Ey | Rust NH ANNALES de la | SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genève tome 110 fascicule 2 2003 E 4 lal GENEVE JUIN 2003 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE REVUE SUISSE DE ZOOLOGIE TOME 110—FASCICULE 2 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 Geneve CHARLES LIENHARD Chargé de recherche au Muséum d’histoire naturelle de Genéve Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis a des experts d’institutions suisses ou étrangères selon le sujet étudié. 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MUSEUM D’HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm SUISSE Fr. 225.— PRIX DE L’ABONNEMENT: (en francs suisses) Les demandes d’abonnement doivent étre adressées a la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse UNION POSTALE Fr, 230.— ANNALES de la SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genéve tome 110 fascicule 2 2003 E 1 EI GENEVE JUIN 2003 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE REVUE SUISSE DE ZOOLOGIE TOME 110—FASCICULE 2 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 Genéve CHARLES LIENHARD Chargé de recherche au Muséum d’histoire naturelle de Geneve Comité de lecture 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, évolution, écologie, éthologie, morphologie et anatomie comparee, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L’ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d’abonnement doivent étre adressées a la rédaction de la Revue suisse de Zoologie, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse REVUE SUISSE DE ZOOLOGIE 110 (2): 247-253; juin 2003 New records of phoretic and soil-living mites from Iran (Acari, Heterostigmata, Scutacaridae) Ernst EBERMANN!, Hamid-reza HAJIQANBAR? & Karim HADDAD IRANI-NEJAD? ! Institut für Zoologie, Karl-Franzens-Universität, Universitätsplatz 2, A-8010 Graz, Austria. E-mail: ernst.ebermann@uni-graz.at 2,3 Department of Plant Protection, Faculty of Agriculture, Tabriz University, Postal Code: 51664, Tabriz, Iran. E-mails: hajiqanbar@ yahoo.com, khaddad@tabrizu.ac.ir New records of phoretic and soil-living mites from Iran (Acari, Heterostigmata, Scutacaridae). - Scutacarus iranicus sp. n. is described from the West-Azarbaijan province (northwestern Iran). The new species was found to be phoretic on the scarabaeid beetle Pleurophorus anatolicus Petr. At the same locality, but in the soil, other scutacarids were found: Scutacarus quadrangularis (Paoli, 1911), Scutacarus serotinus Sevastianov & Chydyrov, 1992, Pygmodispus (Allodispus) latisternus Paoli, 1911 and Pygmodispus (Pygmodispus) calcaratus Paoli, 1911. These species are recorded for the first time from Iran. Key-words: Acari - Scutacaridae - new species - phoresy - scarabaeid beetle - Iran. INTRODUCTION The scutacarid fauna of Iran is almost unknown. In 1970 Mahunka & Rohde described the new species Heterodispus verrucosus and this was the first record of a representative of the mite family Scutacaridae from Iran. Since this record, only a few additional species have been reported (Kamali et al., 2001): These are Imparipes para- picolosimilis Metwaly, 1971, Scutacarus longitarsus (Berlese, 1905), Scutacarus fragariae Rack, 1975 and Scutacarus contiguus Delfinado, Baker & Abbatiello, 1976. Pedobiological collections from sugarbeet-fields in northwestern Iran (by the second author, H. H.) yielded some scutacarid species which are presented here. MATERIAL AND METHODS Locality: Miandoab plain, West-Azarbaijan province (northwestern Iran), sugarbeet fields. Dates of collecting: sample-number 1: April 19, no.2: May 4, no.4: May 15 and May 16, no.5: May 15, no.6: September 14. All soil samples were collected by H. H. in 2000. Mites and beetles were extracted by using Berlese- Tullgren-funnels. Manuscript accepted 01.10.2002 248 E. EBERMANN ET AL. The mites were determinated by the first author (E. E.), the beetles by F.-T. Krell. The material was collected in course of investigations for the master thesis of the second author (H. H.) under the supervision of the third author (K. H.) The following abbreviations are used in the description: ap. = apodeme, Fe = femur, Ge = genu, ITa = length of tarsus, IPrTa = length of pretarsus, PrTa = pretarsus, pstpl = posteriore sternal plate, sol = solenidion, Ta = tarsus, Ti = tibia, TiTa = tibio- tarsus, Tr = trochanter, x= average, = =about the same length, < = shorter than, > = longer than. RESULTS DESCRIPTION OF THE NEW SPECIES Scutacarus iranicus sp. n. (female) Figs 1-3 Material and deposition: 8 ° 2 from sample 6. Holotype specimen and four paratypes in the Muséum d’histoire naturelle Geneve (Switzerland), two paratypes in the Zoologisches Institut und Zoologisches Museum, University of Hamburg (Germany), one paratype in the Acarological Collection, Zoological Museum, College of Agriculture, Tehran University, Karaj (Iran). Diagnosis: Scutacarus iranicus sp. n. is characterized by the comparatively rare feature “setae e and hl tiny“. It shares this feature with little more than 20 species, which have tiny or completely reduced setae e. Scutacarus iranicus sp. n. is most close- ly related to the ,,tackei-species group. Members of this group are Scutacarus tackei Willmann, 1942, S. ellipticus Karafiat, 1959, S. suborbiculatus Rack, 1964 and S. ter- renus Delfinado & Baker, 1976. The most important differences between S. iranicus sp. n. and S. tackei are: iranicus: alveolar canals of setae cl and c2 long, tackei: only in c2 long; iranicus: hl somewhat longer than f, tackei: hl two times longer than f; iranicus: 3b arising far in front of 3a and 3c, tackei: 3a, 3b and 3c nearly in one line; iranicus: ps1 and ps2 < 4a, with only a few barbs, tackei: psl = 4a or psl > 4a, psl and ps2 densely barbed; iran- icus: tip of claw of legl rather blunt, rackei: claw with thin, elongated tip; iranicus: sol wl > gl, tackei: wl = pl; iranicus: tc“TV thick with strong spines, tackei: te“IV thin with fine barbs. The most important differences between S. iranicus and S. ellipticus are: irani- cus: cl, c2 and d distally smooth or sparsely barbed, ellipticus: cl, c2 and d distally densely barbed; iranicus: f slightly thinner than d, e/lipticus: f distinctly thinner than d; iranicus: 4b and 4c being the thickest of all ventral setae, e/lipticus: 4b and 4c very thin; iranicus: psl > ps2, with only a few barbs, ps1 and ps2 < 4a, ellipticus: ps1 = ps2, both setae densely barbed, ps! and ps2 > 4a; iranicus: e and h2 tiny, ellipticus: e and h2 completely reduced; iranicus: claw of leg I large, tip of the claw rather blunt, ellip- ticus: claw small with very thin, elongated tip. The most important differences between S. iranicus and S. suborbiculatus are: iranicus: d > f, suborbiculatus: f > d; iranicus: 3b arising far in front of 3a and 3c, 3a, 3b, 3c shorter than in suborbiculatus, 3c not reaching insertion of 4b, suborbiculatus: 3a, 3b and 3c standing in one line, these setae longer than in iranicus, 3c extending beyond insertions of 4b; iranicus: e and h2 tiny, suborbiculatus: e and h2 completely SCUTACARIDAE FROM IRAN 249 9% 5, Oia FIG. 1 Scutacarus iranicus sp. n. (female holotype) — dorsal view; body length 190 um. reduced; iranicus: tip of claw of leg I rather blunt, suborbiculatus: claw with very thin, elongated tip. The most important differences between S. iranicus and S. terrenus are: iranicus: TiTa I with 4 solenidia, terrenus: TiTa I with 3 solenidia only. SCUTACARIDAE FROM IRAN 251 Description: Body dimensions (measurements in um): Total body length (measured on about one year old microscopic slides) 146 — 190, x = 167 (n = 8), holo- type 190; width pstpl (measured along anterior margin of the plate): 65 — 80, x = 74 (n = 8), holotype 79. Entire surface of the body with tiny pores; cupulae ia and ih roundish. Dorsum (Fig. 1): free margin of tergite C with fine, radiating stripes (not illus- trated in Fig. 1); alveolar canal of all dorsal setae, except e and hl, visible. Dorsal setae smooth or moderately barbed, their relative length: cl >c2eh2,h1 only slightly longer than f, e and h2 tiny. Venter (Fig. 2): ap.1, 2, 3 strongly developed, ap.4 extended, ap.5 reduced. Ventral setae strongly barbed or moderately barbed or smooth, somewhat varying in length. Relative length: la > 1b > 2a or 1b = 2a, 2a = 2b. 2b dagger-shaped, smooth. 3a < 3b < 3c, 3b arising in front of 3a and 3c; 4a < 4b > 4c, 4b standing a short distance in front of 4a; ps1>ps2 >ps3, ps1 and ps2 close to each other. Genital sclerite broader than long, anterior genital sclerite laterally with sclerotized structures. Trichobothrium scl (Fig. 3b): club-shaped, thin-stemmed, with fine scales, outer seta v1 somewhat longer than v2 or reverse. Extremities: Leg I (Fig. 3c): Setal formula: Tr 1, Fe 2, Ge 4, TiTa 16 (4sol), sol 602 < wl > p2 < gl, w2 thin, wl and @2 standing side by side, w1 finger-shaped, @2 thin, pl club-shaped, thin-stemmed; TiTa with large claw, tip of the claw rather blunt. Leg II (Fig. 3d): Setal formula: Tr 1, Fe 3, Ge 3, Ti 4(sol @), Ta 6 (sol w); Ta with 2 claws and pulvillus. Leg III (Fig. 3e): Setal formula: Tr 1, Fe 2, Ge 2, Ti 4 (sol @), Ta 6; Ta with 2 claws and pulvillus. Leg IV (Fig. 3f): Setal formula: Tr 1, Fe 2, Ge 1, TiTa 7; tc° > pv“ > tc“, tc“ with thick spines. Male and larva: Unknown. Bionomics: Specimens of the new species were several times found to be phoretic on the beetle species Pleurophorus anatolicus Petrovitz, 1961 (Scarabaeidae: Aphodiinae: Psammodiini). The mites were clinging to the sutures of the ventral side of the head and the suture between the forelegs and head. OTHER IDENTIFIED SPECIES Pygmodispus (Pygmodispus) calcaratus Paoli, 1911: Sample No. 5. Some records from North America and Eurasia verify the holarctic distribution of this species. Pygmodispus (Allodispus) latisternus Paoli, 1911: Sample No. 4. This species is recorded from Western and Central Europe, as well as from Mongolia and North Africa. Scutacarus quadrangularis (Paoli, 1911): Sample No. 1. This is one of the most common scutacarid species, it is widely distributed in Eurasia and also recorded from Australia and New Zealand. Scutacarus serotinus Sevastianov & Chydyrov, 1992: Sample No.2. Hitherto this species was only known from Turkmenistan. IG. cutacarus iranicus sp. n. (female holotype) — a = trichobothrium, b = leg I (arrows: dors of claw, solenidia @1 and pl), c= leg II, d= leg III, e = leg IV, f = seta tc“ from opposite leg al view IV. SCUTACARIDAE FROM IRAN 253 DISCUSSION The finding of S. iranicus sp. n. on Pleurophorus anatolicus is especially remarkable as there is hardly anything known about associations between Scarabaeids and Scutacarids. Beetles of the family Scarabaeidae have only once been reported as phoretic hosts for Scutacarids: Norton (1973) discovered some phoretic 9 9 of Hetero- dispus sp. on the North American Hermit Flower Beetle (Osmoderma eremicola Knoch). Amongst Coleoptera the family Carabidae is the most thoroughly investigated one. Beetles of this family are known to be used very frequently as phoretic hosts by a high number of scutacarid species, especially those of the genus Archidispus (e.g., Kurosa, 1991). Furthermore, some other beetles‘ families are known to be phoretic hosts (Ebermann, 1988). With the five species recorded in this paper the number of scutacarid species recorded from Iran has been increased to ten. This number most probably still under- estimates the actual number, considering the high number of species already known to occur in Eurasia. The remarkable geographical diversity of Iran and its rich ecological differentiation will deserve more targeted investigations in the future. Soil samples and insects, mainly beetles, ants and other Hymenoptera, can be expected to yield a high number of already described, and also of new species of phoretic and nonphoretic scutacarids. ACKNOWLEDGEMENT We express our sincere thanks to Dr F.-T. Krell, Research Entomologist at The Natural History Museum (Department of Entomology), London, for the determination of the scarabaeid beetles. REFERENCES EBERMANN, E. 1988. /mparipes (Imparipes) pselaphidorum n. sp., a new scutacarid species phoretic upon african beetles (Acari, Scutacaridae; Coleoptera, Pselaphidae). Acarologia 29 (1): 35-42. KAMALI, K., OSTOVAN, H. & ATAMEHR, A. 2001. A catalog of mites & ticks (Acari) of Iran. Scientific Publishing Center of Azad University, Tehran, 192 pp. Kurosa, K. 1991. The Scutacarid mites of Japan. XI. Four new Archidispus associated with ground beetles of the tribes Nebriini, Bembidiini and Patrobini. Japanese Journal of Entomology 59(3): 539-554. MAHUNKA, S. & ROHDE, C. J. jr. 1970. A new scutacarid and some pyemotid mites from Iran. Acarologia 12 (1): 103-105. NORTON, R. A. 1973. Phoretic mites associated with the Hermit Flower Beetle, Osmoderma eremicola Knoch (Coleoptera: Scarabaeidae). The American Midland Naturalist 90 (2): 447-449. SEVASTIANOV, V. D. & CHYDYRov, P. R. 1992. New mite species of the family Scutacaridae (Trombidiformes) from Turkmenistan (in Russian). Vestnik Zoologii 1992 (1): 21-28. REVUE SUISSE DE ZOOLOGIE 110 (2): 255-313; juin 2003 Homalotini del Borneo (Coleoptera, Staphylinidae)* Roberto PACE Via Vittorio Veneto, 13, I-37032 Monteforte d’Alpone (Verona), Italia. Homalotini from Borneo (Coleoptera, Staphylinidae). - The species of the genera Stenomastax Cameron, 1933, Coenonica Kraatz, 1857, Mimopi- salia Pace, 1984, Neosilusa Cameron, 1920, Neocoenonica Cameron, 1950, Neomalota Cameron, 1920 and Linoglossa Kraatz, 1859 are studied for the fauna of Borneo. Stenomastax includes now 21 species (18 species new), Coenonica 31 species (21 species new), Mimopisalia 6 species (4 species new), Neosilusa 4 species (one new) and Linoglossa is recorded for the first time from Borneo with one new species. Based on the study of the types of the species of the five genera already described from Borneo the following new synonyms are proposed: Deralia Cameron is a junior synonym of Coenonica Kraatz and C. soror Cameron, 1933 a junior synonym of C. nigrita Cameron, 1933. New data are given for some already described species. All new species are described, illustrated and compared with simi- lar looking species. Keys to all species of Borneo of the 5 genera mentioned above are presented. Key-words: Coleoptera - Staphylinidae - Aleocharinae - taxonomy - Borneo. INTRODUZIONE I generi della tribù Homalotini Heer, 1839, quali Stenomastax Cameron, 1933, Coenonica Kraatz, 1857, Mimopisalia Pace, 1984, Neosilusa Cameron, 1920 e Neo- coenonica Cameron, 1950 erano finora noti del Borneo. I generi Neomalota Cameron, 1920 e Linoglossa Kraatz, 1859, sono nuovi per questa grande isola. Il genere Stenomastax era rappresentato da tre specie: S. nigrescens (Fauvel, 1905), S. tuber- culicollis (Kraatz, 1859) e S. borneensis Cameron, 1933. Nel presente lavoro questo genere si arricchisce di diciotto nuove specie. Il genere Coenonica nel Borneo era rappresentato da dieci specie: C. vulnerata Bernhauer, 1915, C. abdominalis Cameron, 1930, C. soror Cameron, 1933, C. nigrita Cameron, 1933, C. monticola Cameron, 1933, C. borneensis Cameron, 1933, C. rufiventris Cameron, 1943, C. sarawakensis Cameron, 1943, C. puncticollis Kraatz, 1857 e C, ashei Pace, 1987. Nel presente lavoro questo genere si arricchisce di ventuno nuove specie. Il genere Mimopisalia era rap- presentato da due sole specie: M. borneensis (Cameron, 1933) e M. kinabaluensis * 181° Contributo alla conoscenza delle Aleocharinae. Manoscritto accettato il 15.01.2003 256 R. PACE Pace, 1989. Nel presente lavoro ne sono aggiunte quattro, tutte nuove. All’unica specie del genere Neosilusa, N. borneensis (Cameron, 1928), se ne aggiunge un’altra. N. stric- ticollis Cameron, 1943, non appartiene a questo genere (Pace, 2002). Per il genere Neocoenonica si conferma la presenza di N. antennalis (Cameron, 1936), citata da Hammond (1984), come Drusilla antennalis (Cameron, 1936) (“olim” Astilbus). Le prime Aleocharinae del Borneo sono state raccolte da J.C. Moulton nel 1912. Successivamente furono G.E. Bryant nel 1913-1914 ed E. Mjöberg a contribuire a rac- cogliere nuovo materiale. Le raccolte di H.M. Pendlebury nel 1927 e nel 1929 sono state studiate da Cameron (1930 e 1933). D.H. Kistner e J.M. Pasteel dal 1969 al 1976 descrivono varie specie di Aleocharinae termitofile. Altre raccolte sono state effettuate da M.E. Bacchus nel 1976 e nel 1977-1978 da P.M Hammond, del Museo di Storia Naturale di Londra, e altri membri della Societa Geografica Reale, nel corso della spedizione al Monte Mulu nel Sarawak (Royal Geographic Society Gunung Mulu Expedition). Nel 1990 il collega Guillaume de Rougemont raccolse sul Monte Kina- balu (Borneo) alcune Aleocharinae da me esaminate. Il presente lavoro si basa sulle importantissime e abbondanti raccolte effettuate nelle spedizioni sul Monte Kinabalu e dintorni dal Dr. AlesSmetana di Ottawa, dal Dr. Ivan Löbl e dal Dr. Daniel Burckhardt, entrambi già del museo di Storia Naturale di Ginevra. Sono integrate da alcuni esemplari raccolti dal Prof. Herbert Franz di Mödling (Austria), da una piccola collezione posseduta dal Dr. Volker Assing di Hannover e un esemplare donatomi dall’Ing. Jin Janak di Rtyne nad Bilinou (Repubblica Ceca). Gli olotipi delle nuove specie sono conservati nel Museo di Storia Naturale di Ginevra (MHNG), nel Museo Regionale di Scienze Naturali di Torino (MRSN) e in collezione Franz al Naturhistorisches Museum di Vienna (NHMW). SERIE TIPICHE ESAMINATE, RIGUARDANTI SPECIE DEL BORNEO Stenomastax variventris (Kraatz, 1859) Homalota variventris Kraatz, 1859: 34 266 e 22 2, Ceylon, J. Nietner, sintipi (coll. Kraatz, DEI). Stenomastax tuberculicollis (Kraatz, 1859) Homalota tuberculicollis Kraatz, 1859: 33 1de3 2, Ceylon, J. Nietner, sintipi (coll. Kraatz, DEI). Stenomastax nigrescens (Fauvel, 1905) Homalota nigrescens Fauvel, 1905: 147 1 2, Sumatra, Paja Kombo; 1 de 12, Sukaburmi, sintipi (coll. Fauvel, Bruxelles). Coenonica soror Cameron, 1933 Coenonica soror Cameron, 1933: 354 Holotypus 2, B.N. Borneo, Mt. Kinabalu, Kamborangah, 7200 ft., 23.3.1929, C. soror Cam. TYPE (coll. Cameron, Londra) Coenonica nigrita Cameron, 1933 Coenonica nigrita Cameron, 1933: 354 Holotypus ©, B.N. Borneo, Mt. Kinabalu, Kamborangah, 7500 ft., 23.3.1929, C. ni- grita Cam. TYPE (coll. Cameron, Londra). HOMALOTINI DEL BORNEO 257] Nota. La spermateca degli olotipi di Coenonica soror Cameron, 1933 e di Coenonica nigrita Cameron, 1933 ha forma identica nelle due specie, figg. 178 e 179. Cameron ha descritto due specie differenti probabilmente in base al colore differente del corpo: capo rossiccio in soror, capo nero in nigrita. Tranne queste differenze non se ne osservano altre. Identico è il sistema di reticolazione, uguale il rapporto larghez- za/lunghezza del pronoto e delle elitre. Inoltre le due specie sono state raccolte nello stesso giorno e nella stessa località. Solo la quota altimetrica è di poco differente e nigrita è stata raccolta a quota superiore. Non è una novità che esemplari di una medes- ima specie di Aleocharinae presentino fenomeni di melanismo per le quote altimetriche superiori. In base a questi dati e soprattutto per la forma identica della spermateca è proposta la seguente sinonimia: Coenonica soror Cameron, 1933, Coenonica soror Cameron, 1933: 354, Coenonica nigrita Cameron, 1933: 354, syn. n. La descrizione di Coenonica soror precede quella di Coenonica nigrita nella stessa pagina. Coenonica monticola Cameron, 1933 Coenonica monticola Cameron, 1933: 355 Holotypus ©, B.N. Borneo, Mt. Kinabalu, Pakka, 10,700 ft., 21.Mar.1929, Coenonica monticola Cam, TYPE (coll. Cameron, Londra). Coenonica borneensis Cameron, 1933 Coenonica borneensis Cameron, 1933: 353 Holotypus 2°, B.N. Borneo, Mt. Kinabalu, Lumu Lumu, 5500 ft., 16.4.1929, Coenonica borneensis Cam. TYPE, M. Cameron Bequest BM 1955-147 (coll. Cameron, Londra). Coenonica rufiventris Cameron, 1943 Coenonica rufiventris Cameron, 1943: 41 Holotypus 2, Borneo, Mt. Pais, C. rufiventris Ca., TYPE (coll. Cameron, Londra). Coenonica sarawakensis Cameron, 1943 Coenonica sarawakensis Cameron, 1943: 40 Holotypus 2, Mt. Matang, XII.1913, C. sarawakensis Cam. TYPE (coll. Cameron, Londra). Coenonica vulnerata Bernhauer, 1915 Coenonica vulnerata Bernhauer, 1915: 148 Holotypus d, Mt. Matang, 1000 ft., Sarawak, 3.XII.1913, Moulton (coll. Bernhauer, Chicago). Esaminato da Sawada (1980) che ne ha pubblicato e illustrato edeago e caratteri dis- tintivi. Coenonica ashei Pace, 1987 Coenonica ashei Pace, 1987: 39 Holotypus d, Borneo, Baker, 13449, Coenonica philippina Brnh, det. Bernhauer, Holo- typus Coenonica ashei Pace (coll. Bernhauer, Chicago). Mimopisalia borneensis (Cameron, 1933) Leptusa borneensis Cameron, 1933: 355 Mimopisalia borneensis: Pace, 1984b: 901 Holotypus d, B.N. Mt. Kinabalu, Pakka, 10-200 ft., 21.Mar.1929, Leptusa borneensis Cam. TYPE (coll. Cameron, Londra). Mimopisalia kinabaluensis Pace, 1989 Mimopisalia kinabaluensis Pace, 1989: 6 Holotypus d, Sabah, Mt. Kinabalu, 2600 m, 2.V.1987, leg. Burckhardt & Löbl, (MHNG). 258 R. PACE Apatelieida stricticollis (Cameron, 1943) Neosilusa stricticollis Cameron, 1943: 40 Neosilusa stricticollis: Hammond, 1984: 211 Apatelieida stricticollis: Pace, 2002: 232 Holotypus ©, Borneo, Mt. Poi, 5000 ft., N. stricticollis Cam., TYPE (coll. Cameron, Londra). Neocoenonica antennalis (Cameron, 1936) Astilbus antennalis Cameron, 1936: 18 Drusilla antennalis Hammond, 1984: 209 Neocoenonica antennalis: Cameron, 1950: 94 Holotypus d, The Gap, Selangor, Dr. Cameron, Astilbus antennalis Cam., TYPE, Astilbus antennalis removed to Bolitocharini, tarsi 4,4,5, gen. nov.; 4 es. stessa provenienza; 1 3d, Sumatra, Fort de Koch, leg. Jacobson (coll. Cameron, Londra). Neosilusa ceylonica (Kraatz, 1857) Stenusa ceylonica Kraatz, 1857: 8 Neosilusa ceylonica:Cameron, 1920: 233 Plagiusa ceylonica: Cameron,1939: 167 Neosilusa ceylonica: Blackwelder, 1952: 260 7 22, Ceylon, J. Nietner, Stenusa ceylonica Kraatz, SYNTYPI (coll. Kraatz, DEI). Neosilusa tropica (Bernhauer, 1915) Silusa (Plagiusa) tropica Bernhauer, 1915: 28 Neosilusa tropica: Pace, 1984a: 17 Holotypus 2, Sumatra, Madan Veth, Bolitochara amabilis Motsch., Silusa (Plagiusa) tropica Brnh., TYPUS UNIC.(coll. Bernhauer, Chicago). Neomalota cingulata Cameron, 1920 Neomalota cingulata Cameron, 1920: 245 Holotypus d, Singapore, Bukit Timah, Neomalota cingulata Cam. TYPE (coll. Cameron, Londra). Da questo elenco, purtroppo, mancano tre specie di Homalotini del Borneo quali Stenomastax borneensis Cameron, 1933, Coenonica abdominalis Cameron, 1930 e Neosilusa borneensis Cameron, 1928, non ottenute in esame dal Museo di Storia Naturale di Londra (Dr. M. Brendell), nonostante mia richiesta. La giustificazione € stata la restrizione in corso (1999) per il prestito di materiale tipico. Pertanto tutti 1 riferimenti a queste specie, nel presente lavoro sono basati sulla descrizione originale di Cameron. BEENCO DELLE SPECIE NOTE Stenomastax variventris (Kraatz, 1859) Homalota variventris Kraatz, 1859: 34 Stenomastax variventris Cameron, 1939: 177 1 2, Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg. Burckhardt & Löbl;10 es., Sabah, Mt. Kinabalu, Poring Hot Springs, 480-520 m, 8-9-10.V.1987, leg. A. Smetana; 2 es., Sabah, Mt. Kinabalu, Poring Hot Springs, 510 m, 30.VIII.1988, leg. A. Smetana; 13 es., Sabah, Mt. Kinabalu N.P., Liwagu River, 1490 m, 3.IX.1988, A. Smetana leg.: 2 es., Borneo, Sabah, Crocker Rge. N.P., Hwy. A 3, Km 48 cca, 1000 m, 5.IX.1988, leg. A. Smetana. DISTRIBUZIONE. La Réunion, Sri Lanka, Malaysia, Indonesia, Nuova Guinea e Filippine, Vietnam. Nuova per il Borneo. HOMALOTINI DEL BORNEO 259 Stenomastax tuberculicollis (Kraatz, 1859) Figg. 42-45 Homalota tuberculicollis Kraatz, 1859: 33 Stenomastax tuberculicollis: Cameron, 1939: 177; Pace, 1998: 142 1 de1 ®, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 485 m, 29. VIII.1988, leg. A. Smetana. DISTRIBUZIONE. Sri Lanka, India, Singapore, Cina, Vietnam. Gia nota del Borneo. Questa specie è qui illustrata per la prima volta. Stenomastax nigrescens (Fauvel, 1905) Homalota nigrescens Fauvel, 1905: 147 Stenomastax nigrescens: Cameron, 1939:170; Pace, 1992: 120: Pace, 1998: 142; Pace, 2000: 41 5 es., Sabah, M. Kinabalu N.P., Poring Hot Springs, 485-495 m, 29-30.VIII.1988, leg. A. Smetana. DISTRIBUZIONE. India, Sumatra, Singapore, Malaysia, Giava, Sabah, Cina, Thai- landia, Sumatra, Malaysia, Vietnam. Coenonica soror Cameron, 1933 Coenonica soror Cameron, 1933: 354 54 es., Borneo, Sabah, M. Kinabalu N.P., below Layang Layang, 2600 m, 2-8.V.1987, int. trap, leg. A. Smetana; 35 es., Borneo, Sabah, M. Kinabalu N.P., below Laban Rata, 3150 m, 7.VIIL.1988, leg. A. Smetana. DISTRIBUZIONE. Borneo: Mt. Kinabalu. Coenonica borneensis Cameron, 1933 Coenonica borneensis Cameron, 1933: 353 1 ©, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 485 m, 29.VII.1988, A. Smetana leg. DISTRIBUZIONE. Borneo: Mt. Kinabalu. Mimopisalia borneensis (Cameron, 1933) Leptusa borneensis Cameron, 1933: 355 Mimopisalia borneensis: Pace, 1984b: 899 5 es., Borneo-Sabah, Mt. Kinabalu, 2600 m, 1.V.1987, leg. Burckhardt & Löbl; 5 es., Borneo-Sabah, Mt. Kinabalu, 3150-3200 m, 3.V.1987, leg. Burckhardt & Löbl; 2 es., Sabah, Mt. Kinabalu, 3300 m, Panar Laban, 4.V.1987, leg. Burckhardt & Löbl; 5 3 es., Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 5.V.1987, leg. A. Smetana; 2 es., Sabah, M. Kinabalu N.P., below Laban Rata, 3150 m, 5.V.1987, leg. A. Smetana; 8 es., Sabah, Mt. Kinabalu N.P., above Gunting Lagadan, 3400 m, 6.V.1987, A. Smetana leg. DISTRIBUZIONE. Alte quote del M. Kinabalu. Mimopisalia kinabaluensis Pace, 1989 Mimopisalia kinabaluensis Pace, 1989: 6 54 es., Sabah, Mt. Kinabalu N.P., 2600 m, 1.1V.1987, Burckhardt & Löbl leg.; 13 es., Borneo, Sabah, Mt. Kinabalu N.P., below Layang Layang, 2590 m, 1.V.1987, A. Smetana leg.; 6 es., Borneo, Sabah, Mt. Kinabalu, 2600 m, 2.V.1987, Burckhardt & Löbl leg.; 4 es., Borneo- Sabah, Mt. Kinabalu, 3150-3200 m, 3.V.1987, leg. Burckhardt & Löbl.; 3 es., Sabah, Mt. Kinabalu, 3300 m, Panar Laban, 4.V.1987, leg. Burckhardt & Löbl.; 36 es., Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 5-6.V.1987, leg. A. Smetana. DISTRIBUZIONE. Alte quote del M. Kinabalu. Neosilusa ceylonica (Kraatz, 1857) Stenusa ceylonica Kraatz, 1857: 8 Plagiusa ceylonica: Cameron, 1939: 167 Neosilusa ceylonica: Pace 1984a: 15; Pace 1993: 71 260 R. PACE 1 d e 1 2, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 485 m, 25.VIIL.1988, A. Smetana leg.; 3 dd, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 485 m, 29.VIII.1988, A. Smetana leg.; 1 es., Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 485 m, 30.VIII.1988, A. Smetana leg. DISTRIBUZIONE. Mascarene, India, Malesia, Cina e Giappone. Nuova per il Borneo. Neosilusa tropica (Bernhauer, 1915) Silusa (Plagiusa) tropica Bernhauer, 1915: 28 Neosilusa tropica Pace, 1984a: 17 1 es., Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg. Burckhardt & Löbl; 1 es., Sabah, Poring Hot Springs, 500 m, 13.V.1987, leg. Burckhardt & Löbl. DISTRIBUZIONE. Sumatra, Mascarene (Pace, 1984a) e Madagascar. Nuova per il Borneo. Neomalota cingulata Cameron, 1920 Neomalota cingulata Cameron, 1920: 245 9 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 6.V.1987, Burckhardt & Löbl leg.; 9 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 7.V.1987, Burckhardt & Löbl leg.; 4 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 9.V.1987, Burckhardt & Löbl leg.; 6 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, Burckhardt & Löbl leg.; 10 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 11.V.1987, Burckhardt & Löbl leg.; 1 ea., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 13.V.1987, Burckhardt & Löbl leg. DISTRIBUZIONE. Malaysia, Singapore. Nuova per il Borneo. Neocoenonica antennalis (Cameron, 1936) Astilbus antennalis Cameron, 1936: 18 Drusilla antennalis Hammond, 1984: 209 Neocoenonica antennalis: Cameron, 1950: 94 2 es. Sabah, Crocker Range, 1550-1650 m, 16.V.1987, leg. Burckhardt & Löbl; 9 es., Sabah, Crocker Range, 1200 m, Km 63 r.te Kota Kinabalu-Tambunan, 19.V.1987, leg. Burckhardt & Löbl. DISTRIBUZIONE. Malaysia, Sumatra. Gia nota del Borneo. DESCRIZIONI Stenomastax longacrista sp. n. Figg. 1-5 Holotypus d, Borneo, Sabah, Danum Valley F.S., 85 Km W Lahad Datu, 100 m, 23.VI- 11.1988, D.E. Bright collector (MHNG). Paratypus: 1 ©, stessa provenienza. DESCRIZIONE. Lungh. 1,2 mm. Corpo debolmente lucido e bruno-rossiccio, con elitre brune; antenne bruno-rossicce con antennomero basale giallo-rossiccio; zampe gialle. La punteggiatura del capo é fittissima a punti contigui fra loro. La granulosita del pronoto e delle elitre è indistinta o confusa, quella dell’addome è distinta, rada sugli uroterghi liberi quarto e quinto. Il capo presenta un’impressione occipitale, il pronoto un ampio solco mediano. La reticolazione del pronoto è netta e fine, quella delle elitre e dell’addome è distinta. Edeago figg. 2-3, sesto urotergo libero del d fig. 4, spermateca fig. 5. COMPARAZIONI. Per la taglia corporea minuta e per l’occipite impresso, la nuova specie è ben distinta da quelle note del Borneo, quali S. nigrescens (Fauvel, 1905), HOMALOTINI DEL BORNEO 261 S. variicornis (Kraatz, 1859) e S. borneensis Cameron, 1933. Per la forma dell’edeago non esiste stretta affinita tassonomica con le tre specie citate. Esistono, al contrario, con S. deharvengi Pace, 1990, delle Filippine, ma l’edeago della nuova specie è meno sviluppato, con “crista apicalis” sporgente (bassa in deharvengi) e la spermateca è sub- sferica (ovale in deharvengi). ETIMOLOGIA. Il nome della nuova specie significa “lunga cresta”. È quella prossimale dell’edeago. Stenomastax perpusilla sp. n. Figg. 6-9 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ 1560 m, 23.IV.1987, leg. A. Smetana, (MHNG). Paratypus: 1 2, Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1500 m, 16.V.1987, A. Smetana leg. DESCRIZIONE. Lungh. 1,7 mm. Avancorpo opaco, addome lucido. Corpo bruno, con pigidio bruno-rossiccio; antenne rossicce con l’antennomero basale e l'undicesimo giallo-rossicci; zampe giallo-rossicce. La punteggiatura del capo è composta di punti tra loro contigui. La granulosità del pronoto è confusa, quella delle elitre è superficiale e quella dell’addome è distinta. La reticolazione del pronoto è evidente, quella delle elitre è svanita e quella dell’addome è distinta. Edeago figg. 7-8, sesto urotergo libero del à fig. 9. COMPARAZIONI. Per la forma dell’edeago, la nuova specie è affine a S. /on- gacrista sp. n. sopra descritta, ma l’undicesimo antennomero è giallo-rossiccio (bruno- rossiccio in longacrista) e l’edeago è dilatato in visione ventrale. Quest'ultimo carat- tere non si riscontra nemmeno in S. deharvengi Pace, 1990, delle Filippine, un’altra specie sua affine. ETIMOLOGIA. Il nome della nuova specie significa “piccolissima” e deriva dalla ridotta taglia corporea. Stenomastax bitruncata sp. n. Figg. 10-14 Holotypus d, Sabah, Mt. Kinabalu, 1750 m, 21.1V.1987, leg. Burckhardt & Löbl (MHNG). Paratypi: 2 9 9, Sabah, Poring Hot Springs, 600 m, Bat Cave, 10.V.1987, leg. Burckhardt & Löbl. DESCRIZIONE. Lungh. 1,8 mm. Corpo giallo-rossiccio, con capo, elitre e quarto urite libero bruno-rossicci; antenne bruno-rossicce con i due antennomeri basali e l’undicesimo giallo-rossicci; zampe gialle. La punteggiatura del capo è fitta e confusa. La granulosità del pronoto è indistinta, quella delle elitre è confusa e quella degli uroterghi distinta. La reticolazione del capo e del pronoto è netta e quella delle elitre e dell’addome è distinta. Edeago figg. 11-12, sesto urotergo libero del d fig. 14, sper- mateca fig. 13. COMPARAZIONI. Per la parte dilatata dell’edeago, in visione ventrale, la nuova specie potrebbe essere tassonomicamente affine a S perpusilla sp. n. sopra descritta, ma la forma del margine posteriore del sesto urotergo libero del d separa nettamente le due specie. ETIMOLOGIA. La nuova specie prende nome di “tronca due volte” a motivo della presenza di due lobi tronchi al margine posteriore del sesto urotergo libero del d. 262 R. PACE FIGG. 1-13 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 1-5: Stenmastax longacrista sp. n.; 6-9: Stenomastax perpusilla sp. n.; 10-13: Stenomastax bitrun- cata Sp. n. Stenomastax pseudoliwaguensis sp. n. Figg. 15-17 Holotypus ¢, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 30.IV.1987, leg. A. Smetana (MHNG). HOMALOTINI DEL BORNEO 263 DESCRIZIONE. Lungh. 1,9 mm. Corpo debolmente lucido e rossiccio; antenne rossicce con i due antennomeri basali giallo-rossicci; zampe gialle. La punteggiatura del capo è fitta. Solco frontale profondo. La granulosità del pronoto e delle elitre è in- distinta, quella dell’addome è ben visibile. Il pronoto ha un largo solco mediano. Su tutto il corpo la reticolazione è distinta. Edeago figg. 16-17. COMPARAZIONI. L’edeago della nuova specie è simile a quello di S. variicornis (Kraatz, 1859), dello Sri Lanka. Per questo motivo è possibile un’ affinità tassonomica tra le due specie. Ma mentre S. variicornis presenta il tubulo interno dell’edeago sottile e lungo, quello della nuova specie è largo, robusto e corto. Inoltre il quarto an- tennomero è trasverso nella nuova specie e largo quanto lungo in S. variicornis . ETIMOLOGIA. Il nome della nuova specie significa ‘falsa liwaguensis”. Steno- mastax liwaguensis è la nuova specie sotto descritta, sua affine per la forma dell’edeago. Stenomastax liwaguensis sp. n. Figg. 18-21 Holotypus ¢, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 25.IV.1987, leg. A. Smetana (MHNG). Paratypi: 8 es., stessa provenienza ; 1 2, Sabah, Poring Hot Springs, 600 m, Bat Cave, 10.V.1987, leg. Burckhardt & Löbl; 1 6, Borneo-Sabah, Mt. Kinabalu,1550 m, 23.IV.1987, leg. Burckhardt & Löbl; 1 & e 2 es., Borneo, Sabah, Mt. Kinabalu N.P.,1750 m, 27.IV.1987, Burck- hardt & Löbl leg.; 1 es., Borneo-Sabah, Mt. Kinabalu, 1580 m, 27.IV.1987, leg. Burckhardt & Löbl; 10 es., Borneo-Sabah, Mt. Kinabalu, 1550 m, 29.IV.1987, leg. Burckhardt & Löbl; 3 es., Borneo-Sabah, Mt. Kinabalu, 1500 m, 30.1V.1987, leg. Burckhardt & Löbl; 1 2, Sabah, Crocker Range, 1550-1650 m, 16.V.1987, leg. Burckhardt & Löbl; 3 es. Sabah, Crocker Range, 1600 m, Km 51 rte Kota Kinabalu-Tambunan, 18.V.1987, leg. Burckhardt & Löbl; 1 ¢, Borneo-Sabah, Mt. Kinabalu,1500 m, 21.V.1987, leg. Burckhardt & Löbl; 30 es., Borneo-Sabah, Mt. Kina- balu,1150-1900 m, 21-23-24.V.1987, leg. Burckhardt & Löbl; 5 es., Borneo-Sabah, Mt. Kina- balu Nat. Pk., HQ Silau-Silau Tr., 1560 m, 3.VIIL.1988, leg. A. Smetana; 32 es., Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1490 m, 10.VIII.1988, leg. A. Smetana; 28 es., Bor- neo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1520 m, 11.VIII.1988, leg. A. Smetana. DESCRIZIONE. Lungh. 1,8 mm. Corpo debolmente lucido e rossiccio, con capo, elitre e quarto urite libero bruni; antenne brune con i tre antennomeri basali e l’undice- simo rossicci; zampe gialle. La punteggiatura dell’avancorpo é fittissima. La granu- losita dell’addome é distinta. Una reticolazione superficiale copre la superficie degli uroterghi. Edeago figg. 19-20, spermateca fig. 21. COMPARAZIONI. La nuova specie, per la forma dell’edeago, è affine a S. pseu- doliwaguensis sp. n. sopra descritta e a S. variventris (Kraatz, 1859) dello Sri Lanka. Si distingue da entrambe per la presenza di un robusto sviluppo dell’armatura interna dell’edeago in cui non é distinguibile un tubulo, presente nelle due specie a confronto. L’apice dell’edeago acutissimo e la notevole lunghezza dell’undicesimo antennomero permettono di distinguere ulteriormente la nuova specie da esse. ETIMOLOGIA. La nuova specie prende nome dalla sua localita tipica, Liwagu. Stenomastax ubahensis sp. n. Figg. 22-24 Holotypus 4, Borneo-Sabah, Mt. Kinabalu N.P., Summit Trail Pondok Ubah, 2050 m, 26.IV.1987, leg. A. Smetana, (MHNG). DESCRIZIONE. Lungh. 1,9 mm. Corpo lucido e giallo-rossiccio, con capo ed elitre rossicci; antenne giallo-rossicce; zampe gialle. Il capo presenta un solco discale 264 R. PACE se Ir aa fa ian: Fico. 14-24 Sesto urotergo libero del d, habitus, edeago in visione laterale e ventrale e spermateca. 14: Stenomastax bitruncata sp. n.; 15-17: Stenomastax pseudoliwaguensis sp. n.; 18-21: Steno- mastax liwaguensis sp. n.; 22-24: Stenomastax ubahensis sp. n. HOMALOTINI DEL BORNEO 265 e punteggiatura fittissima e poco profonda. Quella del pronoto è confusa. La granu- losità delle elitre e dell’addome è saliente. Sul corpo la reticolazione è distinta. Edeago figg. 23-24. COMPARAZIONI. Per la forma dell’edeago, la nuova specie si mostra affine a S. pseudoliwaguensis sp. n., sopra descritta. Ma la nuova specie ha apice dell’edeago, in visione laterale, assai largo e non acuto. Inoltre la nuova specie ha pronoto netta- mente trasverso, mentre quello di pseudoliwaguensis lo è appena. ETIMOLOGIA. La nuova specie prende nome dalla sua località tipica Ubah. Stenomastax pugiofora sp. n. Figg. 25-29 Holotypus d, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 485 m, 29.VIII.1988, leg. A. Smetana (MHNG). Paratypus: 1 2, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 495 m, 30. VIII.1988, leg. A. Smetana. DESCRIZIONE. Lungh. 1,9 mm. Corpo lucido e bruno; antenne brune con anten- nomero basale giallo-bruno; zampe gialle. Il capo presenta punteggiatura fitta e molto superficiale e un debole solco mediano. La granulosità del pronoto è indistinta, quella delle elitre è assai poco distinta e quella dell’addome è molto saliente sui tre uroterghi basali e superficiale sui restanti. La reticolazione del capo e dell’addome è distinta, quella del pronoto è netta e quella delle elitre è superficiale. Edeago figg. 26-27, sesto urotergo libero del à fig. 29, spermateca fig. 28. COMPARAZIONI. Per la forma dell’edeago, la nuova specie è sicuramente affine a S. platygaster (Kraatz, 1859) (tipi da me esaminati), a larga diffusione in Oriente. Se ne distingue essenzialmente per il margine posteriore del sesto urotergo libero del à che possiede un larghissimo lobo mediano limitato da incavature superficiali (lobo mediano stretto, limitato da profonda incavatura in platygaster). Questo caratteristico margine posteriore del sesto urotergo libero del & della nuova specie si riscontra pure in S. diogenes Pace, 1998, della Cina, ma questa specie non ha l’apice dell’edeago strettissimo e prolungato come quello della nuova specie. ETIMOLOGIA. Il nome della nuova specie significa “portatrice di pugnale” perché una parte dell’armatura interna dell’edeago ha forma di pugnale. Stenomastax maculiventris sp. n. Figg. 30-32 Holotypus d, Sabah, Mt. Kinabalu,1750 m, 27.IV.1987, leg. Burckhardt & Löbl (MHNG). DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e giallo-rossiccio, con elitre brune e quarto urotergo libero bruno-rossiccio; antenne giallo-rossiccio; zampe gialle. La punteggiatura del capo è fittissima. La granulosita del pronoto e delle elitre è indistinta, quella dell’addome è saliente. La reticolazione è distinta su tutto il corpo. Edeago figg. 31-32. COMPARAZIONI. Per la forma dell’edeago, la nuova specie si pone tassonomica- mente vicino a S. platygaster (Kraatz, 1859) (tipi da me esaminati), a larga diffusione in Oriente. Se ne distingue, oltre per il colore nettamente differente del corpo, per l’apice dell’edeago più robusto in visione laterale e non protratto, in visione ventrale. ETIMOLOGIA. Per la presenza di un anello bruno sull’addome giallo-rossiccio, la nuova specie è chiamata “addome macchiato”. R. PACE 266 0.1 mm 0.1 mm FIGG. 25-32 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 25-29: Stenomastax pugiofora sp. n.; 30-32: Stenomastax maculiventris sp. n. HOMALOTINI DEL BORNEO 267 Stenomastax hawarensis sp. n. Figg. 33-36 Holotypus d, Borneo, Hawar, VI.1998, leg. Hlavac (MRSN). DESCRIZIONE. Lungh. 2,1 mm. Pronoto molto opaco, resto del corpo lucido. Capo ed elitre nero-bruni, pronoto bruno-rossiccio, addome giallo-rossiccio con gli uroterghi liberi terzo e quarto rossicci; antenne brune con i due antennomeri basali e l’apice dell’undicesimo bruno-rossicci; zampe gialle. La punteggiatura del capo è ombelicata fittissima e netta, tranne sulle tempie dove è confusa nella reticolazione vig- orosa, come quella del pronoto. Largo e profondo è il solco mediano del pronoto. La granulosità delle elitre e dei tre uroterghi basali è saliente, quella sugli uroterghi liberi quarto e quinto è superficiale. La reticolazione del capo è netta, quella delle elitre è superficiale e quella dell’addome è distinta, a maglie poligonali irregolari. Edeago figg. 34-35, margine posteriore del sesto urotergo libero del à fig. 36. COMPARAZIONI. La nuova specie presenta edeago simile a quello di S. baliensis Pace, 1986, dell’isola di Bali. Se ne distingue per un evidente tubulo interno dell’edeago, non distinto in baliensis, e per il margine posteriore del sesto urotergo libero del 4 con cinque lobi tra due spine laterali e non tre lobi tra due spine, come in baliensis. ETIMOLOGIA. La nuova specie prende nome dalla sua località tipica Hawar. Stenomastax sericina sp. n. Figg. 37-39 Holotypus d, Sabah, Poring Hot Springs, 500 m, 11.V.1987, leg. Burckhardt & Löbl (MHNG). DESCRIZIONE. Lungh. 1,6 mm. Avancorpo opaco, addome lucido. Corpo bruno, con pronoto rossiccio e addome giallo-rossiccio, con urotergo libero quarto bruno-ross- iccio; antenne rossicce con l’antennomero basale e l’undicesimo giallo-rossicci; zampe gialle. La punteggiatura del capo é distinta e fittissima. La granulosita del pronoto e delle elitre è fittissima, fine e confusa, quella dell’addome è distinta. La reticolazione delle elitre e dell’addome è distinta. Il pronoto presenta un largo solco mediano. Edeago figg. 38-39. ETIMOLOGIA. La pubescenza del corpo, fine come seta, dà il nome di “simile a seta” alla nuova specie. COMPARAZIONI. La nuova specie per l’apice del suo edeago, flesso al lato ven- trale, appartiene al gruppo di specie a cui fa capo S. cribrum (Fauvel, 1878) della Nuova Guinea, ma diffusa anche a Palawan e a Hong Kong. A questo gruppi appartiene anche S. raptoria Pace, 1998 della Cina e S. celebensis Pace, 1986, di Celebes. La nuova specie si distingue da queste specie per i caratteri dati nella seguente chiave. 1 In visione laterale, tra l’apice dell’edeago e la “crista apicalis”, il profi- lo ventrale è concavo; margine posteriore del sesto urotergo libero del 4 con sei lunghi denti tra le spine laterali. Lungh. 3,0 mm. Nuova Guinea, Cina Eiippiner®3rr A Mate etanolo do. S. cribrum (Fauvel) - In visione laterale, tra l’apice dell’edeago e la “crista apicalis”, il profilo ventrale è rettilineo; margine posteriore del sesto urotergo libero del 3 SENDEN. RT ROUE OL. RAS RO SORIA es RE OME AR 2 268 R. PACE 2 In visione laterale, intercapedine apicale dell’edeago stretta............. 3 - In visione laterale, intercapedine apicale dell’edeago larga. Lungh. ROH OMERO S. sericina Sp. n. 3 In visione laterale, intercapedine apicale dell’edeago molto più lunga che larga; apice dell’edeago strettissimo, in visione ventrale. Lungh. DIMMI un S. raptoria Pace - In visione laterale, intercapedine apicale dell’edeago corta e stretta; apice dell’edeago non prolungato e stretto, in visione ventrale. Lungh. POIMMACele DES EN Re e CINI ee ae S. celebensis Pace Stenomastax esuriens sp. n. Figg. 40-41 Holotypus © , Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ 1560-1660 m, 24.IV.1987, leg. A. Smetana, (MHNG). Paratypi: 1 2, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ 1495 m, 12.VIII.1988, leg. A. Smetana; 1 2, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu River, 1500 m, 1.IX.1988, A. Smetana leg. DESCRIZIONE. Lungh. 1,8 mm. Pronoto molto opaco, resto del corpo lucido. Corpo bruno con pronoto rossiccio e addome giallo-rossiccio; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci; zampe gialle. La punteggiatura del capo è fittissima, quella del pronoto è indistinta. La granulosità delle elitre è fittissima. I due uroterghi basali presentano granuli allineati trasversalmente presso il solco basale. La reticolazione del pronoto e dell’addome è distinta. Spermateca fig. 41. ETIMOLOGIA. Il nome della nuova specie significa “affamata”. COMPARAZIONI. Per la forma della spermateca, la nuova specie si colloca tasso- nomicamente vicino a S. parallela Cameron, 1941, delle Filippine (holotypus 2 da me esaminato). Se ne distingue per i caratteri dati nella seguente chiave. i Fossette basali del pronoto strette; elitre giallo-rossicce; base del bulbo distale della spermateca, larga ed emisferica. Lungh. 1,8 mm. Filippine big Sig 6.6.0 8G Sa RE Ie OR PONI Auer S. parallela Cameron - Fossette basali del pronoto larghe; elitre brune; base del bulbo distale della spermateca, stretta e troncoconica. Lungh. 1,8 mm. Borneo En S. esuriens Sp. n. Stenomastax terminicornis sp. n. Figg. 46-47 Holotypus 2, Borneo-Sabah, M. Kinabalu N.P., Summit Trail, 1890 m, (data mancante), leg. A. Smetana, (MHNG). DESCRIZIONE. Lungh. 1,9 mm. Corpo lucido e bruno-rossiccio, con elitre brune e addome rossiccio, con quarto urite libero bruno; antenne brune, con l’antennomero basale rossiccio, il nono e il decimo rossicci e l’undicesimo giallo; zampe giallo- rossicce. La punteggiatura del capo e delle elitre è fittissima e superficiale. La granu- losità del pronoto è confusa nella reticolazione, quella dell’addome è fitta. La retico- lazione del pronoto è evidente, quella dell'addome è estremamente superficiale. Spermateca fig. 47. COMPARAZIONI. La nuova specie presenta il quarto antennomero più lungo che largo, come in S. borneensis Cameron, 1933, pure del Borneo. Ma questa specie HOMALOTINI DEL BORNEO 269 Fico. 33-41 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 33-36: Stenomastax hawarensis sp. n.; 37-39: Stenomastax sericina sp. n.; 40-41: Stenomastax esuriens sp. n. 270 R. PACE presenta antenne brune con l’undicesimo antennomero giallo e non antenne intera- mente giallo-rossicce come in borneensis. Inoltre il capo della nuova specie è bruno- rossiccio e non nero come in borneensis. ETIMOLOGIA. Il nome della nuova specie mette in risalto il suo carattere piü distintivo: il colore giallo dell’undicesimo antennomero, in contrasto con i contigui antennomeri rossicci e bruno-rossicci. Stenomastax pseudonigrescens sp. n. Figg. 48-51 Holotypus d, Sabah, Borneo, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 485 m, 29. VI- 11.1988, leg. A. Smetana (MHNG). Paratypus: 1 2, stessa provenienza. DESCRIZIONE. Lungh. 2,0 mm. Corpo debolmente lucido e bruno-rossiccio, con terzo posteriore delle elitre bruno e addome giallo-rossiccio con il quarto urite libero e la base del quinto bruni; antenne giallo-rossicce; zampe gialle. La punteggiatura del capo e del pronoto è distinta. La granulosita delle elitre è confusa nella distinta retico- lazione. La reticolazione dell’addome è evidente come la sua granulosità. Il capo e il pronoto presentano reticolazione distinta. Spermateca fig. 48, edeago figg. 50-51. COMPARAZIONI. La nuova specie presenta elitre giallo-rossicce come quelle di S. nigrescens (Fauvel, 1905), dell’ India, Bengala, Sumatra e Malesia. Ma quest’ultima specie ha capo e pronoto neri. Per la forma dell’edeago la nuova specie si avvicina tassonomicamente maggiormente a S. cribrum (Fauvel, 1878), della Nuova Guinea, che a nigrescens. Ma la nuova specie, oltre a presentare sei lunghi denti mediani al margine posteriore del sesto urotergo libero del d, presenta l’intercapedine apicale dell’edeago nettamente più lunga e stretta (larga e corta in cribrum). ETIMOLOGIA. Il nome della nuova specie significa “falsa nigrescens”. Stenomastax sphaeratheca sp. n. Figg. 52-53 Holotypus ©, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ Liwagu Riv. trail, 1500-1550 m, 27.IV.1987, A. Smetana leg. (MHNG). DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e bruno-rossiccio, con elitre brune e addome giallo-rossiccio con quarto urite libero rossiccio; antenne giallo-rossicce con undicesimo antennomero giallo: zampe gialle. La punteggiatura del capo è fittissima. La granulosità del pronoto, elitre e addome è fine. Ben visibile è la reticolazione del pronoto e delle elitre. Spermateca fig. 53. COMPARAZIONI. Per la forma della spermateca, la nuova specie sembra affine a S. cribrum (Fauvel, 1878). Ma la dimensione della spermateca è nettamente differente, minuscola e non di grande sviluppo come quella di S. cribrum. Inoltre, nella nuova specie mancano le lunghe setole laterali isolate sulle tibie, pronoto, elitre e addome, presenti in S. cribrum. ETIMOLOGIA. Il nome della nuova specie significa “spermateca sferica”. Stenomastax muluensis sp. n. Figg. 54-55 Holotypus ?, Borneo, Sarawak, Mulu Nat. Park, leg H. Franz (NHMW). DESCRIZIONE. Lungh. 1,7 mm. Avancorpo debolmente opaco, addome lucido. Corpo bruno-rossiccio, con addome giallo-rossiccio; antenne brune con i due anten- HOMALOTINI DEL BORNEO DITA FicG. 42-49 Habitus, edeago in visione laterale e ventrale e spermateca. 42-45: Stenomastax tuberculicollis (Kraatz) del Mt. Kinabalu; 46-47: Stenomastax terminicornis sp. n.; 48-49: Stenomastax pseu- donigrescens sp. n. DIR. R. PACE nomeri basali bruno-rossicci; zampe giallo-rossicce. La punteggiatura dell’avancorpo è fittissima e distinta. La granulosità dell’addome è superficiale. Assente è la retico- lazione dell’addome. Spermateca fig. 55. COMPARAZIONI. La nuova specie, per la forma della spermateca, sembra affine a S. cribrum (Fauvel, 1878). Se ne distingue per la presenza di strette spire prossimali, assai ampie in cribrum. ETIMOLOGIA. La nuova specie prende nome dal Parco Nazionale del Monte Mulu, sua località tipica. Stenomastax myllaenatheca sp. n. Figg. 56-57 Holotypus 2, Mt. Kinabalu, 1580 m, 27.1V.1987, Burckhardt & Löbl leg. (MHNG). DESCRIZIONE. Lungh.1,6 mm. Corpo lucido e giallo-bruno, con addome giallo sporco; antenne brune con i due antennomeri basali e l’undicesimo rossicci, zampe gialle. La punteggiatura del capo è netta e fittissima. La granulosità del pronoto e dell’addome è fine e quella delle elitre è fitta. La reticolazione del pronoto è distinta, quella dell’addome è superficiale. Il solco mediano del pronoto è bruscamente allargato all’indietro. Spermateca fig. 57. COMPARAZIONI. Una spermateca che presenta numerose spire prossimali come quella della nuova specie, si osserva anche in S. nigrescens (Fauvel, 1905). Ma questa specie ha il bulbo distale di quest’organo fortemente oblungo. ETIMOLOGIA. La forma della spermateca è simile a quella di alcune specie del genere Myllaena. Per questo motivo la nuova specie è chiamata “spermateca di Myllaena”. Stenomastax terminalis sp. n. Figg. 58-59 Holotypus 9, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Riv., 1500 m, 25.1V.1987, A. Smetana leg. (MHNG). Paratypi: 2 2 2, Borneo, Sabah, Mt. Kinabalu, 1900 m, 26.1V.1987, Burckhardt & Löbl leg. DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e giallo-bruno; antenne brune con i due antennomeri basali rossicci e i terminali dal nono gialli; zampe gialle. La punteggiatura del capo € ombelicata e fittissima. La granulosita del pronoto e delle elitre è confusa nella netta reticolazione, quella dell’addome è distinta. Spermateca fig 59; COMPARAZIONI. Per la forma della spermateca, la nuova specie si pone tasso- nomicamente vicino a S. myllaenatheca sp. n., sopra descritta e a S. nigrescens (Fauvel, 1905). Se ne distingue per avere i tre antennomeri apicali gialli e per il bulbo distale della spermateca nettamente meno sviluppato di quello di entrambe. ETIMOLOGIA. Il nome “terminale” della nuova specie, allude agli antennomeri apicali di colore nettamente differente da quello dei precedenti. Stenomastax fallax sp. n. Figg. 60-61 Holotypus 2, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480 m, 15.V.1987, leg. A. Smetana (MHNG). DESCRIZIONE. Lungh. 1,9 mm. Corpo lucido e giallo-rossiccio, con capo ros- siccio ed elitre bruno-rossicce; antenne bruno-rossicce con i due antennomeri basali HOMALOTINI DEL BORNEO 273 FicG. 50-57 Edeago in visione laterale e ventrale, habitus e spermateca. 50-51: Stenomastax pseudoni- grescens sp. n.; 52-53: Stenomastax sphaeratheca sp. n.; 54-55: Stenomastax muluensis sp. n.; 56-57: Stenomastax myllaenatheca sp. n. rossicci e apice dell’undicesimo gialli; zampe gialle. La punteggiatura del capo e la granulosità del pronoto sono indistinte. La granulosità delle elitre è poco distinta e quella dell’addome è fine. La reticolazione del capo è netta, quella del pronoto è vigorosa e quella delle elitre è distinta. Spermateca fig. 61. COMPARAZIONI. La nuova specie, per la forma del pronoto, più stretto in avanti che all’indietro, è specie unica nel genere. L’esame della forma della ligula e altre parti 274 R. PACE boccali, oltre a quello della formula tarsale, portano ad attribuire la nuova specie a Stenomastax. ETIMOLOGIA. Il nome della nuova specie significa “ingannevole” perché per la forma del pronoto essa poteva essere attribuita al genere Placusa, ma l’esame delle par- ti boccali esclude tale attribuzione. CHIAVE DEI 6 6 DELLE SPECIE DEL BORNEO DEL GENERE STENOMASTAX 1 10 Quarto antennomero più lungo che largo o lungo quanto largo; decimo antennomero NONO Poco H[aSVEISO' ..... .. 2.2... 2.2... = E 2 Ouarto a/decimo’antennomero trasversi.... ..... 0.020020. 00 Se 3 Quarto antennomero più lungo che largo; elitre brune; tubulo interno dell’edeago corto e robusto; bulbo distale della spermateca molto oblun- go, con parte prossimale descrivente numerose e strette spire. Lungh. 2,4 mmsDallIndiaral’Borneomez. Er... IR S. nigrescens (Fauvel) Quarto antennomero lungo quanto largo; elitre giallo-brune; tubulo interno dell’edeago lungo e sottile; bulbo distale della spermateca sub- sferico, con parte prossimale descrivente poche e larghe spire. Lungh. [FOimms Dalle Mascareneal/Bomeoe ee S. variventris (Kraatz) Parte distale delle antenne. unicolore:...+..... =. nae ole ep RER 7 Parte distalexdelle antenne bicolore . =. ra, 22 SENTE 4 Corpo unicolore bruno, con pigidio bruno-rossiccio. Lungh. 1,7 mm. BS ORME Oe eS ne Sle coer Moet ee S. perpusilla sp. n. Eorpo,bicolore brunore siallo=rossiccio:. ..... .... .)........ A 5) Addome unicolore giallo-rossiccio. Lungh. 1,6 mm. Borneo . S. sericina sp. n. Addome giallo-rossiccio, con fascia posteriore bruna .................. 6 Undicesimo antennomero più lungo dei tre precedenti riuniti; largo solco mediano del pronoto; sesto urotergo libero del 4 senza lobi e spine al margine posteriore. Lungh. 1,8 mm. Borneo........ S. liwaguensis sp. n. Undicesimo antennomero più corto dei tre precedenti riuniti; impres- sione mediana posteriore del pronoto; sesto urotergo libero del d con due lobi tronchi e varie spine al margine posteriore. Lungh. 1,8 mm. BONE RR n n i II S. bitruncata sp. n. Eorpo/unicolorebrunol/orossiccio.. i RE CNN 8 Corpobicolore i. Laine ee LA TE 9 Corpo bruno, con pronoto nettamente trasverso e con lunghe setole iso- late laterali sulle tibie, pronoto e addome. Lungh. 1,9 mm. Borneo soa ee cas ne de Ce te ee CES S. pugiofera sp. n. Corpo rossiccio, con pronoto appena trasverso; assenza di lunghe setole isolate ai lati del corpo. Lungh. 1,8 mm. Borneo . . S. pseudoliwaguensis sp. n. Antenne brune con base rossiccia o giallo-rossiccia ... ... . 2.2.22 meer 10 Antenne unicolori'giallo-rossiecen. 4... MATE Sans SINE Por RER EE 12 Punteggiatura del capo indistinta; elitre giallo-rossicce; margine posteri- ore del sesto urotergo libero del d senza denti o lobi. Lungh. 2,1 mm. Sri Lanka, India, Cina, Vietnam, Singapore, Giava, Borneo SCESE Cds ail col a NE TRIS ehe es LI SI S. tuberculicollis (Kraatz) 11 HOMALOTINI DEL BORNEO DIS Punteggiatura del capo da distinta a forte; elitre brune; margine pos- teriore del sesto urotergo libero del 4 con denti e lobi................. 11 Capo, pronoto e addome bruno-rossicci; margine posteriore del sesto urotergo libero del d con quattro lobi semicircolari tra le spine laterali sınussatesl.unsch!al 21mm rer e S. longacrista sp. n. Solo il pronoto bruno-rossiccio, il capo è nero-bruno; margine posteri- ore del sesto urotergo libero del d con cinque lobi allungati tra le acute spineslaterali. lungh. 2,1lımm. Bomeo. ee e S. hawarensis sp. n. Addome unicolore giallo-rossiccio; un solco discale del capo. Lungh. e9EmmtBormeor Hie ne ore ner LE S. ubahensis sp. n. Addome bicolore giallo-rossiccio, fascia bruna o bruno-rossiccia........ 13 Capo e pronoto giallo-rossiccio; tubulo interno dell’edeago forte. lEunch®l.SımmY Bomeo Au. a We NE S. maculiventris sp. n. Capo e pronoto bruno-rossicci; tubulo interno dell’edeago sottile. Eunehg2.05mm#Borneor.2 is are S. pseudonigrescens sp. n. CHIAVE DELLE 2 £ DELLE SPECIE DEL BORNEO DEL GENERE STENOMASTAX 1 Quarto antennomero piu lungo che largo o lungo quanto largo; decimo AMEINOMETOMONOPOCOLTASVEISORE: MAN EN N 2 Quartorardecimo antennomerorttasyersi nn i ur... 7 Ouarto,antennomero,piulungoichellargor zu... nenn 3 @uartorantennomeroduneorquantoilarso we TI. nn a ee a. a 5 Antenne unicolori bruno-giallicce; bulbo distale della spermateca molto allungato. Lungh. 2,4 mm. Dall’India al Borneo...... S. nigrescens (Fauvel) Parte apicale delle antenne bicolore; bulbo distale della spermateca poco alll aN atone E EINE ULL Sa RETENU AN ER aa) 4 Decimo antennomero lungo quanto largo; parte prossimale della sperma- teca descrivente due sinuosita. Lungh. 1,9 mm, Borneo . S. terminicornis sp. n. Decimo antennomero trasverso; parte prossimale della spermateca de- scrivente numerose spire. Lungh. 1,8 mm, Borneo..... S. terminalis sp. n. Parte apicale delle antenne bicolore; solco mediano del pronoto solo sul- la metà posteriore. Lungh. 1,8 mm. Borneo......... S. sphaeratheca sp. n. Parte apicale delle antenne unicolore; solco mediano del pronoto dal MARINE antenoreralspostemiorer . u u O 6 Capo e pronoto bruno-rossicci. Lungh. 1,9 mm. Dalle Mascarene al BONO RA SRO ORO ann S. variventris (Kraatz) Capo nero; pronoto giallo-rossiccio . Lungh. 2,0 mm. Borneo DO CR PR tea PI RN CR S. borneensis Cameron Antenne unicolori giallo-rossicce. Lungh. 2,0 mm. Borneo 06 RE ale eM er dns ca Ce as te E: S. pseudonigrescens sp. n. Antenne bicolori, con apice e base giallo-rossicci o gialli............... 8 Corpo unicolore bruno; lunghissime setole laterali isolate del corpo e dellesitibresEunsh=2,9:mm? Bomeor ren: S. pugiofera sp. n. Corpo bicolore: assenza di lunghissime setole laterali isolate del corpo e delleitible ne e ROLO SNO 9 276 R. PACE 9 Pronoto;giallo-rossiccio: 0... ee MONNIER 10 - Pronoto bruno-rossiccio o:giallo=bmuno;... RR 13 10 Pronoto più stretto in avanti che all’indietro; bulbo distale della sperma- teca perfettamente sferico e molto sviluppato. Lungh. 1,9 mm... S. fallax sp. n. - Pronoto più stretto all’indietro che in avanti; bulbo distale della spermateca non sferico, se lo e allora. è molto poco;sviluppato ME Rene 11 11 Addome unicolore giallo-rossiccio; bulbo distale della spermateca oyale-Eunseh21,3 ma.Bormeor. an ae S. esuriens sp. n. - Addome bicolore rossiccio o giallo-rossiccio con fascia posteriore bruna; bulbo distale della spermateca sferico e poco sviluppato o piri- forme a grande sviluppo. BS EE RE ARTE RER 12 12 Undicesimo antennomero più lungo dei tre antennomeri precedenti riuniti; bulbo distale della spermateca sferico e poco sviluppato. Lungh. I8ImmNBommeor rn Ri IRA S. liwaguensis sp. n. - Undicesimo antennomero nettamente più corto dei tre antennomeri precedenti riuniti; bulbo distale della spermateca piriforme e molto sviluppato Euneh1F8mmBONMEO FER 1. NE S. bitruncata sp. n. 13 Addome unicolore giallo-rossiccio, bruno-rossiccio o giallo sporco . . . . . . 14 - Addome bicolore bruno-rossiccio con fascia posteriore bruna. Lungh. 2.1 mm. Sri Lanka, India, Cina, Vietnam, Singapore, Giava, Borneo Sa O E ls lsh eee i S. tuberculicollis (Kraatz) 14 Addome giallo-rossiccio; parte prossimale della spermateca descrivente Wiefspire.eungh'Vl:7imm“Bormeo ner er S. muluensis Sp. n. - Addome bruno-rossiccio o giallo sporco; parte prossimale della sper- matecaldescrivente cinque:spire'o;senza' spire) iii CPR Renn 15 15 Capo e pronoto bruno-rossicci; elitre brune; undicesimo antennomero unicolore bruno; parte prossimale della spermateca non descrivente spitesluneh 12mm. Omeon sy ee S. longecristata sp. n. - Capo, pronoto ed elitre giallo-bruni; undicesimo antennomero bicolore bruno con apice rossiccio; parte prossimale della spermateca descrivente Cinque spires Lungin yl :6mm: Borneo S. myllaenatheca sp. n. Coenonica Kraatz, 1857 Coenonica Kraatz, 1857: 45 Deralia Cameron, 1920: 238, syn. n. Per la discussione vedere le comparazioni date per Coenonica cryptospina sp. n. Coenonica fuscipennis (Cameron, 1920), comb. n. Deralia fuscipennis Cameron, 1920: 238 Per la discussione vedere le comparazioni date per Coenonica cryptospina sp. n. Coenonica trilobata sp. n. Figg. 62-66 Holotypus &, Sabah, Mt. Kinabalu, 1550 m, 23.IV.1987, leg. Burckhardt & Löbl (MHNG). Paratypus: 1 2, Borneo-Sabah, Crocker Ra., 1270 m, Km 60, rte. Kota Kinabalu- Tambunan, 17.V.1987, leg. Burckhardt & Löbl. HOMALOTINI DEL BORNEO DAN, 1 mm FicG. 58-66 Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del &. 58-59: Stenomastax terminalis sp. n.; 60-61: Stenomastax fallax sp. n.; 62-66: Coenonica trilobata sp. n. 278 R. PACE DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e giallo-rossiccio; antenne giallo- rossicce; zampe gialle. La punteggiatura del capo è fitta e assente sulla fronte. La granulosità del pronoto è confusa, quella delle elitre e dell’addome è distinta. La reticolazione del capo e dell’addome è distinta, quella del pronoto è netta e quella delle elitre è superficiale. Edeago figg. 63-64, sesto urotergo libero del d fig. 66, sper- mateca fig. 65. COMPARAZIONI. Per avere il corpo non fortemente punteggiato e per la forma del margine posteriore del sesto urotergo libero del 3, la nuova specie si mostra affine a C. puncticollis Kraatz, 1857, largamente diffusa nella zona intertropicale. Ma l’ede- ago ha minore dimensione ed è dilatato, in visione ventrale (a lati paralleli in puncti- collis). Inoltre la nuova specie presenta tre lobi mediani al margine posteriore del sesto urotergo libero del 4, mentre puncticollis ne ha uno solo. ETIMOLOGIA. Il nome della nuova specie è in riferimento ai tre lobi del margine posteriore del sesto urotergo libero del à. Coenonica quadrilobata sp. n. Figg. 67-70 Holotypus d, Sabah, Crocker Ra., 1600 m, Km 51 rte.Kota Kinabalu-Tambunan, 18.V.1987, Burckhardt & Löbl leg. (MHNG). DESCRIZIONE. Lungh. 1,7 mm. Corpo lucido e giallo-rossiccio, con elitre brune con base giallo-rossiccia; antenne brune con i due antennomeri basali e l’undicesimo giallo-rossicci; zampe gialle. La punteggiatura del capo è distinta e assente sulla fronte. La granulosità del pronoto è indistinta, quella delle elitre è confusa e quella dell’ad- dome è saliente. La reticolazione del capo è superficiale, quella del pronoto è netta e quella delle elitre e dell'addome è distinta. Edeago figg. 68-69, sesto urotergo libero del d fig. 70. COMPARAZIONI. La nuova specie è affine a C. trilobata sp. n. sopra descritta. Se ne distingue per il differente colore delle antenne e del corpo, per presentare quattro lobi al margine posteriore del sesto urotergo libero del d e per l’edeago più profonda- mente ricurvo al lato ventrale. ETIMOLOGIA. Il nome della nuova specie è in riferimento ai quattro lobi del margine posteriore del sesto urotergo libero del d. Coenonica pansa sp. n. Figg. 71-75 Holotypus d, Sabah, Mt. Kinabalu,1750 m, 27.IV.1987, leg. Burckhardt & Löbl (MHNG). Paratypus: 1 9, stessa provenienza. DESCRIZIONE. Lungh. 2,0 mm. Corpo lucido e giallo-rossiccio, con elitre brune con base giallo-rossiccia; antenne rossicce con i due antennomeri basali e l'undicesimo giallo-rossicci; zampe gialle. La punteggiatura del capo è fitta e netta. La granulosità del pronoto è confusa, quella delle elitre è distinta e quella dell’addome è saliente. La reticolazione del capo è assente, quella del pronoto è vigorosa e quella delle elitre è netta. Edeago figg. 72-73, sesto urotergo libero del d fig. 75, spermateca fig. 74. COMPARAZIONI. La nuova specie è affine alle due specie immediatamente prece- denti: C. trilobata sp. n. e C. quadrilobata sp. n. Si distingue dalla prima per il colore differente delle antenne e del corpo e da entrambe per il lobo mediano del margine pos- ti st HOMALOTINI DEL BORNEO 279 0.1 mm 68 69 0.1 mm 0,1 mm 0,1 mm 0,1 mm FIGG. 67-75 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 67-70: Coenonica quadrilobata sp. n.; 71-75: Coenonica pansa sp. n. teriore del sesto urotergo libero del ¢ che è lungo e seghettato ai lati e per l’edeago che ha maggiore sviluppo nella nuova specie, con espansioni laterali più accentuate, in visione ventrale. ETIMOLOGIA. Il nome della nuova specie significa “espansa” e allude all’edeago dilatato nella regione preapicale, in visione ventrale. 280 R. PACE Coenonica cryptospina sp. n. Figg. 76-80 Holotypus d, Sabah, Mt. Kinabalu,1550-1650 m, 24.IV.1987, leg. Burckhardt & Löbl (MHNG). Paratypi: 1 d e 8 es., Sabah, Poring Hot Springs, 500 m, 7.V.1987, leg. Burckhardt & Löbl; 3 es., Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt & Löbl; à, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480 m, 10.V.1987, leg. Burckhardt & Löbl; 4 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, Burckhardt & Löbl leg.; 1 es., Sabah, Poring Hot Springs, 500 m, 13.V.1987, leg. Burckhardt & Löbl; 1 9, Borneo, Sabah, Mt. Kinabalu N.P., 1750 m, 27.1V.1987, Burckhardt & Löbl leg. DESCRIZIONE. Lungh. 2,0 mm. Capo e pronoto opachi, resto del corpo lucido. Corpo rossiccio, con capo ed elitre bruni; antenne rossicce con i due antennomeri basali e l’apice dell’undicesimo giallo-rossicci; zampe giallo-rossicce. La punteggia- tura del capo e del pronoto é netta. La granulosita delle elitre & distinta e frammista a forti punti isolati. L’addome presenta granulosita saliente sui tre uroterghi basali e superficiale sui restanti. La reticolazione delle elitre è distinta, assente sul resto del cor- po, tranne nel fondo dei solchi trasversi basali degli uroterghi dove è vigorosa. Alla base del pronoto stanno due impressioni mediane superficiali. Edeago figg. 77-78, sesto urotergo libero del 4 fig. 80, spermateca fig. 79. COMPARAZIONI. La nuova specie presenta due impressioni mediane posteriori del pronoto e un solo lobo mediano al margine posteriore del sesto urotergo libero del d . Per questi caratteri è possibile che la nuova specie sia tassonomicamente affine a C. fuscipennis (Cameron, 1920), di Singapore. Questa specie è il tipo del genere Deralia Cameron, 1920. L’esame dell’olotipo d mi ha permesso di riconoscere che questo genere è sinonimo di Coenonica Kraatz, 1857, essendo i palpi labiali, ligula e altri caratteri generici non molto differenti da quelli della specie di questo genere. Pertanto Deralia Cameron, 1920 è nuovo sinonimo di Coenonica Kraatz, 1857. Di conseguenza Deralia fuscipennis Cameron, 1920 dovrà essere chiamata Coenonica fuscipennis (Cameron, 1920). A conferma di questa sinonimia e nuova combinazione, sottolineo che l’edeago di fuscipennis è molto simile a quello di C. varicornis (Kraatz, 1859) (tipi da me esaminati), descritta in origine come Leptusa varicornis Kraatz, 1859, già trasferita da Cameron (1939) al genere Coenonica. Ritornando alla comparazione della nuova specie con C. fuscipennis, le due specie sono ben distinte. Il pronoto della nuova specie è appena trasverso (molto trasverso in fuscipennis), il lobo mediano del margine posteriore del sesto urotergo libero del & è larghissimo e corto nella nuova specie (molto lungo in fuscipennis) e l’edeago ha sviluppo molto maggiore nella nuova specie, con armatura interna molto più forte di quella dell’edeago di C. fuscipennis. ETIMOLOGIA. Il nome della nuova specie significa “spina nascosta”. È quella, insieme ad altre presenti a ciascun lato del lobo mediano del margine posteriore del sesto urotergo libero del & , visibili solo a forte ingrandimento. Coenonica microincisa sp. n. Figg. 81-85 Holotypus d, Sabah, Mt. Kinabalu, Poring Hot Springs, 550-600 m, 9.V.1987, Burckhardt & Löbl leg. (MHNG). Paratypi: 5 es., stessa provenienza; 1 es., Borneo, Sabah, Mt. Kinabalu N.P.,1550-1650 m, 24.IV.1987, Burckhardt & Löbl leg.; 1 es., Borneo, Sabah, Mt. Kinabalu N.P.,1750 m, HOMALOTINI DEL BORNEO 281 en G 40 mm 0.1 mm Fico. 76-81 Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del ¢. 76-80: Coenonica cryptospina sp. n.; 81: Coenonica microincisa sp. n. 27.IV.1987, Burckhardt & Löbl leg.; 1 2, Borneo, Sabah, Mt. Kinabalu N.P.,1430 m, 22.V.1987, Burckhardt & Löbl leg. DESCRIZIONE. Lungh. 2,2 mm. Corpo lucido e rossiccio, con elitre brune e ad- dome giallo-rossiccio; antenne bruno-rossicce con i due antennomeri basali e 282 R. PACE l’undicesimo gialli, tranne la base di quest’ultimo che & rossiccia; zampe giallo- rossicce. La punteggiatura del capo è ombelicata, superficiale e assente sulla fronte. La granulosità del pronoto è saliente, quella delle elitre è distinta e quella dell’addome è quasi indistinta. La reticolazione del capo è evidente, quella del pronoto e delle elitre è assente e quella dell’addome è distinta solo sugli uroterghi liberi quarto e quinto, sui basali è estremamente superficiale. Edeago figg. 82-83, sesto urotergo libero del d fig. 85, spermateca fig. 84. COMPARAZIONI. L’habitus e il colore del corpo e delle antenne della nuova specie sono molto simili a quelli di C. fuscipennis (Cameron, 1920), di Singapore. La nuova specie se ne distingue per l’assenza di quattro tubercoli sul lobo mediano del margine posteriore del sesto urotergo libero del d, presente in fuscipennis, e per avere lo stesso lobo inciso all’apice (non inciso in fuscipennis). L’edeago è nettamente differente nelle due specie: con bulbo basale molto sviluppato e con “crista proximalis” poco sviluppata in fuscipennis e con bulbo basale meno sviluppato e con “crista proximalis” molto sviluppata nella nuova specie. ETIMOLOGIA. Il nome della nuova specie deriva dalla presenza di una piccola incisione all’apice del lobo mediano del margine posteriore del sesto urotergo libero del d. Coenonica subrhomboidalis sp. n. Figg. 86-90 Holotypus 4, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 480 m, 10.V.1987, leg. A. Smetana (MHNG). Paratypi: 4 es., stessa provenienza; 1 2, Sabah, Poring Hot Springs, Langanan river, 850 m, 14.V.1987, leg. Burckhardt & Löbl. DESCRIZIONE. Lungh. 2.1 mm. Corpo lucido e bruno-rossiccio, con elitre brune e addome giallo-rossiccio con fascia posteriore rossiccia; antenne brune con i due an- tennomeri basali rossicci e l'undicesimo giallo con base bruno-rossiccia; zampe gialle. La punteggiatura del capo è netta sul disco e superficiale ai lati. La granulosità del pronoto è saliente, quella delle elitre e dell'addome è distinta. Profonda è la fossetta mediana posteriore del pronoto. La reticolazione del capo è distinta, quella del pronoto è estremamente superficiale, quella delle elitre è superficiale e quella dell’addome è assente. Edeago figg. 87-88, sesto urotergo libero del & fig. 90, spermateca fig. 89. COMPARAZIONI. La nuova specie è simile a C. varicornis Kraatz, 1859, dello Sri Lanka, per il colore delle antenne, il pronoto trasverso e la struttura dell’edeago. Ma l’edeago della nuova specie presenta forte armatura interna (diafana in varicornis) e non possiede due forti tubercoli sul sesto urotergo libero del à. ETIMOLOGIA. Il nome della nuova specie deriva dalla parte apicale dell’edeago a forma quasi romboidale. Coenonica burckhardti sp. n. Figg. 91-95 Holotypus 3d, Sabah, Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, leg. Burckhardt & Löbl (MHNG). Paratypi: 1 d, stessa provenienza; 2 d d e 1 ©, Sabah, Poring Hot Springs, nr. Bat Cave, 0-600 m, 9.V.1987, leg. Burckhardt & Löbl; 1 é, Sabah, Poring Hot Springs, Langanan Falls, 0 m, 14.V.1987, leg. Burckhardt & Löbl. DESCRIZIONE. Lungh. 2,0 mm. Avancorpo debolmente opaco, addome lucido. Corpo rossiccio; antenne bruno-rossicce con i tre antennomeri basali giallo-rossicci e 55 85 HOMALOTINI DEL BORNEO 283 89 0.1 mm 0,1 mm 0,1 mm Fico. 82-90 Edeago in visione laterale e ventrale, sesto urotergo libero del d, habitus e spermateca. 82-85: Coenonica microincisa sp. n.; 86-90: Coenonica subrhomboidalis sp. n. l’undicesimo giallo con base giallo-rossiccia; zampe giallo-rossicce. La punteggiatura del capo e del pronoto è ombelicata, profonda e fittissima. I granuli che coprono le elitre sono allungati. Una reticolazione vigorosa sta sulla fronte. Assente è la retico- lazione su altri parti del corpo. Edeago figg. 92-93, sesto urotergo libero del d fig 95, spermateca fig. 94. 284 R. PACE COMPARAZIONI. L’habitus della nuova specie ricorda da vicino quello di C. fuscipennis (Cameron, 1920), di Singapore, ma l’edeago è fortemente dilatato nella regione preapicale, in visione ventrale. Ciò non si osserva nell’edeago di fuscipennis. ETIMOLOGIA. La nuova specie è dedicata a uno dei suoi raccoglitori, il dr. Daniel Burckhardt, già del Museo di Storia Naturale di Ginevra. Coenonica lucidula sp. n. Figg. 96-99 Holotypus d, Borneo, Sabah, Mt. Kinabalu N.P., below Layang Layang, 2600 m, 9-20.V.1987, in[terception] trap, leg. A. Smetana (MHNG). Paratypus: 1 d, Borneo, Sabah, M. Kinabalu N.P., Laban Rata, 3200-3250 m, 4.V.1987, leg. A. Smetana. DESCRIZIONE. Lungh. 2,2 mm. Corpo lucido e bruno, con pigidio giallo- rossiccio; antenne rossicce con i due antennomeri basali e l’apice dell’undicesimo giallo-rossicci; zampe giallo-rossicce. La punteggiatura ombelicata del capo è superfi- ciale. La granulosità del pronoto è quasi indistinta, quella delle elitre è distinta. La reti- colazione del capo, del pronoto e dell’addome è netta, quella delle elitre è superficiale. Edeago figg. 97-98, sesto urotergo libero del d fig. 99. COMPARAZIONI. L’edeago e il margine posteriore del sesto urotergo libero del 4 ricordano da vicino quelli di C. vulnerata Bernhauer, 1915, del Mt. Matang (Borneo). Ma mentre in quest’ultima specie il margine posteriore del sesto urotergo libero del 4 è dentellato solo nella parte mediana, quello della nuova specie lo è fino alle spine lat- erali. Inoltre, l’edeago della nuova specie è più ricurvo al lato ventrale. ETIMOLOGIA. Il nome della nuova specie significa “un po’ lucida”. Coenonica kinabaluicola sp. n. Figg. 100-104 Holotypus d, Borneo, Sabah, Mt. Kinabalu N.P., Laban Rata, 3200-3250 m, 4.V.1987, leg. A. Smetana (MHNG). Paratypi: 28 es., stessa provenienza; 1 d, Borneo, Sabah, Mt. Kinabalu N.P., Layang Layang, 2610 m, 2.V.1987, A. Smetana leg.; 1 9, Sabah, Mt. Kinabalu, 2600 m, 2.V.1987, leg. Burckhardt & Löbl; 26 es., Sabah, Mt. Kinabalu, 1750 m, 27.IV.1987, leg. Burckhardt & Löbl]; 13 es., Borneo, Sabah, M. Kinabalu N.P., below Laban Rata, 3150 m, 7.VIII.1988, leg. A. Smetana; | d, Borneo-Sabah, M. Kinabalu N.P., Summit Trail, 1890 m, (data mancante), leg. A. Smetana. DESCRIZIONE. Lungh. 2,7 mm. Avancorpo debolmente opaco, addome lucido. Corpo nero pece, con pigidio rossiccio; antenne nero pece, con i due antennomeri basali rossicci; zampe giallo-rossicce. La punteggiatura del capo è quasi indistinta, quella del pronoto è assente. La granulosità delle elitre è fine, quella dell’addome è distinta. La reticolazione del capo e del pronoto è vigorosa, quella delle elitre e dell’addome è distinta. Spermateca fig. 101, edeago figg. 102-103, sesto urotergo libero del < fig. 104. ETIMOLOGIA. Il nome della nuova specie significa “abitante del Kinabalu”. COMPARAZIONI. La nuova specie è simile a C. soror Cameron, 1933, pure del Borneo, a motivo della forma simile della spermateca. Se ne distingue per i caratteri dati nella seguente chiave. HOMALOTINI DEL BORNEO 285 Fico. 91-98 Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del 3. 91-95: Coenonica burckhardti sp. n.; 96-98: Coenonica lucidula sp. n. 286 R. PACE 1 mm rn X N eh + {} ART 1 1A À 101 | x | 0.1 mm 102 Fico. 99-104 Sesto urotergo libero del 3, habitus, spermateca ed edeago in visione laterale e ventrale. 99: Coenonica lucidula sp. n.; 100-104: Coenonica kinabaluicola sp. n. HOMALOTINI DEL BORNEO 287 1 Addome giallo-rossiccio; edeago arcuato al lato ventrale; spermateca più sferica, senza introflessione apicale del bulbo distale; tra le spine laterali del margine posteriore del sesto urotergo libero del d sono pre- senti dei dentini. Lungh. 3,0 mm. Borneo: Mt. Matang, Mt. Kinabalu 000 à 6. LR E ee ean Ra ie Sam C. soror Cameron - Addome nero pece con pigidio rossiccio; edeago sinuato al lato ventrale; spermateca oblunga, con introflessione apicale del bulbo distale; tra le spine laterali del margine posteriore del sesto urotergo libero del & sono assenti dei dentini. Lungh. 2,7 mm. Borneo: Mt. Kinabalu ME al de nn RÉ e DIDO C. kinabaluicola sp. n. Coenonica bruneiensis sp. n. Figg. 105-107 Holotypus 4, Borneo, Brunei, HW Temburong Kuala, Belalong KBFSC, 10.11.1995, leg. Borcherding (MRSN). DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e nero pece, con addome nero- bruno; antenne nere con antennomero basale nero-bruno; zampe anteriori bruno- rossicce con femori gialli, zampe medie e posteriori nero-brune con tarsi rossicci. La punteggiatura del capo è profonda, quella del pronoto è netta, più densa e più fine sulla fascia longitudinale mediana e quella dell’addome profondissima, ma assente presso il margine posteriore degli urotergo liberi primo a quarto. La granulosità delle elitre è saliente. Ciascuna elitra del 3 presenta una fila di tubercoli salienti presso la sutura. Il quinto urotergo libero del d mostra tubercoli allungati salienti. Edeago figg. 106-107. COMPARAZIONI. La forma dell’edeago e l’habitus della nuova specie sono simili a quelli di C. angusticollis Cameron, 1920, di Singapore (tipi da me esaminati). L’edeago della nuova specie è strettamente arcuato al lato ventrale e la sua armatura interna è forte e di forma complessa, mentre in angusticollis l’edeago è assai ampia- mente arcuato al lato ventrale e la sua armatura interna è composta da un pezzo ricurvo fortemente sclerificato e altri pezzi diafani. esternamente, la nuova specie ha pronoto nettamente trasverso, mentre angusticollis lo ha poco. ETIMOLOGIA. La nuova specie prende nome dal Brunei, dove è stata raccolta. Coenonica mimoleptusa sp. n. Figg. 108-111 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 5.V.1987, leg. A. Smetana (MHNG). DESCRIZIONE. Lungh. 2,1 mm. Corpo lucido e bruno, con addome nero e pigidio giallo-rossiccio; antenne brune con i due antennomeri basali giallo-bruni e metà api- cale dell’undicesimo giallo-rossiccia; zampe giallo-brune. La punteggiatura del capo è fitta e superficiale, assente sulla fronte. La granulosità del pronoto è fitta e superficiale, quella delle elitre è poco saliente e quella dell’addome è evidente. La reticolazione del capo è distinta solo sulla fronte, quella del pronoto è assente, quella delle elitre è netta e quella dell’addome è evidente. Il quinto urotergo libero del d presenta tubercoli allungati a ciascun lato della carena mediana che è poco saliente. Edeago figg. 109-110, sesto urotergo libero del d fig. 111. COMPARAZIONI. La nuova specie presenta il margine posteriore del sesto uro- tergo libero del d dentellato come quello di C. javana Bernhauer, 1914, di Giava, 288 R. PACE 1 mm El Bet Fico. 105-111 Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del d. 105-107: Coenonica bruneiensis sp. n.; 108-111: Coenonica mimoleptusa sp. n. HOMALOTINI DEL BORNEO 289 (holotypus d da me esaminato). Tuttavia, il corpo della nuova specie & superficial- mente punteggiato o granuloso, mentre in javana lo è fortemente. Inoltre, l’armatura interna dell’edeago di javana non è così lunga e forte come quella dell’edeago della nuova specie. ETIMOLOGIA. L’habitus della nuova specie ricorda da vicino delle specie del genere Leptusa Kraatz, 1856. L'esame della ligula esclude l’appartenenza a questo genere. Il suo nome significa “imitatrice di Leptusa”. Coenonica cristamagna sp. n. Figg. 112-116 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ at Liwagu Rv., 1500 m, 30.IV.1987, A. Smetana leg. (MHNG). Paratypi: 1 6 e 1 2, Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1500 m, 25.1V.1987, A. Smetana leg. DESCRIZIONE. Lungh. 2,2 mm. Corpo lucido e bruno, con elitre nero-brune; antenne brune con i due antennomeri basali, la base del terzo e l’apice dell’undicesimo gialli; zampe giallo-rossicce. La punteggiatura del capo è fitta, superficiale e assente sulla fronte. La granulosità del pronoto è distinta, quella delle elitre è forte e saliente e quella dell’addome è molto superficiale, tranne i granuli del margine posteriore degli uroterghi. La reticolazione della fronte e dell’addome è distinta, quella del pronoto è ben visibile, quella delle elitre è assente. Edeago figg. 113-114, sesto urotergo libero del d fig. 115, spermateca fig. 116. COMPARAZIONI. La nuova specie apparentemente è simile a C. microincisa n. sp. sopra descritta, se si osserva la forma del margine posteriore del sesto urotergo libero del 4. Tuttavia, l’edeago della nuova specie non è slanciato come quello di microincisa e il suo apice è largo (acuto in microincisa). ETIMOLOGIA. Il nome della nuova specie significa “grande cresta”. È quella prossimale del suo edeago. Coenonica leptusaesimilis sp. n. Figg. 117-119 Holotypus d, Sabah, [Mt. Kinabalu N.P.], Poring Hot Springs, 550-600 m, 9.V.1987, Burckhardt & Löbl leg. (MHNG). DESCRIZIONE. Lungh. 2,3 mm. Corpo lucido e bruno; antenne bruno-rossicce, coni tre antennomeri basali e l’undicesimo giallo-rossiccio; zampe bruno-rossicce. La punteggiatura del capo e del pronoto è profonda e fitta. La granulosità delle elitre è saliente e assente lungo il margine posteriore delle stesse elitre. Gli uroterghi sono coperti di granulosità evidente. La reticolazione è netta solo sulla fronte e sul pronoto. Assente è sul resto del corpo. Edeago figg. 117-118. COMPARAZIONI. L’habitus della nuova specie è simile a quello di C. rufiventris Cameron, 1943, pure del Borneo (holotypus 9 da me esaminato). Tuttavia, la nuova specie non ha il capo e il pronoto coperti di granulosità forte come in rufiventris. ETIMOLOGIA. L’habitus della nuova specie è simile quello di alcune specie del genere Leptusa Kraatz, 1856. Ma l’esame della ligula esclude questa attribuzione generica. Pertanto la nuova specie è chiamata “simile a Leptusa”. 290 R. PACE FIGG. 112-119 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del 4 e spermateca. 112- 116: Coenonica cristamagna sp. n.; 117-119: Coenonica leptusaesimilis sp. n. Coenonica muluensis sp. n. Figg. 120-124 Holotypus d, Borneo, Sarawak, Mulu Nat. Park, leg. H. Franz, (NHMW). Paratypi: 2 9 2, stessa provenienza. DESCRIZIONE. Lungh. 2,5 mm. Avancorpo debolmente lucido, addome lucido. Corpo giallo-rossiccio, con elitre brune, tranne la base e i lati che sono rossicci, quarto HOMALOTINI DEL BORNEO 291 urite libero bruno-rossiccio; antenne bruno-rossicce con i due antennomeri basali rossicci e l’undicesimo giallo-rossiccio; zampe gialle. La punteggiatura del capo e del pronoto é fitta e profonda. La granulosita delle elitre é fittissima e saliente, quella dei due uroterghi basali è distinta e quella dei restanti uroterghi è assai rada. Il terzo uro- tergo libero del 3 presenta una bozza mediana saliente. Edeago figg. 121-122, sesto urotergo libero del d fig. 124, spermateca fig. 123. COMPARAZIONI. La nuova specie , per la presenza di una bozza mediana sul terzo urotergo libero del d, per la distribuzione della punteggiatura dell’avancorpo, per il margine posteriore del sesto urotergo libero del d, è sicuramente affine a C. angusti- collis Cameron, 1943, di Singapore (tipi da me esaminati). Tuttavia, per la forma dell’edeago, è chiaramente distinta. L’armatura interna dell’edeago è corta e larga nella nuova specie e lunga e sottile in angusticollis. ETIMOLOGIA. La nuova specie prende nome dalla sua località tipica Mulu. Coenonica rhomboidalis sp. n. Figg. 125-129 Holotypus d, Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ 1560-1660 m, 24.IV.1987, leg. A. Smetana, (MHNG). Paratypi: 4 es., stessa provenienza; 4 es., Sabah, Mt. Kinabalu, 1580 m, 27.IV.1987, leg. Burckhardt & Löbl; 2 es., Sabah, Poring Hot Springs, 550-600 m, 9.V.1987, leg. Burckhardt & Löbl;1 es., Borneo-Sabah, Crocker Ra., 1550-1650 m, 16.V.1987, leg. Burckhardt & Löbl; 1 à, Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1560 m, 3.VIIL.1988, leg. A. Smetana. DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e bruno-rossiccio, con addome rossiccio che ha una fascia posteriore bruno-rossiccia; antenne brune con i due anten- nomeri basali, la base del terzo e apice dell’undicesimo giallo-rossicci; zampe giallo- rossicce. La punteggiatura del capo e fittissima e profonda. La granulosità del pronoto e delle elitre è molto saliente, quella dell’addome è fine. Edeago figg. 126-127, sesto urotergo libero del 4 fig. 129, spermateca fig 128. COMPARAZIONI. La nuova specie presenta il lobo mediano del margine posteriore del sesto urotergo libero del 3 inciso all’apice, come in C. microincisa e C. cristamagna nuove specie sopra descritte. Se ne distingue chiaramente per 1 lati dell’edeago espansi, in visione ventrale. I lati dell’edeago delle due specie a compa- razione non lo sono. ETIMOLOGIA. La nuova specie prende nome dalla forma quasi romboidale della parte apicale dell’edeago, in visione ventrale. Coenonica magnilobata sp. n. Figg. 130-133 Holotypus d, Sabah, E Mt. Kinabalu, 1150 m, rte. Ranau-Kota Kinabalu, 24.V.1987, leg. Burckhardt & Löbl (MHNG). DESCRIZIONE. Lungh. 2,5 mm. Corpo lucido e rossiccio, con elitre brune; antenne bruno-rossicce con i due antennomeri basali, la base del terzo e l’undicesimo giallo-rossicci; antenne giallo-rossicce. La punteggiatura del capo è distinta solo sul disco, ai lati è confusa. La granulosità del pronoto è saliente, quella delle elitre è grossolana e fittissima. Gli uroterghi sono coperti da carene longitudinali salienti, tranne il quinto libero. Edeago figg. 131-132, sesto urotergo libero del à fig. 133. COMPARAZIONI. Per la presenza di un grande lobo mediano del margine posteriore del sesto urotergo libero del 4 e per l’edeago molto arcuato al lato ventrale, 292 R. PACE 123 i 128 el È Fico. 120-128 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 120- 124: Coenonica muluensis sp. n.; 125-128: Coenonica rhomboidalis sp. n. la nuova specie è forse affine a C. malayana Cameron, 1936, di Malaya (holotypus é da me esaminato). Se ne distingue per la presenza di carene longitudinali degli uroterghi liberi primo a quarto e per l’edeago espanso nella regione preapicale, in visione ventrale (a lati quasi paralleli in malayana). HOMALOTINI DEL BORNEO 293 0.1 mm Figc. 129-134 Sesto urotergo libero del 6, habitus ed edeago in visione laterale e ventrale. 129: Coenonica rhomboidalis sp. n.; 130-133: Coenonica magnilobata sp. n.; 134: Coenonica obscuripalpi sp. n. ETIMOLOGIA. Per la presenza di un grande lobo mediano al margine posteriore del sesto urotergo libero del à , la nuova specie è chiamata “dotata di un grande lobo”. 294 R. PACE Coenonica obscuripalpi sp. n. Figg. 134-137 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 25.IV.1987, leg. A. Smetana (MHNG). Paratypi: 5 es., stessa provenienza; 12 es., Borneo, Sabah, Mt. Kinabalu N.P., 1750 m, 21.1V.1987, Burckhardt & Löbl leg.; 5 es., Borneo, Sabah, Mt. Kinabalu N.P., 1580 m, 21.1V.1987, Burckhardt & Löbl leg.; 3 22, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 1.IX.1988, leg. A. Smetana. DESCRIZIONE. 3,8 mm. Corpo lucido e bruno-rossiccio, con margine posteriore dei tre uriti basali rossicci; antenne bruno-rossicce con i tre antennomeri basali giallo- rossicci e l’undicesimo giallo; zampe bruno-rossicce con tarsi giallo-rossicci. La pun- teggiatura del capo è ombelicata e grossolana. La granulosità del pronoto e delle elitre è saliente, più grossolana alla base delle elitre. Il quarto urotergo libero presenta a ciascun lato due-tre fossette. Assente è la reticolazione del corpo. Edeago figg. 135-136, spermateca fig. 137. COMPARAZIONI. L’habitus della nuova specie è simile a quello di C. philippina Bernhauer, 1916, delle Filippine (tipi da me esaminati). Tuttavia, l’edeago della nuova specie ha apice molto protratto ( cortissimo in philippina). ETIMOLOGIA. La nuova specie prende nome dal colore oscuro dei suoi palpi mascellari. Coenonica pallipes sp. n. Figg.138-141 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ 1560-1600 m, 24.IV.1987, A. Smetana leg.(MHNG). Paratypi: 2 d d, stessa provenienza. DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e nero-bruno; antenne nero-brune con i tre antennomeri basali giallo-rossicci; zampe brune con tarsi e tibie anteriori gialli. La punteggiatura del capo è composta da punti ombelicati enormi e grossolani. La granulosità del pronoto è grossolana ed è assente nei pressi del margine posteriore, quella delle elitre è composta di granuli allungati e netti. Gli uroterghi sono pressoché nudi, con qualche sparsa setola. Il quinto urotergo libero del d ha una carena mediana concava. Assente è la reticolazione sul corpo. Edeago figg. 139-140, sesto urotergo libero del d fig. 141. COMPARAZIONI. L’habitus della nuova specie è simile a quello di C. philippina Bernhauer, 1916, delle Filippine (tipi da me esaminati). Tuttavia, l’edeago della nuova specie è slanciato e non tozzo come quello di philippina. ETIMOLOGIA. La nuova specie, avendo i tarsi gialli e meso-metatibie brune, è chiamata “dai piedi pallidi”. Coenonica spinae sp. n. Figg. 142-146 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 16.V.1987, leg. A. Smetana (MHNG). Paratypi: 7 es., Borneo, Sabah, Mt. Kinabalu, 1750 m, 21.IV.1987, Burckhardt & Löbl leg.; 1 ?, Borneo, Sabah, Mt. Kinabalu Nat. Pk., Poring Hot Springs, 510 m, 12.V.1987, leg. A. Smetana. DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e nero-bruno; antenne nero-brune con i due antennomeri basali rossicci; zampe nero-brune con i tarsi bruno-rossicci. La HOMALOTINI DEL BORNEO 295 0,1 mm Ficc. 135-141 Edeago in visione laterale e ventrale, spermateca, habitus e sesto urotergo libero del 4. 135-137: Coenonica obscuripalpi sp. n.; 138-141: Coenonica pallipes sp. n. 296 R. PACE Fico. 142-147 Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del d. 142-146: Coenonica spinae sp. n.; 147: Coenonica troglobia sp. n. punteggiatura del capo è ombelicata e profondissima. La granulosità del pronoto è grossolana, quella delle elitre è composta di granuli appena allungati e forti. L’addome è coperto di granulosità saliente distribuita come da fig. 142. Assente è la reticolazione del corpo. Edeago figg. 143-144, sesto urotergo libero del d, fig. 146, spermateca fig. 145. HOMALOTINI DEL BORNEO 297 COMPARAZIONI. L’edeago della nuova specie é simile a quello di C. philippina Bernhauer, 1916, delle Filippine (tipi da me esaminati). Anche per 1 caratteri dell’habitus l’affinità tassonomica delle due specie è sicura. La nuova specie si differenzia da essa per avere il tubulo sporgente dell’edeago nettamente più lungo di quello di philippina e l’apice dello stesso organo della nuova specie è inciso, in visione ventrale, e intero in philippina. Inoltre, le zampe della nuova specie sono nero-brune e in philippina giallo-rossicce. Nel Borneo è nota un’altra specie di questo gruppo di C. philippina. È C. ashei Pace, 1987, che per il colore del corpo è più simile a philippina, ma il suo edeago è nettamente differente, avendo l’apice a paletta, in visione laterale. ETIMOLOGIA. La nuova specie prende nome dal rilievo a forma di spina presente al lato ventrale dell’edeago. Coenonica troglobia sp. n. Figg. 147-150 Holotypus 6, Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2997 m, 5.V.1987, leg. A. Smetana (MHNG). Paratypi: 7 es., stessa provenienza; 3 es., Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 2.V.1987, leg. A. Smetana; 11 es., Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 6.V.1987, leg. A. Smetana. DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e bruno-rossiccio, con addome bruno; antenne brune con i quattro antennomeri basali e l’apice dell’undicesimo giallo- rossicci; zampe giallo-rossicce. La punteggiatura del capo e del pronoto è distinta e poco profonda, quella delle elitre è netta. La granulosità dell’addome è distinta. La reti- colazione del capo, del pronoto e dell’addome è distinta, quella delle elitre è superfi- ciale. Edeago figg. 149-150, spermateca fig. 147. COMPARAZIONI. Un edeago di enormi dimensioni e dentato alla regione pre- apicale ventrale non è mai stato osservato nel genere Coenonica, tranne nella specie seguente, che le è affine. ETIMOLOGIA. Poiché la nuova specie è stata raccolta in una grotta, prende il nome di “cavernicola”. Coenonica perpunctata sp. n. Figg. 151-154 Holotypus d, Borneo-Sabah, Mt. Kinabalu N. P., Silau-Silau Trail, 1558 m, 2.IX.1988, leg. D.E. Bright (MHNG). Paratypi: 23 es., Sabah, Mt. Kinabalu, 1450-1550 m, 23.IV-21.V.1987, leg. Burckhardt & Löbl; 2 es., Borneo, Sabah, Mt. Kinabalu N.P., 1900 m, 26.1V.1987, Burckhardt & Löbl leg.;12 es., Sabah, Mt. Kinabalu, 1550 m, 28.1V.1987, leg. Burckhardt & Löbl; 1 es., Borneo- Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1560 m, 30.1V.1987, leg. A. Smetana; | es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, Burckhardt & Löbl leg. DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e rossiccio; antenne rossicce con i tre antennomeri basali e i due terzi apicali dell’undicesimo giallo-rossicci; zampe giallo-rossicce. La punteggiatura del capo e del pronoto è profonda e fittissima. La granulosità delle elitre è grossolana, quella dell’addome è saliente. Solo gli uroterghi quarto e quinto presentano una reticolazione, che è distinta. Edeago figg. 153-154, spermateca figg. 152. 298 R. PACE 0.1 mm FIGG. 148-152 Habitus, edeago in visione laterale e ventrale e spermateca. 148-150: Coenonica troglobia sp. n.; 151-152: Coenonica perpunctata sp. n. COMPARAZIONI. La nuova specie è sicuramente affine a C. troglobia sp. n. sopra descritta, a motivo della forma dell’edeago e per il simile habitus. Se ne distingue per l’edeago piegato ventralmente ad angolo retto, in visione laterale, (edeago largamente arcuato ventralmente in trog/obia), per il dente preapicale ventrale dello stesso organo > ittici HOMALOTINI DEL BORNEO 299 nettamente piu vicino all’apice di quello di troglobia e per la differente armatura interna dell’edeago. ETIMOLOGIA. L’avancorpo della nuova specie è coperto da forte punteggiatura o granulosità, pertanto porta il nome di “oltremodo punteggiata”. CHIAVE DEI 6 d DELLE SPECIE DEL BORNEO DEL GENERE COENONICA 1 2 (GS) Quarto antennomero lungo quanto largo o più lungo che largo........... 2 Ouartorantennomeroitrasversor. wea teats I Klass II NI 15 OQuantorantennomero piullungoicheslareor.. ME AE Re 3 Quarto antennomero lungo quanto! lango 2). Km. ne ee... 9 Capo, pronoto ed elitre superficialmente punteggiati o finemente granu- losi. Lungh. 2,7 mm. Borneo: Mt. Kinabalu ........ C. kinabaluicola sp. n. Capo, pronoto ed elitre fortemente punteggiati o grossolanamente granu- GET ait LEN et 2e ER PU CES A PRE A ASS ee ne SES NC 4 Pronoto debolmente trasverso, con rapporto larghezza/lunghezza pari a 1,25; quarto urotergo libero con fossette laterali; edeago molto prolun- gato all’apice. Lungh. 3,8 mm. Borneo: Mt. Kinabalu . . C. obscuripalpi sp. n. Pronoto nettamente trasverso, con rapporto larghezza/lunghezza pari a 1,31-1,71; quarto urotergo libero senza fossette laterali; edeago non pro- lungatogalleapicee RENTE ES PRES ke Ameo RE Rd 5 Margine posteriore del sesto urotergo libero del d con un largo lobo medianottrayle:;spineilaterali ie RE 6 Margine posteriore del sesto urotergo libero del d rettilineo ............. 7 Corpo unicolore rossiccio; terzo urotergo libero del 4 senza bozza mediana; lati del lobo del margine posteriore del sesto urotergo libero del 4 semplici; edeago fortemente dilatato nella regione preapicale, in visione ventrale. Lungh. 2,0 mm. Borneo: Mt. Kinabalu . C. burckhardti sp. n. Corpo bicolore giallo-rossiccio, con elitre e fascia addominale posteriore bruna o bruno-rossiccia; terzo urotergo libero del d con una bozza mediana; lati del lobo del margine posteriore del sesto urotergo libero del 4 seghettati; edeago appena dilatato nella regione preapicale, in visione ventrale. Lungh. 2,0 mm. Borneo: Mt. Mulu . . . C. muluensis sp. n. Pronoto meno trasverso, con rapporto larghezza/lunghezza pari a 1,31; granulosità delle elitre mancante lungo il margine posteriore; apice del- l’edeago larghissimo, in visione ventrale. Lungh. 2,3 mm. Borneo: Mt. Kinobaluptggrenzere atene nni C. leptusaesimilis sp. n. Pronoto fortemente trasverso, con rapporto larghezza/lunghezza pari a 1,55-1,71; granulosità delle elitre estesa su tutta la loro superficie; apice dellfedeasofstrettissimo*fmnivisione ventrale ele eee ER A 8 Corpo unicolore rossiccio; pronoto meno trasverso, con rapporto lar- ghezza/lunghezza pari a 1,55 e senza impressione mediana posteriore; edeago piegato ad angolo retto al lato ventrale. Lungh. 2,8 mm. Borneo: MARinabalun ae. ne EEE C. perpunctata sp. n. Corpo bicolore: avancorpo bruno-rossiccio, addome bruno; pronoto fortemente trasverso, con rapporto larghezza/lunghezza pari a 1,71 e con 300 13 14 17 R. PACE | impressione mediana posteriore; edeago arcuato al lato ventrale. Lungh. PAI MMS Bormeo: MtyKanabaluy. een C. troglobia sp. n. Uriti liberi primo a quarto coperti di carene longitudinali salienti. Lungh. 2 Sn BONE OM NIET ee SIR C. magnilobata sp. n. Uriti con la consueta granulosità o punteggiatura .................... 10 Margine posteriore del sesto urotergo libero del & con spinule solo nel terzo mediano. Lungh. 1,7. Borneo: Mt. Matang..... C vulnerata Bernhauer Margine posteriore del sesto urotergo libero del 4 dentellato o lobato Anotalelatie a De pal Solco mediano del pronoto a U o a V; edeago con lunghissimo tubulo sporgente: dallionfiziovapicale vir. e Re 12 Pronoto con depressione posteriore a lati paralleli o con fossetta posteriore 14 Corpo unicolore nero-bruno, con zampe nero-brune o brune; margine posteriore del sesto urotergo libero del d con uno o tre larghi lobi e poco SPOrpEnt sin ee Re Eee PS 13 Corpo tricolore giallo-rossiccio, con capo ed elitre bruno-rossicci e base dell’addome rossiccia; zampe giallo-rossicce; margine posteriore del sesto urotergo libero del d cinque lobi sporgenti. Lungh. 2,5 mm. BornecASandakanirentamaret MI C. ashei Pace Margine posteriore del sesto urotergo libero del 4 con tre lobi tra le spine laterali; edeago con spina ventrale. Lungh. 2,6 mm. Borneo: Mt. Kanal allure risa a NIRO ERE C. spinae sp. n. Margine posteriore del sesto urotergo libero del d con un solo lobo mediano tra i lobi laterali; edeago senza spina ventrale. Lungh. 2,8 mm. Bomeo#MAKmabalu 2. HT IE Rey ee ehe C. pallipes sp. n. Antenne e corpo unicolori nero pece; una fossetta mediana posteriore; margine posteriore del sesto urotergo libero del d con numerosi denti- mipezunghy 2.8mm: Bomeos Brunel: SESSO. C. bruneiensis sp. n. Antenne brune con undicesimo antennomero giallo con base rossiccia; corpo bicolore giallo-rossiccio e bruno; margine posteriore del sesto urotergo libero del d con quattro lobi mediani, i laterali con margine seghettato. Lungh. 2,1 mm. Borneo: Mt. Kinabalu . . . C. subrhomboidalis sp.n. Pronoto poco trasverso, con rapporto larghezza/lunghezza pari a 1,13- LAS sat. cee MORI RR Ae RU ERA re 16 Pronoto molto trasverso, con rapporto larghezza/lunghezza pari a 1,27- ISTE icaro D oo cs co c = 0 = 17 Pronoto con due fossette mediane basali; elitre con forti punti assai ra- di; margine posteriore del sesto urotergo libero del & con largo lobo seghettato ai margini laterali. Lungh. 2,5 mm. Borneo: Mt. Kinabalu a NTO DRE CPE EE M NN REN C. magnilobata sp. n. Pronoto con una fossetta mediana basale: elitre con granulosità distinta; margine posteriore del sesto urotergo libero del d con numerose spine e lobi. Lungh. 2,2 mm. Borneo: Mt. Kinabalu............. C. lucidula sp. n. Margine posteriore del sesto urotergo libero del d con sei piccoli lobi mediani. Lungh. 2,1 mm. Borneo: Mt. Kinabalu...... C. mimoleptusa sp. n. 18 19 20 21 22 HOMALOTINI DEL BORNEO 301 Margine posteriore del sesto urotergo libero del d con un grande lobo MEAN OLE A ET NE SEEN I a ER ES 18 Margine posteriore del sesto urotergo libero del d con lobo mediano in- eisoganmetaralllapice zai LS DEREN 19 Margine posteriore del sesto urotergo libero del d con lobo mediano TS RONA TRI IE A RE IA SRI I E 21 Addome unicolore giallo-rossiccio o bruno; edeago non dilatato ai lati, InWvisIonenventralen UNE LRO TE An ere RER er ANRT AE ee 20 Addome bicolore rossiccio con quarto urite libero bruno-rossiccio; edea- go fortemente dilatato al lati, in visione ventrale. Lungh. 2,6 mm. Bomeo#Mti inabile EN: C. rhomboidalis sp. n. Addome giallo-rossiccio; apice dell’edeago acuto. Lungh. 2,2 mm. Bomeos MtzyKanab ale ps aie: eae ee C. microincisa sp. n. Addome bruno; apice dell’edeago largamente arcuato. Lungh. 2,2 mm. Borneo MtiRinabalu& re asl See eae ee seen C. cristamagna sp. n. Corpo unicolore giallo-rossiccio; margine posteriore del sesto urotergo libero del d con lobo mediano composto da tre piccoli lobi. Lungh. [Samb omeofMtsKnabalif}. aa a a Dame e C. trilobata sp. n. Corpo bicolore giallo-rossiccio, con tre quarti posteriori delle elitre bruni; margine posteriore del sesto urotergo libero del d con un solo lobo seghettato lateralmente o con un lobo composto da quattro piccoli NOIRE EN. ER LE er a RE 22 Pronoto piü trasverso, con rapporto larghezza/lunghezza pari a 1,37; margine posteriore del sesto urotergo libero del d con lobo mediano composto da quattro piccoli lobi. Lungh. 1,7 mm. Borneo: Mt. Kinabalu SA e Te anes, i tali mali» aha (pr C. quadrilobata sp. n. Pronoto meno trasverso, con rapporto larghezza/lunghezza pari a 1,27; margine posteriore del sesto urotergo libero del d con un lungo lobo mediano seghettato lateralmente. Lungh. 2,0 mm. Borneo: Mt. Kinabalu RE O I OE UE A I GS OE C. pansa Sp. n. CHIAVE DELLE 9 2 DELLE SPECIE DEL BORNEO DEL GENERE COENONICA 1 2 Go Quarto antennomero lungo quanto largo o piü lungo che largo........... 2 Ouartorantennomerortrasverso. RI N ae 11 Ouartorantennomero,piunlunsoichelarso rn Er ye ee eee cee ei 3 Ouartorantennomerollungo.quantovlangons i aaa gan. 8 Capo, pronoto ed elitre superficialmente punteggiati o finemente granu- losi; spermateca ovale o piriforme. Lungh. 2,7 mm. Borneo: Mt. KA ARO ay cod etre nee ati an encoun RN! C. kinabaluicola sp. n. Capo, pronoto ed elitre fortemente punteggiati o grossolanamente gran- ULOSTAR HERE SP RE 2 O OA SU I ME ds 4 Pronoto debolmente trasverso, con rapporto larghezza/lunghezza pari a 1,25; quarto urotergo libero con fossette laterali; bulbo distale della sper- mateca molto sviluppato ed ellittico. Lungh. 3,8 mm. Borneo: Mt. iran ra N BER Date C. obscuripalpi sp. n. 302 10 18 R. PACE Pronoto nettamente trasverso, con rapporto larghezza/lunghezza pari a 1,31-1,71; quarto urotergo libero senza fossette laterali; bulbo dustale della spermateca grande e sferico, se ellittico ha dimensioni ridotte....... 5) Bulbo distale della spermateca ridotto ed ellittico; “ductus” conformato a formare un lungo bulbo prossimale. Lungh. 2,5 mm. Borneo: Mt. Mulu BORE NEL SOT NI TIM NAT ORE C. muluensis sp. n. Bulbo distale della spermateca molto sviluppato e sferico; “ductus” a base/larga'e tubuliformes 2). 2.200 EEE 6 Pronoto senza fossetta mediana basale posteriore. Lungh. 2,8 mm. BemeoasMtKmabalusga ete, 1 iis Re C. perpunctata sp. n. Pronoto con due fossette basali mediane posteriori .................... 7 Occhi ridotti, molto pit corti delle tempie; antenne brune con apice dell’undicesimo antennomero giallo-rossiccio. Lungh. 2,9 mm. Borneo: MKinabau EEE U ER C. troglobia sp. n. Occhi molto sviluppati, pit lunghi delle tempie; antenne bruno-rossicce con undicesimo antennomero giallo con base giallo-rossiccia. Lungh. 220 mm Bomeos Mi Kanabalusseier seater ee C. burckhardti sp. n. Punteggiatura o granulosità del capo e del pronoto poco distinte .......... 9 Punteggiatura o granulosità del capo e del pronoto forte ............... 10 Pronoto poco trasverso, con rapporto larghezza/lunghezza pari a 1,13 e con solchi molto prolungati in avanti, oltre il disco dello stesso pronoto; granulosità delle elitre poco distinta; bulbo distale delle spermateca ovale. Lungh. 2,8 mm. Borneo: Mt. Kinabalu ........... C. soror Cameron Pronoto nettamente trasverso, con rapporto larghezza/lunghezza pari a 1,40 e con solchi corti in avanti, non superanti il disco dello stesso pronoto; granulosità delle elitre distinta; bulbo distale delle spermateca sferico. Lungh. 2,1 mm. Borneo: Mt. Kinabalu..... C. subrhomboidalis sp. n. Corpo unicolore nero-bruno, con zampe dello stesso colore; bulbo distale della spermateca con introflessione apicale vestigiale. Lungh. 2,6 mma omeor Mts Kinabalu. SERRE ROLE C. spinae sp. n. Corpo tricolore giallo-rossiccio, con capo ed elitre bruno-rossicci e base dell’addome rossiccia; zampe giallo-rossicce; bulbo distale della sper- mateca senza traccia di introflessione apicale. Lungh. 2,5 mm. Borneo: Sandakan u... Berend RI AIRES C. ashei Pace Pronoto con una fovea o depressione mediano posteriore .............. 12 Pronoto con.due solchi mediani;posteriori +: RESA 13 Capo con una lieve impressione mediana; pronoto con una fovea mediana posteriore; avancorpo nero, con pronoto nero-bruno. Lungh. DIAM Borneo: Samawane: CT C. abdominalis Cameron Capo senza impressione mediana; pronoto con depressione mediana posteriore; avancorpo bruno-rossiccio. Lungh. 2,5 mm. Borneo: Mt. Kinabäli ser te bd C. borneensis Cameron Pronoto poco trasverso, con rapporto larghezza/lunghezza pari a 1,15-1,17 . 14 Pronoto nettamente trasverso, con rapporto larghezza/lunghezza pari a PS PAS pi. wae Vy wis ote ee. ais. don Sa er 15 HOMALOTINI DEL BORNEO 303 14 Capo, pronoto ed elitre coperti da granulosita forte; presenza di solco discale mediano del capo; due solchi obliqui sul pronoto, che presenta un’impressione mediana anteriore. Lungh. 2,2 mm. Borneo: Mt. Pais olen OS e RL C. rifiventris Cameron - Capo e pronoto punteggiati; elitre con forti punti assai radi. Lungh. 25. nm Borneo: Mts Kinabalum. „wa... 22 C. magnilobata sp. n. 15 Bulbordistale-dellasspermatecasreniforme rt... Sram ee eee 16 - Bulbordistale della spermatecarsfenica oipiritorme . "MIN RE 17 16 Capo con fossetta discale; pronoto più trasverso, con rapporto larghezza/lunghezza pari a 1,45; addome bicolore bruno-rossiccio con quarto urite libero bruno; profilo distale della spermateca con un angolo retto. Lungh. 2,1 mm. Borneo: Mt. Matang...... C. sarawakensis Cameron - Capo senza fossetta discale; pronoto meno trasverso, con rapporto larghezza/lunghezza pari a 1,23; addome unicolore bruno-rossiccio con quarto urite libero bruno; profilo distale della spermateca quasi retti- lineo. Lungh. 2,7 mm. Borneo: Mt. Kinabalu....... C. monticola Cameron 177 Granulosita del pronoto poco distinta o confusa nella reticolazione....... 18 - Granwlositaidelgpronotorsalicnte me aa rs een. 19 18 Corpo unicolore giallo-rossiccio; pronoto piü trasverso, con rapporto larghezza/lunghezza pari a 1,46; reticolazione delle elitre superficiale; spermateca piu sviluppata. Lungh. 1,8 mm. Borneo: Mt. Kinabalu 56 0 800 à dot MI OE C. trilobata sp. n. - Corpo bicolore giallo-rossiccio, con tre quarti posteriori delle elitre bruni; pronoto meno trasverso, con rapporto larghezza/lunghezza pari a 1,27; reticolazione delle elitre netta; spermateca meno sviluppata. Euneh22.0/mm> Borneo= Mt=Kanabaluar zn nn C. pansa sp. n. 19 Solchi del pronoto corti, non raggiungenti il disco; bulbo distale della spermateca sferico e molto sviluppato. Lungh. 2,2 mm. Borneo: Mt. | NH oe NU sees RR N deci AM C. cristamagna sp. n. - Solchi del pronoto prolungati fin oltre il disco; bulbo distale della sper- mateca sferico e minuscolo od ovale di media grandezza .............. 20 20 Addome bicolore rossiccio con fascia posteriore bruno-rossiccia; sper- mateca minuscola, con bulbo distale sferico. Lungh. 2,6 mm. Borneo: Miakinaball Eee lare ee ata Ghat C. rhomboidalis sp. n. - Addome unicolore giallo-rossiccio; spermateca con bulbo distale ovale e a pareti ispessite. Lungh. 2,2 mm. Borneo: Mt. Kinabalu scie e ee net C. microincisa sp. n. Mimopisalia leptusoides sp. n. Figg. 155-158 Holotypus d, Sabah, Mt. Kinabalu N.P., above Gunting Lagadan, 3400 m, 6.VIIL.1988, A. Smetana leg. (MHNG). Paratypi: 20 es., stessa provenienza; 11 es., Sabah, Mt. Kinabalu, 3300 m, Panar Laban, 4.V.1987, leg. Burckhardt & Löbl; 1 d, Sabah, Mt. Kinabalu Nat. Pk., Paka Cave, 2995 m, 5.V.1987, leg. A. Smetana. DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e bruno, con pigidio giallo-ros- siccio; antenne rossicce; zampe giallo-rossicce. La punteggiatura del capo & superfi- 304 R. PACE Fico. 153-158 Edeago in visione laterale e ventrale, habitus, e spermateca. 153-154: sp. n.; 155-158: Mimopisalia leptusoides sp. n. Coenonica perpunctata HOMALOTINI DEL BORNEO 305 ciale e assente sul disco. Il pronoto non presenta né punteggiatura né granulosità. La granulosità delle elitre è poco evidente, quella dell’addome è fine. La reticolazione del capo è netta, quella del pronoto è vigorosa, quella delle elitre è distinta e quella del quinto urotergo libero è vigorosa. Edeago figg. 156-157, spermateca fig. 158. COMPARAZIONI. La nuova specie è simile aM. kinabaluensis Pace, 1989, da cui si distingue per la punteggiatura del capo superficiale, il pronoto meno trasverso, le elitre più corte, per l’edeago più sviluppato, con armatura interna più robusta e per il bulbo distale della spermateca meno ovale. ETIMOLOGIA. L’habitus della nuova specie è simile a quello di specie attere del genere Leptusa Kraatz. Per questo è chiamata “con aspetto di Leptusa”. Mimopisalia evoluta sp. n. Figg. 159-162 Holotypus d, Borneo, Sabah, Mt. Kinabalu N.P., below Layang Layang, 2590 m, 1.V.1987, leg. A. Smetana (MHNG). Paratypi: 1 dg, Borneo-Sabah, Mt. Kinabalu N.P., below Layang Layang, 2595 m, 2.V.1987, leg. A. Smetana; 1 d, Borneo, Sabah, M. Kinabalu N.P., below Laban Rata, 3150 m, 7.VII.1988, leg. A. Smetana. DESCRIZIONE. Lungh. 2,2 mm. Corpo lucido e rossiccio; antenne rossicce con i tre antennomeri basali giallo-rossicci; zampe giallo-rossicce. La punteggiatura del capo è indistinta. La granulosità del pronoto e delle elitre è distinta, quella dell’addome è saliente. La reticolazione del capo e delle elitre è distinta, quella del pronoto è nettissima e quella dell’addome è composta da maglie poligonali irregolari chiara- mente visibili. Edeago fig. 160-161, sesto urotergo libero del d fig. 162. COMPARAZIONI. La nuova specie è vicina a M. borneensis (Cameron, 1933). Se ne distingue per le elitre più lunghe, pronoto nettamente più trasverso, per l’intercape- dine apicale dell’edeago stretta e per la notevole differenza di forma del margine pos- teriore del sesto urotergo libero del d. ETIMOLOGIA. Il nome “evoluta” della nuova specie sottolinea la sua differenza rispetto a M. borneensis (Cameron, 1933). Mimopisalia smetanai sp. n. Figg. 163-167 Holotypus d, Borneo, Sabah, Mt. Kinabalu N.P., summit tr. Pondok Lowii, 2300-2400 m, 28.IV.1987, A. Smetana leg. (MHNG). Paratypi: 34 es., stessa provenienza; 3 es., Borneo-Sabah, Mt. Kinabalu N.P., below Layang Layang, 2590 m, 1.V.1987, leg. A. Smetana. DESCRIZIONE. Lungh. 2,4 mm. Avancorpo debolmente opaco, addome lucido. Corpo bruno-rossiccio, con pigidio rossiccio; antenne brune, con i tre antennomeri basali rossicci; zampe giallo-rossicce. La punteggiatura del capo è assente largamente sul disco, ai lati è fittissima, quella dell’addome è assente. La granulosità delle elitre è confusa, quella dei due uroterghi basali è saliente e quella dei restanti uroterghi è superficiale. La reticolazione del capo e del pronoto è vigorosa, quella delle elitre e dell’addome è svanita, solo nel fondo dei solchi trasversi basali degli uroterghi è netta. Edeago figg. 164-165, sesto urotergo libero del d fig. 167, spermateca fig. 166. COMPARAZIONI. La forma dell’edeago della nuova specie è simile a quello di M. kinabaluensis (Cameron, 1933). Ma la nuova specie ha elitre più lunghe del pronoto, mentre nella specie a confronto sono più corte del pronoto. 306 R. PACE FIGG. 159-167 Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del 9 e spermateca. 159-162: Mimopisalia evoluta sp. n.; 163-167: Mimopisalia smetanai sp. n. HOMALOTINI DEL BORNEO 307 ETIMOLOGIA. La nuova specie è dedicata al suo raccoglitore, l’insigne stafilini- dologo Dr. Ales Smetana dell’ “Eastern Cereal and Oilseed Research Centre” di Ottawa, Ontario. Mimopisalia problematica sp. n. Figg. 168-169 Holotypus 2, Borneo, Sabah, Mt. Kinabalu N.P., below Layang Layang, 2600 m, 2- 8.V.1987, int. trap, A. Smetana leg. (MHNG). DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e giallo-bruno; antenne brune con i tre antennomeri basali e l'undicesimo giallo-rossicci; zampe giallo-rossicce. La granulosità del capo e del pronoto è superficiale, quella delle elitre è distinta e quella dell’addome è saliente. Il quinto urotergo libero della 9 presenta granulosità più forte di quella dei precedenti uroterghi. La reticolazione dell’avancorpo è superficiale, quella dell’addome è distinta. Spermateca fig. 169. COMPARAZIONI. La nuova specie si distingue dalle altre del genere per le sue elitre particolarmente lunghe, più lunghe anche di quelle di M. smetanai sp. n., sopra descritta. La larga introflessione apicale del bulbo distale della spermateca distingue ulteriormente la nuova specie da quelle note che presentano bulbo distale della sper- mateca senza introflessione apicale. ETIMOLOGIA. L'assenza del d della nuova specie ha reso problematico il suo inquadramento tassonomico, che in futuro, con il ritrovamento del d potrebbe subire mutamenti. CHIAVE DELLE SPECIE DEL BORNEO DEL GENERE MIMOPISALIA 1 Bhitrespiuscorterdel-pronotorn.2. ERRO A cpm eT ames 2 - Eliterpivmlunshe,delipronoto tesa cee a Eee RS RE + Quarto antennomero trasverso; margine posteriore del sesto urotergo libero del & con sei lobi irregolari, disposti su linea retta, tra le spine lat- erali; intercapedine apicale dell’edeago molto lunga e larga; bulbo dis- tale della spermateca subreniforme. Lungh. 2,4 mm. Borneo: Mt. Kina Dal se ERST. M. borneensis (Cameron) - Quarto antennomero lungo quanto largo o piü lungo che largo; margine posteriore del sesto urotergo libero del d rettilineo; intercapedine api- cale dell’edeago molto stretta; bulbo distale della spermateca ellittico . ..... 5 3 Antenne brune con apice dell’undicesimo antennomero gialli; quarto an- tennomero lungo quanto largo; capo distintamente punteggiato; pronoto più trasverso, con rapporto larghezza/lunghezza pari a 1,25; edeago meno sviluppato, profondamente arcuato al lato ventrale, con armatura interna meno sviluppata; bulbo distale della spermateca ovale allungato. uns mm Borneo= Mt. Kinabalu e eee M. kinabaluensis Pace = Antenne unicolori rossicce; quarto antennomero piü lungo che largo; capo superficialmente punteggiato, senza punteggiatura sul disco; pro- noto meno trasverso, con rapporto larghezza/lunghezza pari a 1,21; edeago piu sviluppato, con profilo ventrale bisinuato, in visione laterale, e con armatura interna più sviluppata; bulbo distale della spermateca ovale corto. Lungh. 2,8 mm. Borneo: Mt. Kinabalu... . M. leptusoides sp. n. (N°) 308 R. PACE 4 Antenne brune, con undicesimo antennomero giallo-rossiccio; elitre molto piu lunghe del pronoto. Lungh. 1,8 mm. Borneo: Mt. Kinabalu nc de i Us M. problematica sp. n. - Antenne rossicce o brune con undicesimo antennomero mai di colore differente: elitre meno lunghe.. .. ....... 2... RR 5 5 Margine anteriore del pronoto arcuato; assenza di fossette del pronoto; margine posteriore del sesto urotergo libero del & con quattro lobi e numerose spine; edeago esile, con un solo tubulo interno. Lungh. ZAMmeBormneog ME Knaben M. evoluta sp. n. 6 Margine anteriore del pronoto bisinuato; presenza di fossette mediane posteriori del pronoto; margine posteriore del sesto urotergo libero del d seghettato; edeago robusto, con forte armatura interna. Lungh. 2ÆmmeBomeoMtE Kinabalu... y=... LN M. smetanai sp. n. Neosilusa terminalis sp. n. Figg. 170-174 Holotypus d, Borneo, Sabah, Mt. Kinabalu Nat.Pk., HQ at Liwagu Rv., 1500 m, 30.1V.1987, A. Smetana leg. (MHNG). Paratypi: 7 es., stessa provenienza; 1 es., Borneo, Sabah, Mt. Kinabalu, 1750 m, 21.1V.1987, Burckhardt & Löbl leg.; 1 es., Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1560-1640 m, 24.1V.1987, A. Smetana leg.; 2 2 2 e 6 es.,, Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1500 m, 25.IV.1987, A. Smetana leg.; 1 es., Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1500-1550 m, 27.IV.1987, A. Smetana leg.; 4 es., , Borneo, Sabah, Mt. Kinabalu N.P.,1750 m, 27.1V.1987, Burckhardt & Löbl leg.;1 es., Borneo, Sabah, Mt. Kinabalu Nat. Pk., HQ at Liwagu Rv., 1500 m, 30.IV.1987, leg. A. Smetana; 2 es., Borneo-Sabah, Crocker Ra., 1550-1650 m, 16.V.1987, leg. Burckhardt & Löbl; 2 es., Borneo, Sabah, Crocker Ra., 1600 m, Km 51 rte. Kinabalu-Tambunan, 18.V.1987, Burckhardt & Löbl leg.; 31 es., Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1560 m, 3. VIII.1988, leg. A. Smetana; 2 es., Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv. tr., 1520 m, 11.IV.1988, A. Smetana leg.; 1 es., Borneo-Sabah, Mt. Kinabalu Nat. Pk., HQ Silau-Silau Tr., 1550 m, 2.1X.1988, leg. A. Smetana. DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e nero, con addome bruno-ros- siccio; antenne rossicce con i tre antennomeri basali giallo-rossicci e l’undicesimo giallo paglierino; zampe bruno-rossicce con tibie e tarsi rossicci. La granulosita del capo e del pronoto é fittissima e forte, quella delle elitre & composta di granuli allun- gati robusti, assenti presso il margine posteriore. Gli uroterghi sono nudi, tranne per la presenza di alcune setole. La punteggiatura del fondo dei solchi trasversi basali degli uroterghi in avanti è limitata da un archetto per ciascun punto. Edeago figg. 171-172, sesto urotergo libero del d fig. 174, spermateca fig. 173. COMPARAZIONI. Per il colore nero del corpo e la granulosità dell’avancorpo, la nuova specie è sicuramente affine a N. borneensis (Cameron, 1928) (“olim” Plagiusa Cameron, 1928: 416; Neosilusa borneensis: Bernhauer & Scheerpeltz, 1926: 540) del Monte Murud. Le due specie si distinguono per i caratteri dati nella seguente chiave. HOMALOTINI DEL BORNEO 309 CHIAVE DELLE SPECIE DEL BORNEO DEL GENERE NEOSILUSA 1 Eorpoltossiccio.con meta posteriore delle.elitre bruna. . 2..........:... 2 - COMO MELO ZI EI MR EL. à. AI EURE 3 2 Pronoto finemente granuloso; decimo antennomero più lungo che largo. Lungh. 2,3 mm. Sumatra, Mascarene, Madagascar, Borneo ME RR RITO ee à, N. tropica (Bernhauer) - Pronoto rugosamente punteggiato o con punteggiatura estremamente fit- ta; decimo antennomero trasverso. Lungh. 2,4 mm. Mascarene, India, Malesia, CinasGiapponesBomeo ih. a. 45.0650... - N. ceylonica (Kraatz) 3 Avancorpo subopaco; margine degli uroterghi oscuramente rossicci; terzo antennomero nero, undicesimo giallo-rossiccio; nono antennomero più lungo che largo. Lungh. 3,3 mm. Borneo: Mt. Murud. site SI a I VITI A AIA IR N. borneensis (Cameron) - Avancorpo lucido; margine degli uroterghi nero; terzo antennomero giallo-rossiccio, undicesimo giallo paglierino; nono antennomero trasverso. Lungh. 2,9 mm. Borneo: Mt. Kinabalu........ N. terminalis sp.n. Linoglossa borneensis sp. n. Figg. 175-176 Holotypus 9, Sabah, Mt. Kinabalu, 3300 m, Panar Laban, 4.V.1987, leg. Burckhardt & Löbl (MHNG). DESCRIZIONE. Lungh. 2,5 mm. Corpo lucido e bruno, con pigidio rossiccio; antenne perdute in fase di raccolta; zampe giallo-rossicce. La punteggiatura del capo è indistinta. La granulosità del pronoto è assente, quella delle elitre è molto superficiale e quella dell'addome è saliente. La reticolazione del capo è netta, quella del pronoto è vigorosa, quella delle elitre è superficiale e quella degli uroterghi è distinta. Spermateca fig. 176. COMPARAZIONI. Finora il genere Linoglossa Kraatz non era noto per il Borneo. La spermateca della nuova specie è simile a quella di L. angustata (Motschulsky, 1858) dell’India. Se ne distingue per i caratteri della seguente chiave. L’esemplare di L. angustata qui a confronto è il tipo ® di L. bifoveolata Kraatz, 1859, sinonimo di L. angustata. 1 Capo con profondo solco longitudinale; depressione posteriore mediana del pronoto divisa da una carena longitudinale mediana; punteggiatura delle elitre netta e rada, irregolarmente distribuita; ognuno dei tre solchi basali dei tre uroterghi basali, con due foveole nel fondo; spermateca iù 7elunch928:mmeIndiaez ar ua va L. angustata (Motschulsky) = Capo senza solco longitudinale; depressione posteriore mediana del pronoto senza carena longitudinale mediana; elitre coperte di granulosita molto superficiale; solchi basali dei tre uroterghi basali, senza foveole nel fondo; spermateca fig. 176. Lungh. 2,5 mm......... L. borneensis sp. n. 310 R. PACE 0.1 mm FiGG. 168-173 Habitus, spermateca ed edeago in visione laterale e ventrale. 168-169: Mimopisalia problemati- ca Sp. n.; 170-173: Neosilusa terminalis sp. n. Se en HOMALOTINI DEL BORNEO 311 176 I mm 174 178 0,1 mm 0,1 mm 179 12777. 0,1 mm FicG. 174-179 Sesto urotergo libero del 4, habitus e spermateca.174: Neosilusa terminalis sp. n.; 175-176: Linoglossa borneensis sp.n.; 177: Linoglossa angustata (Motschoulsky); 178: Coenonica soror Cameron, holotypus; 179: Coenonica nigrita Cameron, holotypus, nuovo sinonimo di Coeno- nica soror Cameron. 312 R. PACE RINGRAZIAMENTI Rivolgo i miei più cordiali ringraziamenti a coloro che mi hanno affidato in studio il raro materiale oggetto del presente lavoro: il Dr. Ales Smetana di Ottawa, e il Dr. Ivan Löbl, già del Museo di Storia Naturale di Ginevra, il Prof. Herbert Franz di Mödling (Austria) e l'Ing. Jin Janak di Rtyne nad Bilinou (Repubblica Ceca). Per il prestito di tipi ringrazio il Dr. PM. Hammond e il Dr. M. Brendell del Museo di Storia Naturale di Londra, il Dr. L. Zerche del DEI di Eberswalde (Berlino), il Dr. D. Drugmand dell’Institut Royal des Sciences Naturelles de Belgique di Bruxelles e il Dr. A.F. Newton del Field Museum of Natural History di Chicago. BIBLIOGRAFIA BERNHAUER, M.1911. Zur Staphylinidenfauna Ostindiens und der Sundainseln (3. Beitrag. Schluss). Entomologische Blätter für Biologie und Systematik der Käfer 7: 86-93. BERNHAUER, M.1914. Neue Staphylinen der indo-malaiischen Fauna. Verhandlungen der zoolo- gisch-botanischen Gesellschaft in Wien 64: 76-109. BERNHAUER, M.1915. Neue Staphyliniden der indo-malaiischen Fauna, insbesondere der Sunda- Insel Borneo. Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 65: 134- 158. BERNHAUER, M.1916. Neue Staphyliniden des indo-malaiischen Faunengebietes, besonders der Philippinen. Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 66: 418- 431. BERNHAUER, M. & SCHEERPELTZ, O. 1926. Coleoperorum Catalogus, pars 82, Staphylinidae VI: 499-988. Berlin. BLACKWELDER, R.E.1952. - The generic names of the beetle family Staphylinidae with an essay on genotypy. Bulletin of the United States National Museum 200: 1-483. BRYANT, G. E.1919. Coleoptera in Borneo. Entomologist’s Monthly Magazine 55: 70-76. CAMERON, M. 1920. New species of Staphylinidae from Singapore, Part III. Transactions of the Entomological Society of London 1920: 212-284. CAMERON, M.1928. Staphylinidae from Mt. Poi & Mt. Penrissen, with descriptions of new species. Sarawak Museum Journal 11: 399-414. CAMERON, M. 1930. Staphylinidae from British North Borneo with Descriptions of New Species. Journal of the Federated Malay States Museums 16 (1-2):160-168. CAMERON, M. 1933. Staphylinidae (Col.) from Mount Kinabalu. Journal of the Federated Malay States Museums 17(2): 338-360. CAMERON, M. 1936. Fauna Sumatrensis. Bijdrage No. 77, Staphylinidae (Col.). Tijdschrift voor Entomologie 76: 1-24. CAMERON, M. 1939a. The Fauna of British India, including Ceylon and Burma. Coleoptera, Staphylinidae, 4 (Part 1). pp. 1-410. London. CAMERON, M. 1941. New Species of Staphylinidae (Col.) from the Philippines. Annals and Magazine of Natural History (11) 8: 379-403. CAMERON, M. 1943. New species of Staphylinidae (Col.) from Borneo. Entomologist’s Monthly Magazine 79: 183. CAMERON, M. 1950. New species of Staphylinidae (Col.) from the Malay Peninsula. Annals and - Magazine of Natural History 3:1-40; 89-131. FAUVEL, A. 1878. Les Staphylinides des Moluques et de la Nouvelle Guinée. Annali del Museo Civico di Storia Naturale di Genova 12:171-315. FAUVEL, A. 1905. Staphylinides exotiques nouveaux (3e Partie). Revue d’Entomologie 24: 113- 147. HOMALOTINI DEL BORNEO 313 HAMMOND, P.M. 1984. An annotated Check-List of Staphylinidae (Insecta: Coleoptera) recorded from Borneo. Sarawak Museum Journal 33: 187-218. HEER, O. 1839. Fauna Coleopterorum Helvetica. Pars I, fasc. 2. Orelii, Fuesslini et Sociorum, Turici (Zurigo), pp. 145-360. KRAATZ, G. 1856. Naturgeschichte der Insekten Deutschlands. Abteilung 1, Coleoptera. Vol. 2, Staphylinii. Nicolai, Berlin, pp. 1-376. KRAATZ, G. 1857. Genera Aleocharinorum Illustrata. Linnaea Entomologica 2: 1-43. Kraatz, G. 1859. Die Staphyliniden-Fauna von Ostindien, insbesonders der Insel Ceylan. Archiv fiir Naturgeschichte 25: 1-196. MOULTON, J. C. 1912. “Where Wallace trod”: being some account of an entomological trip to Mt. Serambu, Sarawak, Borneo. The Entomologist 45: 213-217, plates V-VI, 246-251. MOTSCHULSKY, V. DE. 1858. Enumération des nouvelles espèces de Coléoptères rapportées de ses voyages. Bulletin de la Société impériale des Naturalistes de Moscou 3: 204-264. PACE, R. 1984a. Aleocharinae delle Mascarene, parte I: tribù Myllaenini, Pronomaeini, Oligotini e Bolitocharini (Coleoptera Staphylinidae) (XLV Contributo alla conoscenza delle Aleocharinae). Revue suisse de Zoologie 91: 3-36, 159 figg. PACE, R. 1984b. Due Aleocharinae attere appartenenti a due nuovi generi raccolte in Estremo oriente dal Dr. Ivan Löbl (Coleoptera Staphylinidae) (XLIX Contributo alla conoscenza delle Aleocharinae). Revue suisse de Zoologie 91: 895-901, 30 figg. PACE, R. 1986. Aleocharinae dell’Asia sudorientale raccolte da G. de Rougemont (Coleoptera, Staphylinidae) (LXXII Contributo alla conoscenza delle Aleocharinae). Bollettino del Museo civico di Storia Naturale di Verona 23: 139-237, 291 figg. PACE, R. 1987. Coenonica philippina Bernhauer, 1916 e specie affini (Coleoptera Staphylinidae) (LXXXIII Contributo alla conoscenza delle Aleocharinae). Bollettino dell’ Associazione Romana di Entomologia 42: 37-40, 12 figg. PACE, R. 1989. Aleocharinae attere del Monte Kinabalu (Borneo) (Coleoptera Staphylinidae) (XCVIII Contributo alla conoscenza delle Aleocharinae). Revue suisse de Zooogie 96: 3-8, 8 figg. PACE, R. 1992. Aleocharinae del Vietnam (Coleoptera, Staphylinidae). Nouvelle Revue d’Ento- mologie (N.S.) 9: 119-129. PACE, R. 1993. Aleocharinae della Cina (Coleoptera Staphylinidae). Bollettino del Museo civico di Storia Naturale di Verona 17: 69-126. PACE, R. 1990. Aleocharinae delle Filippine (82° contributo alla conoscenza delle Aleocharinae) (Coleoptera, Staphylinidae). In: BERTI, N. [ed.]. Miscellanées sur les Staphylins. Mémoires du Museum national d’ Histoire naturelle (A) 147: 57-113, 273 figg. PACE, R. 1998. Aleocharinae della Cina: Parte I (Coleoptera, Staphylinidae). Revue suisse de Zoologie 105: 139-220, 234 figg. PACE, R. 2000. Aleocharinae della Thailandia (Coleoptera, Staphylinidae) (144° Contributo alla conoscenza delle Aleocharinae). Bollettino del Museo regionale di Scienze naturali di Torino 17: 39-86. PACE, R. 2002. Nuovi generi di Aleocharinae del Borneo (Coleoptera, Staphylinidae). Revue suisse de Zoologie 109 : 189-240. ana on Wi u Al Tenino i E toa a NX? egal REVUE SUISSE DE ZOOLOGIE 110 (2): 315-324; juin 2003 Redescription and new species of Alexidia (Coleoptera: Staphylinidae: Scaphidiinae) I. LÖBL! & R. A. B. LESCHEN? 1 Muséum d’histoire naturelle, case postale 6434, CH-1211 Geneva 6, Switzerland. E-mail: ivan.lobl@mhn.ville-ge.ch 2 New Zealand Arthropod Collection, Landcare Research, Private Bag 92170, Auckland, New Zealand. E-mail: leschenr@landcareresearch.co.nz Redescription and new species of Alexidia (Coleoptera: Staphylinidae: Scaphidiinae). - The Neotropical genus Alexidia Reitter and its type species A. rogenhoferi Reitter are redescribed. Following additional new species are described: A. carltoni sp. n. from Ecuador, A. dybasi sp. n. from Panama, and A. plaumanni sp. n. from Brazil. A key to the species of Alexidia is provided. Key-words: Coleoptera - Staphylinidae - Scaphidiinae - Alexidia - Neotropics - taxonomy. INTRODUCTION The present classification of Scaphisomatini is unclear with many genera requiring detailed study and redescription. The groupings of genera (Achard, 1924) apparently does not reflect monophyly and have been ignored in recent taxonomic treatments (Löbl, 1971, 1990, 1992). Because of this problem we are in the process of examining the phylogenetic relationships of the genera in an attempt to restructure the classification of the group, especially to place taxa like Neotropical Alexidia, which have not been studied since their description and are rare in collections. In this paper we redescribe the genus and its type species Alexidia rogenhoferi Reitter, 1880, and add three new species from Brazil, Ecuador, and Panama. Several characters of Alexidia rogenhoferi were not examined because only a single type is available for study, and this specimen is in poor state. Material is deposited in the following collections: Field Museum of Natural History, Chicago (FMNH), Museum d’histoire naturelle, Geneve (MHNG), Museum d’Histoire Naturelle, Paris (MHNP), Museo de Zoologia, Pontificia Universidad Catolica del Ecuador, Quito (MZUC), New Zealand Arthropod Collection, Auckland (NZAC) and Snow Entomological Museum, University of Kansas (SEMC). Manuscript accepted 03.01.2003 316 I. LOBL & R. A. B. LESCHEN TAXONOMY Alexidia Reitter, 1880 Alexidia Reitter, 1880: 43. Type species Alexidia rogenhoferi Reitter, 1880. Diagnosis. Maxillary palpus aciculate. Antennomeres 3 and 4 elongate; 7, 9 and 10 each with long and narrow apical stalk and subapical rim of short setae in addition to long setae. Galea wider than long; brush apical and radulate. Surface of mentum with spines. Prothoracic corbiculum present. Metendosternum with stem present. Profemoral ctenidium absent. Mesotibia with a single ventral spine. Empodium bisetose. Description. Dorsal vestiture reduced. Labral setae present and simple (not examined in A. rogenhoferi). Mandible bidentate apically, subapical serrations present (at least on left mandible). Maxillary palpus aciculate; one subapical seta present on palpus 2 (broken off in A. rogenhoferi). Galea wider than long; brush apical and radu- late. Inner and basal setae absent from lacinia, but apical setae extending to subapical area. Hypopharynx with 2 setae on adoral surface; setae spinate. Labial palp 3-seg- mented; terminal labial palpomere not aciculate, inserted apically and strongly curved; subapical palpomere with one seta. Mentum (not examined in A. rogenhoferi) with anterior edge straight; surface with spines. Submaxillary area of head with microtubu- late ducts present (not examined in A. rogenhoferi). Gular area with transverse clusture of pores (not examined in A. rogenhoferi). Frontoclypeal suture present. Eye entire. Antennal insertion below slight ridge and present at midline of eye. Antenna filiform (not seen in A. rogenhoferi which has only the scape and pedicel present in the type specimen but according to Reitter’s description long and as in Baeocera Erichson); antennomeres 3 and 4 elongate; antennomeres 7, 9 and 10 each with long and narrow apical stalk and subapical rim of short setae in addition to long setae. Anterior tentorial tendon absent. Prothoracic corbiculum present. Prosternum poorly developed; anterior margin of procoxal cavity asetose. Hypomeron in lateral view completely visible; apex not projecting beyond pronotum. Prothoracic carina prominent with a bead; not completely visible in dorsal view. Anterior margin of pronotum with a bead. Posterior angle rounded and not extending below ventral edge of elytra; not extending to anapleural suture. Prosternum with spine present. Mesosternum with prepectus (meso- sternal space); secondary and median lines absent. Mesosternal lines present; connecting with mesocoxal cavity, impunctate; not parallel with outer margins of pro- coxal rests. Mesosternal process paxillate. Mesepimeron absent or fused with mesosternum. Mesocoxa wider than intercoxal process; coxa round. Meso- and meta- sternum separate. Submesocoxal lines arcuate, impunctate or punctate. Metasternum without setose patch; primary setae present and located on the disc; transverse premetaxocal lines absent; descrimen absent; intercoxal plate present as a single plate. Metepisternal suture absent or present. Metepisternum visible in ventral view; posterior line absent. Metendosternum with stem present. Pteronotum with scutellum concealed below elytra; width about 1/2 entire width of pteronotum; transverse basal line or carina complete. Metacoxae separate. Brick-wall membranes present on abdominal ventrites 1 to 4. Ventrite one with intercoxal line; submetacoxal lines absent; submetacoxal bead impunctate; primary setae present. Primary setae present on ALEXIDIA (STAPHYLINIDAE: SCAPHIDIINAE) 317 ventrites 2-4; 2 on each segment. Abdominal vestiture absent. Paratergites absent. Hind wings present or absent. Elytron with basal stria present or absent; sutural striae shortened or elongate; basal and sutural striae not connected; epipleural stria present; sutural spines and apical serrations absent. Metacoxal process triangular. Profemoral ctenidium absent. Mesofemora in cross-section rounded; subapical seta present and not spine like. Tibiae smooth. Mesotibia distinctly longer than mesotarsus; one ventral mesotibial spine present, about as long as two thirds of first tarsomere. Metatarsi smooth; mesotarsomere 1 slightly shorter than tarsomeres 2 and 3 combined. Empodium bisetose. Discussion. Alexidia is similar to other members of Scaphisomatini, especially those taxa with aciculate maxillary palpi, elongate antennomeres 3, and basal pronotal angles rounded and not extending to anapleural suture. Whereas other Scaphisomatini, membres of Scaphoxium Löbl excepted, have two ventral mesotibial spines, Alexidia can be distinguished from them by having a single spine (Fig. 4). Scaphoxium has approximate coxal cavities, and is very distinctive from Alexidia. The aedeagus of Alexidia has an elongate and symmetrical internal sac with tripartite basal sclerites that are similar to those seen in species of New Zealand Brachynopus Broun, and a very long, irregularly folded ejaculatory duct inside the median lobe (Fig. 5). Members of Amalocera Erichson have also the ejaculatory duct much longer than the median lobe (Löbl, 1974) but it is coiled, sclerotized and forms a flagellum. A very long, mem- branous ejaculatory duct is present also in the species rich Baeocera group lenta (Löbl, 1971, 1992). In these species the ejaculatory duct is everted outside the median lobe, the sclerites of the internal sac are distinctive, and the basal angles of pronotum extend below the edge of the elytra. Alexidia may be readily distinguished from Amalocera by the abruptly narrowed and long apical part of the antennomeres 7, 8 and 9 (Fig. 1), the aciculate apical segment of maxillary palpi (Fig. 3), the lacinia with setae (Fig. 3) situated apically while in Amalocera (Fig. 2) the setae are present also on the subapical margin of the lacinia. Despite the similarity to Amalocera and Brachynopus, the relationship of Alexidia to other members of Scaphisomatini is uncertain and must await further analysis. KEY TO THE SPECIES OF ALEXIDIA 1 Elytra with sutural striae shortened, not extended to base ................ 2 - Elytra with sutural stria entire, extended from apex to base .............. 3 2 Blyiralackancabasalisttiaes <3 m en A. rogenhoferi Reitter = Elytrawithbasalistii = Ne PR RSR AUTA A. carltoni Sp. n. 3 Elytral punctation much more distinct that pronotal punctation. Hind wines teduced. Metepisteralisuture present. ere À. plaumanni Sp. n. - Elytra and pronotum with similar, very fine punctation. Hind wings developed: Metepisternalsuturerabsent ME ae oe A. dybasi sp. n. Alexidia rogenhoferi Reitter, 1880 Syntype female, labelled: Neugranada Chevrolat (hand written) / Rogenhoferi Rttr / (hand written)/ TYP. REITTER (printed) / TYPE (red, printed) / Muséum Paris Coll. Générale (printed) (MHNP). 318 I. LOBL & R. A. B. LESCHEN Description. Length 1.45 mm. Body strongly convex dorsally, moderately convex ventrally. Body and appendages light reddish-brown, tibiae and tarsi slightly lighter than femora. Head with frons wide, about 2.5 times as broad as eyes long. Eyes flat, shorter than dorso-ventral eye diameter. Pronotum strongly narrowed anteriorly, with lateral edges strongly arcuate and lateral stria concealed (dorsal view). Apical stria entire, not interrupted at middle, at middle finer than laterally. Basal lobe short. Punctation, microsculpture and pubescence completely absent (200 times magni- fication). Hypomera not impressed, impunctate. Elytra strongly narrowed apically, with lateral edges and striae exposed in dorsal view. Apical margins truncate except at angles, edentate. Inner apical angle not prominent. Sutural striae short and shallow, slightly diverging anteriorly, from apex extending almost to sutural mid-length. Adsutural areas flat. Basal striae absent. Epipleura wide at base, gradually narrowed apically. Supra-epipleura oblique, large, lacking basal bead, equally wide in anterior half and about 1.5 times as wide as epipleura near base, gradually narrowed apically. Elytral punctation reduced, indicated by scattered darkened point, microsculpture absent. Hind wings absent. Ventral side of thorax impunctate and lacking micro- sculpture. Mesosternal shield flat, lacking striae or impression. Mesepimera and mesepisterna fused. Width of intercoxal process slightly smaller than mesocoxal width, about as width of metacoxal process. Metasternum short between meso and metacoxae, flattened in middle, lacking impressions. Submesocoxal lines arcuate, impunctate. Length of submesocoxal areas as about as half of shortest interval between submeso- coxal lines and metacoxal margin. Metepisterna concealed, very narrow, with impunc- tate suture. Abdominal ventrites 1 and 2 impunctate, lacking microsculpture. Ventrite 1 lacking impressions. Tibiae almost evenly thick. Pro- and mesotibiae straight, meta- tibiae very weakly curved. Male characters unknown. Alexidia carltoni sp. n. Figs 8-10 Holotype female: Ecuador Azuay 50km NW Cuenca 2470m, 2.Jan. 1992 C. Carlton, R. Leschen # 94 ex: berlese (SEMC). Paratypes: same data as holotype, 1 male 3 females (SEMC, MHNG, MZUC). Description. Length 1.75 mm. Body strongly convex dorsally, moderately convex ventrally, piceous, apex of abdomen light brown to yellowish, appendages slightly lighter than body. Lateral contours of pronotum and elytra continuously arcuate. Head with frons wide, hardly twice as broad as eyes length. Eyes flat, shorter than dorso-ventral eye diameter. Antennae long, with segments 3 to 6 and 8 about equally wide, each bearing scattered, long setae. Apical half of antennal segment 11 with short pubescence, in addition to long, scattered setae. Pronotum strongly narrowed anteriorly, with lateral edges strongly arcuate and lateral stria concealed (dorsal view). Apical stria widely interrupted in middle. Basal lobe short. Punctation very fine, microsculpture absent, pubescence inconspicuous. Hypomera hardly impressed, as finely punctate as pronotum, with flat process posterior procoxae delimited by longitudinal stria. Scutellum concealed. Elytra strongly narrowed api- cally, with lateral edges and striae exposed in dorsal view. Apical margins truncate ALEXIDIA (STAPHYLINIDAE: SCAPHIDIINAE) 319 except at angles, edentate. Inner apical angles not prominent. Sutural striae parallel, shallow, impunctate, shortened, extending up to anterior third of sutural length. Adsutural areas flat. Basal striae entire, very shallow, joined to lateral striae. Epipleura almost equally wide in anterior two thirds, narrowed posteriorly. Supra-epipleura oblique, large, delimited anteriorly by bead, widest at middle and at widest point slightly more than twice as wide as epipleura. Elytral punctation sparse and very fine, about as fine as that on pronotum. Hind wings absent. Mesepimera partly fused to metepisterna, with very fine suture, about as long as half of interval to mesocoxa. Mesepisterna and metasternum lacking microsculpture, very finely punctate. Metasternum narrow between meso- and metacoxae. Submesocoxal lines arcuate, very finely punctate. Submesocoxal areas shorter than interval between lines and meta- coxae. Metepisterna fused to metasternum, suture indicated by straight line. Abdo- minal ventrites distinctly microsculptured, impunctate. Ventrite 1 with very shallow lat- eral impressions. Ventrites 1 and 2 with one pair of primary setae, ventrites 3 and 4 with two pairs of primary setae. Submetacoxal line impunctate. Protibiae straight, narrowed basally, meso-and metatibiae almost evenly thick. Male. Segments 1 to 3 of protarsi widened and bearing tenant setae, segment | almost as wide as apex of tibia. Meso- and metatibiae arcuate. Aedeagus 0.66 mm long, as Figs 8 to 10. Female. Meso- and metatibiae weakly curved. Comments. This species may be easily distinguished from A. rogenhoferi by the presence of elytral basal striae. In addition, it differs by the large body size, the basal part of supra-epipleura more than twice as wide as basal part of epipleura, and the mesepisterna only partly fused. Alexidia plaumanni sp. n. Figs 1, 3-7 Holotype male: Brazil: Santa Catharina; VIII. 1954, leg. F. Plaumann (MHNG). Paratypes: Brazil, Nova Teutonia 27°11’S 52°23’ W, 300-500m, I. 1957, F. Plaumann, 1 male, 2 females (NZAC, MHNG); same data but X.1956, 2 males (MHNG) and X. 1957, 1 fe- male (FMNH); same data but VI. 1952, 2 males 1 female (MHNG). Description. Length 1.8-2.1 mm. Similar to A. carltoni in most characters, differing as follows: Frons 2.5 times as wide as eye length. Pronotal punctation fairly dense and fine, visible at low (20 x) magnification. Elytra with sutural striae not shortened, joined to basal striae. Elytral punctation distinctly coarser than pronotal punctation. Supra-epipleura hardly 3 times as wide as epipleura. Mesosternal process lacking striae. Mesepimera distinct, oblique, very short, not extending beyond mesepis- ternal suture, hardly as long as third of interval to mesocoxa. Middle part of metaster- num coarser punctate than lateral parts of metasternum, and with distinct, very short pubescence. Submesocoxal areas as long as or slightly longer than interval between submesocoxal lines and metacoxae. Mesocoxal process about as wide as 2/3 of meso- coxal width, narrower than metacoxal process. Metepisterna not fused to metasternum, with distinct, slightly curved suture. Abdominal ventrite 1 with lateral impression and distinctly punctate in middle, ventrites 1 and 2 with single pair of primary setae, ven- trites 3 and 4 with 2 pairs of primary setae. Protibiae straight, narrowed basally, meso- and metatibiae almost evenly thick. 320 I. LÖBL & R. A. B. LESCHEN Fics 1-4 1: Alexidia plaumanni sp. n., antennomeres 7 to 11; 2: Amalocera dentipes Löbl, maxilla with palpus; 3: Alexidia plaumanni sp. n., maxilla with palpus; 4: Alexidia plaumanni sp. n., apical part of mesotibia with basal tarsomere. Scale bars = 0.1mm. Male. Protarsi with segments 1 to 3 widened, segment 1 about as wide as apex of protibiae. Meso- and metatibiae arcuate. Aedeagus 0.64 — 0.81 mm long, as Figs 5 LO Female. Mesotibiae hardly curved, metatibiae slightly arcuate. ALEXIDIA (STAPHYLINIDAE: SCAPHIDIINAE) 321 Fics 5-10 Aedeagi in Alexidia. 5 to 7: A. plaumanni sp. n., internal sac (7) in detail; 8 to 10: A. carltoni sp. n., internal sac (10) in detail. Scale bars = 0.2 mm (5, 6, 8, 9) and 0.1 mm (7, 10). 322 I. LOBL & R. A. B. LESCHEN Alexidia dybasi sp. n. Figs 11- 14 Holotype male, labelled: El Valle, Cocle Prov. (tail to Las Minas) PANAMA; II: 23-1959 alt. 2400-2600ft./CNHM Panama Zool. Exped. (1959) H.S. Dybas leg./ground debris Berlese (B-348) (FMNH). Paratype female, with same data as holotype (MHNG). Description. Length 1.4 — 1.55 mm. Body very dark, piceous, elytra lighter at apex, apical abdominal segments and legs rufous, antennae distinctly lighter than legs. Lateral contours of pronotum and elytra continuously arcuate. Head with frons moderate wide, about 1.5 times as wide as eyes length. Eyes convex, about as long as two thirds of dorso-ventral eye diameter. Antennae long, with segments 3 to 6 and 8 about equally wide, each bearing scattered, long setae. Apical half of antennal segment 11 with short pubescence, in addition to long, scattered setae. Pronotum strongly narrowed anteriorly, with lateral edges strongly arcuate and lateral stria visible (dorsal view). Apical stria entire, not interrupted in middle. Basal lobe short. Punctation very fine, microsculpture absent, pubescence very fine but distinct. Hypomera weakly impressed, as finely punctate as pronotum. Elytra strongly narrowed apically, with lateral edges and striae exposed in dorsal view. Apical margins truncate except at angles, edentate. Inner apical angles not prominent. Sutural striae diverging from apex to mid-length, parallel anterior mid-length, shallow, impunctate, extending to and along base to form basal striae joined to lateral striae. Adsutural areas flat. Epipleura almost equally wide in anterior two thirds, narrowed posteriorly. Supra-epipleura oblique, large, delimited anteriorly by bead, widest at middle and at widest point twice as wide as epipleura. Elytral punctation sparse and very fine, about as fine as that on pronotum. Hind wings well developed. Mesepimera with inner part completely fused to metepisterna. Mesepisterna and metasternum lacking microsculpture, very finely punctate. Metasternum narrow between meso- and metacoxae. Submesocoxal lines arcuate, finely punctate mesally, impunctate laterally. Submesocoxal areas as long as half of interval between lines and metacoxae. Mesocoxal process about as wide as metacoxal process and as wide as three fourth of mesocoxal width. Metepisterna fused to metasternum, suture indicated by straight, weak line. Abdominal ventrites distinctly microsculptured, impunctate. Ventrite 1 lacking lateral impressions. Ventrites 1 and 2 with one pair of primary setae, ventrites 3 and 4 with two pairs of primary setae. Submetacoxal line impunctate. Protibiae straight, narrowed basally, meso- and meta- tibiae almost evenly thick. Male. Protarsi with segments | to 3 moderately widened, much narrower than apex of tibiae, with tenant setae. Mesotibiae and metatibiae distinctly curved, meso- tibiae narrowed between mid-length and apical fifth. Aedeagus 0.52 mm long, as Figs 11 to 14. Female. Meso- and metatibiae slightly arcuate. Comments. This species may be distinguished from its congeners by the hind wings well developed. It is similar to A. plaumanni in having elytra with entire sutural striae but divers distinctly by the very fine elytral punctation and fused metepisterna. ALEXIDIA (STAPHYLINIDAE: SCAPHIDIINAE) 323 Fics 11-14 Aedeagus in Alexidia dybasi sp. n., internal sac (13) in detail, apical part of median lobe and paramere (14) in lateral view at higher magnification. Scale bars = 0.2 mm (11, 12) and = 0.1 mm (13, 14). ACKNOWLEDGEMENTS For the loan of material we thank Steve Ashe and Robert Brooks (SEMC), Nicole Berti (MNHP), and Alfred Newton (FMNH). Field work undertaken by RABL in Ecuador was made possible through support from the SEMC, permits issued by Sergio Figueroa (Ministerio de Agricultura y Granaderia, Quito), and logistical support by Luis Coloma, Giovanni Onore, and Gustavo Morejon and his family. 324 I. LOBL & R. A. B. LESCHEN REFERENCES ACHARD, J. 1924. Essai d’une subdivision nouvelle de la famille des Scaphidiidae. Annales de la Société entomologique de Belgique 65: 25-31. LößL, I. 1971. Scaphidiidae von Ceylon (Coleoptera). Revue suisse de Zoologie 78: 937-1006. Losi, I. 1974. New species of the genus Amalocera Erichson from Brazil (Coleoptera, Scaphidiidae). Studies on the Neotropical Fauna 9: 39-45. Los, I. 1990. Review of the Scaphidiidae (Coleoptera) of Thailand. Revue suisse de Zoologie 97: 505-621. LosL, I. 1992. The Scaphidiidae (Coleoptera) of the Nepal Himalaya. Revue suisse de Zoologie 99: 471-627. REITTER, E. 1880. Die Gattungen und Arten der Coleopteren-Familie: Scaphidiidae meiner Sammlung. Verhandlungen des Naturforschenden Vereines in Brünn 18: 35-49. REVUE SUISSE DE ZOOLOGIE 110 (2): 325-353; juin 2003 New records of Pauropoda (Myriapoda) with descriptions of new species from Rwanda and Réunion (Pauropoda and Symphyla of the Geneva Museum XII) Ulf SCHELLER Häggeboholm, Häggesled, S-53194 Järpäs, Sweden. New records of Pauropoda (Myriapoda) with descriptions of new species from Rwanda and Réunion (Pauropoda and Symphyla of the Geneva Museum XII). - A collection of 751 Pauropoda (Myriapoda) specimens has been studied. Three species are new to science and are described: Allopauropus afer sp. n. and Samarangopus rwandaensis sp. n. from Rwanda and Sphaeropauropus reunionensis sp. n. from Réunion and Sri Lanka. Thirty-five more species have been recognised and the species lists for many countries have been lengthened. The following species are here recorded for the first time from: Switzerland, 3 species [Stylopauropus neglectus Remy, Acopauropus consobrinus (Remy), Trachypauropus cordatus (Scheller)]; Greece, 3 species (Allopauropus helophorus Remy, Brachypauropus superbus Hansen, Acopauropus attemsi Hasenhütl); Slovenia, 2 species (Pauropus bagnalli Remy, Stylopauropus limitaneus Remy); the Maltese Islands, 3 species [Allopauropus danicus (Hansen), A. gracilis (Hansen), A. helveticus (Hansen)]; Turkey, 4 species [Aco- pauropus hastatus (Attems), A. tetrastichus Scheller, Trachypauropus cordatus (Scheller), T. glomerioides Tomésvary]; Réunion, 1 species (Sphaeropauropus reunionensis sp. n.); Sri Lanka, 2 species, (Sphaero- pauropus nepalensis Scheller, S. reunionensis sp. n.); Malaysia, 1 species (Allopauropus rastifer Remy); Indonesia, 1 species [Allopauropus danicus (Hansen)]; Rwanda, 2 species (Allopauropus afer sp. n., Samarangopus rwandaensis Sp. n.) and Argentina, 1 species (Stylopauropoides subantarc- ticus Scheller). Key-words: Europe - Asia - Africa - Rwanda - Réunion - South America - Pauropoda - taxonomy - biogeography - soil zoology. INTRODUCTION The study of the taxonomy of Pauropoda was long limited to manually collected and therefore sparse material. During later years automatical extraction methods in combination with skilful sorting out assistants, have been of great importance for the Manuscript accepted 23.01.2003 326 U. SCHELLER studies in several pedozoological fields, among them the biodiversity and distribution of Pauropoda. Thanks to Dr Bernd Hauser, it has been possible to study valuable pauropods from the collections of the Natural History Museum, Geneva. A scattered material of various sizes from many countries and collected mainly by means of automatical extraction methods, a total of 751 specimens, was available for examination. Dr Hauser himself collected most of these specimens and his great faculty in finding valuable collecting sites has resulted in a rich material. Several other soil zoologists have contributed as well: Drs Villy Aellen', Claude Besuchet, Herman Gisinî, Ivan Löbl, Volker Mahnert, Paul Schauenberg, who all have belonged to or are belonging to the staff of the Museum in Geneva. Moreover, some collectors outside the Museum, but in close contact with it, have deposited material studied here. They are: Dr Romano Dallai (Siena), Dr Egon Horak (Zürich), Dr Pierre Strinati (Cologny, Geneva), Dr Konrad Thaler (Innsbruck), and Dr Philippe Werner (Ollon-Chemignon). Three species are new to science and are described. They are Allopauropus afer sp. n. and Samarangopus rwandaensis sp. n. from Rwanda and Sphaeropauropus reunionensis sp. n. from Réunion and Sri Lanka. Thirty-five more species have been recognised and the lists of species of many countries have been lengthened. Several species are here added for the first time to the lists of the following countries: Switzerland, 3 species [Stylopauropus neglectus Remy, Acopauropus consobrinus Remy, Trachypauropus cordatus (Scheller)]; Greece, 3 species (Allopauropus helophorus Remy, Brachypauropus superbus Hansen, Acopauropus attemsi Hasenhütl); Slovenia, 2 species (Pauropus bagnalli Remy, Stylopauropus limitaneus Remy); the Maltese Islands, 3 species [Allopauropus danicus (Hansen), A. gracilis (Hansen), A. helveticus (Hansen)]; Turkey 4 species [Acopauropus hastatus (Attems), A. tetrastichus (Scheller), Trachypauropus cordatus (Scheller), T. glomerioides Tömösväry]; Réunion, 1 species (Sphaeropauropus reu- nionensis sp. n.); Sri Lanka, 2 species, (Sphaeropauropus nepalensis Scheller, S. reunionensis Sp. n.); Malaysia, 1 species (Allopauropus rastifer Remy); Indonesia, 1 species [Allopauropus danicus (Hansen)]; Rwanda, 2 species (Allopauropus afer sp. n., Samarangopus rwandaensis sp. n.) and Argentina, | species (Stylopauropoides sub- antarcticus Scheller). All the material, preserved in alcohol (with one single exception), is deposited in the collections of the Department of Arthropods and Entomology I, Natural History Museum of Geneva. SYSTEMATICS PAUROPODIDAE Genus Allopauropus Silvestri, 1902 Subgenus Allopauropus s. str. 1. Allopauropus (A.) brevisetus Silvestri Allopauropus brevisetus Silvestri, 1902: fasc. 95, no. 12. NEW RECORDS OF PAUROPODA 327 Material examined. CROATIA, Velebit Mountains, at road between Karlobag and Gospi3, on the Gospi3 side of the pass, oak forest, sifting, Berlese extraction, 1 juv. 5!, 2.X.1970 (Loc. Ju-70/1, leg. B. Hauser). Total number. 1 specimen. General distribution. The main range covers southern Europe from France to Romania, Bulgaria and Greece. Known also from Great Britain, Switzerland, Austria and the USA. 2. Allopauropus (A.) danicus (Hansen) Pauropus danicus Hansen, 1902: p. 376-378, pl. III, fig. 4. Material examined. MALTESE ISLANDS, Malta, Dingli, Buskett Forest, soil sample, 1 ad. 9(2), 1 subad. 8( 2), 2.V.1976 (Leg. V. Aellen & P. Strinati). GREECE, Peloponnesus, Achaia, Erymanthos Massif, above Kalusion, alt. 980 m, under Abies cephalonica and Quercus coccifera, soil sample, Berlese extraction, 2 ad. 9( 2), 19 subad. 8(83, 109,1 sex?), 4 juv. 6, 1 juv. 5, 1.V.1980 (Loc. Sam-80/18, leg. B. Hauser); Messenia, near the road Areopolis - Kalamata, before Pygi, alt. 230 m, under Quercus coccifera, near a small stream, soil sample, Berlese extraction, 1 ad. 92), 4 subad. 8(14,39), 2 juv. 6, 2 juv. 5, 18.V.1981 (Loc. Art-81/15, leg. B. Hauser). - Attica, Mount Hymette, near Markopoulon, close to the cave “Vavrona 2”, under Pistacia lentiscus, soil sample, Berlese extraction, alt. 30 m, 5 juv. 6, 1 juv. 5, 20.11.1982 (Loc. Att-82/26, leg. B. Hauser). - Southern Island Arc, Karpathos, Lastos Massif, near the road Aperi — Spoa, alt. 430 m, under Pinus brutia, soil sample, Berlese extraction, 1 ad. 9(d), 2 subad. 8( 2), 1 juv. 6, 10.III.1979 (Loc. Kar-79/8b, leg. B. Hauser). AUSTRIA, Innsbruck, Martinswand, needle litter under pines, soil sample, Berlese extraction, 15 ad. 9(10d, 59), 3 subad. 8(1d, 22), IV-V.1964 (Leg. K. Thaler); ibidem, 5 ad. 918,42), 2 subad. 8(d), 17.V.1969 (Loc. I-69/1, leg. B. Hauser) and 2 ad. 9(4 ), 29.1V.1969 (Loc. I-69/5, leg. K. Thaler). Morocco, Rif Atlas, El-Gouzat, alt. 1050 m, soil sample under evergreen oaks, Berlese extraction, 2 ad. 9(S), 1 juv. 6, 3 juv. 5, 2.VI.1978 (Loc. Mar-78/12, leg. B. Hauser); Smila, alt. 630 m, soil sample under Pinus radiata, Berlese extraction, 1 subad. 8(£), 2. VI.1978 (Loc. Mar- 78/13, leg. B. Hauser). INDONESIA, East Java, Baluran Game Reserve, dry forest, alt. 100 m, in lava soil, 2 juv. 6, 4.VII.1973 (Loc. As-73/3, leg. P. Schauenberg). Total number. 83 specimens. General distribution. A wide range species known from most countries in Europe, from North and East Africa, South Asia and the Americas. Remarks. A. danicus is here reported for the first time from the Maltese Islands and Indonesia. 3. Allopauropus (A.) maroccanus Remy & Moyne Allopauropus (A.) maroccanus Remy & Moyne, 1960: p. 73-76, fig. 1. Material examined. Morocco, Moyen Atlas, Tazzeka District, Bab-Azhar, under cork oaks, soil sample, Berlese extraction, 7 ad. 9(64, 19), 2 subad. 8( 9), 3 juv. 5, 1.VI.1978 (Loc. Mar-78/9, leg. B. Hauser). Total number. 12 specimens. General distribution. Known only from Morocco and Sri Lanka. Taxonomical remarks. After having been able to study specimens from the type series (Morocco, Midelt, alt. 1500 m), it is evident that the temporal organs of this | Abbreviations: ad. ..., subad. ... and juv. ... = an adult, a subadult or a juvenile speci- men with the number of pairs of legs indicated. 328 U. SCHELLER species are provided with large posterior pistils of the shape illustrated by Scheller (1970) in the description of A. (A.) prope maroccanus from Sri Lanka (Southern Province, Hikkaduwa and Deniyaya). These specimens have to be referred to A. (A.) maroccanus Remy & Moyne. The specimens reported above from Bab-Azhar also correspond with Remy & Moyne’s types. 4. Allopauropus (A.) afer sp. n. Figs 1-13 Type material. Holotype: ad. 9(9 ), RWANDA, Rangiro, alt. 1800 m, sifting in forest, 6.VI- 11.1973 (Loc. Rwa-73/8, leg. P.Werner). Paratype: 1 ad. 9(4), same locality and date (Leg. P. Werner). Total number. 2 specimens. Diagnosis. A. (A.) afer sp. n. has many characters in common with A. (A.) jeannelli Remy from Mount Elgon (Remy, 1935b) but is distinguished from it by the shape of the lobes and the appendages of the anal plate (lateral lobes somewhat curved and blunt in A. (A.) afer sp. n., straight and pointed in A. (A.) jeanelli Remy; postero- median lobe large and linguiform, not short, triangular, pointed). A. (A.) ruwenzorien- sis Remy from Mount Ruwenzori (Remy, 1960) is similar in the same way. Good distinguishing characters are e.g. the shape of the posterior part of the anal plate [linguiform in A. (A.) afer sp. n., cleft deeply in A. (A.) ruwenzoriensis Remy], the pubescence of the T; (distal part with branched hairs, not naked and annulate) and the shape of the antennal globulus (almost spherical with narrow stalk, not pear-shaped with thicker stalk). The new species may also have distant relationships with A. (A.) simulator Remy from south-western France (Remy, 1947a) because there are similar- ities in the general plan of the anal plate and the styli, but they are easy to distinguish (e.g. pygidial setae a, very short in A. (A.) afer sp. n., very long in A. (A.) simulator Remy; appendages of the anal plate long, stalked and foliform, not short and clavate without stalk). Etymology. From Latin afer = African. DESCRIPTION Length. 0.90(0.92)? mm. Head. Tergal setae subcylindrical, annulate, blunt. Relative lengths of setae, 1s! row: a, = 10, a, = 11; 2nd row: a, = 11, a, = 11(12), az = 10; 314 row: a, = 13(14), a, = 2(20); 4% row: a, = 9(10), a, = 19, a; = 27, a,= 18; lateral group setae: I, = 25, I, =? 1, = ?. The ratio a,/a,- a, in 1st row 0.8, in 2nd row 0.5, in 3™ row (0.9)1.1 and in 4% row 1.0. Temporal organs ovoid, in tergal view about as long as their shortest distance apart. Head cuticle glabrous. 2 Measurements taken from adult paratypes are given in brackets. Fics 1-13 Allopauropus (A.) afer sp. n., holotype, ad. 9(2). 1, head, median and right part, and right antenna, tergal view; 2, collum segment, median and left part, sternal view; 3, tergite VI, posteromedian and left posterior part: 4, T;: 5, Ty: 6, seta on coxa of leg 9; 7, seta on trochanter of leg 9; 8, tarsus of leg 9; 9, left genital papilla, anterior view; 10, pygidium, median and left part, sternal view; 11, pygidial seta a,; 12, pygidial seta a,; 13, anal plate, lateral view. Pubescence only partly drawn in Fig. 8. Scale a: Figs 2-7, 9; b: Figs 1, 8, 10-13. ee NEW RECORDS OF PAUROPODA 329 330 U. SCHELLER Antennae. Segment 4 with at least 5 cylindrical striate-annulate setae, r thinnest. Relative lengths of setae: p = 100, p’ = 82, p” = 59, p” = 14, r = 44. Tergal seta p 0.8 of the length of tergal branch r. The latter fusiform, (2.8)3.2 times as long as its greatest diameter and 1.1 times as long as sternal branch s which is 1.9(2.2) times as long as its greatest diameter and with its anterodistal corner distinctly truncate. Seta q as tergal- anterior setae of 4th segment but somewhat thinner, as long as s. Relative lengths of flagella (with base segments included) and base segments: F, = 100, bs, = (10)11; F, = (43)46, bs, = 8; F; = 115, bs; = 9(10). F, thinner than F, and F;; F, 4.1(4.3) times as long as f, F, and F; 1.8 and 5.2 times as long as s respectively. Distal calyces hemi- spherical, glabrous, distal part of flagella axes widened only between calyx and first lamella. Globulus g 1.1 times as wide as long and with thin and very short, almost cylindrical stalk, ~ 8 bracts and flattened capsule; diameter of g as long as greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment furcate, primary branch cylindrical, annulate, blunt; secondary branch rudimentary, blunt, glabrous. Sublateral setae (1.6)1.7 times as long as submedian ones; process cleft anteriorly and with very short pubescence; appendages glabrous, subcylindrical, with wide caps. Setae on tergites of about the same length, somewhat lanceolate and with oblique pubescence; 4+4 setae on tergite I, 6+6 on II-V and 4+2 on VI. Submedian pos- terior setae on tergite VI 0.2 of their interdistance and 1.6 times as long as pygidial setae a,. Tergite I somewhat and tergites II-VI distinctly granular. Bothriotricha. Relative lengths of bothriotricha: T, = 100, T, = 2(81), 73 = (92)100, 7, = 108, 7, = 149. Their axes simple, very thin in all but 73. T,, T, and T, similar to each other: short, erect, simple pubescence hairs proximally, somewhat longer outwards, on distal half branched and partly arranged in whorls. T, similar but with thicker axis. 7; thin with short oblique pubescence of simple hairs. Genital papillae (paratype). Glabrous, 1.8 times as long as their greatest diameter, inner margin straight, outer one evenly curved in distal half; seta 0.3 of the length of papilla. Legs. Setae on coxa and trochanter of leg 9 simple, somewhat clavate, densely pubescent with hairs arranged in whorls; more anteriorly these setae are furcate with a rudimentary glabrous secondary branch. Tarsus of leg 9 very little tapering, (3.3)3.8 times as long as its greatest diameter. Proximal seta tapering, pointed, with short oblique pubescence, 0.2 of the length of tarsus and 1.2(1.5) times as long as distal seta which is subcylindrical and annulate. Cuticle of tarsus minutely granular. Pygidium. Tergum. Posterior margin with deep posterior indentation between st, sides steep and bottom almost flat. Relative lengths of setae: a, = 10, a, = (40), a; = ?, st = 8. These setae of very different shape: a, fusiform, blunt, a, lanceolate, both with short oblique pubescence; st subspherical bladders with short thin cylindrical stalk and short erect pubescence; a,. a, and st converging. Distance a, - a, (2.4)2.5 times as long as a,; distance a, - a, 5 times longer than distance a, - a3; distance st - st 4.2 times as long as st and (1.9)2.0 times as long as distance a, - a,. Cuticle minutely pubescent. Sternum. Posterior margin between b, with broad indentation and low median lobe with shallow posteromedian incision. Setae tapering, with short oblique pubes- cence. Relative lengths of setae (a, = 10): b, = 62, b, = (46)48, b; = 16(20). The 5b, NEW RECORDS OF PAUROPODA 331 somewhat curved inwards. b, 0.9 of their interdistance; b, 1.4(1.5) times as long as distance b, - b>; b;0.3(0.4) of distance b; - b3. Anal plate narrowest anteriorly, 1.3 times as long as broad, with lateral, blunt, posterolaterally directed horn-like processes; posterior part of plate extended into a linguiform lobe, from its base two diverging appendages protruding in posterolateral direction; appendages 0.7 of the length of plate, leaf-shaped, 1.4 times as long as wide, with thin cylindrical stalk and a dense oblique pubescence. Subgenus Decapauropus Remy, 1957 (Remy, 1957a) 5. Allopauropus (D.) aristatus Remy Allopauropus (D.) aristatus Remy, 1936a: p.19-22, figs 6-9. Material examined. Morocco, Rif Atlas, El-Gouzat, alt. 1050 m, under evergreen oaks, soil sample, Berlese extraction, 8 ad. 9(2), 41 juv. 6, 7 juv. 3, 2.VI.1978 (Loc. Mar-78/12, leg. B. Hauser). Total number. 56 specimens. General distribution. The species is known in Europe from France, Germany, Switzerland, Bulgaria, Spain, Yugoslavia, Greece; in Africa from Algeria, Morocco, Gambia, Madagascar; and in North America from the USA and Canada. 6. Allopauropus (D.) corsicus Remy Allopauropus (D.) corsicus Remy, 1940: p. 48-50, figs 2-4. Material examined. GREECE, Southern Island Arc, Crete, at the road Sitia - Iraklion, near Sfaka, small ravine, alt. 200 m, under Quercus coccifera, soil sample, Berlese extraction, 1 ad. 9(2), 13.11.1979 (Loc. Kar-79/12a, leg. B. Hauser). Total number. 1 specimen. General distribution. Outside Greece (Scheller, 1977b) this species is known from France, Switzerland and Bulgaria. 7. Allopauropus (D.) cuenoti (Remy) Decapauropus cuenoti Remy, 1931: p. 67-83, figs 1-12. Material examined. GREECE, Elidia, S of Pirgos, forest near the Alfios river, alt. 30 m, under Pinus sp., soil sample, Berlese extraction, 1 juv. 6, 22.11.1982 (Loc. Att-82/32, leg. B.Hauser). - Central Greece, Peloponnesus, Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, soil samples, Berlese extraction, under date palm, 1 juv. 5, and under Eucalyptus, 1 juv. 6, 23.III.1982 (Loc. Att-82/37 and Att-82/39 respectively, leg. B. Hauser); at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, under Abies cephalo- nica and Pinus sp., soil sample, Berlese extraction, 3 ad. 9( 9), 25.1II.1982 (Loc. Att-82/48, leg. B. Hauser). — Attica, Keratea, before the northern entrance of road, alt 160 m, under fig tree, 2 ad. 9(?), 21.11.1982 (Loc. Att-82/30, leg. B. Hauser). — Aegean Islands, Ikaria, near the road Plumarion -Monokampion, alt. 420 m, high water level deposits, Berlese extraction, 1 juv. 5, 24.IV.1980 (Loc. Sam-80/6, leg. B. Hauser). - Samos, near Kosmathei, close to the entrance of the cave “Tsitse Tripa”, alt. 510 m, old pine stumps, Berlese extraction, 2 ad. 9(£), 5 subad. 8(2), 4 juv. 6, 4 juv. 5, 25.1V.1980 (Loc. Sam-80/11, leg. B. Hauser). Morocco, Grand Atlas, “Ifri-El-Kaid”, moist guano sample from the beginning part of the cave, Berlese extraction, 1 juv. 6, 5.VI.1978 (Loc. Mar-78/25, leg. P. Strinati). Total number. 25 specimens. General distribution. Most European countries, North Africa, Madeira, the Canary Islands, Réunion and California. 332 U. SCHELLER 8. Allopauropus (D.) furcula Silvestri Allopauropus furcula Silvestri, 1902: fasc. 95, no. 7. Material examined. GREECE, Elidia, S of Pirgos, forest near the Alfios river, alt. 30 m, under Quercus pubescens, soil sample, Berlese extraction, 1 ad. 9(9), 22.11.1982 (Loc. Att- 82/33, leg. B.Hauser). — Peloponnesus, Laconia, Talanta, at the road Neapolis - Monemvasia, alt. 210 m, under Quercus coccifera, soil sample, Berlese extraction, 3 ad. 911d, 22), 4 subad. 8(1d, 32), 3 juv. 6, 24.11.1982 (Loc. Att-82/45, leg. B. Hauser). - Messenia, at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, soil sample under Abies cephalonica and Pinus sp., Berlese extraction, 1 ad. 9(2), 25.III.1982 (Loc. Att-82/48, leg. B. Hauser). Total number. 12 specimens. General distribution. Known from South Europe, Madeira and the Canary Islands. 9. Allopauropus (D.) gracilis (Hansen) Pauropus gracilis Hansen, 1902: p. 395-397, pl. V, fig. 3. Material examined. MALTESE ISLANDS, Malta, Dingli, Buskett Forest, soil sample, 1 ad. 9(d), 2.V.1976 (Leg. V. Aellen & P. Strinati). GREECE, Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, under Araucaria, soil sample, Berlese extraction, 5 subad. 8(£), 1 juv. 6, 2 juv. 5, 23.III.1982 (Loc. Att-82/38, leg. B. Hauser) and under Eucalyptus, soil sample, Berlese extraction, 1 juv. 5, 23.III.1982 (Loc. Att-82/39, leg. B. Hauser) and under laurel, 2 ad. 9 (6, 9), 23. 11.1982 (Loc. Att-82/40, leg. B.Hauser); at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, soil sample under Abies cephalonica and Pinus sp., Berlese extraction, 1 ad. 99), 25.11.1982 (Loc. Att-82/48, leg. B. Hauser); near the road Areopolis - Kalamata, before Pygi, alt. 230 m, under Ouercus coccifera, near a small stream, soil sample, Berlese extraction, 46 ad. 948, 429), 32 subad. 8(1d, 309, 1 sex?), 27 juv. 6, 6 juv. 5, 1 juv. 3, 18.V.1981 (Loc. Art- 81/15, leg. B. Hauser); ibidem, near Aghios Nikon, alt. 290 m, under Ouercus macrolepis, 1 ad. 9(d), 1 subad. 8(2), 18.V.1981 (Loc. Art-81/13, leg. B. Hauser). - Arcadia, near the road Kandila - Skotini (Nemea), alt. 920 m, under Quercus coccifera, soil sample, Berlese extraction, 2 juv. 6, 17.V.1981 (Loc. Art-81/11, leg. B. Hauser). - Laconia, Taigetos Massif, alt. 960 m, soil sample, Berlese extraction, 12 ad. (23,92, 1 sex?), 4 subad. 8(9), 13 juv. 6, 20.V.1981 (Loc. Art-81/18, leg. B. Hauser). - Achaia, near Patras, Panachaikon Massif, Kastritsion, near the road behind the village, alt. 590 m, soil sample from under a log at the foot of a large plane tree, 1 ad. 9(G ), 22.1V.1980 (Loc. Sam-80/3, leg. B. Hauser); Erymanthos Massif, above Kalusion, alt. 980 m, under Abies cephalonica and Quercus coccifera, soil sample, Berlese extraction, 1 ad. AG), 1.V.1980 (Loc. Sam-80/18, leg. B. Hauser). — Attica, near Megara, Kaki Skala, above the motorway at 48 km sign of the old road, under Pistacia terebinthus, soil sample, Berlese extraction, 1 subad. 8(£), 21.1V.1980 (Loc. Sam-80/2, leg. B. Hauser). — Aegean Islands, Ikaria, near the road Plumarion - Monokampion, alt. 420 m, high water level deposits, Berlese extrac- tion, 2 ad. 9(2), 1 juv. 6, 24.1V.1980 (Loc. Sam-80/6, leg. B.Hauser). Samos, near Kosmathei, close to the entrance of the cave “Tsitse Tripa”, alt. 510 m, old pine stumps, Berlese extraction, 1 subad. 8(2), 3 juv. 6, 1 juv. 5, 2 juv. 3, 25.IV.1980 (Loc. Sam-80/11, leg. B. Hauser). Total number. 171 specimens. General distribution. A. gracilis seems to have a (sub)cosmopolitan range. It is one of the species most often met with in Europe but has also been reported from Africa, South Asia and the Americas. Remarks. It is here reported for the first time from the Maltese Islands. 10. Allopauropus (D.) helveticus (Hansen) Pauropus helveticus Hansen, 1902: p. 390-392, pl. IV, fig. 5. Material examined. MALTESE ISLANDS, Malta, Dingli, Buskett Forest, soil sample, 1 sub- ad. 8(®), 2.V.1976 (Leg. V. Aellen & P. Strinati). ie DE NEW RECORDS OF PAUROPODA 333 GREECE, Peloponnesus, Arcadia, Panachaikon Massif, near the road Kastritsion - Patras, alt. 260 m, under Cupressus sempervirens, soil sample, Berlese extraction, 2 juv. 6, 16.11.1982 (Loc. Att-82/5, leg. B. Hauser). - Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, under Araucaria, soil samples, Berlese extraction, 1 subad. 8( 2 ), 23.1.1982 (Loc. Att-82/38, leg. B. Hauser) and under laurel, 1 juv. 6, 23.11.1982 (Loc. Att-82/40, leg. B. Hauser). — Attica, near Loutropirgos, above the motorway Elefsis — Megara, 1-2 km behind the entrance to the high-way, alt. 90 m, under Pinus halepensis, Berlese extraction, 1 juv. 6, 15.11.1982 (Loc. Att- 82/1, leg. B. Hauser). — Central Greece, Acarnania, near Gavrolimni, S of the road, alt. 170 m, under Pistacia lentiscus, soil sample, Berlese extraction, 1 ad. 9(¢), 17.11.1982 (Loc. Att-82/13, leg. B. Hauser). SWITZERLAND, Vaud, the cave Grande Rolat, Berlese extraction, 1 juv. 6 mounted on slide, 26. VIII.1981 (Leg. P. Strinati, V. Aellen & C. Besuchet). Total number. 8 specimens. General distribution. The species may be Holarctic. It has been collected in most European countries, North Africa, the Azores and North America. Remarks. A. helveticus is here reported for the first time from the Maltese Islands. 11. Allopauropus (D.) helophorus Remy Allopauropus helophorus Remy, 1936b: p. 132-133, fig. 1. Material examined. GREECE, Peloponnesus, Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, under Eucalyptus, soil sample, Berlese extraction, 1 juv. 6, 23.III.1982 (Loc. Att-82/39, leg. B. Hauser). Total number. 1 specimen. General distribution. The known range covers an area from Belgium and northern France in the northwest to Bosnia, Serbia and Romania in the southeast. The locality reported above, the first one from Greece, extends the range considerably in southeastern direction. 12. Allopauropus (D.) multiplex Remy Allopauropus (A.) multiplex Remy, 1936c: p. 75-76 and 1936d: p. 315-316, fig. 3. Material examined. GREECE, Aegean Islands, Samos, near Kosmathei, close to the entrance of the cave “Tsitse Tripa”, alt. 510 m, old pine stumps, Berlese extraction, 1 subad. 8(2), 1 juv. 3, 25.IV.1980 (Loc. Sam-80/11, leg. B. Hauser). SWITZERLAND, Basle-Land, western Sundgau, near small stream Dorrenbach, under Heracleum and Dactylis, moist loess, 1 juv. 6, 16.VIII.1942 (Loc. Ae 254/g, leg. H. Gisin). Total number. 3 specimens. General distribution. A. (A.) multiplex is known from the Western Palaearctic Region only and occurs there from Sweden in the north to Morocco in the south, from Great Britain in the west to Greece in the east. 13. Allopauropus (D.) pectinatus (Hansen) Pauropus pectinatus Hansen, 1902: p. 388-390, pl. IV, fig. 4. Material examined. GREECE, Peloponnesus, Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, under Araucaria, soil sample, Berlese extraction, 1 juv. 6, 23.IIT.1982 (Loc. Att-82/38, leg. B. Hauser). Total number. 1 specimen. General distribution. A. (D.) pectinatus is a West Palaearctic species known from France and Spain in the west to Greece in the east, and also from Morocco and Algeria. 334 U. SCHELLER 14. Allopauropus (D.) productus Silvestri Allopauropus productus Silvestri, 1902: Fasc. 95, no. 9. Material examined. GREECE, Peloponnesus, Laconia, Talanta, at the road Neapolis - Monemvasia, alt. 210 m, under St. John’s bread tree, soil sample, Berlese extraction, 1 ad. (2), 12 juv. 6, 24.11.1982 (Loc. Att-82/46, leg. B. Hauser). - Messenia, at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, under Abies cephalonica and Pinus sp., soil sam- ple, Berlese extraction, 1 ad. 9(9), 25.11.1982 (Loc. Att-82/48, leg. B. Hauser). Morocco, Rif Atlas, El-Gouzat, alt. 1050 m, under evergreen oaks, soil sample, Berlese extraction, 1 juv. 6, 2.VI.1978 (Loc. Mar-78/12, leg. B. Hauser). Total number. 15 specimens. General distribution. A probably Holarctic species with its main distribution around the Mediterranean. 15. Allopauropus (D.) rastifer Remy Allopauropus rastifer Remy, 1948: p. 573-574. Material examined. MALAYSIA, Teman Negara National Park, alt. 240 m, lowland rain forest, soil sample, Berlese extraction, 1 ad. 99), 27.V1.1973 (Loc. As-73/1, leg. P. Schauen- berg). 7 Total number. 1 specimen. General distribution. A. rastifer is a rare species, only 13 specimens were earlier known from 7 sites in Kenya, Madagascar, Réunion, Pondichéry and Sri Lanka. The locality reported here, the first one from Malaysia, extends the range of this species to Southeast Asia. Taxonomical remarks. The species was described from a single female from Kenya (Remy, 1948). In the specimen reported by Remy (1956b) from Madagascar (sub nomine A. rastrifer Remy) the setae a; of the pygidial tergum are proportionally longer than in the type specimen, the posterior margin of the pygidial tergum is different in shape and the anal plate is probably so too. Similar deviations from the original description can be observed in the specimen from Réunion (Remy, 1956c). The anal plate is there proportionately long, its lateral appendages short and the pygidial tergum has a posteromedian semicircular lobe not mentioned by Remy in his description of the species. In the Malaysian specimen reported here, the temporal organs are propor- tionately long and the antennal globulus g more ovoid than in the type specimen. Moreover are the coxal setae of leg 9 simple. Observations on the specimens referred to A. rastifer indicate that the species might not be monotaxic. 16. Allopauropus (D.) vulgaris (Hansen) Pauropus vulgaris Hansen, 1902: p. 392-395, pl. V, fig. 2. Material examined. GREECE, Peloponnesus, Messenia, at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, under Abies cephalonica and Pinus sp., soil sample, Berlese extraction, 1 juv. 6, 25.1II.1982 (Loc. Att-82/48, leg. B.Hauser). — Aegean Islands, Ikaria, near the road Plumarion - Monokampion, alt. 420 m, high water level deposits, Berlese extrac- tion, 1 ad. 92), 1 juv. 5, 24.1V.1980 (Loc. Sam-80/6, leg. B. Hauser). Total number. 3 specimens. NEW RECORDS OF PAUROPODA 335 General distribution. The species is widely distributed in Europe and has also been collected in Africa, Sri Lanka and North America. 17. Allopauropus (D.) zaianus Remy Allopauropus zaianus Remy, 1952: p. 154-155, fig. 4. Material examined. Morocco, Rif Atlas, El-Gouzat, alt. 1050 m, under evergreen oaks, soil sample, Berlese extraction, 1 ad. 9(¢), 2.VI.1978 (Loc. Mar-78/12, leg. B. Hauser). Total number. 1 specimen. General distribution. Known only from the southwestern Palaearctic: Portugal, Spain, Morocco and Algeria. Genus Pauropus Lubbock, 1867 18. Pauropus bagnalli Remy Pauropus Bagnalli Remy, 1935a: p. 1-2. Material examined. SLOVENIA, Julian Alps, Triglav, on Tominskova trail, near Aljazev dom, alt. 1070 m, beech forest, under stone, 1 ad. 9(2), 26.VIII.1967 (Loc. Tg-67/8, leg. B. Hauser). Total number. | specimen. General distribution. Previously known from France, Germany, Austria and the Slovak Republic. It is here reported for the first time from Slovenia. 19. Pauropus huxleyi Lubbock Pauropus huxleyi Lubbock, 1867: p. 182-185, pl. 10, figs 1-19. Material examined. GREECE, Elidia, S of Pirgos, forest near the Alfios river, alt. 30 m, under Pinus sp., soil sample, Berlese extraction, 5 ad. 18,4%), 9 subad. 8(2¢, 59, 2 sex?), 6 juv. 6, 6 juv. 5, 22.11.1982 (Loc. Att-82/32, leg. B. Hauser). — Peloponnesus, Laconia, Taigetos Massif, alt. 960 m, soil sample, Berlese extraction, 1 ad. 9(9 ), 20.V.1981 (Loc. Art-81/18, leg. B. Hauser). Total number. 27 specimens. General distribution. P. huxleyi, in the sense of Lubbock, has been mentioned from many European countries, from North America and New Zealand and may have a wide range. However, it has often been confused with P. lanceolatus Remy, and at present it is impossible to delimit its range. 20. Pauropus numidus Remy Pauropus numidus Remy, 1947b: p. 66-68, fig.1. Material examined. Morocco, Moyen Atlas, Tazzeka District, Bab-Azhar, under cork oaks, soil sample, Berlese extraction, 2 juv. 6, 1.VI.1978 (Loc. Mar-78/9, leg. B. Hauser). Total number. 2 specimens. General distribution. The species is known from Spain and from a dubious record from Austria; outside Europe from Morocco, Algeria and the Canary Islands. Genus Stylopauropoides Remy, 1956 21. Stylopauropoides subantarcticus Scheller Stylopauropoides subantarcticus Scheller, 1974a: p. 62-65, fig. 2. 336 U. SCHELLER Material examined. ARGENTINA, Patagonia, W Ushuaia, Monte Susana, south slope, alt. c. 300 m, in litter of a Nothofagus pumilis-betuloides forest, Berlese extraction, 1 ad. AG), 26.11.1975 (Loc. Am-75/3, leg. E. Horak). Total number. 1 specimen. General distribution. The species is here reported for the first time from Argentina. It was previously known only from one of the Crozet Islands in the French Subantarctic Territory. Its occurrence in southernmost Patagonia may indicate that the species is more or less widespread. Genus Stylopauropus Cook, 1896 Subgenus Stylopauropus s. str. 22. Stylopauropus (S.) neglectus Remy Stylopauropus (S.) neglectus Remy, 1962: p. 80-81, fig. 2. Material examined. SWITZERLAND, Ticino, Grotta del Mago, alt. 350 m, 1 ad. 9(9), 1 subad. 8(¢ ), 24.V1.1975 (Loc. Ju-75/15, leg. B. Hauser). Total number. 2 specimens. General distribution. S. neglectus was previously known from France, Austria and Italy. Remarks. The species is here reported for the first time from Switzerland. 23. Stylopauropus (S.) pedunculatus (Lubbock) Pauropus pedunculatus Lubbock, 1867: p. 185, pl. 10, fig. 20. Material examined. SLOVENIA, Sneznik, alt. 1500 m, beech forest, under stones, 1 ad. 9(2), 11.IX.1969 (Loc. Ju-69/32, leg. V. Mahnert); ibidem, alt. ca. 1100 m, 1 ad. AG), 11.1X.1969 (Loc. Ju-69/33, leg. V. Mahnert). Between Godovic and Crni Vrh, beech forest, 2 ad. 9(d), 1 stad.?, 12.1X.1969 (Loc. Ju-69/35, leg. V. Mahnert). GREECE, Peloponnesus, Achaia, Erymanthos Massif, above Kalusion, alt. 980 m, under Abies cephalonica and Quercus coccifera, soil sample, Berlese extraction, 1 ad. 9(2), 1.V.1980 (Loc. Sam-80/18, leg. B. Hauser). - Messenia, near the road Areopolis -Kalamata, before Pygi, near a small stream, alt. 230 m, under Quercus coccifera, soil sample, Berlese extraction, 1 ad. 92), 3 subad. 8(13, 22), 1 juv. 6, 18.V.1981 (Loc. Art-81/15, leg. B. Hauser). - Arcadia, at a monastery near Kandila, alt. 870 m, 1 subad. 8(sex?), 16.V.1981 (Loc. Art-81/6, leg. B. Hauser). - Laconia, Taigetos Massif, alt. 960 m, under Abies cephalonica, soil sample, Berlese extraction, 6 ad. 9(9), 9 subad. 8(8 2, 1 sex?), 7 juv. 6, 6 juv. 5, 2 juv. 3, 20.V.1981 (Loc. Art-81/18, leg. B. Hauser). — Messenia, Kalamata District, near Messini, Analipsis, alt. 20 m, under laurel, 2 sub- ad. 8(2), 1 juv. 6, 23.11.1982 (Loc. Att-82/40, leg. B. Hauser). Messenia, at the road Sparta - Kalamata, on the Kalamata side of the pass, alt. 1000 m, under Abies cephalonica and Pinus sp., soil sample, Berlese extraction, 1 ad. 9(2), 25.11.1982 (Loc. Att-82/48, leg. B. Hauser). - Corinth, near the road Klenia - Aghionorion, alt. 640 m, in the cave “Tripa tou Kalivaki”, 1 ad. 9(2), 1 subad. 8(2), 13.V.1981 (Loc. Art-81/1, leg. B. Hauser). - Attica, Keratea, before the northern entrance of the road, alt. 160 m, under fig tree, 18 ad. 9(7d, 119), 39 subad 8(124, 252, 2 sex?), 29 juv. 6, 19 juv. 5, 1 stad.?, 21.11.1982 (Loc. Att-82/30, leg. B. Hauser). - Aegean Islands, Ikaria, near the road Plumarion - Monokampion, alt. 420 m, sample from old stump of Platanus orientalis, Berlese extraction, 1 juv. 6, 24.1V.1980 (Loc. Sam-80/7, leg. B. Hauser). Samos, near Kosmathei, the cave “Kako Perato” near the monastery Kimisos Theotoki, alt. 580 m, | juv. 6, 25.IV.1980 (Loc. Sam-80/9, leg. B. Hauser). Samos, near Kosmathei, in the cave “Tsitse Tripa”, alt. 510 m, 1 ad. 9(9), 25.1V.1980 (Loc. Sam-80/10, leg. B. Hauser). Total number. 157 specimens. General distribution. S. pedunculatus occurs in Europe in many countries from Sweden in the north to Spain and Greece in the south. Outside Europe it is known from Morocco, the Canary Islands, and from the USA, Canada and Australia. NEW RECORDS OF PAUROPODA 337 Subgenus Donzelotauropus Remy, 1957 (Remy, 1957a) 24. Stylopauropus (D.) limitaneus Remy Stylopauropus (D.) limitaneus Remy, 1962: p. 76-78, fig 2. Material examined. SLOVENIA, between Godovic and Crni Vrh, alt. ca 600 m, beech forest, 2 ad. 9(2), 12.1X.1969 (Loc. Ju-69/35, leg. V. Mahnert). Julian Alps, Triglav, on Tominskova trail, near Aljazev dom, alt. 1070 m, beech forest, under stone, 1 ad. 9(2), 26.VIII.1967 (Loc. Tg-67/8, leg. B. Hauser). Total number. 3 specimens. General distribution. S. limitaneus 1s a rare species known from a few localities only: one in Austria, some in Romania, one in Italy and one in the USA. It is here reported from Slovenia for the first time. Genus Rabaudauropus Remy, 1953 25. Rabaudauropus cuspidatus (Remy) ?Pauropus cuspidatus Remy, 1939: p. 12-15, figs 4-5. Material examined. GREECE, Peloponnesus, Achaia, Panachaikon Massif, near the road Kastritsion-Patras, alt. 260 m, under Cupressus sempervirens, soil sample, Berlese extraction, 1 ad. 9(4), 1 juv. 5, 6 juv. 3, 16.11.1982 (Loc. Att-82/5, leg. B. Hauser). - Laconia, Talanta, at the road Neapolis - Monemvasia, alt. 210 m, soil samples, Berlese extraction, 24.11.1982, under Quercus coccifera, 1 juv. 6 (Loc. Att-82/45, leg. B. Hauser) and under St. John’s bread tree, 1 ad. 9(2), 1 juv. 6, 1 juv. 5 (Loc. Att-82/46, loc. B. Hauser). Total number. 12 specimens. General distribution. R. cuspidatus has a Mediterranean range: South France, Italy, Bosnia, Romania, Greece and Morocco. BRACHYPAUROPODIDAE Genus Brachypauropus Latzel, 1884 26. Brachypauropus superbus Hansen Brachypauropus superbus Hansen, 1902: p. 410-412, pl. VI, fig. 3. Material examined. GREECE, Peloponnesus, Achaia, Erymanthos Massif, above Kalusion, alt. 980 m, under Abies cephalonica and Quercus coccifera, soil sample, Berlese extraction, | juv. 6, 1.V.1980 (Loc. Sam-80/18, leg. B. Hauser). Total number. 1 specimen. General distribution. The species has not often been met with, it is known from one locality in Poland, some in France, one in Romania, two in Spain and one in Italy. The locality reported above, the first one from Greece, extends the range considerably in southeast direction. EURYPAUROPODIDAE Subfamily Eurypauropodinae Genus Acopauropus Cook, 1896 27. Acopauropus attemsi Hasenhiitl Fig. 14 Acopauropus attemsi Hasenhiitl, 1987: p. 21-24, figs 1-3 on p. 43-47. 338 U. SCHELLER Material examined. GREECE, Peloponnesus, at the road Sparta - Kalamata, alt. 1150 m, under Pinus, soil sample, Berlese extraction, 3 ad. 9(26,19), 20.V.1976 (Loc. The-76/32, leg. B. Hauser). Total number. 3 specimens. General distribution. A. attemsi is known from Carinthia and Styria in Austria only. The locality reported above, the first one from Greece, extends the range to south- east Europe. Taxonomical remarks. Genital papillae with short pubescence, widest in the middle, inner and outer sides evenly curved, 1.4 times as long as their greatest diameter; length of seta 0.7 of the length of papilla. 28. Acopauropus hastatus (Attems) Eurypauropus hastatus Attems, 1895: p.173-175, figs 15-18. Material examined. TURKEY, Istanbul, between Yalova and Orhangazi, 1 ad. 99), 11.V.1976 (Loc. 1, leg. C. Besuchet & I. Löb]). SWITZERLAND, Ticino, Val Mara, alt. 800 m, under old tree stumps, 6 ad. 9(5 49), 4 subad. 8(14,39), 1 juv. 6, 18.1V.1976 (Leg. C. Besuchet). Total number. 12 specimens. General distribution. A. hastatus was previously known from the southern half of Europe only: Switzerland, Austria, Czech Republic, Italy, Slovenia and Romania. The known range of the species is here extended to west Turkey. 29. Acopauropus consobrinus (Remy) Eurypauropus consobrinus Remy, 1937: p. 253-256, figs 1-5. Material examined. FRANCE, Gard, above St. Jean du Gard, hazel, flotation, 1 ad. 9(9), 26.X.1982, and Tornac, olivewood, 2 ad. 9(9), 28.X.1982 (Leg. I.Löbl). Ibidem, Roquedur, flotation, 1 ad. 9(G), 22.X1.1982 (Leg. C. Besuchet). - Pyrénées-Orientales, forest near the cave Pouade, 2 ad. 94,9), 12.1V.1977 (Leg. I. Löbl); ibidem, Parcigoule, 3 ad. 9(1d, 22), 15.1V.1977 (Leg. I. Löbl); ibidem, 16 km W Tech, in ravine near the river Tech, 7 ad. 9(69, 12), 15.IV.1977 (Leg. I. Lobl). SWITZERLAND, Ticino, Rancate, at foot of chestnut stump, 2 ad. 9(9), 27.V.1982; Caprino, compost, 1 ad. 9(2), 29.V.1982; Valle della Crotta, alt. 700 m, in litter, 1 ad. 9(2), 21.V.1982 (Leg. C. Besuchet). Total number. 20 specimens. General distribution. This species has often been collected in South France but is rare Outside. One locality each is known from Austria, Spain and Algeria. A. conso- brinus is here reported for the first time from Switzerland. 30. Acopauropus hispanicus (Scheller) Gravieripus hispanicus Scheller, 1974b: p. 626-630, figs 6-7. Material examined. SPAIN, Gerona Prov., Col de Coubet, alt. 960 m, 22 ad. 9139, 9@), 19. VII.1975 (Leg. I. LODI). Total number. 22 specimens. General distribution. The species was previously known only from the locality of the male holotype in Spain (Gerona Prov., near Olot, Uria). 31. Acopauropus tetrastichus Scheller Acopauropus tetrastichus Scheller, 1981b: p. 167, 169-172, figs 4-5. NEW RECORDS OF PAUROPODA 339 Material examined. TURKEY, above Zonguldak, alt. 500 m, 3 ad. 9(16, 29), 23.V.1976 (Loc. 35, leg. C. Besuchet & I. Löbl). Total number. 3 specimens. General distribution. A. tetrastichus was known only from one specimen from the type locality in the Georgian Republic in the Caucasus Mountains (Scheller, 1981b). Remarks. The specimens studied here correspond in general very well with the type specimen, but the antennal globulus g is proportionately longer and the g’ shorter, both the antennal branches are more slender and the distal part of the genital papillae are more distinctly cylindrical. The species is here reported for the first time from Turkey. Genus Trachypauropus Tömösväry, 1882 32. Trachypauropus cordatus (Scheller) Figs 15-18 Gravieripus cordatus Scheller, 1974b: p. 617-622, figs 1-2. Material examined. ITALY, near Siena, Montalbuccio, 3 ad. 9(@), 1 subad. 8(£), 1 juv. 6, X1.1974 (Leg. R. Dallai). GREECE, Elidia, S of Pirgos, forest near the Alfios river, alt. 30 m, soil samples, Berlese extraction, under Pinus sp., 13 ad. 9(60, 69, 1 sex?), 8 subad. 8 (43, 49), 1 juv. 6, 3 juv. 5, 3 juv. 3 (Loc. Att-82/32, leg. B. Hauser), and under Quercus pubescens, 1 subad. 8(¢), 1 juv. 6, 22.11.1982 (Loc. Att-82/33, leg. B. Hauser). - Peloponnesus, Messenia, near the road Areopolis - Kalamata, before Pygi, near a small stream, alt. 230 m, under Quercus coccifera, soil sample, Berlese extraction, 3 ad. 9(9), 5 juv. 6, 1 juv. 5, 18.V.1981 (Loc. Art-81/15, leg. B. Hauser). — Aegean Islands, Ikaria, near the road Plumarion - Monokampion, alt. 420 m, Berlese extraction, sifting of high water level deposits near small river, 1 juv. 5, 24.1V.1980 (Loc. Sam-80/5b, leg. B. Hauser) and sample from old stump of Platanus orientalis, 1 subad. 8( 2 ), 24.1V.1980 (Loc. Sam-80/7, leg. B. Hauser). Samos, near Kosmathei, close to the entrance of the cave “Tsitse Tripa”, alt. 510 m, old pine stump, Berlese extraction, 1 ad. 9(4), 25.1V.1980 (Loc. Sam-80/11, leg. B. Hauser). — South Island Arc, Crete, at the road Sitia - Iraklion, near Sfaka, small ravine, alt. 200 m, under Quercus coccifera, soil sample, Berlese extraction, 2 ad. 9(2), 13.11.1979 (Loc. Kar-79/12b, leg. B. Hauser). TURKEY, 7 km E Yasilkent, Antakya, alt. 350-400 m, in litter, 1 ad. 9(2), 4.V.1978 (Loc. 22, leg. C. Besuchet & I. Löbl). SWITZERLAND, Geneva, Malval, in moss, 1 ad. 9(2), 8.VII.1983, and Chancy, under old log, 1 ad. 9(2), 13.VII.1983 (Leg. C. Besuchet). Total number. 48 specimens. General distribution. T. cordatus was known earlier from Spain, Italy and Greece, but the localities reported above from Switzerland and Turkey, the first ones from these countries, indicate a wider range both in the north and south-east. Taxonomical remarks. No males were available when 7. cordatus was described (Scheller, 1974b, 1977a) but the discovery of some, both adults and subadults, in the material from Greece makes it possible to describe the genital papillae of both stages. In adults they are glabrous, distinctly conical, 1.4-1.5 times as long as their greatest diameter, with the seta 0.8-0.9 of the length of papilla; in subadults they are also glabrous and without seta, either very small or larger, about half of the length of the papillae in the adults. 340 U. SCHELLER < 15 | 6 di 18 7 VA Er —_____ i 14 Fics 14-18 Genital papillae. 14, Acopauropus attemsi, ad. 9, 15-18, Trachypauropus cordatus, 15-16 ad. 9 and 17-18 subad. 8. Pubescence only partly drawn in Fig. 14. Scale a: Figs 14-16; b: Figs 17-18. 33. Trachypauropus glomerioides Tömösväry Trachypauropus glomerioides Tömösväry, 1882: p. 362-363, figs 4-8. Material examined. TURKEY, above Zonguldak, alt. 500 m, 16 ad. 9(5d, 119), 23.V.1976 (Loc. 35, leg. C. Besuchet & I. Löbl). SWITZERLAND, Geneva, Bellerive, compost, 2 ad. 9(9), 22.VI.1980, and Geneva, Frontenex, at foot of oak, 2 ad. % 2), 11.VI.1981 (Leg. C.Besuchet). Total number. 20 specimens. General distribution. T. glomerioides seems to have a Central European — East Mediterranean distribution from eastern France and Italy in the west to Romania and Turkey in the east. The species is here reported for the first time from Turkey. 34. Trachypauropus latzeli (Cook) Eurypauropus latzeli Cook, 1896: p. 32. Material examined. FRANCE, Drôme, Forêt de Saou, among dead leaves, 1 ad. 9(2), 29.IX.1981 (Leg. C. Besuchet). - Haute-Savoie, Grand Saléve, in moss, alt. 1350 m, 1 ad. 99), 19.X.1980 (Leg. C. Besuchet). SWITZERLAND, Basle, Diegten, in dead leaves, 3 ad. 9(9), 28.IX.1975 and 3 ad. 9(2d, 19), 26.IX.1976 (Leg. C. Besuchet). - Vaud, Onnens, old stump, 1 subad. 8(2), 16.VI.1976 (Leg. C. Besuchet). - Geneva, Chancy, at the foot of old stump, 1 ad. AG), 25.11.1976 (Leg. C. Besuchet); ibidem, Vers-Vaux, at the foot of old stump, 3 ad. 9(16, 29), 16.X.1976 (Leg. C. Besuchet). Total number. 13 specimens. General distribution. The species has not so often been collected but it seems to be widely distributed in the southern half of Europe. NEW RECORDS OF PAUROPODA 341 Genus Samarangopus Verhoeff, 1934 35. Samarangopus molestus Remy Figs 19-25 Samarangopus molestus Remy, 1959: p. 186-189, fig. 14. Material examined. MAURITIUS, Curepipe, Trou-aux-Cerfs, alt. 700 m, 1 ad. 9(9), 20.1.1975 (Loc. Mau-75/26, leg. P. Schauenberg). Total number. | specimen. Remarks. The species was previously known from three sites on Mauritius (Remy, 1959). The specimen reported above agrees well with the description of the type. However, some important details were not studied by Remy and the description is emended in the following respects. COMPLEMENTARY DESCRIPTION Antennae. Neither the antennal setae nor the branch r was drawn by Remy. The main part of the antenna is shown in fig. 20. Base segments of flagella with short oblique pubescence. Trunk. Tergites I-V each with several low elevations (Figs 19, 21) having the largest type of cuticular protuberances. Bothriotricha. All bothriotricha but 7} with very thin axes, glabrous most proximally and outwards covered with erect minute pubescence. T, with distinct pubescence (Fig. 22). Legs (Figs 23, 24). Setae on coxa and trochanter of all legs furcate. The one on coxa of leg 9 with rudimentary, glabrous, pointed secondary branch, the one on trochanter with subequal branches. More anteriorly these setae are similar to those of the coxa of leg 9. Tarsi distinctly tapering, on leg 1 and 9 2.0 times longer than their greatest diameter. Tarsi of leg 1 with one straight and thin seta, almost 0.3 of the length of tarsus. Tarsi of leg 9 with two tergal setae, both strongly tapering, the distal one thickest and distinctly curved; length of proximal seta 1/3 of the length of tarsus and 1.6 times as long as distal seta. Length of the main claw 0.6 of the length of tarsus in leg | and 0.5 in leg 9. Pygidium (Fig. 25). Tergum. Posterior margin with shallow indentation between st and short digitiform appendages behind a;, about as long as a,. Setae a,, a; and st tapering and pointed, a, and st also curved inwards, a; also diverging; a, straight, cylin- drical, blunt and converging, all but st shortly pubescent. Jndex of tergal setae a, = st =5, a, = 3, a, = 8, a, — a, = 10, a, - a, = 22, az — a, = 32, st — st = 9 um; st — st/a, — a, = 0.9, ala, - a, = 0.5, a,/a, - a, = 0.8, a, - ala, — az = 1.0; cuticle between ap, a; and st minutely granular. Sternum. Posterior margin between b, with shallow indentation; setae thin, 5, and b, tapering, pointed, minutely pubescent, 5; cylindrical. Index of sternal setae: b, = 25, b, = 20, b, = 10, b, — b, = 30, b, — b, = 48, b3— bz = 19; b,/b, -b, = 0.8, b,/b,- b, = 0.9; b;/b; — b; = 0.5. Anal plate 1.1 times as broad as long, anterior and posterior ends of the same breadth; lateral margins with a pair of diverging, blunt, cylindrical branches, ~ 0.7 of the length of plate and curved inwards; posterior half of the plate divided longi- 342 U. SCHELLER tudinally by a narrow incision into two somewhat narrowing appendages, which are cut squarely at distal end; their ends with thornlike prolongation of inner margin and large bladders covered with a short but dense pubescence; bladders 3 times longer than wide, 0.8 of the length of plate. 36. Samarangopus rwandaensis sp. n. Figs 24-37 Type material. Holotype: ad. 9(3), RWANDA, Rangiro, alt. 1800 m, sifting in forest, 26. VIII.1976 (Loc. Rwa-73/7, leg. P. Werner). Paratype: 1 ad. 9(¢), same locality as holotype, 10.VII.1973 (Loc. Rwa-73/7, leg. P. Werner). Total number. 2 specimens. Diagnosis. Samarangopus is probably rich in species because new finds almost always have resulted in descriptions of new species. Thus, as several new species re- main to be discovered, every statement on the relationships of S. rwandaensis will be premature. However the structure of the tergites indicates that two New Caledonian species, S. umbraculus Scheller (Scheller, 1993) and S. browni Remy (Remy, 1957b), may be most closely related. Etymology. A latinized adjective of the name Rwanda. DESCRIPTION Length. (1.05)1.10 mm. Head. No vertex setae. Anterior part of temporal organs turned up without con- tact with the head cuticle. Antennae. Chaetotaxy of segments 1-4: 2/2/2/3; no g’; setae thin, tapering, pointed, striate on segment 4, p = 30, p’ = 24(25), p” = (27)29 um; neither p’” nor u and r. Sternal branch subcylindrical but with distinct anterior truncation, anterior margin = (30)32, posterior margin = 45(50), © of base = 12, maximum © = (13)14 um, q thin tapering pointed striate, | = (14)16 um. Posterior margin/length of g (2.6)2.9, posterior margin/maximum © = 3.2(3.8), maximum @/@ of base = (1.1)1.2. Tergal branch subcylindrical, 1 = 40(50), O of base = 8(9), maximum @ = 9.5(12) um; pore not discernible; length of /maximum @ = 4.2. Globulus g, | = 17, maximum © = 10 um; length/maximum © = 1.7; number of bracts 12(13), their length = 7, capsule with flattened bottom, 1 = 3(-4), O = (5)6 um. Relative lengths of flagella (with base seg- ments included): F, = 100, F, = 42, F; = (84)86. Lengths of base segments, bs, = (13)18, bs, = (12)13, bs; = (23)27. F, (2.8) times as long as t, F, and F 1.4 and (2.7)2.8 times as long as s respectively. Antennae glabrous but the helmet-shaped calyces have all a very short pubescence. Trunk. Collum segment hidden. Tergites with two main types of protuberances: 1. rod-shaped with transparent cup protruding in a single row from bulges on anterior and lateral margins of tergite I and from lateral margins of tergites II-VI. 2. round cushion-like structures (in tergal view) with a small opening on the top and a pattern inside similar to a cart wheel with several spokes, which are thickest near a central cavity; cushion-like structures evenly distributed over the tergites, except most anteriorly on tergites II-VI. Posterior margins of tergites I-V lined with a row of such cushion-like structures. On all tergites there is a complicated pattern of other cuticular structures: cavities with canals forming a radiating pattern at the bottom, canals NEW RECORDS OF PAUROPODA 343 Fics 19-25 Samarangopus molestus Remy. 19, body with low elevations, tergal view; 20, left antenna, sternal view; 21, tergite I, median and right part; 22, tergite IV, lateral side with 73, sternal view; 23, leg 9; 24, leg 1; 25, pygidium and left posterolateral part of tergite VI, sternal view. Pubescence only partly drawn in Fig. 25. Scale a: Fig. 21; b: Figs 22-24; c: Figs 20,25. 344 U. SCHELLER connecting the cushion-like protuberances, and bare patches, the latter being few and indistinct. Bothriotricha. All bothriotricha but 7, with very thin axes, glabrous except for their distal parts which have a minute pubescence. 7} with thicker axes, distal third increasing in width distally forming a clavate swelling with very short, almost erect pubescence. Genital papillae. Glabrous, subcylindrical, with almost straight inner margin, (1.7)1.8 times as long as greatest diameter; seta 0.4 of length of papilla. Legs. All legs 5-segmented. Setae on coxa and trochanter of leg 9 furcate, striate, bases pubescent; length of secondary branch 0.5 of primary one; more anteriorly setae with rudimentary pointed glabrous secondary branches. Tarsi tapering; those of leg 9 (2.6)2.8 times as long as their greatest diameter, with two tergal setae, both pointed and glabrous; length of proximal one 20, distal one 15 um. Proximal setae 0.3(0.4) of the length of tarsus and 1.6 times as long as distal seta. Cuticle of tarsus with short pubescence. No proximal seta on tarsi of leg 1. All legs with large main claw and small setose anterior secondary claw; length of main claw 0.5 of the length of tarsus in both leg 1 and 9. Disciform appendage of leg 1 not available for study. Pygidium. Tergum. Posterior margin rounded but with tergal indentation and distinctly pubescent sternal lobe between a,, lobe with small, posteromedian subtrian- gular projection. Setae a,, a, and a; diverging, with oblique pubescence, a, and a, clavate, the latter also curved inwards, a; cylindrical, blunt, diverging, st rudimentary, diverging, glabrous. Index of setae: a, = 10, a, = 8(9), a; = (17)18, st = 1 pm. Distance a, — a, = (11)12, a, - a, = 27(28), a; — a; = 45(47), a, - a, = (9)10, a, — a; = 1112), st — st = 17 um. Distance a, — a, (1.1)1.2 times as long as a,, distance a, — a, (0.8)0.9 of distance a, — 43; distance st - st 17 times as long as st and 1.4 times as long as distance a, — ay. Cuticle glabrous, except in front of a, and a). Sternum. Posterior margin with broad indentation between b,. Setae with distinct oblique pubescence; b, tapering, blunt, with very dense pubescence, b, fusiform, 6.2 times as long as its greatest diameter, pointed, b; tapering, pointed. Index of setae: b, = (40)45, b, = 25(26), bz = 14 um. Distance b, — b, = 43(44), b, — b, = 56(62), b3 — b3 = (29)32, b, — b, = 43, by — b; = 23(26) um. Distance b, — bj 1.0(1.1) of the length of b,, b, 0.6 of distance b, — b>, b 0.4 of distance b; — b;. Anal plate broadest anteriorly, (1.4)1.5 times as long as broad, lateral margins almost straight and with tapering diverging appendages, 0.6 of the length of plate, distal part of plate cleft by a V-shaped incision into two tapering branches; each branch truncate and provided with two appendages: an inner pointed, glabrous one, 0.4 of the length of the plate, and an outer bladder-shaped one with (0.6)0.8 of the length of plate. Plate glabrous, lateral appendages with strong oblique pubescence, bladder-shaped appendages with dense short pubescence. Remarks. This is the first time the genus Samarangopus can be reported from continental Africa. It was earlier known from Madagascar and eastwards from the Oriental Region, as well as from two areas in Notogaea, New Caledonia (Scheller, 1993) and Papua New Guinea (Scheller, 1996). ee NEW RECORDS OF PAUROPODA 345 Fics 26-30 Samarangopus rwandaensis sp. n., holotype ad. 9(d ). 26, left antenna, sternal view; 27, tergite II, bare patch surrounded by cavities in the cuticula and by cushion-like structures; canals between cushion-like structures drawn as dashed lines at the lower part of drawing; 28, tergite IV, anterior part; 29; tergite IV, posteromedian part; 30, tergite VI, lateral part with insertion pit of T;, tergal view. Pubescence only partly drawn in Fig. 26. Scale a: Figs 27-30; b: Fig. 26. 346 U. SCHELLER Fics 31-39 Samarangopus rwandaensis sp. n., holotype ad. 9(4). 31, seta on coxa of leg 9; 32, seta on trochanter of leg 9; 33, seta on coxa of leg 8; 34, seta on trochanter of leg 8; 35, tarsus of leg 9; 36, tarsus of leg 1; 37, T3; 38, right genital papilla and seta on coxa of leg 2, anterior view; 39, pygidium, sternal view. Pubescence only partly drawn in Fig. 39. Scale a: Figs 35, 36, 38; b: Figs 313493750: Fig! 39. nn ner rte NEW RECORDS OF PAUROPODA 347 Subfamily Sphaeropauropodinae Genus Sphaeropauropus Silvestri, 1930 37. Sphaeropauropus nepalensis Scheller Sphaeropauropus nepalensis Scheller, 2000: p. 118-121, figs 71-86. Material examined. SRI LANKA, Central Province, Hakgala, wooded ravine in the north- east, alt. 1700-1800 m, sifting, 1 ad. 9( 2), 28.1.1970 (Loc. 30a, leg. C. Besuchet & I. Löbl). Total number. 1 specimen. General distribution. S. nepalensis is new to Sri Lanka. It was described from Nepal (Scheller, 2000) and is here reported for the first time outside the Himalaya. Taxonomical remarks. The styli on the pygidial tergum of the specimen exam- ined are thinner than in the material from Nepal. 38. Sphaeropauropus reunionensis sp. n. Figs 40-51 Type material. Holotype: ad. 9(9), REUNION, St. Philippe, 14-15.1.1975 (Loc. Mau- 75/60, leg. P.Schauenberg). Paratype: 1 subad. 8(2), same locality and date as holotype. Other material. SRI LANKA, Central Province, Matale, wooded ravine, alt. 400 m, sifting, 1 ad. 9(2), 17.1.1970 (Loc. 7, leg. C. Besuchet & I. Löbl); Kandy, Udawattekele Sanctuary, primary forest, alt. 600 m, sifting, 1 ad. 94), 19.1.1970 (Loc. 11, leg. C. Besuchet & I. Löbl). - Uva Province, about 10 km N Monaragala, forest, sifting, 1 ad. 9(¢), 13.11.1970 (Loc. 64, leg. C. Besuchet & I. Lôbl). Total number. 5 specimens. Diagnosis. The species described here is well delimited from S. malayus Silvestri. Good distinguishing characters can be found in the shape of the antennal globulus g, the T;, the claws and the tarsi and in the cuticular structures of the tergites. Etymology. A latinized adjective of the name Réunion. DESCRIPTION Length. 0.92 mm. Head. Setae hidden. Vertex: ve, rudimentary. Temporal organ, | = 60 um. Antennae. Cuticle of segments 1-4 almost glabrous, branches and flagellae glabrous, calyces small and glabrous. Chaetotaxy of segments 1-4: 2/2/2+g’/3. Setae cylindrical, annulate, g’ small subhemispherical, segment 4, p = 42, p = 35, p” = 15 - pm. Sternal branch with distinct anterodistal truncation, anterior margin = 24, posterior margin = 32, © of base = 9, maximum © = 12.5, upper © = 10, g = 20 pm. Anterior margin/length of g = 1.5; anterior margin/length of g = 1.2; anterior margin/posterior margin = 0.6; anterior margin/maximum © = 2.0; maximum @/@ of base = 1.4. Tergal branch r somewhat fusiform, length = 37, @ of base = 5, maximum © = 10 um, length of t/maximum © = 3.7. Globulus g somewhat curved, stalk conical, length of g = 16, maximum © = 9.5 um; = 10 bracts, their length = 7-8 um; capsule somewhat flattened, length = 5, @ = 5.5 um. Relative lengths of flagella (with base segments included): F, = 100, F, = ?, F; = 102. Lengths of base segments: bs, = 19, bs, = 9, bs; = 17 um. The F, 2.9 times as long as f, F, 3.4 times as long as s. Trunk. Setae of collum segment hidden. Sternite process with narrow and rounded anterior end; appendages with low round caps, © = 25 um. 348 U. SCHELLER Fics 40-46 Sphaeropauropus reunionensis sp. n., holotype 9(9). 40, body, right half, tergal view; 41, left antenna, tergal view; 42, collum segment, sternal view (setae not drawn); 43, tergite I, central part; 44, tergite II, left posterolateral corner; 45, left part of tergite IV with T3, tergal view; 46, left part of tergite VI with base of 7;, sternal view. Scale a: Fig. 42; b: Figs 41, 46; c: Figs 43-45. Tergal sides of tergites with tuft-like setae, those on anterior part of tergite I broadly clavate with thick stalks. Cuticle between tuft-like setae with short rows of granules and very small protuberances covered with great many inward-curved oblique pubescence hairs. Tuft-like setae less clavate on lateral margins, sometimes with almost cylindrical stalks; laterosternal setae often more elongate, sometimes even NEW RECORDS OF PAUROPODA 349 Fics 47-51 Sphaeropauropus reunionensis sp. n., 47, 49-51, holotype 9(?). 48, ad. 9(¢) from Sri Lanka, Kandy. 47, T,; 48, genital papillae, anterior view; 49, leg 1; 50, leg 9; 51, pygidium, sternal view. Scale a: Figs 47-50; b: Fig. 51. with thin pointed stalks. Short pubescence hairs at bases of setae arranged in two whorls, one of them very close to the base. Pubescence longest on lateral parts of ter- gites. The laterosternal furrows strongly bent inwards. Bothriotricha. All bothriotricha but 7} with simple very thin axes and short pubescence of straight almost erect hairs which are branched most distally. 7} with thicker axes and distal fusiform swelling, the latter 0.3 of the length of bothriotrix; distal part of T, almost glabrous, middle part with dense and short pubescence of oblique hairs. Relative lengths of bothriotricha: T, = 100, T, = 166, T; = 85, T, = 126, NER 350 U. SCHELLER Genital papillae (described from the adult male from Kandy, Sri Lanka). 3 times longer than wide, with straight inner side, lateral side evenly rounded, glabrous, seta 0.2 of the length of papilla; base segment well developed, length almost 0.4 of the length of papilla. Legs. All legs 5-segmented. Setae on coxa and trochanter of leg 9 furcate, their main branch annulate and pointed, secondary branch on coxal seta rudimentary, poin- ted, glabrous and on trochanter annulate and pointed; length of main branch = 25 um. More anteriorly these setae with rudimentary, pointed, glabrous secondary branch. On most anterior legs the seta on trochanter is much thinner than coxal seta. Tarsi of legs 1 - 9 slender, tapering, somewhat bow-shaped, distal half almost cylindrical. Tarsi of leg 1 almost glabrous, 4.3 times as long as greatest diameter, those of leg 9 3.8 times as long as greatest diameter; tarsi of leg 9 with minute pubescence. Tarsus of leg 1 with only one distal tergal seta, about 0.1 of the length of tarsus; tarsus of leg 9 with two tergal setae, both tapering, pointed, glabrous, proximal one = 19, distal one = 7 um, proximal seta almost 0.3 of the length of tarsus and 2.4 times as long as distal seta. A triangular pointed appendage on anterior side of femur of leg 1, length = 5 pm. All legs with almost straight thin main claw and small setose anterior secondary claw. On leg 1 main claw = 16, secondary claw = 6 um; on leg 9 main claw = 17, secondary claw = 11 um; main claw not fully 0.3 of the length of tarsus in all legs. Pygidium. Tergum. Posterior margin almost straight, serrate with 6 teeth/lobe. Setae subequal, tuft-like, but with shorter hairs than on the tergites. a, curved outwards, somewhat converging, a, and a; somewhat curved inwards; st cylindrical, blunt, some- what S-shaped, almost glabrous. Index of tergal setae: a, = a, = 7, a, = 6.5; a; — a, = 15, a, — a, = 30, a; — a, = 41, st = 12, st — st = 13 um; st - st/a, — a, = 0.9, a,/a, — a, = 0.5, a,/a, — a, = 1.2, a, — a,/a, — a, = 2.9. Tergum glabrous. Sternum. Posterior margin between 5, straight; setae blunt, b, and b, subcylin- drical, somewhat tapering, the former densely pubescent and the latter almost glabrous; b; cylindrical and with distinct pubescence. Index of sternal setae: b, = 42, b, = 13, b; = 12, b, — b; = 37, b,-— b, = 50, b, — b, = 25, b, — b, = 9, b,/b, — by = 1.1, b/b,= 3.2; b,/b, — by = 1.4, b3/b; — b3= 0.5. Anal plate circular, with two short subcylindrical branches protruding backward from posterior margin. Branches with oblique posterolateral truncation on sternal side, each carrying one distal ovoid bladder-shaped appendage with distinct thin cylindrical stalk; these appendages twice longer than wide, 0.3 of the length of plate and carrying a distinct, dense pubescence of erect hairs. Circular part of plate almost glabrous, cylin- drical appendages with dense, short pubescence. Remarks. Two Sphaeropauropus specimens have been reported previously from Réunion (Remy, 1957a). They were assigned to the only species known at that time, S. malayus Silvestri (Silvestri, 1930). A comparison of Remy’s description of his material with the results of my later redescription of the species (Scheller, 1998) indicates that two species are involved. There are distinct differences in the shape of the antennal globuli g, the claws and genital papillae and in the proportion b,/b, -b | and b,/b, — bo. Because the similarities between S. reunionensis and the other species of the genus are even weaker, the relationships of the species are not yet possible to trace. NEW RECORDS OF PAUROPODA 351 GENERAL REMARKS ON DISTRIBUTION Our knowledge of the taxonomy and distribution of the pauropod species is still very poor and therefore it is not surprising that about half of 38 species reported above are either new to science or have been collected outside their previously known range, sometimes far outside. So e.g., three species are here reported new to Switzerland and Greece, two countries, which are rather well investigated thanks to earlier collecting of zoologists from both Geneva (Scheller, 1976, 1977a, b, 1981a) and France (Scheller, 1973). Including the species reported here, we know now 29 species from Switzerland and 34 from Greece. An earlier review of the occurrence of the genus Samarangopus (Scheller, 1996) shows a range from Madagascar in the west to New Caledonia in the east and from New South Wales in Australia in the south to Thailand in the north. However, the range seems to be much wider. It now includes Nepal (Scheller, 2000) and the present study shows that the genus occurs in continental Africa too. Another genus, Sphaeropauropus, might have a similar range. It was long thought to be restricted to the Australasian islands of Java, the Philippines and Réunion, but it has now been discovered far outside. There are further reports from China (Zang & Chen, 1988), Thailand (Scheller, 1995) and Nepal (Scheller, 2000) and here the genus is reported from Sri Lanka too. REFERENCES ATTEMS, C. 1895. Die Myriapoden Steiermarks. Sitzungsberichte der kaiserlichen Akademie der Wissenschaften in Wien. Mathematisch-naturwissenschaftliche Classe (1)104: 117-238. Cook, O. F. 1896. An enumeration of the Pauropoda. Brandtia 6: 29-32. HANSEN, H. J. 1902. On the genera and species of the order Pauropoda. Videnskabelige Meddelelser fra Dansk Naturhistoriske Forening i Kjobenhavn 1901: 323-424, pls I-VI. HASENHUTL, K. 1987. Neue Zwergtausendfüßer aus Kärnten (Myriapoda, Pauropoda). Carinthia II, 45. Sonderheft: 17-75. LATZEL, R. 1884. Die Myriopoden der östereichisch-ungarischen Monarchie. Alfred Hölder, Wien II: 414 pp., XVI pls. LUBBOCK, J. 1867. On Pauropus, a new type of Centipede. Transactions of the Linnean Society of London 26: 181-190, pl. 10. Remy, P. A. 1931. 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A preliminary study on a new record of the class Pauropoda (Myriapoda) from China. Sinozoologia 6: 39-54. AM rare à REVUE SUISSE DE ZOOLOGIE 110 (2): 355-379; juin 2003 New molecular and morphological data on the “Euscorpius carpathicus” species complex (Scorpiones: Euscorpiidae) from Italy, Malta, and Greece justify the elevation of E. c. sicanus (C. L. Koch, 1837) to the species level Victor FET!, Benjamin GANTENBEIN?*, Michael E. SOLEGLAD3, Valerio VIGNOLT, Nicola SALOMONE,, Elizabeth V. FET! & Patrick J. SCHEMBRIS ! Department of Biological Sciences, Marshall University, Huntington, West Virginia 25755-2510, USA. E-mail: fet@marshall.edu 2 Institute of Cell, Animal and Population Biology, Ashworth Laboratories, The Uni- versity of Edinburgh, Edinburgh EH9 3JT, Scotland. E-mail: B.Gantenbein@ed.ac.uk 3 P.O. Box 250, Borrego Springs, California 92004, USA. E-mail: soleglad@znet.com 4 Department of Evolutionary Biology, University of Siena, I- 53100 Siena, Italy. E-mail: salomone@unisi.it 5 Department of Biology, University of Malta, Msida, MSD 06, Malta. E-mail: patrick.j.schembri@um.edu.mt New molecular and morphological data on the “Euscorpius carpathi- cus” species complex (Scorpiones: Euscorpiidae) from Italy, Malta, and Greece justify the elevation of E. c. sicanus (C. L. Koch, 1837) to the species level. - The first mitochondrial DNA phylogeny (based on 17 unique haplotypes) is presented for a number of scorpion populations from Italy, Malta, and Greece, previously classified under the “catch-all”? name Euscorpius carpathicus (Linnaeus, 1767). A comparative analysis of the mitochondrial gene for 16S (large subunit) ribosomal RNA suggests that at least two clearly separated lineages are present. However, neither of these belongs to E. carpathicus (L.) in a strict sense, which was limited to Romania in a recent morphological study. The first, “western” lineage, found in northern and central Italy (also present in southern France, Slovenia, Croatia, and Austria) corresponds to E. tergestinus (C. L. Koch, 1837) as recently defined by Fet & Soleglad. Another monophyletic, “south- ern” lineage is elevated here to the species rank as E. sicanus (C. L. Koch, 1837). Originally described from Sicily, E. sicanus includes as new syn- onyms E. carpathicus canestrinii (Fanzago, 1872) and six subspecies de- scribed by Caporiacco: E. c. calabriae, E. c. ilvanus, E. c. garganicus, E. c. argentarii, E. c. palmarolae, and E. c. linosae. Morphology confirms the * Corresponding author Manuscript accepted 08.11.2002 356 V. FET ET AL. existence of two lineages: E. sicanus is characterized by a unique tricho- bothrial pattern and number where series eb, and in some populations also series eb,, have 5 trichobothria (all E. tergestinus possess only four of them). E. sicanus is found in southern Italy (including Sicily and Sardinia), northern Africa, Malta, and Greece. The enigmatic “E. mesotrichus Hadzi” from Greece also belongs to E. sicanus. Key-words: Scorpions - trichobothria - 16S mtDNA - biogeography - phylogeny - Southern Mediterranean - Italy - Malta - Greece. INTRODUCTION Scorpions of the genus Euscorpius Thorell, 1876 (Scorpiones: Euscorpiidae) are very common in southern Europe. they are ecologically diverse and they occupy a variety of habitats from xeric to mesic, from the Mediterranean shoreline to the high altitudes of the Alps and of the Balkan. Linnaeus (1767: 1038) described Scorpio carpathicus from the Transylvanian Alps (Romania) (now Euscorpius carpathicus, the type species of the genus). Thorell (1876) introduced the new generic name Euscorpius. Numerous species have been described in Euscorpius; the traditional taxonomy of this genus, based mainly on morphosculpture and coloration characters, is complicated and confusing (Fet & Sissom, 2000). C. L. Koch (1850: 86-87) reviewed all the Euscorpius species in his series “Die Arachniden”, and clearly divided these species into three groups according to the number of trichobothria (“Grübchen”) on the ventral aspect of the pedipalp patella (called tibia by many earlier authors). The large and conspicuous trichobothria (sensory bristles) of Euscorpius were then also used as taxonomic characters by HadZi (1929, 1930, 1931) and by Caporiacco (1950) who utilised total trichobothrial counts on dif- ferent aspects of the pedipalp chela and patella. In the 50 years since Caporiacco’s re- vision (which was mostly confined to the Italian forms, and only touched upon mate- rial from other countries), many authors paid attention to the intricate and intriguing taxonomy of this most common European scorpion genus (Bonacina, 1980; Curkié, 1972; Fet, 1986, 1993, 1997a, 1997b; Fet & Braunwalder, 2000; Kinzelbach, 1975; Lacroix, 1991a, 1991b; Scherabon, 1987; Vachon, 1963, 1975, 1978, 1981; Vachon & Jaques, 1977; Valle, 1975). For almost a century, only four species were listed in Euscorpius: E. carpathi- cus, E. germanus, E. flavicaudis and E. italicus (see e.g., Birula, 1900, 1917; Capo- riacco, 1950; Kraepelin, 1899; Vachon, 1963, 1981; Vachon & Jaques, 1977). Bonacina (1980) reestablished E. mingrelicus (Kessler, 1874). Further revisional work resulted in 12 formally recognized species: E. alpha Caporiacco, 1950, E. balearicus Caporiacco, 1950, E. beroni Fet, 2000, E. carpathicus (Linnaeus, 1767), E. gamma Caporiacco, 1950, E. germanus (C. L. Koch, 1837), E. hadzii Caporiacco, 1950, E. flavicaudis (DeGeer, 1778), E. italicus (Herbst, 1800), E. koschewnikowi Birula, 1900, E. min- grelicus (Kessler, 1874), and E. tergestinus (C.L. Koch, 1837) (Fet, 2000; Fet & Sissom, 2000; Gantenbein er al., 2000a, 2001; Scherabon er al., 2000; Fet & Soleglad, 2002). EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 337, One of the most recent developments in Euscorpius taxonomy was the intro- duction of molecular techniques which started with the pioneering paper of Gantenbein et al. (1999) on application of allozyme and 16S ribosomal RNA gene sequence ana- lyses to assess the phylogeny of this genus. This study provided the first ever published DNA-based phylogeny for scorpions. These data revealed a phylogenetic relationship between four species (E. flavicaudis, E. carpathicus, E. italicus, and E. germanus) which differed from the former, morphology-based views on evolution of this genus (Hadzi, 1931; Kinzelbach, 1975). Further work included more detailed genetic and morphological analysis of E. germanus, E, alpha, and E. gamma (Gantenbein et al., 2000a; Scherabon er al., 2000) as well as E. carpathicus (Gantenbein et al., 2001; Huber et al., 2001). Gantenbein er al. (2001) elevated E. c. balearicus, an isolated E. carpathicus population from the Balearic Islands from subspecific to species status. Most recently, Fet & Soleglad (2002) conducted a morphological analysis of the “E. carpathicus” complex by including a number of new characters (foremost of which are individually mapped patellar trichobothria). They restricted the species E. carpathicus to popul- ations from Romania, elevated one Balkan subspecies to species status (E. hadzii), confirmed species status for E. koschewnikowi from Greece, and assigned a number of Italian subspecies to the species E. tergestinus. However, the five species established by Gantenbein er al. (2001) and Fet & Soleglad (2002) did not yet embrace all diver- sity of populations traditionally placed under E. carpathicus. The goal of this paper is to continue the study of the polymorphic and poorly studied “E. carpathicus” complex in Italy, with additional comparative material now available from Malta and Greece. MATERIAL AND METHODS MATERIAL For this study, we used DNA sequences from 18 specimens belonging to the “E. carpathicus” complex, represented by 14 unique haplotypes. Two additional sequences, belonging to E. tergestinus, have been taken from the GenBank database (Gantenbein et al. 1999, 2001; Huber er al., 2001): EcLM1, AJ389377; EcPR2= EcPR1, AJ309211. GenBank also provided the outgroup sequence of E. italicus, EiTO1 = AJ298067. All 14 new sequences have been deposited in GenBank under accession numbers AY090076 to AY090089 (see below). Specimens used for DNA analysis: E. sicanus (=E. c. ilvanus Caporiacco, 1950; = E. c. argentarii Caporiacco, 1950; =E. c. canestrinii (Fanzago, 1872); =E. c. gar- ganicus Caporiacco, 1950): ECNEI (AY090078), EcNE2 (AY090076), Italy, Nebrodi Mts., beech forest (1500 m), northeastern Sicily, Italy, 30 September 1999 (V. Vignoli); EcTRI (AY090084), San Domino Island, Tremiti Islands (Adriatic Sea), Foggia, Apulia, Italy, 30 August 2001 (V. Vignoli); EcGA1 (AY090085), Vico del Gargano, Foggia, Apulia, Italy, 29 August 2001 (V. Vignoli); EcPUl (AY090077), Punta Ala, Tuscany, Italy, 10 August 2000 (M. Stiegler & P. Weiner) (identical to haplotypes from Follonica and Siena, Tuscany, Italy); EcAR1 (AY090079), Le Piane, Mt. Argentario, Grosseto, Tuscany, Italy, 22 January 2000 (V. Vignoli) (identical to another haplotype 358 V. FET ET AL. from Siena and a haplotype from Orbetello); EcGG1 (AY090081), Campese, Giglio Island, Tuscany, Italy, 30 May 2000 (V. Vignoli) EcGIl (AY090082), Cala Maestra, Giannutri Island, Tuscany, Italy, 30 May 2000 (V. Vignoli) (identical to another haplo- type from Giglio Island); EcSD1 (AY090083), Lu Fraili, northeastern Sardinia, Italy, 16 August 2001 (V. Vignoli); ECMT1 (AY090086), EcMT2 (AY090087), Buskett, Malta, 17 September 2001 (P. J. Schembri); EcOL2 (AY090088), Mt. Olympus, Thessaly, Greece, 15 May 2001 (V. Fet); EcKB1 (AY090089), Kalambaka, Thessaly, Greece, 14 May 2001 (V. Fet). E. tergestinus: ECLMI (AJ389377), La Morra, Piedmont, Italy, 20 September 1996 (I. & B. Gantenbein); EcPR2 (AJ309211), Procchio, Elba, Italy, 4 October 1996 (I. & B. Gantenbein); EcSZ1 (AY090080), Le Grazie, Porto Venere, La Spezia, Liguria, Italy, June 2001 (D. Facheris). Outgroup E. italicus: EtTO1 (AJ298067), Tortoreto, Abruzzo, Italy (M. Bellini). For our morphological analysis we used additional extensive material deposit- ed in zoological museums (see below, under “Material studied”). METHODS DNA techniques: We used a standard protocol as described in Gantenbein et al. (1999, 2000a). For DNA analyses, genomic DNA was extracted from fresh or pre- served (in ethanol 94-98%) muscle tissue (usually pedipalp or metasoma) by using a standard phenol/chloroform precipitation method (Sambrook er al., 1989) or the Qiagen™ DNeasy extraction kit. Extracted DNA was amplified by the polymerase chain reaction (PCR) in the Perkin Elmer 2400 PCR Thermocycler by using conditions and primers as described in Gantenbein er al. (1999). The mitochondrial LSU (large ribosomal subunit) 16S rRNA PCR primers corresponded to the positions 12,867- 12,884 and 13,328-13,308 in the Drosophila yakuba mitochondrial genome, or to the positions 11,173-11,190 and 11,625-11,606 in the Limulus polyphemus mitochondrial genome (Lavrov et al., 2000). The forward primer is a scorpion-specific version of the “universal” primer 16Sbr, or LR-J-12887, while the reverse primer has a scorpion-spe- cific sequence designed by one of the authors (V.F.). The resulting PCR product was verified on 1% agarose electrophoretic gel and purified by Ultrafree MC 30000 cellu- lose filters (Millipore Inc.) or using “Rapid Gel Extraction Systems” columns (Concert™). Automated Sanger dideoxy sequencing of the double-stranded PCR prod- uct was performed at the Molecular Genetics Instrumentation Facility, University of Georgia (Athens, GA), on the ABI 9600 Sequencer (US team) or at the MWG-Biotech automated DNA sequencing service, Esersberg, Germany (Italian team). Phylogenetic analyses: 17 mtDNA sequences representing different haplotypes were aligned using Clustal X 1.81 (Thompson er al., 1997). DNA sequences used in earlier studies were downloaded from the GenBank nucleotide sequence database. Identical haplotypes were not considered in further analyses. The software package PAUP* Version 4.0b10 (Swofford, 1998) was used for se- quence analysis to perform phenetic (Felsenstein, 1984) and cladistic phylogenetic analyses. We are aware that these methods are based on different assumptions, but all of these are expected to estimate the “true” phylogeny in the absence of long-branch- attraction (Page & Holmes, 1998): genetic distance calculation, Maximum Parsimony (MP), and Neighbour-Joining (NJ) analysis (Saitou & Nei, 1987). However, it appears EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 359 that especially for the choice of “correct” tree topology MP is superior to Maximum Likelihood (ML) in the “inverse Felsenstein zone” (Swofford et al., 2001). For choos- ing the most appropriate DNA substitution model for pairwise distances, we initially estimated a NJ tree using JC69 distance (Jukes & Cantor, 1969). This tree was then used for estimating the ML parameters from 56 different substitution models. Maximum likelihood ratio tests were then performed in a nested design by using the program MODELTEST 3.06 (Posada & Crandall, 1998), which is implemented in PAUP*. Details about likelihood ratio tests are given in Huelsenbeck & Crandall (1997) and Huelsenbeck & Rannala (1997). The rate heterogeneity among sites was assumed to follow a gamma distribution (shape parameter a was ML-estimated) with four cate- gories, each represented by its mean (Yang, 1996). In a further step, the molecular clock hypothesis (i.e., equal rates across all sequences) was tested by using the c? approximated likelihood ratio test statistics. The hierarchical LRT statistics suggested the HKY85 + T model (Hasegawa er al., 1985) for the 16S gene, as it was found also by Gantenbein er al. (2001). The test for the molecular clock hypothesis was rejected at the 5% level (2d = 2(/p-1,) = 840.27 - 823.90 = 32.74, P,?= 0.005). Therefore, the NJ clustering algorithm was chosen, which allows for unequal branch lengths (Page & Holmes, 1998). For weighted MP analysis the tree space was explored by 100 heuris- tic tree searches and by randomizing the order of the sequence input in PAUP*. The transition/transversion (Ti/Tv) ratio was ML-estimated to 2.73; therefore we weighted Tv three times over Ti. The four gaps in the data set were treated as the fifth base (McGuire et al., 2001). Confidence limits of individual nodes were assessed by using non-parametric bootstrapping (Felsenstein, 1985; Swofford er al., 1996). ABBREVIATIONS MZUF, Museo Zoologico “La Specola” dell’ Universita di Firenze, Florence, Italy; NMW, Naturhistorisches Museum Wien, Vienna, Austria; ZMH, Zoologisches Museum Universität Hamburg, Hamburg, Germany; NMM, Naturhistorisches Museum, Mainz, Germany; HNHM, Hungarian Natural History Museum, Budapest, Hungary; NMNHS, National Museum of Natural History, Sofia, Bulgaria; VF, private collection of Victor Fet. Specimen numbers from MZUF are given as AAA/BBBB, where AAA is the “collection number” and BBBB is the “catalog number”. RESULTS The resulting 16S rDNA phylogeny in our study reveals two well-separated monophyletic lineages among the material traditionally placed under E. carpathicus (Fig. 8). One of these lineages, recovered by all analyses, is readily identified as E. tergestinus (C. L. Koch, 1837) as defined by Fet & Soleglad (2002). All of its speci- mens possess a “standard” trichobothrial formula (eb,=4, eb=4). In the molecular phylogeny of Gantenbein er al. (2001, Tab. 1, Figs 2, 3), E. tergestinus was represent- ed (as E. carpathicus) by populations from Mathis (Alpes-Maritimes, France), La Morra (Piedmont, Italy), Procchio (Elba, Tuscany, Italy), Vernazza (Liguria, Italy), Hochosterwitz (Austria), and Mala Duba (Croatia), spanning the entire geographic range of this species as currently accepted by Fet & Soleglad (2002). The same species 360 V. FET ET AL. Fics 1-5 Idealized trichobothrial patterns of the external aspect of the pedipalp patella in two Euscorpius species exhibiting eb series = 4 (with E. sicanus populations exhibiting) eb = 5. 1. Euscorpius balearicus, Balearic Islands, Spain; 2. E. sicanus, Sicily, Italy; 3. E. sicanus, Mt. Argentario, Italy; 4. E. sicanus, Mt. Olympus, Greece; 5. E. sicanus, Sardinia, Italy. Series er = external terminal, est = external subterminal, em = external median, esb = external suprabasal, Eby = external basal-a, and eb = external basal; v = ventral. was characterized (as E. carpathicus) by Huber et al. (2001) who studied more popu- lations from Austria and Slovenia. In our data, a new haplotype of E. tergestinus from La Spezia (Liguria) groups together with previously known haplotypes from Piedmont and Elba. Another 16S rDNA clade, also recovered in our study by all analyses (Fig. 8), includes a diverse array of populations, located mainly in Italy (Tuscany, Sicily, Sardinia, Gargano, and Tremiti Islands) but also in Malta and Greece (Thessaly). This clade is treated below as a separate species. The senior synonym available for it is Scorpius sicanus C. L. Koch, 1837 according to Article 23 of the International Code of Zoological Nomenclature (ICZN, 1999). Separation of this second monophyletic clade (phylogenetic species), distinct in the DNA phylogeny, is also supported by our morphological analysis (Figs 1-5). The most important feature separating E. sicanus from E. tergestinus (and from another re- lated species, e.g., E. balearicus), is its unique trichobothrial pattern and number where the external patellar series eb (and in some populations, also series eb,) has 5 tri- chobothria. All specimens examined of E. tergestinus and E. balearicus possess only 4 trichobothria in these series; Fig. 1. A number of Italian subspecies (E. c. canestrinii, E. c. calabriae, E. c. ilvanus, E. c. garganicus, E. c. argentarii, E. c. palmarolae, and E. c. linosae) (Caporiacco, 1950) also belong to this new lineage, and are therefore formally synonymized here with E. sicanus. The closest relative of E. sicanus is E. hadzii, which also has 5 trichobothria in series eb but is distinguished by 6 to 7 trichobothria in series eb, (Fet & Soleglad, 2002). EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 361 ; = 26 cy = 0.095 E.c.calabriae — "° E.c.sicanus nes" °° E.c.ilvanus hs" ” È = 7% = 0. 000 E.c.palmarolae > 00 11 = 16 cv = 0.062 E.c.argentarii ns E.c.garganicus — u E.c.linosae I” = 2 cv = 0.000 6 i 4 = 152 cy = 0.081 E.c. mesotrichus ———_ o E hadzii an = 213 cy = 0.094 E.c.canestrinii 5 6 7 8 3 10 11 12 13 14 Patella: ventral E.c.linosae er ar n= 7 cy = 0,000 E.c.palmarolae | j n = 22 cv = 0.000 E.c.sicanus | H rn = 46 cy = 0.000 E.c.ilvanus | E.c.argentarii i e Gi CALI) j ı = 26 cv = 0.000 E.c.calabriae I snai ce i = 65 = 0112 E.c.garganicus "nen E.c.canestrinii EE" E.c.’mesotrichus’ E. hadzii Patella: eba E.c.calabriae n= 26 cv = 0.116 E.c.palmarolae n = 7 cv = 0.175 E.c.sicanus ee — |" E.c.ilvanus m & = 46 cv = 0.065 E.c.argentarii men 15 cv = 0.072 E.c.garganicus = 65 cy = 0.064 E.c.linosae I° = 2 cx = 0.000 i = 152 cv = D. 089 E.c. mesotrichus’ — er — E. hadzii = = = 216 cy = 0.094 E.c.canestrinii a = 27 cv = 0.074 2 = iq 8 9 10 Patella: et FIG. 6 Trichobothria statistics of Euscorpius carpathicus subspecies now placed in the synonymy of E. sicanus. E. hadzii Caporiacco, 1950 is used as a reference taxon. Series ef = external terminal, eb, = external basal-a, eb = external basal; cv = coefficient of variation. Figs 2-5 show schematized trichobothrial patterns illustrating the five tricho- bothria of the patellar eb external series diagnostic of Euscorpius sicanus. In contrast E. balearicus exhibits the typical four trichobothria in the same series (Fig. 1). Also of 362 V. FET ET AL. n= 521 cy = 0.119 n = 634 cv = 0.032 Within Error: 9.60% €- 80.61% -> 3.79% Within Error: 2.21% €- 97.48% -> 0.32% 100% BE 6 7 8 3 say RI sles) aa) Az Patella: ventral Patella: eb n = 631 cy = 0.114 n = 358 cy = 0,035 Within Error: 0.00% €- 98.42% > 1.58% Within Error: 0.28% €- 98.04% -> 1.68% 351 4 NSOE 3 5 Patella: eb, Patella: em Within Error:09:427 €189 4740311 180 e a] Patella: et FIG. 7 Complete statistics of patellar trichobothria count in all Euscorpius sicanus specimens exam- ined. Series er = external terminal, em = external median, eb, = external basal-a, eb = external basal; cv = coefficient of variation. interest is that E. sicanus populations from Greece and Sardinia (Figs 4-5) exhibit a tendency for five trichobothria in the eb, series, in contrast to some other populations around Italy, including Sicily and Malta, which usually have eb,=4. Also apparent from Figs 1-5 is the larger number of trichobothria in the er and ventral patellar series for Greek and Sardinian populations. Further statistical data for several subspecies defined by Caporiacco (1950) and herein synonymized with E. sicanus are shown in the histograms in Figs. 6-7. EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 363 Malta EcMT1 \ EcMT1 9 EcMT2 Matta EcNE2 Sicily. Sicil EcNE1 EcNE1 : E. c. garganicus EcTRI ECGAI EcPU1 E c. ilvanus E. sicanus clade EcGG1 EcGl1 EcOL2 E. c. 'mesotrichus' EcKB1 EcGl1 98 E. c. canestrinii EcGGI EcSD1 = E. c. argentarii EcAR1 EcAR1 EcLM1 “ EcLM1 E. tergestinus clade 100 E. tergestinus clade EcSZ1 EcSZ1 EcPR27 \ EcPR2 ben ES E. italicus E. italicus EiTO1 EiTO1 — 0.005 substitutions/site A B FIG. 8 DNA phylogeny of E. tergestinus and E. sicanus (branch labels refer to previously recognized subspecies). A, Neighbour-joining (NJ) tree using Maximum Likelihood (ML) distances as an input matrix. HK Y85 + T-ML distances (Hasegawa et al., 1985) were calculated with the fol- lowing parameters (base frequencies: py = 0.36, pr = 0.41, pc = 0.10, Ti/Tv ratio = 2.73 (k = 7.87), gamma shape parameter a = 0.098, respectively. B, Strict consensus tree of Maximum Parsimony (MP) analysis (55 equally parsimonious trees of 79 steps). Clu and RI were 0.72, respectively. Numbers at nodes are bootstrap values. Outgroup, E. italicus. Euscorpius sicanus (C. L. Koch) can be diagnosed as a small to medium sized species with a somewhat thin metasoma; trichobothrial counts: patella ventral = 6-13 (9-11); external, eb = 5, eb, = 4-5, esb = 2, em = 4, est = 4, et = 5-9 (7); pectinal tooth counts, 8-10 (9) in males; 6+-8??+ (7+) in females. Number of trichobothria in series eb (=5) and eb, = (4-5) are key diagnostic characters for this species. E. sicanus is contrasted here with its closest relative E. hadzii, which also has eb = 5 as follows: eb, = 4-5, as opposed to 6-8 (7) in E. hadzii: em = 4, as opposed to 4-5 in E. hadzii. Position of eb; in E. sicanus is in straight line with eb, - eb, - eb,, eb; usually situated below eb»; in E. hadzii, eb, - eb; - eb, is not in straight line, eb; usually situated above eb,. Number of trichobothria in series eb (=5) and em (= 4) distinguishes E. sicanus from E. carpathicus, and eb (=5) distinguishes E. sicanus from E. balearicus, E. tergestinus, and E. koschewnikowi. 364 V. FET ET AL. @ E. sicanus Mediterranean Sea A E. tergestinus \ Mt. Argentario Gia > Giannutri DISCUSSION @ E. italicus Fic. 9 A, Distribution map of E. sicanus, indicating DNA sampling sites. B, Map detail of the boxed area. E. sicanus: 1, Sardinia; 2, Mt. Argentario; 3, Giglio; 4, Giannutri; 5, Punta Ala; 6, Gargano; 7, San Domino, Tremiti; 8, Nebrodi, Sicily; 9, Malta; 10, Mt. Olympus, Greece; 11, Kalambaka, Greece. Outgroup samples: E. tergestinus: 12, La Morra, Piedmont; 13, La Spezia, Liguria; 14, Procchio, Elba; E. italicus: 15, Tortoreto. The phylogenies from the 16S data presented here support our morphological data. One should keep in mind that the partial sequence of this gene represents only a small fraction of the mitochondrial genome. It is also known that different genes may underlie different genealogies and that gene trees may not reflect species phylogenies EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 365 (Pamilo & Nei, 1988). Incomplete lineage sorting processes and introgression of mtDNA between two species/subspecies might be responsible for such inconsistencies (Page & Charleston, 1998; Avise, 2000). Recently, evidence for asymmetric introgres- sion of mtDNA in a buthid subspecies in the nuclear background of another subspecies and its subsequent complete fixation have been reported (Gantenbein & Largiadèr, 2002). However, in euscorpiids, so far, the phylogenetic information of the mitochon- drial 16S fragment has always been in concordance with independent nuclear genetic markers (allozyme data) (Gantenbein ef al., 1999, 2000a, 2001; Scherabon et al. 2000). Therefore, the phylogenetic pattern of the 16S gene is very likely to represent the “species tree” for the genus Euscorpius. For the taxonomic consequences we strongly rely here on the phylogenetic species concept (PSC) (Cracraft, 1989) as we el- evate E. carpathicus sicanus to the species level. Therefore, it seems reasonable to use morphological differentiation, i.e. trichobothrial patterns of the external aspect of patel- la, as a further support for the identified phylogenetic lineages. However, we are also aware that “deep” genetic divergences within morphologically conservative phyloge- netic groups such as scorpions or other arachnids in general have been reported in the absence of obvious morphological differences (Gantenbein et al., 2000a, 2000b; Bond et al., 2001). Scorpius sicanus C. L. Koch, 1837 was described from Sicily, and later syn- onymized with Euscorpius carpathicus (L.). Caporiacco (1950) has also mentioned this form, as a subspecies E. carpathicus sicanus, for the south of peninsular Italy (Apulia, western Calabria), Malta, and the northern African coast. Scorpius canestrinii Fanzago, 1872 was described from Sardinia, and later was synonymized with E. car- pathicus. Caporiacco (1950) treated these two forms as valid subspecies; our data show that they belong to the same species. Caporiacco (1950) also established a number of new subspecies for E. car- pathicus, mainly from Italy. Our morphological investigation of type material for all of these subspecies, and DNA sequences corresponding to some of them, demonstrate that six of Caporiacco’s subspecies (E. c. linosae, E. c. garganicus, E. c. ilvanus, E. c. argentarii, E. c. palmarolae, and E. c. calabriae) belong to E. sicanus (see below). Several other subspecies accepted or described by Caporiacco (1950) (E. c. apuanus, E. c. concinnus, E. c. niciensis, E. c. aquilejensis, E. c. picenus, E. c. oglasae, and E. c. corsicanus) were recently synonymized with E. tergestinus by Fet & Soleglad (2002). Kinzelbach (1975) studied Greek populations of E. carpathicus, but did not dis- cuss external patellar trichobothria at all; as we know now, this character set is crucial for the understanding of the genus Euscorpius (Vachon, 1974, 1975; Soleglad & Sissom, 2001; Fet & Soleglad, 2002). Vachon (1975) applied the patellar trichobothria analysis to all known Euscorpius species. At the same time, the important work of Valle (1975) on E. carpathicus appeared independently; Vachon (1975) in a footnote (p. 645) informed that his paper was already in print when he received Valle’s publi- cation. Valle (1975) attempted to analyse some of Caporiacco’s subspecies described from Italy by using external patellar trichobothria. Probably because Valle used the old system of designations established by Vachon (1963), his data were ignored by subse- 366 V. FETET AL. quent researchers (e.g. Fet, 1986, 1997a, 2000; Scherabon, 1987; Lacroix, 1991a, 1991b), who adhered to the new system by Vachon (1974, 1975, 1978, 1981). Note that Valle scored external patellar trichobothria summarily for both pedipalps, while Vachon’s numbers refer to a single pedipalp. The two systems differ in assignments of individual trichobothria into several groups (“series”). However, comparing Euscorpius trichobothrial schemes in Fig. 1 in Valle (1975), Fig. 3 in Vachon (1963; “old system”), and Fig. 17 in Vachon (1975; “new system”), we can adjust Valle’s sys- tem to Vachon’s conventions used today (Table 1). This rather elaborate adjustment allows one to use Valle’s valuable statistical data, which are based on the analysis of large series from various localities, and which can be interpreted in much more detail than the data of Caporiacco (1950) who used only total counts for patellar external trichobothria. Valle (1975) was the first to identify the most important trend in E. car- pathicus trichobothriotaxy: variation,of the B2 series (i.e. eb). Valle averaged the summary numbers for both pedipalps together (Valle, 1975: Tables 1-4, Figs. 4, 6, 8-10), and his numbers can be established in terms of Vachon (1975) as given in the Table 2. Recently, Fet & Soleglad (2002) restricted the name E. carpathicus to popu- lations from Romania and established the species E. tergestinus (C. L. Koch, 1837) for populations from northern Italy. Valle (1975) specifically noted that all specimens from Piedmont and Liguria have B2=6 (eb=4); this refers to E. tergestinus. In our DNA phylogeny (Fig. 8), one of the monophyletic lineages is referable to E. tergestinus; the other Italian lineage (Euscorpius sicanus) 1s unexpectedly diverse geographically, and includes a number of populations, which are discussed below. Sicily (type locality of E. c. sicanus). For specimens from Sicily, Vachon (1975, Fig. 31) illustrated the character eb=5, and suggested that this could be a good charac- ter, sufficient to elevate this taxon to the species rank. Valle (1975) reported that in specimens from around the island B2 varies from 6 to 8 (eb from 4-4 to 5-5). From his Fig. 6 it is evident that eb=5 specimens are concentrated in the north (corresponding to our DNA samples from Nebrodi Mts.). It remains to be seen if only E. sicanus is present in Sicily, or if in fact there are two sympatric species of Euscorpius as is the case on mainland Italy. Since Koch’s type of E. sicanus originated from Messina (the very northeastern point of Sicily), this name can be associated with Sicilian specimens having eb=5. Apennine Peninsula. E. sicanus is found in central and southern parts of the Apennine peninsula, from Tuscany to Calabria. Our data show that this species is found in Tuscany inland (Siena) as well as on the Tyrrhenian coast (Mt. Argentario, Follonica, Punta Ala), and on the nearby islands of the Tuscany Archipelago: Giannutri, Giglio, Capraia, Gorgona (MZUF; Caporiacco, 1950; our data). Further south we find this species on Palmarola (MZUF) and on the Capri (ZMH) Islands in the Tyrrhenian sea. Caporiacco (1950) listed E. c. sicanus not only for Sicily (type locality) but also for the mainland Italy, from Apulia (Bari area) and western Calabria. In addition, Caporiacco (1950) described several new subspecies, the differences between which, according to our study of his type material, are insignificant and usually amount to EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 367 minor variations in morphosculpture or coloration. E. carpathicus calabriae was described from Calabria; E. c. argentarii, from an isolated massif (ecological refu- gium) of Mt. Argentario in Tuscany; and E. c. garganicus from Apulia (Gargano Peninsula and Tremiti Islands in the Adriatic Sea). Marcuzzi et al. (1963: 81-88) conducted a statistical study on a number of populations from Apulia (using total trichobothrial counts), and could not find characters separating E. c. garganicus from other Apulian populations. Valle (1975, Fig. 8, Table 3) demonstrated B2=8 (eb=5) for most of southern Italy; he specifically noted (p. 228) that this character is not found north of Tuscany and Marches. However, we discovered a population with eb=5 also in Trieste (ZMH); two independent museum records (F. Werner in 1891 and C. Attems in 1901) exclude the possibility of erroneous labeling. It remains to be seen if E. sicanus occurs naturally in the northern Adriatic region or if the Trieste record represents an introduced population. Two other island populations near Italy have been given subspecific names: E. c. palmarolae from the Palmarola Island in the southern Tyrrhenian sea (off the Latium coast) and E. c. ilvanus from the Tuscany Archipelago. Both also exhibit eb=5. Our DNA data shows that all Tuscany populations (mainland and island alike) exhibit very close haplotypes (genetic distance less than 1%), and belong to the same mono- phyletic clade as E. sicanus (Fig. 8, A). The syntype series of E. c. ilvanus originates from several islands of the Tuscany Archipelago (including Elba), and also from Capri. However, our analysis of this material shows that all of Caporiacco’s specimens from Elba and Capri have eb=4, 1.e., they belong to E. tergestinus. This is confirmed by our DNA data (see above). Valle (1975, Fig. 4) also noted Elba populations as having B2=6 (eb=4); Capri may have two sympatric species (which is also possible for Sicily; see above). According to Caporiacco, the rest of the syntype series of E. c. ilvanus origi- nates from Giannutri, Cerboli, Capraia, Giglio, and Gorgona islands. Only syntypes from Giannutri, Cerboli and Capraia are in MZUF, all with eb=5 and eb,=4; among these, we selected a lectotype from Giannutri. Valle (1975) provides statistics for 145 specimens from Giglio and 74 from Giannutri, most of which have B2=8 (eb=5); he also shows this character for specimens from Cerboli, Gorgona, Mt. Argentario, the coast of Tuscany, and Capri (Valle, 1975, Fig. 4). DNA haplotypes from Giannutri (EcGI1) and Giglio (EcGG1) islands are very close to each other and to a haplotype from the mainland (EcPU1) (Punta Ala, identical with Follonica and Siena haplotypes) (Fig. 8). Another haplotype from Tuscany, which also falls in the E. sicanus clade, is found on Mt. Argentario (EcAR1); it is identical to a second haplotype from Siena and to one from Orbetello. The morphological character combination eb=5 and eb,=4, which is typical for Sicily, persists through some Italian populations of E. sicanus, such as those in Tuscany, including the islands of Tuscany Archipelago (type series of E. c. ilvanus), Mt. Argentario (type series of E. c. argentarii); Gargano Peninsula and Tremiti Islands (type series of E. c. garganicus; additional material studied); and Calabria (type series of E. c. calabriae). 368 V. FET ET AL. Sardinia. Our DNA phylogeny (Fig. 8) demonstrates that the Sardinian popul- ation (EcSD1), described originally as a separate species (Scorpius canestrinii), falls within the monophyletic clade of E. sicanus. Caporiacco (1950) studied 75 specimens from this population and treated it as a separate subspecies, E. c. canestrinii.. For the Sardinian population, Vachon (1975) mentioned eb=5. Valle (1975, Table 4) reported B2=8 (eb=5-5), but he also noted that B3 is normally 10 (eb,=5-5). This was confirmed in a detailed analysis and redescription by Vachon (1978) who studied 142 specimens from Sardinia. These specimens consistently possessed eb=5 (94.7% of 284 pedipalps scored), as illustrated in Vachon (1978, Fig. 6; 1981, Fig. 11). Some variation in eb, was observed; while it was predominantly 5 (in Vachon’s series, 71.8% of pedipalps), others (e.g. in a population from Sordogno that we studied) have eb, =4 (25.7%) and even 6 (2.5%). Vachon (1978: 329) noticed that the feature eb=5 was shared with the Sicilian subspecies (E. c. sicanus). Pelagie Islands and Pantelleria. These small islands (belonging to Italy) in the southern Mediterranean Sea between Sicily and Africa are known to have populations of “E. carpathicus”. Caporiacco (1950) described a new subspecies from Linosa Island, E. c. linosae, but our analysis of the type specimen from MZUF clearly indi- cates that it belongs to the E. sicanus clade (eb=5, eb, =4). Valle (1975, Table 2) analysed 13 specimens from Linosa, all with B2=8 (eb=5); he also mentioned a popul- ation from Lampedusa Island but did not characterize it. Crucitti (1993: Fig. I) reported “E. carpathicus” from Pantelleria Island. No DNA data are as yet available from scorpions from the Pelagie Islands or Pantelleria. Malta. Caporiacco (1950) was the first to report E. c. sicanus for Malta. Vachon (1975: 643) mentioned that specimens from Malta have eb=5. At the same time, Kinzelbach (1975), who expanded an alleged “hybrid between E. carpathicus and E. mesotrichus” to many Mediterranean populations, listed it also for Malta. Schembri & Schembri (1990) reported specimens from all of the Maltese islands (Malta, Gozo, Comino, and St. Paul’s Islands) as belonging to E. c. candiota Birula, 1903, following Kinzelbach (1975). This identification was also given by Kritscher (1992) and Schembri (1992). Our analysis of Kritscher’s original specimens deposited at NMW and of new Maltese material clearly indicates that it belongs to the E. sicanus clade (eb=5, eb,=4), as stated by Caporiacco (1950). Trichobothria counts also show simi- larity between populations on Malta and Sicily, with series er and v about 20% lower on Sicily and Malta as compared to Sardinia. Our DNA phylogeny further demon- strates that the Maltese population is closest to the Sicilian E. sicanus (Fig. 8, A). This is to be expected, given the geological history and biogeographic affinities of the Maltese fauna; the Maltese Islands were connected to Sicily, and possibly to North Africa, in Late Miocene (Messinian) times, and may have been connected again during Pleistocene sea-level lows (Thake, 1985; Schembri, 1992; Hunt & Schembri, 1999). As for E. c. candiota Birula, this taxon was originally described from Crete (Birula, 1903), and Kinzelbach’s (1975, 1982) interpretation that this species ranges beyond the Aegean Sea is not justified by either morphology or DNA analysis (Fet et al., in pre- paration). EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 369 North Africa. “E. carpathicus” has been reported many times from isolated localities along the north African Mediterranean coast (Tunisia, Libya, Egypt) (Simon, 1885; Birula, 1909; Caporiacco, 1950; Valle, 1975; Vachon, 1952, 1975, 1978; Fet & Sissom, 2000; Stathi & Mylonas, 2001). Specimens from North Africa were listed un- der E. c. sicanus by Caporiacco (1950) and by Vachon (1975, 1978). Our analysis of museum material shows that these populations conform to the “sicanus” trichobothrial formula. Valle (1975, Table. 4) analysed 15 specimens from Tunisia and one from Libya (Cyrenaica), but his data are unclear since he quoted B2 as mostly 9 or even 10. In this case, Valle might have added up the eb, and eb series, thus eb=eb,=5. Vachon (1978: 328) mentioned specimens from Tunisia (Carthage and Tunis) with eb=eb,=5, and from Tunisia (Sfax) and Libya (Cyrenaica) with eb=5 and eb,=4. We analysed four specimens from Tunisia (Tunis and Djebel bou Kournine), all with eb=5 and eb,=4. However, a single specimen from Tripoli, Libya (Birula, 1909) that we have seen in NMW has eb=eb,=4. Recently, Stathi & Mylonas (2001) reported a new locality at Senebat Lauela Gorge in Kuff National Park, Ruins of Cyrrini, 5 km SE of El Beida, Libya. No DNA is available from the northern African enclaves yet; it remains to be seen if these are true relict populations or if they have been introduced via human activity in historical times. Such transplantation are not unusual in Euscorpius; see for example, the populations of E. flavicaudis in England (Fet & Sissom, 2000), and Austrian populations of E. tergestinus (Huber et al., 2001; as E. carpathicus). Madeira. The single specimen known from the island of Madeira (Portuguese territory in the Atlantic Ocean) was first listed by Kinzelbach (1982) under “E. mesotrichus”. An analysis of this specimen (deposited in the NMM) demonstrated that it has eb=eb,=5, i.e. it is closest to the populations from Tunisia and Sardinia. The Madeira population of Euscorpius is the westernmost known. As with the North African populations, it remains to be shown if the one from Madeira was introduced or represents a true relict. Madeira is known for its relict Mediterranean-type biota. Greece. A nomenclatural controversy surrounds some poorly studied Greek populations belonging to the “E. carpathicus” complex (Fet, 2000; Fet & Braunwalder, 2000; Fet & Sissom, 2000). Kinzelbach (1975) observed sympatry of two morpho- logically different forms in Greece (Thessaly) and separated E. carpathicus (L.) from another species, which he reported as “E. mesotrichus HadZi, 1929”. This name, how- ever, is permanently invalid since it refers to E. carpathicus mesotrichus HadZi, a junior primary homonym of E. italicus mesotrichus Hadzi (Caporiacco, 1950; Fet 1997b) (ICZN Article 57.2). The name “E. mesotrichus Hadzi” was used later by some authors (Michalis & Kattoulas, 1981; Kinzelbach, 1982, 1985; Kritscher, 1993), while others (Vachon, 1978, 1981; Fet, 1986, 1997a) did not comment on Kinzelbach’s division. Fet & Sissom (2000) agreed with the observation by Kinzelbach (1975) that at least two different species of the “E. carpathicus” complex are present in Thessaly, and tentatively accepted the name E. tergestinus for one of them as an available senior synonym of the homonymous E. c. mesotrichus Hadzi (these names were first synonymized by Caporiacco, 1950). Our present study indicates that the Thessalian 370 V. FETETAL. populations, indeed, belong to two species. However, neither of them is E. tergestinus; as defined recently by Fet & Soleglad (2002), this species is not found in Thessaly. One of Thessalian species, referred to as E. carpathicus s.str. by Kinzelbach (1975), is not yet identified; it does not belong to E. carpathicus(L.), which was recently restricted to Romania (Fet & Soleglad, 2002). Another Thessalian form, listed as E. mesotrichus by Kinzelbach (1975) and as E. tergestinus by Fet & Sissom (2000), according to the morphological features, belongs to E. sicanus. The DNA analysis also places the Thessalian populations of “E. mesotrichus” (EcOL2, EcKB1) into the E. sicanus clade. While in MP analysis the branching order is unresolved (Fig. 8, B), in the ML analysis (Fig. 8, A) these Thessalian populations form a sister group to E. sicanus populations from Sardinia, the Adriatic region, Sicilia, and Malta as an outgroup clade. The distribution of E. sicanus in Greece is not limited to the populations from Mt. Olympus massif and northern Pindus Mts. (Kalambaka, valley of the Peneus River), the DNA of which we analysed. According to the museum data studied by V.F. and M.E.S. (which include the ample material previously discussed by Kinzelbach (1975) and Kritscher (1993)), this species is common in Greece found from Macedonia and Thessaly (with the Sporades) to Attica and Peloponnese (see map at Fig. 9, A). The specimens reported by Michalis & Kattoulas (1981) as “E. mesotrichus” from Lakonia, Peloponnese, also probably belong to the same species. Two “Group B” specimens of E. carpathicus reported by Fet (2000, p. 55) from Peloponnese (Mistras) also belong to E. sicanus. Judging from trichobothrial data of specimens (pp. 43-45, Fig. 3) recently reported from many localities from the Peloponnesus (Mistras) under E. car- pathicus by Crucitti & Bubbico (2001) these populations also belong to E. sicanus. The Greek populations are the subject of a continuing study (Fet et al., in preparation). Their morphology varies, but all have the unmistakable “sicanus” signature of eb=5; many populations (Mt. Olympus, Mt. Ossa, Sporades, Taygetos Mts.) also have eb,=5. E. sicanus reaches its highest recorded altitude at Mt. Tsumerka, Pindus Mts., 1500 m a.s.l. (Kritscher, 1993, under “E. mesotrichus”). We thus confirm the presence of more than one form of the “Euscorpius car- pathicus” complex in Thessaly, as first stated by Kinzelbach (1975). While one of these (erroneously cited as “E. mesotrichus HadZi”) belongs to E. sicanus, the nomenclature of the other form, which has eb=4, remains to be established. It cannot be called E. carpathicus, since that species was shown to be restricted to Romania by Fet & Soleglad (2002). A number of former subspecies described in the genus Euscorpius have been recenlty elevated to the species rank using molecular and morphological criteria (E. tergestinus, E. hadzii, E. koschewnikowi, E. balearicus, E. alpha, E. gamma; see Gantenbein er al., 2000, 2001; Scherabon er al., 2000; Fet & Soleglad, 2002); other former subspecies have been synonymized with these species, therefore largely eliminating the need of subspecies category in Euscorpius. It is important to notice that E. sicanus was not found on the majority of the Aegean islands (for trichobothrial data on specimens from 9 islands including Crete, see Fet, 1986, 2000). It was, however, recorded from the Sporades (Skiathos, Alonyssos, Skopelos) under E. mesotrichus by Kritscher (1993). Furthermore, a form with eb=5 was found on Tria Nisia, a small island group in the southwestern Aegean Sea (Fig. 9, A); this specimen was recorded by Werner (1936: 17) as E. carpathicus). EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 371 As a result of our study, one more valid species, E. sicanus, with a complex (and likely ancient) geographic distribution and with considerable intraspecific variation, is added to the genus Euscorpius. Further investigations on the “Euscorpius carpathicus” species complex are desirable, especially for a number of poorly studied (both named and unnamed) populations in the Balkans, Turkey, and Italy. TAXONOMIC CONCLUSIONS Euscorpius sicanus (C. L. Koch, 1837) Scorpius sicanus C. L. Koch, 1837: 106, Pl. CVII-CVIII, Figs 247-248. TYPE MATERIAL. Holotype (female) from Messina, Sicily, Italy (leg. Wagner), formerly in J. Sturm’s collection in Nuremberg (Birula, 1917), now presumed lost (Fet & Sissom, 2000). The designation of a neotype according to the ICZN (Article 75) is desirable for the purposes of taxonomic stability. SYNONYMS Scorpius canestrinii Fanzago, 1872: 78, Fig. 1, la, 1b, syn. n. Lectotype (designated by Kovarik, 1997: 182): female (HNHM 2277-214), Sardinia, Italy (coll. Canestrini, det. Fanzago). Paralectotype: 1 male (pictured by Vachon, 1978: Figs 1-7) (HNHM), Sardinia, Italy. Euscorpius carpathicus linosae Caporiacco, 1950: 184, 227, syn. n. Lectotype (the only avail- able specimen; designated here from the syntype series in order to preserve stability of nomenclature according to the ICZN Article 74), female (MZUF 30/5970), Linosa Island, Pelagie Islands (Mediterranean Sea), Agrigento, Italy. Paralectotypes (deposito- ry unknown): 8 specimens, the same data as for the lectotype. Euscorpius carpathicus garganicus Caporiacco, 1950: 189, 226, syn. n. Lectotype (designated here from the syntype series in order to preserve stability of nomenclature according to the ICZN Article 74): male (MZUF 90/5902), San Domino Island, Tremiti Islands, Foggia, Apulia, Italy, 16 April 1940 (Pomini). Paralectotypes (all from Foggia, Apulia, Italy, coll. Pomini): 4 females, 3 males (MZUF 88/5903-5909), Caprara Island, Tremiti Islands, Foggia, Apulia, Italy, 13 April 1940; 3 females, 1 male (MZUF 86/5892-5895), Bosco Ginestra, Gargano, Apulia, 20 April 1940; 2 females (MZUF 86/5896-5897), Alveo del Lago S. Egidio, 20 April 1940; 1 male (MZUF 87/5910), Foresta Umbra, Gargano, Apulia, September 1940; 1 female (MZUF 92/5910), Sfilze, Gargano, Apulia, 25 April 1940; 3 females (MZUF 91/5899-5901), Sannicandro Garganico, Gargano, Apulia, 25 April 1940. Other paralectotypes (depository unknown): 44 specimens from Gargano and Tremiti. Euscorpius carpathicus ilvanus Caporiacco, 1950: 195, 229 (in part), syn. n. Lectotype (desig- nated here from the syntype series in order to preserve stability of nomenclature accord- ing to the ICZN Article 74): male (MZUF 98/5745), Giannutri Island, Tuscany, Italy, 1878 (G.B. Toscanelli). Paralectotypes (all from Livorno, Tuscany, Italy): 5 females, 1 male (MZUF 98/5744, 99/5746-5751), same label as lectotype; 4 females, 7 males (MZUF 118/5725-5735), Cerboli Island, 1879 (G.B. Toscanelli); 1 female (MZUF 117/5755), Capraia Island, November 1927. Other paralectotypes (depository unknown): 14 males, Gorgona Island; 39 males, Giglio Island. Note: part of the original syntype series, i.e., specimens, from Elba Island (6 females, 2 males, MZUF 120/5736-5743, January 1879 (G.B. Toscanelli)) and from Capri Island (1 female, 2 males, MZUF 141/5752-5754, 31 July 1879 (Corio)) are excluded from this designation since they belong to E. tergestinus. Euscorpius carpathicus argentarii Caporiacco, 1950: 196, 228, syn. n. Lectotype (designated here from the syntype series in order to preserve stability of nomenclature according to the ICZN Article 74): female (MZUF 5977): Mt. Argentario, Grosseto, Tuscany, Italy, 3 November 1874 (Mayor). Paralectotypes: 6 females, 1 juv. male (MZUF 5978-5979, 6256-6260), from the same locality as the lectotype. 372 V. FETETAL. Euscorpius carpathicus palmarolae Caporiacco, 1950: 196, 228, syn. n. Lectotype (designated here from the syntype series in order to preserve stability of nomenclature according to the ICZN Article 74), Palmarola Island, Pontine Islands (Tyrrhenian Sea), Latium, Italy, 1878 (G.B.Toscanelli). Paralectotypes: 2 females, 1 male (MZUF 111/5755, 5758- 5759), from the same locality as the lectotype. Euscorpius carpathicus calabriae Caporiacco, 1950: 200-201, 228 (in part), syn. n. Lectotype (designated here from the syntype series in order to preserve stability of nomenclature according to the ICZN Article 74), 1 female (MZUF 61/5887), Colle di Pizzo, Arena, Catanzaro, Calabria, Italy, 1874 (G. Cavanna); Paralectotypes (all from Calabria, Italy, coll. G. Cavanna): 6 females, 2 males (MZUF 61/5883-5886, 5888-5891), same locality as lectotype; 2 females (MZUF 57/5871, 5873), Filandari, Catanzaro, 13 April 1874; 1 female (MZUF 59/5874), Foresta della Mongiana near Mt. Pecoraro, Catanzaro, 25 June 1874; 1 female (MZUF 172/5868), Palmi, Reggio Calabria, 6 June 1874; 1 female, dam- aged (MZUF 170/5875), Timenovo (mountains near Oppido), Reggio Calabria, 22 May 1874. Other paralectotypes (depository unknown, all from Calabria, Italy): 3 males, Fago del Soldato; 1 female, Monte Consolino (Stilo); 19 males, Monte Oliveto; 17 males, Soveria Manetti; 1 female, Vıbo Valenzia; female, Melossena Cave near Maratea (Potenza, Basilicata). Nore: 5 specimens from the syntype series (MZUF 171/5876, 5879; 172/5869, 5870, 5880; Palmi, Reggio Calabria, 1874 (G. Cavanna)) do not match E. calabriae in morphology (eb=4) and are not included in the paralectotype list (they may belong to E. tergestinus). REFERENCES (SELECTED) Scorpius sicanus: C. L. Koch, 1850: 86; Fanzago, 1872: 83, Fig. 4. Euscorpius carpathicus: Birula, 1909: 518; Werner, 1936: 17; Caporiacco, 1950: 178-209 (in part); Marcuzzi er al., 1963: 81-88; Curcié, 1972: 84-86 (in part); Valle, 1975: 209-234 (in part); Crucitti, 1993: 289-291, Fig. 2 (in part); Fet & Braunwalder, 2000: 19, Fig. 3 (in part); Fet & Sissom, 2000: 357 (in part); Stathi & Mylonas, 2001: 289 (in part). Euscorpius sicanus: Pavesi, 1876: 430; Birula, 1900: 15; Birula, 1917: 105; not Kulczynski, 1887: 349-354 (Tyrol), misidentification. Euscorpius canestrinii: Birula, 1900: 15; Birula, 1917: 105. Euscorpius carpathicus sicanus: Caporiacco, 1950: 183, 227; Valle, 1975: 219, 223; Vachon, 1975: 642-643, Fig. 31; Vachon & Jaques, 1977: 413; Vachon, 1978: 328; Lacroix, 1991a: 19, Fig. 64; Fet & Sissom, 2000: 365. Euscorpius carpathicus canestrinii: Caporiacco, 1950: 184, 227; Valle, 1975: 233; Vachon & Jaques, 1977: 415, 431; Vachon, 1978: 321-230, Fig. 1-7; Vachon, 1981: 199, Fig. 7, 11; Lacroix, 1991a: 19, Fig. 58; Kovarik, 1997: 178, 182, Fig. 1-4; Fet & Sissom, 2000: 363. Euscorpius mesotrichus: Kinzelbach, 1975: 30-38 (in part); Kinzelbach, 1982: 61-63 (in part); Kinzelbach, 1985, Map IV (in part); Michalis & Kattoulas, 1981: 109-110; Kritscher, 1993: 385-386 (in part). Euscorpius carpathicus calabriae: Valle, 1975: 232; Bartolozzi et al., 1987: 296; Lacroix, 1991a: 19 (calabria!); Fet & Sissom, 2000: 362. Euscorpius carpathicus ilvanus: Valle, 1975: 219; Bartolozzi et al., 1987: 296; Lacroix, 199 1a: 19; Fet & Sissom, 2000: 364. Euscorpius carpathicus garganicus: Bartolozzi et al., 1987: 296-297; Lacroix, 1991a: 19. Euscorpius carpathicus argentarii: Bartolozzi et al., 1987: 296; Lacroix, 1991a: 19; Fet & Sissom, 2000: 361. Euscorpius carpathicus palmarolae: Bartolozzi et al., 1987: 296; Lacroix, 1991a: 19; Fet & Sissom, 2000: 362. Euscorpius carpathicus linosae: Bartolozzi et al., 1987: 296; Lacroix, 1991a: 19; Fet & Sissom, 2000: 364. Euscorpius carpathicus candiota (not Euscorpius candiota Birula, 1903, misidentification): Kinzelbach, 1975: 34, 36-37 (in part); Schembri & Schembri, 1990: 20; Kritscher, 1992: 185-188, Figs. 1-4. EUSCORPIUS SICANUS FROM ITALY, MALTA & GREECE 373 Euscorpius carpathicus carpathicus: Kritscher, 1993: 383-384 (in part), misidentification. Euscorpius tergestinus (not Scorpius tergestinus C. L. Koch, 1837, misidentification): Fet & Sissom, 2000: 372 (in part; Greece). MATERIAL STUDIED Specimens used for DNA analysis (see above); type material of E. c. linosae, E. c. gar- ganicus, E. c. ilvanus, E. c. argentarii, E. c. palmarolae, E. c. calabriae (see above) (MZUF); other museum materials: GREECE. 2 females (ZMH), Plataria, Epirus, 20 June 1986 (K. Michalis & P. Dolkeras); 1 juv. female (NMM 0176), road between Rodohori (=Rodochorion) and Naoussa, Macedonia, August 1972 (H. Georgiadis); 1 male (NMW 2130), Chortiatis near Thessaloniki, Macedonia, 9 October 1966 (J. Gruber); 1 male (NMW 16006), Mt. Tsumerka, 1500 m, Pindus Mts., 18 August 1990 (H. Sattmann); 14 females, 7 males (VF), Visitsa, 800 m, Mt. Pilion, Thessaly, 7 May 2001 (V. Fet); 32 females, 11 males (VF), Spilia, 800 m, Mt. Ossa (=Kissavos), Thessaly, 9 May 2001 (V. Fet); 4 females (VF), Kalambaka, 200 m, Thessaly, 14 May 2001 (V. Fet); 3 females, 1 male (NMW 16.035/1-4), Litohoro (=Lithochorion), near Mt. Olympus, Thessaly, 9 June 1984 (E. Kritscher); 1 female (VF), 3 km N of Litohoro, Mt. Olympus, 15 May 2001 (V. Fet); 1 female, 1 juv. male (NMM 0209a), Agios Dionysios, 820 m, Mt. Olympus, Thessaly, 28 August and 10 September 1973 (H. Pieper); 1 female, 1 juv. male (NMM 0206), between Petra and Kokkinopilos, 520 m, Mt. Olympus, Thessaly, 29 September 1973 (H. Pieper); 3 females (NMM 0261), Mt. Olympus, Thessaly, 26 May 1974 (H. Malicky); 6 females, 4 males (ZMH), Karditsa, 560 m, Thessaly, 24 June 1986 (K. Michalis & P. Dolkeras); 4 females (ZMH 30), Sykouri (=Sykourion), Thessaly (K. Michalis & P. Dolkeras); 1 male, 1 female (NMW 16040/1-2), Lazareta, Skiathos Island, Sporades, Thessaly, 24 June 1985 (E. Kritscher); 1 female (NMW 16041), Skopelos Island, Sporades, Thessaly 14 June 1985 (E. Kritscher); 2 females (NMW 16031/1-2), Milia, Alonissos Island, Sporades, Thessaly, 11 June 1985 (E. Kritscher); 1 juv. (NMW), Mt. Parnes, Attica, 1200 m, 18 May 1969 (H. Rausch); 2 females (NMNHS 68), Mistras, Laconia, Peloponnese, 18 September 1983 (P. Beron & V. Beshkov); 1 male (NMW 16039/2), road between Leptokaria and Karia, 14 km, Mani, Peloponnese, 9 June 1984 (E. Kritscher); 4 females, 4 males (NMM 0201) Taygetos Mts., W of Profitis Ilias, 1200 m, spruce forest, Peloponnese, 28 September 1973 (R. Kinzelbach); 1 male (NMW 2193), Tria Nisia (the southernmost island), Dodekanese Islands, Aegean Sea (O. Wettstein). ITALY. 5 specimens (NMW 11299), Messina, Sicily, 1906 (K. Holdhaas); 4 females, 3 males (ZISP 98-913), Sordogno, Sardinia (A. Krausse); 8 females, 3 males (ZMH), Mt. St. Angelo, Gargano, Apulia, 6-8 September 1925; 1 female, 1 male juv. (ZMH), Capri Island, August 1928; 2 females, 1 male (ZMH), Trieste, September 1891 (F. Werner); 3 females, 2 males (ZMH), Trieste, 23 December 1901 (C. Attems). MADEIRA. 1 female (NMM 0204), 2 April 1971 (U. Hecker) (Kinzelbach, 1982; as identified as “E. mesotrichus”). MALTA. 2 specimens (NMW 1857), Wied-il-Babu, near Zurrico [=Wied Babu near Zurrieq], Malta, 16 April 1913; 1 female (NMW), Zurrico [=Zurrieq], Malta, 16 April 1913; 5 females, 1 male (NMW 15.040), Floriana, Malta, 15 June 1990 (E. Kritscher); 1 female (NMW 15.038), Marsalform [=Marsalforn], Gozo Island, 28 May 1990 (E. Kritscher); 6 females, 1 male (NMW 15.039), Marsalform [=Marsalforn], Gozo Island, 31 May 1990 (E. Kritscher). TUNISIA. 2 females (ZMH), Tunis, 23 May 1910 (Wegehn); 1 female, 1 male (NMW 2180), Djebel bou Kournine, 13 April 1913 (F. Werner). LIBYA. 1 male (NMW 2179), Tripoli, August 1906 (B. Klaptocz). DISTRIBUTION Italy (with Sardinia and Sicily), Malta, Greece, northern Africa (Tunisia, Libya, Egypt), Madeira. 374 V. FET ET AL. TABLE |. Correspondence of external patellar trichobothrial designations in Euscorpius by Valle (1975) and Vachon (1975). Valle (1975) (scored for both sides) Vachon (1975 and later) (scored for each side) D1 (variable) (et-right - 1) + (et-left - 1) D2 (constant, always 8, 1.e. 2 from et and 2 from est on each side =4 4 on each side) D3 (constant, always 6, i.e. 3 on [(est-right - 2) + 1] + [(est-left - 2) +1] =6 each side) (est is always 4) D4 (variable) em-right + em-left B4 (constant, always 4, i.e. 2 on (esb-right + esb-left)=4 (esb is always 2) each side) B3 (variable) eb ‚right + eb,-left B2+BI (B2 is variable; eb-right + eb-left Bl is always 2, as it includes only 1 on each side) TABLE 2. Variation of external patellar trichobothrial series B2 (Valle, 1975) corresponding to series eb (Vachon, 1975). Valle (1975) Vachon (1975 and later) B2 eb 6 (3-3) 4-4 7 (3-4 or 4-3) 4-5 or 5-4 (for an asymmetric individual); or 4-4 & 5-5 (when averaged per locality) 8 (4-4) 5-5 ACKNOWLEDGEMENTS We are grateful to Adolf Scholl, Carlo R. Largiadér, Mark Barker, W. Ian Towler, and Pierangelo Crucitti for their enthusiastic cooperation in the initiation and pro- motion of the genetic characterization of Euscorpius. We also are grateful to all those who over the years donated and loaned material and rare literature, and facilitated study of Euscorpius in many European museums. This includes (but is not limited to) Janet Beccaloni, Petar Beron, Alberto Bonacina, Matt Braunwalder, Pierangelo Crucitti, BoZidar Curcié, Hieronymus Dastych, Christo Deltshev, Jason Dunlop, Gérard Dupré, Jiirgen Gruber, Dietmar Huber, Peter Jager, Ragnar Kinzelbach, Frantisek Kovarik, Victor Krivochatsky, Christian Kropf, Matjaz Kuntner, Jean- Bernard Lacroix, Wilson Lourenco, Kirill Mikhailov, Vladimir Ovtsharenko, Guillem Pons, Valentin Popa, Jan Ove Rein, Bernhard Scherabon, Paul Selden, Boris Sket, Iasmi Stathi, Roland Stockmann, Alex Winkler, Marco Valle, and Mark Volkovich. 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E-mail: alain.dechambrier@mhn.ville-ge.ch ; claude.vaucher@mhn. ville-ge.ch The genus Amphoteromorphus (Cestoda: Proteocephalidea), parasite of neotropical catfishes : systematics including morphological data and protein electrophoresis. - The present paper provides a detailed morpho- logical revision of the genus Amphoteromorphus together with a study of isoenzymes. The authors conclude that the concerned genus contains at present the following six species: A. peniculus Diesing, A. parkamoo Woodland, A. piraeeba Woodland, A. piriformis sp. n., A. ovalis sp. n. and A. ninoi sp. n. A. peniculus is unequivocally distinguished by irregularly alterning genital pores. A. ninoi is characterised by the number of testes, by the position of vitellaria and by the number of uterine diverticula; its specific isolation is confirmed by isoenzyme electrophoresis. The remaining four species are closely related. A. piriformis and A. ovalis possess distinctive embryophore shape. A. ovalis is also distinguishable by its electrophoretic profile. Key-words: Cestoda - Monticelliidae - Amphoteromorphus - fish parasites - 1zoenzyme electrophoresis - new species - taxonomy. INTRODUCTION Le genre Amphoteromorphus Diesing, 1850, parasite de poissons chats en Amérique tropicale, comprend selon Freze (1969) et Schmidt (1986), trois espèces: A. peniculus Diesing, 1850 parasite de Brachyplatystoma flavicans, A. piraeeba Wood- land, 1934 parasite de B. filamentosum et de B. flavicans et A. parkamoo Woodland, 1935 parasite de Paulicea luetkeni. Des récoltes effectuées en 1992 et 1995 en Amazonie nous ont fourni un abon- dant matériel, dont quelques spécimens provenant d’un hòte inédit, Brachyplatystoma vaillanti. L’étude morpho-anatomique et génétique par électrophorèse des izoenzymes nous conduit à décrire trois espèces nouvelles dans ce genre. Manuscrit accepté le 21.10.2002 382 M. CARFORA ET AL. MATERIEL ET METHODES Les hötes (récoltés par des pécheurs dans la région de Itacoatiara, dans le fleuve Amazone en 1992 et 1995) ont été disséqués et examinés immediatement apres leur mort. Le tractus digestif, fendu dans toute sa longueur, a été fixé dans une solution de formaldéhyde a 4 % bouillante. Apres le tri au laboratoire, les helminthes ont été conservés dans l’éthanol à 75 %. Les cestodes ont été colorés au carmin chlorhydrique de Mayer, différenciés a l’éthanol acide, déshydratés, éclaircis dans l’eugénol et montés au Baume du Canada. Les coupes sériées transversales ou frontales de 12-15 um d’épaisseur ont été colorées a l’Hématoxyline de Weigert / éosine et montées au Baume du Canada selon le processus récemment exposé par de Chambrier (2001). Les œufs ont été montés extemporanément dans l’eau distillée. Tous les parasites sont déposés au Muséum d’histoire naturelle de Geneve (INVE) et dans la collection d’helminthologie de l’Institut Oswaldo Cruz (CHIOC). Sauf indications contraires, les dimensions sont données en micrometres (um). Abréviations: ag: atrium génital; c: cirre; cd: canal osmorégulateur dorsal; cev: cirre évaginé; cg: cellules a cytoplasme granuleux; CHIOC = Institut Oswaldo Cruz, Rio de Janeiro; cm: musculature circulaire distale des ventouses; cv: canal osmorégu- lateur ventral; de: canal déférent; du: diverticules utérins; eb: embryophore; ebe: embryophore externe; ebi: embryophore interne; ee: enveloppe externe; gm: glande de Mehlis; h: crochets; INVE = Département des invertébrés du Muséum d’histoire naturelle de Geneve; Im: musculature longitudinale interne; MHNG = Muséum d’histoire naturelle, Geneve; mo: membrane oncosphérique; MT = matériel-type; n = nombre de mesures ou d’observations; od: oviducte; on: oncosphere; 00: oocapte; ov: ovaire; pc: poche du cirre; PC = rapport longueur de la poche du cirre sur la largeur du proglottis; pep: paroi épaissie de la poche du cirre; PG = situation du pore génital en % de la longueur du proglottis; rm: musculature radiaire des ventouses; rs: réceptacle séminal; s: coque; sv: sphincter vaginal; te: testicules; tu: tronc utérin; ud: utéroducte; ut: utérus; vb: ventouses bilobées; vc: canal vaginal; vd: vitellogenes dorsaux; vdp: vitellogenes dorsaux paramusculaires; vi: vitelloducte; vv: vitellogenes ventraux; vvp: vitellogènes ventraux paramusculaire; x = moyenne; * = spécimens utilisés pour l’élec- trophorese des protéines. Les morceaux d’individus destinés à l’électrophorèse des protéines ont été places dans un tube Eppendorf et conservés dans l’azote liquide, puis au congélateur (-80°C). Les échantillons ont ensuite été broyés individuellement au laboratoire dans 0.2 ml d’eau distillée. Cette source d’enzyme a été prélevée sur un rectangle de papier Whatman (1.0 x 0.2 mm) et stockées à -80°C. Les électrophorèses ont été réalisées sur gel d’amidon sous une tension d’environ de 80 V et d’une intensité maximale de 80 mA. Les révélations enzymatiques ont été effectuées selon les techniques décrites par Pasteur er al. (1987) ou selon Hillis er al. (1996) pour l’enzyme GCDH. Vingt-sept enzymes ont été testées: AAT (Aspartate amino-transférase, EC 2.6.1.1), ACO (Aconitase, EC 4.2.1.3), ACP (Phosphatase acides, EC 3.1.3.2), AK (Adénylate-kinase, EC 2.7.4.3), AKP (Alkaline-phosphatase, EC 3.1.3.1), ALD LE GENRE AMPHOTEROMORPHUS 383 (Aldolase, EC 4.1.2.13), CK (Créatine-kinase, EC 2.7.3.2), FK (Fructo-kinase, EC 2.7.1.4), GCDH (Glucose-déhydrogénase, EC 1.1.1.118), GLC (NAD-glucose-déhy- drogénase, EC 1.1.1.47), GPI (Glucose-6-phosphate-isomérase, EC 5.3.1.9), GPD (Alpha-glycérophosphate-déhydrogénase, EC 1.1.1.8), G6PD (Glucose-6-phosphate- déhydrogénase, EC 1.1.1.49), 6PGD (Phosphogluconate-déhydrogénase, EC 1.1.1.43), HK (Hexokinase, EC 2.7.1.1), IDH (Isocitrate-déhydrogénase, EC 3.1.1.42), LAP (Leucine-aminopeptidase, EC 3.4.11.1), LDH (L-lactate-déhydrogénase, EC 1.1.1.27), MDH (Malate-déhydrogénase, EC 1.1.1.37), ME (Enzyme-malique, EC 1.1.1.40), MPI (Mannose-6-phosphate-isomérase, EC 5.3.1.8), NP (Purine-nucléoside-phospho- rylase, EC 2.4.2.1), PEP (Peptidases, EC 3.4.-.-: Pep-A, Pep-B, Pep-C and Pep-D, cor- respondant au substrat suivant: Val-Leu, Leu-Gly-Gly, Lys-Leu, Phe-Pro) et PGM (Phosphoglucomutase, EC 2.7.5.1). La mobilité électrophorétique la plus lente est désignée par la lettre “a”. RESULTATS Amphoteromorphus Diesing, 1850 Monticelliidae La Rue, 1911, Zygobothriinae Woodland, 1933. Scolex pourvu de quatre ventouses bilobées. Présence d’un métascolex plissé a la base du scolex. Musculature longitudinale interne plus développée au niveau des marges latérales des proglottis, fibres musculaires formées de petits faisceaux groupés en une couche bien délimitée. Pores génitaux unilatéraux ou irrégulièrement alternes (espèce-type). Poche du cirre allongée, muni d’un cirre non armé. Vitellogènes corti- caux, parfois paramusculaires, en deux bandes dorsales et ventrales, aporales et post- porales (pas de vitellogènes préporaux). Testicules médullaires, antérieurs a l’ovaire et disposés en un champ. Ovaire médullaire bilobé ou compact a contour irrégulier. Vagin toujours antérieur a la poche du cirre. Uterus formé d’un tronc utérin médullaire et de diverticules médullaires. Parasites de poissons siluriformes néotropicaux. Espece-type: Amphoteromorphus peniculus Diesing, 1850. Amphoteromorphus peniculus Diesing Figs 1-4 Amphoteromorphus peniculus Diesing, 1850: 560. Hote: Brachyplatystoma flavicans (Castelnau, 1855) (Pimelodidae); nom vernaculaire: Dourada. Matériel examiné: Brésil, Villa dos Manes, type: 3 préparations totales, 5 préparations de coupes sériées, coll. Institut de Zoologie Neuchatel, collecté par Natterer (numéroté: Wien 613), C 35/82-89, 21.01.1834. Brésil, Etat de Amazonas, Itacoatiara, INVE 19313, 19.09.92; INVE 22214, 13.09.92. Localisation: Intestin. DESCRIPTION Strobiles acraspédotes, jusqu’a 50 mm de long. Tégument peu ou pas plissé. Proglottis matures et gravides larges de 945-1425 (x = 1265, n = 20) et longs de 475- 655 (x =570, n= 10). Scolex long de 1090 (n = 1) et large de 2275 (n = 1). Métascolex peu plissé et légèrement plus large que le cou. Ventouses bilobées, longues de 675-715 (n = 3), pourvues d’une musculature radiaire développée. Rapport entre longueur des ven- 384 M. CARFORA ET AL. ov gm oo od vv Fics 1-4 Amphoteromorphus peniculus Diesing, 1850. 1. 19313 INVE, scolex pourvu de quatre grandes ventouses bilobées. Remarquer la concentration des cellules 4 cytoplasme granuleux 4 l’apex. 2. Croquis d’oeufs “in toto”. 3. 19313 INVE, vue ventrale d’un proglottis mature. Ootype bien -visible 4. 22214 INVE, coupe transversale d’un proglottis mature. Echelles: 1 = 400 mm; 2 = 50 mm.; 3, 4 = 500 mm. LE GENRE AMPHOTEROMORPHUS 385 touses et longueur du scolex: 62-65 % (n = 3) (Fig. 1). Le lobe postérieur est 20-25 % plus petit que le lobe antérieur. Diametre du petit lobe de 380-400 (n = 4), celui du grand lobe de 495-510 (n = 4). Concentration importante a l’apex de cellules a cyto- plasme granuleux (Fig. 1). Musculature longitudinale interne constituée de faisceaux de fibres, avec concentration de ces faisceaux aux marges des proglottis. Canaux osmorégulateurs ventraux atteignant 65 de diamétre et chevauchant la poche du cirre dans sa partie terminale, parfois dans sa partie médiane. Canaux dorsaux d’un diamétre de 20-25 et passant dorsalement a la poche du cirre en position terminale ou subter- minale. Paroi des canaux dorsaux plus épaisse que celle des ventraux. Testicules médullaires ronds à ovoides, au nombre de 105-119 (x = 112, n= 11; MT = 100-136; n = 4), de 45-65 de diametre, disposés souvent en trois couches et répartis plus ou moins uniformément entre les canaux osmorégulateurs ventraux, n’atteignant pas les canaux dorsaux et ventraux (Fig. 3). Follicules vitellins corticaux formant deux bandes dorso-latérales et deux ven- tro-latérales triangulaires (Fig. 3). Rapport entre largeur des vitellogènes et largeur du proglottis: ventraux = 25-32 % (x = 30, n = 15); dorsaux = 10-20 % (x = 15, n = 15). Rapport entre longueur des vitellogenes et longueur du proglottis du côté poral: ventraux, 60-70 % (x = 65, n = 10); dorsaux, 57-71 % (x = 63, n = 10); du côté aporal: ventraux, 78-88 % (x = 82, n = 10); dorsaux, 75-89 % (x = 79, n = 10). Peu ou pas de follicules vitellins, ni de testicules dorsalement et ventralement a la poche du cirre. Poche du cirre allongée, a paroi mince, longue de 295-350 (x = 330, n = 10), PC SU = 26, n = 10; MT = 17-25, x = 22; n = 14)) (Fig. 3). Carre occupant jusqu’a environ 70 % (n = 11) de la longueur de la poche du cirre. Pores génitaux irregulierement alternant, PG = 14-17 % (x = 16, n= 8; MT = 18-25, x = 22, n = 14). Vagin toujours antérieur a la poche (n = 74; MT = 65), pourvu d’un sphincter muscu- laire. Canal éjaculateur très contourné. Canal déférent également très contourné, atteignant le milieu du proglottis et le dépassant méme dans certains cas. Ovaire bilobé, large de 550-875 (x = 735, n = 10), occupant les 55-60 % (x = 57,n = 10; MT = 57-58, n = 3) de la largeur du proglottis. Utérus pourvu de 4-8 (n = 20; MT = 4-7) diverticules utérins allongés latérale- ment (Fig. 3). Oncospheres de 19-21.5 (x = 20, n = 7) de diamétre, pourvues de crochets longs de 6.5-9 (n = 7.5; MT = 7.5-10, x = 9, n = 14); embryophore des oeufs mürs de forme ronde à ovoide (Fig. 2), d’un diamètre de 35-40 (x = 38, n = 7). Enveloppe externe hyaline, peu épaisse, très plissée et asymétrique. REMARQUES Fuhrmann (1934), révisant le matériel original de Diesing (1850), a observé un nombre de testicules plus élevé (120 a 160), contrairement à nos observations sur ce matériel type (100 a 136; x = 118), ce qui correspond aux observations concernant les spécimens récents (105-119, x = 112). Woodland (1933a) redécrit Amphoteromorphus peniculus. A notre avis, cette redescription est fondée sur deux especes mélangées. En raison du nombre figure de testicules et de la disposition des vitellogènes (Figs 48-51), seule la figure 50 corres- pond a A. peniculus. Les figures 48, 49, 51 montrent un nombre nettement plus faible de testicules et des vitellogenes en position plus latérales ce qui correspond a 386 M. CARFORA ET AL. Amphoteromorphus piriformis (cf. Fig. 19). Cette conclusion est confirmee par la forme particuliere de l’embryophore (Woodland, op.cit., Fig. 52) qui est exactement celle observée chez A. piriformis (Fig. 21). De plus, ces deux espèces vivent chez le méme höte. Les pores génitaux de A. peniculus sont irrégulièrement alternes. Cette carac- teristique est unique chez le genre Amphoteromorphus et differencie cette espèce de toutes les autres. Chez Amphoteromorphus, toutes les autres especes du genre ont les pores génitaux unilatéraux ou considérés comme tels (de Chambrier & Vaucher, 1997). A. peniculus est l’espèce du genre qui possède le plus grand nombre de testicules dans le genre Amphoteromorphus (Tableau 1). De plus, cette espèce est bien reconnaissable a la taille de ses ventouses (Fig. 1). Ses vitellogenes ventraux sont plus étendus en direction du milieu du proglottis (Tableau 1 et Figs 3, 4) comme le releve déja Fuhrmann (1934, p. 552). Enfin, il y a une importante concentration de cellules a cyto- plasme granuleux a l’apex du scolex. Amphoteromorphus parkamoo Woodland Figs 5-11 Amphoteromorphus parkamoo Woodland, 1935: p. 214. Hote: Paulicea luetkeni (Steindacher, 1877) (Pimelodidae); nom vernaculaire: Jat. Matériel examiné: Brésil, Etat de Amazonas, Itacoatiara, INVE 19733, 25.09.92; 19852, 19.09.92; INVE 19851, 16.09.92; INVE 22085, 03.10.95; INVE 22240*, 08.10.95; INVE 22241*, 08.10.95; INVE 22242*, 11.10.95; INVE 22243*, 11.10.95; INVE 22244*, 11.10.95. Localisation: Intestin, partie antérieure. DESCRIPTION Strobiles acraspédotes, jusqu’à 50 mm de long, avec de nombreux plis tégu- mentaires transversaux et longitudinaux. Proglottis larges de 935-1300 (x = 1100, n = 15) et longs de 215-485 (x = 345, n = 10). Scolex long de 690-1625 (x = 1275, n = 5), large de 1400-1960 (x = 1680, n = 6) et possédant une musculature longitudinale développée. Métascolex presque deux fois plus large que le cou (Fig. 5). Ventouses bilobées, longues de 430-665 (x = 520, n = 8) et pourvues d’une musculature radiaire puissante, avec une fine couche de mus- cles circulaires du côté interne (Fig. 6). Lobe antérieur de 220-360 de diamètre (x = 295, n = 7), légèrement plus large que le lobe postérieur: de 210-310 de diamètre (x = 270, n = 7). Rapport entre longueur des ventouses et longueur du scolex: 35-50 % (x = 40, n = 8) (Fig. 5). Musculature longitudinale interne constituée de faisceaux de fibres, avec concentration de ces faisceaux aux marges des proglottis (Figs 10, 11). Canaux osmorégulateurs ventraux chevauchant la poche du cirre dans son tiers terminal, de 10- 15 de diamètre (Fig. 7). Canaux dorsaux d’un diamètre de 5 à 10, à paroi plus épaisse que celle des ventraux et passant dorsalement à la poche du cirre en position subter- minale en général (Fig. 7). Testicules médullaires, ronds à ovoïdes disposés en deux couches, exception- nellement trois, au nombre de 34-48 (x = 41, n = 15), de 40-50 de diamètre, organisés en un champ avec une plus forte densité latéralement et atteignant les canaux osmo- régulateurs ventraux et les follicules vitellins, mais sans les dépasser (F1g. 7). Peu ou pas de testicules chevauchant la poche du cirre. Follicules vitellins corticaux et paramusculaires (Fig. 9) (dans 22 % des coupes), formant deux bandes dorso-latérales et deux bandes ventro-latérales. Rapport 387 LE GENRE AMPHOTEROMORPHUS 8-9 S'L-9 S'9-ç 8-59 APIOAO & 9puOI 2PIOAO Sunioyrud 9puoI c Lav Ore GAS Or-0E 59-55 59-55 S9-0S xneipjejun xneipjeplun xne.ipjejiun xneupjejlun 0€-07 SE-ST LEA Oe-ST 07-01 €C-81 vce-SI ST-8I 9Se]eo9p JUOLUIYONVAYI JUILIIYONVAWYI juoulayoneaayo (sadnoo sap 9,6L surp (sores san (sawı san (sadnoo sap 9,6] surp soureynosnuuezed) souejnosnuuted) sole nosnuied) sone nosnuresed) xNBON.1OI XNBIT}IOD xneoniod XNEIIJI09 CELI 58-55 CL-LV CL-6S SEI-00I CLI-VL OLT-S9I SIE-SLT 0S81-0S6 008I-ST6 SBEI-OEII SCyrI-066 OLTT-09EI OT£C-086 Sept O8CI OÿL£E-000€ 06£I-0E6 0081-078 SOLT-O€L STSTOICI Sp-0£ SS-OV Sp-0£ Sp-ST unsojuaun if “g 3 © SUDOIADf umsoouppf 19 HUD]JIDA “g ‘ds nwojskjpjddyov.ig DWOISCID]AKYyDAg puoskjojdcyovig “u ds 10u1u ‘y “u ds sypao y *u ‘ds smusofinnd -y — pqaanaid *Y 85 apuoı SE OL-SS xne.Jojejrun 0€-07 07-81 JUOLIIYONVADYI (sadnoo sap 9,77 suep sole nosnuresed) XNPEIIIOI 8P-re ssr-sIc 00EI-SE6 0961-0071 5791-069 Mayan] DIIYND_ ooupyand y OI-S'L = LN 6-59 OpuoI R IPIOAO Ly = LIN sr 8S-LS = LW 9-55 SOUIO}L ST-LI = LN 0E-ST TEST (sonsuoy snjd Juos SO[UIUOA SO] STRUT) JUOLUAYONEAYI (; some nosnuresed) XNPIIIOI 9€1-00I = LN 611-501 0601 59-09 DULOISAJD]AKYID.AG snynoiuad "V SUDOIADI SJ9490.5 Sap _ININTUO] s.ınu1 Sjnoo sap 3.10ydoA.ıquıa JULIO] suligjn SI[NIAIAIP ap IIJQUION (syyojdo.d ınad.ıe] % ) QJIBAO INITILT] XNJIU98 S2.104 (sto[$oid ına3.ıe] % ) («Id») P4419 np sysod Anonsuo] «Yo, Ud [8014 ınad.ıe] [neu “pra 3 10ddeY SQJB.19Je]-0S10p 19 SA[B19]E]-0.1] UA sapueq sap uoyisodsi( SUITHA SIMAON S$I[NIIS9} Ip 21QUION s1Joj80.1d np inonZuo] sıjojso.d np .ına3.e’] «ui, us X9[09S Np AN98.IU/T um, us X9[09sS Np ININZUOT] «Yo», UI X9]OIS NP ANINSUO]/SISNOJUIA sap Anonsuo] Jıoddey [73 (S)910H sa9ads4 ‘snydiowosajoyduy 31098 np so99dss xıs sop onbiBojoydiow uosmaedwo,) :] NVATAVI, 388 M. CARFORA ET AL. Fics 5-9 Amphoteromorphus parkamoo Woodland, 1935. 5. 22085 INVE, scolex très plissé. 6. 27989 INVE, coupe sagittale d’un scolex montrant ses ventouses bilobées (flèche). 7. 22085 INVE, vue ventrale d’un proglottis mature. L’utérus n’est pas figuré. 8. Oeuf monté dans l’eau distillée. 9. 27989 INVE, coupe transversale partielle d’un proglottis gravide montrant la pénétration de quelques vitellogènes dans la médulla. Echelles: 5 = 300 mm; 6 = 100 mm; 7, 9 = 200 mm; 8 = 25 mm. entre largeur des vitellogènes et largeur du proglottis: ventraux, 18-20 % (x = 19, n= 15); dorsaux, 16-19 % (x = 17, n = 15). Rapport entre longueur des vitellogènes et longueur du proglottis du còté poral: ventraux, 55-65 % (x = 60, n = 10); dorsaux, 54- LE GENRE AMPHOTEROMORPHUS 389 © Een a Fics 10-11 Amphoteromorphus parkamoo Woodland, 1935. 10. 27989 INVE, coupe transversale d’un proglottis prégravide, montrant la disposition de l’utérus et de l’ovaire. Glande de Mehlis bien visible. 11. 27989 INVE, coupe transversale d’un proglottis gravide, au niveau de l’utérus. Tégument tres plisse. Musculature longitudinale interne plus dense aux marges du proglottis. Echelles: 10, 11 = 200 mm. 70 % (x = 56, n = 10); du côté aporal: ventraux, 76-87 % (x = 81, n = 10); dorsaux, 68- 79 % (x = 74,n = 10). Peu ou pas de follicules vitellins dorsalement et ventralement a la poche du cirre (Fig. 7). Poche du cirre allongée, a paroi mince, légèrement plus large dans sa partie po- rale, longue de 220-345 (x = 285, n = 11), PC = 20-30 % (x = 26, n = 20). Cirre occu- pant environ 60 % (n = 7) de la longueur de la poche du cirre (Fig. 7). Pores génitaux unilatéraux, PG = 16-21% (x = 18, n = 9). Vagin toujours antérieur à la poche du cirre (n= 65), avec un sphincter musculaire (Fig. 7). Canal éjaculateur très contourné. Canal déférent très contourné et dépassant le milieu du proglottis (Fig. 7). Ovaire compact a contour irrégulier, large de 705-1415 (x = 1115, n = 10) et oc- cupant les 55-70 % (x = 63, n = 10) du proglottis (Fig. 7). Uterus occupant jusqu’a 70 % de la largeur des proglottis prégravides, avec 3- 5 (n = 18) diverticules utérins peu allongés. 390 M. CARFORA ET AL. Oncosphères de 15-16 (x = 15, n = 9) de diamètre, pourvues de crochets longs de 5-8 (x = 7, n= 17); embryophore des oeufs mûrs de forme ronde, divisée en 2 couch- es. Diametre de la couche externe de 25-34 (x = 29, n = 9). Enveloppe externe hyaline, peu épaisse, très plissée, asymétrique et longue de 48-80 (x = 57,n=9) (Fig. 8). ANALYSE GENETIQUE (Tableau 2 et Fig. 37) Sur les 5 échantillons étudiés, six systemes enzymatiques ont fourni des zymo- grammes dont les mobilités électrophorétiques sont clairement interprétables. Seul l’échantillon INVE 22243 a démontré une faible activité au locus ALD, avec un seul phénotype enzymatique. Aux loci AAT, ACP, AK, ALD, MDH et PGM, les zymo- grammes présentent un seul phénotype enzymatique. Deux allèles existent au locus PGI. A. parkamoo se distingue sur la base des izoenzymes de A. ovalis et A. ninoi, mais pas de A. piraeeba ni de A. piriformis. REMARQUE A. parkamoo se différencie de A. peniculus par ses pores génitaux unilatéraux. En revanche, il est proche de A. piraeeba. Amphoteromorphus piraeeba Woodland Figs 12-16 Amphoteromorphus piraeeba Woodland, 1934: 274. Amphoteromorphus peniculus sensu Woodland, 1933: 189. Amphoteromorphus piraeeba; de Chambrier & Vaucher, 1997: 231. Hote: Brachyplatystoma filamentosum (Lichtenstein) (Pimelodidae); nom vernaculaire: Piraiba. Matériel examiné: Brésil, Etat de Amazonas, Itacoatiara, INVE 19659, 15.09.92: INVE 19660, 17.09.92; INVE 19661, 25.09.92; INVE 22069, 21.10.95; INVE 22070, 15.10.95; 02.10.95: INVE 22221; INVE 22223*; INVE 22225*; INVE 22227; INVE 22228; INVE 22232; INVE 22234; INVE 22235; INVE 22236; INVE 22237; INVE 22239*; 13.10.92: INVE 22242: 16.10.92: INVE 22251*; INVE 22252; INVE 22253: 20.10.95. Localisation: Intestin. DESCRIPTION Strobiles acraspédotes, jusqu’a 65 mm de long. Tégument plisse. Proglottis matures et gravides larges de 990-1425 (x = 1120, n = 22) et longs de 275-365 (x = 330 sni= 110): Scolex long de 1210-2525 (x = 1780, n = 8) et large de 2000-3740 (x = 2920, n = 8). Métascolex plissé et environ deux fois plus large que le cou (Fig. 12). Ventouses bilobées, pourvues d’une musculature radiaire puissante, longues de 495-650 (x = 560, n = 9). Lobe antérieur de 290-400 de diamètre (x = 355, n = 12), légèrement plus large que le lobe postérieur: 260-390 de diamétre (x = 330, n = 9). Rapport entre longueur des ventouses et longueur du scolex: 25-45 % (x = 35, n = 10) (Fig. 12). Présence de cellules à cytoplasme granuleux en faible quantité, toujours situées à l’apex du scolex (Fig. 12). Musculature longitudinale interne constituée de faisceaux de fibres, avec concentration de ces faisceaux aux marges des proglottis (Figs. 14, 16). Canaux osmorégulateurs ventraux de 10-20 de diamétre, chevauchant la poche du cirre dans sa partie subterminale. Canaux osmorégulateurs dorsaux de 8-12 de diamétre, a paroi plus LE GENRE AMPHOTEROMORPHUS 391 Fics 12-13 Amphoteromorphus piraeeba Woodland, 1934. 12. 22249 INVE, scolex; 13. 19659 INVE, vue ventrale d’un proglottis mature. Echelles: 12 = 500 mm; 13 = 200 mm. épaisse que celle des ventraux et croisant également la poche du cirre dans sa partie subterminale (Fig. 13). Testicules médullaires, ronds à ovoides, disposés en deux ou trois couches, au nombre de 59-72 (x = 66, n = 15), de 60-100 de diamétre et organisés en un champ dans les proglottis prématures. Disposition en deux champs dans les proglottis prégra- vides et gravides, atteignant parfois les canaux osmorégulateurs dorsaux mais jamais les canaux ventraux (Fig. 13). Vitellogenes corticaux et paramusculaires (dans 15 % des coupes), formant deux bandes dorso-latérales et chevauchant les deux ventro-latérales. Follicules vitellins dorsaux et ventraux atteignant les canaux dorsaux et en général les canaux ventraux. Rapport entre largeur des vitellogénes et largeur du proglottis, ventraux: 392 M. CARFORA ET AL. Fics 14-16 Amphoteromorphus piraeeba Woodland, 1934. 14. 19659 INVE, coupe transversale montrant la position dorso-ventrale des vitellogènes. 15. 21983 INVE, oeuf monté dans l’eau distillee; 16. 19659 INVE, coupe transversale partielle d’un proglottis mature montrant la pénétration d’un vitellogene dans la musculature longitudinale interne. Echelles: 14 = 500 mm; 15 = 50 mm; 16 = 100 mm. 18-25 % (x = 21, n = 15); dorsaux: 17-26 % (x = 20, n = 15). Rapport entre longueur des vitellogènes et longueur du proglottis, du còté poral: ventraux, 55-80 % (x = 65, n = 10); dorsaux, 53-78 % (x = 64, n = 10); du côté aporal: ventraux, 85-93 % (x = 89, n = 10); dorsaux, 81-90 % (x = 88, n= 10). Poche du cirre allongée, à paroi mince, longue de 230-390 (x = 275, n= 11), PC = 25-30 % (x = 27, n= 11). Cirre occupant jusqu’à environ 60 % de la longueur de la poche du cirre (Fig. 12). Pores génitaux unilatéraux, PG = 12-18 (x = 15,n=9). Vagin toujours antérieur à la poche du cirre (x = 60), avec un sphincter musculaire. Canal éjaculateur très contourné (Fig. 13). Canal déférent également très contourné et atteignant le milieu du proglottis. Ovaire compact, à contour irrégulier, large de 625-860 (x = 735, n= 11), occu- pant les 50-65 % (x = 60, n = 10) de la largeur du proglottis. Utérus pourvu de 3-6 (n = 25) diverticules utérins allongés et minces. Oncosphères de 14-17 (x = 15.5, n = 7), pourvues de crochets longs de 6.5-8 (x = 7, n = 6) (Fig. 15); embryophore des oeufs mürs de forme ronde, divisée en 2 LE GENRE AMPHOTEROMORPHUS 393 couches. Diamètre de la couche externe de 29-32 (x = 30, n = 4). Enveloppe externe hyaline, peu épaisse, très plissée, asymetrique et longue de 54-74 (x = 60, n = 4) (Fig. 15). ANALYSE GENETIQUE (Tableau 2 et Fig. 37) Sur les 3 échantillons étudiés, les zymogrammes présentent un phénotype enzymatique unique aux loci AAT, ACP, AK, MDH et PGI. ALD exprime deux alleles dans un cas. Un exemplaire présente une deuxième tache de très faible activité au locus PGM, que nous n’avons pas prise en considération. Les loci NP et HK ne semblent pas étre exprimés. Cependant, au locus HK, quelques taches parasites apparaissent qui expriment peut-étre l’activité d’une autre enzyme (GCDH) (cf. Hillis er al., 1996). A. piraeeba se distingue sur la base des izo- enzymes de A. ovalis et A. ninoi mais pas de A. parkamoo, ni de A. piriformis. REMARQUES Certaines de nos observations different de ceux de Woodland (1934, p. 274) qui décrit chez A. piraeeba un nombre de testicules par proglottis de trois a quatre fois supérieur: 200 au lieu de 59 a 72. Woodland (1934, p. 274) observe une position postérieure ou antérieure du vagin. Nous constatons que ce dernier débouche toujours antérieurement a la poche du cirre. A. piraeeba et A. parkamoo sont deux espéces voisines morphologiquement et anatomiquement. Plusieurs criteres toutefois permettent de les différencier (Tableau 1). Le scolex de A. piraeeba est plus large, plus long, ses testicules sont plus nombreux (59 a 72 contre 34 a 48 chez A. parkamoo) et d’un diametre supérieur, son sphincter musculaire, entourant le vagin, parait moins développé. La figure 16 montre un décalage des vitellogènes ventraux et dorsaux, mais ne constitue pas un caractere taxonomique. Amphoteromorphus piriformis sp. n. Eies17-21,35336 Amphoteromorphus piraeeba Woodland, 1934: 274. Amphoteromorphus peniculus sensu Woodland, 1933: 189. Amphoteromorphus piraeeba; de Chambrier & Vaucher, 1997: 231. Amphoteromorphus peniculus; Rego, Chubb & Pavanelli, 1999: 251. Hote: Brachyplatystoma flavicans (Lichtenstein) (Pimelodidae); nom vernaculaire: Dourada. Matériel examiné: Etude morpho-anatomique: Brésil, Etat de Amazonas, Itacoatiara, holotype INVE 32923, 3 paratypes INVE 32922, 32924-25, 13.09.92; INVE 19310, 16.09.92; INVE 19311, 13:09°92: INVE 19312; 1'7-09:92: INVE 19314, 21.09:92. INVE 1932071709192: INVE#19326 17209592; INVE= 19624, 21509 92-MINVE 196312, 21.0992 Z21NVE2221985 30.09.95; INVE 22199*, 30.09.95; INVE 22200*, 30.09.95; INVE 22208, INVE 22209*, INVE 222106 /INVE 222117, 21.09.92; INVE 22229* 07.10.95; INVE 27432, 06.10.95. Localisation: Intestin antérieur. DESCRIPTION Strobiles acraspédotes, jusqu’a 50 mm de long. Tégument très plissé. Proglottis matures et gravides larges de 1130-1385 (x = 1240, n = 20) et longs de 165-270 (x = DS AS); Scolex long de 730-1705 (x = 1190, n = 9), large de 1380-2435 (x = 1905, n = 9) et possédant une musculature longitudinale bien développée. Métascolex plissé et M. CARFORA ET AL. 394 qINDd e1Dd = OE CHAN - BMH S'AUMNNd "10d CHAN CAN AH PANDA Add oq EHAMN = n CINDd qd èP NO HAN x i Q'ENDd 9%Dd HAN 5 7 PINDd 9 NO CID qHdUW dN MH WDd [Dd HOW IAA'TV ea IV ea IV q no eq TV ea IV CIV MV MV PV MV MV MV MV edOV EIVV ono gno PdIV BIVV WdOV °IVV QIV AIVV AdDV ALVV GdDV ALVV dOV LVV LCL A | wunsojuawuppf'g ‘ds pwossäpydäyonag SUDIIAD], gl LUNSOJUIUD]Y, °F 1uoyJon] DIIND] SOJOH IOUNU * 1OULU * sıDao ' snutofind * pgood.nd ° oowpy.nd' SSE) SE} TSE, SE, FREE So9ods4 sonbrewAzus SOUIQJSAS JNoU op SOLULUPISOWAT, I] AVATAV I, LE GENRE AMPHOTEROMORPHUS 395 Fics 17-19 Amphoteromorphus piriformis sp. n. 17. 19310 INVE, scolex; 18. Paratype 32922 INVE, coupe sagittale d’un scolex; 19. holotype 32923 INVE, vue ventrale d’un proglottis mature. Echelles: 17 = 600 mm; 18 = 300 mm; 19 = 250 mm. environ deux fois plus large que le cou (Fig. 35). Ventouses bilobées (Figs 17, 18, 36), longues de 340-750 (x = 510, n= 11) et pourvues d’une musculature radiaire puissante, avec une fine couche de muscles circulaires du côté interne (Fig. 18). Lobe antérieur de 195-405 (x = 300, n = 10) étant légèrement plus large que le lobe postérieur: 185- 335 (x = 270, n = 10). Rapport entre longueur des ventouses et longueur du scolex: 30- 45% (x = 35, n = 10) (Fig. 17). Musculature longitudinale interne constituée de fais- ceaux de fibres, avec concentration de ces faisceaux aux marges des proglottis (Fig. 20). Canaux osmorégulateurs ventraux chevauchant la poche du cirre dans son tiers ter- minal, de 25-45 de diamètre. Canaux dorsaux d’un diamètre de 10-20, à paroi plus épaisse et passant également sous la poche dans sa partie terminale (Fig. 19). 396 M. CARFORA ET AL. Fics 20-21 Amphoteromorphus piriformis sp. n. 20. 19310 INVE, coupe transversale d’un proglottis mature; 21. 19311 INVE, œufs montés dans l’eau distillée. Echelles: 20 = 200 mm; 21 = 50 mm. Testicules médullaires, ronds a ovoides, disposés en deux couches, parfois trois, au nombre de 47-72 (x = 56, n = 15), de 40-60 de diamétre et organisés en un champ avec une plus forte densité latéralement, n’atteignant, en général, pas les canaux LE GENRE AMPHOTEROMORPHUS 397 osmorégulateurs dorsaux, mais parfois les canaux ventraux (Fig. 19). Testicules presents dorsalement a la poche du cirre. Follicules vitellins corticaux, parfois paramusculaires, formant deux bandes dorso-latérales et deux ventro-latérales (Fig. 20). Rapport entre largeur des vitellogenes et largeur du proglottis: ventraux = 15-24 % (x = 17, n = 32); dorsaux = 18-23 % (x = 18, n = 15). Rapport entre longueur des vitellogènes et longueur du proglottis, du côté poral: ventraux, 56-66 % (x = 60, n = 10); dorsaux, 50-70 % (x = 57, n = 10); du côté aporal: ventraux, 73-85 % (x = 78, n = 10); dorsaux, 59-83 % (x = 72, n= 10). Peu ou pas de follicules vitellins dorsalement et ventralement à la poche de cirre (Fig. 19). Poche du cirre allongée, à paroi mince, longue de 200-325 (x = 285, n= 14), PC = 17-27 % (x = 23, n= 15). Cirre occupant environ 60 % de la longueur de la poche du cirre (Fig. 19). Pores génitaux unilatéraux, PG = 13-23 % (x = 17, n = 8). Vagin toujours antérieur a la poche du cirre (n = 75), avec un sphincter musculaire sub- terminal (Fig. 19). Canal éjaculateur très contourné. Canal déférent également très contourné et atteignant le milieu du proglottis (Fig. 19). Ovaire compact a contour irrégulier, parfois bilobé, large de 670-900 (x = 750, n= 10) et occupant les 55-65 % du proglottis (x = 60, n = 10). Utérus pourvu de 3-6 diverticules utérins (n = 20) (Fig. 19), contenant de nombreux oeufs avec une faible proportion d’oeufs mürs. Oncosphères de 12-14.5 (x = 13.5, n = 9) de diamètre, pourvues de crochets longs de 5-6.5 (x = 6, n = 13); embryophore des oeufs mürs de forme piriforme (les embryophores des oeufs non mûrs sont de formes rondes), long de 28-33 (x = 30, n = 7) et large de 21-30 (x = 24.5, n = 7) (Fig. 20). Enveloppe externe hyaline, peu épaisse, très plissée, asymétrique et longue de 40-64 (x = 47.5, n = 8) (Fig. 21). ANALYSE GENETIQUE (Tableau 2 et Fig. 37) Sept systemes enzymatiques ont fourni des zymogrammes présentant des mo- bilités électrophorétiques clairement interprétables. Aux loci AAT, ACP, AK, ALD, PGI, PGM, les zymogrammes présentent un seul phénotype enzymatique. MDH présente deux alléles. Les loci HK et NP ne semblent pas exprimes. A. piriformis se distingue sur la base des izoenzymes de A. ovalis et A. ninoi mais pas de A. parkamoo, ni de A. piraeeba. REMARQUES A. piriformis est placé dans le genre Amphoteromorphus en raison de la présence d’un métascolex, d’un scolex pourvu de quatre ventouses bilobées, d’une absence d’organe apical, de proglottis acraspédotes, de vitellogenes principalement corticaux et de pores génitaux unilatéraux (avec quelques alternances irrégulières possibles). A. piriformis présente des embryophores piriformes, qui le différencie de toutes les autres espèces (Fig. 21). Parmi elles, A. piraeeba est la plus proche. Les résultats du présent travail conduisent à revoir les conclusions de de Chambrier & Vaucher (1997): l’oeuf figuré (Fig. 15) n’appartient pas à A. piraeeba mais bien à A. piriformis qui n’avait pas été séparé spécifiquement. Les auteurs figurent 398 M. CARFORA ET AL. deux dessins (Figs 13 et 15) qui ne correspondent pas a A. piraeeba, mais a A. piri- formis. En outre, p. 229 (op. cit.), les hôtes dans la Table 1 sont intervertis : l’hôte de A. peniculus Diesing, 1850 et de A. peniculus sensu Woodland (1933a) étant B flavi- cans, celui de A. piraeeba Woodland, 1934 B. filamentosum. En plus des oeufs, certains caracteres discrets permettent de différencier A. pir- iformis de A. piraeeba (Tableau 1): le scolex de A. piriformis est plus petit, ses ven- touses sont legerement moins longues (510 au lieu de 560 en moyenne), ses proglottis sont plus larges (1240 au lieu de 1120 en moyenne), mais moins longs (225 au lieu de 330 en moyenne), ses testicules ont un diamètre moins important (40-60 au lieu de 60-100) et sont moins nombreux en moyenne (56 au lieu de 66) et le rapport PC est un peu moins élevé (17-27 % au lieu de 25-30%). En conséquence, nous sommes en présence d’une nouvelle espece que nous appelons A. piriformis en référence a la forme particuliere de ses embryophores. L’électrophorèse des protéines ne nous permet pas de distinguer ces deux especes. Amphoteromorphus ovalis sp. n. Figs 22-27 Hote: Brachyplatystoma sp. (Pimelodidae); nom vernaculaire: filhote da capa preta. Materiel examine: Bresil, Etat de Amazonas, Itacoatiara 13.10.95: holotype INVE 22247, 2 paratypes INVE 22249-50, INVE 21742, INVE 22247, INVE 22248, INVE 22249*, INVE 22250. Localisation: Intestin, dans le sixième antérieur. DESCRIPTION Strobiles acraspédotes, avec de nombreux plis tégumentaires transversaux et longitudinaux aux vers. Proglottis larges de 916-1800 (x = 1525, n = 27) et longs de TI Bun 29) Scolex long de 820-1800 (x = 1345, n = 7), large de 980-2310 (x = 1925, n = 7) et possédant une musculature longitudinale développée. Ventouses bilobées, longues de 525-745 (x = 650, n = 14) et pourvues d’une musculature radiaire puissante (Fig. 22). Lobe antérieur de 305-500 de diamètre (x = 420, n = 14), légèrement plus large que le lobe postérieur: de 245-425 de diamètre (x = 360, n = 14). Rapport entre longueur ventouses et longueur du scolex: 40-55 % (x = 45, n = 8) (Fig. 22). Musculature longitudinale interne constituée de faisceaux de fibres, avec concentration de ses faisceaux aux marges des proglottis (Fig. 27). Canaux osmorégulateurs ventraux chevauchant la poche du cirre dans son tiers terminal, de 100-300 de diamètre. Canaux dorsaux d’un diamètre de 7 à 10, à paroi plus épaisse que celle des ventraux et tra- versant également la poche du cirre dans son tiers terminal (Fig. 26). Testicules médullaires, ronds à ovoîdes disposés en deux couches, parfois trois, au nombre de 55-85 (x = 71, n = 22), de 30-50 de diamètre, organisés en un champ, deux dans les proglottis gravides (Figs 23, 26). Follicules vitellins principalement corticaux, formant deux bandes dorso- latérales et deux bandes ventro-latérales (Fig. 27). Rapport entre largeur des vitello- gènes et largeur du proglottis: ventraux: 18-23 % (x = 20, n = 16); dorsaux: 19-26 % (x = 23, n= 17). Rapport entre longueur des vitellogènes et longueur du proglottis: du còté poral: ventraux, 56-79 % (x = 68, n = 20): dorsaux, 60-80 % (x = 72, n = 20); du côté aporal: ventraux, 78-91 % (x = 83, n = 20); dorsaux, 76-87 % (x = 80, n= 15). LE GENRE AMPHOTEROMORPHUS 399 Fics 22-25 Amphoteromorphus ovalis sp. n. 22. Paratype 22249 INVE, scolex. 23. Holotype 22247 INVE, vue dorsale d’un proglottis mature. 24. Holotype 22247 INVE, detail de l’utérus montrant ses di- verticules ramifiés, ainsi que la présence de cellules chromophiles à leur extrémité. 25. 21742 INVE, ceufs montés dans l’eau distillée. Echelles: 22 = 500 mm; 23, 24 = 250 mm; 25 = 50 mm. 400 M. CARFORA ET AL. Poche du cirre allongée, a paroi mince (Fig. 26). Rapport entre longueur de la poche du cirre et largeur du proglottis, PC = 25-35 % (x = 28, n = 22). Cirre allongé, très contourné, à paroi mince et occupant jusqu’a 70% (n = 22) de la longueur de la PC (Fig. 26). Pores génitaux unilatéraux. Vagin toujours antérieur à la poche du cirre (n = 75), pourvu d’un sphincter musculaire bien développé. Canal éjaculateur très con- tourné (Fig. 26). Canal déférent très contourné également et dépassant le milieu du proglottis. Ovaire compact à contour irrégulier occupant les 55-65 % (x = 59, n = 28) du proglottis (Fig. 23). Utérus pourvu de 4-7 (n = 20) diverticules utérins (Fig. 24). Oncosphères de 13-16 (x = 14.5, n = 7) de diamètre, pourvues de crochets longs de 6-7.5 (n= 12); embryophore des oeufs mûrs de forme ovoide , long de 32-37 (x = 33.5, n = 8) et large de 25-27 (x = 26, n = 8) (Fig. 20). Enveloppe externe hyaline, peu épaisse, très plissée, asymétrique et longue de 65-70 (ou plus) (Fig. 25). ANALYSE GENETIQUE (Tableau 2 et Fig. 37) Un seul échantillon a été analysé et ses mobilités électrophorétiques sont inter- prétables dans huit systemes enzymatiques: AAT, ACP, AK, MDH, PGI, PGM, HK et NP. Cette nouvelle espèce possède des mobilités électrophorétiques totalement dif- férentes aux loci AAT et AK. Cette espèce est également la seule à afficher trois phéno- types enzymatiques au locus MDH. Au locus HK, Amphoteromorphus ovalis se dis- tingue de A. ninoi seulement, le locus n’étant pas exprimé dans les autres échantillons. Au locus PGM, A. ovalis présente un phénotype enzymatique qui n’a pas d’équivalent dans les autres échantillons. Il est le seul également à présenter une réaction au locus NP. Ces résultats suggèrent que A. ovalis est un taxon distinct. REMARQUES Amphoteromorphus ovalis se distingue de A. peniculus et de A. parkamoo par le nombre de testicules (Tableau 1) et de A. piriformis par la forme ovoide de ses embryophores. A. ovalis est très proche de A. piraeeba. Certains caracteres permettent tout de méme de les différencier (Tableau 1): chez A. ovalis, le scolex est nettement plus petit, les proglottis sont notablement plus larges et moins longs. Ses ventouses sont plus longues et ses testicules, d’un diamètre inférieur, sont plus nombreux. La forme de ses embryophores est ovoide, alors qu’ils sont ronds chez A. piraeeba. Chez A. ovalis, le sphincter vaginal est plus développé. Par ailleurs, le canal éjaculateur et le canal déférent sont plus contournés que dans les autres espèces. Sur la base de l’analyse génétique par électrophorèse des protéines, A. ovalis se differencie des autres espèces par trois systemes enzymatiques. En conséquence, A. ovalis représente une nouvelle espèce qui est nommée ovalis en raison de la forme de ses embryophores. L’identification de l’hòte de A. ovalis pose problème. John G. Lundberg (Philadelphia) (communication personnelle) déduit de plusieurs photographies qu’il ne s’agit pas de Brachyplatystoma filamentosum (‘‘piraìba’’), mais d’un poisson vraisem- blablement distinct appelé 4 Belem “ filhote da capa preta ”. LE GENRE AMPHOTEROMORPHUS 401 27 + TA Fics 26-27 Amphoteromorphus ovalis sp. n. 26. Holotype 22247 INVE, vue dorsale du vagin et de la poche du cirre. 27. 21742 INVE, coupe transversale partielle d’un proglottis mature. Echelles: 26 = 100 mm; 27 = 250 mm. 402 M. CARFORA ET AL. Amphoteromorphus ninoi sp. n. Figs 28-34 Hôte-type: Brachyplatystoma vaillanti (Valenciennes, 1840) (Pimelodidae); nom ver- naculaire: Piramutaba. Autre höte: B. filamentosum (Lichtenstein); nom vernaculaire: Piraiba. Matériel examine: Etude morpho-anatomique: Brésil, Etat de Amazonas, Itacoatiara, de Brachyplatystoma vaillanti, 02.10.92: holotype INVE 22203, 3 paratypes INVE 32927-28, 32859; INVE 22201; INVE 22202; INVE 22204; INVE 22205; INVE 22206*; INVE 22207*. De Brachyplatystoma filamentosum, 02.10.95: INVE 22232, INVE 22234, INVE 22235*, INVE DDDSCÉAINVNVE 22237 7INVE222398. Localisation: Intestin, partie antérieure. DESCRIPTION Strobiles acraspédotes, jusqu’a 35 mm de long. Tégument peu ou pas plisse. Proglottis larges de 950-1850 (x = 1340, n = 50) et longs de 100-135 (x = 115,n = 15). Scolex long de 930-1390 (x = 1110, n = 10), large de 1360-2270 (x = 1950, n = 10) et possédant une musculature longitudinale tres développée. Métascolex plissé, une fois et demi a deux fois plus large que le cou. Ventouses bilobées (Figs 28, 29), longues de 305-435 (x = 375, n = 15), pourvues d’une musculature radiaire puissante et d’une fine couche de muscles circulaires du còté interne (Fig. 29). Lobe antérieur de 195-400 de diamètre (x = 290, n = 15), légèrement plus large que le lobe postérieur: 160-270 de diamètre (x = 220, n = 11). Rapport entre longueur des ventouses et longueur du scolex: 30-45 % (x = 35, n = 10) (Fig. 28). Présence de cellules a cyto- plasme granuleux situées à l’apex du scolex (Fig. 28). Musculature longitudinale in- terne développée et constituée de faisceaux denses (Figs 32-34). Concentration de ces faisceaux aux marges des proglottis. Canaux osmorégulateurs ventraux chevauchant la poche du cirre dans sa partie médiane, de 25-35 de diamètre (Fig. 30). Canaux dorsaux de 15-20 de diamètre, à paroi plus épaisse que celle des ventraux, chevauchant la poche dans son tiers proximal (Fig. 30). Testicules médullaires, ronds à ovoïdes, disposés en deux couches, exception- nellement trois, au nombre de 17-32 (x = 24, n = 40), organisés en un champ, avec une plus forte densité latéralement, de 40-65 de diametre, atteignant et dépassant parfois les canaux osmorégulateurs ventraux. Testicules dépassant parfois les vitellogenes ventraux, mais jamais les dorsaux (n = 30) et présents dorsalement a la poche du cirre (Fig. 30). Follicules vitellins corticaux, paramusculaires dans 75 % des coupes ou rare- ment médullaires (Figs 32-34). Follicules vitellins formant deux bandes dorso- latérales, décalées par rapport aux deux autres bandes ventro-latérales (Figs 30,32). Rapport entre largeur des vitellogènes et largeur du proglottis: ventraux, 10-20 % (x = 15.5, n = 20); dorsaux, 10-17 % (x = 13, n = 20). Rapport entre longueur des vitel- logènes et longueur du proglottis du côté poral: ventraux, 35-45 % (x = 40, n = 10); dorsaux, 35-46 % (x = 41, n= 10); du côté aporal: ventraux, 40-65 % (x = 45, n = 10); dorsaux, 39-63 % (x = 49, n = 10). Vitellogènes dorsaux plus distaux que les ventraux, n’atteignant pas en général l’ovaire. Follicules vitellins ventraux chevauchant l’extremite de l’ovaire. Peu de vitellogènes dorsalement et ventralement a la poche du cirre (Fig. 30). Poche du cirre allongée, à paroi mince, longue de 230-450 (x = 360, n = 11), PC = 20-30 % (x = 26, n = 25). Cirre très allongé, très mince et occupant entre 60-70 % LE GENRE AMPHOTEROMORPHUS 403 FIGs 28-29 Amphoteromorphus ninoi sp. n. 28. 22202 INVE, scolex avec une importante concentration de cellules à cytoplasme granuleux à l’apex. 29. Paratype 32859 INVE, coupe sagittale d’un scolex. Remarquer l’importante musculature longitudinale interne. Echelles: 28 = 300 mm; 29 = 500 mm. de la longueur de la poche de cirre (Fig. 30). Pores génitaux unilatéraux. Vagin toujours antérieur à la poche du cirre (n = 70), pourvu d’un sphincter musculaire discret (Fig. 30). Canal éjaculateur relativement peu contourné. Canal déférent tres contourné et dépassant le milieu du proglottis (Fig. 30). Ovaire compact, parfois légèrement bilobé, a contour irrégulier, large de 360-775 (x = 440, n = 33) et occupant les 30-40 % (x = 35, n = 31) de la largeur du proglottis (Fig. 30). Uterus pourvu d’un seul diverticule utérin de chaque côté (Fig. 30). Oncosphères de 14-16 de diamétre (x = 15, n = 9), pourvues de crochets longs de 6-8 (x = 7.5, n = 9); embryophore des oeufs mûrs de forme ronde à ovoide, divisée en deux couches. Diametre de la couche externe de 28-35 (x = 31, n = 9). Enveloppe externe hyaline, peu épaisse, peu plissée et asymétrique, longue de 49-55 (x = 50, MI) (er 3): ANALYSE GENETIQUE (Tableau 2 et Fig. 37) Huit systemes enzymatiques sont utilisables, certains difficiles a interpreter. Les six échantillons analysés présentent un seul phénotype enzymatique aux loci AAT, AK, MDH et PGI et deux aux loci ALD et HK. Dans les deux derniers cas, l’un est present chez les individus parasitant B. filamentosum et l’autre chez ceux qui parasitent B. vail- lanti. A. ninoi présente trois phénotypes enzymatiques au locus ACP ; les trois sont présents chez les parasites de B. filamentosum, un seul au contraire est exprimé chez les parasites provenant de B. vaillanti. Au locus PGM, trois phénotypes enzymatiques existent également, les parasites de B. filamentosum en affichent deux, ceux de B. vail- lanti un seul. Tous les échantillons de A. ninoi sont distincts des autres exemplaires analysés au loci AAT, MDH et PGI. 404 M. CARFORA ET AL. Fics 30-32 Amphoteromorphus ninoi sp. n. 30. Holotype 22203 INVE, vue dorsale de proglottis prégra- vides, montrant la disposition décalée des vitellogènes ventraux par rapport aux dorsaux; canal déférent tres contourné et dépassant le milieu du proglottis; glande de Mehlis et réceptacle sémi- nal bien visible; cirre évaginé. 31. 22202 INVE, oeufs montés dans l’eau distillée. 32. Paratype 32928 INVE, coupe transversale partielle d’un proglottis mature montrant la disposition des vitellogènes corticaux et paramusculaires. Tégument peu plissé. Echelles: 30 = 500 mm; 31 = 25 mm; 32 = 100 mm. En conclusion, l’analyse électrophorétique confirme la séparation spécifique de A ninoi. Les résultats aux loci ACP, ALD et PGM montrent des indices d’une Eventuelle séparation plus fine des parasites provenant de B. vaillanti et de B. filamen- tosum. Vu le faible nombre de spécimens analysés et en l’absence de critères morphologiques corrélés, nous ne pouvons pas en tirer des conclusions d’ordre taxonomique. REMARQUES Amphoteromorphus ninoi est nettement différent des autres espèces du genre (Tableau 1). Les testicules sont moins nombreux (Tableau 1). On remarque un décalage constant dans la disposition de ses bandes vitellines ventrales par rapport aux dorsales. LE GENRE AMPHOTEROMORPHUS 405 Fics 33-34 Amphoteromorphus ninoi sp. n. 33. 32860 INVE, coupe transversale au niveau du canal déférent d’un proglottis mature, montrant la musculature longitudinale interne (les faisceaux sont davan- tage concentrés aux marges du proglottis). 34. 22202 INVE, coupe transversale d’un proglottis mature. Noter les vitellogenes paramusculaires. Echelles: 33, 34 = 200 mm. Le rapport entre largeur de l’ovaire et largeur des proglottis est plus faible que chez les autres espèces. Il n’existe qu’un diverticule utérin de chaque côté. Comparativement aux autres especes, les proglottis sont proportionnellement plus larges et moins longs. D’autres caracteres discrets concernent le tégument presque lisse et les nombreux follicules vitellins paramusculaires. En conclusion, nous considérons cette espece comme nouvelle et l’appelons A. ninoi; elle est dédiée à Monsieur Aniello Carfora (“Nino”), pere du premier auteur. 406 M. CARFORA ET AL. Po Fics 35-36 Amphoteromorphus piriformis sp. n. Photographies au microscope électronique à balayage (MEB). 35. 19310 INVE, scolex. Noter le métascolex plissé. 36. Agrandissement d’une ventouse bilobee. Echelles: 35 = 200 mm; 36 = 50 mm. DISCUSSION GENERALE Le genre Amphoteromorphus se compose donc de six espèces, toutes parasites de pois- sons-chats néotropicaux: Amphoteromorphus peniculus Diesing, 1850, höte-type: Brachyplatystoma flavicans, A. piraeeba Woodland, 1934, höte-type: B. filamentosum, A. parkamoo Woodland, 1935, höte-type: Paulicea luetkeni, A. piriformis sp. n., höte-type: B. flavicans, A. ovalis sp. n., höte-type: Brachyplatystoma sp. et A. ninoi sp. n., hôte-type: B. vaillanti, autre hôte: B. filamentosum. Cinq espèces du genre Amphoteromorphus présentent une spécificité oioxène, alors que A. ninoi parasite deux espèces hôtes (spécificité sténoxène). Deux espèces d’hötes hébergent chacune deux espèces de parasites: B. filamentosum est parasité par A. piraeeba et par A. ninoi, B. flavicans par A. peniculus et par A. piriformis Les six espéces présentent un fort degré de ressemblance morphologique. A part A. peniculus, qui peut étre séparé par l’alternance des pores génitaux, les cing autres espèces se différencient par des caractères plus discrets. L’analyse de huit izoenzymes interpretables n’a pas apporté une réponse décisive dans tous les cas. A. parkamoo, A. piraeeba et A. piriformis présentent des mobilités électrophorétiques semblables pour les systemes étudiés (Tableau 2 et Fig. 37). Méme si le nombre d’échantillons examinés par électrophorése est peu élevé, cette approche biosystématique s’est révélé LE GENRE AMPHOTEROMORPHUS 407 AAT @idegeesr>chered | |, ACP ee i reale | Be . ii PEGS Es 59 Cd, beat i i ii i | I 1 age ! I Of i ! | Dra ii fe AK ANT EN ILE CE © 3 sw IE 22 BE 4 li à È * | RS i h IRE Peer Ss AA) igi ESTA 22297 aA2r een ave iO? gen Lui Li Be ha NN] e u ee co 2 RL AP i EC È d © ap, Db c d e MDH a ne: episssgii ‘is UL oo POME Se ı ı rat ity vel it 28 4 reat siens $9,262 oo? —? eee $ DS eS eS Se ZN po dl DS e NE AVR a ENG SE I P| be EN a b c d e a b c d e PGI I 1 ER BESO ig & #99: 56! “n i | 06838 I I I 1 1 x tt MNEENEZZ SZ SIE] ES a b c d e FIG. 37 Zymogrammes de sept systèmes enzymatiques des cinq espèces du genre Amphoteromorphus. a = A. parkamoo; b = A. piraeeba; c = A. piriformis; d = A. ovalis; e = A. ninoi (hötes = B. vail- lanti: 2 premières pistes, B. filamentosum. précieuse en association avec les critères utilisés en taxonomie traditionnelle. Néanmoins, les trois espèces peuvent étre séparées par plusieurs caractères morpho- anatomiques, métriques et méristiques (Tableau 1) d’importance taxonomique sùre (nombre de testicules, taille du scolex, forme de l’embryophore). Au contraire, A. ovalis et A. ninoi sont séparables, en plus de leurs profils électrophorétiques, par le nombre de testicules, la largeur comparative de l’ovaire, le nombre de diverticules utérins. Chez toutes les espèces du genre, la longueur comparative de la poche du cirre, la longueur des crochets des oncosphères et la largeur des proglottis est très semblable. En dépit de leur ressemblance morphologique générale, certaines espèces présentent des apomorphies utiles pour l’identification. A ninoi se distingue par ses vitellogènes peu nombreux, en bandes transversales, par la faible longueur de ses proglottis et par 408 M. CARFORA ET AL. la présence de deux diverticules utérins seulement. A. piriformis est le seul a posséder des embryophores mürs piriformes. A. piraeeba possede un scolex en moyenne de fort diametre. CLE DE DETERMINATION 1 Pores génitaux irrégulièrement alternes; hôte: Brachyplatystoma flavi- GOISERN REN ENO SUR Re RAP Ans A. peniculus Diesing, 1850 - Pores génitaux unilatéraux (ou avec une ou deux alternances dans de très RATES ICAS) EN i lie ws eee à Se Lo LIRE 2 2 Moins de 50 testicules par proglottis "Ne Ne PE PEER 5 - Pluside 50 testicules par proglottis .. PRE SERRE 4 3 Moins de 33 testicules par proglottis; hötes: Brachyplatystoma filamen- tosumset Be VQUIANI ps RI ANSIA DINAR A. ninoi sp. n. - Entre 34 et 48 testicules par proglottis; höte: Paulicea luetkeni RAR A ER I MORE SE Ver A. parkamoo Woodland, 1935 4 Embryophore des oeufs mürs de forme piriforme; höte: B. flavicans N cranny A usa ORIO A. piriformis Sp. n. - Embryophore des oeufs mûrs de forme ronde à ovoïde ................. 5 9 Embryophore des oeufs mürs de forme ovoide. Scolex 980-2310 de diametre shoteBrachyplatystoma Sp. MC SME SERE A. ovalis sp. n. - Embryophore des oeufs mürs de forme ronde. Scolex 2000-3740 de diametre hôte: Bi filamentosum ©... 2.562). 2 a): A. piraeeba Woodland, 1934 REMERCIEMENTS Nous tenons tout particulièrement à remercier le Dr Jean Wuest, Genève, (mi- croscope électronique à balayage), le Dr François Renaud, Montpellier, (interprétation des électrophorèses), le Dr John Lundberg, Philadelphia (identification des hötes) et le Prof. Amilcar Arandas Rego, Rio de Janeiro, (participation au travail de terrain et a l’obtention des autorisations nécessaires). Une version antérieure de ce travail constitue le diplôme du premier auteur présenté à l’Institut de Zoologie de Neuchâtel sous la direction du Pr Bruno Betschart. RÉFÉRENCES DE CHAMBRIER, A. 2001. A new tapeworm from the Amazon, Amazotaenia yvettae gen. n., sp. n., (Eucestoda: Proteocephalidea) from the siluriform fishes Brachyplatystoma filamen- tosum and B. vaillanti (Pimelodidae). Revue suisse de Zoologie 108 (2): 303-316. DE CHAMBRIER, À. & VAUCHER, C. 1997. Révision des cestodes (Monticelliidae) décrits par Woodland (1934) chez Brachyplatystoma filamentosum avec redéfinition des genres Endorchis Woodland, 1934 et Nomimoscolex Woodland, 1934. Systematic Parasitology 373219233. DIESING, C.M. 1850. Systema Helminthum. Vol. I. Vindobonae, xiii + 679 pp. EUZET, L. & COMBES, C. 1980. Les problèmes de l’espèce chez les animaux parasites. Mémoires de la Societe Zoologique de France 40 (3): 239-285. FREZE, V.I. 1965. Essentials of cestodology. Vol. V. Proteocephalata in fish, amphibians and reptiles. Moskva, Izdatel’stvo “Nauka”, 538 pp. (En russe, traduction anglaise: Israel Program of Scientific Translation, 1969, Cat. No. 1853, v + 597 pp). LE GENRE AMPHOTEROMORPHUS 409 FUHRMANN, O. 1934. Vier Diesing’sche Typen (Cestoda). Revue suisse de Zoologie 41: 545-564. HırLıs, D.M, MORRITZ, C. & MABLE, B. K. 1996. Molecular Systematics. Sinauer, 655 pp. PASTEUR, N., BONHOMME, G., CATALAN, J. & BRITTON-DAVIDIAN, J. 1987. Manuel technique de génétique par électrophorèse des protéines. Paris, Lavoisier, 217 pp. REGO, A.A. & PAVANELLI, G.C. 1999. The metascolex in proteocephalids. Acta Scientarum 21(2): 247-254. REGO, A.A. 1994. Order Proteocephalidea Mola, 1928 (pp. 257-293). In: KHALIL, L.F, JONES A. & Bray R.A. (eds). Keys to the Cestode Parasites of Vertebrates, Wallingford, CAB International. ScHMIDT, G.D. 1986. Handbook of Tapeworm identification. Boca Raton, Florida, C.R.C. Press Inc., 675 pp. WOODLAND, W.N.F. 1933. On the anatomy of some fish cestodes described by Diesing from the Amazon. Quarterly Journal of Microscopical Science 76: 175-208. WOODLAND, W.N.F. 1934. On some remarkable new cestodes from the Amazon siluroid fish, Brachyplatystoma filamentosum (Lichtenstein). Parasitology 26: 268-277. WOODLAND, W.N.F. 1935. Some more remarkable cestodes from Amazon siluroids fish. Para- sitology 27: 207 —225. de x ESS a) tire REVUE SUISSE DE ZOOLOGIE 110 (2): 411-426; juin 2003 The presence of Soprano pipistrelle Pipistrellus pygmaeus (Leach, 1825) in Switzerland: first molecular and bioacustic evidences Barbara WICHT!, Marco MORETTF,, Damiano PREATONE, Guido TOSB & Adriano MARTINOLI3* ! Institut de Zoologie, Faculté des Sciences de l’Université de Neuchatel, Chantemerle 22, CH-2007 Neuchatel, Switzerland. 2 Centro Protezione Chirotteri Ticino, CH-6714 Semione, Switzerland. E-mail: marco.moretti@bluewin.ch 3 Dipartimento di Biologia Strutturale e Funzionale, Universitä degli Studi dell’Insubria, Via J. H. Dunant 3, I-21100 Varese, Italy. E-mail: adriano.martinoli@uninsubria.it The presence of Soprano pipistrelle Pipistrellus pygmaeus (Leach, 1825) in Switzerland: first molecular and bioacustic evidences. - The two widespread sibling species of pipistrelle bats, the soprano pipistrelle (Pipistrellus pygmaeus), and the common pipistrelle (Pipistrellus pipistrellus), that emit at distinct call frequencies (maximum energy at around 55 and 45 kHz respectively), have been found in sympatry over much of Europe, but little is known about their relative abundance and possible habitat use. In this study, we provide the first record of P. pygmaeus in southern Switzerland, confirmed by echolocation analysis and mitochon- drial DNA sequencing. During 70 catching sessions in the southern region of the Swiss Alps, echolocating bat calls were recorded to define the distri- bution/presence of the two species. Habitat characteristics of capture sites suggested that the sibling species have a high degree of niche overlap (0.88). Nevertheless, P. pygmaeus seemed to select agricultural areas and riversides more frequently than P. pipistrellus, which is found more often in urban ar- eas. Divergence in the cytochrome b gene of mitochondrial DNA of 37 bats confirmed the echolocation data for 31 of 33 P. pipistrellus and for two of four P. pygmaeus, while four animals, two of each species, had been mis- classified based on echolocation only. A canonical discriminant function using morphological data was not reliable to distinguish the sibling species. Key-words: Chiroptera - Pipistrellus pygmaeus - Sibling species - mtDNA - echolocation - Switzerland. INTRODUCTION The study of sibling species has great importance in community ecology, zoo- geography and conservation. Sibling species exist in many zoological taxa (insects and Vertebrates as fishes, reptiles and rodents). Among European bats, this has been * Corresponding author Manuscript accepted 25.11.2002 412 B. WICHT ET AL. demonstrated for Myotis myotis and Myotis blythii (Arlettaz et al., 1993, 1997; Arlettaz, 1996) and for P. pipistrellus and P. pygmaeus (Barlow, 1997; Barlow & Jones, 1997, 1999). Populations of sibling bat species living in sympatry are defined as cryp- tic species, because they consist of individuals that are morphologically similar, or even identical, that can only be recognised by biomolecular or bioacoustical methods. The case of the common pipistrelle bat (Pipistrellus pipistrellus Schreber, 1774) is probably one of the most surprising and interesting discoveries among European mammals: until 1999, this species was regarded as a single biological unit. In 1993 two different phonic types were discovered among British pipistrelles: individuals emitting two distinct search-phase echolocation calls were recognized, with frequencies of max- imum energy at around 45 kHz and at around 55 kHz (Jones & Van Parijs, 1993). Subsequently, genetic analysis proved that the Europe’s most widespread and well- studied bat actually exist as two cryptic species (Barratt er al., 1997), separated by a sequence divergence of >11% in the cytochrome b gene of mitochondrial DNA. Despite the fact that a definitive taxonomic classification is not yet defined (von Helversen et al., 2000; Mayer & von Helversen, 2001) we have chosen to adopt the name proposed by Jones & Barratt (1999), based on suggestions made by Hutson and Jones (von Helversen et al., 2000: pp.115, 116). Jones and Barratt (1999) proposed the nomenclature Pipistrellus pygmaeus (Leach, 1825) for the phonic type “55 kHz” - popularly called soprano pipistrelle, thanks to its high-pitched call - while the phonic type “45 kHz” remains the common pipistrelle, Pipistrellus pipistrellus. The two phonic types can be readily identified by using a bat detector, and have been found in sympatry in Switzerland (Zingg, 1990; Letard & Tupinier, 1997), Great Britain (Jones & Van Parijs, 1993), North of Ireland (Russ, 1996), central Spain (Ruedi er al., 1998; Mayer & von Helversen, 2001), Germany (Haussler et al., 1999; Mayer & von Helversen, 2001), France (Lustrat, 1999) and in the South of Italy and Sardegna (Russo & Jones, 2000). In other countries the presence of the P. pygmaeus is recorded also for Greece (Weid & von Helversen, 1987; Mayer & von Helversen, 2001), Denmark, Norway and Portugal (Jones, 1997). In the northern part of the Swiss Alps, a first bioacoustical record of two forms of echolocating common pipistrelles was made by Zingg already in 1990 even though Zingg didn’t identify them as two different species but as two different phonotypes. The presence of two species, after the classification of Jones & Barratt (1999), has never been confirmed in Switzerland by biomolecular evidence. In this study, we investigate the occurrence of the two sibling species of Pipistrellus in the Southern region of the Swiss Alps, that includes the Cantone Ticino (Switzerland) and two northern provinces of Italy (Como and Varese), using biomolecular and bioacoustical analyses, and analysing environmental and morphological data. MATERIALS AND METHODS STUDY AREA AND CAPTURE METHODS The study was carried out from May to October 2001. We visited 70 sites (Fig. 1), 52 in Cantone Ticino (Switzerland), 10 in the province of Como (Italy) and 8 in the province of Varese (Italy), where we captured different bat species. The study PIPISTRELLUS PYGMAEUS IN SWITZERLAND 413 N ® df i > Gordevio > are ns one hee b n oe" . 4 D N \ / Lugano | jo © (DI 2 di a © fi À vs. I Verbania 7 ) Varese Ir | Lecco Han Ÿ Como ® ® e°° 10010220 km SI se Bil Study area. Dots indicate sampling sites. area extends over 7152 km? (coordinates range: 46°33’46.8”N; 9°15’43.2”E; 45°35°16.8"S; 8°22’33.6”W) and the landscape consists of 20% lowlands, 45% hills and 35% mountains, with elevation ranging from 200 m a.s.l., at Lake Maggiore, Varese province, to over 3000 m in the Swiss Alps (Rheinwaldhorn, 3348 m), in the northern part of the study area. The continental climate is tempered by the large lakes, with relatively mild winters, and abundant precipitation ranging from 1500 to 2200 mm per year. 414 B. WICHT ET AL. Most sites visited in the study area (n = 45) were maternity roosts located in houses that had been previously signalled by local people to the Centre for the Protection of Bats of the Canton Ticino, a local bat conservation group founded in 1990. A church, two factories and one cave were also checked for bats. Additionally, 19 foraging sites along rivers and water-courses (random points) and 115 bat boxes in two deciduous woodlands were monitored. Bats were captured during evening emergence at roosts, using hand-nets, which allowed us to catch only part of the individuals in each of the colonies. Of all caught bats, morphological measurements were taken and their echolocation calls were recorded, while skin samples were taken from a maximum of five animals per site. The total number of animals emerging was counted. At foraging sites, mist-nets were placed along ponds and water-courses, as described by Kunz (1988), that remained activated from dusk to midnight, and all the data described above were taken for each captured bat. Bats roosting in boxes, mainly males in mating groups, were taken by hand. SAMPLING PROCEDURE Each captured bat was sexed and aged. The age-class, (sub-adult or adult), was determined based on epyphyseal growth plates closure in the metacarpal-phalangeal joint of the fifth finger (Anthony, 1988). Reproductive condition was assessed check- ing testes development in males and lactation in females (Racey, 1988). Females were defined lactating when milk appeared while gently squeezing the nipples, or when the fur surrounding nipples was absent. Twelve morphological measurements of bats were taken using a precision callipers (+ 0.01 mm): forearm length, wing span (from the wing extremity to the shoulder, gently stretching out the left wing,), III and V finger (from thumb insertion to III finger extremity and V finger extremity respectively, gently stretching out the left wing), II and III phalanx of III finger, thumb (from thumb inser- tion to thumb extremity without nail), foot (from heel to the third finger extremity with- out nail), tibia, tail (from anus to tail extremity), and ear length (from tragus insertion to ear extremity). Bats were weighed to the nearest 0.5 g with a 50 g Pesola dy- namometer. In a first step, bats were determined as belonging to one of the sibling species using their echolocation characteristics, morphological measurements were compared by multivariate analysis of variance (MANOVA, Sokal & Rohlf, 1995). Ina second step, the morphological parameters that were significantly different between the two species were analyzed with the SYSTAT statistical package (SPSS, 1997) in order to calculate a Discriminant Function with Jackknife procedure (Hinkley, 1977) to determine the percentage of cases classified correctly (significance was tested with a F-transformed Wilk’s lambda test, Tukey, 1977). Finally, a canonical discriminant function was calculated to distinguish between the two species. For each bat, echolocation calls were recorded in time expansion mode with a Pettersson D980 bat detector, while releasing it from the hand in open habitat near the roost. Echolocation calls spectrograms were subsequently obtained (1024 points Fast Fourier Transform, Parzen window) and analysed using BatSound software (Pettersson, 1999). For statistical analysis, to avoid sample size problems, a subset of 18 randomly chosen recordings of P. pipistrellus were compared with the total data set PIPISTRELLUS PYGMAEUS IN SWITZERLAND 415 of 18 P. pygmaeus. For each recording, six characteristic frequency-domain parameters were measured: start frequency (F,.„), maximum frequency (F,,,), minimum fre- quency (Fin) frequency at maximum intensity (Fm), end frequency (F.,4), and frequency at half the duration of the search phase call (F,,). Finally, call duration (in ms) was also measured. Each parameter was compared between the two sibling species with a non-parametric Mann-Whitney U-test. Significant parameters were then used in a Discriminant Function Analysis with Jackknife procedure, to obtain a classification function based on these echolocation parameters. GENETIC ANALYSIS Two tissue samples of 4 mm diameter were taken from the tail membrane (uropatagium) of each skin-sampled bat using a biopsy punch, and placed in a 0.2 ml eppendorf vial filled with 70% ethanol and stored at —20 °C until the moment of genetic sequencing. Genetic analyses were carried out on a total of 137 bats, for which also bioacoustic and morphologic data were available. All bats were handled with care and kept as little as possible to avoid stress, and no animals were injured or died during handling. DNA from tissue samples was extracted using the QIAmp DNA Mini kit (Qiagen) following supplier’s instructions. Each sample was amplified by PCR (Saiki et al., 1988) using primers L14841 and H15149 (Kocher er al., 1989), targeting a 307 base pairs (bp) portion of the mitochondrial DNA (mtDNA) cytochrome b (cyt b) gene. Double-stranded cycle sequencing was conducted using the ABI PRISM™ BigDye Terminator Cycle Sequencing Ready Reaction kit, Version 3.0 (Applied Biosystems). Purified sequencing products were loaded on a ABI PRISM™ 377 DNA sequencer (Applied Biosystems). Both strands have been sequenced with the same primers used in the PCR amplification. For species identification, the DNA sequences obtained from the samples were compared with homologous sequences of other Chiroptera available in DNA databases, in particular with the Pipistrellus spp. sequences from Barratt et al. (1997) (GenBank accession nr. U95499, U95501, U95503, U95505, U95507, U95509). Distance matrix calculation (Kimura 2-parameter distances; Kimura, 1980) and phylogenetic analysis (maximum parsimony, exhaustive search with Rhinolophus ferrumequinum (U95513) as outgroup) were performed with PAUP* 4.0b8 (Swofford, 2001). Resolution of internal nodes was evaluated using 10000 bootstrap replications with the PAUP* package. HABITAT ANALYSIS A Geographical Information System was set up with ESRI ArcView GIS version 3.2 (ESRI, 1999), using scanned 1:25000 paper maps from the Swiss Federal Office of Topography and Regione Lombardia CT10 1:10000 Technical Regional Digital Cartography as reference coverages. Land cover data was derived from the GEOSTAT data sets (Swiss Federal Statistical Office, 1997) for the Swiss portion of study area, and from CORINE Land Cover coverage (Commission of the European Communities, 1993) for the Italian part. Sampling sites were digitized heads-up using the above cited reference maps. Due to the different projection systems used in Switzerland and Italy, all the Italian geographic datasets were converted using ESRI 416 B. WICHT ET AL. ArcGIS ARC/INFO 8.1 PROJECT routine (Booth, 1999) into Hotine Oblique projec- tion (Swiss Reference Grid) and harmonised with GEOSTAT geodataset. Furthermore, CORINE Land Cover coverage has been converted to raster format and downsampled at the same spatial resolution of GEOSTAT, that is at a pixel size of 0.1 km2. Because of different classification criteria in the two land cover datasets, classes were pooled and standardized according to Land Cover Classification System standards (Di Gregorio & Jansen, 2000), yielding the land cover classes shown in Table 1. TABLE 1. The percentage of habitat use by the sibling species Pipistrellus pipistrellus (N = 3984 locations) and P. pygmaeus (N = 404 locations), and the percentage of available habitat deter- mined for the entire study area based on 0.1 km? grid cells. Habitat types Habitat use Available P. pipistrellus P. pygmaeus habitat Woodland 46.4 46.0 44.5 Shrubland 0.4 0.5 3.9 Orchards, vineyards 6.3 2 5.3 Cultivated farmland 0.8 DU 4.5 Meadows, alpine meadows 10.8 6.7 16.0 Water 4.2 9.9 5.8 Sterile land 0.6 0.5 Mol Urbanised land 30.4 233 12.4 Finally, in order to obtain habitat class frequency for all the sampling sites, each point site was buffered using a fixed radius of 500 m, and converted into raster, and habitat class frequency was recorded as number of 0.1 km? pixels intersecting or con- tained in the buffered areas. Habitat use and habitat overlap were both evaluated using a normalized version of the Proportional Similarity Index (PS,,), that is _PS-min(g;). “4 1—min(q; ) PS n 0< PS, <1], where PS is the Czekanowski’s Proportional Similarity Index (Feinsinger ef al., 1981): PS 1; Zn seg; aan) The symbols p; and q; indicate respectively the proportion of used and available land use class, when PS, is used to estimate niche breadth, whereas the same notations indicate the proportion of habitat used by each species when PS, is used to estimate niche overlap; min(g;) represents the minimum observed resource item frequency. Habitat preference by each species was evaluated using Ivlev’s Electivity Index (Jacobs, 1974). RESULTS A total of 371 bats of different species were captured and analysed, 240 of which belonged to one of the two Pipistrellus sibling species. In a first step, distinction between P. pipistrellus and P. pygmaeus was based on acoustical data analysis. For a PIPISTRELLUS PYGMAEUS IN SWITZERLAND 417 subset of 18 individuals of each species, there was no difference in call duration (Mann-Whitney U-test: U = 105.5, p = 0.07), and in call maximum frequency (Fig. 2b, U = 121.0, p = 0.20). All other call parameters differed significantly between the two sibling species (Fig. 2, Mann-Whitney U-test all p < 0.01) and the largest difference was found for the frequency at maximum amplitude, on average 46.48 kHz for P. pipi- strellus and 57.52 kHz for P. pygmaeus (Fig. 2e, U = 0, p < 0.01). Using only signi- ficant call frequency variables, a canonical discriminant function analysis (DFA) was calculated (F.pprox. = 35.0; df =5, 30; p < 0.001), which permitted us to determine 181 of the 201 recorded individuals’ as P. pipistrellus and 20 as P. pygmaeus. The discri- minant function, in its canonical form is 2110100336... O.071F,,, OA0TE vac, — 0.081. +002. with a total discriminant capacity of 94%. Negative y values refer to P. pygmaeus, while positive ones refer to P. pipistrellus (Fig. 3). Reconstruction of the phylogenetic tree by maximum parsimony and 10000 bootstrap replications showed that 33 of the 37 samples of Pipistrellus, belonged to the species P. pipistrellus, and four to P. pygmaeus. Of the four bats classified erroneously based on echolocation data, two genetically determined P. pygmaeus had been first determined as P. pipistrellus based on frequencies at maximum call intensity (Fin) of 45.40 and 45.94 kHz respectively, while two genetically determined P. pipistrellus had been first classified as P. pygmaeus based on (Fin) Of 57.05 and 61.90 kHz. Analysis of the 307 base pairs of the mtDNA cyt b gene confirmed DFA results, except for four animals that had been misclassified. The comparison of DNA sequences obtained with DNA reference sequences resulted in a K2P genetic distance of 0.4% between the two P. pipistrellus, 0.4% between the two P. pygmaeus, and 12.8 + 0.5% between the two species. The MANOVA on the 12 morphological traits was statisti- cally significant (F = 4.40; df = 12, 171; p < 0.001) and seven out of 12 morphological parameters differed significantly between the two sibling species at the 1% level (one- way ANOVA, level of significance p < 0.01, Table 2). A discriminant function for each species was calculated using only the four parameters that had the highest significance level (single one-way ANOVA, p < 0.001, Table 2). For P. pipistrellus, the slightly larger species, the discriminant function was described by y = -499.91 + 20.58thumb + 3.93tibia + 2.24tail + 25.35 forearm while for P. pygmaeus it was y = -463.34 + 18.66thumb + 3.02tibia +1.84tail + 25.15 forearm The Jackknife procedure classified correctly 141 out of 175 (81%) P. pipistrellus, and 16 out of 20 (80%) P. pygmaeus (probability of uncorrect assessment: Wilk’s lambda F.pprox. = 9.99; df = 4, 190; p < 0.0001). The canonical discriminant function obtained was described by y = -24.92 + 1.28thumb + 0.61 tibia +0.26tail + 0.13 forearm with y < 0 indicating P. pipistrellus and y > 0 indicating P. pygmaeus (Fig. 5). Thus, taking into account that about 19% of animals were misclassified, morphological measurements alone are not sufficient to distinguish between the two pipistrelle sibling 418 B. WICHT ET AL. 90 90 e | > 0 | 3 LIUTO 4 = cla Sr | Isl à | x 60+ 1 | + | e E Ta 50 + al LESC | 40 - 4 40 - 30 30 | 90 90 c | d 80 + - 80 - 4 70 + - = 70+ 4 FIT N x È TE = È i 4 60 a * n £ 0. | ie CRB =. en 50 - Alk ° + E50), | | + ns) A aL ie) | = e | 40 + = 40 È Ai e 30 30 | 90 90 e | f 80 + = 80 F = N 70} UE | E se a — ra x | je 60) ALS) - AGE 5. - x | = = (00) e © LE 50! Ber 4 L 50+ Si a - 40 + 7 - 40 - 3 { | | 30 30 =] P. pipistrellus P. pygmaeus P. pipistrellus P. pygmaeus BiG Box-and-whisker plot comparison between Pipistrellus pipistrellus and P. pygmaeus ultrasonic search-phase call parameters: a: Fin, b: Fax, €: Fin, di Fin €: FE EU min? maxint? end: species in our study area. Finally, following Haussler et al. (1999) we also calculated the mean differences between the second and third phalange of the 3"4 finger, but found no significant difference between the sibling species (Mean difference + sd in mm: P. pipistrellus 1.05 + 0.70; P. pygmaeus 1.10 + 0.91; t-test t = 0.27; df = 187; p = 0.79). PIPISTRELLUS PYGMAEUS IN SWITZERLAND 419 ie 10 © pipistrellus x pygmaeus Frequency oe 0 = LC “3 5.2.4, 53.2 120, 1 23145 Discriminant function value Fic. 3 Discriminant Function Analysis (DFA) canonical scores plot for ultrasonic search-phase call parameters in Pipistrellus spp. Of the 33 P. pipistrellus examined, 23 (70%) had been captured at 10 different roosts in houses, eight (24%) along foraging sites in the Cantone Ticino, Switzerland, and two (6%) at one foraging site in the province of Como, Italy. Two P. pygmaeus had been captured at Gordevio, along the river Maggia, and two in bat boxes at Bosco Isolino, Cantone Ticino, Switzerland (Fig. 1). Using both echolocation and genetic determination of the two sibling species, we compared the habitat characteristics of the sites where they had been found (N = 3984 for P. pipistrellus, N = 404 for P. pygmaeus, Table 1). Spatial niche breadth val- ues were high and similar for both species (PS, = 0.78 for P. pipistrellus; PS, = 0.79 for P. pygmaeus), and there was a high degree of niche overlap (PS, = 0.88). Each species used habitat types in a significantly different fashion if compared with habitat availability (Table 1; x? = 41.4; df = 7; p < 0.001 for P. pipistrellus; = x? = 29.2; df = 7; p < 0.001 for P. pygmaeus). The Ivlev’s electivity index showed avoidance for sterile land (non-vegetated areas, bare rock and glaciers), shrubland and meadows, and posi- tive selection for orchards and urbanised areas in both species. Farmland and water bodies were avoided by P. pipistrellus, but positively selected by P. pygmaeus (Fig. 4). 420 B. WICHT ET AL. 1.00 I P. pipistrellus BR 0.75 . pygmaeus 0.50 0.25 0.00 Electivity Index -0.25 -0.50 -0.75 -1.00 Sterile land Shrubland Meadows Woodland Farmland Orchards Water Urbanised land Habitat types Fic. 4 Ivlev’s Electivity Index values by habitat type for Pipistrellus pipistrellus and P. pygmaeus. DISCUSSION Morphological data from the two sibling species from the Rhine valley in Southwest Germany suggested that P. pipistrellus tends to be slightly larger than P. pygmaeus (Haussler et al., 1999). In particular, there was a significant difference in mean length of the forearm, fifth finger and average tail length. This was confirmed by our data of external morphological measurements taken on live animals from alpine and prealpine areas of southern Switzerland and the provinces of Varese and Como, northern Italy. However, although differences in mean size existed, individual variation was large and there was considerable overlap in all measurements that were taken. Consequently, the best canonical discriminant function still misclassified about 20% of bats of each sibling species in our study area. Haussler er al. (1999) suggested that the difference in length between the 2nd osseous phalange and the terminal 3" cartilaginous phalange of the 3" finger, almost the same length in P. pygmaeus, while in P. pipis- trellus the 3" phalange is generally 2-3 mm shorter than the 24, is a useful diagnostic morphological characteristic that can be measured in the field. However, they do not produce any statistical evidence for this statement. Moreover, in our study area there was no such difference, indicating that it can not be used as a reliable diagnostic criteria. Thus, morphological measurements alone are not sufficient to investigate the presence/absence or distribution of the pipistrelle sibling species, at least in the southern Swiss Alps and confining areas in northern Italy. PIPISTRELLUS PYGMAEUS IN SWITZERLAND 42] 15 © pipistrellus x pygmaeus 10 Frequency O1 AIA Discriminant function value FIG. 5 Discriminant Function Analysis (DFA) canonical scores plot for biometric parameters in Pipistrellus spp. In contrast, a detailed analysis of echolocation calls allowed us to calculate a canonical discriminant function which resulted in correct determination of 35 out of 36 bats (97%). In agreement with previous studies in different parts of Europe (Weid & von Helversen, 1987; Jones & Van Parijs, 1993; Vaughan et al., 1996, 1997; Barlow, 1997; Barlow & Jones, 1999; Russo & Jones, 2000; Mayer & von Helversen, 2001), the frequency at maximum intensity of the search phase call was the call parameter with the lowest inter-specific overlap (see Fig. 2). However, we must underline that definitive species determination using the 307 bp sequence of the cyt-b gene of mtDNA, demonstrated that in some cases, species classification based on echolocation calls had failed. In part this might be caused by the individual variation in search phase call frequency in both sibling species, found also in British colonies (Barlow & Jones, 1999), with some individuals using frequencies 5-7 kHz above or below the mean value (see Fig. 2e). Other sources of erroneous classification can be linked with recording conditions. Therefore, we want to stress that a low error rate when discriminating bat species by means of their echolocation calls can only be ensured when the following conditions are met: (1) high-quality equipment (bat detector and sonagraph or 422 B. WICHT ET AL. computer-based Digital Signal Processing system); (2) ideal call recording conditions (no nearby obstacles, no interference by other bats with the recorded individual, limited distance between recorder and animal); (3) released bats in good condition, not stressed or suffering cold; (4) recording of the correct search phase call, not of other call types (e.g. social calls); (5) correct choice of call parameters for data analyses; and (6) appropriate statistical analysis. Although other studies on sympatric pairs of cryptic bat species have shown there can be considerable habitat niche differentiation (Arlettaz, 1996; Arlettaz er al., 1997), this was not the case for the sibling pipistrelle species in our study area which both had a large niche width, and a high degree, about 88%, of niche overlap. Nevertheless, some differences in habitat selection could be noted. As ın south-west England, the foraging habitat of the soprano pipistrelle (P. pygmaeus) was more asso- ciated with water and riparian habitats, probably related to its diet consisting largely of insects with aquatic larval stages, while it clearly avoided urbanised areas (Barlow, 1997; Vaughan er al., 1997). Also, in the landscape around maternity roosts, water courses and riparian habitat belts were present, as well as farmland, and no roosts were found in houses. This contrasted findings from Scotland, were P. pygmaeus selected house roosts that had a higher degree of nearby cover (large trees) than random houses (Jenkins er al., 1998). However, these house roosts were surrounded by a greater area of deciduous and coniferous woodland than random buildings and nearly always within 500 m from a major river (Jenkins er al., 1998), which underlines the importance of cover and the presence of nearby riparian habitats for this species (Oakeley & Jones, 1998). This was confirmed by our data with soprano pipistrelles roosting in nest box- es in deciduous woods, thus in a habitat with good cover, and foraging often along rivers and lakes. In contrast, most roosts of the common pipistrelle (P. pipistrellus) were in buildings. The latter species, commonly found in woodlands, also foraged in- tensively in cities and small villages as well as in nearby farmland. Although some- times found at water or riparian foraging sites and roosts, it used these habitats less frequently than the soprano pipistrelle. Thus, habitat requirements are slightly different for the two sibling species (Vaughan er al., 1997), with the common pipistrelle being a habitat generalist and the soprano pipistrelle selecting riparian habitats and farmland. However, overall, there were only small differences in our measurements of habitat use by the sibling species (see Table 1), resulting in a high degree of habitat niche overlap, and differentiation may be more accentuated for other niche components. In fact, both in our study and in others (Park er al., 1996; Barlow, 1997; Oakeley & Jones, 1998; Jenkins er al., 1998), there were no roosts occupied by both species simultaneously. Moreover, diet studies of the sibling pipistrelles in the British Isles showed that although both species ate mostly the dipteran suborder Nematocera and there was no difference between the phonic types in dietary breadth, there were some differences in food choice (Barlow, 1997). The main prey groups in the diet of P. pipistrellus were the families Psychodidae, Anisopidae and Muscidae, whereas the families Chironomidae and Ceratopogonidae occurred most frequently in the diet of P. pygmaeus (Barlow, 1997). Thus, stable co-existence of sympatric populations of P. pipistrellus and P. pyg- maeus in heterogeneous landscapes seems possible because of differential selection of roost sites and preferred prey. Further studies are needed to test this hypothesis and to PIPISTRELLUS PYGMAEUS IN SWITZERLAND 423 TABLE 2. Morphological characteristics (mean + sd) of 164 Pipistrellus pipistrellus and 20 P. pygmaeus. Determination of sibling species based on echolocation call frequency. Differences between means for each single parameter tested by one-way ANOVA (df = 1, 182 in all cases). Parameter P. pipistrellus P. pygmaeus E p Body mass (g) 4.95 + 0.59 3213, 0/92 122 027 Tumb (mm) 4.34 + 0.36 4.02 + 0.35 15.8 < 0.001 Tibia (mm) 11.74 + 0.50 1122) +067 17.8 < 0.001 Foot (mm) 6.53 + 0.62 6.46 + 0.36 0.15 0.70 Tail (mm) 31.9832. 29.54 + 1.84 DS < 0.001 Ear length (mm) 9.50 + 0.88 9.36 + 0.72 0.24 0.62 Forearm (mm) 31.19 + 1.01 30.28 + 1.29 14.1 < 0.001 II Finger (mm) 54.34 + 2.56 32.27,+3.06 11:5 0.001 V Finger (mm) 40.00 + 1.96 38.84 + 1.87 6.37 0.012 III Phalanx (mm) Wee 0.53 6.82 + 0.71 8.48 0.004 II Phalanx (mm) 8.27 + 0.60 7:92 2058 6.59 0.011 Wing length (mm) 93.34 3,39 91.26 + 4.97 NEID. 0.006 investigate whether the realised niche width of allopatric populations of P. pygmaeus differs from that of conspecifics in sympatry with the sibling species. Collecting further information on habitat composition of foraging sites and lo- cation of preferred roosting sites of P. pygmaeus will be essential for landscape man- agement and planning of specific conservation measures. For example, modification of certain riparian habitats and water courses by constructing sewage plants and conduc- tors could cause marked changes of the invertebrate fauna of rivers, torrents and small lakes downstream of the sewage output, and thus alter food availability for, and activ- ity of, foraging bats (see also Vaughan et al., 1996). Since the soprano pipistrelle feeds primarily on insects with aquatic larval stages, such modifications might have serious consequences for the local survival of soprano pipistrelle populations. We conclude that our genetic data definitively confirm that both P. pipistrellus and P. pygmaeus occur in Switzerland. Furthermore, our results indicate that correct monitoring of echolocation calls of pipistrelle bats is the most appropriate technique for large-scale studies on presence/absence and distribution of the two sibling species, but larger samples of genetically identified P. pygmaeus are needed to verify how reli- able echolocation analyses are for correct species determination, as well as careful ul- trasound recording. ACKNOWLEDGEMENTS We thank the Cantonal Museum of Natural History, Lugano, for financial sup- port and the Cantonal Nature Conservation Office for the permission to capture and study bats. The Centro Protezione Chirotteri Ticino allowed us access to their GEOSTAT data base and helped with the fieldwork. We are also grateful to Sabrina Mattiroli, Riccardo Pierallini, Roberta Chirichella, Tiziano Maddalena, Marzia Roesli, Mosé Nodari for helping with the fieldwork, to Luca Fumagalli for genetic analyses and to Claude Mermod for his encouragement throughout the project. 424 B. 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Akustische Artidentifikation von Fledermäusen (Mammalia: Chiroptera) in der Schweiz. Revue suisse de Zoologie 97: 263-294. REVUE SUISSE DE ZOOLOGIE 110 (2): 427-435; juin 2003 Description of a new species of Tityus (Scorpiones, Buthidae) from Serra do Cipo in the State of Minas Gerais, Brazil 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 Description of a new species of Tityus (Scorpiones, Buthidae) from Serra do Cipo in the State of Minas Gerais, Brazil. - Tiryus adrianoi sp. n., belonging to the Tityus bahiensis species group (Scorpiones, Buthidae) is described on the basis of one female specimen collected in the region of the Serra do Cipo, State of Minas Gerais, Brazil. Comments on the taxo- nomic position of the new species and on the environmental characteristics of the type locality are given. An identification key of the species belonging to the Tityus bahiensis group distributed in the Cerrados is provided. Some comments on their patterns of geographical distribution are added. Key-words: Scorpiones - Buthidae - Tityus adrianoi sp. n. - Serra do Cipo - Cerrados - Brazil. INTRODUCTION Tityus species which inhabit the Cerrados landscape formations of Central Brazil and belong to the Tityus bahiensis species group (as defined by Lourengo, 2002), have been the subject of several taxonomic studies in the last 20 years. Noteworthy among these species are: Tityus fasciolatus Pessôa and Tityus charreyroni Vellard (Lourenço, 1980); Tityus stigmurus (Thorell) and the related species Tityus serrulatus Lutz & Mello (Lourenco, 1981); Tityus bahiensis (Perty) and related species (Lourengo, 1982). In subsequent contributions, precise patterns of distribution and differentiation have been synthesised (Lourenco, 1986, 1994, 1996), and I assumed that the Tityus species living in the Cerrados were largely known. Several regions within the Cerrado formations of Central Brazil have been intensively surveyed, mainly in connection with scorpionism (Lourenço & Cloudsley-Thompson, 1996; Lourenço et al., 1996). This is the case in particular for the states of Minas Gerais and Goias. The recent discovery and description of new Tityus species from the Cerrado formations of Central Brazil (Lourenco, 2001a,b) attests, however, that the inventory work is far from being complete. In the present paper a new species, Tityus adrianoi sp. n., belonging to the Tityus bahiensis species group, is described from a female specimen collected in the region of the Serra do Cipo, State of Minas Gerais. The Serra Manuscript accepted 10.12.2002 428 W. R. LOURENCO do Cipo region is part of the Central Brazil Cerrado formations, and separates the Atlantic forest zone from the Cerrado region (Eiten, 1978, 1982). The taxonomic position of the new species, and the environmental characteristics of the type locality are discussed. RESULTS DESCRIPTION Tityus adrianoi sp. n. Figs 1-10 Type. Brazil, State of Minas Gerais, Serra do Cipo region (Campo Rupestre formation), female holotype, 28/11/1963 (J. Lacroix leg.), deposited in the Natural History Museum, Geneva. Etymology. Patronym in honor of Dr Adriano Monteiro de Castro Pimenta of the Universidade Federal de Minas Gerais, Belo Horizonte, for his continuous interest in the study of scorpions. Diagnosis. Scorpion of medium size, with 55 mm of total length. Coloration reddish yellow to yellowish, with the carapace and tergites blackish brown. Granul- ation strong throughout the body. Fixed and movable fingers of pedipalps with 15/16 rows of granules. All carinae complete. Pectinal teeth count 21. Description. Coloration. Basically reddish yellow. Prosoma: carapace blackish brown in the anterior and lateral regions; the anterior region with an inverted triangular blackish spot stretching from the median eyes to the lateral eyes; regions behind the oc- ular tubercle and lateral eyes yellowish; eyes strongly marked with black pigment. Mesosoma: tergites I-VI blackish brown; tergite VII yellowish, with carinae brownish. Metasoma: segments I to IV yellowish; V reddish. Vesicle: same colour as segment V; extremity of aculeus darker than vesicle. Venter yellow. Chelicerae yellowish at their base; anterior region with variegated dark pigmentation; fingers dark reddish brown. Pedipalps: yellowish; fingers slightly reddish. Legs yellowish without any diffuse spots. Morphology. Carapace moderately to strongly granular; anterior margin with a median concavity. Anterior median superciliary and posterior median carinae moder- ately developted. All furrows moderately deep. Median ocular tubercle anterior to the center of the carapace. Three pairs of lateral eyes. Sternum triangular. Mesosoma: tergites strongly granular. Median carina strong on all tergites. Tergite VII penta- carinate. Venter: genital operculum wider than long. Pectines: pectinal teeth count 21/21; basal middle lamellae of the pectines slightly dilated. Sternites with a fine granulation and with elongate stigmata; VI without keels; VII with 4 moderately developed carinae. Metasoma: segments I-II with 10 carinae; segments III-IV with 8 carinae; segment V with 5 carinae. Intercarinal spaces weakly granular. Telson with vestigial granulation on ventral and lateral surfaces and with a long and strongly curved aculeus; subaculear tooth strong and spinoid, with two dorsal teeth. Cheliceral dentition characteristic of the family Buthidae; ventral teeth on movable finger reduced and almost fused (cf. Vachon, 1963); ventral aspect of both fingers and of manus den- sely covered with long setae. Pedipalps: femur pentacarinate; patella with 7 carinae; chela with 9 carinae; all carinae moderately to strongly developed; entire surface A NEW SPECIES OF TITYUS FROM BRAZIL 429 Fics 1-2 Tityus adrianoi sp. n., female holotype. Dorsal and ventral aspects. weakly granular. Fixed and movable fingers with 15/16 oblique rows of granules. Trichobothriotaxy; orthobothriotaxy A-a (Vachon, 1974, 1975). Legs: tarsus with numerous short fine setae ventrally. 430 W. R. LOURENCO ann aan, © mann. re Le SNS Sore, II ES es a N nNAN AADTANAIANA An aan NANNINI Fics 3-10 Tityus adrianoi sp. n., female holotype. 3-8. Trichobothrial pattern of pedipalp. 3-4. Chela, dorso-external and ventral aspects. 5-6. Patella, dorsal and external aspects. 7-8. Femur, detail of the internal aspect and dorsal aspect. 9. Metasomal segment III-V and telson, lateral aspect. 10. Chelicera. A NEW SPECIES OF TITYUS FROM BRAZIL 431 Measurements (in mm). Total length 55.1. Carapace: length 6.6; anterior width 4.4; posterior width 7.3. Metasoma, segment I: length 4.6; width 3.5. Metasoma, segment V: length 8.4; width 3.3; depth 3.2. Vesicle: width 2.4; depth 2.5. Pedipalp: Femur: length 6.6; width 1.9. Patella: length 7.5; width 2.5. Chela: length 13.1; width 2.6; depth 2.4. Movable finger: length 8.8. RELATIONSHIPS The new species 7. adrianoi sp. n. belongs to the Tityus bahiensis species group. In its general coloration pattern the new species shows affinities with Tityus blaseri Mello-Leitäo from Chapada dos Veadeiros in the State of Goiäs. Both species are known from localities in between 1000 and 2000 m altitude and they are isolated by the Cerrado formations (sensu strictu) of Central Brazil. The following characters are distinctive: - The pigmentation of carapace and tergites is intensely dark, almost blackish in the new species, whereas in 7. blaseri it is yellowish brown. - In the new species fixed and movable pedipalp fingers show 15/16 rows of granules, whereas T. blaseri possesses 17/18 rows. - The new species has 21 pectinal teeth, whereas T. blaseri posseses 23-25. - The basal middle lamella of the pectines is weakly dilated in the new species, but not dilated in 7. blaseri. The new species can also be easily distinguished from Tityus bahiensis (Perty) and Tiryus serrulatus Lutz & Mello, both found in the State of Minas Gerais. All three species show different patterns of pigmentation. Moreover, the new species lacks posterior spinoid granules on the dorsal carinae of the metasoma (see Lourenço, 2002 for details). ENVIRONMENTAL CHARACTERISTICS OF THE TYPE LOCALITY The Serra do Cipo is part of the Cerrado formations of Central Brazil. Accord- ing to Eiten (1978, 1982), the type of cerrado found in Serra do Cipo is the one defined as Campo rupestre (or rocky field). These occur on certain highlands and on mountain tops and ridges at moderate altitudes, usually between 1000 and 1800 m. These campos have very high species endemicity. The soils are derived from special rock types, usually meta-quartzite (such as itacolomite), some also from fine grain ortho-quartzite, iron ore, etc. Soil derived from quartzites is a fine white or light gray sand, usually 10- 30 cm thick, sometimes with humus. The bedrock usually outcrops in blocks so that the soil may vary in depth, humus content and drainage over distances of a few cen- timetres. The physiognomy of the vegetation may be low-tree and scrub woodland, open scrub, closed scrub, savanna or grassfield. According to Eiten (1978, 1982), the flora of the Campos rupestres can be divided into at least four categories. The Serra do Cipo region is characterised by species with a growth form typical of white sand campos in central Brazil and eastern Amazonia. This growth form is characterized by branches or their ends having closely-set, short, four-ranked leaves perpendicular to the stem, that is, they are squarrose or short-cruciate. Herbs, semi-shrubs and shrubs have this growth form; the larger woody plants have the same leaf-arrangement and a candelabra branching. Altogether, the plants are quite distinct from Cerrado plants. 432 W. R. LOURENCO KEY TO THE CERRADO SPECIES OF THE TITYUS BAHIENSIS GROUP 1 Metasomal segments III and IV with 1 to 5 granules modified as moderate or weak spines; coloration yellowish ....................... 2 (1) Metasomal segments III and IV without granules modified as spines; coloration from yellow to blackish brown EER RAS 3 2 One longitudinal dark stripe running over all tergites ee nn T. stigmurus (Thorell, 1876) (2) @ontluenfspots.ontersites. >... aaa T. serrulatus Lutz & Mello, 1922 3 Coloration yellowish to reddish yellow with confluent pale brown or blackish'spots'- innate RR E, 4 (3) Coloration generally dark, brown to blackish brown or with 3 dark lon- gitudinal’ stripes on tersites tt. ei RT E 5 4 Coloration reddish yellow with confluent pale brown spots; distribution: STAUCIOMGOAS Te tars ans RINO Eee T. blaseri Mello-Leitao, 1931 (4) Coloration yellowish with confluent blackish spots; distribution: State of NfinasaGeralse e hee eee cid LOI T. adrianoi sp. n. 5 Coloration brown to blackish brown; tergites dark brown; spots present OMPEAPALPs anale gs". E. ci nn ne SI EE IE 6 (5) All tergites with 3 dark longitudinal stripes; spots present or absent on pedipalpssand legs... wee sh. bes ole 408 oh ace EE U 6 Coloration blackish brown; spots on pedipalps blackish; distribution: States or Minas Gerais and Sao Paulo” oe T. bahiensis (Perty, 1833) (6) Coloration brown; spots on pedipalps brown with small white circles; distribution: Federal District and the State of Goiäs PER ft ele cs OT os ey PO II T. jeanvellardi Lourenco, 2001 7 Pedipalps and legs densely spotted; distribution: Federal District and the SLACIORGOIAS SLR ne ect DIE T. fasciolatus Pessôa, 1935 (Mrz Bedipalps’and less with spots reduced! or absent 5. CN CLS 8 8 Pedipalps and legs without spots; distribution: State of Mato Grosso do SU 22.378 Ser se ln et i er SÌ T. trivittatus Kraepelin, 1898 (8) Pedipalps with reduced spots; legs with diffuse spots; distribution: westem part of the State of Goiäs............ T. charreyroni Vellard, 1932 BIOGEOGRAPHY 1. Most scorpion population of Tityus consist of monomorphic species. In the region of Cerrados two closely related species, Tityus fasciolatus Pessôa and Tityus charreyroni Vellard, occur in parapatry areas. A statistical study of traits, such as the patterns of pigmentation, showed no variability within either population. Moreover, pigmentation patterns within each population showed no variability during post- embryonic development. Therefore these species are clearly distinct. These two possible sibling species are both savannicolous, but they occupy different, quite spe- cific microhabitats. 7. fasciolatus is a termitophilous scorpion which lives exclusively in termite mounds built by Armitermes euamignathus Silvestri, whereas T. charreyroni lives under stones in a region west of that occupied by T. fasciolatus (Fig. 11). The A NEW SPECIES OF TITYUS FROM BRAZIL 433 9 Tityus adrianoi sp. n. ® Tityus bahiensis bahiensis O Tityus bahiensis eickstedtae A Tityus blaseri % Tityus charreyroni * Tityus fasciolatus % Tityus jeanvellardi 1000 Km ed ad aa Lan, nari —______mm OT r—————@Lcmmer=e| Fic. 11 Map showing the Central region of Brazil with the areas of distribution of some species discussed in this study. occurrence of each species depends upon the presence of its specific microhabitat. A. euamignathus is associated with particular grasses, its staple food. With variation in the soil gradient, the vegetation also changes. Mounds of A. euamignathus are only present in association with a grass cover. When grass is no longer present, termite mounds and 7. fasciolatus disappear. In areas where termite mounds are common, stones are rare, thus making the habitat unfavourable for 7. charreyroni. 2. The study of Tityus bahiensis (Perty) (Lourenco, 1982), first suggested several morphological differences between northern and southern populations. This pattern was first interpreted as a multi-taxon situation by Lourenco (1982, 1994), with populations of different species and subspecies involved. This situation also illustrates the difficulties of clearly defining the biogeographic status of a given population. T. bahiensis is a very common species in the southeastern region of Brazil, and it represents an opportunistic species often living in disturbed habitats. Differences in the patterns of body pigmentation of individuals belonging to what appeared to be two 434 W. R. LOURENCO different allopatric populations first led to the recognition of two different species (Lourenco, 1982): T. bahiensis with a southern distribution and 7. eickstedtae Lourengo, with a northern distribution. Moreover, additional data from the contact zone between the two populations indicated what could be interpreted as hybrid forms. More detailed studies, however, later suggested the existence of a single polytypic species (Lourenco, 1986) with two sub-species (and two sub-populations), one in the south (T. bahiensis bahiensis), the other in the north (T. bahiensis eickstedtae). 3. The biogeographic pattern presented by Tiryus adrianoi sp. n. and by Tityus blaseri Mello-Leitao, suggests that these are allopatric species. The two species inhabitat similar landscape formations of the type Campo rupestre which, however, belong to different categories according to Eiten (1978, 1982 — see also paragraph “Environmental characteristics of the type locality”). Besides, these two areas are separated by almost 1000 km of typical Cerrado vegetation which acts as a barrier. This present pattern of geographical separation of the two species could suggest that in past geological times a contact zone existed between the two populations. The present pattern of distribution of the two species may be the result of the palaeoclimatological vicissitudes in tropical South America during the late Cenozoic and the Pleistocene (Lourenco, 1996). ACKNOWLEDGEMENTS I am very grateful to Dr Peter Schwendinger, Natural History Museum of Geneva, for reviewing the manuscript. REFERENCES EITEN, G. 1978. A sketch of the vegetation of Central Brazil. Resumos dos trabalhos, II Congresso Latino-Americano de Botanica, Brasilia, 1978: 1-37. EITEN, G. 1982. Brazilian “Savannas” (pp. 25-47). In: HUNTLEY, B.J. & WALTER, B.H. (eds). Ecological Studies, vol. 42: Ecology of Tropical Savannas. Springer-Verlag, Berlin, Heidelberg, New York. LOURENCO, W. R. 1980. Contribution a la connaissance systématique des Scorpions appartenant au “complexe” Tityus trivittatus Kraepelin, 1898 (Buthidae). Bulletin du Museum na- tional d'Histoire naturelle, Paris 4e sér., 2 (A-3): 793-843. LOURENCO, W. R. 1981. Sur la systématique des scorpions appartenant au complexe Tityus stig- murus (Thorell, 1877) (Buthidae). Revista brasileira de Biologia 41 (2): 351-362. LOURENCO, W. R. 1982. La véritable identité de Tiryus bahiensis (Perty, 1834). Description de Tityus eickstedtae n. sp. (Scorpiones, Buthidae). Revue arachnologique 4: 93-105. LOURENCO, W. R. 1986. Les modeles de distribution géographique de quelques groupes de scor- pions néotropicaux. Compte-rendu des Séances de la Société de Biogéographie 62 (2): 61-83. LOURENCO, W. R. 1994. Biogeographic patterns of Tropical South American scorpions. Studies on Neotropical Fauna and Environment 29 (4): 219-231. LOURENCO, W. R. 1996. The biogeography of scorpions. Revue suisse de Zoologie, vol. hors se- rie Il: 437-448. LOURENCO, W. R. 2001a. The Brazilian scorpion Tityus stigmurus (Chelicerata, Buthidae) and its complex of morphos. A new model is needed. Biogeographica 77 (1): 21-34. LOURENCO, W. R. 2001b. Sur les pas de Jean A. Vellard. A propos de sa contribution a l’étude des scorpions (Chelicerata). Revista Iberica de Aracnologia 3: 25-36. A NEW SPECIES OF TITYUS FROM BRAZIL 435 LOURENCO, W. R. 2002. Scorpions of Brazil. Les Editions de I’ IF, Paris, 320 pp. LOURENCO, W. R. & CLOUDSLEY-THOMPSON, J. L. 1996. Effects of human activities on the envi- ronment and the distribution of dangerous species of scorpions (pp. 49-60). In: BON, C. & GOYFFON, M. (eds). Envenomings and their treatments. Editions Fondation Marcel Merieux, Lyon. LOURENCO, W. R., CLOUDSLEY-THOMPSON, J. L., CUELLAR, O., EICKSTEDT, V. R. D., BARRAVIERA, B. & Knox, M. B. 1996. The evolution of scorpionism in Brazil in recent years. The Journal of Venomous Animals and Toxins 2 (2): 121-134. LOURENCO, W. R., Knox, M. B. & MAGALHAES, E. D. 1997. Redescription of Tityus blaseri (Scorpiones, Buthidae) from Goiäs Brazil. Revista de Biologia Tropical 44 (4): 1579- 1582. VACHON, M. 1963. De l’utilité, en systématique, d’une nomenclature des dents des chélicères chez les Scorpions. Bulletin du Museum national d’ Histoire naturelle, Paris 2e sér., 35 (2): 161-166. VACHON, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum national d Histoire naturelle, Paris 3e sér., 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 281 (D): 1597-1599. ar pa = Ù # HUE REVUE SUISSE DE ZOOLOGIE 110 (2): 437-452; juin 2003 Amara pulpani Kult, 1949 - eine valide Art in den Ostalpen (Coleoptera: Carabidae) Wolfgang PAILL Ökoteam, Institut für Faunistik und Tierökologie, Bergmanngasse 22, A - 8010 Graz. E-mail: oekoteam@sime.com Amara pulpani Kult, 1949 - a valid species in the Eastern Alps (Coleo- ptera: Carabidae). - Based on extensive material from the Eastern Alps, the taxonomic position of Amara pulpani Kult, 1949, formerly often discussed and mostly considered synonymous with Amara communis (Panzer, 1797), is investigated. Based on morphological and ecological features the two taxa are considered to represent distinct species. The distri- bution of A. pulpani, recorded from Austria, Italy, Slovenia and Croatia for the first time, is shown. The following nomenclatural decisions are made; Amara communis var. alpicola Heer, 1837 = nomen dubium (contra Hieke, 1995); Amara communis var. aenea Heer, 1837 = Amara nitida Sturm, 1825, nov. syn. (contra Hieke, 1995). Key-words: Coleoptera - Carabidae - Amara - taxonomy - ecology - Alps. 1. EINLEITUNG Untersuchungen zur Systematik des Amara communis-Komplexes wurden bere- its in größerer Zahl durchgeführt. Doch selbst ohne Berücksichtigung der zahlreich beschriebenen Synonyme (vgl. Hieke, 1995) zieht sich ein roter Faden interpretativer Widersprüchlichkeiten um die Stellung einiger Taxa durch die wissenschaftliche Literatur des 20. Jahrhunderts. Die heute auf breiter Basis anerkannte Differenzierung von Amara communis (Panzer, 1797) und Amara convexior Stephens, 1828 wurde noch vor wenigen Jahrzehnten in Frage gestellt; selbst der ausgezeichnete Carabiden- Systematiker Karl Lindroth synonymisierte beide Taxa (Lindroth, 1943), um einige Jahre später seinen Fehler einzugestehen (Lindroth, 1953). Der Status der beiden weiteren Arten des communis-Komplexes ist jedoch nach wie vor umstritten. Hieke (1973, 1978) stellte Amara pulpani Kult, 1949 und Amara makolskii Roubal, 1923 als Synonyme zu A. communis, während Hejkal und Hürka (in Hürka, 1996), aber auch bereits Gersdorf & Kuntze (1957) sowie Burakowski (1967) Bestimmungsschlüssel für alle vier Taxa publizierten, ohne jedoch zufriedenstellende Abbildungen zur Differenzierung der Arten zu liefern. Meldungen von A. pulpani außerhalb Tschechiens und der Slowakei fanden bisher keinen Eingang in die faunis- tische Literatur (z. B. Bonadona, 1971; Kryzhanovskij et al., 1995; Trautner er al., 1997). Manuskript angenommen am 16.10.2002 438 W. PAILL Neueste Erkenntnisse auf dem Gebiet der vergleichenden Ei- und Larval- morphologie unterstützen die von Hurka (1996) vorgelegten Interpretationen (Hürka & Ruzickova, 1999). Dem wird von Kritikern jedoch nach wie vor die hohe Variabilität imaginaler Merkmale entgegengehalten (mündl. Mitt. Hieke). Anhand von Belegtieren zweier kürzlich entdeckter österreichischer Populationen von A. pulpani sollen nun weitere Differenzierungsmerkmale beschrieben und dargestellt werden. Diese sind geeignet, um den Artstatus von A. pulpani zu untermauern sowie weitere Hinweise zur Bestimmbarkeit zu liefern. Zusätzlich werden Daten zur Verbreitung von A. pulpani präsentiert und durch Angaben zum Lebensraum, der saisonalen Aktivität sowie zur Begleitfauna ergänzt. 2. MATERIAL UND METHODEN 2.1. FUNDORTE Das untersuchte Material von A. pulpani stammt von nachfolgend aufgelisteten Lokalitäten. Soweit nicht anders erwähnt, befinden sich alle Tiere am Museum für Naturkunde der Humboldt-Universität Berlin. Österreich: Kärnten, Karawanken, SW Eisenkappel, Trögerner Klamm (PU1, 46°27/N, 14°29°E, 850 m), Schneeheide-Kiefernwald, Barberfallen: 12.5.1999-5.8.1999, 14.10.1999-14.11.1999, 14.11.1999-10.9.2000 (7 Fallen, jeweils mehrere Fang- perioden), 14 88,13 2 2,2 Larven leg., det. & coll. Paill, 1 6 6,3 2 2 im Muséum d’histoire naturelle, Genf. Kärnten, Gailtal, NW Kötschach/Mauthen, Mussen (PU2, 46°42°N, 12°54 E, 1850-1900 m), Barberfallen: 1.7.2000-26.8.2000 (2 Fallen, 1 Fang- periode), Handfang: 14.9.2000, 11 dd, 13 9 2, leg. Komposch, Komposch & Paill, det. & coll. Paill, 3 dd, 1 22 im Muséum d’histoire naturelle, Genf. Kärnten, Karawanken, Rosenbach, 29.5.1931, 1 d, leg. Wirthumer, det. Hejkal. Kärnten, Karawanken, Koschuta, Waidischbach, 12.6.1963, 1 d, leg. Kirch, det. Hejkal. Nieder- österreich, Lunz, Lunzberg, 1.7.1966, 1 4, 1 9, leg Malicky, det. Paill. Oberösterreich, Steyr, Schoberstein, 23.5.1909, 1 d, leg. Petz, det. Hejkal. Oberösterreich, Linz, 20.5.1923, 1 d,1 ©, leg. Kogler, det. Paill. Oberösterreich, Engelhartszell, Nieder- ranna, 1893, 1 ©, leg. Rupertsberger, det. Paill. Tirol-Stubaital, Telfeser Wiesen, 22.6.1975, 1 2, det. Paill. Deutschland: Bayern, Berchtesgaden, Watzmann (700 m), 7.9.1994, 1 9, leg., det. & coll. Hurka. Bayern, Fränkische Schweiz, Umgebung Pottenstein, 14.-18.8.1978, 1 d, leg. Zwick, det. Paill. Slowenien: Nova Gorica, Ternowaner Wald, 1902, 1 G, leg. Ganglbauer, det. Paill. Postojna, Nanos, 20.5.1923, 1 ©, leg. Lona, det. Paill. Remschnigg, Sp. Kapla (46°39°N, 15° 24 E, 900 m), 16.5.2001, 1 9, leg., det. & coll. Paill. AMARA PULPANI IN DEN OSTALPEN 439 Kroatien: Istrien, Utka, 26.5.1965, 2 9 ©, leg. Freude, det. Paill. Istrien, Ucka, 2.- 8.6.1931, 1 2, leg. Stöcklein, det. Paill. Rijeka, ViSevica (1200 m), 4.6.1939, 1 6, leg. Svircev, det. Hejkal. Rijeka, Risnjak (1500 m), 8.7.1962, 1 9 , leg. Malicky, det. Paill. Velebit, Senj, Krasanska duliba, 21.4.1979, 1 d, leg. Feller, det. Hejkal. Italien: Friaul Julisch Venetien, Cividale, Monte Matajur, 5.7.1922, 1 9, leg. Schatz- mayr, det. Paill. Friaul Julisch Venetien, Pontebba, Monte Cavallo (1570 m), 7.1972, 1 3, leg. Schmidt, det. Hejkal. Friaul Julisch Venetien, Tolmezzo, Monte Festa, 4.9.1962, 1 ©, leg. Springer, det. Paill. Friaul Julisch Venetien, Tolmezzo, Monte Festa, 23.6.1968, 1 d, leg. Springer, det. Hejkal. Friaul Julisch Venetien, Lago di Gavazzo (Monte Festa, 500-600 m), 16.6.1957, 1 d, leg. Springer, det. Paill. Friaul Julisch Venetien, Tarcento, Monte Bernadia, 11.5.1952, 1 2, leg. Busulini, det. Paill. Friaul Julisch Venetien, Lusevera, Musi (650 m), 17.6.1982, 1 2, leg. Delfabbro, det. Paill. Friaul Julisch Venetien, Maniago, Monte Raut (S-Seite, 1000 m), 7.1976, 1 ®, leg. Schmidt, det. Paill. Friaul Julisch Venetien, Passo della Mauria, 20.6.1956, 1 6, leg. Fabris, det. Hejkal. Südtirol, Dolomiten, 6.1955, 1 ?, leg. Vogt, det. Paill. Südtirol, Kastelruth, Seis am Schlern, 1.-13.6.1964, 1 à , leg. Abraham & Deeming, det. Hejkal. Südtirol, Bozen, 19.5.1890, 1 &, det. Paill. Südtirol, Bozen, Val d’arsa, 22.6.1923, 1 2, det. Paill. Südtirol, Grödnertal, St. Ulrich, 6.6.1962, 2 © 2, leg. Kirch, det. Paill. Südtirol, Valle del Fèrsina, Palai, 1 9, det. Paill. Lombardei, Gardasee, Tremosine, Vall di Bondo (600 m), 6.-7.6.1974, 1 9, leg. Schawaller, det. Paill. Drei Populationen von A. communis lieferten Vergleichstiere; vom steirischen Fundort (CO2) wurden außerdem Tiere von Amara lunicollis Schiödte, 1837 zur Vermessung der Flügel herangezogen. Österreich, Kärnten, Ossiacher See, Bleistätter Moos (COI, 46°41°N, 14°01°E, 510 m), 38 Exemplare. Österreich, Steiermark, W Liezen, Wörschacher Moos (CO2, 47°33’N, 14°11’E, 640 m), 32 Exemplare, 1 Larve. Deutschland, Baden-Württemberg, Stuttgart, Sindelfingen (CO3), 20 Exemplare. 2.2. MORPHOMETRIE UND GENITALMORPHOLOGIE Die Körpermaße von A. pulpani wurden mit Hilfe eines Okularmikrometers ermittelt und mit jenen der nahe verwandten A. communis sowie im Falle der Flügel- ausbildung zusätzlich mit A. lunicollis verglichen. Die Länge der Elytren wurde von der Flügeldeckenbasallinie bis zur Flügeldeckenspitze gemessen, die häutigen Flügel wurden abpräpariert, aufgespannt und an ihrer breitesten Stelle sowie als leicht stan- dardisierbares Maß ihrer Länge von der Adernaufzweigung zwischen den Haupt- längsadern Radius und Cubitus bis zur Flügelspitze vermessen (vgl. Abb. 4). Die männlichen Genitalien wurden präpariert und in Kanadabalsam eingebettet. Die äußere Form des Aedeagus, die Innenstrukturen und die Ausbildung der linken Paramere wurden analysiert und den entsprechenden Ausbildungen bei A. communis gegenübergestellt. Im weiblichen Geschlecht erfolgte das Studium äußerer Merkmale am 8. und 9. Abdominalsegment. 440 W. PAILL Die Studien zur Genitalmorphologie, Flügelausbildung und zur Form des Halsschildvorderrandes basieren auf Tieren (5 & d und 5 © © je Population) aus den beiden aktuell in Österreich aufgefundenen Populationen (PU1, PU2). Die weiteren Vermessungen sowie die Registrierung von Färbungs- und Punktierungsmerkmalen wurden hingegen an allen aufgezählten Tieren durchgeführt. 3. ERGEBNISSE UND DISKUSSION 3.1. MORPHOLOGISCHE MERKMALE 3.1.1. Körperlänge Die durchschnittliche Größe der männlichen A. pulpani lag bei 7.3 mm (6.3-8.1 mm), jene der weiblichen Tiere bei 7.7 mm (6.4-8.5 mm). Eine Häufigkeitsverteilung der Längenmaße geht aus Abb. 1. hervor. Von minimaler Körperlänge erwiesen sich ein Männchen mit 6.3 mm und ein Weibchen mit 6.4 mm, maximale Körperlänge erreichte ein 8.1 mm messendes Männchen bzw. ein Weibchen mit 8.5 mm. Während Kult (1949) bzw. Burakowski (1967) eine Körpergröße von 6.1-7.2 mm angeben, entspricht die von Hürka (1996) ermittelte Spanne von 5.9-8.1 mm etwa den im Rahmen der vor- liegenden Arbeit vermessenen Tieren (Abb. 1). Deutliche Unterschiede konnten zwischen den Tieren der beiden näher unter- suchten Populationen festgestellt werden. Bei durchschnittlich 7.4 mm gegenüber 7.1 mm im Falle der Männchen (t=2.7; p<0.05) und 8.1 mm gegenüber 7.2 mm bei den Weibchen (t=7.6; p<0.01) waren die Tiere der hochmontanen Population (PU1) signifikant größer als jene der subalpinen (PU2). Letztere Tiere waren damit so groß bzw. klein wie durchschnittliche Vertreter von A. communis. 3.1.2. Körperform Der Körperumriss von Amara s. str. ist äußerst uniform. So unterscheiden sich A. pulpani und A. communis weder hinsichtlich des Quotienten aus Flügeldeckenlänge und -breite (1.37 bei A. pulpani bzw.1.35 bei A. communis), aus Flügeldeckenlänge und Halsschildlänge (2.52 bei A. pulpani bzw. 2.55 bei A. communis) sowie aus Flügel- deckenbreite und Halsschildbreite (1.08 bei beiden Arten). Signifikante Unterschiede sind jedoch im Breiten-Längen-Verhältnis des Halsschildes zu finden, welches im Falle von A. pulpani durchschnittlich 1.69 und bei A. communis 1.75 beträgt (t=6.23; p<0.01). Kult (1949) nennt für A. pulpani sogar einen Wert von 1.6. Differential- diagnostisch bedeutend ist schließlich die Form des Halsschild-Vorderrandes. Die zur Differenzierung von A. convexior und A. communis relevante Abwinkelung der Vor- derecken (Hieke, 1973) ist nämlich auch zwischen A. communis und A. pulpani signifikant unterschiedlich (t=20.32; p<0.01). Zwar liegt der Mittelwert für A. commu- nis mit 34.9° aufgrund möglicherweise etwas anderer Meßmethode 5° über dem von Hieke (1973) angegebenen Wert, doch ist der Unterschied zu A. pulpani von etwa 10° bzw. 30% jedenfalls als konstant anzunehmen (Abb. 2). AMARA PULPANI IN DEN OSTALPEN 441 Individuen i SN N oO on oO O1 6 6.5 7 7.5 8 8.5 9 Körperlänge [mm] ABB. | Häufigkeitsverteilung der Körperlängen bei männlichen und weiblichen Amara pulpani. & > on (©) Winkel [°] & oO 25 ABB. 2 Winkel der Halsschildvorderecken bei Amara pulpani (Populationen PU1, PU2) im Vergleich zu A. communis (CO1, CO2, CO3). 3.1.3. Färbung Sowohl die Färbung der Körperoberfläche (2 der 86 normalerweise kupfer- färbigen A. pulpani zeichneten sich durch bläulichen Glanz aus) als auch die Aus- dehnung der Aufhellung der Antennenbasis bilden keine geeigneten Unterscheidungs- merkmale zwischen beiden Taxa. So ist das dritte Antennenglied bei A. pulpani zwar meist stärker verdunkelt als bei A. communis, doch treten bei der hinsichtlich dieses Merkmales sehr variablen A. communis auch Exemplare mit fast gänzlich verdun- keltem dritten Glied auf (vgl. Hieke, 1978, 1988). Auch der Färbung der Schienen kann 442 W. PAILL entgegen den Angaben von Kult (1949) und Burakowski (1967) kein differenzial- diagnostischer Wert beigemessen werden, da kein Unterschied im Ausmaß der braun- rötlichen Aufhellung im Vergleich zwischen A. pulpani und A. communis feststellbar war. 3.1.4. Flügelausbildung Die Länge der Hinterfliigel von A. pulpani entspricht etwa der Länge der Flügeldecken. Weder die Geschlechter noch die beiden untersuchten Populationen un- terschieden sich in diesem Merkmal; die Relation zwischen Flügel- und Elytrenlänge bei den Tieren aus der Trögerner Klamm betrug 0.98 und bei jenen von der Mussen 1.0. Auch die relative Breite der Flügel war mit Werten von 1.16 (PU1) und 1.17 (PU2) zwischen beiden Populationen identisch (Abb. 3). Vergleiche mit A. communis offenbarten einen hochsignifikanten Unterschied in der Flügelausbildung, der als differenzialdiagnostisches Merkmal zur Unterscheidung beider Arten zumindest im mitteleuropäischen Raum anwendbar ist (Abb. 3, 4). Demnach sind die Flügel von A. communis nicht nur in Relation zur Körpergröße deut- lich länger (t=27.46; p<0.01) und breiter (t=24.82; p<0.01) als bei A. pulpani, sondern auch aufgrund qualitativer Unterschiede (stärkere Sklerotisierung, größeres Analfeld) offenbar besser für die Fortbewegung per Flug geeignet. Auch die von Drioli (1987) vermessenen A. communis wiesen Hinterflügel von 1.5-1.6 facher Elytrenlänge auf. Dennoch scheint A. communis trotz zahlreicher Nachweise (z. B. van Huizen, 1980) nur selten tatsächlich flugfähig zu sein; diese Individuen besitzen neben einer gut aus- gebildeten Flugmuskulatur, die sonst oft degeneriert ist (Tietze, 1963), auch längere Flügel (Desender, 1989). 3.1.5. Punktierung und Beborstung Der Halsschild von A. pulpani ist in den meisten Fällen relativ schwach punk- tiert. Allerdings ist dieses tendenziöse Merkmal nicht geeignet, um eine sichere Differenzierung von der zumeist deutlich kräftiger punktierten A. communis vor- zunehmen; immerhin treten bei beiden Arten Individuen mit weitgehend fehlender bis sehr kräftiger Punktierung auf (Abb. 2). Die Ausbildung der Porenpunkte am Ende des 7. Flügeldeckenstreifens ist hingegen von größerem Interesse. Die Anzahl dieser Punkte hat sich nämlich trotz hoher Variabilität zu einem allgemein anerkannten Merkmal in der Differenzial- diagnose innerhalb der Gattung Amara etabliert (z. B. Lindroth, 1943). Hürka (1996) führt das Auftreten von zwei bei A. pulpani gegenüber drei Punkten bei A. communis als differentialdiagnostisches Merkmal zwischen den beiden Taxa an, während Hieke (1973) den Wert dieses Merkmales anhand seiner Analysen bei mehreren Populationen von A. communis und auch des Typenmaterials von A. pulpani kritisch hinterfragt. Die eigenen Untersuchungen zeigten zwar den bedeutenden Wert dieses Merkmals, konnten jedoch auch deren Variabilität festhalten. Demnach kann es bei beiden Arten zu Veränderungen der Borstenzahlen kommen, die aber zumeist nur einseitig ausfallen. A. pulpani scheint eine wesentlich höhere Konstanz dieses Merkmales zu besitzen, nachdem lediglich eines der 86 untersuchten Individuen eine einseitig erhöhte Porenzahl aufwies. Demgegenüber ist die Porenzahl bei A. communis als vergleichs- AMARA PULPANI IN DEN OSTALPEN 443 OT. È : | i TD © 14 © rel. L & © & © L À Sii oy + de) iL 0.8 ABB. 3 Relative Flügellängen (rel. L) und -breiten (rel. B) bei Amara pulpani (Populationen PU1, PU2) und A. communis (CO1, CO2, CO3). Als Vergleich dient die häufig mit A. communis verge- sellschaftete A. /unicollis (LU). ABB. 4 Rechter Hinterflügel von Amara pulpani (a) und A. communis (b). Maßstab = 1 mm. Die Flügeldecken beider Tiere waren mit 4,0 mm von identischer Länge. Die Klammer verdeutlicht die Methode der Längenmessung. 444 W. PAILL weise variabel zu bezeichnen, zumal nicht weniger als 13% der 90 untersuchten Individuen Abweichungen von der „Norm“ zeigten. 10 Tiere hatten einseitig verän- derte Borstenzahlen (2 Individuen mit 4 und 8 Individuen mit 2 Borsten) und bei zwei Individuen waren an beiden Flügeldecken nur noch zwei Porenpunkte sichtbar, die bei alleiniger Verwendung dieses Merkmales zu Verwechslungen mit A. pulpani führen könnten. Berücksichtigt man jedoch die Tatsache, dass der Abstand zwischen dem größten, am nächsten zur Flügeldeckenbasis gelegenenen Porenpunkt und dem mittel- großen, zweiten Porenpunkt, bei A. pulpani wesentlich größer als bei A. communis ist (Abb. 5), so läßt sich eine mögliche Reduktion des kleinsten, apikalen Porenpunktes bei A. communis relativ leicht erahnen. Zu bedenken gilt zusätzlich, dass das Auffinden des dritten Porenpunktes bei abgebrochener Borste nicht nur aufgrund seiner geringen Größe sondern auch wegen der nicht selten veränderten Lage (vgl. Lindroth, 1943) am apikalen Ende des ersten oder zweiten Flügeldeckenstreifens schwer fällt. 3.1.6. Genitalmorphologie Innerhalb der Gattung Amara sind die Genitalien wenig differenziert. Dennoch ermöglicht die hohe Konstanz der Merkmale deren Einsatz in der Differenzial- diagnostik. Der Aedeagus von A. pulpani ist geringfügig größer als jener von A. communis und sein Apex verjüngt sich abweichend von der leicht konischen Divergenz bei A. communis fast parallel (Abb. 6, 7). Dabei liegt das Apex-Längen-Breiten Verhältnis beider Arten bei durchschnittlich 1.5 und entspricht damit den von Hürka (1996) angegebenen Maßen. Im Innensack fällt die Chitinisierung bei ausgehärteten Tieren deutlich kräftiger als bei A. communis aus. Die linke Paramere ist etwas länger und die Spitze als etwas kräftigerer Haken differenziert. Dieser liegt bei A. pulpani in der vd ABB. 5 Charakteristische Borstenstellung am Ende des 7. Fliigeldeckenstreifens bei Amara pulpani (a) und A. communis (b). Maßstab = 1 mm. AMARA PULPANI IN DEN OSTALPEN 445 ABB. 6 Genitalmorphologie männlicher Amara pulpani. Aedeagus in Aufsicht, linke Paramere in Seitenlage, “Genitalring”. Maßstäbe (Aedeagus groß, Paramere und Genitalring klein) = 1 mm. LO ABB. 7 Genitalmorphologie männlicher Amara communis. Aedeagus in Aufsicht, linke Paramere in Seitenlage, “Genitalring”. Maßstäbe (Aedeagus groß, Paramere und Genitalring klein) = 1 mm. Ebene der seitlich abgeflachten Paramere, während er bei A. communis deutlich zur Seite abgewinkelt ist und in der Abbildung daher vergleichsweise klein erscheint. Schließlich ist auch der als “Genitalring” differenzierte Teil des 9. Abdominalsegments im basalen Teil kräftiger ausgebildet und kann trotz relativ großer Variabilität als Differenzialmerkmal zwischen den beiden Arten gelten. Das weibliche Genital liefert ebenfalls diffizile Unterschiede zwischen beiden Arten. Zwar ist die zur Differenzierung von A. communis und A. convexior her- anziehbare Form der Gonocoxite (Hieke, 1976) zwischen A. communis und A. pulpani nur geringfügig abweichend. Auffallend ist jedoch die unterschiedliche Beborstung des apikalen Häutchens (Ventrit) am 9. Abdominalsegment: diese ist bei A. pulpani durch wesentlich kräftigere Borsten als bei A. communis gekennzeichnet. Hinzu kommt die unterschiedliche Form in der Sklerotisierung am 8. Abdominaltergit (Abb. 8, 9). 446 W. PAILL ABB. 8 Genitalmorphologie weiblicher Amara pulpani. Gonocoxite und Ventrit (8. Abdominalsegment), Sklerotisierung am 9. Abdominalsegment. Maßstab = 0,5 mm. ABB. 9 Genitalmorphologie weiblicher Amara communis. Gonocoxite und Ventrit (8. Abdominal- segment), Sklerotisierung am 9. Abdominalsegment. Maßstab = 0,5 mm. 3.2. PHÄNOLOGIE Die über einen Zeitraum von zwei Jahren mit Hilfe von Barberfallen unter- suchte Population CO1 in der Trögerner Klamm in 850 m Seehöhe bot die Gelegenheit, standardisierte Daten zur jahreszeitlichen Aktivität von A. pulpani zu gewinnen. In beiden Jahren trat A. pulpani bereits sehr früh im Jahr auf, erreichte im Mai bzw. Anfang Juni die maximale Laufaktivität und wurde danach kaum noch im imaginalen Stadium nachgewiesen. Der sommerlarvale Entwicklungsmodus konnte durch den Fund zweier Larven im Juli des zweiten Untersuchungsjahres sowie zweier immaturer, frisch geschlüpfter Käfer im darauffolgenden August bestätigt werden (Abb. 10). Korrespondierende Angaben zur Saisonalität von A. pulpani macht Kult (1949). Mit zunehmender Höhenlage dürften Erstauftreten und Populationsmaximum in den Hochsommer verschoben sein, wie die Erhebungen auf der Mussen (PU2) in 1900 m Seehöhe andeuten. Immature Käfer wurden hier erst im September gefangen. AMARA PULPANI IN DEN OSTALPEN 447 3.3. LEBENSRAUM UND HÖHENVERBREITUNG Die Standorte der beiden untersuchten Populationen von A. pulpani sind durch extrem trockene und auch thermisch günstige Verhältnisse gekennzeichnet. In der strukturreichen Trögerner Klamm bewohnt die Art ausschließlich steile, südexponierte Flanken. Die Standorte sind durch geringe Bodenbildung und mäßige Nährstoffver- sorgung gekennzeichnet. Die in der Baumschicht mehr oder weniger lückigen Schnee- heide-Kiefernbestände (Erico-Pinetum sylvestris) sind durch mosaikartig eingestreute vegetationslose Offenstellen mit Dolomitgrus-Auflage (kleinflächige Schuttfluren) gekennzeichnet. Auch auf der Mussen bewohnt A. pulpani vegetationsarme Schutt- fluren. Diese liegen hier inmitten ausgedehnter, traditionell bewirtschafteter skelett- bodenreicher Kalkmagerrasen (Seslerio-Caricetum sempervirentis) im Bereich besonders steiler oder anthropogen beeinflusster (Mahd, Weganschnitte) Stellen. Neben der Schneeheide (Erica carnea) erreichen Alpen-Steinquendel (Acinos alpinus), Hufeisenklee (Hippocrepis comosa), Wilde Nelke (Dianthus sylvestris), Alpen- Labkraut (Galium anisophyllum), Kriech-Quendel (Thymus praecox), Alpenhelm (Bartsia alpina) und Herzblatt-Kugelblume (Globularia cordifolia) höhere Deckung (Wieser & Komposch, 2002). Die Lebensräume der beiden österreichischen Populationen korrespondieren mit den Angaben aus Tschechien und der Slowakei, wo A. pulpani in trockenen Biotopen wie Schutthalden, Heiden (oftmals Calluna), Waldsteppen und Waldrändern auftritt (Kult, 1949; Hürka, 1996). Indifferenz besteht offenbar hinsichtlich der Deckung der Baumschicht. In den genannten Ländern gilt die Art als Verteter der Gebirgsregion, was ebenfalls gut zu den aktuellen Funden aus den Alpen passt. Die Population von der Mussen mit einzelnen Tieren aus 1900 m dürfte sogar den bisher höchsten Fund von A. pulpani darstellen. Die stenöke A. pulpani unterscheidet sich im ökologischen Verhalten grund- legend von A. communis. Letztere Art hat ein deutlich breiteres Lebensraumspektrum und kommt überwiegend auf frischen bis mäßig feuchten Wiesen und Hochstauden vor. Nur selten werden leicht bewaldete Standorte und ausgesprochen trockene Biotope be- siedelt (z. B. Dülge et al., 1994; Marggi, 1992; Tietze, 1973; Turin, 2000; Lindroth, 1986; Luff et al., 1992). Außerdem vikariiert A. communis offenbar auch hinsichtlich der bewohnten Höhenstufe mit A. pulpani. So kennt Marggi (1992) in der Schweiz kaum A. communis-Populationen oberhalb von 600 m und in Südosteuropa werden Höhengrenzen von 1400 m genannt (Hieke & Wrase, 1988; Guéorguiev & Guéorguiev, 1995). Allerdings sind diese Angaben zu hinterfragen, zumal es sich bei den Meldungen aus höheren Lagen durchaus um A. pulpani handeln könnte. Konkrete Vermutung dafür besteht jedenfalls für Tiere, die vom Glocknermassiv in den Hohen Tauern aus über 1600 m Höhe gemeldet wurden (Franz, 1943; Jung, 1981) und ebenso für Individuen aus Felsenheiden und subalpinen Rasen in den Südalpen (Brandmayr & Zetto Brandmayr, 1988). 448 W. PAILL D (cb) .£ 9 10 je (= o LI 2000 1999 2000 1999 2000 Frühjahr Sommer Herbst ABB. 10 Jahrezeitliches Auftreten (1999 und 2000) von Amara pulpani in der Trégerner Klamm. Imagines (schwarze Balken), Larven (weißer Pfeil) und immature Käfer (grauer Pfeil). 3.4. BEGLEITFAUNA An beiden österreichischen Standorten zählte A. pulpani (zumindest über den jeweils bearbeiteten Zeitraum) zu den häufigsten Laufkäferarten. Im Schneeheide- Kiefernwald in der Trögerner Klamm stellte A. pulpani sogar 50% der gefangenen Individuen der insgesamt aus 8 Arten bestehenden arten- und individuenarmen Lauf- käferzönose. Als weitere autochthone Arten traten unter anderem Carabus cancellatus dahli Heer, 1841, Carabus arcensis Herbst, 1784, Molops ovipennis Chaudoir, 1842 und Abax ovalis (Duftschmid, 1812) auf. In der Schuttflur auf der Mussen erreichte A. pulpani 40% der Aktivitäts- dominanz und wurde nur von Harpalus solitaris Dejean, 1829 als häufigste Art der Zönose übertroffen. Auch hier wird die Gemeinschaft aus wenigen weiteren subrezen- denten Vertretern wie Carabus germarii Sturm, 1815 und Prerostichus jurinei (Panzer, 1803) gebildet. An keinem der beiden Standorte konnte eine Vergesellschaftung von A. pulpani mit A. communis, noch mit einer anderen Amara-Art, sieht man von einem Einzeltier von A. convexior am Standort PU1 ab, festgestellt werden. 3.5. VERBREITUNG Im Nordosten Mitteleuropas ist die Verbreitung von A. pulpani aufgrund der in Tschechien und der Slowakei durchgehenden Anerkennung als eigenständiges Taxon relativ gut bekannt. Hier ist die Art zwar relativ selten aber weit verbreitet (Kult, 1949; Hürka, 1996). Darüberhinaus waren bisher nur vereinzelte gesicherte Nachweise bekannt. So erwähnt Kult (1949) in seiner Beschreibung weiteres Material aus Deutschland, Frankreich (ohne genaue Lokalisierung) und der Ukraine (Karpaten, Galicia: Lwöw). Horion (1951) gab A. pulpani auch für Polen an, offenbar in fälsch- AMARA PULPANI IN DEN OSTALPEN 449 licher Zuordnung der Meldung aus “Galicia” (vgl. Burakowski et al., 1974), und aus Österreich, ohne jedoch auf Fundorte oder Belegtiere einzugehen. Anhand des vorliegenden Materials wird A. pulpani nun erstmals für Österreich, Slowenien, Kroatien und Italien sowie nach Kult (1949) und Gersdorf & Kuntze (1957) bzw. Horion (1960) abermals für Deutschland gemeldet (Abb. 11). Eine weitere Verbreitung der mittel-südosteuropäischen Art im nördlichen Alpenvorland bis Mittel- deutschland (mündl. Mitteilung H. Terlutter) und in den Gebirgszügen des Balkans ist wahrscheinlich. ABB. 11 Verbreitung von Amara pulpani. Mit Ausnahme der zwar in der Literatur genannten, aber nicht exakt lokalisierbaren Funde sind alle bekannten Vorkommen verzeichnet; die Häufung an Nachweisen in Tschechien und der Slowakei ist durch ein helles Oval angedeutet. Der Typus- Fundort ist durch ein schwarzes Quadrat und die aktuell untersuchten Populationen in Österreich (PU1 und PU2, siehe Text) sind durch große Kreise symbolisiert. 3.6. DETERMINATION Die sichere Bestimmung von Einzeltieren von A. pulpani kann Schwierigkeiten bereiten. Jedenfalls müssen mehrere Merkmale, nämlich zumindest Anzahl und Lage der Porenpunkte am terminalen Ende des 7. Flügeldeckenstreifen, der Winkel des Halsschildvorderrandes und die relativen Flügelmaße berücksichtigt werden. Ein dichotomer Bestimmungsschlüssel soll zum gegenwärtigen Zeitpunkt noch nicht dar- gestellt werden, zumal auch im zentralen mitteleuropäischen Raum mit dem Vor- kommen einer weiteren sehr ähnlichen Art, A. makolskii, zu rechnen ist. 3.7. NOMENKLATORISCHE ANMERKUNGEN Untersuchtes Material: Amara communis var. alpicola Heer, 1837 (ETH Zürich): etikettiert mit “Typus ?”, “f’ (Fundortkiirzel “f’ = Kanton Glarus laut Catalog 450 W. PAILL der Schweizer Sammlung Coleoptera, schriftl. Mitt. Schmid, ETH Zürich), “Amara (Amara) curta Dej. det. F. Hieke 1984”. Amara communis var. aenea Heer, 1837 (ETH Zürich): etikettiert mit “Syntypus”, “e” (Fundortktirzel “e” = Rheinwald laut Catalog der Schweizer Sammlung Coleoptera, schriftl. Mitt. Schmid, ETH Zürich), “Amara (Amara) nitida Sturm det. F. Hieke 1984”. Amara brunnicornis Heer, 1837 (ETH Zürich): Holotypus. Amara aubryi Schuler, 1964 (Musée Zoologique de Strasbourg): Holotypus und weitere von Schuler als A. aubryi determinierte Tiere. Hieke (1995) fiihrt in seinem Namensverzeichnis der Gattung Amara zahlreiche Synomyme von A. communis an. Die Giiltigkeit des derzeit verwendeten Namens von A. pulpani musste daher hinterfragt werden. Mit Amara communis var. alpicola Heer, 1837 wurde ein aufgrund der Fundumstände in subalpiner Lage am ehesten in Frage kommendes Taxon aus der Liste von Hieke untersucht. Allerdings fallt die Inter- pretation des einzig vorhandenen Tieres schwer. So enthält der mit “Typ ?” versehene Beleg ein sich auf den Kanton Glarus beziehendes Fundortkürzel, während Heer (1837) in der Originalbeschreibung Rheinwald und später präziser (Heer, 1841) Nufenen im Rheinwald (Anmerkung: im Kanton Graubünden auf etwa 1600 m Höhe gelegen) als Lokalität nennt. Nach Vorlage dieses möglicherweise nicht richtig zuge- ordneten Typus stellte sich außerdem heraus, dass das Tier von Hieke als Amara curta (richtig) determiniert und bezettelt worden war, ein Umstand der jedoch keine Fort- führung im zitierten Verzeichnis fand, wo Amara communis var. alpicola Heer, 1837 als Synonym von A. communis geführt wird. Amara communis var. alpicola Heer, 1837 kann nach derzeitigigem Wissensstand daher weder A. communis noch A. curta zuge- ordnet werden, sondern muss als nomen dubium abgelegt werden. Amara communis var. aenea Heer, 1837 ist gleichfalls kein Synonym von A. communis, wie fälschlicher- weise trotz anderslautender Revision des Lectotypus durch Hieke im Namens- verzeichnis geführt, sondern dem Taxon Amara nitida zuzuordnen. Überdies sind beide Namen jüngere Homonyme (von Amara alpicola Dejean, 1828 bzw. Amara aenea (De Geer, 1774) und daher nicht verfügbar. A. brunnicornis Heer, 1837 wurde aufgrund des “verdächtigen” Namens ebenfalls untersucht, ist jedoch ein Synonym von Amara curta, wie von Hieke (1995) bereits dargestellt. Die ebenfalls aus dem Rheinwald beschriebene Amara communis var. atrata Heer, 1837 fehlt in der Typensammlung Heer an der ETH Zürich (schriftl. Mitt. Schindler) und konnte daher leider nicht untersucht werden. Weitere Namen wurden auf Basis ausreichend begründeter Synonymisierungen mit A. communis (A. aemilina Fiori, 1903 und A. manevali Jeannel, 1942 nach Hieke, 1973; A. nigrita Chaudoir, 1844 und A. viatica Motschulsky, 1845 nach Hieke, 1975; Carabus vagabundus Duftschmid, 1812 und A. atrocoerulea Sturm, 1825 nach Schaum, 1846) oder aufgrund tiergeografischer Überlegungen (A. latescens Stephens, 1828, A. obtusa Stephens, 1828, A. plebeja Stephens, 1828, A. dalei Rylands, 1841 jeweils aus planaren Lagen in Großbritannien beschrieben; A. impressa Motschulsky, 1848 aus Kasachstan beschrieben) ausgeschieden. Die Synonymisierung von A. aubryi Schuler, 1964 mit A. communis (Hieke, 1988) konnte nach Vorlage des Holotypus und weiteren Materials bestätigt werden, nachdem auf Basis der von Hieke ausführlich beschriebenen Merkmale eine Syno- nymie mit A. pulpani nicht ausgeschlossen werden konnte. AMARA PULPANI IN DEN OSTALPEN 451 4. DANK Fiir zahlreiche wertvolle Hinweise bin ich Herrn Dr. Fritz Hieke, Berlin zu großem Dank verpflichtet. Er stellte mir wie auch Dr. Alexander Dostal, Wien umfang- reiches Material von A. pulpani zum Studium zur Verfügung; Dr. Karel Hürka, Prag teilte mir einen Fund aus Bayern mit; Dr. Christian und Mag. Brigitte Komposch, Graz betreuten die Barberfallen auf der Mussen. Vergleichstiere von A. communis aus einer deutschen Population erhielt ich dankenswerterweise von den Herren Jürgen Trautner und Dr. Michael Bräunicke, Filderstadt. Dr. Jose Matter, Strasbourg und Dr. Andreas Müller, Zürich liehen mir Typen-Material aus den jeweiligen Sammlungen; Dr. Charles Huber, Bern und Werner Marggi, Thun unterstützten mich bei der Literatur-Recherche. Joachim Schmidt und Dr. Christian Komposch verdanke ich die kritische Durchsicht des Manuskripts. Die laufkäferkundlichen Grundlagenerhebungen in den Naturschutzgebieten Trögerner Klamm, Mussen und Bleistätter Moos wurden von der Kärntner Landes- regierung Abt. 20/Unterabt. Naturschutz (Dr. Thusnelda Rottenburg & Dr. Christian Wieser, Klagenfurt) gefördert. 5. LITERATUR BONADONA, P. 1971. Catalogue des Coléoptères Carabiques de France. Nouvelle Revue d’Entomologie, Supplement, 177 pp. BRANDMAYR, P. & ZETTO BRANDMAYR, T. 1988. Communitä a coleotteri Carabidi delle Dolomiti Sudorientali e delle Prealpi Carniche. Studi Trentini Scienze Naturali, Acta Biologica Supplemento 64: 125-250. BURAKOWSKI, B. 1967. Biology, ecology and distribution of Amara pseudocommunis Burak. 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Naturwissenschaftlicher Verein für Kärnten, Klagenfurt, 290 pp. REVUE SUISSE DE ZOOLOGIE Tome 110 — Fascicule 2 EBERMANN, Ernst, HAJIQANBAR, Hamid-reza & HADDAD IRANI-NEJAD, Karim. News records of phoretic and soil-living mites from Iran (Acari Heterostigmata: Scutacaridae). anne in AE PACE, Roberto. Homalotini del Borneo (Coleoptera, Staphylinidae)...... LÔBL, I. & LESCHEN, R. A. B. Redescription and new species of Alexidia (Coleoptera: Staphylinidae: Scaphidimae) 457 + 0e et en = SCHELLER, UIf. New records of Pauropoda (Myriapoda) with descriptions of new species from Rwanda and Réunion (Pauropoda and Symphyla of ineGeneva NUTI IL FET, Victor, GANTENBEIN, Benjamin, SOLEGLAD, Michael E., VIGNOLI, Valerio, SALOMONE, Nicola, FET, Elizabeth V. & SCHEMBRI, Patrick J. New molecular and morphological data on the “Euscorpius carpa- thicus” species complex (Scorpiones: Euscorpiidae) from Italy, Malta, and Greece justify the elevation of E. c. sicanus (C. L. Koch, 1837) to incispecie steve n 0. 0 nl celle a ee eieauy Sima CARFORA, Michaël, DE CHAMBRIER, Alain & VAUCHER, Claude. Le genre Amphoteromorphus (Cestoda: Proteocephalidea), parasite de poissons- chats d’ Amérique tropicale: étude morphologique et approche biosys- tematique,parielectrophorese des proteines........... ee gee WICHT, Barbara, MORETTI, Marco, PREATONI, Damiano, Tosi, Guido & MARTINOLI, Adriano. The presence of Soprano pipistrelle Pipistrellus pygmaeus (Leach, 1825) in Switzerland: first molecular and bioacustic BMIdENEESER N rele bea eared IT LOURENCO, Wilson R. Description of a new species of Tityus (Scorpiones, Buthidae) from Serra do Cipo in the State of Minas Gerais, Brazil... PAILL, Wolfgang. Amara pulpani Kult, 1949 — eine valide Art in den Ostalpeny(Coleoptera: Carabidae) ee ER ene eet An Pages 247-253 255-515 315-324 323-353 355-379 381-409 411-426 427-435 437-452 REVUE SUISSE DE ZOOLOGIE Volume 110 — Number 2 EBERMANN, Ernst, HAJIQANBAR, Hamid-reza & HADDAD IRANI-NEJAD, Karim. News records of phoretic and soil-living mites from Iran (Acaniprleterostigmatas Seufacaridae) EEE EEE ee PACE, Roberto. Homalotini from Borneo (Coleoptera, Staphylinidae)...... LOBL, I. & LESCHEN, R. A. B. Redescription and new species of Alexidia (Coleoptera::Staphylinidac:Scaphidimae) +P. =o ays ae ee SCHELLER, Ulf. New records of Pauropoda (Myriapoda) with descriptions of new species from Rwanda and Réunion (Pauropoda and Symphyla of thexGeneva Museum xa) retti ttt RE FET, Victor, GANTENBEIN, Benjamin, SOLEGLAD, Michael E., VIGNOLI, Valerio, SALOMONE, Nicola, FET, Elizabeth V. & SCHEMBRI, Patrick J. New molecular and morphological data on the “Euscorpius carpa- thicus” species complex (Scorpiones: Euscorpiidae) from Italy, Malta, and Greece justify the elevation of E. c. sicanus (C. L. Koch, 1837) to thekspecies level. pra LA PES MOMIE PER Se eee CARFORA, Michaél, DE CHAMBRIER, Alain & VAUCHER, Claude. The genus Amphoteromorphus (Cestoda: Proteocephalidea), parasite of neo- tropical catfishes : systematics including morphological data and pro- teinkeleetrophoresise. cc.) vias TTT RIO TE WICHT, Barbara, MORETTI, Marco, PREATONI, Damiano, Tosi, Guido & MARTINOLI, Adriano. The presence of Soprano pipistrelle Pipistrellus pygmaeus (Leach, 1825) in Switzerland: first molecular and bioacustic ÉVIDENCE SRE seria nine dena LS Seb bee as re LOC LOURENÇO, Wilson R. Description of a new species of Tityus (Scorpiones, Buthidae) from Serra do Cipo in the State of Minas Gerais, Brazil... PAILL, Wolfgang. Amara pulpani Kult, 1949 — a valid species in the Eastern Alpsi(ColeopteraCarabidae)f. tata LEI CRE Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 247-253 255-313 315-324 325-353 333-319 381-409 411-426 427-435 437-452 PUBLICATIONS DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N9S 1-17 (1908-1926) ........ serie Fr. (prix des fascicules sur demande) REVUE DE PALEOBIOLOGIE sats wm jase. ea: Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris)....... par fascicule Fr. COLLEMBOLENFAUNA EUROPAS ÉRGISIN 2 12) peel 960/(Reimpressromsl9 SA) ee eo ne Fr. THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION, WITH KEYS FOR DETERMINATION ), NOTIEREN Sup tO Sees AN ORIO e re Fr. CLASSIFICATION OF THE DIPLOPODA RGIMETOEEMANS2376Pt TINO Ne ER vey oration sey or le Aal one Rene Fr. LES OISEAUX NICHEURS DU CANTON DE GENEVE P. GEROUDET, C. GUEX & M. MAIRE Sp nombreuses Cartesyet MIP UTES. 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PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... Fr. CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) Unstrumenta Biodiversitatis I), I. LOBL, xii + 190 p., 1997. LA Fr. CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis U), H. G. DIRICKX, x +187 p., 1998 ................. Fr. A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis III), S. BOWESTEAD, 203 p., 1999 ................... Fr. THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE SOKOTRA ARCHIPELAGO (Instrumenta Biodiversitatis IV), B. SCHATTI & A. DESVOIGNES, MPA N N nt EEE Fr. PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY (Instrumenta Biodiversitatis V), C. LIENHARD & C. N. 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