 35 Androdeloscia poeppigi sp. n. holotype 4 2.5mm. Anl antennula; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx2 coxal plate 2; Had habitus in dorsal view; Hal habitus in lateral view; UR uropod. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 875 Mx2 Fic. 36 Androdeloscia poeppigi sp. n. holotype d 2.5mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral enditein caudal and rostral view; Mx2 maxillula. 876 ANDREAS LEISTIKOW 100um Scl Fic. 37 Androdeloscia poeppigi sp. n. holotype d 2.5mm. Dac dactylus 4 in rostral view; PE1-7 pereopods I(rostral view), 2, 6, 7 (caudal view); Scl ornamental and longest sensory spine of carpus 1; Sc2 sensory spine of carpus 2; Sp! distal sensory spine of propus 1. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 877 Fic. 38 Androdeloscia poeppigi sp. n. holotype d 2.5mm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and laterorostral view. 878 ANDREAS LEISTIKOW Fic. 39 Androdeloscia malleus sp. n. holotype ¢ 3mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 879 SOum Mx2 Fic. 40 Androdeloscia malleus sp. n. holotype d 3mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx! maxillula with detail of apex of lateral endite; Mx2 maxilla. 880 ANDREAS LEISTIKOW Fic. 41 Androdeloscia malleus sp. n. holotype d 3mm. Dac dactylus 3 in rostral view; PE1-7 pere- opods I(rostral view), 4, 6, 7 (caudal view); Scl ornamental and longest sensory spine of carpus 1; Spl distal and medial sensory spine of propus 1. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 881 m 100um PL4 100um Fic. 42 Androdeloscia malleus sp. n. holotype d 3mm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with details of endopodite | in caudal and rostral view. 882 ANDREAS LEISTIKOW — == 100um PL2 PLI BIE2 Fic. 43 Androdeloscia malleus sp. n. holotype ¢ 3mm. PL1-2 pleopods 1-2, rostral view, with details of endopodite | in caudal and mediorostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 883 Fic. 44 Androdeloscia opercularis sp. n. holotype d 3mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx1 coxal plate 1; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 884 ANDREAS LEISTIKOW Fic. 45 Androdeloscia opercularis sp. n. holotype d 3mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 885 Fic. 46 Androdeloscia opercularis sp. n. holotype d 3mm. Dac dactylus 3 in rostral view; PEI-7 pereopods 1, 2, 6, 7 (caudal view), detail of carpus 1 (rostral view); Scl ornamental and longest sensory spine of carpus 1; Spl distal and medial sensory spine of propus 1. 886 ANDREAS LEISTIKOW FIG. 47 Androdeloscia opercularis sp. n. holotype d 3mm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 887 Fic. 48 Androdeloscia silvatica (Lemos de Castro & Souza, 1986) 3 3.5mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 888 ANDREAS LEISTIKOW Fic. 49 Androdeloscia silvatica (Lemos de Castro & Souza, 1986) ¢ 3.5mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 889 Fic. 50 Androdeloscia silvatica (Lemos de Castro & Souza, 1986) 4 3.5mm. Dac dactylus 1 in rostral view; PEI-7 pereopods 1 (rostral view), 2, 6, 7 (caudal view), with detail of merus 7; Scl ornamental and longest sensory spine of carpus 1; Spl distal and medial sensory spine of propus |. 890 ANDREAS LEISTIKOW Fic. 51 Androdeloscia silvatica (Lemos de Castro & Souza, 1986) 4 3.5mm. Gen genital papilla; PL1- 5 pleopods 1-5, rostral view, with detail of endopodite I in caudal view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 891 FiGy a2 Androdeloscia pseudosilvatica sp. n. holotype ¢ 3mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 892 ANDREAS LEISTIKOW Fic. 53 Androdeloscia pseudosilvatica sp. n. holotype ¢ 3mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 893 RES INN SAS ST SES PVP C Fic. 54 Androdeloscia pseudosilvatica sp. n. holotype d 3mm. Dac dactylus 3 in rostral view; PE1-7 pereopods 1 (rostral view), 4, 6, 7 (caudal view), with detail of merus 7; Scl ornamental and longest sensory spine of carpus 1; Sp! distal and medial sensory spine of propus 1; Sp7 sensory spine of propus 7. 894 ANDREAS LEISTIKOW Fic. 55 Androdeloscia pseudosilvatica sp. n. holotype d 3mm. Gen genital papilla; PL1-5 pleopods 1- 5, rostral view, with detail of endopodite 1 in caudal view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 895 Fic. 56 Androdeloscia dalensi sp. n. holotype ¢ 3.5mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 896 ANDREAS LEISTIKOW FIG. 57 Androdeloscia dalensi sp. n. holotype 4 3.5mm. Mdl/r left and right mandible; Mxp maxilliped with detail of endite in rostral view; Mx1 maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 897 Fic. 58 Androdeloscia dalensi sp. n. holotype d 3.5mm. Dac dactylus 1 in rostral view; PEI-7 pereopods 1 (rostral view), 2, 5, 7 (caudal view), with detail of carpus 7 in rostral view; Scl ornamental sensory spine of carpus 1; Spl distal sensory spine of propus 1. 898 ANDREAS LEISTIKOW Fic. 59 Androdeloscia dalensi sp. n. holotype & 3.5mm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with detail of endopodite | in caudal view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 899 Fic. 60 Androdeloscia digitata sp. n. holotype d 3mm. An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Cxp coxal plates with position of noduli laterales; Cx3 coxal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 900 ANDREAS LEISTIKOW 7 SOum — Fic. 61 Androdeloscia digitata sp. n. holotype d 3mm. Anl antennula; Mdr right mandible; Mxp maxilliped with detail of endite in rostral view; Mx! maxillula with detail of apex of lateral endite; Mx2 maxilla. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 901 FIG. 62 Androdeloscia digitata sp. n. holotype d 3mm. Dac dactylus 1 in rostral view; PE1-7 pere- opods 1 (rostral view), 2, 6, 7 (caudal view), with detail of merus 7 in rostral view; Scl ornamental sensory spine of carpus 1; Spl distal sensory spine of propus 1. 902 ANDREAS LEISTIKOW Fic. 63 Androdeloscia digitata sp. n. holotype ¢ 3mm. Gen genital papilla; PL1-5 pleopods 1-5, rostral view, with detail of endopodite | in caudal and rostral view. A NEW GENUS OF SOUTH AMERICAN ONISCIDEA 903 Fic. 64 Androdeloscia ferrarai sp. n. Above left: male pleopod 5 exopodite from caudal; above right: guide slot of pleopod 5 with pectinate scales. - Androdeloscia merolobata sp. n. Below left: antennula, below right: nodulus lateralis of coxal plate 3; scale bars: 50 um. 904 ANDREAS LEISTIKOW A. dalensi A. hamigera A. digitata A. opercularis A. silvatica 6 Vv q dnoin 5 4 7 A. pseudosilvatica A. ferrarai 8 A. taitii 9 A. merolobata v dnoip 3 10 A. longiunguis A. conipus 1 A. feistae 13 A. plicatipus 11 A. malleus 14 A. poeppigi FIG. 65 Phylogenetic relationships within the genus Androdeloscia gen. n., explanation of apomorphic characters see text. qdnoip | I dnoip | a dnoıy REFERENCES ERHARD, F. 1997. Das pleonale Skelet-Muskel-System von Titanethes albus und weiterer Taxa der Oniscidea mit Schlußfolgerungen zur Phylogenie der Landasseln. Stuttgarter Beiträge zur Naturkunde, Serie B (Biologie) 550: 1-70. LEISTIKOW, A. 1998. Redescriptions of terrestrial Isopoda from Chile and Peru (Crustacea: Isopoda: Oniscidea). Spixiana 21(3): 215-225. LEMOS DE CASTRO, A. 1984. Uma nova espécie de Prosekia de uma florestia inundavel na Amazonia Central. Amazoniana 8: 441-445. LEMOS DE CASTRO, A. & SouzA, L. A. 1986. Trés espéces novas de isopodes terrestres do género Prosekia Vandel da Amazonia Brasileira. Revista Brasileira de Zoologia 46: 429-438. Lima, I. M. B. 1997. Uma nova espécie de Prosekia Vandel, 1968 da Amazonia Brasileira (Crustacea: Isopoda: Philosciidae). Amazoniana 14: 101-104. SCHMALFUSS, H. 1980. A revision of the neotropical genus Ischioscia Verhoeff, with descrip- tion of four new species. Studies on Neotropical Fauna and Environment 15: 125-139. SCHMALFUSS, H. 1990. Die Landisopoden Griechenlands. 11. Beitrag: Gattung Chaeto- philoscia. Revue suisse de Zoologie 97: 169-193. SIMPSON, B. B. & HAFFER, J. 1978. Speciation patterns in the Amazonian forest biota. Annual Revue of Ecology and Systematics 9: 497-518. STURM, H. 1994. Diskontinuierliche Gene und Evolution. Zeitschrift fiir zoologische Systematik und Evolutionsforschung 32: 241-263. VANDEL, A. 1952. Etude des isopodes terrestres récoltes au Vénézuela par le Dr. G. Marcuzzi. Memorias del Museo civico di Storia Naturale in Verona 3: 59-203. VANDEL, A. 1968. Isopodes terrestres. Jn: LELEUP N. and J. (eds). Mission zoologique belge aux Iles de Galapagos et Ecuador 84: 35-168. VANDEL, A. 1972. Les isopodes terrestres de la Colombie. Studies on Neotropical Fauna and Environment 7: 147-172. VERHOEFF, K. W. 1908. Neue Isopoden-Gattungen. Zoologischer Anzeiger 33: 520-525. REVUE SUISSE DE ZOOLOGIE 106 (4): 905-912; décembre 1999 Sphaeropsocopsis myrtleae sp. n., a blind subterranean psocid from St Helena (Psocoptera: Sphaeropsocidae) Charles LIENHARD! & N. Philip ASHMOLE? | Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. 2 University of Edinburgh, Division of Biological Sciences, West Mains Road, Edinburgh EH9 3JT, U. K. Sphaeropsocopsis myrtleae sp. n., a blind subterranean psocid from St Helena (Psocoptera: Sphaeropsocidae). - The new species is described and illustrated, based on a female captured by a modified pitfall trap ("boot trap") in a lava tube on the mid-Atlantic island of St Helena. The species is adapted to subterranean life in being depigmented and blind. This is the first case of complete reduction of compound eyes known in the insect order Psocoptera. Key-words: Psocoptera - Sphaeropsocidae - Sphaeropsocopsis - eye reduction - cave species - St Helena island. INTRODUCTION The Psocoptera of St Helena are poorly known. 12 species have been recorded by BADONNEL (1976); one of them has also been described under a synonym by NEw (1977). All this material has been collected on vegetation or in light traps, none from subterranean habitats. Among the several psocids collected by N. P. & M. J. Ashmole in such habitats one troglobitic species of the suborder Troctomorpha is of particular interest and will be described in the following. During the period December 1994 to May 1995 N. P. Ashmole and M. J. Ashmole carried out intensive field work on the subterranean fauna of the island of St Helena, in mid-Atlantic in 5°W, 15°S. This work was stimulated by earlier studies on dispersal of island arthropods (ASHMOLE & ASHMOLE 1988) and on the fauna of lava and cave habitats on other volcanic Atlantic islands (ASHMOLE & ASHMOLE 1987, OROMI et al. 1990, ASHMOLE et al. 1992). Geologically youthful volcanic islands such as Ascension sometimes have closely related endemic species respectively in barren lava habitats on the surface and in volcanic caves and smaller underground spaces (ASHMOLE 1994, ASHMOLE & ASHMOLE 1997, CHRISTIANSEN 1998). A degree of troglomorphy [morphological Manuscript accepted 25.08.1999 906 C. LIENHARD & N. P. ASHMOLE specialization for subterranean life (CHRISTIANSEN 1962)] may be shown by caverni- colous species on such islands [e.g. pseudoscorpions (MAHNERT 1993); spiders (MERRETT & ASHMOLE 1997); psocids (LIENHARD 1996)] but is typically more extreme in caves on older islands, where true troglobites (species obligately asso- ciated with the deep cave zone) are often found (OROMI er al. 1991). However, underground caves and cracks formed at the time of volcanic eruptions are relatively ephemeral (HOWARTH 1981, ASHMOLE er al. 1992, HocH & ASCHE 1993). Animals may evolve specializations to life in them, but as the caves collapse and the cracks become filled with soil, troglobites on islands may become extinct or survive in only a few localities. The recent work on St Helena was designed to find any specialized caverni- colous arthropods that might have survived the period of about seven million years since volcanic activity on the island ceased (BAKER 1970); the species here described is the only unequivocal example encountered. Few lava tubes survive and near-sur- face subterranean spaces are inhabited largely by introduced soil animals (ASHMOLE & ASHMOLE, unpublished). METHODS AND STUDY SITES The traps used for sampling the fauna of underground spaces on St Helena were of three types: "pipe traps" (described in ASHMOLE & ASHMOLE, in press a), conventional plastic straight-sided pitfall traps (ASHMOLE et al. 1992) and specially designed "boot traps"; it was in one of the latter that the blind psocopteran was captured. The boot traps are a form of pitfall trap modified for use in caves and other situations where it is impractical to bury traps with the rim flush with the substrate; they are ceramic and were commissioned from a local potter. The trap weighs a little under 400 g and is boot-shaped in profile, with base dimensions c. 125 mm long, c. 70 mm wide and c. 80 mm high. The "heel" of the boot is in the form of a straight-sided pitfall trap with internal dimensions c. 70 mm deep and c. 45 mm diameter; the "toe" forms a ramp up which arthropods can climb. The ramp is coarsely roughened to mimic the surface of lava or cinders but the interior of the trap is smooth and glazed; a broad glazed lip slopes down from the top of the ramp into the trap; on the other side of the trap opening the narrow roughened rim meets the glazed interior at a right- angle. Each trap contains c. 25 ml of modified Turquin's liquid consisting of 10 g chloral hydrate, 5 ml formalin, 5 ml glacial acetic acid, 1 ml liquid detergent, plus beer (preferably including one third stout) to make 1 litre. In addition a bait of "Danish Blue" cheese is placed in a tiny plastic cup formed by an inverted specimen tube lid impaled on a nail with its point resting on the bottom of the trap, thus holding the bait dry above the level of the liquid. Most of the work on St Helena involved setting traps in the network of small underground cracks and crevices characteristic of basaltic lava and termed the "MSS" or mesocavernous shallow stratum (JUBERTHIE et al. 1980, MEDINA & OROMI 1990, SPHAEROPSOCOPSIS MYRTLEAE SP. N. 907 ASHMOLE 1994). However, traps were also placed in the largest among the few accessible lava tubes, and it was in this cave that the blind psocid was captured. The cave, referred to as Rupert's Battery Cave, is at UTM Grid Zone 30, TH O88 386 in northwest St Helena. The opening is at a height of roughly 50 m beside a path traversing a very steep and rocky slope above the shore between Rupert's Bay and James Bay. The opening had been blocked by a large rock which was moved in order to gain access to the cave. The cave slopes downwards very steeply to the north and may continue to sea level, but has been blocked with rubble at about four metres from the top, probably by people trying to make it less dangerous. The traps were set on the top of the rubble, mainly close to the cave walls, and were in semi-darkness; further down, the cave is presumably completely dark and also humid. Other arthropods caught in the cave included an unidentified and possibly endemic species of mogoplistine cricket, together with introduced species of blaniulid Diplopoda and lepismatid Thysanura (Zygentoma). The following abbreviations are used in the description: B = body length; V = width of head capsule on vertex; FW = forewing length; F+tr = length of femur and trochanter (hindleg); T = length of tibia (hindleg); tl, t2, t3 = length of tarsomeres, measured from condyle to condyle (hindleg); MHNG = Muséum d'histoire naturelle de Genève. DESCRIPTION Sphaeropsocopsis myrtleae sp. n. (9) Figs 1-8 Material. Holotype © (slide mounted, MHNG). St Helena: Rupert's Battery Cave, 13- 17 March 1995, modified pitfall trap ("boot trap"), leg. N. P. & M. J. Ashmole (sample: 680 SH). Etymology. The species is dedicated to Myrtle Ashmole, who in 1984 suggested studying the fauna of barren volcanic terrain in the Canary Islands and who participated in all the field work on St Helena. Coloration. Body white to very slightly yellowish, no subcuticular pigment, cuticle transparent, the light brown mandibules visible through the head capsule. Wings, legs and antennae colorless. Sculpture. Vertex with fine granular tubercles, their diameter distinctly smaller than the diameter of the alveoli of the small vertical hairs. Tubercles evenly dis- tributed, no distinct spindle-shaped or polygonal areas recognizable. Sculpture of thoracic tergites similar to head. Morphology. Compound eyes absent, no ocelli. Head capsule normally shaped but very slightly concave laterally where in the other species of the genus the compound eyes are located (fig. 1). Pilosity of head short, with a pair of distinctly longer hairs near antennal base and another pair on postclypeus, three of these hairs are broken but localizable by their large alveoli (fig. 1). Vertical suture well deve- loped, frontal suture only visible as a slight interruption of sculpture. Both antennae broken, only a few basal segments left. Maxillary palps lacking (broken). Pilosity of labial palps badly damaged, but a particularly well developed club-like sensillum 908 C. LIENHARD & N. P. ASHMOLE Fics 1-8 Sphaeropsocopsis myrtleae sp. n., female: 1, head in frontal view; 2, forewing; 3, reticulate sculpture in the middle of forewing (same magnification as fig. 2); 4, labial palpus (pilosity not represented except large lateral sensillum); 5, lacinial tip; 6, spermapore; 7, claw; 8, T-shaped sclerite of subgenital plate. present on outer side (fig. 4). Lacinial tip largely bifurcate, outer denticle with two secondary denticles on half-length on its inner side (fig. 5). Thoracic tergites not divided into lobes. Forewings shortened, reaching about the middle of the abdomen, hindwings absent. Venation of forewing strongly reduced, two longitudinal rows of hairs indicate the presence of two faint longitudinal main veins, wing margins also rather densely pubescent, all hairs short (fig. 2). Wing membrane with a very faint network of fine sculptural lines delimiting more or less hexagonal areas (only visible at high magnifications under phase contrast) (fig. 3). SPHAEROPSOCOPSIS MYRTLEAE SP. N. 909 Legs relatively long (index T/V = 1.4), tibiae with two apical spurs, claws slender, with one preapical denticle and some microtrichia on outer side (fig. 7). No coxal organ present. Terminalia without particular diagnostic characters, T-shaped sclerite of subgenital plate well developed (fig. 8), spermapore on both sides with a field of microtrichia (fig. 6). The spermatheca is full of sperm, therefore we can affirm that the species is bisexual, even if the male is not yet known. Measurements (um). B = 1350; V = 315; FW = 580; F+tr = 360; T = 450; tl = 5052043, — 60! DISCUSSION At present the new species has to be placed in the genus Sphaeropsocopsis Badonnel, 1963 because mesothoracic sternites are not subdivided into lobes and because of the eye reduction. In the very closely related genus Badonnelia Pearman, 1953 females always have compound eyes with 7 (exceptionally 6 or 8) ommatidia and the mesothoracic sternites show some lobation. The generic characters of the forewing (shape and venation) can not be observed in the new species because of its strong brachyptery. According to BADONNEL (1963), the form of the terminal segment of the maxillary palp is also of some value to decide about generic attribution; unfortunately both palps are broken in the unique female of the new species. Within the genus Sphaeropsocopsis the number of ommatidia in females can vary from 3 to 10, but intraspecific variability is usually low. Even in S. microps, the only species of the genus with 3 ommatidia (sometimes 4), these remaining omma- tidia are well developed and situated on a hemispherical protuberance of the head capsule, as usual in the species with higher numbers of ommatidia. The complete reduction of the compound eyes in S. myrtleae clearly represents an adaptation to cave life and is the first case of blindness known in Psocoptera. Several troglophilic psocids have been mentioned by BADONNEL & LIENHARD (1994), some of them show strong reduction of compound eyes but a blind species has never been observed up to now. Even in the edaphic species of the apterous genus Liposcelis Motschulsky, 1852 (belonging to the Liposcelididae, family closely related to the Sphaeropsocidae), where strong eye reduction occurs, at least two ommatidia always persist, forming together with some underlying pigment a reduced but probably functional compound eye (cf. LIENHARD 1998). The only known troglobitic liposcelidid, Troglotroctes ashmoleorum Lienhard, 1996, has usually 3 (rarely 4 or 2) ommatidia (cf. LIENHARD 1996). In S. myrtleae however, the head capsule shows no subcuticular pigmentation or irregularity of sculpture at the place where eyes are usually located. Unfortunately the antennae and the maxillary palps of the single specimen known are broken. Therefore it is impossible to say if in S. myrtleae the optical deficiency could eventually be compensated by tactile sense organs, as long antennae and palps with particularly well developed sensilla. But it is interesting to see that the club-like outer sensillum of the labial palp is extremely well developed in this species, significantly greater than is usual in Troctomorpha. In the troglobitic Troglotroctes 910 C. LIENHARD & N. P. ASHMOLE ashmoleorum, known from subterranean habitats on Ascension Island (ASHMOLE & ASHMOLE 1997) an augmented number of sensilla on maxillary and labial palps has been observed, compared with the usual number in the family Liposcelididae (LIENHARD 1996). The relatively long legs (index T/V = 1.4) of S. myrtleae could probably also be interpreted as an adaptation to cave life, in the other species of the genus this index is <1.0 (cf. BADONNEL 1962, 1963, 1967, 1971, 1972). Another evident adaptation to subterranean life is the lack of pigmentation, which has never been observed to such an extent in the other species of the genus, which are usually litter-dwellers (cf. BADONNEL, papers cited above). The presence of a species of Sphaeropsocopsis on the mid-Atlantic island of St Helena is not really surprising. The genus is known from South America, Ascension Island (mid-Atlantic), Africa, South Australia and Tasmania. A single species is known in each case from the Australian region [S. recens (Hickman, 1934); cf. SMITHERS 1984] and continental Africa [S. reisi Badonnel, 1971, only recorded from Angola (BADONNEL 1971)]. The highest diversity has been observed in South America, where one species is known from Argentina [S. argentina (Badonnel, 1962)] and 5 species from Chile: S. chilensis Badonnel, 1963; S. microps Badonnel, 1963; S. spinosa Badonnel, 1972; S. valdiviensis Badonnel, 1972; S. valeriae Badonnel, 1967. A single damaged female of Sphaeropsocopsis cf. microps has also been recorded from a subterranean habitat on Ascension Island (ASHMOLE & ASHMOLE 1997). St Helena, which is more than 14 million years old (BAKER 1970) has been colonised during its long life by a wide variety of arthropod groups (BASILEWSKY 1985; ASHMOLE & ASHMOLE, in press b). Psocoptera, including even apterous forms such as Liposcelis species, are effective aerial dispersers (THORNTON 1964, THORNTON & HARRELL 1965) and the ancestors of Sphaeropsocopsis myrtleae seem likely to have colonised St Helena by air. There are endemic psocids on many oceanic islands, and on Ascension Island (the closest land to St Helena) there are several apparently indigenous species including a troglobitic generic endemic (7roglotroctes Lienhard, 1996, see above and ASHMOLE & ASHMOLE 1997). ACKNOWLEDGEMENTS Field work by N. P. & M. J. Ashmole on St Helena was made possible by support from the Carnegie Trust for the Universities of Scotland and the British Ecological Society. It is a pleasure also to thank the many people on the island who helped to make the visit so productive and enjoyable. REFERENCES ASHMOLE, M. J. & ASHMOLE, N. P. 1987. Arthropod communities supported by biological fallout on recent lava flows in the Canary Islands. Entomologica Scandinavica, Supplement 32: 67-88. ASHMOLE, N. P. 1994. Colonization of the underground environment in volcanic islands. Mémoires de Biospéologie 20: 1-11. SPHAEROPSOCOPSIS MYRTLEAE SP. N. 9]] ASHMOLE, N. P. & ASHMOLE, M. J. 1988. Insect dispersal on Tenerife, Canary Islands: high altitude fallout and seaward drift. Arctic and Alpine Research 20: 1-12. ASHMOLE, N. P. & ASHMOLE, M. J. 1997. The land fauna of Ascension Island: new data from caves and lava flows, and a reconstruction of the prehistoric ecosystem. Journal of Biogeography 24: 549-589. ASHMOLE, N. 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Air-borne Psocoptera trapped on ships and aircraft. 2. Pacific ship trappings 1963-1964. Pacific Insects 7(4): 700-702. REVUE SUISSE DE ZOOLOGIE 106 (4): 913-927; décembre 1999 New species of Agathidium Panzer from China (Coleoptera, Leiodidae, Agathidiini) Fernando ANGELINI S.S.7 per Latiano, Km. 0,500, I-72021 Francavilla Fontana (Brindisi), Italy. New species of Agathidium Panzer from China (Coleoptera, Leiodidae, Agathidiini). - Records and/or descriptions are given for 13 species of Agathidium from China (Sichuan), deposited in the Geneva Museum and collected by Dr. S. Kurbatov. New species are: Agathidium (Agathidium) modestum sp. n., A. (A.) armatum Sp. n., A. (A.) rubiginosum sp. n., A. (A.) hani sp. n., A. (A.) luojiense sp. n., A. (A.) pseudouniforme sp. n., A. (Microceble) xilingense sp. n., A. (Macroceble) kurbatovianum sp. n., A. (M.) megacephalum sp. n. New records for Sichuan are: Agathidium (Agathidium) brunneipenne Ang. & Dmz. and A. (A.) lugubre Ang. & Dmz. Key-words: Coleoptera - Leiodidae - Agathidium - China - new species - new records. INTRODUCTION A large amount of new data on the Agathidium fauna of China results from a study of the material deposited in the Geneva Museum. The collection treated consists of 102 specimens and 13 species collected in 4 localities of the Sichuan Province. The new data concern: a) 9 new species. b) 2 new records for Sichuan: Agathidium (Agathidium) brunneipenne Ang. & Dmz. and A. (A.) cf. lugubre Ang. & Dmz. The Chinese Agathidiini are moderately known. With the new species are known 80 species, collected mainly in Sichuan, Yunnan and Zhejiang Province. The majority of species are endemic; only 6 specie are present in Taiwan or Oriental Region. The specimens are deposited in the Geneva Museum (MHNG) and in Angelini's collection in Genoa Natural History Museum (AC-GNHM). For methods see ANGELINI & DE Marzo (1981: 188-192). I am indebted to Dr Ivan Löbl for making the material available for study and to Jonathan Cooter (Hereford) for comments on the earlier version of the manuscript. Manuscript accepted 30.04.1999 914 FERNANDO ANGELINI Agathidium Panzer, 1797 Agathidium (Neoceble) dundai Ang. & Svec Agathidium (Neoceble) dundai Angelini & Svec, 1994: 12. Material: China, Sichuan, Xiling Mt., 1600-2400 m, 30.VII-4.VIIL.1996, leg. S. Kur- batov, 1 ex. 2 in MHNG. Distribution: China (Sichuan). Agathidium (Agathidium) modestum sp. n. Figs 1, 7-9, 16 Holotype d: China, Sichuan, env. Xichang, M. Luoji, 2300-2400 m, 16-24.VII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 7 d and 8 © in MHNG, 3 d and 2 © in AC-GNHM. Length 2,6-3,0 mm (holotype d 2,90 mm). Dorsum reddish-brown, venter lighter; antennae uniformly testaceous; legs reddish-brown. Microreticulation absent; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures very small, superficial, separated from each other by 2-10 times their own diameter. Widest at the eyes; eyes flattened (fig. 1); antero-lateral margins distinctly raised; clypeus moderately emarginate; clypeal line absent. 3rd antennal segment 1,27 times as long as 2nd and shorter than 4th + Sth together. Pronotum: Punctures as those on head, separated from each other by 2-15 times their own diameter. 1,59 times as broad as head, moderately broader than long (width/length= 1,55) and very convex (width/height= 1,4); anterior margin weakly curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 0,90 mm, width 1,40 mm and height 1,00 mm. Elytra: Punctures as those on head, separated from each other by 2-5 times their own diameter. As broad as pronotum, a little broader than long (width/length= 1,03) and moderately convex (width/height= 1,79); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,35 mm, width 1,40 mm and height 0,78 mm. Metathoracic wings absent. Meso- and metasternum: median carina absent, lateral lines absent, femoral lines complete, a small tubercle present between the metacoxae; metasternum short, meso- and metacoxae almost touching. Legs: Male hind femora curved at posterior margin (fig. 16). Tarsal formula: d 5-5-4, 2 4-4-4. Aedeagus as in figures 7, 8. Spermatheca as in figure 9. Discussion: Species of the madurense group; Agathidium modestum sp. n. is similar to A. alesi Ang. & Dmz. (from Taiwan) and A. distinguendum Ang. & Dmz. (from Taiwan); from A. distinguendum it differs in shape of eyes, greater length, and in having the pronotum less transverse; from A. alesi it differs only in shape of pronotum, which is less transverse and male hind femora, without tooth at posterior margin. Distribution: China (Sichuan). NEW SPECIES OF AGATHIDIUM 915 Fics 1-6 Head of: 1, Agathidium modestum sp. n., 2, A. armatum sp. n.; 3, A. rubiginosum sp. n.; 4, A. hani sp. n.; 5, A. luojiense sp. n.; 6, A. pseudouniforme sp. n. 916 FERNANDO ANGELINI Agathidium (Agathidium) armatum sp. n. Figs 2, 10-12, 17 Holotype d: China, Sichuan, M. Xiling, 1600-2400 m, 30.VII.-4.VIII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 2 d and 5 © in MHNG, 2 d and 1 2 in AC-GNHM. Length 2,85-3,10 mm (holotype 4 3,00 mm). Dorsum reddish-brown, venter lighter; antennae uniformly testaceous; legs reddish-brown. Microreticulation present only on the elytra, impressed; puncturation fine and sparse on head and pronotum, absent on elytra. Sutural striae absent. Head: Microreticulation absent; punctures small, impressed, separated from each other by 2-5 times their own diameter. Widest at the eyes; eyes flattened (fig. 2); antero-lateral margins uniformly raised; clypeus moderately emarginate; clypeal line absent. 3rd antennal segment 2,36 times as long as 2nd and longer than 4th + Sth together. Pronotum: Microreticulation absent; punctures smaller and more superficial than those on the head, hardly visible, separated from each other by 2-10 times their own diameter. 1,33 times as broad as head, moderately broader than long (width/ length= 1,47) and moderately convex (width/height= 1,47); anterior margin moderately curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 0,95 mm, width 1,40 mm and height 0,95 mm. Elytra: Microreticulation impressed and uniform; without pucturation. Slightly narrower than the pronotum, as broad as long and weakly convex (width/height= 1,87); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,35 mm, width 1,35 mm and height 0,72 mm. Metathoracic wings absent. Meso- and metasternum: median carina sharp, lateral lines absent, femoral lines complete, rounded at middle. Legs: Male hind femora with a sharp tooth at the posterior margin (fig. 17). Tarsal formula: ¢ 5-5-4, © 5-4-4. Aedeagus as in figures 10, 11. Spermatheca as in figure 12. Discussion: Species of the /aevigatum group; Agathidium armatum sp. n. is similar to A. rubiginosum sp. n. (from Sichuan) and A. rufescens Ang. & Dmz. (from Sichuan) in colouring of dorsum, length ratio 3rd/2nd and in microreticulation only on elytra; it differs in shape of eyes, very long and flattened. The male and female genitalia also exhibit marked differences and reference to these is necessary for precise deter- mination. Distribution: China (Sichuan). Agathidium (Agathidium) procerum Ang. & Dmz. Agathidium (Agathidium) procerum Angelini & De Marzo, 1998: 359. Material: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VIII.1996, leg. S. Kur- batov, 5 ex. in MHNG and AC-GNHM. Distribution: China (Sichuan). NEW SPECIES OF AGATHIDIUM 917 aac «Su: om Aedeagus (lateral and dorsal view of its apex) and spermatheca of: 7-9, Agathidium modestum sp. n.; 10-12, A. armatum sp. n.; 13-15, A. rubiginosum sp. n. Agathidium (Agathidium) rubiginosum sp. n. Figs 3, 13-15, 18 Holotype d: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VIII.1996, leg. S.Kurbatov, in MHNG. Paratypes: Same data as holotype, 1 d in MHNG, 1 d and 1 © in AC-GNHM. Length 3,05-3,15 mm (holotype d 3,15 mm). Dorsum reddish-brown; meso- sternum testaceous, metasternum reddish-brown; antennae uniformly testaceous; legs 918 FERNANDO ANGELINI reddish-brown. Microreticulation present only on the elytra, superficial, more im- pressed on female; puncturation fine and sparse on head and pronotum, absent on elytra. Sutural striae absent. Head: Microreticulation absent; punctures small, superficial, separated from each other by 2-6 times their own diameter. Widest at the eyes; eyes flattened (fig. 3); antero-lateral margins uniformly raised; clypeus slightly emarginate; clypeal line absent. 3rd antennal segment 2,4 times as long as 2nd and longer than 4th + 5th together. Pronotum: Microreticulation absent; punctures smaller and more superficial than those on the head, separated from each other by 1-15 times their own diameter. 1,59 times as broad as head, moderately broader than long (width/length= 1,42) and mode- rately convex (width/height= 1,56); anterior margin sharply curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 1,05 mm, width 1,50 mm and height 0,96 mm. Elytra: Microreticulation superficial and uniform; puncturation absent. Slightly narrower than the pronotum, as broad as long and weakly convex (width/height= 1,86); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,40 mm, width 1,40 mm and height 0,75 mm. Metathoracic wings absent. Meso- and metasternum: median carina weak, lateral lines absent, femoral lines incomplete, rounded at middle. Legs: Male hind femora with a sharp tooth at the posterior margin (fig.18). Tarsal formula: 4 5-5-4, 2 5-4-4. Aedeagus as in figures 13, 14. Spermatheca as in figure 15. Discussion: See Agathidium armatum sp. n.; from A. rufescens Ang. & Dmz. (from Sichuan) it differs in greater width ratio pronotum/head and in shape of pro- notum, less transverse. Distribution: China (Sichuan). Agathidium (Agathidium) brunneipenne Ang. & Dmz. Agathidium (Agathidium) brunneipenne Angelini & De Marzo, 1998: 363. Material: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VII.1996, leg. S. Kur- batov, 1 2 in MHNG. Distribution: China (Hubei, Sichuan). New record for Sichuan. Agathidium (Agathidium) cf. lugubre Ang. & Dmz. Agathidium (Agathidium) lugubre Angelini & De Marzo, 1998: 365. Material: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VIII.1996, leg. S. Kur- batov, 3 © in MHNG (tentative determination based on female specimens). Distribution: China (Guangxi, Sichuan). New record for Sichuan. Agathidium (Agathidium) hani sp. n. Figs 4, 19, 25-27 Holotype à : China, Sichuan, M. Emei, VII.1980, leg. Jäch in AC-GNHM. Paratypes: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VII.1996, leg. S. Kur- batov,1 2 in MHNG, 1 © in AC-GNHM. NEW SPECIES OF AGATHIDIUM 919 18 ~ | 19 DA 20 ta 24 Fics 16-24 Male hind femora of: 16, Agathidium modestum sp. n.; 17, A. armatum sp. n.; 18, A. rubiginosum sp. n.; 19, A. hani sp. n.; 20, A. luojiense sp. n.; 21, A. pseudouniforme sp. n.; 22, A. xilingense sp. n.; 23, A. kurbatovianum sp. n.; 24, A. megacephalum sp. n. 920 FERNANDO ANGELINI Length 3,8-4,4 mm (holotype d 4,40 mm). Dorsum dark reddish-brown, venter reddish-brown, mesosternum lighter; antennae uniformly testaceous; legs reddish- brown. Microreticulation almost absent, present only in traces on the elytra; punc- turation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures very small, superficial, separated from each other by 4-10 times their own diameter. Widest at the eyes; eyes flattened (fig. 4); antero-lateral margins uniformly raised; clypeus moderately emarginate; clypeal line absent. 3rd antennal segment 2 times as long as 2nd and longer than 4th + 5th together. Pronotum: Punctures smaller and more superficial than those on the head, hardly visible, separated from each other by 4-15 times their own diameter. 1,62 times as broad as head, weakly broader than long (width/length= 1,34) and very convex (width/height= 1,43); anterior margin sharply curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 1,55 mm, width 2,08 mm and height 1,45 mm. Elytra: Microreticulation present only in traces; punctures as on head but separated from each other by 1-5 times their own diameter; slightly narrower than the pronotum, a little longer than broad (width/length= 0,97) and weakly convex (width/ height= 1,9); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,95 mm, width 1,90 mm and height 1,00 mm. Metathoracic wings absent. Meso- and metasternum: median carina weak, lateral lines absent, femoral lines incomplete, rounded at middle. Legs: Male hind femora with two tooth at the posterior margin (fig.19). Tarsal formula: & 5-5-4, 2 5-4-4. Aedeagus as in figures 25, 26. Spermatheca as in figure 27. Discussion: Species of the dentatum group; Agathidium hani sp. n. is similar to A. lugubre Ang. & Dmz. (from Guangxi); it differs in shape of male hind femora, aedeagus and in greater width ratio pronotum/head. Distribution: China (Sichuan). Agathidium (Agathidium) luojiense sp. n. Figs 5, 20, 28-30 Holotype d: China, Sichuan, Xichang, M. Luoji, 2300-2500 m, 16.-24.VII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 5 d and 8 2 in MHNG, 2 6 and 2 © in AC-GNHM. Length 3,25-3,50 mm (holotype d 3,45 mm). Dorsum reddish-brown; meso- sternum reddish-brown, metasternum black; antennae uniformly testaceous; legs reddish-brown. Microreticulation almost absent, present only in traces on the elytra; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures very small, superficial, separated from each other by 1-10 times their own diameter. Widest at the eyes; eyes flattened (fig. 5); antero-lateral margins uniformly raised; clypeus moderately emarginate; clypeal line absent. 3rd antennal segment 1,3 times as long as 2nd and as long as the 4th + 5th together. Pronotum: Puncturation as that of head. 1,42 times as broad as head, moderately broader than long (width/length= 1,57) and very convex (width/height= 1,4); anterior NEW SPECIES OF AGATHIDIUM 921 e 28 29 VE | 31 32 Fics 25-33 Aedeagus (lateral and dorsal view of its apex) and spermatheca of: 25-27, Agathidium hani sp. n.; 28-30, A. luojiense sp. n.; 31-33, A. pseudouniforme sp. n. 922 FERNANDO ANGELINI margin sharply curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 1,03 mm, width 1,62 mm and height 1,15 mm. Elytra: Microreticulation almost absent, only traces; puncturation as that of head. As broad as pronotum, as broad as long and moderately convex (width/height= 1,8); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,62 mm, width 1,62 mm and height 0,90 mm. Metathoracic wings absent. Meso- and metasternum: median carina absent, lateral lines absent, femoral lines short, rounded at middle; metasternum short, meso- and metacoxae almost touching. Legs: Male hind femora curved at posterior margin (fig.20). Tarsal formula: à 5-5-4, 9 5-4-4. Aedeagus as in figures 28, 29. Spermatheca as in figure 30. Discussion: Species of the dentatum group; Agathidium luojiense sp. n. 1s similar to A. huaense Ang. & Dmz. and A. pseudouniforme sp. n.; it differs only in lesser length ratio 3rd/2nd antennal segments, in shape of male hind femora and aedeagus. Distribution: China (Sichuan). Agathidium (Agathidium) pseudouniforme sp. n. Figs 6; 21-31-33 Holotype d: China, Sichuan, Xichang, M. Luoji, 2300-2500 m, 16.-24.VII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 4 d and 9 © in MHNG, 3 d and 2 2 in AC-GNHM; Sichuan, env. Xichang, 1600 m, 28.VII.1996, leg. S. Kurbatov, 4 © in MHNG. Length 2,5-3,2 mm (holotype 4 2,90 mm). Dorsum reddish-brown; venter reddish-brown, mesosternum lighter; antennae uniformly testaceous; legs reddish- brown. Microreticulation present only in traces at sides of pronotum and on the elytra; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures very small, superficial, separated from each other by 1-10 times their own diameter. Widest at the eyes; eyes flattened (fig. 6); antero-lateral margins uniformly raised; clypeus moderately emarginate; clypeal line absent. 3rd antennal segment 1,7 times as long as 2nd and longer than 4th + 5th together. Pronotum: Microreticulation present only in traces at sides of pronotum; puncturation as that of head. 1,37 times as broad as head, moderately broader than long (width/length= 1,45) and very convex (width/height= 1,42); anterior margin weakly curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 0,93 mm, width 1,35 mm and height 0,95 mm. Elytra: Microreticulation present only in traces; punctures as those on head, separated from each other by 10-15 times their own diameter; slightly narrower than the pronotum, little longer than broad (width/length= 0,94) and weakly convex (width/ height= 1,73); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,32 mm, width 1,25 mm and height 0,72 mm. Metathoracic wings absent. Meso- and metasternum: median carina sharp, lateral lines absent, femoral lines complete, rounded at middle. NEW SPECIES OF AGATHIDIUM 923 34 35 36 Fics 34-36 Head of: 34, Agathidium xilingense sp. n.; 35, A. kurbatovianum sp. n.; 36, A. megacephalum sp. n. Legs: Male hind femora with a sharp tooth at the posterior margin (fig.21). Tarsal formula: & 5-5-4, £ 5-4-4. Aedeagus as in figures 31, 32. Spermatheca as in figure 33. Discussion: See Agathidium luojiense sp. n., from A. huaense Ang. & Dmz (from Shaanxi) it differs in shape of tooth of male hind femora, in shape of aedeagus, in the greater length ratio 3rd/2nd antennal segments and width ratio pronotum/head. Distribution: China (Sichuan). Agathidium (Microceble) xilingense sp. n. Figs 22, 34, 37, 38 Holotype d: China, Sichuan, Mt. Xiling, 1600-2400 m, 30.VII.-4.VII.1996, leg. S. Kur- batov, in MHNG. Paratypes: Same data as holotype, | d in MHNG, 1 d in AC-GNHM. Length 2,2-2,3 mm (holotype d 2,3 mm). Dorsum reddish-brown; mesosternum testaceous, metasternum reddish-brown; antennae testaceous but with segments 9-10 darker; legs reddish-brown. Microreticulation almost absent, present only in traces on the pronotum and elytra; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Microreticulation absent; punctures small, impressed, separated from each other by 5-8 times their own diameter. Eyes hemispherical, head widest at the eyes (fig 34); antero-lateral margins distinctly raised; clypeus weakly emarginate, with a weakly impressed short groove and small pit either side. 3rd antennal segment 1,6 times longer than 2nd and as long as the 4th + Sth together. Pronotum: Microreticulation almost absent, present only in traces; punctures larger than those of the head but more clearly impressed, separated from each other by 5-10 times their own diameter. 1,75 times as broad as head, moderately broader than long (width/length= 1,55) and moderately convex (width/height= 1,64); anterior margin 924 FERNANDO ANGELINI weakly curved; lateral outline broadly rounded. Measurements of pronotum of holo- type: length 0,72 mm, width 1,12 mm and height 0,68 mm. Elytra: Microreticulation almost absent, present only in traces; punctures similar to that of the pronotum but more superficial, separated from each other by 10-15 times their own diameter. Slightly narrower than the pronotum, as broad as long and moderately convex (width/height= 1,77); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,10 mm, width 1,10 mm and height 0,62 mm. Metathoracic wings present. Meso- and metasternum: median carina sharp, lateral lines absent, femoral lines incomplete, rounded at middle. Legs: Male hind femora rounded at posterior margin (fig.22). Tarsal formula: d 5-5-4, ? unknown. Aedeagus as in figures 37, 38. Discussion: Species of the andrewesi group; Agathidium xilingense sp. n. is similar to A. venustum Ang. & Dmz (from Taiwan, Guangxi, Hong Kong, Shaanxi) and A. taiwanense Ang. & Dmz. (from Taiwan); it differs in greater length ratio 3rd/2nd antennal segment, the less body length and in less width ratio pronotum/head and aedeagus. Distribution: China (Sichuan). Agathidium (Macroceble) kurbatovianum sp. n. Figs 23, 35, 39-41 Holotype d: China, Sichuan, Xichang, M. Luoji, 2300-2500 m, 16.-24.VII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 3 2 in MHNG, 1 d and I © in AC-GNHM. Length 2,60-2,75 mm (holotype d 2,65 mm). Dorsum dark reddish-brown, venter lighter; antennae testaceous but antennal club darker; legs reddish-brown. Microreticulation absent; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures very small, superficial, separated from each other by 2-8 times their own diameter. Widest at the eyes; eyes slightly flattened (fig. 35); antero-lateral margins uniformly raised; clypeus slightly emarginate; clypeal line absent; left mandible with a tooth. 3rd antennal segment 1,09 times as long as 2nd and shorter than 4th + 5th together. Pronotum: Microreticulation absent; punctures smaller and more superficial than those on the head, hardly visible, separated from each other by 2-15 times their own diameter. 1,46 times as broad as head, weakly broader than long (width/length= 1,14) and very convex (width/height= 1,34); anterior margin moderately curved; lateral outline broadly rounded. Measurements of pronotum of holotype: length 1,00 mm, width 1,14 mm and height 0,85 mm. Elytra: Puncturation as that of pronotum. As broad as pronotum, a little broader than long (width/length= 1,07) and weakly convex (width/height= 1,9); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,10 mm, width 1,18 mm and height 0,62 mm. NEW SPECIES OF AGATHIDIUM 925 TG ch Fics 37-44 Aedeagus (lateral and dorsal view of its apex) of: 37-38, Agathidium xilingense sp. n.; 39-40, A. kurbatovianum sp. n.; 42-43, A. megacephalum sp. n. — Spermatheca of: 41, Agathidium kurbatovianum sp. n.; 44, A. megacephalum sp. n. Metathoracic wings absent. Meso- and metasternum: median carina weak, lateral lines absent, femoral lines absent, a small tubercle present between the meta- coxae; metasternum short, meso- and metacoxae almost touching. 926 FERNANDO ANGELINI Legs: Male hind femora curved at posterior margin (fig.23). Tarsal formula: & 5-5-4, 9 4-4-4. Aedeagus as in figures 39, 40. Spermatheca as in figure 41. Discussion: Agathidium kurbatovianum sp. n. is similar to A. sherpa Ang. & Dmz. (from Nepal) and A. schawalleri Ang. & Dmz. (from Nepal); it differs in lesser length ratio 3rd/2nd antennal segments; from A. schawalleri it differs also in shorter body length and in shape of the pronotum which is less transverse. From A. oblitum Ang. & Dmz. (from Taiwan) and A. narusawae Hoshina (from Ryukyus Is.) it differs in greater body length, greater width ratio pronotum/head and length ratio 3rd/2nd antennal segments. From A. megacephalum sp. n. (Sichuan) it differs in greater length ratio 3rd/2nd antennal segments and in shape of pronotum, less transverse and more convex. Distribution: China (Sichuan). Agathidium (Macroceble) megacephalum sp. n. Figs 24, 36, 42-44 Holotype d : China, Sichuan, Xichang, M. Luoji, 2300-2500 m, 16.-24.VII.1996, leg. S. Kurbatov, in MHNG. Paratypes: Same data as holotype, 2 d and 1 © in MHNG, 1 d and 1 © in AC-GNHM. Length 2,7-3,0 mm (holotype d 2,90 mm). Dorsum dark reddish-brown, venter lighter; antennae testaceous with club darker; legs reddish-brown. Microreticulation absent; puncturation fine and sparse on whole dorsum. Sutural striae absent. Head: Punctures microscopic, hardly visible, separated from each other by 2-6 times their own diameter. Widest at the eyes; eyes slightly flattened (fig. 36); antero- lateral margins uniformly raised; clypeus slightly emarginate; clypeal line absent; left mandible of male with a tooth. 3rd antennal segment 0,88 times as long as 2nd and shorter than 4th + 5th together. Pronotum: Punctures similar or smaller and more superficial than those of the head, separated from each other by 5-10 times their own diameter. 1,4 times as broad as head, slightly broader than long (width/length= 1,29) and moderately convex (width/ height= 1,65); anterior margin weakly curved; lateral outline broadly rounded. Measu- rements of pronotum of holotype: length 1,05 mm, width 1,36 mm and height 0,82 mm. Elytra: Punctures as those of head, separated from each other by 1-10 times their own diameter. Slightly narrower than the pronotum, as broad as long and moderately convex (width/height= 1,84); lateral outline with humeral angle very broadly rounded. Measurements of elytra of holotype: length 1,20 mm, width 1,20 mm and height 0,65 mm. Metathoracic wings absent. Meso- and metasternum: median carina absent, lateral lines absent, femoral lines absent, a small tubercle present between the meta- coxae; metasternum short, meso- and metacoxae almost touching. Legs: Male hind femora curved at posterior margin (fig.24). Tarsal formula: 3 5-5-4, 2 4-4-4. Aedeagus as in figures 42, 43. Spermatheca as in figure 44. Discussion: See Agathidium kurbatovianum sp. n. Distribution: China (Sichuan). NEW SPECIES OF AGATHIDIUM 927 REFERENCES ANGELINI, F. & DE Marzo, L. 1981. Reports of Agathidium from Himalaya: expeditions of Basel Natural History Museum and Prof. H. Franz (Coleoptera, Leiodidae). Entomologica Basiliensia 6: 187-294. ANGELINI, F. & DE MARZO, L. 1998. Agathidiini from China, with description of 14 new species. Revue suisse de Zoologie 105 (2): 351-373. ANGELINI, F. & Svec, Z. 1994. Review of Chinese species of the subfamily Leiodinae. Acta Societas Zoologica Bohemoslava 58: 1-31. an , ARM TENERA i la } Le TRI ais ci 1 u 3 Î 7 N REIT Te cn a walk hinten nant u ia ah teas lt ie REVUE SUISSE DE ZOOLOGIE 106 (4): 929-938; décembre 1999 Considérations taxonomiques sur le genre Hadogenes Kraepelin, 1894: création des Hadogeninae subfam. n. et description d’une espèce nouvelle de I’ Angola (Scorpiones, Scorpionidae, Hadogeninae) Wilson R. LOURENCO Laboratoire de Zoologie (Arthropodes), Muséum National d'Histoire Naturelle, 61, rue de Buffon, F-75005 Paris, France; e-mail: arachne @ mnhn.fr Taxonomic considerations on the genus Hadogenes Kraepelin, 1894: proposition of the Hadogeninae subfam. n., and description of a new species from Angola (Scorpiones, Scorpionidae, Hadogeninae). - The study of a new species of Hadogenes Kraepelin, 1894, leads to the reana- lysis of the taxonomic position of this genus, with the proposition of a new subfamily Hadogeninae subfam. n. The new subfamily is placed in the family Scorpionidae, with therefore the transfer of the genus Hadogenes from the Ischnuridae to the Scorpionidae. The new species Hadogenes angolensis sp. n. is described from Angola. Key-words: Scorpionidae - Hadogeninae - Hadogenes - taxonomy - Angola. INTRODUCTION Le genre Hadogenes a été établi par KRAEPELIN (1894) ayant comme espece- type Scorpio (Ischnurus) trichiurus Gervais, 1843 [= Hadogenes trichiurus (Gervais 1843)]. KRAEPELIN place ce genre dans la famille des Scorpionidae et dans la sous- famille des «Ischnurini». Cette classification a été maintenue par KRAEPELIN (1899) dans sa monographie «Das Tierreich» et dans son analyse phylogénétique et biogéo- graphique des scorpions publiée en 1905. BIRULA (1917) propose une «tribu» Ischnuraria avec les sous-familles Ischnurinae, Hormurinae, Hemiscorpiinae et Hetero- scorpioninae, avec le genre Hadogenes toujours placé dans la sous-famille des Ischnu- rinae. Tous les auteurs subséquents (e.g. HEWITT 1918; LAWRENCE 1955; LAMORAL 1979; LAMORAL & REYNDERS 1975) suivent cette classification. Lors de ma révision des genres appartenant aux Ischnurinae, désormais élevée au rang de famille des Ischnuridae (LOURENÇO 1985, 1989), un certain nombre de décisions n'ont pu étre prises de manière satisfaisante. Ainsi, les statuts des genres Chiromachetes Pocock, 1899 et Hormiops Fage, 1933, considérés comme douteux, position d'ailleurs suivie par SISSOM (1990), ont été révisés et revalidés a l'appui de nouvelles données (LOURENÇO 1997; LOURENÇO & MONOD 1999). Le genre Hadogenes a recu peu d’attention, et est resté placé chez les Ischnuridae. Il est suggeré seulement Manuscrit accepté le 06.04.1999 930 WILSON R. LOURENCO une certaine affinité entre Hadogenes et Heteroscorpion Birula, 1903, endémique de Madagascar, pour lequel déja KRAEPELIN (1905) avait proposé une sous-famille Heteroscorpioninae. Des études approfondies sur la faune des scorpions de Madagascar (LOURENÇO 1996) ont permis une réanalyse précise de la position du genre Hetero- scorpion, avec élévation de cette sous-famille au rang de famille (Heteroscorpionidae). TAXONOMIE REANALYSE DU STATUT TAXONOMIQUE DU GENRE HADOGENES AVEC PROPOSITION D’ UNE NOUVELLE SOUS-FAMILLE Dans son travail sur les scorpions de la Namibie, LAMORAL (1979) dresse une liste des caractères différentiels entre le genre Hadogenes et les autres genres appar- tenant aux Ischnuridae, en particulier ceux tirés de la structure des hémispermatophores et du modele trichobothriotaxique, caractères globalement négligés par les auteurs précédents déjà cités. Ces caractères sont réanalysés lors de l’étude réalisée par LOURENÇO (1985, 1989), sans pour autant aboutir a une décision sur la position systé- matique des Hadogenes. A present, un ensemble de caracteres m’amene a proposer une nouvelle sous- famille monotypique avec genre-type Hadogenes. Cette sous-famille est placée, par prudence, au sein de la famille des Scorpionidae. Hadogeninae subfam. n. Diagnose: Scorpion de grande taille (10 à 20 cm). Prosoma et mesosoma très aplatis dorso-ventralement. Hémispermatophore armé d’un crochet a double apex (Figs 16 a 18); les genres d’Ischnuridae et de Scorpionidae présentent, sans exception, des hémispermatophores armés d’un crochet avec un seul apex. Yeux médians situés au centre de la carapace, plus proches de l’arrière que de l’avant; enfouis avec un tubercule oculaire obsolete. Trichobothriotaxie du type C, néobothriotaxique majorante, plétho- taxique (Vachon, 1973) (Figs 11 à 15). La pléthotaxie est absente chez les genres d’Ischnuridae, mais présente chez certains genres de Scorpionidae tels Pandinus Thorell. Glandes a venin des types lisse, pré-lobé ou semi-lobé (Figs 23 et 24), alors que les genres d’Ischnuridae présentent généralement la structure lisse et ceux des Scorpionidae la structure lobée. DESCRIPTION D'UNE NOUVELLE ESPECE LAMORAL (1979) signale que le genre Hadogenes doit faire l’objet d’un travail de révision. Il se contente cependant de caractériser les trois espèces jugées valıdes présentes en Namibie. Déjà HEWITT (1918), dans sa monographie sur les scorpions de l’Afrique du Sud, fait une remarque semblable à propos du genre, et soulève les difficultés de classification des espèces, en grande partie en raison d’une certaine «variabilite» des caracteres étudiés. Cette situation amène la description de variétés pas toujours satisfaisantes. I] est vrai que les auteurs anciens ont utilisé dans leur études des LE GENRE HADOGENES arme commen | | (lh Fics 1 à 10 Hadogenes angolensis sp. n. 1. Chélicère (mâle). 2. Tarse, vue latérale (mâle). 3. Telson (mâle). 4. Tranchant du doigt mobile (mâle). 5. Hanche, sternum, opercule génital et peigne (mâle). 6 Sternum, opercule génital et base des peignes (femelle). 7 et 8. Région proximale des peignes chez le mâle et la femelle. 9 et 10. Yeux latéraux chez le mâle et la femelle. 93] 932 WILSON R. LOURENCO caracteres tirés exclusivement de la morphologie externe, en négligeant la structure des hémispermatophores qui joue actuellement un rôle majeur dans la systématique du genre Hadogenes, ainsi que dans celle des genres d’Ischnuridae et de Scorpionidae. Le genre Hadogenes est réparti sur une grande partie du sud de |’ Afrique, plus précisément, en Angola, Botswana, Congo (?), Mozambique, Namibie, Afrique du Sud et Zimbabwe. Sa présence en Angola est limitée a une seule citation de LAWRENCE (1959), qui signale Hadogenes taeniurus (Thorell) dans ce pays (Vila Arriaga pres de Lungo: aujourd’hui Lubango, et Lucira). NEWLANDS (1972), LAMORAL (1979) et LAMO- RAL & REYNDERS (1975) mentionnent a nouveau Hadogenes taeniurus pour la partie sud du pays, sans pour autant préciser avoir examiné le matériel en question. La faune des scorpions d’Angola a été jusqu’a present tres peu connue. Les publications se limitent a celles de MONARD (1929, 1937), suivies de LAWRENCE (1949, 1959, 1961) et VACHON (1950). Seul LAWRENCE (1959) mentionne le genre Hadogenes, mais il m’a été impossible de localiser le matériel en question. L’étude de trois spécimens collectés dans la région du Plateau de l’Huila, pres de Capangombe, permet la description d’une nouvelle espece, qui présente des simi- litudes avec H. taeniurus. Ceci souleve la question d’une possible erreur d’identifi- cation de la part de LAWRENCE (1959), non vérifiée par LAMORAL (1979). Hadogenes angolensis sp. n. Figs) 17271701982 093 Matériel-type. Holotype mâle, Angola, Région de l’Huila, «Fazenda Bumbo» près de Capangombe (15°10’S - 13°09’E), 21-26/VI/1954 (W. Kisker & H. Barmann leg.). Une femelle allotype et un mâle (juvenile) paratype, mêmes données que pour l’holotype. Muséum d'histoire naturelle, Geneve. Etymologie. Le nom spécifique fait référence au pays ou la nouvelle espece a été collectée. Diagnose. Hadogenes angolensis sp. n. peut étre distingué des autres especes du genre, et en particulier d’H. taeniurus (Thorell), par des différences dans la structure du VIleme sternite (Figs 19 a 22) et dans celle des hémispermatophores, en particulier celle de la lame distale (Figs 16 a 18), et par un nombre plus élevé de dents aux peignes, 22/23 et 15/16 chez les males et femelles de H. angolensis contre 17/19 et 13/15 chez H. taeniurus. Description fondée sur l’holotype mâle. Mensurations (en mm) dans le tableau I. Coloration. La couleur de base est jaune-rougeatre. Prosoma: plaque proso- mienne de couleur rouge-jaunatre avec des zones latéro-médianes plus foncées, rou- geatres. Tubercule oculaire et zones des yeux latéraux plus foncées, allant vers le noiratre. Mesosoma de méme couleur que la plaque prosomienne, mais plus unifor- mément jaunatre, avec des zones rougeatres peu marquées. Metasoma avec les cing anneaux de couleur jaune-rougeatre, avec quelques taches brunatres, tres estompées. Vésicule jaunätre avec quelques zones brunätres estompées; l'aiguillon rougeätre. Ster- Fics 11 à 15 Hadogenes angolensis sp. n. Trichobothriotaxie du type C, néobothriotaxie majorante pletho- taxique (holotype male). 11 à 13. Pince, vues dorsale, externe et ventralo-externe. 14 et 15. Tibia, vues dorsale et ventrale. m INES LE GENRE HADOGE aoe goco sco rss TF e Coe 934 WILSON R. LOURENCO nites jaunatre foncé; le VIlème légèrement plus foncé postérieurement. Peignes et opercule génital jaunätre foncé; sternum, hanches et processus maxillaires presque entièrement rougeatres. Pattes jaunatres avec des zones rougeatres sur les articulations et sur les carenes internes. Pédipalpes jaune-rougeätre; carenes internes et doigts noir- cis. Chéliceres jaune-rougeatre avec absence de toute trame plus foncée; doigts rougeatres. Morphologie. Front de la plaque prosomienne legerement échancré. Tubercule oculaire presque au centre de la plaque prosomienne, mais légèrement en arrière de 0,6 mm; yeux médians séparés par plus d'un diametre oculaire. Trois yeux latéraux (Fig. 9). Sillon interoculaire peu profond, s'allongeant jusqu'à l'arrière des yeux médians où il se divise en deux, devenant très estompé, et entoure une fossette triangulaire. Toute la plaque prosomienne est couverte de granulations très fines. Les sillons et la fossette triangulaire sont dépourvus de granulations. Mesosoma: tergites couverts d'une très fine granulation, moins marquée dans leur région postérieure. Carene axiale pratiquement absente. Metasoma à anneaux très aplatis latéralement, en particulier les anneaux II à V; toutes les carènes sont très peu marquées à l’exception des dorsales sur les anneaux II à V. Anneaux II à V à face ventrale pourvue de granules spiniformes, bien plus marqués sur le Veme. Telson en forme de poire avec la vésicule dépourvue de carenes; aiguillon proportionnellement tres court; présence d’une importante chetotaxie sur la vésicule (Fig. 3). Peignes avec 23-22 dents (Fig. 5). Sternites lisses, a grands stigmates linéaires. Opercule génital fait de deux plaques semi-ovales (Fig. 5). Pédipalpes: fémur a 5 carènes pourvues de granules très gros; tégument intercarénal avec des granules fins. Tibia et pince avec quatre carenes completes formées par des gros granules. Présence sur la face interne du tibia d'un éperon tres développé dans la région basale, avec deux granules majeurs a l'extrémité (Figs 14 et 15). Pince grande et aplatie; main bien plus longue que les doigts (Figs 11 a 13); face dorsale pourvue de granulations plutöt fines; faces latérales avec des granulations moyennes; face ventrale brillante et presque réticulée. Doigts lisses; granulations du tranchant des doigts se disposant en deux lignes longitudinales (Fig. 4); granules accessoires absents; présence d’un mamelon sur le doigt mobile. Pattes: dernier article tarsal avec six épines plus importantes disposées en trois séries. Chéliceres avec la dentition caractéristique de la famille des Scorpionidae (Fig. 1; Vachon, 1963). Trichobothriotaxie du type C, néobothriotaxique majorante, plethotaxique (Figs 11 à 15 ; VACHON 1973). Hémispermatophores: les figures 16 et 17 montrent l'hémispermatophore en vue d'ensemble, faces interne et externe, avec des détails de la région médiane, ou les différentes parties sont illustrées, en particulier le crochet a double apex. Glande a venin du type simple (Fig. 23). Fics 16 à 24 16 a 18. Hémispermatophores. 16 et 17. Hadogenes angolensis sp. n., vue externe et interne. 18. Hadogenes taeniurus, vue interne (redessiné d’après LAMORAL 1979). — 19 à 22. Structure du VIlème sternite. 19 et 20. Hadogenes angolensis sp. n., male et femelle. 21 et 22. Hadogenes taeniurus, male et femelle (redessiné d’apres Lamoral 1979). — 23 et 24. Structures des glandes a venin chez le genre Hadogenes. 23. H. angolensis sp. n. (simple). 24. H. gracilis (semi-lobée). 935 LE GENRE HADOGENES (RE ar sl 936 WILSON R. LOURENCO Femelle. Coloration semblable a celle du male, avec les tergites plus clairs et les anneaux du metasoma davantage rougeatres. Corps globalement plus trapue. Pédipalpes plus robustes, anneaux du metasoma nettement plus courts et moins aplatis latéralement (cf. Tableau I). Yeux latéraux disposés plus en courbe que ceux du mâle (Fig. 10). Opercule génital couvert par une plaque unique de forme pentagonale, fendue a l’extremite (Fig. 6). Peignes plus petits avec 16-15 dents (Fig. 6). Male juvenile (paratype), avec 22-22 dents aux peignes. TABLEAU I Male Femelle Prosoma - Longueur 16,2 17,4 - Largeur antérieure 9,4 10,5 - Largeur posterieure 16,9 18,4 Anneau caudal I - Longueur 10,0 dol - Largeur 31 3,6 Anneau caudal V - Longueur 15,4 12,5 - Largeur 22 2,3 - Hauteur 3,4 32 Vésicule - Largeur 2,8 2,8 - Hauteur 3,6 3% Pédipalpe - Fémur longueur 16,1 16,2 - Fémur largeur 5,8 6,5 - Tibia longueur 15,1 16,5 - Tibia largeur 349) 6,4 - Pince longueur 29:9 31,9 - Pince largeur 9,8 112; - Pince hauteur 4,4 5,6 Doigt mobile - Longueur 12,4 14,7 REMERCIEMENTS Je suis très reconnaissant à Philippe Bouchard et Maurice Gaillard, Muséum National d’Histoire Naturelle, Paris pour leur contribution à la réalisation des dessins illustrant le présent travail, aux Drs Jacqueline Kovoor et Sabine Jourdan (Paris) pour la lecture du manuscrit et au Dr A. de Barros Machado (Oeiras) pour m’avoir fourni des renseignements sur les spécimens décrits. LE GENRE HADOGENES 937 Lee 3x Hadogenes angolensis | @ Hadogenes taeniurus (?) | \ 7 AES | e + — | | 260kM X. NAMIBIE — LD _ LT SO zz ı BOTSWANA ~ Ÿ FRE & 9% NE — PAS: È aaa FIG. 25 Carte avec indication de la localité-type d’ Hadogenes angolensis sp. n. et les stations citées pour Hadogenes taeniurus par LAWRENCE (1959). BIBLIOGRAPHIE BIRULA, A. 1917. Arthogastric Arachnids of Caucasia. Part I. Scorpiones. Mémoires du Museum du Caucase, ser. A, 5 : 253 pp. (English translation : Israel Program for Scientific Translations, Jerusalem, 1964 : 170 pp.). Hewitt, J. 1918. A survey of the scorpion fauna of South Africa. Transactions of the Royal Society of South Africa 6 : 89-192. KRAEPELIN, K. 1894. Revision der Scorpione. II. Scorpionidae und Bothriuridae. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 11 : 1-248. KRAEPELIN, K. 1899. Scorpiones und Pedipalpi. /n: Das Tierreich, 8 : 1-265. Friedländer Verlag, Berlin. KRAEPELIN, K. 1905. Die Geographische Verbreitung der Scorpione. Zoologische Jahrbücher, Abteilung Systematik 22 : 321-364. LAMORAL, B. H. 1979. The scorpions of Namibia (Arachnida : Scorpionida). Annals of the Natal Museum 23 (3) : 497-784. LAMORAL, B. H. & REYNDERS, S. 1975. A catalogue of the scorpions described from the Ethio- pian Faunal region. Annals of the Natal Museum 22 (2) : 489-576. LAWRENCE, R. F. 1949. A collection of Opiliones and Scorpions from North-East Angola made by Dr. A. de Barros Machado in 1948. Publicaçôes Culturais da Companhia de Diamantes de Angola: 1-20. LAWRENCE, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with check-lists and keys to South African families, genera and species. South African Animal Life 1 : 152-262. 938 _ WILSON R. LOURENCO LAWRENCE, R. F. 1959. A collection of Arachnida and Myriapoda from the Transvaal Museum. Annals of the Transvaal Museum 23 : 363-386. LAWRENCE, R. F. 1961. New Scorpions and Solifugae from S.W.A. and Angola. Kungl. Fysio- grafiska Sällskapet i Lund Forhandlingar 31 (15) : 147-160. LOURENCO, W. R. 1985. Essai d'interprétation de la distribution du genre Opisthacanthus (Arach- nida, Scorpiones, Ischnuridae) dans les régions Néotropicale et Afrotropicale. Etude taxinomique, biogéographique, évolutive et écologique. Thèse de Doctorat d'Etat, Uni- versité Pierre et Marie Curie, 287 pp. LOURENCO, W. R. 1989. Retablissement de la famille des Ischnuridae, distincte des Scorpionidae Pocock, 1893 a partir de la sous-famille des Ischnurinae Pocock, 1893. Revue Arachno- logique 8 (10) : 159-177. LOURENCO, W. R. 1996. Scorpions. Faune de Madagascar, M.N.H.N., 87 : 1-102. LOURENCO, W. R. 1997. Considérations taxonomiques sur le genre Chiromachetes Pocock, 1899 (Chelicerata, Scorpiones, Ischnuridae). Zoosystema 19 (1) : 81-89. LOURENCO, W. R. & Monon, L. 1999. Confirmation de la validité du genre Hormiops Fage, 1933 avec redescription d’Hormiops davidovi Fage, 1933 (Scorpiones, Ischnuridae). Zoo- systema 21 (2): 337-344. MONARD, A. 1929. Matériaux de la mission scientifique suisse en Angola. Scorpions. Bulletin de la Société neuchäteloise de Sciences naturelles 54 : 37-43. MONARD, A. 1937. Scorpions, Solifuges et Opilions d’ Angola. Revue suisse de Zoologie 44 (13) : 251-258. NEWLANDS, G. 1972. A description of Hadogenes lawrencei sp. nov. (Scorpiones) with a checklist and key to the South West African species of the genus Hadogenes. Madoqua, ser. 2, 1 (54-62) : 133-140. SISSOM, W. D. 1990. Systematics, Biogeography, and Paleontology. Jn: POLIS, G.A. (ed.). The Biology of Scorpions. Stanford University Press, Stanford, pp. 64-160. VACHON, M. 1950. Remarques sur les Scorpions de l’Angola. Publicacöes Culturais da Companhia de Diamantes de Angola: 1-18. 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, 2e ser., 35 (2) : 161-166. VacHon, M. 1973. 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 Museum National d'Histoire Naturelle, Paris, 3e sér., n° 140, Zool. 104: 857-958. REVUE SUISSE DE ZOOLOGIE 106 (2): 939-943; décembre 1999 Studies on tiger beetles. XCVIII. A new Cylindera from Chubut, Argentina (Coleoptera: Cicindelidae) Fabio CASSOLA Via F. Tomassucci 12/20, I-00144 Roma, Italy. Studies on tiger beetles. XCVIH. A new Cylindera from Chubut, Argentina (Coleoptera: Cicindelidae). - A new tiger beetle species, Cylin- dera (Plectographa) chubuti sp. n., closely related to C. (P.) gormazi (Reed, 1871), is described and figured, based on four specimens collected at El Maidén, Chubut, Argentina. Key-words: Tiger beetles - Cicindelidae - Cylindera - new species - Argentina. INTRODUCTION The tiger beetle fauna of Argentina is probably still uncompletely known. A checklist had been presented long ago by BRUCH (1911), who listed 42 species in all, while additional information was later given in general catalogues of the family such as those by Horn (1905, 1908-15, 1926), BLACKWELDER (1944), and WIESNER (1992). Afterwards, a single new species was described by MANDL (1963), and VIDAL SARMIENTO (1965) mentioned the occurrence of a Ctenostoma species in the northern part of Argentina. Later, VIDAL SARMIENTO (1966) recognized the occurrence of 19 species belonging to the genus Cicindela s. |. in Argentina, while SUMLIN (1979) sub- sequently described four additional new species and raised up to 27 the Cicindela s. I. species known from the country. The same author (SUMLIN 1993) added later one more new species and, by adding also a previously overlooked species, he brought up to 29 the known number of the Argentine species of subtribe Cicindelina (sensu RIVALIER 1971). Finally, SUMLIN (1994) has recorded from Argentina one /resia species as well. Present-day knowledge has been summarized by CassoLa & PEARSON (2000), who recorded for Argentina a total of 65 species, 13 of which endemic of the country. Several years ago, among other tiger beetle material of the Muséum d’histoire naturelle of Geneva (Switzerland), which was kindly submitted for identification by Dr. Ivan Lobl, I found four specimens of a remarkable Cylindera species of the subgenus Plectographa Rivalier, 1954, which appeared to be new to science. This species (from Chubut, northern Patagonia) was obviously a near relative of C. (P.) gormazi (Reed, 1871), but it showed unusual elytral markings which made it to be easily distin- guishable from any other known related species. However, so far I have refrained Manuscript accepted 10.06.1999 940 FABIO CASSOLA myself from describing the new species because a form of C. (P.) gormazi (Reed, 1871) from Chile, having enlarged partially confluent elytral markings, had been described by VARAS ARANGUA (1921) as “/zquierdoi n. var.”, and it was not clear from his descrip- tion whether my specimens could well be the same form. Moreover, two additional names (Cicindela vidali Philippi & Juliet, 1871, and Cicindela gormazi var. dolorosa W. Horn, 1896, both described from Chile) are available in the entomological literature which appear to belong to C. (P.) gormazi (VARAS ARANGUA 1921, Horn 1926, WIESNER 1992). Therefore, to not take the risk of creating a useless junior synonym, I felt it better to wait for additional material and information. Luckily enough, thanks to the courtesy of Dr. Lothar Zerche of the Deutsches Entomologisches Institut (DEI) of Eberswalde, Germany, I was recently able to borrow two interesting female specimens (labelled respectively “Santiago/ de Chile/ Dr. Puelma” and “Chile/ ex cab/ Brown”) which appear to be izquierdoi. As to vidali, it 1s just a junior synonym of gormazi, as PHILIPPI himself did recognize (PHILIPPI 1887, VARAS ARANGUA 1921), while the original description of dolorosa alone makes it clear that this name applies to a gormazi population having unusually narrow elytral markings (HORN 1896, VARAS ARANGUA 1921). Thus, my last doubts about the status of the Chubut species can be finally set aside, and therefore I give below its description. Cylindera (Plectographa) chubuti sp. n. Fig. la-b Material examined: Male holotype and three male paratypes from ARGENTINA (Chubut), El Maitén, 8 February 1966, A. Kovacs leg.; holotype and one paratype in the collection of the Muséum d’histoire naturelle (Geneva, Switzerland), two paratypes in authors collection. The type-locality lies in northern Chubut, along the eastern base of the Andes Mountains, just near the border with Rio Negro. Diagnosis. A small black Cylindera species of the subgenus Plectographa, closely allied to C. (P.) gormazi (Reed, 1871) (Fig. Ic-d), however easily distinguished from it by the different elytral markings. Female unknown. Description. Head dull black with some slight metallic bluish-green to violet reflections on vertex and eyes; front, clypeus and cheeks rather strongly pilose, with many sparse, white, semi-erect hairs; some sparse, fine, erect long hairs on vertex too. Eyes dark yellowish, relatively small. Labrum testaceous (narrowly darkened on mar- gins), short, transverse, feebly unidentate in the middle, with 10-11 white short hairs or setigerous punctures near forward edge. Mandibles relatively short, externally testa- ceous from base to middle, shining black with greenish reflections internally and in apical half. Maxillary palpi metallic dark to green, the basal joint with long white hairs especially on the outer side; labial palpi testaceous with the last joint metallic green. Antennae relatively short and thick, reaching approximately the first third of the elytral length in male (expectedly shorter in female); articles 1-4 metallic dark green to violet, nearly glabrous, a single seta or setigerous puncture on tip of scape; joints 5-11 equally long, dull brownish-black, finely and evenly pubescent. Thorax: pronotum obviously wider than long, sub-rectangular shaped, dull black with metallic bluish-green reflections in transversal grooves, sparsely pubescent A NEW CYLINDERA FROM ARGENTINA 94] Fic. | Cylindera (Plectographa) chubuti sp. n., male paratype: a, habitus; b, labrum. C. (P.) gormazi (Reed, 1871), male specimen from “Llanquihue, Chile”: c, right elytron. C. (P.) gormazi, female specimen from “Santiago de Chile” (m. izquierdoi Varas Arangua, 1871): d, right elytron. Scale lines: 1 mm. at sides, a few hairs also sparsely on disc (expectedly often rubbed-off in old speci- mens). Episterna metallic greenish-black with some cupric reflections at base near the coxae, sparsely covered with long erect white hairs. Elytra relatively large, together much wider than head with eyes, slightly rounded at sides, dull brownish-black with some greenish reflections on shoulders; a few poorly apparent green punctures near the shoulders and in a irregular longitudinal row parallel to, but some distance from, the suture; lateral margins and base of elytra 942 FABIO CASSOLA shining metallic green to violet. Hind margins evenly rounded, non-microserrated, sutural angle right with a very small apical tooth. Elytral markings yellowish-white, conspicuous, constituted by a long complete humeral lunule, a large wide middle band (widely expanded along the lateral border and obliquely crossing the elytral disc until a short distance from the suture), and a short complete apical lunule, whose subapical dot is narrowly connected on disc with the hind margin of the middle band. Epipleura metallic greenish-bronze, more or less rufescent in apical half. Underside metallic dark bronze with green to bluish metallic reflections, strongly covered with white long hairs on sternal pieces and coxae, as well as in the outer third of the abdominal sternites; trochanters dark blackish-brown. Legs metallic green with some cupric to violet reflections; a few rows of white spiniform hairs on femora and tibiae, tarsal claws rufescent apically. Male aedeagus similar to that of C. (P.) gormazi (VIDAL SARMIENTO 1966), short, bulky, arc-shaped, with a short blunt protruding apex followed by a conspicuous lateral crest on both sides. Female unknown. Length: 9.3-9.5 mm (without labrum). Etymology. This small new Plectographa species is named from the geogra- phical region where it comes from, the province of Chubut in northern Argentinian Patagonia. Remarks. C. (P.) gormazi and C. (P.) chiliensis (Audouin & Brullé, 1839) - both described from Chile (REED 1871, AUDOUIN & BRULLE 1839, PENA & BARRIA 1973) - have also been recorded from Chubut (BRUCH 1911, SUMLIN 1979), but unfor- tunately no precise locality data are available. However, the apparent overlapping of ranges seems to strenghten the concept of C. (P.) chubuti sp. n. as specifically distinct. AKNOWLEDGMENTS I would wish to thank Dr. Ivan Löbl (Muséum d’histoire naturelle, Geneva, Switzerland) for having submitted for identification many tiger beetle materials. Best thanks are also to be given to Dr. Lothar Zerche (DEI, Eberswalde, Germany), who kindly allowed me to borrow and study several specimens from Walther Horn’s important collection. LITERATURE CITED AUDOUIN, J. V. & BRULLE, A. 1839. Descriptions des espéces nouvelles ou peu connues de la famille des Cicindélètes, faisant partie de la collection du Muséum. Archives du Muséum d Histoire naturelle, Paris 1: 115-142, 3 pls. BLACKWELDER, R. E. 1944. Checklist of the Coleopterous Insects of Mexico, Central America, the West Indies, and South America. Part I. Bulletin of the United States national Museum, No. 185: 1-188. BRUCH, C. 1911. Catalogo sistemätico de los Coleöpteros de la Republica Argentina. Pars I. Famiglia Carabidae (Cicindelinae, Carabinae). Revista del Museo de la Plata 17 (2, 4): 143-180. A NEW CYLINDERA FROM ARGENTINA 943 CassoLa F. & PEARSON, D. L. 2000. Global patterns of tiger beetle species richness (Coleoptera: Cicindelidae): Their use in conservation planning. Biological Conservation 92 (in press). Horn, W. 1896. Die Cicindeliden der Dohrn’ schen Sammlung. Stettiner entomologische Zeitung 57: 164-177. Horn, W. 1905. Systematischer Index der Cicindeliden. Deutsches entomologische Zeitschrift, Beiheft: 1-56. Horn, W. 1908-15. Coleoptera, Fam. Carabidae, Subfam. Cicindelinae. Jn: WYTSMAN P. (ed.). Genera Insectorum, 82A/C: 1-487, pl. 1-23. Horn, W. 1926. Carabidae: Cicindelinae. /n: JUNK, W. (ed.). Coleopterorum Catalogus, Pars 86, Berlin, 345 pp. ManpL, K. 1963. Uber neue Cicindeliden-Aufsammlungen aus tropischen Ländern (Col.). Entomologischen Arbeiten aus dem Museum G. Frey, Tutzing 14 (2): 575-592, pl. HILL PENA, L. E. & BARRIA P., G. 1973. Revision de la Familia Cicindelidae (Coleoptera), en Chile. Revista chilena de Entomologia 7: 183-191. PHILIPPI, F. 1887. Catalogo de los Coleöpteros de Chile. Anales de la Universidad de Chile 71: 619-806. REED, E. C. 1871. Notes on some Chilean Cicindelae, with description of a new species. Entomologist’s Monthly Magazine 8: 76-77. RIVALIER, E. 1954. Démembrement du genre Cicindela Linné. II. Faune américaine. Revue francaise d’Entomologie 21: 249-268. RIVALIER, E. 1971. Remarques sur la tribu des Cicindelini (Col. Cicindelidae) et sa subdivision en sous-tribus. Nouvelle Revue d’Entomologie 1: 135-143. SUMLIN, W. D. 1979. A brief review of the genus Cicindela of Argentina (Coleoptera: Cicindelidae). Journal of the New York entomological Society 87 (2): 98-117. SUMLIN, W. D. 1993. Studies on the Neotropical Cicindelidae IV: Notes on Argentine Cicindela with the description of a new species (Coleoptera). Cicindela 25 (3/4): 29-39. SUMLIN, W. D. 1994. Studies on the Neotropical Cicindelidae V: A review of the genus /resia (Coleoptera). Cicindelidae: Bulletin of worldwide Research 3 (1): 1-32. VARAS ARANGUA, E. 1921. Contribuciön al estudio de los Cicindelidae. Los Cicindelidae de Chile. Revista chilena de Historia natural pura y applicada 25: 28-61. VIDAL SARMIENTO, J. 1965. Genero Ctenostoma Klug nuevo para la Argentina (Coleoptera, Cicindelidae). Revista de la Sociedad entomolögica argentina 27 (1-4): 29-32. VIDAL SARMIENTO, J. 1966. Las especies argentinas de los generos Cicindelidia Riv., Brasiella Riv. y Cylindera Westwood. Estudio de las estructuras genitales del macho. Revista del Museo de la Plata (Nueva Serie), Seccion Zoologia 9: 25-46. WIESNER, J. 1992. Verzeichnis der Sandlaufkäfer der Welt. Checklist of the Tiger Beetles of the World. Verlag Erna Bauer, Keltern, 364 pp. af BIA MEIN GBM TE Oe A 7 i ns CAM Aie ul | 8 . ni a > OVALI REVUE SUISSE DE ZOOLOGIE 106 (4): 945-982: décembre 1999 A taxonomic revision of the family Oncopodidae II. The genus Gnomulus Thorell (Opiliones, Laniatores) Peter J. SCHWENDINGER! & Jochen MARTENS2 l'Institut für Zoologie und Limnologie, Universität Innsbruck, Technikerstr. 25, A-6020 Innsbruck, Austria. Present address: Muséum d'histoire naturelle, Case postale 6434, CH-1211 Geneve 6, Switzerland. 2 Institut für Zoologie, Johannes Gutenberg-Universitat Mainz, Saarstr. 21, D-55099 Mainz, Germany. A taxonomic revision of the family Oncopodidae II. The genus Gno- mulus Thorell (Opiliones, Laniatores). - The 27 formally recognized species of Gnomulus Thorell, 1890 are listed and placed in six provisional species groups. Poorly known species are rediagnosed. Gnomulus segnipes (Loman) is placed in synonymy with G. armillatus (Thorell). Both nominal species occur only in western Sumatra; corresponding records from outside Sumatra are attributed to other species, some of them undescribed. For G. aborensis (Roewer), G. rostratus Thorell, G. insularis (Roewer), G. piliger (Pocock), G. pulvillatus (Pocock), G. sumatranus Thorell and G. annulipes (Pocock) penis morphology is illustrated for the first time. Additional illustrations are presented for G. sundaicus (Schwendinger), G. laevis (Roewer) and G. goodnighti (Suzuki). Intraspecific variation is discussed. The original designation of G. sumatranus as the type species of Gnomulus is maintained, despite being in conflict with the International Code of Zoological Nomenclature. Key-words: Opiliones - Oncopodidae - taxonomy - Asia. INTRODUCTION Following part I of our revision of the Oncopodidae (MARTENS & SCHWEN- DINGER 1998), where seven new and unusual species of Gnomulus were described, we now continue with a more comprehensive study of this species-rich genus. Consi- dering the large material available, including several additional new species, it appears more practical to split our treatment of Gnomulus into two parts. In this paper we present a revision of the already known species, provisionally assigned to species Manuscript accepted 04.08.1999 946 P. J. SCHWENDINGER & J. MARTENS groups and listed in geographical order roughly from northwest to southeast. Hereby we focus on species described before 1969, for many of which male genital characters can be shown. For a few species described more recently, new specimens are illustrated to provide additional information on intraspecific variation. In our next article descriptions of several new Gnomulus species shall follow. Abbreviations and terms used in the text: AMNH American Museum of Natural History; NHML Natural History Museum, London (formerly British Museum of Natural History); MAR collection of J. Martens, Mainz; MSNG Museo Civico di Storia Naturale, Genova; MHNG Muséum d'histoire naturelle, Genève; MHNP Muséum National d'Histoire naturelle, Paris; SMF Naturmuseum und Forschungs- institut Senckenberg, Frankfurt; ZMA Zoölogisch Museum, Universiteit van Amsterdam; ZMB Museum für Naturkunde der Humboldt-Universität, Berlin; ZMC Zoologisk Museum, Kgbenhavn; ZMH Zoologisches Institut und Museum, Univer- sıtät Hamburg; ZMT Zoological Museum, University of Turku. Body measurements refer to the distance between the anterior margin of the carapace and the posterior margin of the dorsal scutum. Leg articles were measured on their dorsal sıde, from joint to joint. All measurements are given in mm. Termino- logy of penis morphology follows that of MARTENS & SCHWENDINGER (1998: fig. 1 - showing dorsal and lateral but not ventral view as given in the corresponding legend). The dorsal connection between the prosomal and opisthosomal part of the dorsal scutum, formed by paired or unpaired tooth-like processes or lobes (e.g. Figs 28-30), is here called the "carapace-abdomen bridge". TAXONOMY Gnomulus Thorell, 1890 Gnomulus Thorell (1890: 378), type species by original designation, G. sumatranus Thorell, 1891!. - Pocock (1897: 285). - LoMAN (1902: 182). - ROEWER (1923: 60-61). - SCHWENDINGER (1992: 197). - MARTENS & SCHWENDINGER (1998: 526). Pelitnus Thorell (1891: 757), type species by original designation, P. armillatus Thorell, 1891. - Pocock (1897: 285). - LOMAN (1902: 182). - ROEWER (1923: 62). - SORENSEN (1932: 213). - SCHWENDINGER (1992: 197). - MARTENS & SCHWENDINGER (1998: 501, 526). Diagnosis: See MARTENS & SCHWENDINGER (1998: 526). | The original designation of G. sumatranus Thorell, 1891 as the type species of Gno- mulus Thorell, 1890 was never questioned by subsequent authors, although it does not accord with the International Code of Zoological Nomenclature (INTERNATIONAL TRUST FOR ZOOLO- GICAL NOMENCLATURE 1985). Gnomulus sumatranus was then not a nominal species (but a “nomen nudum”) and G. rostratus Thorell, 1890 was the only species available in this genus. Hence article 68c (monotypy) of the code would have to be applied, making G. rostratus the type species. However, there was no such code in 1890 for Thorell to obey and we think that he had good reasons to favour G. sumatranus above G. rostratus. Therefore the original desig- nation should be maintained. A corresponding proposal (case 3116) has been submitted to the International Commission on Zoological Nomenclature (SCHWENDINGER & MARTENS 1999). THE GENUS GNOMULUS 947 Species account and distribution: 27 nominal species, here placed in six provisional species groups, are known from the Himalayan region and from Southeast Asia (Fig. 1). süß A ni 5 nr o> nl EI Ian si Ze] Fic. | Distribution of Gnomulus in the Himalayan region and in Southeast Asia. - 1 Ulleri (G. hyatti), 2 Rotung (G. aborensis), 3 Doi Inthanon (G. lannaianus), 4 Langkawi Island (G. insularis), 5 Penang Island (G. insularis, G. rostratus), 6 Bukit Besar (G. piliger), 7 Maxwell Hill (G. laruticus), 8 Chenderiang (G. rostratus, G. asli), 9 Batu Caves (G. pulvillatus), 10 Templer Park, Ulu Gombak, Genting Highlands (G. hirsutus), 11 Dolok Banol - location unknown (G. drescoi), 12 Gunung Singgalang (G. sumatranus, G. armillatus), 13 Bukittinggi, Anai, Ajer Mancior, Kayutanam (G. armillatus), 14 Gunung Kerinci (G. armillatus), 15 Cibodas (G. thorelli), 16 Kuching, Matang (G. laevis), 17 Kuching, Bau, Serian (G. sundaicus), 18 Baram River, Gunung Mulu (G. annulipes), 19 Andulau, Sungai Lubang Barus (G. baharu), 20 Amo (G. imadatei), 21 Kabili-Sepilok, Kolapis (G. conigerus), 22 Mt. Matalingajan (G. palawanensis), 23 Sagada (G. maculatus), 24 Crystal Caves (G. coniceps), 25 Mt. Santo Thomas (G. crucifer), 26 Mt. Makiling (G. minor), 27 Puerto Galera (G. maculatus - doubtful record), 28 Visca (G. leyteensis), 29 Mt. Katanglad (G. goodnighti), 30 Mt. McKinley (G. goodnighti). 948 P. J. SCHWENDINGER & J. MARTENS THE ABORENSIS-GROUP Diagnosis: Medium-sized to large species with robust cheliceral hands in males, low interocular areas, medially divided dorsal scutal areas and short, blunt lateral sclerites on glans penis. This group comprises three species from Nepal, NE-India and N-Thailand. Gnomulus hyatti (Martens, 1977) Fig. 18 Pelitnus hyatti Martens (1977: 296-298, figs 1-8); description of d and ®. - SCHWENDINGER (1992: 178, 197). Gnomulus hyatti (Martens): MARTENS & SCHWENDINGER (1998: 526). Types: NEPAL, Ulleri, 1800-2200 m, 4 holotype (NHML, examined), 1 2 paratype (SMF 29229, examined), leg. K. H. Hyatt, 19.V.1954. Remark: No additional material of this species is available although several new and unusual oncopodid specimens have since been collected at other localities in Nepal. Relationships: Gnomulus hyatti is close to G. lannaianus and G. aborensis. Unlike the latter two species, G. hyatti has a fairly short (about 2.2 times longer than wide) distitarsus II (Fig. 18) and a less pronounced external sexual dimorphism. Distribution: Known only from the type locality, in the southern part of the Annapurna massif, central Nepal (Fig. 1). Gnomulus aborensis (Roewer, 1913) Figs 2-17 Pelitnus aborensis Roewer (1913: 207); description of species, without distinction of sexes. Gnomulus aborensis (Roewer): Roewer (1923: 62, figs 65 a, b). - SCHWENDINGER (1992: 197). - MARTENS & SCHWENDINGER (1998: 526). Types: INDIA, Abor District, Rotung, ca. 400 m, Upper Rotung, ca. 620 m, 2 syntypes (Museum Calcutta, not examined), leg. S. W. Kemp, 21./31.XIL1911. Other material: Abor District, 2 4, 1 9 (SMF 1257), not mentioned in the original description but recorded as "cotypes" by ROEWER (1923: 62), possibly part of the original series of specimens examined by Roewer. Diagnosis: Fairly large species, characterized by: arched carapace without distinct eye tubercle, with pronounced lateral tubercles below carapace-abdomen bridge (Figs 8-11; ROEWER 1923: fig. 65b); dorsal scutum high, areas I-VII elevated, medially divided by a pronounced longitudinal furrow and separated from each other by transversal furrows (Figs 8-11; ROEWER 1923: fig 65a); coxa I ventrally with distinct anterio-lateral process, coxae II and III with anterio-proximal one, coxa II with additional posterio-proximal one; chelicerae fairly robust, proximal article with a distad-inclined proventral process (Figs 12-15); palp with conical ventral process on trochanter, a low rounded ventral process and a distinct dorsal boss proximally on femur (Fig. 16); distitarsus of leg II about 2.7 times longer than wide (Fig. 17). External sexual dimorphism in shape and size of carapace and chelicerae, being distinctly larger in 6 d (Figs 8-9, 12-13); ventral scutal elevation of d only little swollen and covered with plenty of fine hairs (Fig. 8). Truncus penis fairly slender, ORIBATES D'UNE TOURBIERE JURASSIENNE 949 \\\ \ \\\ || \l Fics 2-19 Gnomulus aborensis (Roewer) (2-17), G. hyatti (Martens) (18), G. lannaianus (Schwendinger) (19). - Penis, dorsal (2) and lateral view (3); apex of penis of 2 à, dorsal (4, 6) and lateral view (5, 7). Body of 4 (8) and of ® (10), lateral view; anterior part of body and chelicerae of d (9) and of ® (11), dorsal view; left chelicera of 4 (12) and ® (14), retrolateral view; proximal article of right chelicera of d (13) and of 2 (15), prolateral view; trochanter and femur of left palp of d, retrolateral view (16); distal part of left (17, 19) and of right leg II (18), retrolateral view. - Scale lines 0.1 mm (4-7), 0.5 mm (2, 3, 12-19), 1.0 mm (8-11). 950 THE GENUS GNOMULUS with strongly arched distal margin. Glans penis with more or less pentagonal median plate not covering outer side of membraneous tubes; lateral sclerites stout, with prominent outer margin, strong wrinkles on lower face and blunt tip; stylus slender (Figs 2-6). Variation: Measurements (d, in brackets 2): body 5.88-6.78 (6.13) long, 4.44-4.88 (4.53) wide; carapace region 1.51-1.87 (1.25) long, 2.54-2.94 (2.48) wide; é m=2), 2 (19) Relationships: Gnomulus aborensis is closest to G. lannaianus. Both share sexual dimorphism in the size of carapace and chelicerae and possess a long (more than 2.5 times longer than wide) distitarsus II. Due to differences in penis morphology and scutal surface structure, Gnomulus hyatti appears more distant to these two species. Distribution: Known only from the foothills of the Himalaya in Arunachal Pradesh, northeastern India (Fig. 1). Gnomulus lannaianus (Schwendinger, 1992) Fig. 19 Pelitnus lannaianus Schwendinger (1992: 178-180, 197, figs 1-13); description of d and 9. Gnomulus lannaianus (Schwendinger): MARTENS & SCHWENDINGER (1998: 527). Types: THAILAND, Chiang Mai Province, Doi (= Mount) Inthanon, 2300-2530 m, & holotype, 1 4,7 2 paratypes (MHNG, examined), leg. P. Schwendinger, X.1986-X.1987. Remark: No new material available. Distitarsus II is about 2.8 times longer than wide (Fig. 19). Relationships: A close relationship between G. lannaianus and G. hyatti was already pointed out by SCHWENDINGER (1992: 197). Distribution: Known only from the summit area of Doi Inthanon (northern Thailand), where several other remarkable animal and plant species with affinities to the Himalayan region occur (Fig. 1). THE ROSTRATUS-GROUP Diagnosis: Medium-sized species with distinctly anteriad-inclined pointed eye tubercles and unique penis morphology, i.e. median plate of glans long and narrow, stylus distally very wide, shaped like a depressed bell. This group includes two species from the west of peninsular Malaysia and nearby islands. Gnomulus insularis (Roewer, 1927) Figs 20-37 Pelitnus insularis Roewer (1927: 268); description of d . - SCHWENDINGER (1992: 197). Gnomulus insularis (Roewer): MARTENS & SCHWENDINGER (1998: 526). Type: MALAYSIA, Pulu Pinang (= Pulau Pinang = Penang Island) - doubtful indication of locality, 4 holotype (SMF 301, examined). Other material: MALAYSIA, Kedah, Langkawi Island, 3 d, 1 2 (NHML), leg. P. D. Hillyard, II.1983. THE GENUS GNOMULUS 951 N 30 = bo E] Fics 20-37 Gnomulus insularis (Roewer), 3 holotype (25, 28-29, 31-33). - Penis, dorsal (20) and lateral view (21); apex of penis, dorsal (22), lateral (23) and ventral view (24); apex of penis (25) and glans penis (26-27) of 3 5, dorsal view. Anterior body of 3, dorsal (28) and lateral view (29), anterior body of ?, lateral view (30); left chelicera (31), left palp (32) and distal part of left leg II (33), retrolateral view; eye tubercles of 3 4 (34-36) and 1 2 (37) from Langkawi Island, dorsal view. - Scale lines 0.1 mm (22-27), 0.5 mm (20-21), 1.0 mm (28-37). 952 P. J. SCHWENDINGER & J. MARTENS Diagnosis: Medium-sized species with pointed, anteriad-inclined eye tubercle (Figs 29-30; ROEWER 1927: fig. 4b), pentagonal in dorsal view, with lateral margins converging from base to tip (Figs 28, 34-37); carapace-abdomen bridge composed of two opposing pairs of conical processes (Fig. 28); scutal areas only indistinctly elevated, ventral ones of & 4 more densely covered with fine hairs than in 9 9; chelicerae weak, with disto-dorsal boss and low ventro-median hump (Fig. 31); palp with distad-inclined process ventrally on trochanter and conical proximo-ventral process on femur (Fig. 32); distitarsus II about 1.6 times longer than wide (Fig. 33); coxa I ventrally with small anterio-lateral process, coxa II ventrally with anterio- and posterio-proximal processes, coxa III ventrally with anterio-proximal one. Penis fairly stout; truncus with a constriction behind proximal 1/4, apex very narrow, clearly separated from rest of truncus (Fig. 24). Glans penis with very large membraneous socket obscuring apex of truncus in dorsal view; lateral sclerites forceps-like, its apices split into a scoop-like and a filiform part; median plate very narrow, split into 2-4 pointed tips; membraneous tubes visible below subbasal constriction of lateral sclerites; stylus very wide, shaped like a flattened bell, with a quadrangular extension projecting from the inner side behind the opening of the sperm duct (Figs 20-27, see also Fig. 69c-e). Variation: Measurements (4, in brackets 9): body 5.36-5.83 (6.17) long, 3.20-3.52 (3.70) wide; carapace region 1.25-1.41 (1.44) long, 1.82-1.93 (1.90) wide; leg II (measured in its entirety) 7.67-8.29 (8.23) long; d (n = 4), 2 (n = 1). The median plate of the glans penis is distally split into 2-4 tips (Figs 22, 25-27). The observed variation in the shape of the eye tubercles is only slight (Figs 28, 34-37). Relationships: Gnomulus insularis is closest to G. rostratus. Distribution: Apparently present on two islands off the northern west coast of peninsular Malaysia (Fig. 1). However, the report of the type specimen from Penang Island appears doubtful, since all other conspecific specimens available were collec- ted on Langkawi Island, about 120 km to the north. Similar oncopodids, collected on Penang Island quite recently, all belong to the closely related G. rostratus (see below). Therefore, either both species occur on Penang Island (with the possibility of G. rostratus being introduced from the mainland), or the single record of G. insularis from Penang is incorrect. Considering that closely related species of Oncopodidae usually occur in allopatry and taking into account the unreliability of Roewer's locality indications (see HELVERSEN & MARTENS 1972), we are in favour of the latter explanation. Gnomulus rostratus Thorell, 1890 Figs 38-69 Gnomulus rostratus Thorell (1890: 378-381); description of species, without distinction of sexes. - ROEWER (1923: 61). - SCHWENDINGER (1992: 197). Type: MALAYSIA, Pulo Pinang (= Penang Island), 2 holotype (MSNG, examined), leg. L. Fea & L. Loria, 1889. Other material: MALAYSIA, Penang Island, Penang Hill (= Bukit Bendera), 650-760 m, 1 & (decayed, penis complete), 1 2 (19.1.1995), 710 m, 2 & (8.-9.XII.1997); - Perak, forest ca. 5 km northeast of Chenderiang, 290-330 m, 4 4,1 2 (22.-31.1.1994), 330-400 m, 1 8,12 (15.-22.1.1995); all specimens leg. P. Schwendinger. - Perak, without locality, 1 2 (SMF 5104, "Pelitnus segnipes 3", Roewer det. 1932). THE GENUS GNOMULUS 953 FIGs 38-67 Gnomulus rostratus Thorell, © holotype (55-57). - Penis of d from Penang, dorsal view (39), lateral view (40), apex of penis, dorsal (41), lateral view (42); apex of penis of other d from Penang (43); total penis (38) and apex of penis (44) of d from Chenderiang; median plate of glans of 3 d from Penang (45-47) and of 5 & from Chenderiang (48-52). Anterior body of 6. lateral view (53) and dorsal view (54); left chelicera (55), left palp (56) and distal part of right leg II (57), retrolateral view; eye tubercle of 2 2 (58-59) and 4 & (60-63) from Chenderiang, of 1 © from Perak (64), of 2 & (65-66) and 1 © (67) from Penang, dorsal view. - Scale lines 0.1 mm (41-52), 0.5 mm (38-40), 1.0 mm (53-67). 954 P. J. SCHWENDINGER & J. MARTENS Diagnosis: Very similar to G. insularis, distinguished by eye tubercle wider in dorsal view, with lateral margins parallel or divergent (from base towards tip) in basal part (Figs 54, 58-66); dorsal scutum of 9 lower (Fig. 68c); distitarsus II about 1.8 times longer than wide (Fig. 57). Penis different in: apex of truncus penis much wider; membraneous socket of glans smaller, not covering apex of truncus in dorsal view; lateral sclerites less distinctly sigmoid, covering lateral parts of stylus; median plate entire, about as long as stylus, distal margin evenly truncate, or widened and invaginated (Figs 38-52, Fig. 69c-e). Variation: Measurements range (d, in brackets 9 ): body 5.42-5.86 (5.42-6.26) long, 3.21-3.62 (3.27-3.90) wide; carapace region 1.45-1.65 (1.38-1.68) long, 1.79- 2.09 (1.79-2.14) wide; leg II (measured in its entirety) 7.89-9.10 (7.39-8.91) long; & (n= 8), 2 (n = 5). The specimens from Chenderiang (n = 7) all have a slightly wider body and a longer leg I than the others. The truncus penis of 4 & from Chenderiang (Fig. 38) is stouter than in those from Penang (Fig. 39). The median plate of the glans is distally widened and invaginated in all 4 & from Chenderiang (Figs 48-52) and in one from Penang (Fig. 47), in 2 é from Penang it is parallel-sided and evenly truncate (Figs 45-46). Relationships: Gnomulus rostratus and G. insularis are close to each other but quite distinct from the other species in Gnomulus. Distribution: Known from Penang Island and from two localities (one of them not specified) in Perak, on the Malay Peninsula (Fig. 1). A juvenile specimen from Maxwell Hill (= Bukit Larut, near Taiping, 1320 m, leg. P. Schwendinger), which lies in between Penang and Chenderiang, probably also belongs to G. rostratus. Remarks: It first appeared that the © type of G. rostratus and the d type of G. insularis are conspecific (see MARTENS & SCHWENDINGER 1998: 526-527) since both are supposed to originate from Penang Island. After thorough examination and comparison with the newly collected specimens from Penang and Langkawi Islands, however, it turned out that the two types really do belong to different species. The G. rostratus type (Fig. 68) has the same basally wide eye tubercle (seen in dorsal view) as all new specimens from Penang and Perak (Figs 54, 58-67), but it has a distinctly less elevated dorsal scutum than the G. insularis © from Langkawi (Fig. 30). The G. insularis type, on the other hand, has a basally narrow eye tubercle (Fig. 28) and a penis (Fig. 25) as in the specimens from Langkawi (Figs 22, 26-27). The pubescence on the ventral scutal areas of d d is more conspicuous than in ? 2, best seen in dry specimens. The hair shafts are coated by a crust arranged in longitudinal furrows (Figs 69a, b), which makes these hairs appear stronger and lighter in colour than the bare hairs on other parts of the d body or on the ventral scutal areas of © ©. The crust is made of a crystallized secretion, which seemingly originates from glands at the hair bases. Such modified hairs have already been recognized in G. piliger and P. pulvillatus by Pocock (1903: 413 - "transverse bands of coarse pubescence") and G. insularis by ROEWER (1927: 268 - "Querband kurzer Haare") and they occur in a number in other species as well. What significance these hairs have (presumably before or during mating), has yet to be learnt from histolo- gical studies and biological observations. THE GENUS GNOMULUS FIG. 68 Gnomulus rostratus Thorell, 2 (holotype, with colour pattern drawn from syntopic cimen), body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm. 2 spe- 956 P. J. SCHWENDINGER & J. MARTENS FIG. 69 Scanning electron micrographs of Gnomulus rostratus 3 .- Hairs on ventral scutal areas coated in a crust of crystallized secretion (a), the same with crust partly broken off (b). Distal part of penis with lateral sclerites on left side removed, dorsal view (c), lateral view (d); stylus, disto- lateral view (e). - Scale lines 20 um (a, b, e), 50 um (d), 100 um (ce). THE ASLI-GROUP Diagnosis: Small to very small species with rounded eye tubercles and curved, narrow, acutely pointed lateral sclerites on glans penis. This group comprises three species from the Malay Peninsula. Gnomulus laruticus Martens & Schwendinger, 1998 Gnomulus laruticus Martens & Schwendinger (1998: 539-542, figs 105-113); description of d. Type: MALAYSIA, Perak, Taiping, Maxwell Hill, 1320 m, & holotype (MHNG, examined), leg. P. Schwendinger, 26.1.1995. THE GENUS GNOMULUS 957 Remark: No new material available. Relationships: Despite its unusual tarsal formula (2-2-2-2; shared only by G. crucifer), genital morphology shows that G. laruticus is closely related to G. asli and G. hirsutus. Distribution: Known only from the summit area of Maxwell Hill (= Bukit Larut) in peninsular Malaysia (Fig. 1). Gnomulus asli Martens & Schwendinger, 1998 Gnomulus asli Martens & Schwendinger (1998: 542-544, figs 114-122); description of d and 9. Types: MALAYSIA, Perak, forest 5 km northeast of Chenderiang, 290-400 m, 4 holo- type (MHNG, examined), 2 d, 7 © paratypes (MAR, MHNG, examined), leg. P. Schwen- dinger, 1.1994, 1.1995. Remark: No new material available. Distribution: Known only from the type locality in peninsular Malaysıa (Fig. 1), where G. asli occurs syntopically with G. rostratus, with a third Gnomulus species (carrying a conical eye tubercle; | juvenile available) and with an Oncopus species (1 ® available). Gnomulus hirsutus Martens & Schwendinger, 1998 Gnomulus hirsutus Martens & Schwendinger (1998: 545-548, figs 123-133); description of d and 9. Types: MALAYSIA, Selangor, Templer Park, d holotype (MHNG, examined), leg. C. Deeleman-Reinhold, 3.XII.1990; Templer Park and Ulu Gombak, 200 m, road to Genting Highlands, 500-800 m, 2 4, 1 2 paratypes (MAR, MHNG, examined), leg. D. Agosti - C. Deeleman-Reinhold - H. Ono. Remark: No new material available. Relationships: Gnomulus hirsutus and G. asli are closest relatives; G. laruticus appears to be more distant. Distribution: Known from the area north of Kuala Lumpur, peninsular Malaysia (Fig. 1). THE SUMATRANUS-GROUP Diagnosis: Large species with: massive chelicerae (especially in males), pro- ximal and subdistal processes ventrally on palpal femora, low interocular areas, transversal keels on dorsal scuta and widely separated narrow, acutely pointed lateral sclerites on glans penis; anterio-proximal processes on ventral side of coxa II absent. At present, this group is represented only by G. sumatranus from W-Sumatra; a related, undescribed species occurs in N-Sumatra. Gnomulus sumatranus Thorell, 1891 Figs 70-87 Gnomulus sumatranus Thorell (1891: 759-763, figs 37-40); description of species, without distinction of sexes. - LOMAN (1893: 25). - LoMAN [1903: figs O, V-f (illustration of penis), pl. 11, figs 19, 21]. - ROEWER (1923: 61-62, figs 64a-c). - SCHWENDINGER (1992: 197). 958 P. J. SCHWENDINGER & J. MARTENS Types: SUMATRA, West Sumatra Province (Sumatera Barat), Mt. Singalang (= Gunung Singgalang); 1 4,3 9, 2 juv. syntypes (MSNG, examined), leg. O. Beccari, VII.1878. 4 here designated lectotype, because illustrated in the original description and carrying most relevant characters. Other material: From the type locality, 1 d with missing penis (removed by Loman; ZMA), leg. M. Weber; Gunung Singgalang, 2100-2600 m, 1 juv. (MAR), leg. A. Riedel, 16.X.1990. Diagnosis: Distinguished by high carapace with indistinct eye tubercle; dorsal scutal areas with transversal keels (Figs 74-77); ventral side of coxa I with quite large, outwards-directed process, coxa II ventrally without anterio-proximal process but with short, broadly rounded posterio-proximal process, coxa III ventrally with indis- tinct anterio-lateral process (Fig. 78); chelicerae very strong, with a triangular pro- ventral process on proximal article (Figs 79-81); palpal femur ventrally with a large, strongly distad-inclined proximal process and a short, conical subdistal process; palpal trochanter with a distad-inclined ventral process and a small, pointed dorsal process (Figs 82-86); distitarsus II about 2.7 times longer than wide (Fig. 87). Exter- nal sexual dimorphism in: & d with longer, higher carapace (Figs 74-75), stronger chelicerae (Fig. 79), stronger processes ventrally on palpal femur (Figs 82-83) and more elevated ventral scutal areas (Fig. 74). Penis slender, with circular fold around subbasal truncus; glans penis quite remote from rounded apex of truncus; one seta on each side of membraneous socket of glans; lateral sclerites widely apart, very slender and acutely pointed, only little bent and running parallel to each other; median plate rounded, not covering tips of membraneous tubes; stylus long and slender (Figs 70-73). Variation: Measurements (¢, in brackets 9 ): body 8.81-8.94 (8.19-8.47) long, 5.81-6.28 (5.56-5.81) wide, carapace region 3.44-3.69 (2.75-2.88) long, 4.44-4.47 (3.56-3.81) wide; d (n= 2), £ (n = 3). The larger juvenile from the type series (7.79 long) has 3-segmented posterior tarsi; in the smaller one (6.17 long) they are still 2- segmented (THORELL 1891: fig. 40). The penis (probably lost) of the d in ZMA, portrayed by LOMAN (1903: fig. V- f), seems to have a pronounced swelling at the position where the circular fold is found on the penis of the lectotype. Relationships: Externally G. sumatranus appears very distinct from all other congeners, but penis morphology is quite similar to species of the asli-group. Distribution: Known only from the surroundings of Gunung Singgalang, a mountain ca. 20 km south of Bukittinggi, West Sumatra Province (Fig. 1). THE ARMILLATUS-GROUP Diagnosis: Medium-sized to large species with more or less distinctly elevated conical (often pointed) eye tubercles and lateral glans sclerites, which are either paddle-shaped and bent, or narrow and pointed with a sigmoid or strongly curved course. This group includes 11 species from peninsular Malaysia, Sumatra, Java, Borneo and Palawan Island (Philippines). THE GENUS GNOMULUS 959 Fics 70-87 Gnomulus sumatranus Thorell, 3 lectotype (70-75, 78-80, 82, 87). - Penis, dorsal (70) and lateral view (71); apex of penis, dorsal (72) and lateral view (73). Body of & (74) and of 9 (76), lateral view; anterior body and proximal article of chelicerae of d (75) and of © (77), dorsal view; prosoma and genital operculum, ventral view (78); left chelicera of & (79) and of 2 (81), retrolateral view; proximal article of & chelicera, proventral view (80); left palp, retrolateral view (82); trochanter and femur of left palp of other & (83) and of 3 2 (84-86; 86 showing right palp), retrolateral view; distal part of left leg II, retrolateral view (87). - Scale lines 0.1 mm (72-73), 0.5 mm (70-71), 1.0 mm (74-87). 960 P. J. SCHWENDINGER & J. MARTENS Gnomulus piliger (Pocock, 1903) Figs 88-97 Pelitnus piliger Pocock (1903: 413); description of species, without distinction of sexes. - ROEWER (1923: 64). - SCHWENDINGER (1992: 197). Gnomulus piliger (Pocock): MARTENS & SCHWENDINGER (1998: 527). Type: THAILAND, Bukit Besar, 750 m, ¢ holotype (NHML, examined), leg. Annan- dale & Robinson, 4.V.1901. Diagnosis: Medium-sized species with rounded eye tubercle, wide, indistinctly divided carapace-abdomen bridge and moderately elevated dorsal scutal areas (Figs 92-93); proximal article of chelicerae with dorso-distal boss and wide retro-ventral tubercle (Fig. 95); large distad-inclined processes ventrally on trochanter and ventro- basally on femur of palp (Fig. 92), palpal femur fairly large in dorsal view (Fig. 96); distitarsus II about 2.3 times longer than wide (Fig. 97); coxa I ventrally with small anterio-lateral process, coxae II and III with pronounced anterio-proximal processes (no posterio-lateral one on coxa II; Fig. 94); ventral scutal areas swollen and covered by fine hairs. Penis slender, widening towards the apex, distal margin slightly invaginated; lateral sclerites of glans penis thin, sigmoid, median plate broadly U- shaped, covering membraneous tubes; stylus slender (Figs 88-91). Measurements: Body 5.92 long, 4.36 wide; carapace region 1.25 long, 2.24 wide; leg II (measured in its entirety) 12.45 long. Relationships: This species is very close to G. pulvillatus. Distribution: Known only from Bukit Besar (now called Khao Sankalakhiri), a mountain of 890 m altitude, situated in the Nam Tok Sai Khao National Park, about 20 km northwest of Yala city, where the borders of the provinces Songkhla, Pattani and Yala meet. Remarks: It was somewhat difficult to correctly locate "Bukit Besar", the type locality of G. piliger. In ROEWER (1923: 64) we find it located in "Malakka" (= peninsular Malaysia) and as it is a Malay name (meaning "big hill"), it seemed likely that it lies in Malaysia. Moreover, there are at least three mountains called "Bukit Besar" and another three called "Gunung Besar" (meaning "big mountain") in peninsular Malaysia. Only after receiving from the NHML the itinerary and map of the corresponding expedition (ANNANDALE & ROBINSON 1903), it became clear that the holotype was actually collected in an area which in 1901 was a Malay state under Siamese rule and today is part of Thailand. The same locality was also visited by the "Skeat-Expedition" in 1899-1900, in which Annandale took part as well (SKEAT 1954). Gnomulus pulvillatus (Pocock, 1903) Figs 98-107 Pelitnus pulvillatus Pocock (1903: 413); description of species, without distinction of sexes. - ROEWER (1923: 64). - SCHWENDINGER (1992: 197). Gnomulus pulvillatus (Pocock): MARTENS & SCHWENDINGER (1998: 527). Type: MALAYSIA, "Selangore, in cave" or "The Caves; Selangore" (as written on two labels in the vial with the type), d holotype (NHML, examined), leg. H. N. Ridley, 1897. Diagnosis: Similar to G. piliger but distinguished by a more elevated, narrow eye tubercle; ventral processes on palpal femur and trochanter smaller, less distinctly THE GENUS GNOMULUS 961 FIGs 88-97 Gnomulus piliger (Pocock), 4 holotype. - Penis, dorsal (88) and lateral view (89), apex of penis, dorsal (90) and lateral view (91). Anterior body and proximal palp, lateral view (92); anterior body and chelicerae, dorsal view (93); prosoma and genital operculum, ventral view (94); left chelicera, retrolateral view (95); trochanter and femur of left palp, dorsal view (96): distal part of left leg II, retrolateral view (97). - Scale lines 0.1 mm (90-91), 0.5 mm (88-89), 1.0 mm (92-97). distad-inclined (Fig. 102), palpal femur slightly smaller in dorsal view (Fig. 106); distitarsus II about 2.6 times as long as wide (Fig. 107); coxae II with posterio- proximal process (Fig. 104); truncus penis stouter, with an isolated pair of setae on each side of narrower membraneous socket of glans; lateral sclerites of glans slightly stronger, median plate very widely V-shaped, covering membraneous tubes; stylus slender (Figs 98-101). Measurements: Body 6.11 long, 4.36 wide; carapace region 1.25 long, 2.24 wide; leg II (measured in its entirety) 12.58 long. Relationships: Gnomulus pulvillatus and G. piliger appear closely related to G. armillatus from Sumatra. Distribution and bionomics: The caves, where the type specimen was collec- ted, are most likely the famous Batu Caves, north of Kuala Lumpur (Fig. 1). Ridley was the first to carry out a zoological investigation of these caves, which he then called the "Selangore Caves" (RIDLEY 1898). 962 P. J. SCHWENDINGER & J. MARTENS Fics 98-107 Gnomulus pulvillatus (Pocock), 4 holotype. - Penis, dorsal (98) and lateral view (99); apex of penis, dorsal (100) and lateral view (101). Anterior body and proximal palp, lateral view (102); anterior body and chelicerae, dorsal view (103); prosoma and genital operculum, ventral view (104); left chelicera, retrolateral view (105); trochanter and femur of left palp, dorsal view (106); distal part of left leg II, retrolateral view (107). - Scale lines 0.1 mm (82-83), 0.5 mm (80-81), 1.0 mm (84-89). It is, however, fairly improbable that G. pulvillatus is a troglobite. The type specimen shows no adaptations to cave life and no other oncopodids are so far known to live in caves. The animal was presumably collected from the leaf litter at the cave entrance or in the nearby rain forest. Remark: 1 ® and 2 juv. specimens, reported from Ulu Gombak (just a few km away from the Batu Caves) and identified as G. drescoi by SUZUKI (1983), possibly also belong to G. pulvillatus. Both species are very similar in external morphology. Gnomulus drescoi (Silhavy, 1962) Pelitnus drescoi Silhavy (1962: 464-466, figs 1-9); description of 2. - SUZUKI (1982: 186, figs 8-14); description of d, here considered conspecific with G. armillatus. - SCHWEN- DINGER (1992: 188, 197, figs 54-55). Gnomulus drescoi (Silhavy): MARTENS & SCHWENDINGER (1998: 526). Type: SUMATRA, Dolok Banol or Telok Belong (SILHAVY 1962: 466) or Dolok Barol (ROEWER 1927: 268) (locality unknown), 2 holotype (MHNP 24219, examined). Diagnosis: Medium-sized species with elevated, conical eye tubercle; wide, indistinctly divided carapace-abdominal bridge; moderately elevated scutal areas; anterio-proximal processes on ventral sides of coxae II and III distinct, posterio- THE GENUS GNOMULUS 963 proximal process on ventral side of coxa II indistinct; small ventral processes on palpal trochanter and femur (SILHAVY 1962: figs 1-9). Remarks: & unknown. Silhavy obviously compared his type specimen with ROEWER'S (1923: 63, figs 66a-d) incorrect redescription and illustrations of "G. segnipes" (placed in synonymy with G. armillatus in here; see below). Had he examined 9 specimens from close to the type locality of "G. segnipes", he might not have described G. drescoi. The 2 holotype of G. drescoi accords with © 9 of G. armillatus in all relevant characters (cf. SILHAVY 1962: 466). However, as the type locality of G. drescoi - and more importantly, the d from the type locality - is not known, this species cannot be formally placed in synonymy with G. armillatus. Moreover, additional 9 © spe- cimens were found near Medan (North Sumatra Province), which are similar to, but seemingly not conspecific with, 2 £ of G. armillatus. A 6 from "Lumban, Gavt" (locality unknown; ZMC), identified as G. drescoi and illustrated by SUZUKI (1982: 186, figs 8-14), clearly belongs to G. armillatus. Relationships: Gnomulus drescoi is very close to G. armillatus, possibly conspecific with it. Distribution: Unknown; type locality not identified. The specimens recorded from Ulu Gombak (near Kuala Lumpur, Malaysia) by SUZUKI (1983), presumably belong to G. pulvillatus (see above). Gnomulus armillatus (Thorell, 1891) Figs 108-134 Pelitnus armillatus Thorell (1891: 758-759); description of juvenile. - LOMAN (1902: 182-183). - Roewer (1923: 63). - ROEWER (1927: 267-268, doubtful record from Penang Island). - SCHWENDINGER (1992: 197-198). Gnomulus armillatus (Thorell): MARTENS & SCHWENDINGER (1998: 527). Pelitnus segnipes Loman (1893: 25-26, pl. 1 , figs 13-15), nov. syn.; description of juvenile. - LoMAN (1902: 182-183, pl. 9, figs 20-21; 1903: figs P-e, Q-b); description of adults from allospecific (and conspecific?) specimens. - ROEWER (1923: 63, figs 66a-d); illustration of allospecific (and conspecific?) specimens. - ROEWER (1927: 267-268). - SCHWENDINGER (1992: 187, 197-198, figs 57-61); description of é from allospecific specimen. Gnomulus segnipes (Loman): MARTENS & SCHWENDINGER (1998: 526). Type: SUMATRA, West Sumatra Province, Ajer Mancior (= Ajir Mancior = Ajer Mantcior = Ajer Mantjor = Ajer Mantjur), 360 m, ca. 7 km west of Padangpanjang, on the road from Padangpanjang to Padang, juvenile holotype (MSNG, examined), leg. O. Beccari, VI.1878. Other material: SUMATRA, West Sumatra Province, Kajoetanam (= Kayutanam = Kaju Tanam), ca. 12 km southwest of Padangpanjang, juvenile holotype of "G. segnipes" (placed in synonymy with G. armillatus in here) (ZMA), leg. M. Weber. - Lumban, Gavt (locality unknown), 1500 m, 1 d [ZMC; illustrated under G. drescoi by Suzuki (1982: 186, figs 8-14)], leg. O. Hagerup, X1.1916. - Fort de Cock in Bukittingi, 3 2 (SMF 300). - Anai (valley at the foot of Gunung Singgalang), 400-440 m, secondary forest, 1 d (collection of C. Deeleman-Reinhold), leg. C. Deeleman-Reinhold, 10.-22.V1.1994. - Gunung Singgalang, 1500- 1750 m, in grass, 1 d (ZMT), leg. P. Lehtinen, 25.-27.IX.1978. - Padang (exact locality unknown), | d (SMF 5754). - Jambi Province, Gunung Kerinci, lower montane forest, in leaf litter, 1 d (NHML), leg. P. D. Hillyard, 11.1985. 964 P. J. SCHWENDINGER & J. MARTENS Fics 108-126 Gnomulus armillatus (Thorell). - Penis of 4 from Padang, dorsal (108) and lateral view (109); apex of penis, dorsal (110) and lateral view (111); apex of penis, dorsal view, of dd from: Anai (112), Lumban (113), Gunung Singgalang (114, truncus penis dorso-ventrally collapsed THE GENUS GNOMULUS 965 Diagnosis: Characterized by elevated, more or less pointed eye tubercle; lateral tubercles on posterior carapace small or absent (Figs 116-123); carapace-abdomen bridge wide, medially indistinctly divided or entire; scutal areas moderately elevated, ventral ones pale and swollen in d d; distinct anterio-lateral processes ventrally on coxa I, anterio-proximal ones on coxae II and III, posterio-proximal process on coxa II very indistinct or absent; chelicerae weak, proximal article slightly stronger in d d (Fig. 124), with dorso-distal boss and short, rounded ventro-median tubercle (Figs 124-125); ventral processes on palpal trochanter and femur strong in dd (Figs 119- 123), generally weaker in 9 © (Figs 116-118); distitarsus of leg II ca. 2.7 times longer than wide (Fig. 126). Penis fairly slender, distal margin of truncus straight or slightly invaginated; glans penis with sigmoid, pointed lateral sclerites; median plate very widely V-shaped, covering membraneous tubes; stylus slender (Figs 108-115). Variation: Measurements (d, in brackets 9 ): body 5.53-6.09 (6.31-6.62) long, 3.90-4.51 (4.45-4.73) wide; carapace region 1.21-1.42 (1.17-1.21) long, 2.13-2.41 (2.29-2.41) wide; leg II (measured in its entirety) 11.07-12.29 (11.68-13.87) long; d (n=5), 2 (n= 3). The juvenile type of G. armillatus (Figs 127-130; tarsi 1-1-2-2) has body length 1.91, width 1.80, the juvenile type of "G. segnipes" (placed in synonymy with G. armillatus in here) (Figs 131-133; tarsi 2-2-2-2) 4.13 and 3.27 respectively. Eye tubercle shapes in mature specimens vary from high and acutely pointed to fairly low and broadly rounded; the tubercle runs straight down to the carapace front margin or is separated from it by an indistinct step (Figs 116-123). The posterior part of the carapace-abdomen bridge is entire or medially divided by a small invagination. Lateral tubercles on posterior carapace are more or less distinct in the specimens from the north of the distribution area (Figs 116-121, 123) and absent in the one from the south (Fig. 122). Variation in penis morphology is small (Figs 110, 112-115). The penis in Fig. 114 is deformed (truncus collapsed at the height of the glans) and therefore appears different from the others. In external characters, the 3 from Gunung Kerinci (the most southern locality; Figs 115, 122) looks somewhat different from the other specimens examined but penis morphology shows conspecifity. Its low rounded eye tubercle (Fig. 122) is possibly due to individual variation and not to geographical variation. Quite different eye tubercle shapes are also seen in 3 © from Fort de Cock (Figs 116-118). External sexual dimorphism in G. armillatus is expressed in dd having a slightly stronger proximal cheliceral article (Fig. 124), longer ventral processes on their palps (Figs 119-123) and swollen ventral scutal areas. Remarks: The juvenile types of G. armillatus (Figs 127-130) and of "G. segnipes" (placed in synonymy with G. armillatus in here) (Figs 131-133) both show no distinctive characters, apart from an elevated, conical eye tubercle and a ventral behind penis), Gunung Kerinci (115). Anterior body and proximal palp, lateral view, of: 3 © from Fort de Cock, Bukittinggi (116-118), d from Lumban (119), & from Padang (120), & from Anai (121), d from Gunung Kerinci (122) and d from Gunung Singgalang (123); left chelicera of 4 (124) and of ? (125), retrolateral view; distal part of left leg II, retrolateral view (126). - Scale lines 0.1 mm (110-115), 0.5 mm (108-109), 1.0 mm (116-126). 966 P. J. SCHWENDINGER & J. MARTENS + D DR ORE 136 | Se Din. RE a Fics 127-138 Gnomulus armillatus (Thorell) (127-134), Gnomulus sp. (135-138). - Juvenile holotype (1.91 mm long, tarsi 1-1-2-2), body, dorsal (127) and lateral view (128); left chelicera (129) and left palp (130), retrolateral view. Juvenile holotype of "Gnomulus segnipes" (= G. armillatus; 4.13 mm long, tarsi 2-2-2-2), body, dorsal view (131); anterior body and proximal palp, lateral view (132); left chelicera, prolateral view (133). Juvenile from Gunung Singgalang (5.31 mm long, tarsi 2-2-3-3), anterior body and proximal palp, lateral view (134). Eye tubercles in 3 juveniles (135-137) and 1 © (138) of an undescribed species from Aceh, Sumatra, lateral view. - Scale lines 1.0 mm. process on palpal trochanter. This, however, is also present in many other Gnomulus species. Additional characters, which were previously considered significant [i.e. eye tubercle clearly set back from carapace front margin (thus forming a distinct clypeus) and palpal femur without ventro-proximal process], appear to be juvenile character states which change during postembryonic development. In all available juveniles of Gnomulus species which possess a conical eye tubercle, this is small and set back from the carapace front margin in early instars. As the animals grow, the eye tubercle becomes larger and its base usually extends to the carapace front margin, in the course of which the clypeus disappears. These gradual changes in 3 juveniles (of different size) and 1 £ of an undescribed species from the northern part of Sumatra are illustrated in Figs 135-138 (see also Fig. 134, showing a large G. armillatus juvenile). THE GENUS GNOMULUS 967 In small juveniles the ventro-proximal process on the palpal femur is a low dome, which (together with a small wart-like process on palpal trochanter) becomes increasingly prominent during postembryonic development. Three steps of this development in juveniles of G. armillatus are illustrated in Figs 130, 132 and 134. The types of G. armillatus and "G. segnipes" and the mature specimens that we attribute to this species are all very similar to each other (juvenile characters taken into account) and they originate from the same area in western Sumatra. Although this area has been relatively well sampled for oncopodids since 1878, no other congeneric species were collected there, apart from the very distinct (even in juveniles) G. sumatranus. We consider this as a sufficient reason for placing "G. segnipes" in synonymy with G. armillatus and to attribute them to mature specimens from the same area. By doing so, two of the most doubtful oncopodid species can be united and placed on a satisfactory taxonomic basis. Much confusion about the identity of "G. segnipes" (placed in synonymy with G. armillatus in here) was caused by Roewer's redescription and illustrations (ROEWER 1923: 63, figs 66a-d), which were mostly based on a d specimen from Bandjermasin (= Banjarmasin in southern Kalimantan). Among the specimens Roewer examined and considered conspecific, unfortunately this one is morpho- logically and geographically most distant to the type. Penis morphology now clearly shows that this d is distinct from G. armillatus; it will be formally described as a new species in our subsequent paper. Misguided by Roewer's unlucky choice of a representative for "G. segnipes", SCHWENDINGER (1992: 187, figs 62-66) incorrectly (but with reservation) described the male of this species from the Bandjermasin specimen. Moreover, it seemingly also led to the controversial description of G. drescoi by SILHAVY (1962; discussion see above) and to an incorrect distinction between "G. segnipes" and G. drescoi by SUZUKI (1977: table 3). It also appears that ROEWER (1923: 63) took the characters for his distinction of "G. segnipes" from at least two specimens from outside Sumatra, which belong to different species. His figure 66b obviously refers to the & from Banjarmasin, whereas the median furrow between areas I-II in fig. 66a (between areas I-IV given in the corresponding text) is not visible in the Banjarmasin specimen but in the ones from Telang or Talang (locality unknown), Borneo and from Mt. Gede, Java instead. None of the specimens mentioned have an eye tubercle as distinctly set back from the carapace front margin as shown by ROEWER (1923: fig. 66c). One of Roewer's diagnostic characters is non-existent: "4. Coxa dorsal apical lateral mit 1 stumpfen Dorn" (coxa IV dorsally with a blunt disto-lateral spur; ROEWER 1923: 63, fig 66a, also referred to by SUZUKI 1977: table 3). This is neither present in any of the specimens from Roewer's collection (now in SMF), nor in any other Gnomulus specimens examined. He seemingly had confused the upper part of two opposing distal spurs, which occur pro- and retrolaterally on all leg coxae of all oncopodid species, with a single dorsal spur. Relationships: Gnomulus armillatus is close to G. drescoi from Sumatra (possibly conspecific) and to G. piliger and G. pulvillatus from peninsular Malaysia. 968 P. J. SCHWENDINGER & J. MARTENS Distribution: Known from the surroundings of Bukittinggi and Padang in West Sumatra Province and from Gunung Kerinci at the border between the provinces of West Sumatra and Jambi (Fig. 1). The type locality, Ajer Mancior (THORELL 1891: 95), is a waterfall located in the Lembah Anai Nature Reserve, in the Anai Valley, near Gunung Singgalang (BECCARI 1930: 581; LOOSE 1996: 240). The male specimen collected by O. Hagerup at "Lumban, Gavt" (locality not identified) and reported by SUZUKI (1982: 186) is accompanied by a hand-written (probably by Suzuki) label, which reads "Lumban Gayl". This could possibly corres- pond with "Lumban Gaol", a village near Balige, at the southern end of Lake Toba, in Northern Sumatra Province, about 340 km north of the type locality. However, according to information by Dr Ib Friis from the Botanical Museum in Copenhagen where Hagerup was working (N. Scharff, personal communication), the locality in question is possibly situated in the "Padang Highlands". Previous records of "G. segnipes" from Java and Borneo (LOMAN 1902: 182; ROEWER 1923: 63) belong to different, yet undescribed species (see also SCHWEN- DINGER 1992: 187, 197-198). The record of G. armillatus from Penang Island by ROEWER (1927: 268; specimen in Museum Calcutta, not examined) is most doubtful, since he also has misidentified a G. rostratus 9 from the same locality for a "G. segnipes" 3 (see above). Gnomulus armillatus hence appears to have a fairly small distribution area, as it is the case in all other oncopodid species (also to be shown for Oncopus). Gnomulus thorelli (Sgrensen, 1932) Figs 139-143 Pelitnus thorelli Sorensen (1932: 213-215); description of species, without distinction of sexes. Not P. thorelli Schwendinger (1992: 180-182, 197, figs 14-26). Gnomulus thorelli (S@rensen): MARTENS & SCHWENDINGER (1998: 527). Types: JAVA, without locality, 2 9 syntypes (both with missing ovipositor; ZMC, examined), ded. E. Simon, "M. 27.VI.1898". Other material: JAVA, Tjıbodas (= Cibodas, near Bogor), 1400 m, 1 juv. (AMNH), leg. Dammermann, V.1922. Diagnosis: Medium-sized species with moderately elevated, conical eye tu- bercle; small lateral tubercles present on posterior carapace (Figs 140-141); carapace- abdomen bridge wide, indistinctly divided (Fig. 139); scutal areas slightly elevated; anterio-lateral process on coxa I and anterior-proximal process on coxa II distinct, posterio-lateral one on coxa II and anterio-lateral one on coxa II indistinct; chelicerae weak, proximal article with dorso-distal boss, no ventral tubercle (Fig. 142); palp with small, knob-shaped process ventrally on trochanter; ventral process on palpal femur short and slightly distad-inclined, distinctly remote from proximal margin (Figs 140- 141); distitarsus II about 2.5 times longer than wide (Fig. 143). Variation: Measurements of both © 9: body 6.42-6.67 long, 4.30-4.46 wide; 10.03 long. The eye tubercle in one © (Fig. 141) is distinctly more pointed than in the other (Fig. 140). THE GENUS GNOMULUS 969 Fics 139-143 Gnomulus thorelli (Sgrensen). - Body and chelicerae of © syntype, dorsal view (139); anterior body and proximal palp of first 2 syntype (140) and of second © syntype (141), lateral view; left chelicera (142) and distal part of left leg II (143), retrolateral view. - Scale lines 1.0 mm. Remarks: 3 unknown. Three other specimens from Java are available, which clearly differ from G. thorelli in palpal armament and eye tubercle shape. One of them is a d from Mt. Gede (near Cibodas; ZMH), identified as "G. segnipes" (placed in synonymy with G. armillatus in here) by LOMAN (1902: 182) and ROEWER (1923: 63). The others are 2 2 (without exact locality; SMF), labeled by Roewer in 1929 as "Pelitnus javanus n. sp., Typus". For all we know, such a species has neither been described, nor was it ever mentioned in the literature. These specimens will be described in our next paper. Relationships: From external characters G. thorelli appears closest to the unnamed d from Banjarmasin (Southern Kalimantan), which was previously mis- identified as "G. segnipes". Distribution: The exact type locality is unknown but the find of a conspecific juvenile (with typical sub-proximal, distad-inclined process on ventral femur) indicates that G. thorelli occurs in the mountains southeast of Bogor, western Java (Fig. 1). Gnomulus laevis (Roewer, 1915) Figs 144-158 Pelitnus laevis Roewer (1915: 128, fig. 72); description of species, without distinction of sexes. - ROEWER (1923: 64, fig, 67). - SCHWENDINGER (1992: 188-190, 197, figs 62-66); description of d. Gnomulus laevis (Roewer): MARTENS & SCHWENDINGER (1998: 526). Types: BORNEO, Mt. Tiloeng (on Roewer's label: "Tilung Berg") (locality unknown, probably in Sarawak), 2 © syntypes (SMF 1260, examined). Other material: BORNEO, without locality, 1 5 [illustrated by Schwendinger (1992: figs 62-66); MHNG]; SARAWAK, Kuching, 1 d (AMNH), leg. Krishna, 4.1V.1978: Mt. Matang (= Gunung Serapi, near Matang, west of Kuching), 2 2 (AMNH), leg. Mjöbere. Diagnosis: Distinguished by palpal femur without distinct ventro-basal process (at best represented by a low rounded hump) (Figs 148, 152-154) but with a pro- 970 P. J. SCHWENDINGER & J. MARTENS Fics 144-158 Gnomulus laevis (Roewer), 2 syntype (146, 150, 154-156). - Apex of penis, dorsal (144) and lateral view (145). Body and chelicerae, dorsal view (146); anterior body, chelicerae and proximal palp of d from Kuching, dorsal view (147), the same (without chelicera), lateral view (148); right chelicera of d (149) and of 2 (150), retrolateral view; distal part of left leg II, retrolateral view (151); trochanter and femur of left palp of d (152) and of 2 © (153-154), lateral view, the same of ©, dorsal view (155); eye tubercles of 2 2 (156-157) and of é from "Borneo" (158), lateral view. - Scale lines 0.1 mm (144-145), 1.0 mm (146-158). dorsal, median boss (Figs 147, 155); ventral process on palpal trochanter peg-like (Figs 148, 152-154); proximal article of chelicerae with dorso-distal and dorso- median bosses, hand slender (Figs 149-150); distitarsus of leg II about 2.9 times longer than wide (Fig. 151); dorsal side of leg coxa IV with two small tubercles below lateral margin of dorsal scutum (Fig. 148); eye tubercle more or less raised and conical (Figs 148, 156-158); carapace-abdomen bridge wide, composed of a pair of slightly recurved lobes (Figs 146-147); small lateral tubercles on posterior carapace present (Figs 146-148); dorsal scutal areas only slightly elevated; coxa I ventrally with small anterio-lateral process, coxa II with anterio- and posterio-proximal pro- cesses, the latter overlapping anterio-proximal process on coxa III. External sexual dimorphism in: d d with stronger proximal article of chelicerae (Fig. 149), more developed prodorsal boss on palpal femur (Fig. 147) and pale, swollen ventral scutal areas. Penis fairly slender, apex of truncus carrying plenty of lateral setae from tip to below glans; distal margin of truncus penis broadly arched; glans short, wider than THE GENUS GNOMULUS 971 truncus; lateral sclerites strongly convex laterally, their tips paddle-shaped, pointing away from the truncus; median plate rounded, with lateral teeth, covering membra- neous tubes; stylus slender (Figs 144-145; see also SCHWENDINGER 1992: figs 63-66). Variation: Measurements (d, in brackets £): body 6.91-7.88 (6.94-7.88) long, 4.63-5.25 (4.63-5.13) wide; carapace region 1.44-1.75 (1.34-1.53) long, 2.59-2.94 (2.41-2.66) wide; leg II (measured in its entirety) 14.52-15.08 (12.36-14.58) long; d (n= 2), 2 (n= 4). The eye tubercle is high and pointed in one 9 syntype (Fig. 156) and low and rounded in the é from Kuching (Fig. 148); the other specimens are intermediate (Figs 157-158). The second 9 syntype and one 9 from Mt. Matang have a narrowly rounded posterior margin on the dorsal scutum, making the body appear almost elliptical (Fig. 146); in the other specimens the dorsal scutum is more rounded behind. Relationships: In penis morphology, G. /aevis is more similar to the unnamed dé from Banjarmasin, Kalimantan (SCHWENDINGER 1992: figs 58-61, sub "G. seg- nipes") than to the sympatric G. sundaicus. Distribution: Known from the surroundings of Kuching, Sarawak (Fig. 1). The type locality, Mt. Tiloeng or Mt. Tilung, however, remains unknown. There is a Mt. (Gunung) Tilong, mentioned by BECCARI (1904: 254, 303), which apparently is one of the major mountains between Sarawak and Kalimantan ["... in the interior from which the principal rivers in Borneo flow."]. This, however, would-be in the central part of Borneo, quite far away from Kuching, and therefore may not correspond with the type locality of G. laevis. Gnomulus sundaicus (Schwendinger, 1992) Figs 159-160 Pelitnus sundaicus Schwendinger (1992: 185-187, 197, figs 42-53); description of d and ®. Gnomulus sundaicus (Schwendinger): MARTENS & SCHWENDINGER (1998: 527). Types: SARAWAK, environs of Bau, Serian and Kuching, 10-50 m, d holotype, 2 6, 2 © paratypes (MHNG, examined), leg. C. Lienhard, 3.-13.X11.1987. Other material: BORNEO, Mus Twin (locality unknown), 1 & (with label: "Pelitnus modestus n. sp.", ZMC); SARAWAK, without exact locality, 1 $ (SMF 5336, det. Pelitnus laevis by Roewer in 1935). Remarks: "Pelitnus modestus" has never been described (collection name). The penis of this specimen (Figs 159-160) clearly shows that it belongs to G. sundaicus (cf. SCHWENDINGER 1992: figs 48-53). Relationships: Externally G. armillatus resembles the long-legged G. annu- lipes, whereas from penis morphology it appears closer to G. armillatus. Distribution: Known from the area south of Kuching, western Sarawak (Fig. 1). Gnomulus annulipes (Pocock, 1897) Figs 161-171 Pelitnus annulipes Pocock (1897: 283-285); description of species, without distinction of sexes. - ROEWER (1923: 63-64). - SCHWENDINGER (1992: 188, 197, fig. 56). Gnomulus annulipes (Pocock): MARTENS & SCHWENDINGER (1998: 527). Type: SARAWAK, Baram, 9 holotype (NHML, examined), leg. C. Hose. Other material: Baram River, 1 2 (SMF 5103, det. "Pelitnus segnipes" by Roewer in 1932); Gunung Mulu National Park, Pinnacle Camp (Camp 5), 1200 m, limestone scrub, 1 d (NHML), leg. F. Wanless, 29.V.1978. 972 P. J. SCHWENDINGER & J. MARTENS ZUR yr Fics 159-160 Gnomulus sundaicus (Schwendinger), 4 specimen from ZMC, labeled "Pelitnus modestus". - Apex of penis, dorsal (159) and lateral view (160). - Scale line 0.1 mm. Diagnosis: Characterized by a narrow, more or less distinctly elevated, slightly forward-inclined eye tubercle (Figs 166-168); carapace-abdomen bridge wide, sub- division indistinct (Fig. 165); lateral tubercles on posterior carapace present; dorsal scutal areas markedly elevated; ventro-proximal process on palpal femur more or less distinct, ventral process on trochanter peg-like (Figs 166-168); proximal article of chelicerae with dorso-distal and dorso-median boss, hand slender (Figs 169-170); legs quite long; distitarsus of leg II 3.4-3.9 times as long as wide (Fig. 171); coxa I ventrally with distinct anterio-lateral process, coxa II with anterio- and posterio-pro- ximal processes, coxa III with indistinct anterio-proximal one. External sexual dimor- phism in: d with a stronger proximal cheliceral article (Fig. 169), with proximal tarsalia on legs HII and IV distinctly wider than distal one and with pale, swollen ventral scutal areas. Penis fairly slender, truncus constricted at height of glans, distal margin only slightly arched and medially invaginated, apex with plenty of lateral setae (not reaching below glans); lateral sclerites of glans with dorsal margins eleva- ted above median plate, tips of sclerites drawn into small sigmoid hooks; median plate widely V-shaped, covering short membraneous tubes; stylus slender (Figs 161-164). Variation: Measurements (d, in brackets 9): body 7.35 (6.73, holotype 7.03) long, 4.86 (4.68) wide; carapace region 1.49 (1.28) long, 2.62 (2.40) wide; leg II (measured in its entirety) 17.36 (15.00) long; d (n= 1), £ (n = 2). Variation in the shape of the eye tubercle, see Figs 166-168. The tubercle on the right side of the posterior carapace of the © holotype is unusually large (Fig. 167); on its left side it is of the same size as in the other specimens (see Figs 166, 168). In the 2 © examined, the carapace-abdomen bridge is more distinctly divided than in the d (Fig. 165). The dé has a smaller proximal process ventrally on its palpal femur (Fig. 166) and a much larger anterio-lateral process ventrally on its coxa I than the © ©. This is more likely due to individual variation than to sexual dimorphism. THE GENUS GNOMULUS 973 Fics 161-171 Gnomulus annulipes (Pocock), holotypus (167, 170). - Penis, dorsal view (161) and lateral view (162); apex of penis, dorsal view (163) and lateral view (164). Body and chelicerae of d, dorsal view (165); anterior body and proximal palp of & (166) and of 2 2 (167-168), lateral view; left chelicera of & (169) and of (170), retrolateral view; distal part of left leg II, retro- lateral view (171). - Scale lines 0.1 mm (163-164), 0.5 mm (161-162), 1.0 mm (165-171). Relationships: This species is close to congeners from Borneo. Its penis is most similar to that of an undescribed species from eastern Sarawak and Sabah, which will be described in our next paper. Distribution: Known from the catchment area of the river Baram, northeastern Sarawak, near the border to Brunei (Fig. 1). Gnomulus baharu Schwendinger, 1998 Pelitnus thorelli Schwendinger (1992: 180-182, figs 14-26, pl. 1); description of d and ®. Not P. thorelli Sgrensen, 1932: 213-215. Gnomulus baharu Schwendinger in MARTENS & SCHWENDINGER (1998: 527); transferred from "Pelitnus" and renamed because of homonymy. Types: BRUNEI, Andulau Forest Reserve and Sungai Lubang Barus, & holotype, 1 à, 1 2 paratypes (MHNG, examined), leg. B. Hauser, 16./26.X1.1988. 974 P. J. SCHWENDINGER & J. MARTENS Remark: No new material available. Relationships: This species is close to G. conigerus and probably also to G. palawanensis (3 unknown). Distribution: Known from the area west of Bandar Seri Begawan, western Brunei (Fig. 1). Gnomulus conigerus (Schwendinger, 1992) Pelitnus conigerus Schwendinger (1992: 183, 197, figs 27-41); description of d. Gnomulus conigerus (Schwendinger): MARTENS & SCHWENDINGER (1998: 527). Types: SABAH, Kabili-Sepilok Forest Reserve and Kolapis, both near Sandakan, 30-60 m, 4 holotype, 1 d, 1 juv. paratypes (MHNG, examined), leg. B. Hauser, 12.11.1983. Remark: No new material available. Relationships: Gnomulus conigerus is closest to G. baharu. Distribution: Known from the area north of the Sandakan Bay, eastern Sabah (Fig. 1). Gnomulus palawanensis (Suzuki, 1982) Pelitnus palawanensis Suzuki (1982: 184-186, figs 1-7); description of 9. - Schwendinger (1992: 198). Gnomulus palawanensis (Suzuki): MARTENS & SCHWENDINGER (1998: 527). Type: PHILIPPINES, Palawan, Mt. Mantalingajan, 1150 m, west of Brooke's Point, 9 holotype (ZMC, not examined), leg. Noona Dan Expedition, 15.IX.1961. Remark: No new material available; d unknown. Relationships: According to external morphology, G. palawanensis is closest to G. baharu and G. conigerus. Distribution: Known only from a mountain in the southwest of Palawan Island (Fig. 1): THE GOODNIGHTI-GROUP Diagnosis: Small species with low, rounded or conical, pointed eye tubercles; penes with more or less distincly enlarged stylus. This group comprises six species from the Philippines and one (assignment uncertain) from northern Borneo. Gnomulus imadatei (Suzuki, 1969) Pelitnus imadatei Suzuki (1969: 12-16, figs 1-4); description of d. - SCHWENDINGER (1992: 197). Gnomulus imadatei (Suzuki): MARTENS & SCHWENDINGER (1998: 526). Type: BRUNEI, Amo (= Sungai Amoh), ¢ holotype (Zoological Laboratory, Univer- sity of Hiroshima, not examined), leg. G. Imadaté, 24.11.1962. Remark: No new material available. Relationships: Small size (3.77 m body length) and a seemingly slightly enlarged stylus (SUZUKI 1969: fig. 4d) indicate close relationship with G. goodnighti and other species from the Philippines. THE GENUS GNOMULUS 975 Distribution: Known only from the type locality, about 40 km southeast of Bandar Seri Begawan, eastern Brunei (Fig. 1). Gnomulus maculatus Martens & Schwendinger, 1998 Gnomulus maculatus Martens & Schwendinger (1998: 530-533, figs 82-89); description of d and ©. Types: PHILIPPINES, Luzon, environs of Sagada, and a doubtful record from Puerto Galera, Mindoro, 9 holotype (MHNG, examined), | d, 2 2 paratypes (MAR, MHNG, examined), leg. L. Deharveng, 21.X1.1979-9.1.1980. Remark: No new material available. Relationships: This species 1s externally similar to G. goodnighti but internally distinct by a penis with tongue-shaped median plate and markedly enlarged, bifid stylus. Distribution: Known from the area around Sagada, in the central Cordilleras of northern Luzon (Fig. 1). Gnomulus coniceps Martens & Schwendinger, 1998 Gnomulus coniceps Martens & Schwendinger (1998: 533-536, figs 90-96); description of d. Type: PHILIPPINES, Luzon, environs of Crystal Caves near Baguio, 1500 m, d holotype (MHNG, examined), leg. L. Deharveng, 12.1.1980. Remark: No new material available. Relationships: The penis of G. coniceps superficially resembles that of G. rostratus and G. insularis, but its slightly enlarged stylus and various external charac- ters point towards a close relationship with other congeners from the Philippines. Distribution: Known only from the mountains north of Baguio, northern Luzon (Fig. 1). Gnomulus crucifer Martens & Schwendinger, 1998 Gnomulus crucifer Martens & Schwendinger (1998: 527-530, figs 71-81); description of à. Type: PHILIPPINES, Luzon, Mt. Santo Thomas near Baguio, 1850 m, & holotype (MHNG, examined), leg. L. Deharveng, 14.1.1980. Remark: No new material available. Relationships: Gnomulus crucifer is unusual in its genital and external mor- phology. Congruence in tarsal formula (2-2-2-2) with G. laruticus is obviously convergent. The strongly enlarged and modified stylus penis shows relationship with other species of the goodnighti-group. Distribution: Known only from a mountain south of Baguio, northern Luzon (rs 01): Gnomulus minor Tsurusaki, 1990 Gnomulus minor Tsurusaki (1990: 59-62, figs 1-14); description of 2. - SCHWENDINGER (1992: 198). - MARTENS & SCHWENDINGER (1998: 527). 976 P. J. SCHWENDINGER & J. MARTENS Type: PHILIPPINES, Luzon, Laguna Province, Los Banos, Mt. Maquilin (= Mt. Makiling), 2 holotype (National Science Museum, Tokyo, not examined), leg. R. Morse, 10.VI.1966. Remark: No new material available. A similar, yet undescribed species with peculiar penis morphology occurs at the same locality. Relationships: In external morphology G. minor accords with congeners from the Philippines. Distribution: Known only from the type locality, south of Lake Laguna de Bay, central Luzon (Fig. 1). Gnomulus leyteensis Martens & Schwendinger, 1998 Gnomulus leyteensis Martens & Schwendinger (1998: 536-539, figs 97-104); description of à. Type: PHILIPPINES, Leyte, Visca near Baybay, 200-500 m, d holotype (MHNG, examined), leg. Martens & Schawaller, 10.11.1991. Remark: No new material available. Relationships: In external and genital morphology this species most closely resembles G. goodnight. Distribution: Known from the mountainous area close to the west coast of southern Leyte (Fig. 1). Gnomulus goodnighti (Suzuki, 1977) Figs 172-178 Pelitnus goodnighti Suzuki (1977: 9-12); description of d and 2. - SCHWENDINGER (1992: 196, 198). Gnomulus goodnighti (Suzuki): MARTENS & SCHWENDINGER (1998: 526). Types: PHILIPPINES, Mindanao, Davao Province, Mt. McKinley (close to Mt. Apo, east of Davao City), 2520 m, collected from tree ferns in a mossy forest, 4 holotype, 1 4,1 9 paratypes (Chicago Field Museum of Natural History; not examined), leg. H. Hoogstraal & D. Heyneman, 9.X1.1946. Other material: PHILIPPINES, Mindanao, Bukidnon, Mt. Katanglad, 1480 m, ca. 15 km east of Malaybalay, 2 d (Bishop Museum, Honolulu), leg. L. Quate & C. Yoshimoto, 27.- 3932: Diagnosis: Small species with low, widely conical eye tubercle (Figs 176- 177); carapace-abdomen bridge formed by two opposing pairs of pointed processes; dorsal scutal areas elevated, with paramedian pairs of small tubercles in areas II-VII (Fig. 175); chelicerae very weak and slender; palp with fairly short femur bearing a small conical ventro-basal process; palpal trochanter ventrally with strongly distad- inclined process (Figs 176-177); distitarsus II 2.3-2.4 times longer than wide, densely covered by fine curved hairs dorsally (Fig. 178); coxa I ventrally with broadly rounded anterio-lateral process, coxa II with distinct conical anterio- and posterio- proximal processes, coxa III with indistinct anterio-proximal one. No external sexual dimorphism apparent. Penis with distinctly invaginated distal margin; lateral sclerites of glans with long pointed tips (bearing denticles on the lower side) bent towards each other and away from the truncus: dorsal margins of lateral sclerites elevated above short, widely W- or U-shaped median plate; membraneous tubes large, distal part not THE GENUS GNOMULUS 977 DU i Fics 172-178 Gnomulus goodnighti (Suzuki). - Apex of penis, dorsal (172) and lateral view (173); apex of penis of other d, dorsal view (174). Body and chelicerae, dorsal view (175); anterior body and proximal palp of 2 à, lateral view (176-177); distal part of left leg II, retrolateral view (178). - Scale lines 0.1 mm (172-174), 1.0 mm (175-178). covered by median plate; stylus markedly incrassate (Figs 172-174, see also SUZUKI 1977: fig. 2f-1). Variation: Measurements [d, in brackets 9; measurements of d holotype (paratype?) and © paratype taken from SUZUKI (1977: 9)]: body 2.91-3.27 (3.34) long, 2.13-2.38 (2.41) wide; leg II (measured in its entirety) 5.13-5.16 (5.14) long; 3 (n = 3), 2 (n= 1). Variation in the shape of eye tubercles and palpal processes is illustrated in Figs 176-177, variation in glans structures in Figs 172, 174. One & has its carapace-abdomen bridge (usually formed by two opposing pairs of teeth touching each other at their tips) not closed (Fig. 176). In contrast to the type illustrated by SUZUKI (1977: fig. la, b), the specimens examined have a dorsal scutum with a more pointed posterior margin (Fig. 175) and a longer genital operculum. Remark: Unlike in larger Gnomulus species, G. goodnighti 3 $ have no swollen ventral scutal elevations and also their proximal cheliceral article seems to be indistinguishable from that of the 2 (SUZUKI 1977: 12). Relationships: Gnomulus goodnighti 1s closest to G. leyteensis. A close rela- tionship between "G. segnipes" (placed in synonymy with G. armillatus in here) and G. drescoi on one hand and G. goodnighti on the other, as suggested by SUZUKI (1977: 6, 12), is not apparent. Distribution: Known from two mountains in Davao and Bukidnon Province on Mindanao Island (Fig. 1). 978 P. J. SCHWENDINGER & J. MARTENS DISCUSSION Most Gnomulus species described before 1998 correspond with the definition of "Pelitnus" (placed in synonymy with Gnomulus by MARTENS & SCHWENDINGER 1998: 526) sensu THORELL (1891: 93), LOMAN (1902: 182), ROEWER (1923: 62) and SORENSEN (1932: 213). They are mostly 5.5-8 mm long, possess a conical, more or less elevated eye tubercle and the tarsal formula 2-2-3-3. There were only few exceptions to this. Gnomulus imadatei is the first small species (3.8 mm) described in this genus. The second small species, G. minor (3.4 mm), 1s unusual in completely lacking an eye tubercle. However, as this species is known only from a single ©, its generic placement has yet to be confirmed by dd. Gnomulus sumatranus, on the other hand, stands apart by its large size (almost 9 mm long) and by the presence of keeled dorsal scutal elevations, which in Gnomulus is otherwhise only found in the undescribed 4 from Bandjermasin and in G. sundaicus (larger but less acute). In general appearance G. sumatranus resembles species in the genus Oncopus, but tarsal formula (2-2-3-3) and penis morphology prove that it belongs in Gnomulus. Although at present no major distinctions in penis morphology between Gnomulus and Oncopus are known (but both genera clearly differ exter- nally), the penis of G. sumatranus is unquestionably a typical Gnomulus penis, very similar to those of congeners from the same geographic region. The remaining deviating species are Gnomulus rostratus and G. insularis (both about 6 mm long). They are readily distinguishable from all other congeners by a strongly anteriad-inclined eye tubercle and a unique penis morphology. Particularly in the shape of the stylus (Fig. 69c-e), the penes of G. rostratus and G. insularis differ considerably from those of other Gnomulus species. A superficial resemblance in penis form with G. coniceps (stylus tube-like; MARTENS & SCHWENDINGER 1998: figs 90-93) from the Philippines is probably the result of convergence. Due to the widespread use of sifting techniques for collecting soil mesofauna during the last decades, many more small and untypical species were discovered and described recently, which considerably extended the traditional limits of Gnomulus (see MARTENS & SCHWENDINGER 1998). The genus now covers a much wider range of external characters, e.g., body length 2.34-8.94 mm, tarsal formula 2-2-2-2 (in G. laruticus and G. crucifer) and 2-2-3-3, eye tubercle present or absent. Variation in penis morphology is also pronounced. The most aberrant species, G. crucifer, possesses a strongly enlarged stylus (MARTENS & SCHWENDINGER 1998: figs 71-75), which shows superficial resemblance with the hypertrophical styli in the genus Caenoncopus Martens & Schwendinger (1998: figs 5, 9-10, 18-19). SPECIES GROUPS With the number of species in Gnomulus increasing, an internal grouping becomes necessary. Genital morphology hereby best reflects relationships among oncopodid species and provides the most useful criteria for a grouping. At present we can provisionally divide Gnomulus into the following six species groups: THE GENUS GNOMULUS 979 1. The rostratus-group comprises Gnomulus rostratus and G. insularis from Malaysia. Three new species will be described from Thailand and Malaysia in our next paper; many more species probably exist (1 juv. from Narathiwat Province, southern Thailand, 1 juv. from Kuala Tahan, Taman Negara, Pahang and 1 © from Bukit Timah, Singapore examined). Due to its unique penis morphology and unusual eye tubercle, this is the most clearly distinguished group within Gnomulus, obviously forming a monophyletic group. It may need to be re-evaluated for generic distinction after additional species (and d d) have become known. 2. The aborensis-group is represented by G. hyatti, G. aborensis and G. lannaianus from the northwest of the known distribution of the family. These species possess similar penes, quite strong chelicerae in males, a low interocular area and dorsal scutal areas divided by a distinct median furrow. The aborensis-group is not as clearly defined as the rostratus-group, but it also appears to be derived from a common ancestor. 3. The asli-group comprises G. laruticus, G. asli and G. hirsutus from penin- sular Malaysia. These are small to very small species with a rounded eye tubercle and with a similar penis morphology. Despite the unusual tarsal formula (2-2-2-2) of G. laruticus, they are probably close relatives. Externally these species resemble those of the goodnighti-group from the Philippines, but penis morphology indicates closer affinities with congeners from peninsular Malaysia and Sumatra. 4. The sumatranus-group presently contains only the very large (almost 9 mm body length) G. sumatranus from Sumatra, which is distinguished by a unique armament on the palpal femur, by keeled dorsal scutal areas and by a pronounced sexual dimorphism. Another species to be placed in this group shall be described in our next paper. 5. The goodnighti-group comprises small (about 3 mm long) species from the Philippines, which show a tendency towards enlargement of the stylus and modi- fication of the other parts of the glans penis. These species are: G. maculatus, G. coni- ceps, G. crucifer, G. leyteensis and G. goodnighti. Gnomulus imadatei from Brunei seemingly and Gnomulus minor (generic placement uncertain) presumably belong here too. In terms of genital morphology, this 1s clearly the most diverse group. 6. The armillatus-group is an assemblage of the remaining species, 1. e. G. piliger, G. pulvillatus, G. armillatus, G. drescoi, G. thorelli, G. laevis, G. sundai- cus, G. annulipes, G. baharu, G. conigerus and G. palawanensis. These are medium- sized species (5.8-8.6 mm) with a conical eye tubercle, distributed over peninsular Malaysia and the islands of Sumatra, Java, Borneo and Palawan. Additional species shall be described also from Thailand. External morphology in this group is fairly uniform but variation in penis morphology is more pronounced. It possibly does not represent a monophyletic group. At the present state of knowledge, only the rostratus-group appears reasonably distinct in external and genital characters. All other groups are more or less clearly linked by intermediate forms and no major morphological discontinuities can be observed which may warrant generic separation. As further untypical Gnomulus species await description, a re-evaluation of this grouping will be taken into account afterwards. 980 P. J. SCHWENDINGER & J. MARTENS INTRASPECIFIC VARIATION Intraspecific variation in Oncopodidae was previously little known because of the rarity of conspecific material available for examination. From the species decribed recently, we learned that sexual dimorphism in Gnomulus is expressed in the shape of several external characters. Males have: more elevated ventral scutal areas [present in all larger species (cf. SCHWENDINGER 1992: 196), but apparently not in the small G. laruticus, G. asli, G. maculatus, G. coniceps, G. crucifer, G. leyteensis, G. good- nighti], a denser pubescence on ventral scutal areas [distinct in G. aborensis, G. lannaianus, G. insularis, G. rostratus, probably also in G. piliger, G. pulvillatus (2 unknown); less distinct in G. asli, G. hirsutus], a longer carapace and a less elevated interocular area (in G. aborensis, G. lannaianus, G. sumatranus), stronger chelicerae (in G. sumatranus, G. aborensis, G. lannaianus, G. laevis, G. annulipes, to a lesser extent in G. armillatus) and longer processes on ventral palps (in G. armillatus). Apart from this, we find considerable variation in external characters between conspecific animals of the same sex within the same population. Eye tubercles and palpal processes often have quite individual shapes, as, e.g., seen in 3 @ of G. armillatus from Fort de Cock (Figs 116-118). The same also holds true for variation among different populations. Whenever more than just a few conspecific specimens are available for examination, we usually find fairly different eye tubercle shapes [see G. rostratus (Figs 54, 58-67), G. armillatus (Figs 116-123), G. laevis (Figs 148, 156- 158) and G. annulipes (Figs 166-168)]. This character should therefore be used with caution. A certain degree of intraspecific variation is also present in genital characters, as seen in some details (especially the median plate) of the penes of G. insularis (Figs 22, 25-27), G. rostratus (Figs 41, 43-52) and G. armillatus (Figs 110-115). For other species see SCHWENDINGER (1992) and MARTENS & SCHWENDINGER (1998). ACKNOWLEDGEMENTS Dr Bernd Hauser (MHNG) ignited this study and continuously fuelled it with specimens, literature and help in tracing localities. Further material was kindly made available by the following colleagues: Dr Ben Brugge (ZMA), Dr Hieronymus Dastych (ZMH), Dr Jason Dunlop (ZMB), Dr Christa Deeleman-Reinhold (Ossen- drecht), Dr Giuliano Doria (MSNG), Dr Manfred Grasshoff (SMF), Dr Ivan Löbl (MHNG), Mrs Janet Margerison and Dr Paul Hillyard (both NHML), Dr Pekka Lehtinen (ZMT), Dr Norman I. Platnick (AMNH), Dr Christine Rollard (MHNP), Dr Nicola) Scharff (ZMC). Dr William A. Shear (Hampden-Sydney College) provided information and specimens. Dr Jiirgen Gruber (Naturhistorisches Museum, Wien) critically read the manuscript of this paper. Dr C. Deeleman-Reinhold, Janet Margerison, Liz Price (Kuala Lumpur), Dr Robert Cunningham (Surat Thani) and Dr N. Scharff helped locating places in SE-Asia. Mrs Käthe Rehbinder (Mainz) skillfully produced the whole-animal figures. Dipl.-Biol. Peter Jäger (Mainz) drew our attention THE GENUS GNOMULUS 98] to the formally incorrect type species designation in Gnomulus and Dr Philip Tubbs (NHML) further commented on it. Dr Barbara Thaler-Knoflach assisted in raising live opilionids, Prof. Dr Konrad Thaler (Universitat Innsbruck) provided literature and research facilities. Mr Konrad Eller, Mr Willi Salvenmoser and Mr Karl Schatz (Uni- versität Innsbruck) helped producing SEM-micrographs. Dr Stephen Elliott (Chiang Mai University) checked the English text. The German Academic Exchange Service (Deutscher Akademischer Austauschdienst, Bonn) supported P. J. Schwendinger with a l-month research grant to stay in Mainz at the beginning of the oncopodid project. The Feldbausch Foundation at "Fachbereich Biologie" of Mainz University provided travel funds to J. Martens. We are grateful to all friends, colleagues and institutions mentioned. REFERENCES ANNADALE, N. & ROBINSON, H. C. 1903. Itinerary in Perak, Selangor, and the Siamese Malay States. Fasciculi Malayenses (Supplement) 1: 1-43, 1 map. BECCARI, N. 1930. Odoardo Beccari in Sumatra e la scoperta dell’ "Amorphophallus Titanum" (Frammenti di diario inediti transcritti ed ordinati). Bolletino della R. Società Geografico Italiana (serie 6) 7: 569-595. BECCARI, O. 1904. Wanderings in the great forests of Borneo. Archibald Constable & Co. Ltd., London. HELVERSEN, O. von & MARTENS, J. 1972. Unrichtige Fundort-Angaben in der Arachniden- Sammlung Roewer. Senckenbergiana biologica 53 (1/2): 109-123. INTERNATIONAL TRUST FOR ZOOLOGICAL NOMENCLATURE, 1985. International Code of Zoological Nomenclature (3" edition). University of California Press, Berkley and Los Angeles. Loman, J. C. C. 1893. Opilioniden von Sumatra, Java und Flores. Jn: WEBER, M. (ed.). Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien 3: 1-26. Leyden. LOMAN, J. C. C. 1902. Neue aussereuropäische Opilioniden. Zoologische Jahrbücher, Abteilung für Systematik 16: 163-216. LOMAN, J. C. C. 1903. Vergleichend anatomische Untersuchungen an chilenischen und anderen Opilioniden. Zoologische Jahrbücher (Supplement 6) 3: 117-200, plates X-XIH. Loose, S. 1996. Sumatra. Travel Handbuch, Band 15. Stefan Loose Verlag, Berlin. MARTENS, J. 1977. Opiliones aus dem Nepal-Himalaya. HI. Oncopodidae, Phalangodidae, Assamiidae (Arachnida). Senckenbergiana biologica 57: 295-340. MARTENS, J. & SCHWENDINGER, P. 1998. A taxonomic revision of the family Oncopodidae I. New genera and new species of Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de Zoologie 105 (3): 499-555. Pocock, R. I. 1897. Descriptions of some new Oriental Opiliones recently received by the British Museum. Annals and Magazine of Natural History, including Zoology, Botany and Geology 19: 283-292. Pocock, R. I. 1903. On some new harvest-spiders of the order Opiliones from the southern continents. Proceedings of the Zoological Society of London 1902: 392-413. RIDLEY, H. N. 1898. The white snake of the Selangore Caves. Journal of the Straits Branch of the Royal Asiatic Society 1898: 99-101. ROEWER, C. F. 1913. Zoological Results of the Abor Expedition: XIV. Arachnida, II: Opiliones. Records of the Indian Museum 8 (3): 203-207. ROEWER, C. F. 1915. 106 neue Opilioniden. Archiv für Naturgeschichte 81 (3): 1-152. 982 P. J. SCHWENDINGER & J. MARTENS ROEWER, C. F. 1923. Die Weberknechte der Erde. Fischer, Jena. ROEWER, C. F. 1927. Weitere Weberknechte I. Abhandlungen herausgegeben vom naturwissen- schaftlichen Verein zu Bremen 26 (2): 261-402. SCHWENDINGER, P. J. 1992. New Oncopodidae (Opiliones, Laniatores) from Southeast Asia. Revue suisse de Zoologie 99 (1): 177-199. SCHWENDINGER, P. J. & MARTENS, J. 1999. Case 3116. Gnomulus Thorell, 1890 (Arachnida, Opiliones): proposed designation of G. sumatranus Thorell, 1891 as the type species. Bulletin of Zoological Nomenclature 56 (3): 171-173. SILHAVY, V. 1962. Un opilion nouveau, Pelitnus drescoi n. sp. (Fam. Oncopodidae). Bulletin du Muséum national d'histoire naturelle, Paris (série 2) 34 (6): 464-466. SKEAT, W. W. 1954. Reminiscences of the Cambridge University expedition to the north- eastern Malay states, 1899-1900. In: GIBSON-HILL, C. A. (ed.). The Cambridge Univer- sity expedition to the north-eastern Malay states, and to upper Perak, 1899-1900. Journal of the Malayan Branch of the Royal Asiatic Society 26 (4): 9-147. SORENSEN, W. 1932. Descriptiones laniatorum (Arachnidorum Opilionum Subordinis) fecit William Sgrensen opus posthumum recognovit et edidit Kai L. Hendriksen. Mémoires de l'Académie Royale des Sciences et des Lettres de Danemark, Copenhague (Section des Sciences) (série 9) 3 (4): 199-422. SUZUKI, S. 1969. On a collection of opilionids from Southeast Asia. Journal of Science of the Hiroshima University (series B, division 1, zoology) 22: 11-77. SUZUKI, S. 1977. Report on a collection of opilionids from the Philippines. Journal of Science of the Hiroshima University (series B, division 1, zoology) 27: 1-120. SUZUKI, S. 1982. Contributions to the taxonomy and zoogeography of the Opiliones of the Philippines, Bismarck and Solomon Islands. With an appendix on some related species from the Moluccas and Sumatra. Steenstrupia 8 (8): 181-225. SUZUKI, S. 1983. Additional notes on the Malaysian harvestmen (Arachnida: Opiliones). Acta arachnologica 32: 1-4. THORELL, T. 1890. Aracnidi di Pinang raccolti nel 1889 dai signori L. Loria e L. Fea. Annali del Museo civico di storia naturale di Genova (2) 10: 269-383. THORELL, T. 1891. Opilioni nuovi o poco conosciuti dell'Arcipelago Malese. Annali del Museo civico di storia naturale di Genova (2) 10: 669-770, plates VIII - IX. TSURUSAKI, N. 1990. Gnomulus minor, a new species of oncopodid harvestmen from Luzon, the Philippines. Acta arachnologica 39: 59-62. REVUE SUISSE DE ZOOLOGIE 106 (4): 983-1003: décembre 1999 Weitere Angaben zur Regenwurmfauna Frankreichs mit Beschreibung fünf neuer Arten (Oligochaeta: Lumbricidae) Andräs ZICSI & Csaba CSUZDI Zootaxonomische Forschungsgruppe der Ungarischen Akademie der Wissenschaften am Lehrstuhl für Tiersystematik und Ökologie der Eötvös-Loränd Universität, Puskin utca 3, H-1088 Budapest, Ungarn. Further contribution to the earthworm fauna (Oligochaeta: Lumbri- cidae) of France, with description of five new species and one sub- species. - The study of a larger collection of earthworms from different regions of France enables comments on genera described from this country and on some other recently described genera: Orodrilus Bouché, 1972, Prosellodrilus Bouché, 1972, Kritodrilus Bouché, 1972, Satchellius Gates, 1978, Murchieona Gates, 1978, Allolobophoridella Mrsic, 1990. Five spe- cies and one subspecies are described as new to science : Lumbricus klarae sp. n., Kritodrilus tetryae sp. n., Kritodrilus micrurus sp. n., Aporrectodea haymoziformis sp. n., Prosellodrilus albus sp. n., and Lumbricus friendi bouchei ssp. n. Key-words: Earthworms - Lumbricidae - taxonomy - France. EINLEITUNG Seit der 1972 von BOUCHE veröffentlichten Monographie über die Regen- würmer Frankreichs sind ausser den vom Erstautor (1977 a, b) erschienen kleinen Beiträgen keine weiteren Angaben über französische Vorkommen von Lumbricidae veröffentlicht worden. Offensichtlich war die Zahl der beschriebenen neuen Taxa (75) und die der wiedergefundenen bekannten Regenwürmer so überwältigend gross, dass sich ein weiteres Suchen nach dieser Tiergruppe erübrigte. Zweifelsohne hat BOUCHE (1972) durch Aufstellung von Gattungen, Beschreibung neuer Arten bzw. Unterarten eine Vielfalt neuer Kenntnisse für die Wissenschaft erbracht, die eben wegen ihrer Vielfältigkeit kaum kritisch gewertet werden konnten. Deswegen wurden seine taxonomischen Ergebnisse vorwiegend auch kritiklos übernommen, wenn nicht, dann ohne die Arten Frankreichs zu kennen, wie dies im Falle von OMODEO (1988), OMODEO & RoTA (1989) und Mrsic (1991) geschah. Da die Begrenzungen der Gattungen nur auf Grund der in Frankreich vorkommenden Arten erfolgten, die bisher beschriebenen übrigen Arten nicht berücksichtigt und bei den neuaufgestellten Gattungen keine Differenzialdiagnosen gegeben wurden, kann ohne Kenntnis dieser Taxa zu den im System der Lumbricidae vollzogenen Veränderungen auch nicht mit Manuskript angenommen am 17.06.1999 984 ANDRAS ZICSI & CSABA CSUZDI Sicherheit Stellung genommen werden. Dies ist der Grund dafiir, dass wir uns bisher keiner dieser Veränderungen angeschlossen haben. Obwohl seit der Revision des Erstautors (Zıcsı 1982) die Zahl der Taxa auf mehr als das Doppelte gestiegen ist, wird unsererseits die Aufstellung neuer Gattungen ebenfalls befürwortet. Aber dies darf unseres Erachtens nach nur in Kenntnis sämtlicher betroffener Arten durch- geführt werden und nicht nur allein aufgrund der Fauna eines Landes. Da dies die ganze Paläarktis betrifft, müssten noch zahlreiche Arten aus den verschiedensten Teilen dieser Region überprüft und Beschreibungen revidiert werden, bis ein ein- heitlicher Überblick möglich ist und eine kritische Beurteilung der Familie Lumbri- cidae erfolgen kann. Die vorliegende Arbeit hat zum Ziel, die von verschiedenen Fundorten stam- menden, in der Sammlung des Naturhistorischen Museums Genf aufbewahrten Arten aus Frankreich bekanntzugeben. Zusätzliche Aufsammlungen wurden durch den Erstautor (1982) und beide Autoren (1997) in den Pyrenäen durchgeführt. Zusätzlich zu den Beschreibung neuer bzw. zur Meldung bekannter Arten wird zu den von verschiedenen Autoren aufgestellten Gattungen Stellung genommen. Das bearbeitete Material wird in der Sammlung des Naturhistorischen Museums Genf (G), Belegexemplare auch am Tiersystematischen und Ökologischen Lehrstuhl der Universität Budapest (Z) aufbewahrt. BESPRECHUNG DER ARTEN FUNDORTLISTE Sämtliche Fundorte ohne Angabe des Sammlers beziehen sich auf die Sammeltatigkeit von P. Haymoz (Naturhistorisches Museum, Genf), die Abkürzung DFK steht für Dözsa-Farkas Klära. Dept. de la Haute-Savoie LE Carrieres de Veyrier au Pas-de-l'Echelle, 1/1. 470 m, 26.1.1974. 1/2. 9.2.1974. 1/3. 192219742 1/4. 4 97S. 1/5. 430°m, 1792197368 10197817001 0659700738 600 m, 16.9.1979. 1/9. 28.10.1979. 1/10. 9.12.1979 Chambéry le Vieux, 300 m, 26.8.1978 Fontaine, 28.7.1973 Grotte du Diable, 4/1. 19.8.1973. 4/2. 1.9.1973 La Leysse, 30.9.1979 La Motte-Servolex, 2.7.1978 Lucinges, 7/1. 730 m, 3.9.1979. 7/2. 800 m, 13.10.1979. 7/3. 750 m, 20.10.1979. 7/4 730 m, 27.10.1979. 7/5. 700 m, 27.3.1982. 7/6. 750 m, 3.10.1982 8. Monnetier, 8/1. 9.2.1975. 8/2. 750 m, 17.4.1979 9 Petit-Salève, 9/1. 700 m, 26.1.1973. 9/2. 750- 800 m, 14.4.1974. 9/3. 700 m, 26.1.1975. 9/4. 750 m, 23.2.1975. 9/5. 750 m, 18.9.1977. 9/6. 750-800 m, 25.9.1977. 9/7. 1250 m, 15.7.1981 10. Vouglans, 30.9.1979 Dept. de l'Ain lil. Bellegarde, 11/1. 8.7.1973. 11/2. 500 m, 11.4.1982. 11/3. 28.5.1983 43. Narderant - Reculet, 1500 m, Jura, 30.5.1982 NDNA WD REGENWURMFAUNA FRANKREICHS 985 [22 Beriaz, 467 m, 28.7.1973 13. Ceignes, 600 m, 10.8.1973 14. Champfromier, 700 m, 4.10.1980 [SE Grottes du Seznet, 15/12" 700: mi 8.971973. 15/2..18009m)16:941973.7215/32 700m) 11.11.1973 16. Mollou-Ravin, 16/1. 250 m, 27.9.1975. 16/2. 1.10.1972 WE Pont-d'Ain, 25.9.1974 18. Pyrimont, 28.7.1973 19: Soblay, 307 m, 10.8.1973 20. Thoiry, 16.5.1982 De Vallon d'Arlod 21/1. 5.4.1980. 21/2. 13.4.1980. 21/3. 7.9.1980. 21/4. 11.7.1981. 21/5. 26.7.1981 Dept. Saöne-et-Loire DAR Rocher du Solutré, 25.8.1973 Dept. de l'Isère 23% Verpilliere, 10.8.1974 44. St. Joseph de Riviere, 19.9.1982 45. St. Lattier les Fauries, 19.9.1982 Dept. de l'Essone 24. Auver-Saint-Georges, 26.7.1974 Dept. du Puy-de-Döme 23: Colombier des Roys, 11.4.1977 26. Orciens, 850 m, 25.6.1982 DT: Pont de Chäteau, 26.6.1982 Dept. de la Marne 28. Chavot 28/1. 1.7.1981. 28/2. 3.7.1981 29: Cuis, 1.7.1981 30. Eperney, 6.8.1979 ole Mt. de Reims, 30.6.1981 32: Rilbyla Mt., 150 m, 7.8.1981 Dept. de l'Indre 33. Buzangais, 19.5.1980 34. Grand Chaventon, 23.5.1980 35: Pointe de la Touche, 21.5.1980 Dépt. de la Gironde 36. St. Jean-d'Etampes, 26.7.1980 STE Saucats, 26.7.1980 Dept. de la Dröme 38. Eglise d'Hauterives, 1.6.1985 59) Tersanne, 39/1. 400 m, 39/2. 3.10.1981. 26.5.1985, leg. P. Haymoz und A. Rivand 40. Valras, 20.6.1984 41. Vinsobres, 21.6.1984 Dept. de la Charente-Maritime 22. Saujon, 26.6.1983 Dept. des Alpes de Haute Provence 46. Montagne de Lure, 1000 m, 3.4.1978, leg. Löbl und Bouchet Dept. de l'Hérault 47. Béziers, 9.9.1982 - 10.9.1982, leg. Zicsi 986 ANDRAS ZICSI & CSABA CSUZDI Dépt. de l'Ariège 48. Castillon, 13.9.1982, leg. Zicsi 49. Lac de Bemale, 13.9.1982, leg. Zicsi 74. Moulis - Remilassé, 230 m, (42°56'51"N: 1°05'38"O), 27.6.1997, leg. Zicsi - Csuzdi - DFK 73; Audressein, 500 m, (42°55'34"N: 1°01'10"O), 27.6.1997, leg. Ziesi - Csuzdi - DFK 76. St. Lary, 700 m, (42°45'10"N: 0°53'07"O), 27.6.1997, leg. Zicsi - Csuzdi - DFK Dept. des Pyrenees-Atlantiques 50. Camou-Cihigue, 14.9.1982, leg. Zicsi Sl, Louvie-Juzon, 15.9.1982, leg. Zicsi 2% Orolon, 14.9.1982, leg. Zicsi 33: Ossas-Suhare, 14.9.1982, leg. Zicsi 54. St. Etienne, 6.9.1982, leg. Zicsi De Ossas, 200 m, (43°08'16"N: 0°53'32"O), 22.6.1997, leg. Zicsi - Csuzdi - DFK 58. St. Engrace, 500 m, (43°08'16"N: 0°53'32"O), 22.6.1997, leg. Zicsi - Csuzdi - DFK 59! Haux, 300 m, (43°04'35"N: 0°53'57"O), 22.6.1997, leg. Zicsi - Csuzdi - DFK 60. Esterencubi, 250 m, (43°06'44"N: 1°12'28"W), 23.6.1997, leg. Zicsi - Csuzdi - DFK 61. Esterencubi - Iraty, 600 m, (43°04'11"N: 1°10'57"W), 23.6.1997, leg. Zicsi - Csuzdi - DFK 62. Beherobie, 650 m, (43°03'58"N: 1°12'57"W), 23.6.1997, leg. Zicsi - Csuzdi - DFK 63. Beherobie, Orion Bach, 600 m, (43°04'09"N: 1°12'43"W), 23.6.1997, leg. Zicsi - Csuzdi - DFK 64. St. Michel, 300 m, (43°07'50"N: 1°13'37"W), 23.6.1997, leg. Zicsi - Csuzdi - DFK 65. Col d' Osquich, 350 m, (43°13'33"N: 0°22'52"W), 24.6.1997, leg. Zicsi - Csuzdi - DFK 66. Osses, 350 m, (43° 14'10"N: 1°18'27"W), 24.6.1997, leg. Zicsi - Csuzdi - DFK 67. Col d' Ispequy, 550 m, (43° 10'54"N: 1°22'52"W), 24.6.1997, leg. Zicsi - Csuzdi - DFK 68. Arette Bach, 260-300 m, (43°05'50"N: 0°43'16"W), 25.6.1997, leg. Zicsi - Csuzdi - DFK 69. Issor, 500 m, (43°05'28"N: 0°42'09"W), 25.6.1997, leg. Zicsi - Csuzdi - DFK 70. Asasp, leg. Zicsi - Csuzdi - DFK 70/1. Le Lourdios Bach, 450 m, (43°06'26"N: 0°8'23"W), 25.6.1997. 70/2. 380 m, (43°07'08"N: 0°32'49"W) Dept. des Hautes-Pyrenees II Castelnau, 12.9.1982, leg. Zicsi 56. Loucrup, leg. Zicsi, 56/1. 15.9.1982. 56/2. 500 m, (43°07'06"N: 0°04'40"W), 26.6.1997, leg, Ziesi - Csuzdi - DFK Zul. Lanne, 400 m, (43°09'50"N: 0°0'11"W), 26.6.1997, leg. Zicsi - Csuzdi - DFK 122 Visker, Eches Bach, 400 m, (43°08'43"N: 0°02'43"W), 26.6.1997, leg. Zicsi - Csuzdi - DFK Ws Arcizac - Adour, 500 m, (43°08'51"N: 0°04'56"W), 26.6.1997, leg. Zicsi - Csuzdi - DFK Dept. de la Haute-Garonne Ul Aspet, 530 m, (42°57'57"N: 0°47'33"W), 27.6.1997, leg. Zicsi - Csuzdi - DFK Dept. Territoire-de-Belfort 78. Cesarhof, Sundgaurand, 5.6.1981, leg. Bono LUMBRICIDAE Rafinesque-Schmaltz, 1815 Da nicht nur die Zahl der Arten seit der letzten Revision (Zıcsı 1982) gestiegen ist, sondern auch die der Gattungen bzw. Untergattungen von 17 auf 44, fassen wir diese vorausgehend, nach Autoren geordnet, zusammen. REGENWURMFAUNA FRANKREICHS 987 Bouche: Diporodrilus, 1970, Ethnodrilus, 1972, Orodrilus, 1972, Prosello- drilus, 1972, Kritodrilus, 1972, Nicodrilus, 1972, Rhodonicus, 1972, Scherotheca, 1972, Opothedrilus, 1972, Spermophorodrilus, 1975. Easton: Perelia, 1983 (nom. nov. pro Svetlovia Perel, 1976). Eisen: Dendrobaena, 1873, Allolobophora, 1874. Gates: Eisenoides, 1969, Satchellius, 1978, Murchieona, 1978. Hoffmeister: Helodrilus, 1845. Linnaeus: Lumbricus, 1758. Malm: Eisenia, 1877. Michaelsen: Eiseniella, 1900. Moore: Bimastos, 1893. Mrsic: Alpodinariella, 1987, Dinariella, 1987, Creinella, 1987, Meroandriella, 1987, Allolobophoridella, 1990. Mrsic & Sapkarev: Zicsiona, 1987, Italobalkaniona,1988, Serbiona, 1988, Panoniona, 1988, Karpatodinariona, 1988. Omodeo: Cernosvitovia, 1956, Dendrodrilus, 1956, Eiseniona, 1956, Microeophila, 1956, Octodrilus, 1956, Healyella, 1989. Orleyi: Aporrectodea, 1885, Octolasion, 1885. Rosa: Eophila, 1893. Zicsi: Fitzingeria, 1978, Proctodrilus, 1985, Octodriloides, 1986. Da fiir einen Teil der angefiihrten Gattungen Merkmale nicht bekannt sind, die neuerdings zur Begrenzung der supraspezifischen Taxa ebenfalls herangezogen werden, kann ein endgiiltiger Gattungsschliissel nicht vorgelegt werden. Wir ver- suchen die Arten der fiir uns bislang unbekannten Gattungen eindeutig zu definieren und, soweit dies erforderlich ist, die Arten aus anderen Teilen der Paläarktis einzuordnen. Lumbricus L., 1758 Die meisten Arten der Gattung Lumbricus sind aus den westlichen Teilen Europas beschrieben worden, von wo eine Verschleppung nach Osten erfolgte. Wie aus den Verbreitungskarten der Arten dieser Gattung aus Frankreich (BOUCHE 1972: p. 354, 375; Abb. 50 u. 54) hervorgeht, kommen einige Arten, wie L. friendi nur in den südlichen Teilen Frankreichs, vorwiegend im Gebiet der Pyrenäen vor, wo die weitverschleppte Art L. terrestris nicht auftritt. Diese Verbreitung in Frankreich wird durch unsere Fundorte bestätigt. Das Fehlen von L. terrestris in südlicheren Teilen Europas (Diaz Cosın et al. 1992) kann auch durch unsere Erfahrungen aus Serbien (Grenze von Kosovo) bestätigt werden, wo sie nur bis Pecs anzutreffen war, südlich davon und in Griechenland konnte sie nicht gesammelt werden (Zicsi & MICHALIS 1981). Lumbricus terrestris L., 1758 Fundorte: Nr. 1/2. G/1106 2 Ex., Z/9338 1 Ex., Nr. 1/4. G/1449 1 Ex., Nr. 1/6. G/1462 1 Ex., Nr. 1/7. G/1444 2 Ex., Nr. 1/9. G/1644 3 Ex., Nr. 3. G/969 1 Ex., Nr. 7/1. G/1589 4 Ex., 988 ANDRAS ZICSI & CSABA CSUZDI Z/9507 3 Ex., Nr. 7/2. G/1608 4 Ex., Nr. 7/3. G/1770 4 Ex., Nr. 7/4. G/1607 4 Ex., Nr. 8/1. G/994 1 Ex., Nr. 8/2. G/1456 2 Ex., Nr. 9/4. G/1116 2 Ex., Nr. 9/5. G/1422 9 Ex., Nr. 9/6. G/1428 2 Ex., Nr. 11/1. G/970 1 Ex., Nr. 15/1. G/977 4 Ex., Nr. 16/1. G/1096 1 Ex., Nr. 17. G/1095 1 Ex., Nr. 19. G/981 7 Ex., Z/7824 4 Ex., Nr. 21/1. G/1658 1 Ex., Nr. 21/2. G/1669 1 Ex., Nr. 21/3. G/1654 1 Ex., Nr. 21/4. G/1788 1 Ex., Nr. 23. G/985 2 Ex., Nr. 25. G/1418 1 Ex., 7493453 Ex Nr 26) G/1915 Ex, Nr, 28/2 G/l787 MEX Nr 29 GHASOTMIRE ANT: G/2357 1 Ex., Z/10774 2 Ex., Nr. 39/2. G/2350 1 Ex., Nr. 41. G/2061 1 Ex., Nr. 42. G/2037 4 Ex., Z/9814 5 Ex., Nr. 44. G/2053 1 Ex., Z/9824 2 Ex., Nr. 45. G/2059 1 Ex. Lumbricus friendi friendi Cognetti, 1904 Fundorte: Nr. 16/1. G/1097 1 Ex., Nr. 20. G/1879 3 Ex., Nr. 42. G/2031 1 Ex., Z/9808- 09 27 Ex., Nr. 43. G/1845 9 Ex., Nr. 51. G/2065 1 Ex., Z/9841 1 Ex., Nr. 55. G/1943 1 Ex., Nr. 56/1. Z/12550 3 Ex., Nr. 57. Z/12490 7 Ex., Nr. Z/59. G/2539 3 Ex.,Z/12572 5 Ex., Nr. 69. Z/12538 3+2 Ex., Nr. 70/1.G/2543 2 Ex,,Z/12546 4+4 Ex., Nr. 71. Z/12558 1 Ex., Nr. 72. G/2541 2 Ex.,Z/12525 2 Ex., Nr. 73. Z/12560 2+1 Ex., Nr. 74. Z/12564 3+2 Ex., Nr. 75. Z/12568 3+1 Ex., Nr. 76. G/2540 1 Ex., Z/12573 3 Ex. Dies ist die häufigste in den Pyrenäen vorkommende Lumbricus Att. Lumbricus friendi bouchei ssp. n. Lumbricus friendi var. lineatus Bouché, 1972: 357 Fundorte: Holotypus Z/12558 Dept. des Hautes-Pyrénèes Lanne, 400 m leg. Zicsi- Csuzdi-DFK. Paratypen: Nr. 48 G/1945 2 Ex., Z/9783 2 Ex., Z/9785 2 Ex., Nr. 57.G/2542 4 Exe Z/ 12437 HER, NC 27112538342 Exe Nr IE 2112538 DEZ? Diagnose: L. 70-90 mm, B. 5-6 mm, Segmentzahl 102-113. Rotbraun. Kopf tanylobisch. Gürtel vom 33.-37., Pubertätsstreifen linienförmig vom 34.-36. Segment. Innere Organisation wie bei L. friendi. Bei der Bestimmung unseres Materials sind wir dem Unterschied in der Ausbildung der Pubertätsorgane, den Bouché veranlasste, die Varietät lineatus zu beschreiben, ebenfalls begegnet. Dieser Unterschied ist nicht mehr oder weniger bedeutend als der, der bei A. caliginosa caliginosa und A. caliginosa trapezoides zur Aufstellung einer Unterart führte. Da bei L. friendi und L. friendi v. lineatus auf dem 34.-36. Segment die gleichen Unterschiede bestehen stellen wir fiir v. lineatus eine neue Unterart auf. Beschreibung vergl. BOUCHE (Lumbricus friendi var. lineatus 1972: p. 357). Lumbricus klarae sp. n. Fundort: Holotypus. Z/12947. Esterencubi, 250 m, (43°06'44"N: 1°12'28"W), 23.6. 1997., leg. Zicsi+Csuzdi+D6zsa-Farkas. Paratypen. Z/12504 2+3 juv. Ex., G/2544 2 Ex., Fundort wie beim Holotypus. Diagnose: L. 39-71 mm, B. 3-4 mm, Segmentzahl 61-122. Rotbraun. Kopf tanylobisch. Borsten eng gepaart. Gürtel vom 32.-35., Pubertätsstreifen vom 33.-35. Segment. Verdickte Dissepimente fehlen. Herzen im 7.-11., Kalkdriisen im 10. Segment. Unpaarige Testikelblasen im 10. und 11. Segment. 3 Paar Samensäcke im 9., 11., 12. Segment. Typhlosolis einfach. Nephridialblasen J-förmig. Muskulatur gefiedert. 2 Paar Samentaschen im 10. und 11. Segment. REGENWURMFAUNA FRANKREICHS 989 Linge des Holotypus 71 mm, Breite 4 mm, Segmentzahl 112. Paratypen. Länge 39-68 mm, 3-4 mm, Segmentzahl 61-122. Farbe rotbraun. Kopf tanylobisch. Borsten am ganzen Körper eng gepaart. Borstendistanz hinter dem Gürtel aa:ab:bc:cd:dd wie 5:1,6:4,3:1:12. Rückenporen vor dem Gürtel undeutlich, wenn vorhanden so auf Intersegmentalfurche 6/7 oder 7/8 beginnend, hinter dem Gürtel deutlich zu erkennen. Nephridialporen oberhalb der Borstenlinie d. Segmente 8-11, 14-16, 26-31 auf der Ventralseite in der Borsrenlinie ab etwas drüsig. Borsten ab des 37. Segment von Drüsenpapillen umgeben. Samen- taschenporen auf Intersegmentalfurche 9/10 und 10/11 in der Borstenlinie c. Gürtel sattelförmig vom 32.-36. Segment, Pubertätsstreifen vom 33.-35. Segment. Innere Organisation. Dissepimente 6/7-13/14 nicht verdickt. Schlunddrüsen bis ins 6. Segment reichend. Herzen im 7.-10. Segment gross, im 11. klein. Klalkdrüsen im 10. Segment, sind abstehende, nach vorne gerichtete Ausbuchtungen. Hoden und Samentrichter im 10. und 11. Segment, in unpaarige Testikelblasen eingeschlossen. Samensäcke im 9., 11. und 12. Segment, die des 12. gross, bis ins 19. Segment reichend. Ovarien im 13. Segment, Ovarientrichter auf Intersegmentalfurche 13/14. Kropf im 15.-16. Segment, Muskelmagen im 17.-18. Segment, Thyphlosolis im 19. Segment beginnend. Nephridialblase J-förmig. Muskulatur vom gefiederten Typus. Die neu Art steht L. friendi, L. centralis und L. terrestris am nächsten, unter- scheidet sich von allen dreien in der Lage des Gürtels und der Pubertätsstreifen. Die neue Art wird zu Ehren von Frau Prof. Dr. Klara Dözsa-Farkas benannt, die uns bei den Aufsammlungen in den Pyrenäen sehr behilflich war. Bei der Beschreibung von L. improvisus (Zicsi, 1963) wurde bereits darauf hingewiesen, dass die Vertreter der Gattung Lumbricus in der Gesamtheit ihrer taxonomischen Merkmale sich äusserst nahe stehen. Für die Aufstellung einer neuen Art genügt die Verschiebung der Gürtel -und Pubertätsorgane um ein Segment. Die vorliegenden Exemplare zeigen eine solche Verschiebung, so dass wir uns ge- zwungen sehen, eine neue Art zu beschreiben. Das Vorkommen von L. improvisus in Frankreich (BOUCHE 1972: p. 373) erscheint uns als sehr fraglich, da von jedem Fundort nur Einzelstücke vorlagen. Eine Anomalie kann somit nicht ausgeschlossen werden. Lumbricus castaneus (Savigny, 1826) Fundorte: Nr. 1/1. G/1108 3 Ex., 7/9329 3 Ex., Nr. 1/2. G/1101 16 Ex., Z/ 9336 1 Ex. Nr. 1/5. G/1453 1 Ex., Nr. 1/10. G/1678 2 Ex., Nr. 3/7. G/1768 2 Ex., Nr. 4/2. G/962 4 Ex., Nr. 4/2. G/983 2 Ex., Nr. 6. G/1439 4 Ex., Nr. 7/1. G/1590 2 Ex., Z/9505 2 Ex., Nr. 7/2. G/1609 2 Exe Nr. 7/4G/1606 5 Ex., Nr. 8/1. G/995 3 Ex., Mr. 8/2. G/1457 12 Ex., Z/99322 5 Ex., Nr. IMAENEO3Z LER UNTEI2IE/IT2 REX N11: G/IEINBTER. Nr. 15/1.2G/975 3 Ex. INT: IS 2 /G'OMMPIEX Z11822 Diexts Nr, 15/511G/974A EX... ZN182373 Ex., Nr 16/2. G/1437 1 EX, N22187G/968-17 EX INLI204G/11878-167Ex., No 2 G/16S9P Ex 1Z/9SSOUNEX Nr 21/2. G/1670 5 Ex., Nr. 21/4. G/1789 1 Ex., Nr. 25. G/1417 17 Ex., Z/9346 9 Ex., Nr. 26. G/1916 3 Ex., Nr. 30. G/1470 1 Ex., Nr. 31. G/1799 4 Ex., Nr. 33. G/1598. 3 Ex., G/1627 3 Ex., G/1633 MERIZIDSIS Ex. Z/9 51 4 Exe, Nr 4 G/1625M2 ex 21055215 ex Nes IG/ISTOMS Be Z/9539 195 x NUN 372 GHICISMMEX INT 8416/2352 MExXANr B9/2NG/23462/EX Z/10768 6 Ex., Nr. 41. G/2162 8 Ex., Z/10652 9 Ex., Z/12122 30 Ex., Nr. 43. G/1847 4 Ex., Z/12123 27 Ex., Nr. 44. G/2052 3 Ex., Z/9821 7 Ex., Nr. 56/1.Z/12551 5 Ex. 990 ANDRÄS ZICSI & CSABA CSUZDI Lumbricus festivus (Savigny, 1826) Fundorte: Nr. 33. G/1597 1 Ex., G/1625 1 Ex., G/1634 5 Ex., G/1677 1 Ex., Z/9526 6 Ex., Z/9545 1 Ex., Nr. 34. G/1617 8 Ex., Z/9554 7 Ex., Nr. 35. G/1582 8 Ex., Z/9542 6 Ex. Lumbricus polyphemus (Fitzinger, 1833) Fundort: Nr. 78. Z/10725 | Ex. Lumbricus rubellus Hoffmeister, 1843 Fundorte: Nr. 1/8. G/1522 4 Ex., Nr. 3/7. G/1769 3 Ex., Nr. 4=1. G/963 6 Ex., Nr. 7/1. G/1591 EX Z/9508 ITEX. Nr. 772. G/1610 1 Ex., Nr 7/4..G/1605 2 Ex ZI N 135,G/966 2 EX Nr. 22-'G/967-5-Ex.,, Nr. 33. G/11626 4Ex:, G/1632 HEXFTZIIZISIERING 34. G/1621 5 Ex., Nr. 35. G/1580 6 Ex., Z/9540 9 Ex., Nr. 38. G/2353 3 Ex., Nr. 39/2. G/ 2347 6 Ex., Z/10769 1 Ex., Nr. 43. G/1846 1 Ex. Murchieona Gates, 1978 Allolobophora (part.), ROSA 1906: 38, MULDAL 1952: 463, MURCHIE 1959: 329, ZICSI 1968: 404, 1973: 101, 1981: 176, 1982: 433, BOUCHÉ 1972: 452, GATES 1975b: 7, Zıcsı & MICHALIS 1981: 251, TRIGO et al. 1990: 534, DIAZ COSIN et al. 1992: 1352. Bimastos (part.), PICKFORD 1926: 97, OMODEO 1956: 179, GERARD 1964: 34, MRSIC & SAPKAREV 1988: 31. Murchieona Gates, 1978: 114, EASTON 1983: 483, Sims & GERARD 1985: 109, MRSIC 199175332 Durch die ungewöhnliche Ausbildung der Kalkdrüsen im 10. Segment (ab- stehende, nach vorne gerichtete U-förmige Gebilde) und die würstchenförmigen Nephridialblasen sowie durch die Muskulatur vom gefiederten Typ, ist die Gattung in der Familie Lumbricidae gut charakterisiert. Nach Überprüfung von Exemplaren aus Griechenland (Zıcsı 1973, Zıcsı & MICHALIS 1981), Kroatien (Zıcsı 1986), Italien, (Zıcsı 1981) wird die neue Gattung auch unserseits anerkannt. Typusart: Allolobophora muldali Omodeo, 1956. Weitere Art: M. minuscula (Rosa, 1906). Murchieona muldali (Omodeo, 1956) Fundorte: Nr. 57. Z/12489 1 Ex., Nr. 62. Z/12508 1 Ex., Nr. 69. 1 Ex., G/2545 1 Ex., Nr. 70. Z/12544 1 Ex. Nach Überprüfung von Exemplaren aus verschiedenen südeuropäischen Lädern (vergl. weiter oben) konnten Populationen mit verschiedener Gürtellage unterschieden werden. Bei den Tieren aus Griechenland, Kroatien und Italien liegt der Gürtel am 26., 1/2 26.-32. Segment (M. minuscula), bei denen aus Frankreich, Portugal und den offen- sichtlich Verschleppungsformen in England und den Vereinigten Staaten von Amerika vom 27., 1/2 27.-33. Segment (M. muldali). In Anbetracht dieser konstanten Unterschiede wird die Synonymisierung von muldali widerrufen (Zıcsı 1981), beide Arten werden hiermit anerkannt. Helodrilus Hoffmeister, 1845 Helodrilus Hoffmeister, 1845: 88, MICHAELSEN 1900: 495, OMODEO 1953: 75, 1956: 171, PEREL 1976: 833, Zicsi 1985: 277, Mrsic 1991: 101. Typusart: H. oculatus Hoffmeister, 1845. REGENWURMFAUNA FRANKREICHS 99] Die von Pop (1941) synonymisierte Gattung Helodrilus wurde von Omodeo (1953) wieder ins Leben gerufen, ohne sie aber eindeutig zu begrenzen. Sie ist jedoch durch das Fehlen von Nephridialblasen charakterisiert (PEREL 1976). Derzeit sind 19 Arten bekannt (PEREL 1976, 1979, Zicsi 1985, Mrsic 1991). Bei der Bestimmung des Materials aus den Pyrenäen sind wir auf drei Arten bzw. Unterarten dieser Gattung gestossen. Helodrilus oculatus Hofmeister, 1845 Fundorte: Nr. 37. G/1694 1 Ex., Z/9603 1 Ex., Nr. 70. Z/12545 1 Ex. Von dieser Art, die zusammen mit A. putricolus gesammelt wurde, sind nur wenige Exemplare erbeutet worden. Da von A. oculatus oft nur juvenile oder prae- adulte Tiere gefangen werden (die Zeit der vollen Geschlechtsreife hält nur kurz an), ist das Variieren der Gürtel und Pubertätsorgane schwer zu beurteilen. Bei den von uns bestimmten Exemplaren liegt der Gürtel vom Fundort Nr. 37 auf dem 21.-32., die Pubertätsstreifen auf dem 1/4 28. -1/4 31. Segment, bei denen vom Fundort Nr. 70 erstreckt sich der Gürtel vom 22.-32., die Pubertätstreifen vom 1/2 28.-1/2 31. Segment. Helodrilus putricola putricola (Bouché, 1972) Allolobophora putricola putricola Bouché, 1972 : 442 Eundorte= Nr. 37.2.6/169573 Ex., Z/9604 1 Ex. Nr: 70.212542 juv Ex. Nr. #2: Z/12523, 12524, 18+3 praead.+2 juv. Ex., G/2546 3+3 juv. Ex. Vom Fundort Nr. 72 liegen uns zahlreiche Exemplare vor, die alle am Bachrand unter Steinen gesammelt wurden. Obwohl unsere Tiere einwandfrei als A. putricola identifiziert werden konnten, muss das starke Variieren der Pubertätsstreifen innerhalb einer Population hervorgehoben werden ( Lage des Gürtels vom 23., 1/2 23.-29., 30., die der Pubertätsstreifen vom 23., 24., 25.-28., 29., 1/2 29., 30. Segment). Da die Lage der Pubertätsstreifen, mit Ausnahme der Gattung Helodrilus, bei den übrigen Gattungen der Familie Lumbricidae im allgemeinen ein konstantes Artmerkmal ist und nicht mit der Geschlechtsreife der Tiere zusammenhängt, so wie dies beim Gürtel der Fall ist, muss dies bei dieser Art unbedingt erwähnt werden, damit bei Einzelfängen nicht weitere neue Taxa aufgestellt werden. Bei Schwanzregeneraten am Körperende ist der After rückenständig gelegen. Von A. putricola tebra (Bouché, 1972), die sich durch den Besitz von 3 Paar Samentaschen von der Stammform unterscheidet, wurden keine Exemplare gefangen. Helodrilus putricola orionensis (Zicsi, 1977) stat. n. Allolobophora orionensis Zicsi, 1977 : 682 Fundorte: Nr. 57 Z/12482 14 Ex., G/2547 3+1 juv. Ex., Nr. 61. Z/12505 3 Ex. In der Originalbeschreibung wurde die Form der Nephridialblasen nicht erwähnt. Eine Nachbestimmung des Holotypus und des reichen neuen Materials erbrachte den Nachweis eines Fehlens von Nephridialblasen, sodass diese Art ebenfalls zur Gattung Helodrilus gestellt werden muss. Sie ist nahe verwandt mit A. putricola, 992 ANDRAS ZICSI & CSABA CSUZDI von der sie sich durch die 5 Paar Samentaschen unterscheidet. Die Lage des Giirtels und der Pubertätsstreifen, die jetzt an mehreren Exemplaren bestimmt werden konnte, zeigt ein grösseres Variieren (Gürtel vom 23., 1/2 23., 1/2 24.- 29., 30., 31., Pubertätsstreifen vom 23., 1/2 24.- 29., 1/2 30., 30. Segment), sodass wir orionensis als Unterart von putricola betrachten. Ein rückenständiger After konnte bei einigen Exemplaren ebenfalls beobachtet werden. Kritodrilus Bouché, 1972 Kritodrilus Bouché, 1972: 250, PEREL 1976: 642, EASTON 1983: 482, OMODEO & ROTA 1989: 185. Typusart: Octolasion calarensis Tetry, 1944. Weitere Arten: K. osellai (Zicsi, 1970), K. ruffoi (Zicsi, 1970), K. pseudorosea (Moreno, 1983). Von PEREL (1976) wurde die Gattung durch die Form der Nephridialblasen (doppellappig=bilobous) eindeutig definiert. Sie reihte auch Dendrobaena auriculata (Rosa, 1897) zu Kritodrilus. Später wurden dieser Gattung von OMODEO & ROTA (1989) noch die Arten D. crassa (Michaelsen, 1900), D. clujensis Pop, 1938, D. pseudorosea Moreno, 1983, D. ruffoi Zicsi, 1970, D. osellai Zicsi, 1970, D. mrazeki (Cernosvitov, 1935) zugeordnet. Obwohl D. auriculata und D. mrazeki doppellappige Nephridialblasen besitzen, können sie wegen des gefiederten Muskeltyps nicht in diese Gattung gestellt werden, da calarensis, die Typusart, den bündelförmigen Typ besitzt. D. clujensis und D. crassa verfügen unseres Wissens über keine zweilappigen Nephri- dialblasen. Im vorliegendem Material liegen von zwei verschiedenen Fundorten einige Exemplare vor, die in diese Gattung eingereiht werden können. BOUCHE (1972) meint zur Beschreibung von K. calarensis Tetry zwar, dass es sich um eine polymorphe Art handeln soll, führt jedoch klare Bestimmungsmerkmale an, allein die Drüsenpapillen scheinen stark zu variieren. Unsere Tiere, die etwas nördlich von seinen Fundorten gesammelt wurden, weichen so deutlich von K. calarensis ab, dass wir sie als neue Arten für die Wissenschaft betrachten. Kritodrilus tetryae sp. n. Fundort: Holotypus. G/2660. Dept. de l'Ain, 1 km vor d Oncieu, 330 m.,25. 3. 1989 leg Haymoz. Paratypen. AF/12298 2 +1 juv. Ex., G/2667 2Ex. Diagnose: L. 75-115 mm, B. 7,8-8,4 mm, Segmentzahl 238-278. Unpigmen- tiert, weiss. Kopf prolobisch. Borsten eng gepaart. Gürtel vom 26., 27.-37., Puber- tätsstreifen vom 1/2 31., 32.-36., 1/2 37. Segment. Dissepimente 5/6-9/10 verdickt. Herzen im 6.-11., Kalkdrüsen im 11. Segment. Perioesophageale Testikelblasen im 10. und 11. Segment. 2 Paar Samensäcke im 11. und 12. Segment. Typhlosolis dreifach gelappt. Nephridialblasen doppellappig. Muskulatur bündelförmig. 2 Paar Samentaschen im 11. und 12. Segment. Länge des Holotypus 75 mm, Breite 8 mm, Segmentzahl 245. Paratypen. Länge 70 -115 mm, Breite 7,8-8,4 mm, Segmentzahl 238-278. REGENWURMFAUNA FRANKREICHS 993 Farbe weiss, unpigmentiert. Kopf prolobisch. I. Segment längsgefurcht, Seg- mente vom 9. doppelt oder mehrfach gefurcht. Borsten ungepaart, Borstenverhältnis hinter dem Gürtel aa:ab:be:cd:dd wie 3:2,5:2,4:1:4. Erster Rückenporus auf Inter- segmentalfurche 5/6. Nephridialporen oberhalb der Borstenlinie b, nicht alternierend. Vier grosse Drüsenpölster auf dem 11. und 30., auf dem 14. und 29. drei, auf dem 16. und 28. Segment zwei, sie umgeben entweder die Borsten ab, oder nur a. Allein auf dem 11. Segment sind die mächtigen Pölster konstant, auf den ürigen kommen sie variierend vor. Samentaschenporen auf Intersegmentalfurche 10/11 und 11/12 in der Borstenlinie c. Weibliche Poren auf dem 14. Segment, männliche Poren auf dem 15. Segment von grossen Drüsenhöfen umgeben, die benachbarten Segmente ausbreitend, ohne auf diese überzugehen. Gürtel sattelförmig vom 26., 37.-37. Segment, Puber- tätsstreifen vom 1/2 31., 32.-36., 1/2 37. Innere Organisation. Dissepimente 5/6-9/10 sehr stark, 10/11-11/12 nur schwach verdickt. Schlundkopf bis in 5. Segment reichend. Herzen im 6.-11. Segment. Kalkdrüsen im 11. Segment mit kleinen Ausbuchtungen. Hoden und Samentrichter im 10. und 11. Segment in perioesophageale, aufgefranzte Testikel- blasen eingeschlossen. 2 Paar Samensäcke im 11. und 12. Segment, ungefähr gleich- grosse Gebilde, die auf ihre Segmente beschränkt sind. Ovarien im 13. Segment, mit reifen Eiern gefüllt, grosse Ovarientrichter auf Dissepiment 13/14. Kropf im 15.-16. Segment, Muskelmagen im 17.-19. Segment. Thyphlosolis vom 20. Segment be- ginnend, dreifach gelappt, weit nach hinten reichend. Nephridialblasen im Hinter- körper doppellappig. Samentaschen 2 Paar im 11. und 12. Segment kleine, runde, sessile Gebilde. K. tetryae unterscheidet sich von K. calarensis durch die Lage des Gürtels und der Pubertätsstreifen sowie durch die Zahl der Samentaschen. Die neu Art wird zu Ehren von Frau A. Tetry benannt, die sich grosse Verdienste in der Erkundung der Regenwurmfauna Frankreichs erworben hat. Kritodrilus micrurus sp. n. Fundort: Holotypus. G/2661. Dept. de la Dröme. Coriencon pres de Vinsobres, 21. 6. 1984. Paratypus Z/12145 1 praead. Ex. Fundort wie beim Holotypus. Diagnose: L. 45-54 mm, B. 1,5-2 mm, Segmentzahl 138-148. Unpigmentiert, weiss. Kopf prolobisch. Borsten eng gepaart. Gürtel vom 26.-35., Pubertätsstreifen vom 1/4 30.-1/2 35. Segment. Dissepimente 5/6-9/10 verdickt. Herzen im 6.-11., Kalkdrüsen im 11. Segment. Perioesephageale Testikelblasen im 10. und 11. Segment. 2 Paar Samensäcke im 11. und 12. Segment. Typlosolis dreifach gelappt. Nephridialblasen doppellappig. Muskulatur bündelförmig. 3 Paar Samentaschen im WERD ESesment. Länge des Holotypus 54 mm, Breite 2 mm, Segmentzahl 148. Paratypus. Länge 45 mm, Breite 1,5 mm, Segmentzahl 138. Farbe weiss, unpigmentiert. Kopf prolobisch. Vordere Segmente ungeringelt, vom 9. doppelt geringelt. Borsten ungepaart. Borstenverhältnis hinter dem Gürtel aa:ab:bc:cd:dd wie 3,3:2:1,3:1:6,6. Grosse Polsterdriisen auf den Borsten ab des 13., 994 ANDRAS ZICSI & CSABA CSUZDI auf der Borste a des 16., und 17., auf den Borsten aa des 25. Segments. Auf dem Gürtel fehlen diese Pölster, die Borsten aa des 28.-30. Segments werden nur von kleinen runden Papillen umgeben. Beim praeadulten Exemplar sind keine Pölster vorhanden. Rückenporus auf Intersegmentalfurche 11/12 beginnend. Nephridialporen in der Borstenlinie b nicht alternierend. Samentaschenporen auf Intersegmentalfurche 9/10, 10/11, 11/12 in der Borstenlinie c. Gürtel sattelförmig vom 26.-35. Segment, Pubertätsstreifen vom 1/4 30.-1/2 35. Segment. Weibliche Poren auf dem 14. Segment, männliche Poren auf dem 15. Segment, von grossen Drüsenhöfen umgeben. Innere Organisation. Dissepimente 5/6-9/10 stark verdickt. Herzen im 6.-11. Segment. Kalkdrüsen im 11. Segment mit kleinen Ausbuchtungen. Hoden und Samen- trichter im 10. und 11. Segment in perioesophageale Testikelblasen eingeschlossen. Samensäcke im 11. und 12. Segement. Ovarien im 13. Segment, Ovarientrichter auf Dissepiment 13/14. Kropf im 15. und 16., Muskelmagen im 17.-1/2 20. Segment. Typhlosolis im 20. Segment beginnend, dreifach gelappt. Nephridialblasen hinter dem Gürtel doppellappig. Muskulatur bündelförmig. Samentaschen 3 Paar im 10., 11., 12. Segment als kleine, runde, sessile Gebilde, von vorne nach hinten grösser werdend. Die neue Art steht K. calarensis und K. tetryae sp. n. am nächsten, unter- scheidet sich von ihnen durch die Lage des Gürtels, und der Pubertätsstreifen, von K. tetryae auch durch die Zahl der Samentaschen. Dendrodrilus Omodeo, 1956 Dendrodrilus Omodeo, 1956: 175 Durch die weitgepaarten Borsten, durch die Ausbuchtungen der Kalkdrüsen im 10. Segment, den gefiederten Muskeltyp und die U-förmigen Nephridialblasen, deren Ausführungen nach vorne gerichtet sind, unterscheidet sich Dendrodrilus von allen bisher beschriebenen Gattungen der Familie Lumbricidae. Dendrodrilus rubidus rubidus (Savigny, 1826) Fundorte: Nr. 1/1. G/1109 1 Ex., Nr. 1/6. G/1463 1 Ex., Nr. 1/7. G/1445 1-Ex., Nr. 4/1. G/961 1 Ex., Nr. 4/2. G/982 9 Ex., Nr. 6. G/1440 1 Ex., Nr. 8/2. G/ 1458 2 Ex., Nr. 9/7. G/1779 1 Ex., Nr. 16/2. G/1438 1 Ex., Nr. 21/5. G/1808 2 Ex., Nr. 28/1. G/1775 1 Ex., Nr. 30. G/1469 2 Ex., Nr. 39/1. G/1781 1 Ex., Nr. 44. G/2057 2 Ex., Z/9830 3 Ex., Nr. 63. 12514 1 Ex. Dendrodrilus rubidus subrubicundus (Eisen, 1874) Fundorte: Nr. 1/2. G/1105 4 Ex., Z/9334 5 Ex., Nr. 7/5. G/1884 1 Ex., Nr. 10. G/1672 1 Ex., Z/9566 2 Ex., Nr. 25. G/1416 1 Ex., Nr. 26. G/1913 8 Ex., Nr. 33. G/1596 8 Ex., G/1630 1 Ex... G/11636 2. Ex., Z/9546 2 Ex., Nr: 37. G/1689 13 Ex., Z/9597715.Ex., Nr 39/1 G/1782531 Ex., Nr. 42. G/2040 1 Ex. Dendrobaena Eisen, 1873 Dendrobaena attemsi (Michaelsen, 1902) Fundorte: Nr. 37. G/1691 1 Ex., Z/9599 1 Ex. REGENWURMFAUNA FRANKREICHS 995 Dendrobaena octaedra octaedra (Savigny, 1826) Fundorte: Nr. 9/7. G/1802 6 Ex., Nr. 26. G/1914 2 Ex., Nr. 34. G/1619 1 Ex., Z/9553 2 Ex., Nr. 36. G/1652 2 Ex., Nr. 37. G/1692 1 Ex., Z/9600 2 Ex., Nr. 56/1. Z/12554 2 Ex., Nr. 57. MIDAS Sale xe. NL,6022112502,2 Exe, Nr 62, ZU2SL] IFEx., Nr, 69, ZU 233, EX Nr 70/4 2125433 Ex., Nr 74. 2712562 I Fx., Nr. 77. 2/1257645+5 Ex. Dendrobaena octaedra quadrivesiculata Pop, 1938 Fundorte: Nr. 49. G/1944 10.Ex., Nr. 56/1. Z/12554 2 Ex., Nr. 57. Z/12493 1 Ex., Nr. 38.82412500,2 Ex. Nr. 60. 212502 2 Ex Nr. 627 21251 1 EX NII632 BP IB 2 Exe NG 69 Z/12537 2 Ex., Nr. 70. Z/12543 1 Ex., Nr. 77. Z/12576 45 Ex. Diese bisher nur aus Siebenbürgen, Moldawien und Muntenien (Rumänien ) aus höheren Gebirgslagen gemeldete Unterart (Pop 1938) wurde von BOUCHE (1972) in verschiedenen Teilen Frankreichs gesammelt. Wir haben sie nur in den Pyrenäen gesammelt. Sie unterscheidet sich eindeutig durch die 4 Paar Samensäcke von der Stammform. Dendrobaena cognettii (Michaelsen, 1903) Fundort: Nr. 74. Z/12561 5 Ex. Bei den von uns angetroffenen Exemplaren liegt der Giirtel vom 33.-37. Seg- ment, wie dies bei den meisten von uns früher erbeuteten Tieren der Fall ist (ZICSI 1981). Satchellius Gates, 1975 Satchellius Gates, 1975a: 4, EASTON 1983: 484, Sims & GERARD 1985: 118, Mrsic 1991: 667. Typusart: Enterion mammale Savigny, 1826. Weitere Arten: Satchellius madeirensis (Michaelsen, 1891), S. alvaradoi Moreno, 1982. Diese Gattung umfasst rotpigmentierte Arten mit weitgepaarten Borsten und Kalkdrüsen im 10. Segment sowie würstchenförmigen Nephridialblasen und gefiederter Muskulatur. Sie ist innerhalb der Familie Lumbricidae gut abgegrenzt. Satchellius mammalis (Savigny, 1826) Fundorte: Nr. 25. G/1419 5 Ex., Z/9344 5 Ex., Nr. 33. G/1639 50 Ex., Z/9534 50 Ex., 11933333 EX Nr. 34. Gills 3 Exe Z/9550!5 Exe Nro7 GS NS IE ISIS IA Xe Nie 42. G/2032 6 Ex., Z/9810 9 Ex. Aporrectodea Orley, 1885 Aporrectodea Orley, 1885: 22. Allolobophopra Pop, 1941:20. Allolobophora (Allolobophora), OMODEO 1956: 180, Zıcsı 1982: 444. Nicodrilus (part.), BOUCHE 1972: 315, PEREL 1979: 206. Aporrectodea, GATES 1975a: 4, SIMS & GERARD 1985: 53, Mrsic 1991: 27. Typusart: Aporrectodea trapezoides Ant. Duges, 1928. 996 ANDRAS ZICSI & CSABA CSUZDI Die von Gates wieder ins Leben gerufene Gattung Orleys hat das Problem der Sammelgattung Allolobophora nicht gelöst, es werden ihr Arten mit J-bis U-förmigen Nephridialblasen mit nach hinten gerichtetem Ausführungsgang zugeordnet, ohne Rücksicht auf andere, grundlegende Merkmale wie Typ der Muskulatur und Lage der Kalkdrüsen. Wir reihen einstweilen nur die bislang zur Gattung Allolobophora gestellten Arten zu Aporrectodea, die wie die Typusart gefiederte Muskulatur besitzen und bei denen die Nephridialblasen J-U-förmig und nach hinten gerichtet sind.. Aporrectodea caliginosa caliginosa (Savigny, 1826) Fundorte: Nr. 1/2. G/1103 3 Ex., Nr. 1/3. G/988 2 Ex., Nr. 1/5. G/1452 2 Ex., Nr. 1/6. G/1467 4 Ex., Nr. 1/7. G/1448 1 Ex., Nr. 1/9. G/1649 5 Ex., G/1650 4 Ex., Nr. 6. G/1442 2 Ex., Ne27EENB5I2IB EX; G/1593 8Ex 77/9508, 9 Ex., Z/9509 3 Ex. Nr 12 IGNEOTBEOHER® G/1614 2 Ex., Z/ 95233 Ex., Nr: 7/3. G/1767 13 Ex., G/177210Ex,G/1778101ExSZ09 64430] Bx 2/9647, 107, Ex., Nr. 7/4. G/1599.7 Ex., G/1600 20 Ex Z/I61 5, 9EX& ZOO aii Sexe Nie LO! G/2010 2 Ex., Nr. 8/1. G/996 6 Ex., G/997 7 Ex., Nr. 8/2: G/1460 21 Ex., ZI32SSAEX INT 91G/1415 5 Ex., Nr. 9/2. G/1427 1 Ex., Nr. 9/3.,G/1112 7 Ex., Nr) 9/4.'GAA SEIREXKANTSIIBN G/1423’ A Ex., Nr: 9/7. G/1777 3 Ex., Nr. 10. G/1674 1 Ex.) Z/9565 1FEx., Nr21137@2029 1Ex., Nr. 16/1. G/1099 1 Ex., Nr. 19. G/980 2 Ex., Nr. 21/1. G/1664 1 Ex., Z/9559 2 Ex., Nr. 21/2. G/1671 1 Ex., Nr. 21/3. G/1655 1 Ex., G/1657 2 Ex., Nr. 23. G/987 2 Ex., N. 24. G/989 1 Exe, Nr. 25. G/1420'2 Ex., 2/9342 7 Ex., Z/9343 1 Ex. Nr. 26. G/1918 3 EX NLZ2AGH 82 Exe) Nr 33.27@/1641 3° Ex: -Z/9530"2 Ex:.; Nr. 38 G/2355 7’ EXS NT. 39/23 Gi23843e2a xe. Z/10770 6 Ex., Nr. 41. G/2165 4 Ex., Z/10655 3 Ex., Nr. 42. G/2038 2 Ex., Z/9812 15 Ex., Nr. 43..G/1853 1 Ex., Nr. 44. G/2055 4 Ex., Z/9826 1 Ex., Z/ 9827 13\Ex. Nr. 492G/20602 FE 7.198337 2 Ex., Nr. 47. G/1939 3 Ex, Z/ 9773 43 ex. Z/9771 2 EX Nr S33G 2007 axe ZI9849 3 Ex., Nr. 54. G/1937 3) Ex; Z/9765-4.Ex.; Nr. 55. G/1942 1 Ex. Z/9776 1x 50: G/2061 7 Ex., Z/9837 10 Ex., Nr. 57. Z/12483 14 Ex., Nr. 63. Z/12516 1 Ex., Nr. 66. Z/12520 la Bxe Nr 68: 212530 2 Ex, Ne 70/1,Z/12541 4 Ex. Nr 76. Z/12575 Vex Aporrectodea caliginosa trapezoides (Ant. Duges, 1828) Fundorte: Nr. 41. G/2166 2Ex., Z/10656 2Ex., Nr. 43. G/1854 1 Ex., Nr. 56. G/2062 2 BxeiZ/9836)2 Exe Nr 58: Z/12498 Exe Aporrectodea longa (Ude, 1885) Fundorte: Nr. 1/6. G/1464 3 Ex., Nr. 1/9. G/1651 1 Ex., Nr. 9/1. G/1414 1 Ex., Nr. 9/3. G/1113 2 Ex., Nr. 9/4. G/1117 3 Ex., Nr. 1661. G/1100 1 Ex., Z/9340 2 Ex., Nr. 19. G/978 1 Ex., Nr. 42. G/2039 2 Ex., Z/9813 4 Ex. Aporrectodea terrestris (Savigny, 1826) Fundorte: Nr. 7/3. G/1771 35 Ex., Z/9640 5 Ex., Z/9641 5 Ex., Z/9642 5 Ex., Nr. 21/1. G/1663 2 Ex., Z/9560 1 Ex., Nr. 31. G/1801 2 Ex., Nr. 44. G/2054 3 Ex., Z/9825 2 Ex. Aporrectodea cupulifera Tétry, 1937 Fundorte: Nr. 21/1. G/1662 1 Ex., Nr. 33. G/1640 4 Ex., Nr. 35. G/1576 15 ex., G/1577 IOEX 2/9537, 26, EX Ne 4156/2160 S1Exe94/ W065) DilaExs REGENWURMFAUNA FRANKREICHS 997 Aporrectodea haymozi Zicsi, 1977 Fundorte: Nr. 58. G/2557 1+1 juv. Ex., Z/12496 4+1 Ex., Nr. 60. 12501 2+1 Ex., Nr. 70/1. 12540 7 Ex. Die Uberpriifung des Typenmaterials zeigte, dass die Lage der Samentaschen in den Originalbeschreibung falsch angefiihrt wurde: sie legen im 12., 13. und 14. Segmnet und miinden auf Intersegmentalfurche 11/12, 12/13 und 13/14 aus. Die Nephridialblasen sind fast U-förmig und nach hinten gerichtet. Bei den jetzt erbeu- teten Tieren erstreckt sich der Gürtel vom 33.-1/2 40., 40. Segment, die Pubertäts- streifen liegen am 35., 36. - 40. Aporrectodea haymoziformis sp. n. Fundorte: Holotypus. Z/12949. Dep. Pyrénées-Atlantiques. Camou-Cihigue, 14. 9. 1982. leg. A. Zicsi. Paratypen. Z/12950 1Ex., Z/9845 3 Ex., G/2066 1 +1 juv. Ex. Diagnose : L. 49-56 mm, B. 3,5-42 mm, Segmentzahl 102-117. Unpigmentiert, weiss. Kopf prolobisch. Borsten eng gepaart. Gürtel vom 1/2 33., 33., 34.-40., 1/2 41. Segment. Dissepimente 6/7-11/12 verdickt. Herzen im 6.-11., Kalkdrüsen im 10. Segment. Hoden und Samentrichter frei. 4 Paar Samensäcke im 9.-12. Segment. Typhlosolis einfach gelappt. Nephridialblasen U-förmig. Muskulatur gefiedert. 4 Paar Samentaschen im 11., 12., 13., 14. Segment. Länge des Holotypus 52 mm, Breite 4 mm, Segmentzahl 109. Paratypen. Länge 49-56 mm, Breite 3,5-42 mm, Segmentzahl 102-117. Farbe weiss. Kopf prolobisch. 1. Segment gefurcht, die übrigen ungeringelt. Borsten eng gepaart, Borstenverhältnis hinter dem Gürtel aa:ab:bc:cd:dd wie 6:1:3:1:9. Borsten ab und cd auf dem 11. Segment, cd auf dem 12., 13., 14. Segment, sowie ab auf dem 31. und 32. Segment von kleinen Papillen umgeben. Die Borsten ab entlang der Pubertätsstreifen sind ebenfalls von kleinen Papillen umgeben. Erster Rückenporus auf Intersegmentalfurche 9/10. Nephridialporen in der Borstenlinie cd, nicht alternierend. 4 Paar Samentaschenporen auf Intersegmentalfurche 10/11, 11/12, 12/13 und 13/14. Gürtel sattelförmig vom 1/2 33., 33., 34.-40., 1/2 41., Segment. Pubertätsstreifen 1/2 34., 34. -40., 1/2 41. Segment. Innere Organisation. Dissepimente 6/7-9/10 stark, 10/11-11/12 weniger stark verdickt. Schlund bis ins 6. Segment reichend. Herzen im 6.-11, Segment. Kalkdrüsen nur kleine Ausbuchtungen im 10. Segment. Hoden und Samentrichter im 10. und 11. Segment frei. 4 Paar Samensäcke 1m 9.-12. Segment, die im 9. und 10. klein, im 11. und. 12. Segment grösser und gelappt. Ovarien im 13. Segment, Ovarientrichter auf Dissepiment 13/14. Nephridien hinter dem Gürtel U-förmig nach hinten gebogen. Kropf im 15.-16. Segment, Muskelmagen im 17.-18. Segment, Typhlosolis einfach gelappt. Muskulatur gefiedert. Samentaschen kleine runde Gebilde ohne Stiel im 11., 12., 13. und 14. Seg- ment, mit Samenmassen prall gefüllt. Die neue Art steht A. haymozi am nächsten. Unterscheidet sich von ihr durch die Lage der Pubertätsstreifen und die höhere Zahl der Samentaschen. 998 ANDRAS ZICSI & CSABA CSUZDI Aporrectodea riparia (Bretcher, 1901) Fundort: Nr. 44. G/2056 1 Ex. Die Art ist neu fiir die Fauna Frankreichs. Aporrectodea icterica (Savigny, 1826) Fundorte: Nr. 1/9. G/1647 1° Ex.) Nr. 7/1. GAS88 20 Ex: Z/9512) iE xe Nica G/1611 5 Ex., Nr. 7/3. G/1763 16 Ex., Z/9643 16 Ex., Nr. 7/4. G/1602 16 Ex. Z/9521F Exe; Nr. 7/6. G/2008 1 Ex., Z/9807 2 Ex., Nr. 9/7. G/1776 1 Ex. Aporrectodea rosea (Savigny, 1826) Fundorte. Nr. 1/6. G/1466 1 Ex., Nr. 1/7. G/1446 1 Ex., Nr. 1/9. G/1648 1 Ex., Nr. 7/1. G/1586 13 Ex., Nr. 7/2. G/1616 6 Ex., Nr. 7/3. G/1765 4 Ex., Z/9645 3 Ex., Nr. 7/4. G/1604 2 Ex., Nr. 7/6: G/2009' 1 Ex., Nr. 8/2. G/14594 Ex., Z/9324 3Ex., Nr. 93. GI III Ex Nr 0/7 GNA V Ex., Nr. SA G/1122 1 Ex., Nr. 21/1.°G/1660) 2 Ex.” Nr-21237 GN SSCHEER ENT 21/4. G/1790 2 Ex., Nr. 43. G/1848 1 Ex., Nr. 56/1. Z/12553 3 Ex., Nr. 57. Z/12485 1 Ex., Nr. 388 2AITZIER., Nr. 7522/12565. 6 Ex. Nr. 76. 712574 Ex Nie i 2257 RE Allolobophora Eisen, 1874 Typusart: Enterion chloroticum Savigny, 1826 Allolobophora cholorotica chlorotica (Savigny, 1826) Fundorte: Nr. 1/1. G/1471 1 Ex., Nr. 1/2. G/1102 5 Ex., Z/9337 6 Ex., Nr. 1/5.G/1451 2 Ex., Nr; 1/6, G/1471 1) Ex., Nr: 1/7. G/1447 3 Ex., Nr: 1/9. G/1645 10/exs Nr=17104G@167928 EX Nr 6. G/1441 Ex. Nr 7/1.)G/1587 12 Ex: 2/9513 9. Ex. Nr T/2AG/0ISMOIES Z/9524 3 Ex., Nr. 7/3. G/1764 10 Ex., G/1766 1 Ex., Z/9646 6 Ex., Nr. 7/4. G/1603 10 Ex., Z/9519 12 Ex., Nr. 7/6. G/2007 4 Ex., Nr. 8/1. G/993 2 Ex., Nr. 8/2. G/1461 6 Ex., Z/9323 4 Ex., Nr. 9/1. G/1413 2 Ex., Nr. 9/3. G/1110 2 Ex., Nr. 9/4. G/1114 4 Ex., Nr. 10. G/1673 1 Ex., Z/9564 5 Ex., Nr. 11/2. G/1887 1 Ex., Nr. 11/3. G/2030 1 Ex., Nr. 13. G/964 1 Ex., Nr. 14. GTA Ex. Nr. 19. G/979 9 Ex., Nr. 20. G/18803 Ex., Nr..21/1. G/1661 Ex 2727/95372 E53 Nr 21/42. G/1791 1 Ex., Nr. 24. G/990 1 Ex., Nr. 25. G/1421:6 Ex, Z/93416RExSSNk89)18 G/1800 4 Ex., Nr. 32 vn 1 Ex., Nr. 33. G/1629 1 Ex., G/1638 16 Ex., Z/9532 14 Ex., Nr. 34.6/1620 2 Ex., Nr 35 G/1583 10. Ex., Z/9541 29 Ex., Nr 38" G/2354 Ex NEw oe G/2349 2 Ex., Z/1077 1 Be Nr. 41. G/2164 7 Ex., Z/10654 4 Ex., Nr. 42. G/2035 6 Ex., Z/9816 12 Ex., Nr. 43. G/1849 3 Ex., Nr. 44. G/2051 2 Ex., Z/9819 3 Ex., Nr. 47. G/1938 7 Bx, 2/9770) TEx? Nr 48..G/1946 1 Ex., 7/9784 5 Ex., Nr. 55. G/1941 1 Ex ZINN 56. G/2063 1 Ex., Z/10654 4 Ex., Nr. 56/1. 12555 1 Ex. Allolobophora chlorotica postepheba Bouché, 1972 Fundorte: Nr. 20. G/1881 8 Ex., Nr. 35. G/1585 1 Ex., Nr. 43. G/1850 8 Ex. Allolobophoridella Mrsic, 1990 Lumbricus (part.), LEVINSEN 1884: 241. Dendrobaena (part.), OMODEO 1956: 156. Bimastus (part.), OMODEO 1956: 178. Eisenia (part), BOUCHE 1972: 378, Zıcsı 1982: 428. Allolobophora (part.), PEREL 1979: 186, EASTON 1983: 475, Zıcsı 1994: 69. Allolobophoridella Mrsic, 1990: 252. Typusart: Lumbricus eiseni Levinsen, 1884. REGENWURMFAUNA FRANKREICHS 999 Allolobophoridella eiseni (Levinsen, 1884) Fundorte: Nr. 4/2. G/984 1 Ex., Nr. 44. G/2058 1 Ex., Z/9820 3 Ex., Nr. 57. Z/12515 4 Ex., Nr. 63. Z/12484 2 Ex. Die zeitweise in 5 verschiedene Gattungen gestellte Art zeigt die Unsicher- heit, mit der einige Arten der Familie Lumbricidae behandelt wurden. Wir schliessen uns der von MRSIC (1991) vorgeschlagenen Gattungsdefinition (rotpigmentierte Arten mit Nephridien, deren Ausführungsgang nach vorne gerichtet ist) an, mit der Bemerkung, dass allein die Typusart, die über gefiederten Muskeltyp verfügt, in dieser Gattung bekannt ist. Zur zweiten Art von Mrsic mit bündelartigem Muskeltyp (A. parva Eisen, 1874) können wir vorerst nicht Stellung nehmen. Eisenia Malm, 1877 Typusart: Enterion fetidum Savıgny, 1826. Eisenia fetida (Savigny, 1826) Fundorte: Nr. 1/2. G/1104 1 Ex., Nr. 2. G/1454 21 Ex., Nr. 26. G/1917 7 Ex., Nr. 27. G/2028 1 Ex., Nr. 33. G/1595 10 Ex., G/1635 1 Ex., Z/9544 10 Ex., Nr. 34. G/1622 15 Ex., Nr. 36. G/1653 1 Ex., Nr. 42. G/2036 6 Ex., Z/9815 10 Ex., Nr. 68. Z/12529 1+2 Ex. Der Artenname fetidum wurde von MICHAELSEN (1900) in foetidum emendiert und Jahrzehnte hindurch so benützt. Da dies die Regeln der Nomenklatur nicht zulassen, muss die alte Schreibweise wieder eingeführt werden. Eisenia lucens (Waga, 1857) Fundort: Nr. 63. Z/12512 3 Ex. Eiseniella Michaelsen, 1900 Eiseniella tetraedra tetraedra (Savigny, 1826) Fundorte: Nr. 5. G/1680 1 Ex., Nr. 11/2. G/1886 25 Ex., Nr. 16/1. G/1098 2 Ex., Nr. 33. G/1631 5 Ex., G/1637 10 Ex., Z/9533 1 Ex., Z/9548 1 Ex., Nr. 34. G/1624 9 Ex., Z/9551 1 Ex., Nr. 35. G/1578 15 Ex., Z/9536 50 Ex., Nr. 41. G/2163 10 Ex., Z/10653 8 Ex., Nr. 57. 12488 5 Ex., Nr. 58. 12499 1 Ex., Nr. 61. 12506 1 Ex., Nr. 68. 12528 7 Ex., Nr. 72. 12527 1 Ex. Eiseniella tetraedra hercynia (Michaelsen, 1890) Fundort: Nr. 33. G/1643 1 Ex. Octolasion Örley, 1885 Octolasion cyaneum (Savigny, 1826) Fundorte: Nr. 1/2. G/1107 1 Ex., Z/9335 1 Ex., Nr. 1/6. G/1468 1 Ex., Nr. 1/7. G/1443 DEAN 946/1604601 Ex, Nr.,8/22@1455 53, Ex. Z/932 1 EX, Nr. 9/7 \G/1780, BER INT. BAARGOTOMME ENT 0AG/ISSSM EX CANTA 34/2 59009 REX NL 39/2 G2 35) 2s ERS, Z/10773 1 Ex., Nr. 41. G/2159 2 Ex., Nr. 42. G/2034 4 Ex., Z/9811 6 Ex., Nr. 43. G/1851 1 Ex., Nr. 44. G/2050 1 Ex. 1000 ANDRAS ZICSI & CSABA CSUZDI Octolasion lacteum (Orley, 1881) Fundorte: Nr. 11/2. G/1888 1 Ex., Nr. 20. G/1882 3 Ex., Nr. 23. G/986 1 Ex., Nr. 33. G/1628 1 Ex., G/1642 1 Ex., Z/9527 1 Ex., Nr. 35. G/1584 4 Ex., Z/9543 3 Ex., Nr. 43. G/1852 5 Ex., Nr. 44. G/2049 1 Ex., Z/9817 1 Ex., Nr. 56. G/2064 1 Ex., Z/9838 1 Ex., Nr. 56/1. 71125932 1S)Ex., Nr. 37. Z/12486 3’Ex., Nr. 60. 2/12503 MEX Nr. 63. 212 S A BERN EOS ZMN2SSI2HEPEX:, Nr. 6972112536 2 Ex:, Nr. 74. Z/12563 3 Ex., Nr 75. Z/12567) Sx. Orodrilus Bouché, 1972 Orodrilus Bouché, 1972: 219, EASTON 1982: 484 Typusart: Allolobophora doderoi Cognetti, 1904. Diese auf Meroandrie der männlichen Geschlechtsorgane aufgestellte Gattung, lässt sich durch die eng bis weitläufig gepaarten Borsten, durch den gefiederten Muskeltyp und die nach hinten gerichteten J-förmigen Nephridialblasen von den übrigen Gattungen unterscheiden. Für die meroandrischen Arten der Gattungen Octodrilus und Octodriloides sind keine gesonderten Gattung aufgestellt worden, da hier andere Begrenzungsmerkmale, wie die Lage der männlichen Poren, ungepaarte Borsten und die Zahl der Samentaschen, ausschlaggebend waren. Wir schliessen uns der Meinung von Bouché an und betrachten die meroandrischen Arten, mit den weiter oben angeführten Merkmalen, als dieser Gattung angehörend. Orodrilus gavarnicus (Cognetti, 1904) Fundort: Nr. 69. Z/12948 1 Ex. Prosellodrilus Bouché, 1972 Prosellodrilus Bouché, 1972: 231, EASTON 1983: 484, Mrsıc 1991: 662. Typusart: Prosellodrilus idealis Bouché, 1972. Durch den stark nach vorne gertickten Giirtel (am 19., 20. Segment be- ginnend), die eng gepaarten Borsten, die fehlende Pigmentation, die hinter dem Giirtel doppellappigen Nephridialblasen und der bündelförmigen Muskulatur sowie durch die doppeltgelappte Typhlosolis unterscheidet sie sich von allen übrigen Gattungen. Prosellodrilus pyrenaicus (Cognetti, 1904) Fundort: Nr. 42. G/2033 1 Ex. Prosellodrilus albus sp. n. Fundort. Dept. Haute-Garonne, Holotypus. Z/13003 Audressein (42° 55' 34" N: 1° O1' 1" O) 500 m, 27. 6. 1997 leg. Zicsi-Csuzdi-DFK. Paratypen, Z/12569 3+5 praed.+1 juv. Ex.- G/2662 2+ 1 praed, Ex. Fundort wie beim Holotypus. Diagnose: L. 78-83 mm, B. 3,2-4 mm, Segmentzahl 134-184. Unpigmentiert, weiss. Kopf prolobisch. Borsten eng gepaart. Gürtel vom 20.-27., Pubertätsstreifen vom 20.-1/4 25. Segment. Dissepimente 5/6-8/9 verdickt. Herzen im 6.-11., Kalkdrüsen im 11. Segment. Perioesophageale Testikelblasen im 10. und 11. Seg- REGENWURMFAUNA FRANKREICHS 1001 ment. 2 Paar Samensäcke im 11. und 12. Segment. Typhlosolis doppelt gelappt. Nephridialblasen doppellappig. Muskulatur bündelförmig. 2 Paar Samentaschen im 14. und 15. Segment. Länge des Holotypus 83 mm, Breite 4 mm, Segmentzahl 184. Paratypen. Länge 78-81 mm, Breite 3,2-3,8 mm, Segmentzahl 134-178. Farbe weiss, unpigmnetiert. Kopf prolobisch. 1. Segment längsgefurcht, vom 7. doppelt und mehrfach gefurcht. Borsten eng gepaart. Borstenverhältnis hinter dem Gürtel aa:ab:bc:cd:dd wie 20:1:5:1:30. Borsten ab des 12., 14., 16. sowie die des 25.- 33. Segments von winzigen Papillen umgeben. Erster Rückenporus auf Interseg- mentalfurche 4/5. Nephridialporen oberhalb der Borstenlinie b, nicht alternierend. Samentaschenporen auf Intersegmentalfurche 13/14 und 14/15 unterhalb der Borsten- linie cd. Weibliche Poren auf dem 14. Segment, männliche Poren auf dem 15. Seg- ment, von einem kleinen Hof umgeben, der nicht auf das 16. Segment tibergeht. Gürtel sattelförmig auf dem 20.-27. Segment, Pubertätsstreifen vom 20.-1/4 25. Segment. Innere Organisation. Dissepimente 5/6-8/9 sehr stark, 9/10 nur schwach verdickt. Schlund bis ins 5. Segment reichend. Herzen im 6.-11. Segment. Kalkdrüsen im 11. Segment. Hoden und Samentrichter im 10. und 11. Segment von perieoso- phagealen Testikelblasen umgeben. Ausführungsgang der Samenrinne nicht gewun- den. 2 Paar Samensäcke im 11. und 12. Segment. Ovarien im 13. Segment. Ova- rientrichter auf Dissepiment 13/14. Kropf im 15.-16. Segment, Muskelmagen im 17- 19. Segment. Mitteldarm im 20. Segment mit einer doppeltgelappten Typhlosolis versehen. Nephridien hinter dem Gürtel doppellappig. Muskulatur bündelförmig. Samentaschen 2 Paar im 14. und 15. Segment, kleine gebogene, birnförmige Ampulle mit kurzem Ausführungsgang. Der Ausführungen der Samentaschen dicht von Drüsen umgeben. Die neue Art steht P. pyrenaicus am nächsten. Sie unterscheidet sich von ihr durch die Lage des Gürtels und der Pubertätsstreifen. Prosellodrilus fragilis fragilis Bouché, 1972 Fundort: Nr. 69. Z/12533-34 38 juv. Ex. Keines der zahlreichen Tiere besass einen entwickelten Giirtel, allein die Pubertätsstreifen liessen sich erkennen, aufgrund deren Lage ist das Material mit Vorbehalten zu fragilis fragilis gestellt worden. DANKSAGUNG Für die Ermöglichung der Sammelreise in die Pyrenäen (1997) sowie für einen Arbeitsplatz im Naturhistorischen Museum von Genf, wo auch ein Teil des neueren Materials bearbeitet wurde, sprechen wir der Direktion des Museums und Herrn Dr. Cl. Vaucher unseren besten Dank aus. 1002 ANDRAS ZICSI & CSABA CSUZDI LITERATUR BoucHE, M. B. 1970. Remarques sur quelques Lumbricina de France et conséquences de la découverte des nouveaux taxons Vignysinae (Subfam. nov.) et Diporodrilidae (Fam. nov.). Pedobiologia 10:246-256. BoucHé, M. B. 1972. Lombriciens de France. Ecologie et systématique. Institut National de la Recherche Agronomique, Articles de Zoologie-Ecologie animale (Numéro hors-série). 671 pp. Diaz Cosin, D., TRIGO, D. & MASCATO, R. 1992. Earthworms of the Iberian Peninsula. Species list and some biographical considerations. Soil Biology and Biochemistry 24:1351- 1356. Easton, E. G. 1983. A guide to the valid names of Lumbricidae (Oligochaeta) (pp. 475-485). In: SATCHELL, J. E. (ed.). Earthworm ecology from Darwin to vermiculture. Chapman and Hall, London. Gates, G. E. 1975a. Contributions to a revision of the earthworm family Lumbricidae. XII. Enterion mammale Savigny, 1826 and its position in the family. Megadrilogica 2(1): 1-8. Gates, G. E. 1975b. Contributions to a revision of the earthworm family Lumbricidae. XVII. Allolobophora minuscula Rosa, 1906 and Enterion pygmaeum Savigny, 1826. Mega- drilogica 2(6): 7-8. GATES, G. E. 1978. The earthworm genus Lumbricus in North America. Megadrilogica 3(6): 81-116. GERARD, B. M. 1964. A synopsis of the British Lumbricidae Synopses of the British Fauna No. 6. Lumbricidae (Annelida). The Linnean Society of London, 58 pp. HOFFMEISTER, W. 1845. Ubersicht aller bis jetzt bekannten Arten aus der Familie der Regenwürmer. Friedrich Vieweg und Sohn, Braunschweig. 43 pp. LEVINSEN, G. M. R. 1884. Systematik-geografisk oversigt over de nordiske Annulata, Gephy- rea, Chaetognathi, og Balanoglossi. Videnskabelige Meddelelser fra Dansk Natur- historisk Forening i Kjobenhavn, 1883: 92-350. MICHAELSEN, W. 1900. Oligochaeta. Das Tierreich 10: 1-575. Mrsic, N. 1990. Description of a new subgenus, three new species and taxonomic problems of the genus Allolobophora sensu Mrsic and Sapkarev, 1988 (Lumbricidae, Oligochaeta). Bioloski Vestnik, Ljubljana 38(1): 49-68. Mrsic, N. 1991. Monografija o dezevnikih (Lumbricidae) Balkana I-II. Academia Scientarium et Artium Slovenica, Classis IV: Historia Naturalis, Opera 31: 1-757. MULDAL, S. 1952. A new species of earthworms of the genus Allolobophora. Proceedings of the Zoological Society, London 122: 463-465. MURCHIE, W. R. 1959. Redescription of Allolobophora muldali Omodeo, 1956 (Lumbricidae: Oligochaeta). The Ohio Journal of Science 59(6): 329-332. OMODEO, P. 1953. Nota su alcuni lombrichi delle Alpi e considerationi sul Gen. Helodrilus di Hoffmeister. Memorie del Museo Civico di Storia Naturale di Verona 4: 75-85. OMODEO, P. 1956. Contributo alla revisione dei Lumbricidae. Archivio Zoologico Italiano 41: 129-143. OMODEO, P. 1988. The genus Eophila (Lumbricidae, Oligochaeta). Bolletino di Zoologia Napoli 55: 73-84. OMODEO, P. & Rota, E. 1989. Earthworms of Turkey. Bollettino di Zoologia Napoli 56: 167- 1993 OrLEY, L. 1885. A palearktikus övben é16 Terricoläknak revizidja és elterjedése. Ertekezesek a termeszettudomanyok Köreböl, Budapest 15: 1-34. PEREL, T. S. 1976. A critical analysis of the Lumbricidae genera system (with key to the USSR fauna genera). Revue d'Ecologie et de Biologie du Sol 13(4): 635-643. REGENWURMFAUNA FRANKREICHS 1003 PEREL, T. S. 1979. Range and regularities in the distribution of earthworms of the USSR fauna (with keys to the Lumbricidae and other Megadrili). Nauka, Moskow. 272 pp. PICKFORD, G. E. 1926. On a new species of earthworm belonging to the subgenus Bimastus from Wicken Fen. The Annals and Magazine of Natural History London 17: 96-98. Pop, V. 1938. Neue Lumbriciden aus Rumänien. Buletinul Societatii de Stiinte din Cluj 9: 134- 152: Pop, V. 1941. Zur Phylogenie und Systematik der Lumbriciden. Zoologische Jahrbücher 74 (5-6): 487-522. Rosa, D. 1906. L'Allolobophora minuscula n. sp. Atti della Società dei Naturalisti e Matematici di Modena (4)7: 38-39. Sims, R. & GERARD, B. M. 1985. Earthworms Keys and notes for the identification and study of the species. Synopses of Brtitish Fauna 31: 171 pp. Trico, D., MASCATO, R., IGLESIAS BRIONES, M. J. & DIAZ-Cosin, D. J. 1990. Lombrices de tierra de Portugal Continental. Inventario y Cytas. Arquivos do museu Bocage, Nov. Sema 21-567: ZICSI, A. 1968. Neuere Angaben zur Kenntnis der Lumbricidenfauna Jugoslawiens. Annales Universitatis Scientiarum Budapestiensis de Rolando Eötvös Nominatae, Sectio Biologica, Budapest 9-10: 401-405. ZICSI, A. 1973. Regenwürmer (Oligochaeta: Lumbricidae) aus Griechenland. Opuscula Zoolo- gica, Budapest 12: 99-103. Zicsi, A. 1977a. Wiederbeschreibung zweier Arten aus der Familie Lumbricidae (Oligochaeta). Opuscula Zoologica, Budapest 13: 107-110. Zıcsı, A. 1977b. Neue Regenwürmer (Oligochaeta: Lumbricidae) aus den Atlantischen Pyre- näen. Revue suisse de Zoologie 84: 681-685. Zıcsı, A. 1981. Weitere Angaben zur Lumbricidenfauna Italiens (Oligochaeta: Lumbricidae). Opuscula Zoologica, Budapest 17-18: 157-180. ZICSI, A. 1982. Verzeichnis der bis 1971 beschriebenen und revidierten Taxa der Familie Lumbricidae (Oligochaeta). Acta Zoologica Hungarica 28: 421-454. Zicsi, A. 1985. Uber die Gattungen Helodrilus Hoffmeister, 1845 und Proctodrilus gen. n. (Oligochaeta: Lumbricidae). Acta Zoologica Hungarica 31: 275-289. Zıcsı, A. 1986. Über die taxonomischen Probleme der Gattung Octodrilus Omodeo, 1956 und Octodriloides gen. n. (Oligochaeta: Lumbricidae). Opuscula Zoologica, Budapest 22: 103-112. Zicsi, A. 1994. Die Regenwürmer Österreichs (Oligochaeta: Lumbricidae) mit Bestimmungs- tabellen der Arten. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Oster- reich 131: 37-74. Zicsi, A. & MICHALIS, K. 1981. Übersicht der Regenwurm-Fauna Griechenlands (Oligochaeta: Lumbricidae). Acta Zoologica Hungarica 27: 239-264. ” 4 i È Ra | E x L se: SPOT ke The (th ce u HE ét pi: 0 ‘np 1 4¥ ad à Lab de | Pre REVUE SUISSE DE ZOOLOGIE 106 (4): 1005-1012; décembre 1999 A review of the Himalayan Thorectes (Coleoptera: Geotrupidae), with description of a new species from northern India Giuseppe M. CARPANETO Universita degli Studi “Roma Tre”, Dipartimento di Biologia, Viale G. Marconi, 446, I-00146 Roma, Italia. Roberto MIGNANI Universita degli Studi “Roma Tre”, Dipartimento di Fisica “E. Amaldi”, Via della Vasca Navale, 84, I-00146 Roma, Italia. A review of the Himalayan Thorectes (Coleoptera: Geotrupidae), with description of a new species from northern India. - Thorectes shankara sp. n. from West Himalayas is described and illustrated. A checklist of the Himalayan species of Thorectes is given, including detailed records. A key to these species is provided and their diagnostic characters are summarized. Metathoracic wings are strongly but not completely reduced in these species. The wing reduction in Thorectes is discussed. Key-words: Coleoptera - Geotrupidae - Himalayas - India - wing size - taxonomy. INTRODUCTION In the framework of a worldwide generic revision of Geotrupinae, ZUNINO (1984) raised Thorectes Mulsant, 1842 from subgenus of Geotrupes Latreille, 1796 to genus, with 36 species included. The genus has discontinuous distribution, with 32 Mediterranean species, one (7. banghaasi Reitter, 1893) endemic in Gissar and Darvaz Mountains in Tajikistan (NIKOLAJEV 1987), and three from Nepal Himalayas (BARAUD 1974, KRIKKEN 1981). The Nepalese species are: T. nepalensis Baraud, 1974, T. martensi Krikken, 1981 and T. stellosus Krikken, 1981. Two subspecies are recognized in the latter species. New collections from the Indian part of the western Himalayas include an additional species of Thorectes which is described below. Thorectes shankara sp. n. Holotype: 3, India, Uttar Pradesh, Kedarnath 3300 m a.s.l. (14 kms north of Rambara), 26-29 July 1994, M. Snizek leg. Paratypes: 13 6 d and 22 9 9, all from the type locality. Holotype in Zoological Museum of Rome University (MZUR); 5 paratypes in the Carpaneto collection; 2 paratypes (1 d, 1 9) in the “Museum d’histoire naturelle de Genève”; 2 paratypes (1 d, 1 9) in the “Forschungsinstitut Senckenberg, Frankfurt am Main”; remaining specimens in the Mignani collection. Manuscript accepted 07.06.1999 1006 G. M. CARPANETO & R. MIGNANI Diagnosis: Allied to T. nepalensis (figs 1, 3, 15) but distinguished by the following characters: (1) third external tooth of protibia directed downwards (fig. 2); (2) 7 teeth, instead of 6, along outer margin of protibia (fig. 2); (3) 8-9 denticles, instead of 7, along ventral face of protibia (fig. 2); (4) pronotum more strongly convex; (5) pronotal surface shining, elytral surface sericeous; (6) elytral striae partially effaced but all moderately distinct; (7) mesosternal apophysis pointed anteriorly (on lateral view); (8) disc of metasternum and of abdominal sternites glabrous and almost impunctate; (9) apex of parameres wider and more rounded, slightly turned downwards (fig. 4). Description of the holotype: Male, length 17.5 mm, greatest width 8.7 mm. Entirely black. Moderately shining but elytra opaque with sericeous aspect. Antennal club brownish black. Clypeus (length 3.7 mm) widely U-shaped. Clypeal surface densely and roughly punctate. Clypeal margin slightly raised. Disc with obsolete central elevation. Eye normally shaped, eye-canthus broadly rounded, with anterolateral angle subdistinct. Clypeo-frontal sulcus V-shaped, fronto-lateral ridge moderately pronounced. Labrum with anterior margin feebly emarginate and sides rounded. Left mandible with shallow apical external emargination. Right mandible with protruding lobe near apical-external emargination. Pronotum (length 5 mm) strongly convex, with maximum width (8.7 mm) at posterior half, entirely marginate, except for posterior edge on both sides of antescu- tellar section of base. Pronotal contours with anterior angle ca. 100°, posterior angle rounded. Pronotal punctation very fine, formed by small and minute punctures (as in T. nepalensis), and widely scattered all over surface. Scutellum broadly triangular (length 1 mm). Elytra convex (maximum length 8.7 mm; maximum width 8.7), humeral angle rounded, with umbone distinct. Elytral striae irregular and moderately distinct. Surface opaque with sericeous aspect due to microreticulation. Wings strongly reduced (micropterous): wing length 2.6 mm, WL/EL (wing length/elytral length) 0.3. Mesosternal apophysis pointed anteriorly (on lateral view); mesosternal cavity deep. Metasternum with very shallowly impressed midline. Disc of metasternum and of abdominal sternites glabrous and almost impunctate. Protibia with 7 teeth along outer margin, apical tooth bifid; third external tooth directed downwards; inferior longitudinal crest with 9 denticles (fig. 2). Metafemur with irregular series of 6-7 denticles, some notched, decreasing in size towards apex (fig. 5). Apex of parameres wide and rounded, slightly turned downwards (in T. nepa- lensis, parameres have apex narrower, more pointed, and distinctly directed upwards) (figs 3-4). Paratype variability: Male length: 15-18.4 mm; male width: 7.7-9.1 mm. Female length: 12.5-18.6 mm; female width: 6.1-9.5 mm. In males, the shape and the number of the denticles on the posterior margin of the metafemur are highly variable (figs 5-12); a certain degree of variability is also observed in the inferior denticulation of protibia. Sexually-dimorphic characters are as those usual in other Himalayan Thorectes species. Females differ by: (1) apical tooth of protibia simple (not bifid); (2) protibia HIMALAYAN THORECTES 1007 (ee) Fics 1-4 Thorectes nepalensis (holotype): 1, underside of protibia; 3, ventral view of parameres. Thorectes shankara sp. n. (holotype): 2, underside of protibia; 4, ventral view of parameres. The arrows in figs 1-2 indicate the third external tooth of tibia. without inferior crest of denticles; (3) metafemur without denticles; (4) pronotum less convex and narrower, with a slightly denser punctation. Ecological notes: According to the collector, Miroslav Snizek, the specimens were captured in pitfall traps (with fish as bait), during several rainy days, on the western slope of a mountain near to Kedarnath. The slope was covered by a mixed forest, slightly damaged by cutting. Etymology: This species is named to honour the Indian philosopher Shankara (dead at Kedarnath about 750 AD). CHECKLIST OF THE HIMALAYAN THORECTES A checklist of the Himalayan species and subspecies of the genus Thorectes is here given together with the material examined and distribution, including the altitudinal range. The main diagnostic characters are compared in Table 1. G. M. CARPANETO & R. MIGNANI 1008 ayeyuepHy nu 950125 ‘ayejound asojas ‘arezound payurod pooeyyo Ajpen.red popyunım ‘onbedo juasaid 6108 910 G poppurim ‘onbedo possoidiur A[qooy} ‘oTIe] smruindeuue ‘S'IL SEJUSPHINW as0]9s ‘ayejound as0]9s ‘aezound paurod Po9oPJJo popyunım ‘onbedo juasoid 6108 iE popyurim ‘onbedo possaidiur [[aM ‘9318] ‘MS ‘S SNSO[[9IS € L ayequaphynu 9s0]9s ‘aezound dsojas ‘arezound padeys-mold poorjja AUIUS Juosge L 9 AUIYS our ‘asieds sisuajedou * |, ayejuapHy nu snoge]3 ‘ayeJoundut snoiqe[s ‘aezoundui paurod pootzjo Apenzed SNODIIIIS juasqr 6108 L X9AUO9 A[SUONS ‘AUIYS auly ‘asieds BIBYUEYS ‘], possaldWi [[9M ‘osuop ojejuapiun snoIge]$ ‘aejound INUIDJRIOU EIN SONUI9]S [eurwopge JO ISI(q snoiqr|s ‘ayejyound A]3UONS SIP JEUINISEION podeys-moid sısAydode [eu1sIsosaW PUnsIp aes [enA[H AUIUS 29PJINS [PHAIA juasqe viqnoid few JO 4100) [eMuoA ex 9 10 G viqnoid ojeu JO sajonuap [e.HU9A i eiqnold [RU JO Y]99] [RUIOIX| wp AIOLISJuR ‘Aurys (ape) 29BJINS [PJOUOI sainjound |e}0UOIq ISU9JIEUU * SPPEAIEY,) ‘52192401/J, JO Satoadsqns pue soroods ueAejewig oY) JO SIAJO8IEU9 Insouserg - | AVL HIMALAYAN THORECTES 1009 Fics 5-12 Variation of denticles on posterior margin of metafemur in males of Thorectes shankara sp. n. (12, holotype). T. martensi Krikken, 1981 Material examined: Paratype d, Nepal, S-Dhaulagiri. Dhorpatan, 3000-3200 m, 7-25 May 1973, Expedition Jochen Martens (Forschungsinstitut Senckenberg, Frankfurt am Main, Germany). Distribution: Nepal, S-Dhaulagiri, 3000-3600 m a.s.l. T. shankara sp. n. Material examined: See description, type material. Distribution: India, Uttar Pradesh, 3300 m a.s.l. 1010 G. M. CARPANETO & R. MIGNANI 18 14 15 2mm Fics 13-16 Wing size in Himalayan species of the genus Thorectes: 13, T. martensi; 14, T. shankara; 15, T. nepalensis; 16, T. stellosus. T. nepalensis Baraud, 1974 Material examined: Holotype d, Nepal, Katmandu, Miss. USHQ, 1953 (Muséum d’histoire naturelle de Genève). 1 4, Nepal, Karnali zone, Churchi Lagna, N Jumla, 3400 m, 26 June-2 July 1995, Ahrens & Pommeranz leg. (G. Carpaneto collection). Distribution: W and C Nepal, 2300-3400 m a.s.l. T. stellosus stellosus Krikken, 1981 Material examined: Paratype d, Nepal, S-Dhaulagiri, Bobang, S of Dhorpatan, 2500 m, April 1970, Expedition Jochen Martens; Paratype 9, (idem) (both in For- schungsinstitut Senckenberg, Frankfurt am Main, Germany). Distribution: Nepal, S-Dhaulagiri (only known from type locality). T. stellosus annapurnicus Krikken, 1981 Material examined: Paratype d, Nepal, S-Annapurna, Gorapani Pass, 2750- 2900 m, 24-28 July 1970, Expedition Jochen Martens; Paratype 2 (idem) (both in Forschungsinstitut Senckenberg, Frankfurt am Main, Germany). Distribution: Nepal, S-Annapurna (only known from type locality). HIMALAYAN THORECTES 1011 KEY TO HIMALAYAN SPECIES AND SUBSPECIES OF THE GENUS THORECTES Ie Elytra without distinct striae. Inferior crest of male metafemora multi- dentate (Rigs 202) Pronotum nommnalliy:CONVEX RR, ost N 2 - Elytra with moderately distinct striae. Inferior crest of male metafemora at most with a single small tooth. Pronotum flattened anteriorly ...... martensi 2 Upper surface smooth. Pronotum shiny, with fine punctures. Male pro- tibiae withimuladentate inrenioreresta(Eiss l 2). e eee 3 - Upper surface wrinkled. Pronotum opaque, with large punctures. Male protibiae with multidentate inferior crest and one non-aligned additional tooth 4 3% Third external tooth of male protibiae directed downwards (Fig. 2); ely- tral surface sericeous; disc of metasternum and of abdominal sternites elabrousgandlalmosgimpunetater sn. Se ee Er shankara - Third external tooth of male protibiae directed outwards (Fig. 1); elytral surface shining like pronotum; disc of metasternum and abdominal ster- mitesipungtatesandssetosern iu en 2) Bra a ak GES RIE nepalensis 4. Dorsum coarsely wrinkled, pronotum with well impressed, star-like punctures; western form (Dhaulagiri area)................ stellosus stellosus - Dorsum finely wrinkled, pronotum with feebly impressed punctures; easternktornml(Annapumasarea)pe rd Eee stellosus annapurnicus DISCUSSION According to ZUNINO (1984) two lineages occur within the genus Thorectes: the first including most of the Mediterranean/south-European species and T. banghaasi from Tajikistan; the second one including the Himalayan species and two western Medi- terranean species (7. geminatus Gene, 1839 and 7. punctatissimus Chevrolat, 1840). Subsequentely, LOPEZ-COLON (1989) splitted the genus Thorectes into four subgenera: Thorectes Mulsant, 1942; Zuninoeus Lopez-Colon, 1989; Jekelius Lopez-Colon, 1989; Silphotrupes Jekel, 1866. The last was originally conceived by JEKEL (1866) to group species having the apical tooth of male protibia simple and clypeus semielliptic (T. punctatissimus and T. escorialensis Jekel, 1866), but LoPEZ-COLON placed here species which ZUNINO (1984) considered related to 7. punctatissimus, i.e. T. geminatus and the Himalayan species. However, the latter ones were not examined by LOPEZ-COLON (1989), and therefore his conclusions were based on descriptions only. Thus, his subgeneric arrangement needs to be confirmed after a more comprehensive revision. The new species is closely related to T. nepalensis, as suggested by the shape of male genitalia. The presence of teeth on posterior margin of male metafemora is a character shared by some geotrupid genera which are very distant among them as well as belonging to different tribes (e.g., Thorectes among Geotrupini, Chromogeotrupes Bovo & Zunino, 1983, and Phelotrupes Jekel, 1866, among Chromogeotrupini). Within the genus Thorectes, this character also occurs in T. geminatus, T. valencianus, and T. distinctus; but only 5 species (T. valencianus, T. geminatus, T. nepalensis, T. stello- sus and T. shankara) show a true denticulation (not a single tooth), i.e. a series of small teeth along the posterior margin of metafemur. 1012 G. M. CARPANETO & R. MIGNANI The genus Thorectes was so far considered to comprise wingless species (BOUCOMONT 1905; BARAUD 1966a, 1966b). In order to check this character state, we dissected several species, including all members of the subgenus Si/photrupes, and the type species of other subgenera. Actually, all members of Thorectes show wing reduction and sometimes complete absence of wings. We have calculated the ratio WL/EL (wing length/elytral length) to evaluate the meiopterism degree according to the classification used by BIONDI (1993) for Chrysomelidae Alticinae. The WL/EL may vary from micropterous to subapterous condition: only T. geminatus resulted to be apterous; 7. shankara shows a 0.3 value of the ratio and is considered micropterous; all other species examined have extremely reduced hind wing (WL/EL < 0.3) and so belong to the subapterous group. Wings of Himalayan species are shown in figs. 13-16. ACKNOWLEDGEMENTS We are grateful to the following specialists and museum curators: I. Löbl, Museum d’histoire naturelle de Geneve (Switzerland); D. Kovac and A. Vesmanis, Forschungsinstitut Senckenberg, Frankfurt am Main (Germany); E. Sprecher, Natur- historisches Museum, Basel (Switzerland): M. Snizek and Ales Bezdek (Praha); D. Ahrens, Staatliches Museum für Tierkunde, Dresden (Germany). Special thanks are due to M. Bologna (Department of Biology, University “Roma Tre”) for useful discussions and a critical reading of the manuscript. REFERENCES BARAUD, J. 1966a. Revision du sous-genre Thorectes Mulsant (Col. Scarabaeoidea) (le note): les especes ibériques. Eos 41: 181-198. BARAUD, J. 1966b. Revision du sous-genre Thorectes Mulsant (Col. Scarabaeoidea) (3e note): les especes d’Europe (sauf Péninsule ibérique) et du Moyen-Orient. Annales de la Société Entomologique de France (n.s.) 2 (3): 563-575. BARAUD, J. 1974. Révision du sous-genre Thorectes Mulsant (Col. Scarabaeoidea) (7e note). Les especes asiatiques. Bulletin de la Société Entomologique de France 79: 123-125. BOUCOMONT, A. 1905. Etude sur les Enoplotrupes et Geotrupes d Asie. Revue d’Entomologie (1904): 209-252. BionpI, M. 1993. Revisione del sottogenere Asiorestia Jacobson s. str. (Coleoptera, Chryso- melidae, Alticinae). Bollettino del Museo Civico di Storia Naturale di Verona 17 (1990): 1-56. JEKEL, H. 1866. Essai sur la classification naturelle des Geotrupes Latreille et description d’ especes nouvelles. Annales de la Société Entomologique de France (4) 5: 513-618. KRIKKEN, J. 1981. Geotrupidae from the Nepal Himalayas. New flightless species of Geotrupes Latreille 1796, with a biogeographical discussion (Insecta: Coleoptera). Senckenbergiana Biologica 61 (1980): 369-381. Lopez CoLon, J. I. 1989. Algunas consideraciones sobre la morfologia de la armadura genital masculina en el género Thorectes Mulsant, 1842 y sus implicaciones filogenéticas (Col. Scarabaeoidea, Geotrupidae). Boletin Grupo Entomologico de Madrid 4: 69-82. NIKOLAJEV, G. 1987. The scarab beetles (Coleoptera, Scarabaeoidea) of Kazahstan and Central Asıa. Academy of Sciences of Kazahastan, Alma Ata, 232 pp. (in Russian). ZUNINO, M. 1984. Sistematica generica dei Geotrupinae (Coleoptera, Scarabaeoidea: Geotru- pidae). Filogenesi della sottofamiglia e considerazioni biogeografiche. Bollettino del Museo Regionale di Scienze Naturali di Torino 2 (1): 9-162. REVUE SUISSE DE ZOOLOGIE 106 (4): 1013-1024; décembre 1999 The identity of Glomeris quadrifasciata C. L. Koch (Diplopoda: Glomeridae) René HOESS & Adolf SCHOLL University of Berne, Institute of Zoology, Division of Population Biology, Baltzerstr. 3, CH-3012 Berne. The identity of Glomeris quadrifasciata C. L. Koch (Diplopoda: Glome- ridae). - The synonymy of Glomeris quadrifasciata C. L. Koch, 1847, with G. undulata C. L. Koch, 1844, recognized by VERHOFFF (1931), is confir- med. Glomeris oblongoguttata Verhoeff, 1894, stat. n. is the valid name for Glomeris quadrifasciata sensu VERHOEFF (1911) from South Tyrol. This taxon is classified as a species closely related to G. transalpina C. L. Koch, 1836, by VERHOEFF (1911). Allozyme electrophoretic data agree with mor- phological results. Possible glacial refugia and postglacial colonization, which might explain the actual distribution of G. oblongoguttata and G. transalpina, are discussed. Specific separation of both taxa is concluded to have happened before the Pleistocene (> 1.6 mio years BP). Key-words: Diplopoda - Glomeris - revision - allozymes - postglacial colonization - refugia. INTRODUCTION In most diplopods, the gonopods as highly differentiated sclerotized structures have successfully been used for species separation and distinction. Glomeris, however, do not have gonopods. The telopods with their syncoxite that are used for copulation (HAACKER 1969) show only minor differences among species and even show variation within some species (e.g. VERHOEFF 1932, 1936a and SCHUBART 1934, G. undulata nominate form and f. conspersa). In addition, they have never been tested for their qualification as isolating mechanism. Molecular data instead, might be more useful for taxonomic studies in these animals. In fact, allozyme electrophoretic data have recently been successfully used to analyse whether the Glomeris-taxa hexasticha and intermedia are species or subspecies (HOESS er al. 1997). Taxonomic studies and revisions in Glomeris are complicated by the fact that no type material was indicated in the Glomeris literature until the mid of the 20th century. In any case, however, type material cannot be used in allozyme studies because it is usually preserved in ethanol that destroys enzyme activity. We therefore collected new material in the areas of the taxa concerned and we depend for identification on morphological characters, such as Manuscript accepted 11.05.1999 1014 R. HOESS & A. SCHOLL the colour pattern, the grooves on the shield and the shape of the last tergite, that are usually sufficiently outlined in the original descriptions. For our allozyme-based genetic analysis of the Central European Glomeris species, we also sampled a population of Glomeris quadrifasciata sensu VERHOEFF (1911) (nec Koch, 1847). We found that the oldest available name for Verhoeff's taxon is G. oblongoguttata which was first described as a local variety of G. transalpina (VERHOEFF 1894) and later classified as a variety of his G. quadrifasciata (VERHOEFF 1911). VERHOEFF (1911) considered his taxon as closely related to G. transalpina. This relationship is here confirmed by allozyme data. Furthermore, based on this allozyme data set, we postulate glacial refugia for and postglacial colonization of G. oblongo- guttata and G. transalpina. MATERIAL AND METHODS One population sample of G. oblongoguttata (endemic for South Tyrol and the Bergamo Pre-Alps) and eight population samples of G. transalpina from large parts of its range were analyzed (number of specimens in brackets): G. oblongoguttata: Latsch (17); G. transalpina: Pfynwald (19), Simplon (2), Loucherhorn (2), Airolo (13), Brugnasco (16), Albula (11), Lago di Poschiavo (2), Glurns (19). The small samples have been included especially in order to analyse ways of postglacial colonization, they are problematical as population samples for electrophoretic studies, however. The main colour characters which separate G. oblongoguttata and G. transalpina are shown in Fig. 1, and the distribution of both taxa (compiled from literature) and the localities of the sample sites are given in Fig. 2. We used routine enzyme electrophoretic methods of our laboratory (cf. SCHOLL et al. 1978). Vertical starch gel electrophoresis was conducted using the same buffer systems as before (HOESS er al. 1997). 18 enzyme loci were analyzed. The enzymes investigated and the loci scored (in brackets) are: aspartate aminotransferase (Aat-1, Aat-2), glyceraldehyd-3-phosphate dehydrogenase (Gapdh), glucose-6-phosphate iso- merase (Gpi), hexokinase (Hk), leucine aminopeptidase (Lap), L-lactate dehydrogenase (Ldh-1, Ldh-2), malate dehydrogenase (Mdh-1, Mdh-2), malic enzyme (Me), mannose- 6-phosphate isomerase (Mpi), peptidase (Pep), 6-phosphogluconate dehydrogenase (Pgd6), phosphoglucomutase (Pgm), superoxide dismutase (Sod-1, Sod-2) and sorbitol dehydrogenase (Sodh). The zymograms were photographed (Polaroid) for reference. We refer to observed electromorphs as alleles which are identified by their electrophoretic mobility (in mm) relative to previously studied species (HoEss er al. 1997). We used the BIOSYS-1 programme package (SWOFFORD & SELANDER 1989) for calculation of allele frequencies and data treatment. Using the matrix of Nei's distance (NEI 1978), a pheno- gram of the populations was created by average linkage cluster analysis (UPGMA). With the allozyme data, we also tried to test the biological species concept by MAYR & ASHLOCK (1991) where species are defined as groups of interbreeding natural populations that are reproductively isolated from other such groups. For the usability of small samples cf. GORMAN & RENZI (1979). IDENTITY OF GLOMERIS QUADRIFASCIATA 1015 Heterozygosity was not estimated and tested with Hardy-Weinberg-expectations because most samples were collected in a very small sampling area. Glomeris have low individual mobilities. This increases the chance of picking up siblings. The assumptions that underly the Hardy-Weinberg-principle are therefore not given for these samples. 4 mm Fic. | Colour patterns of Glomeris transalpina (a, b) and G. oblongoguttata (c, d). Shield (2nd) and 3rd tergite (a, c) (dorsal view) and last (12th) tergite (b, d) (posterior view) (females). Scale: 4 mm. TAXONOMY Glomeris oblongoguttata Verhoeff, 1894 stat. n. Fos NC ds? Glomeris transalpina C. L. Koch var. oblongoguttata Verhoeff, 1894: 17, Sulden (South Tyrol); VERHOEFF 1902: 180. Glomeris transalpina C. L. Koch var. spinalemontis Verhoeff, 1902: 180, Mt. Spinale (South Tyrol); VERHOEFF 1906: 220. Glomeris quadrifasciata C. L. Koch sensu VERHOEFF 1911: 113-114, 139 (nec Koch, 1847); ATTEMS 1927: 255; VERHOEFF 1929: 563; VERHOEFF 1930: 655; VERHOEFF 1931: 429, 433, pl. 7, figs 33, 34 [part]; VERHOEFF 1932: 641; VERHOEFF 1936a: 430; VERHOEFF 1936b: 231; VERHOEFF 1937: 168; STRASSER & MINELLI 1984: 197; FODDAI et al. 1995: 12: Glomeris quadrifasciata sensu VERHOFFF (loc. cit.) (nec Koch, 1847) var. sevini Verhoeff, 1931: 431, Bergamo Pre-Alps; VERHOEFF 1937: 167. Glomeris quadrifasciata sensu VERHOFFF (loc. cit.) (nec Koch, 1847) var. brixiensis Verhoeff, 1931: 432, Brescia (Lombardia). 1016 R. HOESS & A. SCHOLL Diagnosis: Shield with broad yellow-red margins at the outer parts of front and hind edge. Central dark spots on tergites 2-11 parallel-sided or divergent posteriorly. Dark spot on the last tergite triangular, almost reaching the hind edge with the tip. No principal groove but 3-7 accessory grooves on the shield. Last tergite in the male with a notch in the hind margin. Dark pigment always in well defined spots. Speckles often present. VERHOEFF (1911) designated, with modified and additional characters, G. qua- drifasciata as a species closely related to G. transalpina Koch. G. quadrifasciata sensu Verhoeff differs from G. quadrifasciata Koch by the following characters: the former has a large, triangular, dark spot on the last tergite that hardly separates the two yellow- red spots, and the shield has a broad yellow-red margin at the outer parts of the hind edge, too (Fig. 1). These characters do not lie within the range of variation of G. undu- lata Koch (see below) to which Koch's G. quadrifasciata belongs. This implies that Verhoeffs taxon is a different species. Verhoeffs use of the name G. quadrifasciata C. L. Koch is therefore incorrect. VERHOEFF (1931) actually recognized this fact but he did not explicitely rename his taxon. Glomeris transalpina var. oblongoguttata Verhoeff, 1894, is the oldest available name for Verhoeff’s taxon. 100 km O G. transalpina —-- D G. oblongoguttata a | A On ) \ << Pfnw O = Lago di Poschiavo \ HE Ehynwald Simplon ye EI 5 a Fic. 2 Distribution (compiled from literature) and sample sites of Glomeris transalpina and G. oblongo- guttata. IDENTITY OF GLOMERIS QUADRIFASCIATA 1017 Glomeris undulata C. L. Koch, 1844 The synonymy of G. undulata presented here is restricted to the names that are needed within the scope of this paper: Glomeris undulata C. L. Koch, 1844: H. 40, pl. 8, South Germany. Glomeris conspersa C. L. Koch, 1847: 89, South Germany [synonymized with G. undulata by HoEss & SCHOLL 1999: 657, 659]. Glomeris quadrifasciata C. L. Koch, 1847: 91, origin unknown, probably South Germany; C. L. KocH 1863: 108, pl. 49, figs. 98 a, b, c, one specimen indicated from the museum at Bamberg; VERHOEFF 1931: 433 [part]. Glomeris tridentina Latzel, 1884: 118, South Tyrol [synonymized with quadrifasciata by VERHOEFF 1902: 177]. Glomeris conspersa quadrifasciata C. L. Koch sensu VERHOEFF 1902: 180; VERHOEFF 1906: 166, IE Diagnosis: Shield with broad yellow to red margins only at the outer parts of the front edge. Central dark spots on tergites 2-12 variable: parallel-sided or divergent posteriorly in the nominate form, convergent posteriorly in f. conspersa. No principal groove, except some specimens of f. conspersa from the South-eastern Alps which have one, but with 3-4 accessory grooves on the shield. Only some males of nominate form with a notch in the hind margin of the last tergite. Dark pigment next to the central spots usually scattered in speckles, seldom without speckles. KocH described (1847, 1863) and illustrated (1863) Glomeris quadrifasciata. On the shield, this taxon shows no complete groove (character 1), but it has a broad yellow band (character 2) along the front edge. The last tergite has two large yellow-red spots that are separated by a broad dark band (character 3) which widens towards the front edge. The middle segments show four bands (character 4) of yellow-red spots that are separated in the middle by a dark band made of parallel-sided spots (character 5). Characters 1, 2, 3 and 5 are typical for what is presently recognized as G. undulata Koch, whereas character 4 lies within the range of variability of G. undulata which is known to be one of the most variable species of the genus. Typically, G. undulata shows speckles which are absent in G. quadrifasciata. However, there are some varieties of G. undulata, that do not express the speckles (see e.g. VERHOEFF 1931: confirmed by own data from electrophoretically analyzed specimens). Consequently, G. quadrifasciata does not provide any character for its own. We therefore agree with VERHOEFF (1931) who recognized G. quadrifasciata Koch as a synonym of G. undulata Koch. LATZEL (1884) synonymized G. quadrifasciata Koch with G: connexa Koch. This is not acceptable because the significant characters 1 and 2 are different in G. connexa (see also VERHOEFF 1902). VERHOEFF (1902) first classified quadrifasciata as a subspecies of G. conspersa; but characters 3, 4 and 5 are different in the latter. Glomeris transalpina C. L. Koch, 1836 Figs la, b; 2; 4 This species shall also be diagnosed here. It was classified as closest related to G. oblongoguttata by VERHOEFF (1911) which we confirm by allozyme data later in this paper. 1018 R. HOESS & A. SCHOLL TABLE | Allele frequencies of the 14 polymorphic loci for all populations of Glomeris oblongoguttata (Latsch) and G. transalpina (other populations). Number of examined specimens in brackets. A = average number of alleles per locus over all 18 loci. Locus/ Population Allele Latsch Glurns Lago di Albula Brugnasco Airolo Loucher- Simplon Pfynwald Poschiavo horn (17) (19) (2) (11) (16) (13) (2) (2) (19) Aat- | 105 0.53 0.55 0.25 0.22 0.43 0.69 0.75 I 0.32 104 0.06 - - - - - - - - 100 - - - 0.11 - - - - - 97 0.38 0.45 0.75 0.67 0.57 0.31 0.25 - 0.68 Ome ys 0:03 - - - - - - - - Gapdh 111 - 0.08 - - - - - - - 109 l 0:92 | 1 l | | 1 1 Gpi 100 I | | I | | | 1 0.92 96 - - - = - - - - 0.08 Hk 103 0.74 0.39 0.25 0.25 0.12 0.50 0.75 0.75 0.21 100 0.26 0.61 0875 OFS 0.8 0.50 0.25 0.25 0.79 Ldh-2 100 0.68 - - - - - - - - 97 0.23 - - - - - - - - 93 0.09 - - - - - - - - 82 - l I l | | | I 1 Mdh-1 105 - - - - 0.28 0.37 - 0.50 0.63 100 I | ] | 0.72 0.63 | 0.50 0.37 Mdh-2 102 - - 075 - - - - - - 100 0.66 0.82 0.25 0.80 0.88 0.88 0.50 il 0.94 9930.09 - - - - 0.06 - - - 98 0.09 0.06 - 0.20 0.04 - 0.50 - - 97 0.04 0.03 - - 0.04 - - - - 95 0.04 0.06 - - - 0.06 - - 0.03 94 0.04 - - - - - - - - 9] 0.04 0.03 - - 0.04 - - - 0.03 Me 102 - 0.94 | | 0.09 0.25 - - - 100 I 0.06 - - 0.82 0.56 - I I 98 - - - - 0.09 0.19 ] - - Mpi 104 0.18 I l I I I 0.75 1 1 100 0.41 - - - - - = = = 97 - - - - - - 0.25 - - 95 0.09 - - - - - = - = IDENTITY OF GLOMERIS QUADRIFASCIATA 1019 Pep 104 0.29 - - - - - - - - 102 0.18 - - 0.05 0.14 027 - - - OOPS 90:58 0.10 - 0.18 0.80 0.61 | 0.50 0.74 98 - 0.40 | - 0.03 0.04 - - 0.05 96 - 0.50 - 0.77 0.03 0.08 - 0.50 0.21 Pgd6 106 - - - - 0.10 0.14 I - - 104 0.06 - - - - - - - - 100 0.94 I | | 0.90 0.86 - | | Pgm 100 - 0.14 - 0.05 0.11 0.35 0.50 - 0.61 97, - 0.86 | 0.95 0.89 0.65 0.50 | 0.39 95 0.05 - - - È - - - - 93 0.50 - - - - - - - - 89 0.45 - - - - - - - - Sod-2 100 0.77 0.40 0.50 0.41 0.47 0.46 - - 0.45 98 - 0.05 0.25 0.36 0.28 0.31 0.75 0.50 0.08 95 0.23 0.29 - 0.05 0.06 0.23 0.25 0.50 0.31 93 - 0.26 0.25 0.18 0.19 - - - 0.16 Sodh 2 - 0.11 - 0.21 0.87 0.62 1 - 0.47 109 0.03 0.52 0.50 - - - - - 0.03 106 - 0.26 0.50 0.79 013 0.38 - | 0.29 103 - 0.11 - - - - - - 0.21 OO 0:97, - - - - - - - - A 2.22 1.94 133 Mol 1.94 1.83 2.33 1,22 1.78 Val Venosta Grisons Ticino Bernese Alps Valais Diagnosis: Shield with broad (yellow-)red margins at the outer parts of the front and hind edge. Central dark spots of tergites 2-11 confluent with the remaining dark pigment, convergent posteriorly but not reaching the hind edge. Dark spost on the last tergite triangular but ending far away from the hind edge. No principal groove but 3-4 accessory grooves on the shield. Never a notch in the hind margin of the last tergite. Dark pigment always in well defined spots. Speckles rarely present. RESULES The alleles observed and their frequencies are listed in Table 1 except for the four loci Aat-2, Lap, Ldh-1 and Sod-1 that were monomorphic, and all populations of both taxa were fixed for the same alleles, viz. Aat-2100 Lap!00, Ldh-1!99 and Sod-199, respectively. Three other loci were monomorphic except for one or two populations, and the same allele was found in both taxa: Locus Gapdh, allele Gapdh!" (except the G. transalpina population from Glurns with one additional allele), locus Gpi, allele Gpi!® (except the G. transalpina population from Pfynwald with one additional allele), 1020 R. HOESS & A. SCHOLL and locus Mpi, allele Mpi!* (except the G. transalpina population from Loucherhorn with one additional allele and the G. oblongoguttata population with three additional alleles). Locus Ldh-2 was monomorphic in all populations of G. transalpina (allele Ldh-282), the G. oblongoguttata population, however, had three alleles that were not observed in G. transalpina. The other loci showed high genetic polymorphism. Exclu- ding the populations with small sample sizes (N < 10), the average number of alleles per locus (total: 18 loci) was 2.22 in G. oblongoguttata and 1.61-1.94 in the G. transalpina populations. G. oblogoguttata and G. transalpina were clearly genetically distinct by alter- native alleles at the loci Ldh-2 and Pgm (Table 1). Additionally, the two geographically closest populations of each taxon namely Latsch and Glurns (distance about 25 km, both localities are in the same valley) showed remarkable differences in the allele frequencies at five other loci. These loci were Hk, Me, Mpi, Pep and Sodh (Table 1). Of the 59 alleles recorded at the 18 loci, 21 alleles were found in both taxa, 15 alleles were only observed in G. oblongoguttata, and 18 alleles were only observed in G. transalpina, 13 of these 18 alleles of G. transalpina were widespread in the range of that species. Several alleles of G. transalpina were restricted in their distribution to certain regions (Table 1). At locus Mdh-1, the allele Mdh-1!9 was only found in Ticino (populations Brugnasco and Airolo) and Valais (populations Simplon and Pfynwald). At locus Me, the allele Me! was fixed or nearly fixed in the three eastern populations (Glurns, Lago di Poschiavo and Albula), it was rare in Ticino and was not present in the three western populations (Loucherhorn, Simplon and Pfynwald). This allele is substituted by alleles Me! and Me?8 in the western populations. At locus Pgd6, the allele Pgd6!° was only found in the Ticino populations (Brugnasco and Airolo) and in Loucherhorn. TABLE 2 Nei-distances D in pairwise comparisons of populations of Glomeris transalpina and G. oblon- goguttata (sample sizes > 10). G. transalpina Pfynwald Airolo Brugnasco Albula Glurns G. transalpina Airolo .037 Brugnasco .038 .020 Albula .158 .098 .120 Glurns .149 .086 121 .034 G. oblongoguttata Latsch .264 .238 .268 375 .324 Table 2 shows the genetic distances D (Nei 1978) in pairwise comparisons of all populations (sample sizes > 10) using all 18 loci. These distances varied from 0.02 - 0.16 among different populations of G. transalpina and from 0.24 - 0.38 among populations of the two taxa. IDENTITY OF GLOMERIS QUADRIFASCIATA 1021 —— Pfynwald Airolo Brugnasco G. transalpina —— Albula L____ Glurns Latsch G. oblongoguttata Tg TU Um RESET IF ZARA T El 0.3 0.2 0.1 0.0 Nei-distance D FIG. 3 UPGMA dendrogram using Nei’s coefficient of genetic distance for populations of Glomeris transalpina and G. oblongoguttata (sample sizes > 10). The Nei distances were graphically transformed into an UPGMA dendrogram (Fig. 3). G. oblongoguttata 1s separated from G. transalpina at a distance level of 0.29. The G. transalpina populations are found in two subclusters that separate at a distance level of D = 0.12 and contain the Valais and Ticino populations on the one hand and the eastern populations on the other hand. DISCUSSION The most distant populations of G. transalpina, Glurns and Pfynwald, that are about 225 km apart, share at least one allele at each locus. On the other hand, the closest populations of both taxa (G. transalpina population from Glurns and G. oblongoguttata population from Latsch) that are found only 25 km apart in the same valley have alternative alleles at two loci (Ldh-2 and Pgm). This indicates that the two taxa belong to different gene pools. This is also supported by five other loci where the allele frequencies between the Latsch and Glurns populations are very different (Hk, Me, Mpi, Pep and Sodh). Specimens with transition in morphological characters that would indicate hybridisation are not known. We therefore conclude that G. transalpina and G. oblongoguttata are in fact different species. However, these species are closely related. In a previous study (HoEss er al. 1997) we found that G. undulata (sub G. conspersa), which shares characters 1 and 2 (see above) with G. transalpina, is genetically more distant to G. transalpina than G. oblongoguttata. The locally observed alleles, the allele substitution observed at the Me locus, and the Nei-D values that are graphically interpreted in the dendrogram show two subgroups within G. transalpina, an eastern subgroup from Grison and its adjacent valleys, and a western subgroup from Ticino to Valais and the Bernese Alps, respectively. However, there is no reason to assume that these subgroups are specifically distinct. The subgroups are not correlated with the variation of the colour pattern. Instead, each population has some own trends in altering the basic colour pattern. It is therefore not appropriate to rank the subgroups as subspecies. 1022 R. HOESS & A. SCHOLL present range of G. transalpina ~ sampling sites \ (cf. Fig. 2) \ le PU € &2I \ | maximum ı | glaciation e i | during the last ı | Ice Age ı | (Würm) Adda Glacier à 7 nunataks aay DS om oa È ©) probable refugia VÀ Bergamo Fic. 4 Supposed glacial refugia and ways of postglacial colonization of Glomeris transalpina (see text for explanation). The geographically restricted alleles suggest that there must have been at least two glacial refugia for G. transalpina from where colonization of the presently inhabited localities has started (Fig. 4). The eastern subgroup with Albula, Glurns and Lago di Poschiavo probably had a refuge in the mountains north of Bergamo and/or Brescia. Lago di Poschiavo may have been colonized from the Bergamo refuge through the Adda Valley, while Glurns was colonized from the Brescia refuge. The origin of the Albula population is not clear, the allozyme data are not conclusive in this case. All western populations may originate from the nunataks in Ticino. The Valais most pro- bably was colonized only once over the Nufenen, and the Bernese Alps independently from that over the Nufenen and the Grimsel. The alleles that are restricted to Valais and Ticino do not favour a separate colonization of the Simplon from the south. Loucherhorn was a Nunatak and might have been inhabited by G. transalpina during the Pleistocene. The allozyme data, however, show that this population is very close to the Valais and Ticino populations which suggests that Louchernhorn was colonized from the south. Taking into consideration its present distribution, we assume that G. oblongo- guttata had its glacial refuge on both sides of the Oglio Glacier like the eastern populations of G. transalpina. This, of course, requires genetic isolation during the Pleistocene. We, therefore, conclude that specific separation of G. oblongoguttata and G. transalpina occurred before the Pleistocene (> 1.6 mio years BP). IDENTITY OF GLOMERIS QUADRIFASCIATA 1023 At present, both species inhabit woods of the mountainous, subalpine and the adjacent levels. G. transalpina prefers higher regions compared to G. oblongoguttata. Thus, especially in the pre-alps, colonization would not be confined to the valleys. This renders the interpretation of postglacial colonization more difficult, but shows that mountain chains are not necessarily a barrier for dispersal. The limited present ranges of both species, as compared to e.g. the widespread G. marginata, may be due to the crooked topography of the Alps. The scarcity of records of G. transalpina in the Northern Alps also supports the hypothesis of southern refugia and a northward expansion. ACKNOWLEDGEMENTS M. E. Braunwalder, B. Gantenbein, P. Marmet and P. Sonderegger helped us gather the glomerids used in this study. L. Beer, L. Frauchiger and V. Siegfried assisted in the laboratory work. H. Baur gave nomenclatorial advice. REFERENCES ATTEMS, C. 1927. 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REVUE SUISSE DE ZOOLOGIE 106 (4): 1025-1034; décembre 1999 Redescription of Electrogena galileae (Demoulin, 1973) (Ephemeroptera, Heptageniidae) Carlo BELFIORE! and Michel SARTORI? | Dipartimento di Zoologia, Universita di Napoli Federico II, via Mezzocannone 8, I-80134 Napoli. 2 Musée de Zoologie, Palais de Rumine, Case postale 448, CH-1000 Lausanne 17. Redescription of Electrogena galileae (Demoulin, 1973) (Ephemerop- tera, Heptageniidae). - Male imagines and larvae of Electrogena galileae (Demoulin, 1973), from Israel and Lebanon, are redescribed using the stan- dard diagnostic characters, both quantitative and qualitative, recently proposed for the species of the genus Electrogena. Three new larval characters (One meristic and two multistate ones), useful for the taxonomy of E. galileae, are added to the former set. E. galileae proved to be distinctly separated from the other species described by the standard set. Key-words: Ephemeroptera - Heptageniidae - Electrogena galileae - redescription - taxonomy - Lebanon - Israel. INTRODUCTION A lot of Electrogena species were described from Near and Middle East (BRAASCH 1978, 1980a, b, 1981, 1983a, b; KAZANCI 1986, 1987, 1990; KAZANCI & BRAASCH 1986) and many other entities belonging to this genus are possibly still unknown. As pointed out by SARTORI (1992), most of the species from those areas were only partially and summarily described. Therefore any attempt of identification is still very difficult. Recently several diagnostic characters for the separation of Electrogena species were proposed (BELFIORE 1994, 1995, 1996, 1997; BELFIORE & DEsıo 1995; BELFIORE et al. 1997; MALZACHER 1996), which proved to be very useful: for the identification and discrimination of most European species. Male imagines can be identified mainly by the markings on abdomen, some characteristics of genitalia (i.e. shape of penis lobes and titillators, denticulation of outer sclerites (MALZACHER 1996)), wing and body colouration. Diagnostic characters for female imagines are abdominal markings, wing colouration, shape of the subgenital plate and morphology of eggs. Many more characters are available for the identification of larvae: a set of sixteen diagnostic features, both quantitative and qualitative, was proposed by BELFIORE (1994). Redescriptions of species using these new taxonomic tools is a necessary step for checking the validity of Electrogena species and understanding the relationships among them. This would be particularly useful for those areas, like Near and Middle East where the diversification of the genus seems very high. Manuscript accepted 15.06.1999 1026 C. BELFIORE & M. SARTORI In this paper we begin the revision of the Middle Eastern species with a redescription of Electrogena galileae, formerly described by DEMOULIN (1973) and SARTORI (1992). REDESCRIPTION Electrogena galileae (Demoulin, 1973) Ecdyonurus golanicus Samocha, 1972, nomen nudum Ecdyonurus galileae Demoulin, 1973 Ecdyonurus galileae: Dia 1983; MOUBAYED 1986; KOCH 1988 Electrogena galileae: SARTORI 1992 Material. ISRAEL (Museum of Zoology, Lausanne; Michel Sartori leg. et det.): Hula Valley, Nahal Dan, Tel Dan, 170 m, 8.V.1990, 1 male imago (reared) with nymphal skin, 3 larvae; Hula Valley, Nahal Dan, 180 m, 7.V.1990, 3 larvae; Hula Valley, Enot Layish (Tel Springs), 200m, 12.V.1990, 5 larvae. LEBANON (C. Belfiore Collection; A. Dia leg.): Aouali River, 710m, 4.V11.1980: 1 male imago; Nabaa Mourched, 800 m, 13.V1.1981, 2 male imagines; Santa Yahfoufah, 8.1V.1985, 1 male imago. MALE IMAGO (in alcohol) Body 9.5 mm; cerci: 23 mm. Eyes grey, ventral margin bordered with a dark grey band. Thoracic tergites and sternites light brown, pleurae pale yellowish. Fore legs light brown: femora with a wide dark band in the middle and a smaller one near the junction with the tibia; the last two segments of tarsi are lighter. Mid and hind legs yellowish with the same markings as fore legs; tarsi slightly darker. Wings uniformly tinged with yellowish brown; veins light brown, cross veins darker, bordered with brownish violet in the fore part of fore wing. First costal cross vein bold, coloured with violet brownish. Abdomen yellowish, with violet brown markings (Fig. 1). Cerci greyish brown, slightly darker at joints. Genitalia (Figs 2-3): styliger yellowish, hind margin with two flat and widely rounded projections. Gonopodes greyish, last segment lighter. Penis stem very short. Penis lobes asymmetrical, outer edge rounded, inner edge straight. 2-4 small denticles on outer sclerite, near basis of lobes. LARVA (in alcohol) Body length (full-grown larvae): 8 mm (male); 8.5 mm (female). Cerci: 14-15 mm. General aspect and colouration. Larvae of Æ. galileae are small with long and slender legs. General colouration is brown with lighter markings. Two light rounded spots are at sides of median line, near the fore margin of head (Fig. 4). Imaginal markings are visible also in half grown larvae. Diagnostic characters (number of specimens examined=11). The set of larval diagnostic characters is fully explained and figured in BELFIORE (1996, 1997). A. Quantitative characters Mean, range and variance are reported. E. galileae is compared with the Elec- trogena species previously examined by the standard set of diagnostic characters: REDESCRIPTION OF ELECTROGENA GALILEAE 1027 1mm 0.2 mm Fics 1-3 Electrogena galileae, male imago: III-V tergites and sternites (from a slide) (1); genitalia from ventral view (2); penis from dorsal view (3). 1028 C. BELFIORE & M. SARTORI 0.2mm Fics 4-6 Electrogena galileae, larva: head (4); fore femur from dorsal view (5); magnification of hind side of femur (long bristles are truncated). E. calabra Belfiore, 1995; E. fallax (Hagen, 1864); E. grandiae (Belfiore, 1981); E. gridellii (Grandi, 1953), E. hyblaea Belfiore, 1994; E. lateralis (Curtis, 1834); E. malickyi (Braasch, 1983); E. ujhelyii (Sowa, 1981), E. zebrata (Hagen, 1864) (BELFIORE 1996) and E. lunaris Belfiore & Scillitani, 1997 (BELFIORE et al. 1997). Phenetic affinities between species are discussed by comparison of mean values and ranges. a) Meristic characters: 1. N_PLP : 11.09; 7-19; 10.9909. The number of hairs on fore side of first segment of maxillary palpus is moderately low. Hairs are often distributed in two groups, near basis and at apex of segment (Fig. 15). Closest species are E. calabra (mean: 9.97) and E. zebrata (12.52). Range overlaps with all other species. REDESCRIPTION OF ELECTROGENA GALILEAE 1029 0.2 mm 0.5 mm 11 13 0.2 mm 0.1 mm Fics 7-14 Electrogena galileae, larva: bristles on the ventral surface of labrum (7); labrum (9); Ist gill plate (10); 7th gill (11); bristles on upper surface of fore femur (12); Sth comb shaped bristle of galea-lacinia (13); apex of hypopharynx lobe (14). E. ujhelyii, larva: bristles on the ventral surface of labrum (8). 1030 C. BELFIORE & M. SARTORI 0.2 mm e a TATESE SS in ag 17 {Was ES Fics 15-17 First segment of maxillary palpus of larvae: Electrogena galileae (15); E. zebrata (16) E. gridellii (17). In (16) and (17) the long hairs on fore margin are not figured. REDESCRIPTION OF ELECTROGENA GALILEAE 1031 2. N_OUT: 0; 0; 0; 0. The only species with no hairs on outer margin of galea-lacinia and zero variance are E. malickii and E. zebrata. Species with such hairs always present are E. calabra, E. fallax, E. gridellii and E. ujhelyii. 3. N_CBS: 19.77; 19-21.5; 0.7182. The number of comb-shaped bristles on the fore margin of galea-lacinia is moderately high. Closest species are E. calabra (19.12) and E. ujhelyii (20.38). Range is overlapping with all species. 4. N_TCB (Fig. 13): 14.55; 13-16; 0.7227. The number of teeth on the 5th comb- shaped bristle (fore margin of galea-lacinia) is the highest among species considered. The closest species is E. malickyi (13.98). Non-overlapping are E. grandiae (6.5-11.5) and E. lunaris (5-7). 5. N_CLW: 2.36; 2-4; 0.4545. The number of teeth on the tarsal claw is generally two, with the exception of only one specimen which has 3 to 5 (modal value=4). Closest species is E. ujhelyii (2.52), which share the same range and the unusually high variance. The only non-overlapping species are those with an invariant single tooth (E. lateralis and E. lunaris). 6. N_BVF: 22.45; 6-36; 90.8727. The number of bristles on the ventral side of femora, near the hind margin, shows an unusual wide range. However E. galileae can be considered belonging to the group with «many» such bristles (E. ujhelyii (27.90), E. zebrata (36.65), E. fallax (41.59)). The maximum number of bristles shared by all other species is 3. A very distinctive feature of E. galileae is the shape of these bristles (Fig. 6), blunt or rounded at apex (pointed in other species). One more meristic character proved to be relevant to taxonomy of E. galileae. It can be added to the standard set: 7. N_HFF: Number of setae (long and tiny hairs) on the fore (ventral) margin of fore femur. Hairs counted are those at least 1.5 times long as bristles along the fore margin of femora. E. galileae has generally three long hairs near basis of the femora (Fig. 5). None of the other species share this character. b) Ratio characters: 8. R_IGI (Ist gill length/width): 2.116; 1.798-2.411; 0.0284. First gill is moderately short, intermediate between E. lateralis (2.137) and E. zebrata (1.992). Range 1s overlapping with all species. The shape of the first gill (Fig. 10) is relatively constant among individuals. 9. R_7GI (7th gill length/width): 2.613; 2.304-2.807; 0.0247. Seventh gill is moderately long (Fig. 11), intermediate between E. grandiae (2.627) and E. malickyi (2.423). It cannot be separated from other species by range. 10. R_LBR (total width of labrum/width of lateral projections): 4.157; 3.835-4.511: 0.0352. Labrum is very slender. R_LBR is the lowest among species considered. Closest species is E. malickyi (4.331). The only non-overlapping species is E. lunaris (4.786-6.744). The shape of labrum is very peculiar (Fig. 9): it recalls the shape of labrum in the Ecdyonurus species. The tips of lateral projections are often bent backwards. 1032 C. BELFIORE & M. SARTORI 11. R_GLA (outer distance/inner distance between glossae): 3.213; 2.958-3.778; 0.0710. Glossae are relatively close each other. Closest species are E. lateralis (3.174) and E. hyblaea (3.237). This character and the next one are the less discriminating among the Electrogena species. 12. R_GLB (outer distance between glossae/mean width of glossae): 2.812; 2.537- 3.056; 0.0463. Glossae are moderately wide. Closest species are E. fallax (2.777) and E. ujhelyii (2.837). B. Shape and multistate characters 13. S_HLB - the apex of lateral lobes of hypopharynx is covered with short hairs, about 1/4 long as hairs along the fore margin (Fig. 14). This character state is intermediate between species with very short hairs (1/8 of fore hairs: E. lateralis, E. lunaris, E. ujhelyii) and species with long hairs all around the apex (all other species). 14. S_PGL - paraglossae are narrow, somewhat pointed. 15. S_PNT - hind corners of pronotum are smoothly rounded. 16. S_BFE - distal bristles on upper surface of fore femora are short and rounded, with diverging sides (Fig. 12). 17. S_TAR - tarsi are darkened at apex only. 18. S_7GI - 7th gill is gradually narrowing at apex. Two more characters are relevant to taxonomy of E. galileae and are to be added to the previous list. Even if they could be considered quantitatively, two discrete character states can be recorded throughout the species here considered. We provi- sionally treat them as multistate characters. 19. S_PLB: length of bristles on hind margin of the first segment of maxillary palpus. These bristles are very short in E. galileae (1/4 to 1/5 as width of segment) (Fig. 15). The other species have longer bristles (at least 1/3 as width of segment) (Fig. 17) with the exception of E. zebrata (about 1/4) (Fig. 16). 20. S_LBB: arrangement of paramedian bristles on ventral side of labrum, near the fore margin (a character used for the discrimination of helveticus and venosus group within the genus Ecdyonurus (BELFIORE & BUFFAGNI 1994). In E. galileae these bristles are arranged in two irregular rows (Fig. 7), unlike other Electrogena species, which share slender bristles arranged in a single row (Fig. 8). DIAGNOSIS E. galileae definitely appears phenetically far from the other considered species. It could be more usefully compared with other Electrogena species from the same area, when complete redescriptions of them will be available. At this moment the larval diagnosis 1s very easy because of some unique features: labrum with lateral projections often bent backwards; bristles on ventral surface of labrum (S_LBB) arranged in two rows; some long hairs at basis of fore margin of femora (N_HFF); apex of lobes of hypopharynx with hairs 1/4 as long as hairs along the fore margin of lobes (S_HLB). REDESCRIPTION OF ELECTROGENA GALILEAE 1033 Useful characters for identification are also the following: labrum very slender (R_LBR low); very short bristles on the hind side of the first segment of maxillary palpus (S_PLB), lack of hairs on outer margin of galea-lacinia (N_OUT), a very high number of teeth on the Sth comb-shaped bristle (N_TCB). Male imagines can be identified by markings of abdomen, colouration of wings, pigmentation of wing veins and shape of penis stem and penis lobes. ACKNOWLEDGMENTS We are indebted to Prof. A. Thomas (Toulouse) fot the kind gift of specimens from Lebanon. LITERATURE BELFIORE, C. 1994. Taxonomic Characters for Species Identification in the Genus Electrogena Zurwerra & Tomka, with a Description of Electrogena hyblaea sp. n. from Sicily (Ephe- meroptera, Heptageniidae). Aquatic Insects 16(4): 193-199. BELFIORE, C. 1995. Description of Electrogena calabra n. sp., a new species from Southern Italy (Ephemeroptera, Heptageniidae). Annales de Limnologie 31(1): 29-34. BELFIORE, C. 1996. Identification and discrimination of Electrogena species by numerical methods (Ephemeroptera, Heptageniidae). Systematic Entomology 21: 1-13. BELFIORE, C. 1997. Taxonomic characters and discrimination of species in the genus Electrogena Zurwerra & Tomka (Ephemeroptera, Heptageniidae), pp. 427-433. In: LANDOLT, P. & SARTORI, M. (eds). Ephemeroptera & Plecoptera: Biology, Ecology, Systematics. MTL Fribourg, X1+569 pp. BELFIORE, C. & BUFFAGNI, A. 1994. Revision of the Italian species of Ecdyonurus helveticus group: taxonomy of nymphs (Ephemeroptera, Heptageniidae). Bulletin de la Societe entomologique suisse 67: 143-149. BELFIORE, C. & Desio, F. 1995. Taxonomy and distribution of Electrogena ujhelyii (Sowa, 1981) (Insecta: Ephemeroptera: Heptageniidae). Annalen des Naturhistorisches Museum in Wien 97 B: 151-154. BELFIORE, C., SCILLITANI, G., PICARIELLO, O. & CATAUDO, A. 1997. Morphological and electrophoretic evidence for a new species of Electrogena from Central Italy: description of E. lunaris sp. n. Aquatic Insects 19(3): 129-140. BRAASCH, D. 1978. Neue Ecdyonurus-Arten (Ephemeroptera, Heptageniidae) aus dem Kaukasus und aus Transkaukasien (Sowjetunion). 1. Entomologische Nachrichten 22(10): 145-160. BRAASCH, D. 1980a. Neue Ecdyonurus-Arten (Heptageniidae, Ephemeroptera) aus dem Kaukasus und Transkaukasien (Sowjetunion). II. Entomologische Nachrichten 23(2): 23-28. BRAASCH, D. 1980b. Neue Ecdyonurus-Arten (Heptageniidae, Ephemeroptera) aus dem Kaukasus und Transkaukasien (Sowjetunion) (3). Entomologische Nachrichten 24(7): 103-110. BRAASCH, D. 1981. Eintagsfliegen aus Anatolien und Iran (Insecta, Ephemeroptera). Faunistische Abhandlungen, Staatliches Museum fiir Tierkunde Dresden 8(6): 75-79. BRAASCH, D. 1983a. Ecdyonurus bothmeri n. sp. aus dem Iran (Ephemeroptera, Heptageniidae). Entomologische Nachrichten 27(4): 177. BRAASCH, D. 1983b. Ecdyonurus armeniacus n. sp. aus der Armenischen SSR (UdSSR). Reichenbachia 21: 87-90. DEMOULIN, G. 1973. Contribution a l’etude des Ephémèroptères d'Israel. Introduction et 1. Heptageniidae. Bulletin de l'Institut royal des Sciences naturelles de Belgique 49(8): 1-19. 1034 C. BELFIORE & M. SARTORI Dia, A. 1983. Recherches sur l’écologie et la biogéographie des cours d’eau du Liban méridional. These Doctorat d’Etat, Marseille, 302 pp. Kazancı, N. 1986. A new Ephemeroptera (Heptageniidae) species from Anatolia. Doga Tu Bio. 10(3): 391-393. Kazancı, N. 1987. Ecdyonurus necatii, a new Ephemeroptera (Heptageniidae) species from Turkey. Aquatic Insects 9 (1): 17-20. Kazancı, N. 1990. Turkiye Heptageniidae (Insecta: Ephemeroptera) faunasi uzerine arastir- malar.Il. Cins Electrogena Zurwerra et Tomka, 1985. Hacettepe Fen. Muhendislik Bilimleri Derg. Ser. A 11: 169-180. KAZANCI, N. & BRAASCH, D. 1986. Zwei neue Heptageniidae (Ephemeroptera) aus Anatolien. Bulletin de la Société entomologique suisse 59: 365-368. Kocu, S. 1988. Mayflies of the Northern Levant (Insecta: Ephemeroptera). Zoology of the Middle East 2: 89-112. MALZACHER, P. 1996. Genitalmorphologische Merkmale zur Unterscheidung der in Baden- Württemberg vorkommenden Electrogena-Arten (Heptageniidae, Ephemeroptera). Lau- terbornia 25: 81-93. MOUBAYED, Z. 1986. Recherches sur la faunistique, l’écologie et la zoogéographie de trois réseaux hydrographiques du Liban: |’ Assi, le Litani et le Beyrouth. Thèse Doctorat d'Etat, Univ. Toulouse, 496 pp. SAMOCHA, M. 1972. Ephemeroptera of Israel. Thesis, Tel-Aviv University, 112 pp. SARTORI, M. 1992. Mayflies from Israel (Insecta; Ephemeroptera). I.- Heptageniidae, Ephe- merellidae, Leptophlebiidae & Palingeniidae. Revue suisse de Zoologie 99(4): 835-858. REVUE SUISSE DE ZOOLOGIE Tome 106 — Fascicule 4 ZOOLOGIA ET BOTANICA ‘99, Zurich, 17-19 February 1999 (Joint meeting of the Swiss Society of Zoology and the Swiss Society of Botany). ... . BESUCHET, Claude. Psélaphides paléarctiques nouveaux ou méconnus (Cole- Opiera Staphylinidae Bselaphinae)e. 2 are ROIO LEISTIKOW, Andreas. Androdeloscia gen. n., anew genus of South American terrestrial isopods with description of 13 new species (Crustacea: Onis- eideasphılosendae Ds Aare RE LIENHARD, Charles & N. Philip ASHMOLE. Sphaeropsocopsis myrtleae sp. n., a blind subterranean psocid from St Helena (Psocoptera: Sphaeropso- CAS) EL SAI LIO AR CARI I ra ANGELINI, Fernando. New species of Agathidium Panzer from China (Cole- OPiCra e C10dida e ACUM IRINA (RISSA, LOURENCO, Wilson R. Considérations taxonomiques sur le genre Hadogenes Kraepelin, 1894: création des Hadogeninae subfam. n. et description d’une espèce nouvelle de 1’ Angola (Scorpiones, Scorpionidae, Hado- SMIRNE NI O RE CASSOLA, Fabio. Studies on tiger beetles. XCVIII. A new Cylindera from @hubut Argentina, (Coleoptera: (Ciemdelidae) Era ee. SCHWENDINGER, Peter J. & Jochen MARTENS. A taxonomic revision of the family Oncopodidae II. The genus Gnomulus Thorell (Opiliones, Pantatore Ss) N ee IE An Zıcsı, Andras & Csaba CsuzpI. Weitere Angaben zur Regenwurmfauna Frankreichs mit Beschreibung fiinf neuer Arten (Oligochaeta: Lumbri- CICERO RR OR CARPANETO, Giuseppe M. & Roberto MIGNANI. A review of the Himalayan Thorectes (Coleoptera: Geotrupidae), with description of a new species MOMMA OCEMEN TMM GNA RE O IS Hogss, René & Adolf SCHOLL. The identity of Glomeris quadrifasciata C. L. Kochi(DiplopodaxGlomenidae): ran. te eee eee BELFIORE, Carlo & Michel SARTORI. Redescription of Electrogena galileae (Demoulin, 1973) (Ephemeroptera, Heptageniidae)............... Pages 745-787 789-811 813-904 905-912 913-927 929-938 939-943 945-982 983-1003 1005-1012 1013-1024 REVUE SUISSE DE ZOOLOGIE Volume 106 — Number 4 ZOOLOGIA ET BOTANICA ‘99, Zurich, 17-19 February 1999 (Joint meeting of the Swiss Society of Zoology and the Swiss Society of Botany)... .. BESUCHET, Claude. New or poorly known Palaearctic pselaphids (Coleop- (eras StaphylinidaePselaphinae)). u... se aerate LEISTIKOW, Andreas. Androdeloscia gen. n., anew genus of South American terrestrial isopods with description of 13 new species (Crustacea: Onis- Cilea SPhillosciidaey) i tata. ies stk RIE SOR RER LIENHARD, Charles & N. Philip ASHMOLE. Sphaeropsocopsis myrtleae sp. n., a blind subterranean psocid from St Helena (Psocoptera: Sphaeropso- CICALE ANGELINI, Fernando. New species of Agathidium Panzer from China (Cole- OpteraylwerodidaesAcathidimi). ....... ... 2 IRR En NNNREr LOURENCO, Wilson R. Taxonomic considerations on the genus Hadogenes Kraepelin, 1894: proposition of the Hadogeninae subfam. n., and des- cription of a new species from Angola (Scorpiones, Scorpionidae, Hadogennao)g sen in Aa ARA RE CASSOLA, Fabio. Studies on tiger beetles. XCVIII. A new Cylindera from Chubut Arsentma(Coleoptera Cicindelidac) FER eee SCHWENDINGER, Peter J. & Jochen MARTENS. A taxonomic revision of the family Oncopodidae H. The genus Gnomulus Thorell (Opiliones, PANTALONE Re ne a) OE OREO ERNE eR RE Zıcsı, Andras & Csaba Csuzpi. Further Contribution to the earthworm fauna (Oligoschaeta: Lumbricidae) of France, with description of five new SPECIESFANAFONE SUDSPÈCIES. 22. eue RIA IE CARPANETO, Giuseppe M. & Roberto MIGNANI. A review of the Himalayan Thorectes (Coleoptera: Geotrupidae), with description of a new species ombnorthemIndian... „ums spe se ne e ee ERE HoEss, René & Adolf SCHOLL. The identity of Glomeris quadrifasciata €. L. Kochi (Diplopoda:,Glomeridae)..: ..........2....5 Mon. ERO BELFIORE, Carlo & Michel SARTORI. Redescription of Electrogena galileae (Demoulin, 1973) (Ephemeroptera, Heptagentidae)............... Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 745 789 813 905 ONE 929 930, 945 983 PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N°8 1-17 (1908-1926) ..... série Fr. 285.— (prix des fascicules sur demande) REVUEDE PALEOBIOLOGIE® . A "UN RENT RL, Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris)........ par fascicule Fr. COLLEMBOLENFAUNA EUROPAS von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Er. THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION WITH KEYS FOR DETERMINATION byAMINOSEKs)346 pages, 11 lstieuresnmitext; LOTS! a5 a Wa aise Ne PET LIE Fr. CLASSIFICATION OF THE DIPLOPODA par Richard L. HOFFMAN, 237 pages, 1979 LES OISEAUX NICHEURS DU CANTON DE GENEVE par P. GEROUDET, C. GUEX et M. MAIRE 3slpagessnombreusesieantesteltlgures, MOSS Me AE ee Fr. CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ A LA CONNAISSANCE DES ECHINODERMES ACTUELS PawMichelANGOUXG.67 pages, Ilsplanchesy 1989 LR cee tetany Fr. RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) pale LINED IAWCEST et KeMAREDANO2 paces lOO ait Rn Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN; SHELLS AND RADULAS bYANBENERTAWUESTANdKMAREDA 1992 ee Fr. O. SCHMIDT SPONGE CATALOGUE par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992.......... Fr. ATLAS DE RÉPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr. THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST PATRMAVE SEINE Ale Dipases MONS En ee OTO Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE pawlean-@laude!@ATELIEZ AO pages MOIS ASIE RIN Fr. PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr. CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) (Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997 .................... Fr. CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis II) par Henri G. DIRICKx, 188 pages, 1998 ............ Fr. A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Fr. 35.— 35.— 30.— 30.— 30.— 45.— 60.— l'al REATO a EN aad Paty; mi gti Pe MER. 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