“A D Qu: ALES NH | SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genéve tome 112 fascicule 2 2005 g 2 kl GENEVE JUIN 2005 ISSN 0035 - 418 X REVUE SUISSE DE ZOOLOGIE SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE TOME 112—FASCICULE 2 Publication subventionnée par: ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENÈVE SOCIÉTÉ SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d’histoire naturelle de Genève Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L’ ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d’abonnement doivent étre adressées a la rédaction de la Revue suisse de Zoologie, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Genéve 6, Suisse ANNALES de la SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genéve tome 112 fascicule 2 2005 ARISONIANA, 3 5 \ ah. \ 2 2009 } = VA — grate g 2 FL GENEVE JUIN 2005 ISSN 0035 - 418 X REVUE SUISSE DE ZOOLOGIE SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE TOME 112—FASCICULE 2 Publication subventionnée par: ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENÈVE SOCIÉTÉ SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d’histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d’histoire naturelle de Genève Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d’ institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration 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 112 (2): 311-312; juin 2005 Honorary Membership to Prof. S. C. Stearns One of the rewarding things about life is the chance to meet someone who makes a difference — who is ready to share knowledge, experience, ideas, advice, and constructive criticism. It is in this spirit of gratefulness, felt not only individually by many of its members, but also as a representative of the scientific community in our country, that the Swiss Zoological Society awards a honorary membership to Stephen C. Stearns, Professor at the Department of Ecology and Evolutionary Biology in the Faculty of Arts and Sciences at Yale University. Stephen C. Stearns was professor of zoology at the University of Basel from 1983 to 2000, when he moved to Yale University, the alma mater where he had gra- duated in 1967, before earning a M.Sc. from the University of Wisconsin and a Ph.D. from the University of British Columbia. The focus of Prof. Stearns' research interests is life history evolution, which connects ecology and evolutionary biology. His work in the area of life history evo- lution has substantially contributed to raise scientific awareness for this field (“The Evolution of Life Histories”, Oxford 1992). He has most likely been the first orga- nismic biologist in Switzerland to appreciate the importance of the upcoming genomics techniques for evolutionary biology, and to start to use them in his work. His research interests also cover evolutionary medicine (“Evolution in health and disease”, Oxford 1998). During his nearly two decades in Switzerland, Prof. Stearns enormously enriched research and teaching in zoology, well beyond the walls of the atheneum on the Rhine shores. His successful research, his dedication to expose students to the international scientific community - resulting in innovative forms of teaching such as the Guarda seminars, as well as in memorable wine tastings organized by Steve and his wife Beverly at their house in Arlesheim -, and his commitment to promote organismic biology and evolutionary ecology at the national and international level have been exemplary. He founded and has served as the president of the European Society for Evolutionary Biology and the Tropical Biology Association, was founding editor of the Journal of Evolutionary Biology, and has been a vice president of the Society for the Study of Evolution. In Switzerland, Steve has been the driving force in strengthening evolutionary biology. In this spirit the annual meeting of Swiss organismic biologists, now called BIOLOGY, has been invigorated, rejuvenated, becoming thriving and of high academic standards in recent years. He and his students (Jacqui Shykoff in particular) first established Darwin’s birthday party in Basel in 1992, an event that has become an essential highlight of the BIOLOGY conferences. Above all, the success of his efforts is reflected by the brilliant academic careers of many of his former colla- borators and students, to whom he has been a wise advisor and a source of inspiration. Steve's impact is further witnessed by the fact that his view of teaching and education continues — the Guarda seminars, where Steve always attracted brilliant faculties, have now become an international tool for student education, and this formula has spread from the peaks of Graubiinden to those of Valais, where similar workshops are held in 312 HONORARY MEMBERSHIP TO PROF. S. C. STEARNS the framework of the “3ème cycle romand”. Steve's and his assistants' courses in eco- logy, evolution and behaviour (EOV) finally changed the life of many young biologists. Steve's contribution to zoology and evolutionary biology in our country, to our scientific standards and academic performance, to our views and values, has been and continues to be very substantial and a point of reference, and is the gift of a very committed and generous scientist. Those who attended Darwin's birthday party held during the meeting of the Swiss Zoological Society in 1992 in a castle near Basel will never forget that in those days Darwin himself seemed to have paid us a visit. Basel, February 2005, The board and general assembly of the Swiss Zoological Society. REVUE SUISSE DE ZOOLOGIE 112 (2): 313-327; juin 2005 Gammarus species from River Jumahe, China (Crustacea, Amphipoda, Gammaridae) Zhong-E HOU & Shugiang LI Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, P. R. China. E-mail: lisg@ioz.ac.cn (all correspondence to Shugiang Li) Gammarus species from River Jumahe, China (Crustacea, Amphipoda, Gammaridae). - Three Gammarus species are reported from River Jumahe. Gammarus madidus sp. n. is characterized by antenna 2 with groups of long setae along the anterior and posterior margins, inner ramus of uropod 3 reaching about 40% of length of outer ramus and both rami fringed with long simple setae. G. lacustris Sars, 1863 is distinguished by epimeral plates 2 and 3 with sharp posterodistal corners, inner ramus of uropod 3 reaching about 75% of length of outer ramus and both rami armed with plumose setae. G. nekkensis Uchida, 1935 differs from congeneric species by pereopod 3 with long curled setae on the posterior margin and inner ramus of uropod 3 reaching 50% of outer ramus. Distribution data of these gammarids are presented. Keywords: Amphipoda - Gammarus - new species - taxonomy - China. INTRODUCTION On December 27, 2002, China started its gigantic South-to-North Water Transfer Project, which is expected to take 50 years to complete and will cost 58 billion US dollars. The project involves three canals running 1,300 kilometers across the eastern, middle and western parts of China, linking four major rivers, the Yangtze, Yellow, Huaihe and Haihe River (Anonymous, 2002). This project is thought to be an important infrastructural work for the optimal allocation of water resources in China. Beijing is one of the target cities of the Middle Line Project within the South- to-North Water Transfer Project. To better understand the ecological effects of this gigantic project, we have investigated the current Gammarus-fauna in most water systems around Beijing. In this paper we present a Gammarus inventory of the River Jumahe. River Jumahe, 254 km long, is one of five water systems running through Beijing City. Its source is located in Laiyuan County of Hebei Province and its mid- upper stretch runs through Fangshan District of Beijing, about 100 km away from urban Beijing. In Zhangfang Town it divides into the Rivers South-Jumahe and North- Jumahe, and then runs into the River Daginghe and further into Bohai Sea. Manuscript accepted 13.10.2004 314 Z.E. HOU & S. LI Gammarus is among the most species-rich groups of epigean freshwater am- phipods (Hou & Li, 2002a, b, 2003a; Hou, Li & Morino, 2002). Our intensive investi- gation of the Gammarus fauna of the River Jumahe began in May 2000, although previous collection is also used in the current study. Collecting was done at possible locality on both sides of the river where we could stop our car. Although most samples taken in 2001-2004 contained no Gammarus, we continued our search all along the river. This yielded three amphipod species: Gammarus lacustris, G. nekkensis and G. madidus sp. n., the latter of which is new to science. A detailed description of this new species, as well as the localities of all three species (Map 1) is given. MATERIAL AND METHODS Specimens were collected by a net and then preserved in 75% alcohol. For each species three to five specimens of each sex were dissected and appendages were mounted on slides according to the methods described by Holsinger (1967). The drawings were made with the aid of a drawing tube mounted on an Olympus BX-41 compound microscope. Holotype of the new species is deposited at the Institute of Zoology, Chinese Academy of Sciences (IZCAS), Beijing, China. Additional specimens, including paratypes of the new species are deposited at the same institute and at the Muséum d’histoire naturelle, Geneva (MHNG), Switzerland. DESCRIPTIONS Gammarus lacustris Sars, 1863 Gammarus lacustris Sars, 1863: 207; Pinkster, 1972: 166-169, Figs 1-2: Karaman & Pinkster, 1977: 32-34, Fig.12; Barnard & Dai, 1988: 92-94, Figs 1-5. Gammarus pulex Dahl, 1915: 1-32, Fig. 1. Gammarus scandinavicus S. Karaman, 1931: 101, Fig. 6a. Gammarus bolkayi S. Karaman, 1934: 325, Fig.1. Gammarus (Rivulogammarus) lacustris Schellenberg, 1937: 490, Figs 2-6. Rivulogammarus lacustris Dussart, 1948: 101-102; Straskraba, 1967: 208. Gammarus lacustris lacustris Bousfield, 1958: 80. Gammarus wigrensis Micherdzinski, 1959: 598-599, Fig. 81. Material examined: 97 33, 163 99 and 29 juv. (No. 1), a spring near Shidu (39°38 .521°N, 115°38.273’E), Fangshan District, Beijing, altitude 160-190 m, water temperature 6°C, 12 May 2000, collected by Shugiang Li; 38 ¢ 4,40 2 © and 7 juv. (MHNG, No. 2), same data as No. 1; 46 6 d, 87 © © and 10 juv. (No. 3), same data as No. 1 except for the collecting day 13 May 2000; 75 88,108 £ and 16 juv. (No. 4), same data as No. 3; 23 66,28 22 and 10 juv. (No. 5), spring near Cave Yugudong (39°45.434°N, 115°24.312° E), Yesanpo Scenic Area, Laishui County, Hebei Province, altitude 340 m, 13 May 2000, collected by Shugiang Li; 8 33,7 2 and 2 juv. (No. 6), same data as No. 5; 55 dd and 50 9 2, a fishery without name, Laiyuan County (39°18° N, 114°42’E), Hebei Province, 8 August 1988, collected by Shugiang Li; 32 Sd and 32 2, headwaters of River Jumahe, Laiyuan County, Hebei Province, 5 September 2004, collected by Zhong-E Hou and Yucheng Lin. Diagnosis: Body large and stout; peduncle of antenna 2 with few short setae, calceoli present in male, pereopods 3 and 4 with long straight setae on posterior mar- gins, epimeral plates 2 and 3 with sharp posterodistal corners, inner ramus of uropod 3 reaching about 75% of length of outer ramus, both rami set with plumose setae. GAMMARUS FROM RIVER JUMAHE 315 17.4 26.1 34.8km MAP 1 Localities of freshwater amphipods along the River Jumahe. Triangle = Gammarus lacustris Sars, 1863; square = G. nekkensis Uchida, 1935; pentangle = G. madidus sp. n.. Remarks: G. lacustris was redescribed by Karaman & Pinkster (1977) based on samples from all over the distribution area, while Barnarad & Dai (1988) redescribed it in detail with material from Yunnan, China. The specimens examined accord well with the figures and description of G. lacustris given by Barnard & Dai (1988), except that the bases of pereopods 5-7 is a little more elongate, epimeral plates 2 and 3 without very sharp posterodistal corners and telson with 1 basolateral spine. Distribution: Karaman & Pinkster (1977) recorded the known distribution of the species as follows: Afghanistan, Austria, Balkan region, Canada, Czech Republic, Denmark, Finland, France, Germany, India, Ireland, Italy, Norway, north Poland, Scotland, Slovakia, Sweden, Switzerland, Spain, Turkey, northern parts of U.S.A. According to the material deposited in IZCAS, G. lacustris also occurs in Nepal (Parbat Distr.) and China (Yunnan Province, Hebei Province and Xinjiang Uygur Autonomous Region). Gammarus nekkensis Uchida, 1935 Gammarus nekkensis Uchida, 1935: 1-6, pls 1-4; Barnard & Barnard, 1983: 463; Karaman, 1984: 147-148; Karaman, 1989: 19-35, Figs 1-5. Gammarus (Rivulogammarus) nekkensis Barnard & Dai, 1988: 90. Material examined: 20 85,21 22 and 5 juv., Mt. Baicaopan, Beibiangiao Village, Jiulong Town, Yesanpo Scenic Area, Laishui County, Hebei Province, 13 May 2000, collected 316 Z. E. HOU & S. LI by Shugiang Li; 2 dd and 2 9 9, a sump under Baicaopan Glacier, Yesanpo Scenic Area, gla- cier water temperature 5.5°C, glacier surface temperature -1.5°C, altitude 1180 m, 13 May 2000, collected by Shugiang Li; 1 4,2 2 2 and 5 juv., a brook on the left hand of the glacier, 13 May 2000, collected by Shugiang Li; 13 dd, a sump in the middle hill of Mt. Baicaopan, 13 May 2000, collected by Shugiang Li; 8 dd, 10 9 © and 5 juv. (MHNG), a drinkable water spring near Wanglaopu Village, Liudu Town, Fangshan District, Beijing City, opening about 0.5 square meter, 24 November 2003, collected by Shugiang Li, Lihong Tu and Kaibaryer Meng. Diagnosis: Antennae 1 and 2 with few setae, calceoli present in male, merus to propodus of pereopod 3 with long curled setae on posterior margin, pereopods 5-7 with few setae on anterior margin, epimeral plates 1-3 with blunt posterodistal corners, inner ramus of uropod 3 reaching about 50% of length of outer ramus, outer margin of outer ramus in uropod 3 with long simple setae only. Remarks: G. nekkensis was first described on the basis of material from Wuling Mountain, Hebei, then it was reviewed in detail by Karaman (1989) on the basis of specimens from Beijing. The present material accords well with the figures and description given by Karaman (1989), except for a little shorter inner ramus of uropod 3 and long setae on the dorsal surface of the telson. These features may be variable within the species. Distribution: Known from the River Jumahe and Mt. Wuling. River Jumahe runs along Mts. Taihangshan, which is the west boundary of North China Plain (33.0- 40.5°N, 113.0-119.5°E), while Mt. Wuling is the highest peak in Mts. Yanshan, the north boundary of North China Plain. Gammarus madidus sp. n. Figs 1-6 Material examined: Holotype (IZCAS-I-A0111), 4, a sump on the middle hill of Mt. Baicaopan, Yesanpo Scenic Area, Laishui County, Hebei Province, 13 May 2000, collected by Shugiang Li; paratypes, 195 dg 4,172 9 © and 13 juv. IZCAS), 20 dd and 20 2 2 (MHNG), same data as holotype. Etymology: The latin epithet “madidus” (= wet, moist) refers to the habitat of the species. Diagnosis: Peduncular articles 4 and 5 with groups of long setae along anterior and posterior margins, calceoli present in male; gnathopods 1 and 2 with straight setae; pereopod 3 with long, weakly curled setae on posterior margin; inner ramus of uropod 3 reaching about 40% of length of outer ramus, both rami fringed with long simple setae. Description of male holotype: Body length 14.0 mm. Head (Fig. 1A): Inferior antennal sinus deep, eyes medium in size. Antenna 1 (Fig. SA): Peduncular articles 1-3 with length ratio 1: 0.7 : 0.5, bearing some distal setae; flagellum with 30 articles, most articles with aesthestascs, accessory flagellum with 5 articles. Antenna 2 (Figs 5B, F, G): Peduncular article 4 about as long as article 5, both with 5-7 groups of long setae on inner margin, 6-7 groups of long setae on inner sur- face and 3-5 groups of long setae on outer margin; flagellum with 15 articles, calceoli present. Upper lip (Fig. 1C): Convex, with minute setae. Lower lip (Fig. 1J): Inner lobe absent. GAMMARUS FROM RIVER JUMAHE 317 Fic. 1 Gammarus madidus sp. n., holotype, male. A, head; B, urosomites 1-3 (dorsal view); C, upper lip; D, left mandible; E, maxilliped; F, incisor and lacinia mobilis of right mandible; G, maxilla - 2; H, left maxilla 1; I, palp of right maxilla 1; J, lower lip. 318 Z.E. HOU & S. LI Mandibles (Figs 1D, F): Left incisor with 5 teeth; lacinia mobilis with 4 denti- tions; spine row with 9 plumose setae; molar bearing 1 plumose seta; article 2 of palp bearing 21 stiff setae, article 3 about 75% of length of article 2, bearing a group of 5 A-setae on outer surface, 4 B-setae on inner surface, a row of plumose D-setae and 5 long E-setae. Right incisor with 4 teeth, lacinia mobilis bifurcate, each with crenu- lations at edge. Maxilla 1 (Figs 1H, I): Asymmetrical, inner margin of inner plate bearing several setules basally and 17 plumose setae evenly distributed between base and apex; outer plate bearing 11 serrated spines; palp with 2 articles, left second article falcate, bearing 8 slender spines apically accompanied by 4 apico-facial setae; right second article bearing 5 blunt spines associated with 2 stiff setae. Maxilla 2 (Fig. 1G): Inner plate bearing a diagonal row of 15 plumose setae on inner margin, some pubescent setae on outer margin and many stiff setae apically; outer plate with long stiff setae apically and some setules on outer margin. Maxilliped (Fig. 1E): Inner plate bearing 1 subapical spine and 3 apical blunt spines; outer plate with row of 16 blade spines and 4 pectinate setae; palp with 4 articles. Coxal plates: Coxal plate 1 (Fig. 2A) weakly dilated distally, bearing 3 setules on anteroventral corner and 2 setules on posteroventral corner; coxal plates 2 and 3 subrectangular (Figs 2B, 3B), bearing 2 setules on anteroventral corner and 1 setule on posteroventral corner; coxal plate 4 excavated on posterior margin (Fig. 3A), bearing 2 setules on anteroventral corner and 5 setules on posterior margin; coxal plates 5 and 6 with small anterior lobe (Figs 4A, B), bearing 2 setules on posterior margins; coxal plate 7 shallow (Fig. 4C), bearing 4 setules on posterior margin. Coxal gills present on gnathopod 2 and pereopods 3-7 (Figs 2B, 3A, B, 4A-C). Gnathopod 1 (Figs 2A, C): Basis bearing long setae on antero-proximal and posterior margin; carpus about 75% of length of propodus, densely setose, with long setae on posterior margin; propodus ovate, palm oblique, bearing 1 palmar medial spine, 4 pairs of spines and 1 spine on posterior margin and 6 spines on inner surface, these associated with groups of long, naked setae on posterior margin; dactylus with 1 seta on outer margin and 2 setules at hinge of nail. Gnathopod 2 (Figs 2B, D): Basis similar to that of gnathopod 1; carpus reaching about 75% of length of propodus; propodus subrectangular, palm weakly oblique, bearing 1 medial palmar spine, 4 spines on inner posterodistal corner and 2 spines on outer posterodistal corner, these associated with groups of setae on posterior margin; dactylus bearing 1 seta on outer margin and 2 setules at hinge of nail. Pereopod 3 (Figs 3B, C): Basis with groups of long setae on posterior margin; merus to propodus densely set with long, weakly curled setae on posterior margins, carpus and propodus accompanied by 4-5 spines; dactylus bearing 1 plumose seta on outer margin and 2 setae at hinge of nail. Pereopod 4 (Figs 3A, D) shorter than pereopod 3; basis with long setae on pos- terior margin; merus with 5 groups of long setae on posterior margin; carpus bearing 2-3-2-1 spines accompanied by long setae on posterior margin; propodus bearing 2-1- 1-1 spines accompanied by some long setae on posterior margin; dactylus bearing 1 plumose seta on outer margin and 2 setae at hinge of nail. GAMMARUS FROM RIVER JUMAHE 319 Fic. 2 Gammarus madidus sp. n., holotype, male. A, gnathopod 1; B, gnathopod 2; C, propodus of gnathopod 1; D, propodus of gnathopod 2. Z. E. HOU & S. LI 320 KVP 4 NS N Ne WG / \ à \ 3 ARS NETZ — 7 y > N È y LE ono AL, AA 4 ZZ SNS = 1% G TL 4, Cf RS (LZ SSS L,) DELI TEA 7 LH ZZ Fic. 3 Gammarus madidus sp. n., holotype, male. A, pereopod 4; B, pereopod 3; C, dactylus of pere- opod 3; D, dactylus of pereopod 4; E, uropod 1; F, uropod 2; G, uropod 3; H, pleopod 1. GAMMARUS FROM RIVER JUMAHE 321 Pereopod 5 (Figs 4A, D): Basis with almost straight posterior margin, bearing 5 long setae and 5 single short spines on anterior margin, a row of 7 setules on posterior margin; merus bearing 3 groups of spines accompanied by short setae on anterior margin and 3 spines on posterior margin; carpus bearing 4 groups of spines associated with short setae on anterior margin and 3 pairs of spines on posterior margin; propodus bearing 5 groups of spines on medial surface, some setae and 1 spine on posterior margin; dactylus bearing 1 plumose seta on outer margin and 2 stiff setae at hinge of nail. Pereopod 6 (Figs 4B, E) longer than pereopod 5, basis elongate, attenuated distally, bearing a group of 4 long setae and 5 single spines on anterior margin and a row of 11 setules on posterior margin; merus and carpus bearing 3-4 groups of spines accompanied by a few short setae on anterior margin and 2-3 groups of spines on posterior margin; propodus bearing 4 groups of spines on medial surface, 1 spine on posterior margin; dactylus similar to that of pereopod 5. Pereopod 7 (Figs 4C, F) similar to pereopod 6, basis weakly rounded on pro- ximal posterior margin, inner surface bearing 1 spine accompanied by 3 setae on posterodistal corner. : Epimeral plates: Plates 1-3 progressively acute on posterodistal corners, bearing 3-5 setules on posterior margins; plate 1 (Fig. SC) bearing 11 long setae on antero- ventral corner associated with 1 spine; plate 2 (Fig. 5D) with nearly straight posterior margin, bearing 1 setule and a pair of spines on ventral margin; plate 3 (Fig. SE) bearing 2 setae on anterior margin and 3 spines on ventral margin. Pleopods (Fig. 3H): Pleopods 1-3 subequal, peduncle bearing 1-2 retinacula on anterodistal corner and some long setae on dorsal surface; outer ramus a little shorter than inner ramus, both rami armed with plumose setae. Urosomites 1-3 dorsally flat (Fig. 1B), urosomite 1 bearing 2-1-1-1 spines accompanied by some setae on posterodorsal margin from left to right; urosomite 2 bearing 2-1-1-2 spines accompanied by setae, setae longer than spines; urosomite 3 bearing 2 pairs of spines on both sides and 2 pairs of setae on medial dorsal margin. Uropod 1 (Fig. 3E): Peduncle bearing 1 basofacial spine, 1 spine on outer margin, 2 single spines on inner margin, 2 spines on anterodistal corner and 1 spine on posterodistal corner; outer ramus reaching 57% of length of peduncle, bearing 1 spine on each side; inner ramus a little shorter than outer ramus, bearing 1 spine on inner margin; both rami bearing 5 distal spines. Uropod 2 (Fig. 3F): Peduncle bearing 2 spines on outer margin, 3 spines on inner margin, | spine on anterodistal corner and 1 spine on posterodistal corner; outer ramus reaching 90% of length of peduncle, bearing 1 spine on outer margin; inner ramus a little longer than outer ramus, bearing 1 spine on inner margin and 1 spine on outer margin. Uropod 3 (Fig. 3G): Peduncle bearing 2 spines on outer margin, a few setae on inner margin, 2 spines on mid-ventrodistal margin, 1 spine apico-laterally and 1 spine apico-medially; inner ramus reaching 40% of length of outer ramus, bearing 1 long distal spine; article 1 of outer ramus bearing 1-2-2-2 spines on outer margin and 2 pairs of distal spines; article 2 shorter than adjacent spines; both rami densely fringed with long simple setae. 322 Z.E. HOU & S. LI FIG. 4 Gammarus madidus sp. n., holotype, male. A, pereopod 5; B, pereopod 6; C, pereopod 7; D, dactylus of pereopod 5; E, dactylus of pereopod 6; F, dactylus of pereopod 7; G, telson. GAMMARUS FROM RIVER JUMAHE 323 Telson deeply cleft (Fig. 4G), each lobe bearing 1 lateral spine accompanied by long setae, 3-4 distal spines accompanied by long setae and some facial setae. Description of female: Body length 13.5 mm. Antenna 2 (Fig. SH): Peduncular articles 4 and 5 bearing groups of long setae along anterior and posterior margins; flagellum with 12 articles, calceoli absent. Gnathopod 1 (Fig. 6A): Propodus ovate, palm not as oblique as that of male, bearing 8 spines on posterodistal corner; dactylus bearing 1 seta on outer margin and 2 setules at hinge of nail. Gnathopod 2 (Fig. 6B): Propodus subrectangular, palm truncate, bearing 5 spines on posterodistal corner; dactylus bearing 1 seta on outer margin and 2 setules at hinge of nail. Pereopods 5-7: Bases shorter than those of male (Figs 6C-E). Uropod 3 (Fig. 6F): Inner ramus reaching 44% of length of outer ramus, bear- ing 1 lateral and 1 distal spines; outer ramus with 1-2-2 spines on outer margin and 1 spine on inner margin, both rami fringed with simple setae. Oostegites (Fig. SI) present on gnathopod 2 and pereopods 3-5. Remarks: Gammarus madidus sp. n. is similar to G. electrus Hou & Li (see Hou & Li, 2003c), which is distributed in and around Beijing City and to G. curvativus Hou & Li (see Hou & Li, 2003b), which occurs in Yunnan and Sichuan Province. All three species possess (1) antenna 2 with calceoli, (2) inner ramus of uropod 3 reaching about 40% of length of outer ramus and both rami armed with long simple setae, (3) uro- somites 1-3 flat, with 4 groups of spines and setae. G. madidus sp. n. differs from G. electrus (character states of G. electrus in parentheses) in (1) peduncular articles 4 and 5 of antenna 2 bearing groups of long setae along anterior and posterior margins (short setae), (2) gnathopods 1 and 2 with long straight setae (gnathopod 1 with long curled setae on posterior margin of propodus, gnathopod 2 with long curled setae on dorsal margins of carpus and propodus), (3) pereopod 3 with long straight setae on posterior margin (long curled setae). G. madidus sp. n. differs from G. curvativus (character states of G. curvativus in parentheses) by (1) accessory flagellum of antenna 1 with 4 or 5 articles (2 articles); (2) gnathopod 2 with straight setae (gnathopod 2 with long curled setae on dorsal margin of carpus and weakly curled setae on dorsal margin of propodus); (3) telson with 1 lateral spine (no spines). Gammarus madidus sp. n. is also similar to G. nekkensis in (1) antenna 2 with calceoli, (2) epimeral plates 1-3 progressively acute, (3) inner ramus not more than half of outer ramus. The new species can be distinguished from G. nekkensis (character states of G. nekkensis in parentheses) by (1) antenna 2 with long setae on peduncular articles 4 and 5 (short setae), (2) pereopod 3 with long straight setae (curled setae), (3) both rami of uropod 3 bearing long simple setae (outer margin of outer ramus bearing long simple setae, but inner margin of outer ramus and both margin of inner ramus bearing plumose setae). Gammarus madidus sp. n. coexists with G. nekkensis at Mt. Baicaopan. Distribution: Known only from Mt. Baicaopan, Hebei Province. 324 Z. E. HOU & S. LI FIG. 5 Gammarus madidus sp. n., male: A-G: female: H, I. A, antenna 1; B, antenna 2; C, epimeral plate 1; D, epimeral plate 2; E, epimeral plate 3; F, peduncular article 5 of antenna 2; G, one flagel- lomere of antenna 2; H, peduncular articles 3-5 of antenna 2; I, oostegite 2. GAMMARUS FROM RIVER JUMAHE 325 Fic. 6 Gammarus madidus sp. n., female. A, propodus of gnathopod 1; B, propodus of gnathopod 2; C, basis of pereopod 5; D, basis of pereopod 6; E, basis of pereopod 7; F, uropod 3. 326 Z. E. HOU & S. LI DISCUSSION Gammarus lacustris and G. nekkensis are very common species in China. They occur at several places in upper reaches of River Jumahe. They live specially in river- side springs, but we could also find them in the main rivers in wintertime. In the low- er reaches of River Jumahe, including Rivers South-Jumahe and North-Jumahe, we did not find any Gammarus species. The reason could be that the river often dried. In case that there is enough water, water pollution may be another reason. Gammarus madidus sp. n. occurs in a small stretch of the River Jumahe, while G. lacustris and G. nekkensis are widespread. G. lacustris and G. nekkensis are ecological generalists that have been able to colonize many kinds of new, disturbed and marginal habitats, whereas related species with only a narrow distribution, as G. madidus, are ecological specialists that competitively dominate specific kinds of rela- tively stable biotopes (Glazier & Eckert, 2002). ACKNOWLEDGEMENTS The authors would like to express their thanks to Dr Dirk Platvoet (University of Amsterdam, the Netherlands) for providing advice in the course of this study and for critically reading the manuscript. This study was supported by the National Natural Sciences Foundation of China (NSFC-30270183, 30370263, 30310464, 30470213, 30499341), by the National Science Fund for Fostering Talents in Basic Research (NSFC-J0030092), by the Beijing Natural Science Foundation (6052017) and partly also by the Kadoorie Farm and Botanic Garden, Hong Kong Special Administrative Region, China. REFERENCES ANONYMOUS, 2002. Beijingers able to drink water from Yangtze River in 2007. People’s Daily, 9 February. BARNARD, J. L. & BARNARD, C. M. 1983. Freshwater Amphipoda of the World I, II. Hayfield Associates, Mt. Vernon, Virginia, 1-830. BARNARD, J. L. & DAI, A. Y. 1988. Four species of Gammarus (Amphipoda) from China. Sinozoologia 6: 85-112. BOUSFIELD, E. L. 1958. Freshwater amphipod crustaceans. Canadian Field Naturalist 72 (2): 55- 113% DAHL, K. 1915. En studie over Grundaatense eller Matfloens (Gammarus pulex) [sic] biologi og utbredelse I Norge. Norsk Jaeger Fiskerforenings Tidsskrift 44: 1-32. DUSSART, B. 1948. Sur la présence en Haute-Savoie de Gammarus (Rivulogammarus) lacustris Sars. Comptes Rendus de la Societe de Geographie 25: 101-103. GLAZIER, D. S. & ECKERT, S. E. 2002. Competitive ability, body size and geographical range size in small mammals. Journal of Biogeography 29: 81-92. HOLSINGER, J. R. 1967. Systematics, speciation, and distribution of the subterranean amphipod genus Stygonectes (Gammaridae). Bulletin of United States National Museum 259: 1-176. Hou, Z. E. & LI, S. Q. 2002a. A new cave amphipod of the genus Sinogammarus from China. Crustaceana 75 (6): 815-825. Hou, Z. E. & LI, S. Q. 2002b. Descriptions of two new species of genus Gammarus (Crustacea: Amphipoda: Gammaridae) from Yunnan, China. The Raffles Bulletin of Zoology 50 (1): 37-32: GAMMARUS FROM RIVER JUMAHE 327 Hou, Z. E. & Li, S. Q. 2003a. Terrestrial talitrid amphipods (Crustacea: Amphipoda) from China and Vietnam: studies on the collection in IZCAS. Journal of Natural History 37 (20): 2441-2460. Hou, Z. E. & LI, S. Q. 2003b. Two new freshwater gammarids (Crustacea: Amphipoda: Gammaridae) from Lake Lugu, China. Revue suisse de Zoologie 110 (3): 547-564. Hou, Z. E. & LI, S. Q. 2003c. Gammarus electrus, a new species of Amphipoda from Beijing, China (Crustacea: Amphipoda: Gammaridae). Acta Zootaxonomica Sinica 28 (2): 241-248. Hou, Z. E., Li,S.Q. & Morino, H. 2002. Three new species of the genus Gammarus (Crustacea, Amphipoda, Gammaridae) from Yunnan, China. Zoological Sciences 19 (8): 939-960. KARAMAN, G. S. & PINKSTER, S. 1977. Freshwater Gammarus species from Europe, North Africa and adjacent regions of Asia (Crustacea-Amphipoda). Part I. Gammarus pulex-group and related species. Bijdragen tot de Dierkunde 47 (1): 1-97. KARAMAN, G. S. 1984. Remarks to the freshwater Gammarus species (Fam. Gammaridae) from Korea, China, Japan and some adjacent regions (Contribution to the knowledge of the Amphipoda 134). The Montenegrin Academy of Sciences and Arts Glasnik of the Section of Natural Sciences 4: 139-162. KARAMAN, G. S. 1989. One freshwater Gammarus species (Gammaridea, Fam. Gammaridae) from China (Contribution to the knowledge of the Amphipoda 189). Poljoprivreda I Sumarstvo 35 (1-2): 19-36. KARAMAN, S. 1931. Beitrag zur Kenntnis der Süsswasser-Amphipoden. Glasnik Skopso Naucnog Drustva 9: 93-107. KARAMAN, S. 1934. Beitrag zur Kenntniss jugoslawischer Siisswasser-Amphipoden. Zoo- logischer Anzeiger 107: 325-333. MICHERDZINSKI, W. 1959. Die Gammarusarten (Amphipoda) Polens. Acta Zoologica Cracoviensia 4 (10): 527-637. PINKSTER, S. 1972. On members of the Gammarus pulex-group (Crustacea-Amphipoda) from Western European. Bijdragen tot de Dierkunde 42:164-191. SARS, G. O. 1863. Beretning om en i Sommeren 1862 foretagen zoologisk Reise i Christianias og Trondhjems Stifter. Nyt Magazine for Naturvidenskaberne, Christiania 12: 193-340. SCHELLENBERG, A. 1937. Kritische Bemerkungen zur Systematik der Süsswassergammariden. Zoologische Jahrbiicher, Abteilung fiir Systematik 69: 469-516. STRASKRABA, M. 1967. Amphipoda (pp. 202-209). In: ILLIES J. (ed.). Limnofauna Europaea. Gustav Fischer Verlag, Stuttgart, 474 pp. UCHIDA, H. 1935. Crustacea of Jehol — Orders: Phyllopoda, Decapoda, Isopoda and Amphipoda. Freshwater Amphipoda. Report of the First Scientific Expedition to Manchoukuo, section 5, division 1, part 2, article 9: 1-6. PC SMO SON AMAR ET | = | a‘ a) BEN ih ) eboqriqena bitte being EURE as ieh ann Yu à L PAR ! u malate u Cape; fcr SET! SENS Pinon A Tila | M n ; i i 10 ee 19 = { = di #3 7 x ; a A Tann | faina Bin PL À A 1 u Te a i i) je DEEE © D Mi (15 30e a ul Thee Sid ra 44 7 n a * = L REVUE SUISSE DE ZOOLOGIE 112 (2): 329-331; juin 2005 Über eine neue Octodrilus-Art aus Frankreich (Oligochaeta: Lumbricidae) Andras ZICSI! & Gerard CUENDET? ! Lehrstuhl für Tiersystematik und Ökologie der Eötvös Loränd Universität, Zootaxonomische Forschungsgruppe der Ungarischen Akademie der Wissenschaften, Pàzmany Péter sétany 1/C, H-1117 Budapest, Ungarn. E-mail: zicsi@cerberus.elte.hu 2 CH-1675 Vauderens, Switzerland. E-mail: gerard.cuendet@ worldcom.ch On a new Octodrilus species from France (Oligochaeta: Lumbricidae). - The new earthworm species Octodrilus juvyi sp. n. is described and the taxonomic differences among the red, small bodied Octodrilus species are emphasized. Keywords: Earthworms - Oligochaeta - Lumbricidae - taxonomy - new species - France. EINLEITUNG Aus Frankreich war bislang als einzige Art der Gattung Octodrilus complanatus (Dugès, 1828) gemeldet, die in den südöstlichsten Teilen Frankreichs und in Korsika verbreitet ist (Bouché, 1972). Octodrilus complanatus ist in den südlichen Teilen Europas, vorwiegend jedoch im mediterranem Gebiet und Nordafrika, weit verbreitet und gehört zu den tiefgra- benden, nie rot pigmentierten Grossformen dieser Gattung. Die kleinen, in der obersten Bodenschicht oder Laubstreu lebenden, rotpigmentierten Octodrilus-Arten, die haupt- sächlich im Alpengebiet Italiens, der Schweiz, Osterreichs und Sloweniens und in den Dinarischen Alpen anzureffen sind, waren bisher aus dem westlichen Alpengebiet Frankreichs nicht gemeldet. Jetzt liegen 6 Exemplare einer kleinen, rotpigmentierten Art vor, die Herr Bernard Juvy in den Westalpen im Massiv Grande Chartreuse gesammelt hatte und die nachstehend als neue Art beschrieben wird. BESCHREIBUNG DER ART Octodrilus Omodeo, 1956 Octolasium (Octodrilus) Omodeo, 1956: 177, part. Octodrilus; Bouché, 1972: 311. Octodrilus; Zicsi, 1986: 108. Octodrilus; MrSic, 1991, 356. Octodrilus; Qui & Bouché, 1998: 203. Octodrilus; Csuzdi & Zicsi, 2003: 206. Typus-Art: Lumbricus complanatus Dugès, 1828 Manuskript angenommen am 13.10.2004 330 A. ZICSI & G. CUENDET Nach Abtrennung der Gattung Octodriloides Zicsi, 1986 verblieben in der Gattung Octodrilus 36 Arten (Mrsic, 1991), die über ganz Europa und Nordafrika ver- breitet sind. Der Hauptunterschied zwischen den beiden Gattungen besteht in der Lage der männlichen Poren, die bei Vertretern von Octotrilus auf dem 15. Segment, bei denen von Octodriloides jedoch vor oder auf dem Giirtelsegment liegen. Die Arten beider Gattungen sowie die von Lumbricus Linnaeus, 1758 und Octolasion Orley, 1885 sind durch die Lage des Gürtels und der Pubertätsstreifen ausschlaggebend gekennzeichnet, die Verschiebung dieser Merkmale um ein einziges Segment reicht zur Trennung der Arten aus. Octodrilus juvyi sp. n. Holotypus: HNHM 15201. Frankreich, Massif Grande Chartreuse, Saint Pierre de Chartreuse, Rocher de la Petite Vache, 1195 m, 17.6.2003, leg. B. Juvy. Paratypen: HNHM 12502 2 Ex., MHNG - INVE 35901 1 Ex. Fundort wie beim Holotypus. HNHM 15203 2 Ex. Bois de Valombré, 1460 m, 20.09.2002, leg. B. Juvy. Typenmaterial wird im Naturhistorischen Museum Budapest (HNHM) und im Naturhistorischen Museum Genf (MHNG) aufbewahrt. Etymologie: Die neue Art wird zu Ehren des Sammlers, Herrn Umweltingenieur Bernard Juvy (CEMAGREF - Unité de Recherche E.P.M., Saint Martin d’Heres, Frankreich) benannt. Diagnose: L. 55-63 mm, B. 3-4 mm, Segmentzahl 76-113. Rotviolett. Kopf epi- lobisch. Borsten ungepaart. Gürtel vom 29.-35., Pubertätsstreifen vom 29.-35. Segment. Dissepimente 11/12-13/14 schwach verdickt. Letztes Paar Herzen im 11., Kalkdrüsen im 10-12. Segment. Testikelblasen im 10. und 11. Segment. 4 Paar Samen- säcke im 9.-12. Segment. Typhlosolis einfach. Nephridialblasen ocarina-förmig. Mus- kulatur gefiedert. 5 Paar Samentaschen in Segment 6/7-10/11. Die neue Art unter- scheidet sich von allen rotpigmentierten, kleinen Arten mit männlichen Poren auf dem 15. Segment durch die Lage des Gürtels in Kombination mit der Lage und Zahl der Samentaschen (vergl. auch Tabelle 1). Holotypus: Länge 63 mm, Breite 3,5 mm, Segmentzahl 123. Paratypen: Lange 55-63 mm, Breite 3-4 mm, Segmentzahl 76-113. Farbe dorsal, bis zur Borstenlinie c rotviolett, ventral weiss. Kopf epilobisch, halb offen. Segmente nicht geringelt. Borsten ungepaart, hinter dem Gürtel aa:ab:bc:cd:dd wie 2,7:1,3:1,3:1:4. Nephridialporen etwas oberhalb der Borstenlinie b, regelmässig angeordnet. Erster Riickenporus auf Intersegmentalfurche 14/15. Weibliche Poren winzig kleine Offnungen auf dem 14. Segment, dicht neben der Borstenlinie b. Männliche Poren ebenfalls winzig kleine Offnungen auf dem 15. Segment, in der Mitte zwischen der Borstenlinie b und c. 5 Paare kleiner Samentaschenöffnungen auf Intersegmentalfurche 6/7-10/11, zwischen den Borstelinien b und c. | Gürtel vom 29. - 35. Segment, sattelförmig. Pubertätsstreifen vom 29. - 35. Segment. Innere Organisation: Dissepimente 6/7-9/10 dünn, 11/12 - 13/14 etwas kräftiger ausgebildet. Schlunddriisen bis zum 6. Segment reichend. Letztes Paar Herzen im 11. Segment, eine unpaarige Schlinge im 12. Segment. Kalkdriisen im 10-12. Segment, mit Ausbuchtungen im 10. Segment. Kropf im 15.-16. Segment; Muskelmagen im 17.-18. EINE NEUE OCTODRILUS-ART 331 Segment; Mitteldarm im 19. Segment beginnend, Typhlosolis vorhanden, sackförmig. Nephridialblase mit einer nach hinten gebogenen, verschmolzenen Schlinge (ocarina- shaped). Hoden und Samentrichter im 10. und 11. Segment von mächtigen Testikel- blasen umgeben. Vier Paar Samensäcke im 9.-12. Segment. Ovarien im 13. Segment. Samentaschen im 6.-11. Segment, die beiden vorderen vor den Dissepimenten, die drei hinteren hinter den Dissepimenten liegend. TABELLE |. Wichtige Trennungsmerkmale kleiner, rotpigmentierter Octodrilus-Arten. Art Giirtel Pubertats- Samen- Zahl der Samen- streifen taschen Samen- säcke öffnungen taschen O. argoviensis (Bretscher, 1899) 28-34 28-34,35 6/7-11/12 6 9-12 O. croaticus (Rosa, 1895) 28-35 28-35 5/6-10/11 6 9-12 O. lissaensis (Michaelsen, 1891) 29-36 29-36 5/6-10/11 6 9-12 O. bretscheri Zicsi, 1969 29,30-36 29,30-36 6/7-10/11 5 9-12 O. hemiander (Cognetti, 1901) 28-36,37 28-38,39 6/7-12/13 if 10, 12 O. illyricus MrSic, 1987 28, 29-36 29-36 6/7-10/11 5 10, 12 O. lissaensioides Zicsi, 1971 29-36 29-37 5/6-10/11 6 910212 O. juvyi sp. n. 29-35 29-35 6/7-10/11 5, 9-12 DANKSAGUNG Für die Unterstützung der Ungarischen Wissenschaftlichen Stiftung (OTKA No. T42745) wird auch an dieser Stelle gedankt. LITERATUR BOUCHÉ, M. B. 1972. Lombriciens de France. Ecologie et systématique. Institut National de la Recherche Agronomique, Articles de Zoologie-Ecologie animale (Numéro hors-série), 671 pp. Csuzpi, Cs. & Zıcsı, A. 2003. Earthworms of Hungary. Pedozoologica Hungarica. No. 1. Hungarian Natural History Museum, Budapest, 272 pp. MRSIC, N. 1991. Monografija o dezevnikih (Lumbricidae) Balkana I-II. Academia Scientarium et Artium Slovenica, Classis IV: Historia Naturalis, Opera 31: 1-757. OMODEO, P. 1956. Contributo alla revisione dei Lumbricidae. Archivio Zoologico Italiano 41: 129-143. Qiu, J.-P. & BoucHÉ, M. B. 1998. Révision des taxons supraspécifiques de Lumbricoidea. Documents pédozoologiques & intégrologiques 3: 179-216. Zicsi, A. 1986. Uber die taxonomischen Probleme der Gattung Octodrilus Omodeo, 1956 und Octodriloides gen. n. (Oligochaeta: Lumbricidae). Opuscula Zoologica, Budapest 22: 103-112. end i. page ner Fissi pu sad. Pr FH) mult fel In : “al pP om ABs 3s rune SEX ME ca ì be: £ I) PAP Ren di n Sade) An (tye i sf gi AU VAR > VER ali “ail we { 01 A Ne | F, san A zu pmi le ae à ì Tay tis) Rio: LL (LOC PTE PTE a ar: | A de À à dus | All Des u An e. Wine Sy | È Pube th AD ME Ù 7. { Ato av; ee Kr = REVUE SUISSE DE ZOOLOGIE 112 (2): 333-349; juin 2005 Description of a new beetle-like psocid (Insecta: Psocoptera: Protroctopsocidae) from Turkey showing an unusual sexual dimorphism Charles LIENHARD Muséum d’histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. E-mail: charles.lienhard@mhn.ville-ge.ch Description of a new beetle-like psocid (Insecta: Psocoptera: Protrocto- psocidae) from Turkey showing an unusual sexual dimorphism. - Reticulopsocus besucheti gen. n., sp. n. is described and illustrated from a series of specimens of both sexes collected under stones in southern Turkey. The new genus is closely related to the Mediterranean genera Chelyopsocus Lienhard and Philedaphia Lienhard. Some illustrations of the type genus of the family, Protroctopsocus Mockford, are also provided. Both sexes of the new genus are elytropterous and beetle-like in habitus but the fore wing venation is strongly sexually dimorphic: males have a normal venation, though somewhat simplified, females however, have a highly apomorphic, reticulate venational pattern. A key to the genera of the family Protrocto- psocidae is provided. Keywords: Reticulopsocus - Protroctopsocus - Chelyopsocus - Philedaphia - new genus - new species - wing venation - elytroptery - soil fauna - Mexico. INTRODUCTION Sometimes spectacular discoveries within a particular insect group are not due to the targeted collecting of a specialist but to the quick-witted reaction of a non-spe- cialist encountering an unusual representative of a group, which he did not really search for. So it is not surprising that litter-dwelling beetle-like psocids are more effectively collected by coleopterists interested in soil-fauna than by psocidologists. This is the case in the Oriental genus Coleotroctellus Lienhard, 1988 (Electrento- moidea: Troctopsocidae) where three of the four known species have been discovered by entomologists searching for soil Coleoptera by sifting plant debris in forests. C. burckhardti and C. loebli were collected by my Swiss colleagues Daniel Burckhardt and Ivan Löbl, C. venosus by the Russian coleopterist Sergei Kurbatov. A similar and very recent collecting success is due to my colleague Claude Besuchet, well known specialist of pselaphine Coleoptera and former curator at the Natural History Museum of Geneva. When turning stones in a pine forest in southern Turkey in May 2004, he immediately recognized some fast-moving dark brown “peculiar beetles” on the undersurfaces of the stones and identified them as unusual Manuscript accepted 24.01.2005 334 C. LIENHARD psocids. Remembering my interest in these insects, he made a special effort to collect a representative series of these intriguing creatures. Back to Geneva he immediately gave them to me, commenting that he had apparently found two kinds (two species?) of very particular psocids, both of them with dark brown elytriform fore wings, one of them with a characteristically reticulate structure, the other with a relatively simple venation. What he initially thought to be two different species are actually males and females of a new electrentomoid psocid exhibiting one of the most intriguing cases of sexual dimorphism ever observed in the order Psocoptera. This species is here consi- dered as a representative of a new genus of the family Protroctopsocidae Smithers, which is already represented in the Mediterranean region by the genera Chelyopsocus Lienhard and Philedaphia Lienhard. Mockford (1967) described the very particular monotypic Mexican genus Protroctopsocus in the family Troctopsocidae. Smithers (1972: 338) later placed Protroctopsocus Mockford in a subfamily of its own: Protroctopsocinae Smithers. Based on the short phylogenetic discussion presented by Lienhard (1995), two other troctopsocid genera, Chelyopsocus Lienhard and Philedaphia Lienhard, have been assigned to this subfamily by Lienhard & Mockford (1997). Subsequently elevated to family rank, this taxon has been assigned to the superfamily Electrentomoidea Enderlein (see Enderlein, 1911) (suborder Troctomorpha, infraorder Amphientometae) by Lienhard & Smithers (2002). The superfamily Electrentomoidea sensu Lienhard & Smithers (2002) contains four relatively small families comprising 20 genera, most of them monotypic (12 genera), the others containing only few species (6 genera with 2-5 spp., 2 with 7-8 spp.) (see Lienhard & Smithers, 2002 and Lienhard, 2003). All genera are characterized by striking autapomorphies but their relationships are still poorly understood (see Lienhard & Mockford, 1997). It is not the purpose of this paper to analyse these rela- tionships but to introduce a new species representing a new genus and to present some morphological characters, which could be of phylogenetic significance for future analyses. A sample of the new species has also been sent to Kazunori Yoshizawa for DNA extraction (see “Material and Methods”). The family Protroctopsocidae is characterized by the following synapomorphies of its genera: presence of larval glandular hairs on thorax and abdomen (at present only confirmed in Protroctopsocus and Philedaphia, see Lienhard, 1995; nymphs of other genera not known); AP joined to vein m; endophallus with a conspicuous eversible pouch, sac-shaped (Figs 10, 11, 29) or tube-shaped (Lienhard, 1995: fig. 24) when everted and characteristically folded in resting position (Fig. 9 and Lienhard, 1995: fig. 23 and Garcia Aldrete, 1982: fig. 10) (this character not confirmed in Chelyopsocus, the male of which is not known). The main diagnostic characters of protroctopsocid genera are mentioned in the key below. At present the following four genera and five species are known: Protroctopsocus Mockford, 1967: P. enigmaticus Mockford, 1967 (Mexico), Sd 2.- See Mockford (1967) and Garcia Aldrete (1982). Chelyopsocus Lienhard, 1980: C. garganicus Lienhard, 1980 (Italy), 9 (d un- known). - See Lienhard (1980, 1998). NEW BEETLE-LIKE PSOCID 335 Philedaphia Lienhard, 1995: P. aphrodite Lienhard, 1995, type species (Cyprus), 4 2. - P. hauseri (Lienhard, 1988) (Greece), 2 (male unknown, partheno- genetic). - See Lienhard (1988, 1995, 1998). Reticulopsocus gen. n.: R. besucheti sp. n. (Turkey), d 9. - See description below. MATERIAL AND METHODS The following protroctopsocids have been examined for this study: Reticulopsocus besucheti gen. n., sp. n.: type material (MHNG), see description below. NOTE. Thorax (except fore wings) and basal half of abdomen of one paratype male (MHNG 7625) were sent to Kazunori Yoshizawa (SEHU) for DNA extraction in June 2004; after their return, the exoskeleton of these parts was mounted on a per- manent slide together with the other parts of the specimen. Chelyopsocus garganicus Lienhard: type material (2 2 , MHNG), see Lienhard (1980). Philedaphia aphrodite Lienhard: type material (dd and 2 2, MHNG), see Lienhard (1995). Philedaphia hauseri (Lienhard): type material (9 2, MHNG), see Lienhard (1988). Protroctopsocus enigmaticus Mockford: 14,19 (both slightly brachypterous, with elytriform fore wings) (MHNG), MEXICO: Mexico, K. 87, Toluca - Tejupilco, 1750m, 4.V1.1988, leg. A. Cadena, L. Cervantes. Habitus photographs were made with the AutoMontage® system using a JVC® video camera mounted on a Leica MZ APO stereomicroscope and slightly reworked with ADOBE Photoshop®. Abbreviations used in the descriptions: A = antenna length; AP = areola postica (cubital loop in fore wing); BL = body length (in alcohol); f1, f2, f3, f4 = length of basal four antennal flagellomeres; F = hind femur length; FW = fore wing length; IO/D = shortest distance between compound eyes divided by antero-posterior diameter of compound eye in dorsal view of head; P4 = fourth (apical) article of maxillary palpus; PS = pterostigma (fore wing); T = hind tibia length; t1, t2, t3 = tarsomeres of hind tarsus (lengths measured from condyle to condyle). For standard abbreviations concerning wing venation, see Lienhard (1998). Other abbreviations: BISU = Biological Sciences, Illinois State University, Normal; MHNG = Muséum d’histoire naturelle de la Ville de Genève, Switzerland; SEHU = Systematic Entomology, Hokkaido University, Sapporo; UNAM = Universidad Nacional Autönoma de México. DESCRIPTION AND DISCUSSION Reticulopsocus gen. n. Diagnosis (see also key to the genera of Protroctopsocidae below). Brachy- pterous and flightless, general habitus somewhat beetle-like (Pl. 1 AB) due to blackish brown elytriform fore wings with thickened veins; venation slightly simplified in male, forming a highly apomorphic, reticulate pattern in female. Hind wings strongly 336 C. LIENHARD PLATE 1 Reticulopsocus besucheti gen. n., sp. n., habitus in alcohol, dorsal view, legs not visible on photograph. A: female paratype. B: male holotype [epiproct and paraprocts not visible, they were hidden by the extruded phallic structures (cf. Figs 10, 11) that have been deleted from the photograph]. NEW BEETLE-LIKE PSOCID 337 PLATE 2 Habitus, in alcohol. A: Philedaphia aphrodite Lienhard, female paratype (dorsal view). B: Chelyopsocus garganicus Lienhard, female paratype (dorso-lateral view; with right/left reversal of photograph for publication; left wing of the same specimen illustrated in Lienhard, 1998: pl. 9g). 338 C. LIENHARD reduced, veinless (Fig. 3). Antenna 14-segmented. Frons about equal in length to post- clypeus in anterior view. Ocelli present in male, close to each other, absent in female. Outer cusp of lacinial tip distinctly bidenticulate (Fig. 19), incisive region of mandibles acute (Fig. 21). Fore femur with a longitudinal ventral row of particularly well-deve- loped sculptural denticles on anterior side (Fig. 22). Pretarsal claw with one preapical denticle and some ventral and lateral microtrichia (Fig. 23). Principal veins of fore wing not clearly recognizable in female (PI. 1A, Fig. 1). Venation of male fore wing (Pl. 1B, Figs 4-6) similar to that of other genera of the family, but slightly simplified and somewhat variable; membrane of wing cells slightly concave. Epiproct and paraproct (Figs 8, 14) with some simple stout marginal setae in both sexes, setae on weakly differentiated sense cushion of paraproct without basal rosettes. Lateral hind margin of male clunium simple, roughly right-angled, without densely pilose posterior lobe (Fig. 8). Hypandrium simple (Fig. 7). Phallosome V- shaped (Figs 9, 11), lacking medio-apical sclerite; lateral struts simple but curved to the middle at their apex; latero-apical sclerites elongated, articulated to the apical angle of lateral struts close to a pore-bearing membranous zone. Endophallic pouch sac-shaped in everted position (Fig. 10), centro-ventral pore-bearing lobe of endophallus apically bilobed, its position varying according to the degree of eversion of the endophallic sac (cf. Figs 9, 11). Female subgenital plate (Fig. 12) with 2 stout apical setae and with sclerotized latero-dorsal zones on margin, these joined to T-shaped sclerite by small sclerotized bridges. External valve of gonapophyses (Fig. 13) deeply bilobed. Sperma- thecal duct (Figs 15, 16) relatively short, straight and of same width throughout its length; spermapore situated on a slightly sclerotized oval zone surrounded by fine wrinkles; spermathecal sac with a distinct basal zone bearing a field of inward-directed papillae near opening of duct; wall of spermathecal sac loosely covered with small pores each bearing a short outward-directed microtubular appendix (Fig. 17). Type species. Reticulopsocus besucheti sp. n. Etymology. The genus name is of masculine gender and refers to the reticulate venational pattern of female fore wing. Remarks. See discussion of the type species. Reticulopsocus besucheti sp. n. Fics 1528 PIS MATERIAL Holotype ¢. TURKEY (Mersin Province): Toros mountains, valley from Mersin town up to village Arslanköy, close to village Aladag, 850 m, forest of Pinus sp., under stones, 2.V.2004, leg. C. Besuchet (no. 34); the specimen has been dissected and mounted on two slides (MHNG 7645). Allotype 9, same data as for holotype; dissected and mounted on two slides (MHNG 7646). Other paratypes: 4d ,52, same data as for holotype. ETYMOLOGY The species is dedicated to Dr Claude Besuchet who collected the type material (see also “Introduction”). DESCRIPTION MALE. Coloration. See Pl. 1B. Body generally medium to dark brown, vertex light brown with some darker spots. Compound eye uniformly dark brown (observed after some months in alcohol). Antenna and maxillary palpus light brown; legs light to NEW BEETLE-LIKE PSOCID 339 Fics 1-6 Reticulopsocus besucheti gen. n., sp. n.: 1, right fore wing (9 allotype); 2, idem, detail of wing sculpture (in apical half of wing); 3, hind wing (© allotype, coloration not shown); 4, right fore wing (d holotype); 5, left fore wing (6 paratype MHNG 7625), venation (5a) and detail of sculpture (5b) in centre of wing (coloration not shown); 6a-c, schematic presentation of variability of male fore wing venation (see text). Scale bars: 0.2 mm (Figs 1, 4, 5a; 3). medium brown, with dark brown coxae and femora. Membrane of fore wing uniformly dark brown, glossy, veins blackish brown. Abdomen with dense red-brown hypodermal pigmentation on dorsal and ventral side, terminalia brown. Morphology. See generic diagnosis, with the following complements. Vertex antero-posteriorly rounded, outline slightly undulated in frontal view (cf. Garcia Aldrete, 1982: fig. 1), due to the presence of a shallow dimple on each side, about half 340 C. LIENHARD way between compound eye and vertical suture; frontal suture also visible. Vertex with distinct tubercular sculpture, locally forming a more or less scale-like pattern, pilosity very short. Antennal flagellomeres with annulate sculpture and relatively long slightly wavy hairs (length several times the width of flagellomere). Maxillary palpus as in Fig. 20; sensilla on distal margin of labrum as in Fig. 18, the placoid sensilla lacking central cones. Internal microtrichial field of galea (mentioned for Philedaphia by Lienhard, 1998: fig. 44b) absent or only weakly developed. Prodorsum short, dorsally covered by prominent and almost sharp-edged anterior margin of mesonotum fitting into the occipital margin of the head-capsule (contracted condition of the body). Sculpture of mesonotum similar to that of vertex. Elytriform fore wings in resting position joined along posterior margin to form a hemispherical sclerotized capsule covering the soft abdomen. Veins of fore wing with distinct rounded tubercles, dorsal membrane glossy, only finely granular and locally with fine reticulate pattern (Fig. 5b). Vein an2 absent. Observed variability of wing venation (often asymmetrical on same individual; totally 9 wings of 5 males examined; right wing of paratype male MHNG 7625 lost during capture): PS basally closed (Fig. 4) in 7 wings, open (Fig. 5a) in 2 wings; rudimentary second section of sc present in 3 wings, bifurcating from r proximally to rl-rs bifu- cation (Fig. 5a: 1 wing) or bifurcating from basal section of sc near its meeting point with r (Fig. 6a: 2 wings); rs and m basally meeting in a point (Fig. 4) or fused for a short length (Fig. 5a) and in 1 wing distally joined by a crossvein (Fig. 5a: cell R5 sub- divided in middle), the latter completely absent (5 wings) or rudimentary (3 wings) (PI. 1B and Fig. 4); in 1 wing r4+5 fused to ml for the portion before the wing margin, resulting in a closed cell RS (Fig. 6b); cul (1. e. AP) distally fused to m3, resulting in a large AP apparently joined to m by a short crossvein and m appearing only 2- branched (Figs 4, 5a: 6 wings), or proximal section of cul completely absent, resulting in an apparently 3-branched m and absence of AP due to simple vein cu (Fig. 6c: 3 wings). Hind wing strongly reduced (cf. Fig. 3) and almost vertically protruding from thorax (transparent through fore wings in PI. 1B). Pearman-organ of hind coxa incom- plete, only tympanum visible, coxal rasp not differentiated; tibiae with two apical spurs; number and position of microtrichia of pretarsal claws somewhat variable, usually three relatively long seta-like ventral microtrichia at base and on ventral edge (cf. Fig. 23). Phallosome in resting position as in Fig. 9, with everted endophallic sac as in Fig. 11, hook-shaped lateral struts medially separated (Fig. 11) or joined by a small sclerotized bridge (Fig. 9); membrane near apex of latero-apical sclerite on each side with a basally directed, slightly sclerotized longitudinal zone (Figs 9, 11). Measurements (3 holotype, in um). BL = 1450; FW = 1280; A = 1510; fl = MSP = 150; 3) = 155; f4 = 160: F = 365; = ST UE 289; 12) 5373-8 = Ds FEMALE. Coloration. Same as in male, but membrane of fore wing dull (PI. 1A). Morphology. See generic diagnosis and description of male, with the following complements. Sculpture of vertex with large rounded tubercles (their diameter about twice the diameter of the alveoli of the fine hairs on vertex), usually not forming scale- like pattern. Pilosity of antennal flagellum much sparser and shorter than in male. NEW BEETLE-LIKE PSOCID 341 Fics 7-11 Reticulopsocus besucheti gen. n., sp. n., male (paratype MHNG 7625: Figs 7-9, holotype: Figs 10-11): 7, hypandrium (pilosity not shown); 8, epiproct, right paraproct, right angle and hind margin of clunium (coloration not shown, brown cuticular pigmentation of epiproct and para- proct delimited by dotted lines); 9, phallosome (ventral view, endophallic sac not everted); 10, phallosome with everted endophallic sac, in situ (lateral view, schematic, hypandrium hatched); 11, phallosome with everted endophallic sac, slide-mounted (dorsal view, centro-ventral pore- bearing lobe of endophallus visible by transparency, lower magnification than in Fig. 9). 342 C. LIENHARD Fics 12-17 Reticulopsocus besucheti gen. n., sp. n., female (allotype: Figs 12-15, paratype MHNG 7648: Figs 16-17): 12, subgenital plate (general coloration of ventral side not shown, brown cuticular pigmentation anteriorly delimited by dotted line); 13, gonapophyses; 14, epiproct and right paraproct (coloration not shown, brown cuticular pigmentation delimited by dotted lines); 15, spermatheca (empty spermatheca of virgin female); 16, spermatheca of mated female, with spermatophore (sperm packet) in basal zone of sac; 17, pore with microtubular appendix of spermathecal membrane (scale bar 0.02 mm). NEW BEETLE-LIKE PSOCID 343 FA yon ; mg MA 23 Fics 18-24 Figs 18-23. Reticulopsocus besucheti gen. n., sp. n., female allotype: 18, sensilla on distal mar- gin of labrum; 19, lacinial tip; 20, maxillary palpus (pilosity not shown); 21, mandibles (anteri- or view); 22, anterior side of fore femur (base left, apex right; general sculpture not shown); 23, pretarsal claw. - Fig. 24. Chelyopsocus garganicus Lienhard, female holotype: mandibles (ante- rior view, same magnification as Fig. 21). Principal veins of fore wing not clearly recognizable, except vein ani and basal parts of r and m+cu visible as particularly well-differentiated straight venational structures within the reticulate pattern covering the whole wing (Pl. 1A, Fig. 1) and delimiting small cells of irregular shape, these cells sometimes partially open due to incomplete veins (see Fig. 1). Veins with distinct, rounded tubercles; dorsal membrane slightly rugose and reinforced by sclerotized circular plaques (Fig. 2). Dorsal valve of gonapophyses particularly broad (Fig. 13). The terminalia of two female paratypes have been dissected, one of them (MHNG 7646 allotype) proved to be virgin, its spermatheca being shrivelled and empty (Fig. 15), in the other one (MHNG 7648) a simple spermatophore (sperm packet) could be observed in the basal zone of the spermathecal sac (Fig. 16). Measurements (9 allotype, in um). BL = 1730; FW = 1470; A = 1460; fl = 120; 2450 15 = 155,14 = 170; F= 40057. = 610; tl = 285312 = 55313, 887I0/D 1.7. 344 C. LIENHARD BIOLOGY The type series was collected under stones lying on the soil of a relatively open forest of Pinus sp., situated in a mountainous region of southern Turkey, about 30 km from the Mediterranean coast. When disturbed, these black, hemispherical, beetle-like insects moved quickly on the underside of the stones (C. Besuchet, pers. comm.). DISCUSSION The definition of the family Protroctopsocidae and its generic and specific com- position have already been presented in the “Introduction”. Within this family the three Mediterranean genera Philedaphia, Chelyopsocus and Reticulopsocus gen. n. undoubtedly form a monophyletic group defined by the following synapomorphies: antenna 14-segmented, female subgenital plate with sclerotized latero-dorsal zones on margin, endophallus with pore-bearing central lobe. Its sister-group is the Mexican genus Protroctopsocus, which is characterized by several plesiomorphic features and by the autapomorphic basal enlargement of P4 and the complex sclerotizations around the spermapore (Fig. 26). The three Mediterranean genera can be assigned to two phylogenetic clades, one comprising Philedaphia, characterized by the complex phallosome (see key below), and the other comprising Chelyopsocus and Reticulo- psocus. The latter two genera have a very similar spermatheca structure (simple spermapore, membrane of spermathecal sac with pores bearing microtubular appen- dices; but spermatheca lacking inward-directed basal papillae near opening of duct in Chelyopsocus) which can be interpreted as a probable synapomorphy. In both other protroctopsocid genera (i.e. Protroctopsocus and Philedaphia) the membrane of the spermathecal sac lacks basal papillae and pores with microtubular appendices; however, in Philedaphia this membrane bears regularly distributed inward-directed spine-like structures (see Lienhard, 1988: fig. 7 and 1995: fig. 14), which are probably homologous to the inward-directed, small, rugose plates present in Protroctopsocus (Figs 27, 28). Chelyopsocus is characterized by the clearly apomorphic blunt incisive region of the mandibles (Fig. 24; this newly observed character state was confirmed for both specimens known of this genus, holotype and paratype females of C. garganicus); in all other protroctopsocid genera this region is acute, corresponding to the plesio- morphic condition within Psocoptera (cf. Fig. 21). The broad and only weakly sub- divided outer cusp of the lacinial tip can also be interpreted as an autapomorphy of this genus; all other protroctopsocids show a distinct indentation (cf. Fig. 19). Occasional brachyptery with a tendency towards elytriform fore wings can be observed in Protroctopsocus, but it is without sexual dimorphism. Elytroptery seems to be genetically fixed in the female of Chelyopsocus (male not known) and in both sexes of Reticulopsocus, where a strong sexual dimorphism occurs. The male exhibits a “moderate elytroptery” (with essentially normal wing venation, similar to that present in the female of Chelyopsocus and in the brachypterous morph of Protroctopsocus), whereas the female shows a “perfect elytroptery” with densely reticulate fore wings forming a regularly vaulted and uniformly structured, hemispherical covering of the abdomen strongly resembling elytra in Coleoptera. NEW BEETLE-LIKE PSOCID 345 Fics 25-31 Figs 25-30. Protroctopsocus enigmaticus Mockford (female: Figs 25-28; male: Figs 29-30): 25, subgenital plate (general coloration of ventral side not shown, brown cuticular pigmentation an- teriorly delimited by dotted line); 26, gonapophyses and sclerotizations around spermapore; 27, spermatheca; 28, rugose areas of spermathecal membrane (scale bar 0.02 mm); 29, phallosome with everted endophallic sac, in situ (lateral view, schematic, hypandrium hatched); 30, phallo- some (ventral view). - Fig. 31. Philedaphia aphrodite Lienhard, male paratype: medio-apical sclerite of phallosome (coloration not shown) and membranous pore-bearing central lobe of en- dophallus (ventral view, endophallic sac not everted). The presence of morphologically so different brachypterous morphs within the same species has never been observed before in Psocoptera. In the other electrento- moid genus with beetle-like females, the litter-dwelling Coleotroctellus Lienhard (see “Introduction”), males are always macropterous, with normal venation and fully 346 C. LIENHARD developed hind wings. In that genus the mobility of the males apparently represents a more important evolutionary advantage than it could be offered by an efficient abdominal protection in both sexes (Lienhard, 2002). In Rericulopsocus this is appa- rently not the case. Males have also become flightless and perhaps they are on the way to evolve towards a “perfect elytroptery” as it has already been genetically fixed in females. The many venational aberrations observed in males (see description) may be interpreted as an indication of a diminishing selection pressure for “normal” venation in these brachypterous and flightless animals. In Coleotroctellus the females show a beetle-like habitus similar to that of the female of Reticulopsocus. In both genera shape and length of the dark brown female fore wings are about the same. However, in Coleotroctellus the sclerotized vaulted wing membrane is stabilized by a series of thickened longitudinal veins derived from the normal venation of the family Troctopsocidae, which is very similar to the normal venation of Protroctopsocidae. In some species these veins are relatively easy to homologize with the normal venation (C. burckhardti and C. loebli, see Lienhard, 1988 and Lienhard & Mockford, 1997), but in C. venosus the multiplication of longitudinal veins makes this homologization very difficult. C. venosus apparently is the most derived form resulting from an evolutionary trend towards “perfect elytroptery”, where the vaulted fore wing membrane is mechanically stabilized by numerous parallel longitudinal ribs (see Lienhard & Mockford, 1997: figs 8-10). As we have seen above, another type of “perfect elytroptery” has evolved in Reticulopsocus, based on a different constructional principle, which consists of “wire-netting” the vaulted fore wing membrane by a dense reticulate venational pattern (Pl. 1A and Fig. 1). Reticulate fore wings are very rare in Psocoptera. Some distinct but very fine reticulate sculpture can be observed in the strongly reduced scale-like fore wings of Lepinotus reticulatus Enderlein (Trogiidae, suborder Trogiomorpha) and in the parchment-like elytriform fore wings of Badonnelia titei Pearman (Sphaeropsocidae, suborder Troctomorpha) (L. reticulatus: see Lienhard, 1998: fig. 26d; B. titei: see Pearman, 1958: fig. 6, New, 1974: fig. 130, Lienhard, 1982: fig. 3). But in these cases the reticulation is not directly derived from the wing venation. The only family, where extensive venational reticulation in fore wings is currently known are the Calopsocidae (suborder Psocomorpha; see Thornton & Smithers, 1984). In some species the reti- culate venation is especially well-developed in the apical half of the wing, but species with almost completely reticulate wings are also known (e. g., Calopsocus reticuloides Thornton & Smithers). In Calopsocidae this reticulate condition is usually also combined with some wing shortening and a slight tendency towards elytroptery, but hind wings are always well-developed, with normal venation, and probably these pso- cids are all able to fly. Sexual dimorphism in wing structure is not known in Calopsocidae (see male and female of C. reticuloides, Thornton & Smithers, 1984: figs 116 and 121). Calopsocids usually live on vegetation and with a fore wing length of several millimeters they are all much larger than Reticulopsocus. Probably selection pressure leading to reticulate fore wings differs considerably in these phylogenetically very distant groups. NEW BEETLE-LIKE PSOCID 347 KEY TO GENERA OF PROTROCTOPSOCIDAE NOTE. Figures of Protroctopsocus have been published by Mockford (1967, female) and Garcia Aldrete (1982, male) and additional information is given in Figs 25-30 of the present paper. For Chelyopsocus and Philedaphia see Figs 24 and 31 and figures in Lienhard (1998). The male of Chelyopsocus is not known. All known species of Protroctopsocidae are mentioned in the “Introduction”. 1 Antenna 15-segmented. Head capsule clearly elongated in anterior view, frons distinctly longer than postclypeus (Garcia Aldrete, 1982: fig. 1). P4 basally enlarged, its maximal width near base (Mockford, 1967: fig. 46 and Garcia Aldrete, 1982: fig. 8). Fore femur with a longitudinal ventral row of short articulated spines on anterior side (Mockford, 1967: fig. 29; Garcia Aldrete, 1982: fig. 4). Female subgenital plate (Fig. 25) with 2-5 (usually 4)!) stout apical setae, lacking sclerotized latero-dorsal zones on margin. Phallosome (Fig. 30) V-shaped, without medio-apical sclerite, lateral struts straight, simple; endophallus lacking pore-bearing central lobe. Fore wing not sexually dimorphic, normally shaped in macropterous morph (Mockford, 1967: fig. 26), slightly elytriform with concave wing cells but with normal venation in brachypterous morph (Garcia Aldrete, 1982: fig. 2); this morph known in both sexes NET, CC ORR: PUMA), BEINE Protroctopsocus Mockford - Antenna 14-segmented. Head capsule weakly elongated, nearly circular in anterior view, frons not or only slightly longer than postclypeus. P4 nearly spindle-shaped, its maximal width roughly in the middle (Fig. 20). Fore femur with a longitudinal ventral row of particularly well-developed sculptural denticles (not articulated!) on anterior side (Fig. 22). Female subgenital plate (Fig. 12) with 2 (rarely 3) stout apical setae and with sclerotized latero-dorsal zones on margin, these more or less distinctly joined to T-shaped sclerite by small sclerotized bridges. Phallosome relatively complex, with medio-apical sclerite (Lienhard, 1998: fig. 45d) or lateral struts curved inward (Figs 9, 11); endophallus with a pore-bearing central lobe (Figs 9, 11, 31). Fore wing normal (Pl. 2A) or strongly elytriform and with modified venation (Pls 1AB, ZB) sometimes;sexuallyzdimorphier. RR 2 2 Incisive region of mandibles blunt (Fig. 24). Apex of outer cusp of lacinial tip broad, only with a slight marginal incision (Lienhard, 1998: fig. 43d). Brachypterous female with elytriform fore wing, venation modified to form concave “honeycombed” cells (Pl. 2B and Lienhard, OSS TE AS ap OR Chelyopsocus Lienhard = Incisive region of mandibles acute (Fig. 21). Apex of outer cusp of lacinial tip with distinct indentation (Fig. 19). Brachypterous female 1) Mockford (1967: fig. 37) illustrated a subgenital plate with only 2 stout apical setae; the spe- cimen examined has 5 such setae (Fig. 25). In 1979 Mockford (in litt.) stated: “4 stout setae is the usual situation for both forms” [i. e. for macropterous and brachypterous morphs]. 348 C. LIENHARD with normally shaped fore wing and slightly reduced venation (Lienhard, 1998: fig. 46a, pl. 3a) or with reticulate elytriform fore wing (Eisslkand Bl. VA) RASE CAL ee PROIETTI NP 3 3 Fore wing normally shaped in macropterous morph (male and female) (Pl. 2A) and in brachypterous morph (female), venation normal in both morphs but slightly reduced in brachypterous females. Phallosome U-shaped (Lienhard, 1998: fig. 45d), with medio-apical sclerite; lateral struts of phallosome doubled, not curved to the middle at their apex. Angle of lateral hind margin of clunium with a densely pilose posterior lobes(Ieienhard 998 show 4 5a). ee seer ie Philedaphia Lienhard - Only brachypterous morph known (male and female), fore wing elytri- form with somewhat simplified venation in male (Figs 4-6; Pl. 1B) and with reticulate venational pattern in female (Fig. 1; Pl. 1A). Phallosome V-shaped (Figs 9, 11), lacking medio-apical sclerite; lateral struts simple but curved to the middle at their apex. Lateral hind margin of clunium simple, roughly right-angled, without densely pilose posterior lobe (Riso) case oa ono rn ao oro ae MAME core eer ale Reticulopsocus gen. n. ACKNOWLEDGEMENTS Many thanks go to Claude Besuchet, former curator at the MHNG, for the gift of the Reticulopsocus specimens. A detailed comparison with Protroctopsocus enig- maticus was possible due to the donation of two specimens of this species by Alfonso Garcia Aldrete (UNAM). I am also very grateful to Edward Mockford (BISU) for dis- cussion on electrentomoids and to him and Peter Schwendinger (MHNG) for critical reading of the manuscript. I thank the following colleagues for assistance in producing the AutoMontage® photographs and for preparing them for publication: Florence Marteau, Bernard Landry, Peter Schwendinger (all MHNG). REFERENCES ENDERLEIN, G. 1911. Die fossilen Copeognathen und ihre Phylogenie. Palaeontographica 58: 279-360. GARCIA ALDRETE, A. N. 1982. Descripcion del macho de Protroctopsocus enigmaticus Mockford y registros de distribucion de la especie (Psocoptera: Troctopsocidae). Anales del Insti- tuto de Biologia, Universidad Nacional Autonoma de Mexico, Ser. Zool.52(1): 119-122. LIENHARD, C. 1980. Chelyopsocus garganicus n. gen., n. sp., eine neue lapidicole Psocoptere aus Süditalien (Psocoptera: Troctopsocidae). Mitteilungen der Schweizerischen Entomo- logischen Gesellschaft 53: 209-214. LIENHARD, C. 1982. Les Psoques de nos maisons ou les tours malicieux des appareils Berlese. Musées de Genève 222: 16-21. LIENHARD, C. 1988. Three new extra-neotropical species of Troctopsocidae (Insecta: Psocoptera). Journal of Natural History 22: 575-587. LIENHARD, C. 1995. Psocoptères (Psocoptera) nouveaux ou peu connus d’Italie, de Chypre et du Yémen. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 68: 335-361. LIENHARD, C. 1998. Psocoptères euro-méditerranéens. Faune de France 83: XX + 517 pp. LIENHARD, C. 2002. Three extraordinary new species of Psocoptera (Insecta) from Colombia, Malaysia and Thailand (Epipsocidae, Lachesillidae, Ectopsocidae). Revue suisse de Zoologie 109(2): 383-395. NEW BEETLE-LIKE PSOCID 349 LIENHARD, C. 2003. Additions and Corrections (Part 1) to Lienhard & Smithers, 2002: “Psocoptera (Insecta) - World Catalogue and Bibliography” (pp. 2-24). In: YOSHIZAWA, K. (ed.). Psocid News. The Psocidologists’ Newsletter. No. 4 (February 17, 2003). Systematic Entomology, Faculty of Agriculture, Hokkaido University, Sapporo, 24 pp. Online at http://insect3.agr.hokudai.ac.jp/psoco-web/psoco-net/index .html LIENHARD, C. & MOCKFORD, E. L. 1997. New taxa and records of Troctopsocidae (Psocoptera). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 70: 361-385. LIENHARD, C. & SMITHERS, C. N. 2002. Psocoptera (Insecta): World Catalogue and Bibliography. Instrumenta Biodiversitatis 5. Muséum d’histoire naturelle, Genève, XLI + 745 pp. MocKFORD, E. L. 1967. The electrentomoid psocids (Psocoptera). Psyche 74(2): 118-165. NEW, T. R. 1974. Psocoptera. Handbooks for the identification of British Insects 1(7): 1-102. PEARMAN, J. V. 1958. Augmented description of Badonnelia titei Pearman (Psoc., Sphaero- psocidae), with notes on the genus Sphaeropsocus. Entomologist's Monthly Magazine 94: 49-52. SMITHERS, C. N. 1972. The classification and phylogeny of the Psocoptera. Australian Museum Memoirs 14: 1-349. THORNTON, I. W. B. & SMITHERS, C. N. 1984. Systematics of the Calopsocidae, an Oriental and Melanesian family of Psocoptera. Systematic Entomology 9(2): 183-244. RE Le i Ù co wi on | page Dent ul i. "+ Literie > SO Ù du è don Alle Ù à | 7 L È : i | ; i | bll > ; A | pa VERI REVUE SUISSE DE ZOOLOGIE 112 (2): 351-369; juin 2005 Revision of the Oriental Genus Loeblites Franz (Coleoptera, Scydmaenidae) Pawet JALOSZYNSKI Wichrowe Wzgörze 22/13, 61-678 Poznan, Poland. E-mail: japawel@man.poznan pl Revision of the Oriental genus Loeblites Franz (Coleoptera, Scydmae- nidae). - The Oriental scydmaenid beetle genus Loeblites Franz is revised. Loeblites mastigicornis Franz from Thailand and L. sabahensis Franz from Borneo are redescribed, and a new species, L. minor sp. n. from Borneo is described. The general morphology of the genus is described and illustrated in details, including mouthparts, wings, female and male genitalia, and an identification key to the species of Loeblites is given. Keywords: Coleoptera - Scydmaenidae - Loeblites - revision - morphology - new species - Oriental region - taxonomy. INTRODUCTION The genus Loeblites (Scydmaeninae, Cyrtoscydmini) was established by Franz (1986) for a single species, L. mastigicornis, on the basis of four individuals collected in Thailand. A few years later L. sabahensis was described by the same author from Borneo (Franz, 1992). These two species were the only members of the genus known so far. As mentioned previously (Jatoszyfiski, 2004), Loeblites shows a high degree of similarity to Syndicus Franz and Horaeomorphus Schaufuss; the three genera may form a monophyletic group within the Cyrtoscydmini. Details of morphology have been described only in Syndicus (Jatoszyñski, 2004); therefore, a comprehensive comparison of all three genera will be possible when respective structures have been described in Horaeomorphus (Jatoszyñski, in preparation). The purpose of the present paper is to provide a clear diagnosis of Loeblites and a detailed description of the morphology. Existing descriptions of the two known species lack important details (e.g., spermathecae, mouthparts, thoracic morphology), and during the present study I discovered that the type series of L. sabahensis consisted of two different taxa. Therefore, herein L. mastigicornis and L. sabahensis are redescribed, the detailed mor- phology of the genus is given (including previously unknown characters), and a new species, L. minor sp. n., from Borneo is described. METHODS Illustrated anatomical structures were disarticulated, dehydrated in isopropanol, transferred to xylene, and mounted in Canada balsam, or dehydrated in ethanol and mounted in euparal. The measurements are as follows: length of the head is from the Manuscript accepted 05.11.2004 352 P. JALOSZYNSKI occipital constriction to the anterior margin of clypeus; width maximum includes eyes; length of pronotum is measured along midline; length of elytra is measured along su- ture; width of elytra combined, measured at greatest width; elytral index (El): length/width. Mounted specimens often have head and pronotum bent ventrally to a various extent, and therefore a total length measured in dorsal view might be mis- leading; for this reason the total length given in descriptions is the sum of individual measurements of the head, pronotum and elytra. The length of spermatheca is the longest measurement of the capsular part. The nomenclature of morphological details follows that used in Jatoszyfiski, 2004, which was partly adopted from O’Keefe & Monteith (2000), Bordoni & Castellini (1973), and Nomura (1991). Studied specimens are deposited in Muséum d’histoire naturelle, Geneva, Switzerland (MHNG), Natural History Museum, Vienna, Austria (NMW), private collection of the author, Poznañ, Poland (PCPJ), and private collection of Peter Hlaväë, KoSice, Slovakia (PCPH). TAXONOMY Loeblites Franz Loeblites Franz, 1986: 965. Type species: Loeblites mastigicornis Franz, 1986 (des. orig.); Newton & Franz, 1998: 147. Diagnosis. Members of Loeblites can be identified on the basis of the following characters: body of moderate size (2.31-2.82 mm), relatively slender, very convex, light to dark brown, covered with moderately dense and long setation; head wider than long, widest at eyes; mandibles nearly planar, subtriangular with long prostheca, slender and sharp apical tooth and subapical tooth; maxillary palpi with broadened, elongate palpomere III and fusiform, slender palpomere IV; antenna without club, very long and slender, barely broadened toward apex, antennomere XI not modified, well separated from X; pronotum widest near anterior 1/5-!/3, with rounded anterior margin and sides, lateral margins distinctly narrowing posteriorly, without lateral carina or sharp edge, with constriction separating short posterior collar, in dorsal view posterior collar demarcated from disc by transverse row of four pits, each side of pronotum bears two additional, shallow and indistinctly delimited pits or impressions; scutellum not visible; elytra entire, oval, each elytron with two basal foveae located very close to elytral insertion, so that pits are partly covered by posterior margin of pronotum, base of elytra usually with shallow internal humeral impressions; all femora with slender basal stalk-like part and strongly clavate distal part with narrow dorsal femoral groove and very small circular or oval pore, presumably a gland opening. Aedeagus sym- metrical or with minimally asymmetrical internal armature, elongate, with long, asetose parameres and relatively complicated, darkly sclerotized internal sac; sper- matheca well sclerotized, usually dark brown, ovoid with tubular or funnel-like exca- vation and approximate insertions of very thin ductus spermathecae and thicker duct of accessory gland; bursa copulatrix variously developed, in one case missing, in remaining two known species relatively large and dark, asymmetrical to symmetrical. Redescription. The following detailed description of the general morphology is based on disarticulated male and female specimens of Loeblites sabahensis (remarks REVISION OF LOEBLITES 353 on the remaining species are included, if described structures are significantly different). This species was selected because of the abundance of specimens available for dissections. Head (Figs 1-4, 19). In dorsal view head wider than long, widest at eyes; occiput broad, only slightly narrower than vertex, distinctly separated from vertex by constriction and relatively deep transverse groove, tempora long, rounded; vertex about 2.5 times as wide as long, slightly convex, with small and indistinctly delimited median tubercle adjacent to occipital constriction; supraantennal tubercles relatively large and strongly raised; frons relatively short, gradually lowering anteriorly, demar- cated from clypeus by transverse groove. Punctation of vertex and frons relatively fine and sparse; setation sparse and moderately long. Clypeus wider than long, with uneven, granulated surface; anterior tentorial foveae barely visible, located in supraclypeal groove, in anterior margin of head; eyes located near middle of head length, convex, relatively large, in lateral view with emarginate postero-ventral margin, finely faceted; gena and postgena strongly convex, with moderately long setae directed laterally and dorsally. In ventral view gula large, subtriangular with rounded lateral margins; sub- mentum subtriangular, with shallow groove parallel to posterior margin of mentum; posterior tentorial pits not visible; hypomera (sensu Jatoszyfiski, 2004; hypostomae of O’Keefe, 2000) well developed. Antennal cavities broadly separated, located in latero- anterior part of head, under supraantennal tubercles; antenna (Figs 36-38) without club, very long, antennomeres with well delimited basal rings, covered with long, modera- tely dense, suberect to erect setae. Mouthparts (Figs 5-8). Labrum (Fig. 5) wider than long, subtrapezoidal, with straight posterior margin, rounded sides and minimally concave anterior margin; lateral and anterior margins with narrow, lightly sclerotized velum. Dorsal surface of labrum with seven pairs of symmetrically located setae of various lengths. Mandible (Fig. 6) subtriangular, nearly planar, with broad base and curved, sharp apical tooth and single subapical tooth. External margin of mandible near posterior condyle bears group of several setae. Prostheca well developed, relatively broad, composed of short setae nearly from base of mandible to about middle of internal margin, which between prostheca and subapical tooth is sharp and irregularly, very finely serrated. Maxilla (Fig. 7) with subtriangular stipes bearing two basal setae; elongate palpifer with two long subapical setae and three or four short setae on ventral surface; elongate galea with row of long and dense setae along internal apical margin and additionally two shorter and thinner setae on apex and on external margin; and long lacinia with dense and long setae along internal margin and two additional thinner sensory setae near middle of internal margin. Palpus maxillaris relatively large, palpomere I very small, subtrapezoidal, with single subapical seta; II elongate and curved, pipe-like, with slender basal part, gradually broadening toward truncate apex, covered with relatively sparse, suberect setae confined to distal 2/;; palpomere III distinctly enlarged, with narrow base, broadened between middle and distal third, then narrowing toward trun- cate apex, with moderately dense, suberect setation; palpomere IV small and long, nar- rowing from base to pointed apex, setation relatively short and dense, recumbent to suberect, apical third to fourth asetose. Labium (Fig. 8) with darkly sclerotized, sub- rectangular mentum with numerous tiny pores on ventral surface; palpomere I about 354 P. JALOSZYNSKI III, EXTER S IE ubmentum n TTT NI SO ifrons! | vertex postgena——> hypomeron occiput frons anterior tentorial On fovea | 3 clypeus 4 Fics 1-4 Loeblites sabahensis Franz: 1 — head, dorsal view, 2 — head, ventral view, 3 — head, anterior view, 4 — head, left lateral view. Scale 0.2 mm. twice as wide as long, broadened in distal half, asetose; palpomere II large, subcylin- drical with rounded sides, nearly three times longer than I, with several long setae; palpomere III long and slender, slightly more than half length of II, narrowing toward pointed apex, asetose. Anterior margin of labium between insertions of palpi bears pair of long bristles in middle; hypostom well developed, with short and dense setation. Prothorax (Figs 9-10, 19). In dorsal view (Fig. 19), pronotum elongate, anterior margin rounded, lateral margins without sharp edges or carinae, rounded, posterior margin arcuate, hind angles indistinct. Pronotum widest near anterior third or fourth; posterior third or less demarcated by transverse row of eight variously developed foveae. In strictly dorsal view, four foveae fully visible and additional pair of lateral foveae partly visible; external, very shallow pits or rather impressions located laterally, visible only in lateral view (Fig. 10). Dorsal surface of pronotum moderately or strongly convex, except for posterior collar, which is flatter than disc. Dorsal surface covered with very dense, round granules (L. mastigicornis) or distinct, but relatively small and sparse punctures (L. sabahensis and L. minor); setation moderately dense, composed of curved, suberect to erect setae. Ventral side of pronotum (Fig. 9) very short, with relatively well developed basisternum; procoxal cavities large, nearly REVISION OF LOEBLITES 355 Pla HO arts mi LARA ANT ow Fics 5-8 Loeblites sabahensis Franz: 5 — labrum, dorsal view, 6 — left mandible, dorsal view, 7 — right maxilla, ventral view, 8 — labium, ventral view. Scale 0.2 mm. circular, procoxal insertions very widely separated; subcoxal foveae small, located close to posterior margin of prosternum, near coxal insertions, in natural position covered by coxae. Hypomera large and well demarcated, elongate, impressed in middle. Ventral surface of pronotum covered with relatively dense, short suberect setation, except for asetose areas under coxae. Mesothorax (Figs 11-12). Mesonotum (Fig. 12) elongate; prescutellum rela- tively short and broad, subtriangular, with slightly concave anterior margin and convex, rounded posterior margin, each lateral margin terminated at short anterior notal wing process; scutellum long and narrow, with rounded apex; postscutellum very short, with deeply emarginate posterior margin; posterior notal wing processes long 356 P. JALOSZYNSKI and slender, extending latero-posteriorly toward latero-anterior margins of metatergal prescutum. Mesosternum (Fig. 11) wider than long, subtrapezoidal; mesepimeron elon- gate, with shallow subtriangular impression; mesepisternum elongate, distinctly convex, partly visible in dorsal view; basisternal area large, broadly and deeply im- pressed in middle, posteriorly basisternum expanded in middle into raised, moderately narrow mesosternal process extending to about anterior third of metasternum, gra- dually raising to posterior !/,-!/;, then lowering toward rounded apex. Coxal insertions small, oval, very broadly separated. Surface of basisternal area uneven, densely covered with long setae; convex parts of mesepimera glabrous and asetose, glossy, concave area on each mesepimeron with dense, short setae; mesepisterna covered with short and dense setae. Metathorax (Figs 11-12). Metanotum (Fig. 12) distinctly wider than long. Prescutum relatively small, transverse, with rounded posterior margin, distinctly demarcated from scutum, laterally connected to wing insertion, postero-laterally to scutum; scutum large, nearly completely divided into two lateral parts by large, trian- gular scutellum, distinctly separated from scutum and from postnotum by sutures; post- notum relatively small, narrow, with convex, rounded posterior margin. Metasternum (Fig. 11) elongate, basisternal area with biemarginate anterior margin, deeply de- pressed between mesosternal process and mesocoxal insertions; meso-metasternal foveae (metasternal foveae in Jatoszynski, 2004; the foveae are actually located between meso- and metasternum) small, in natural position covered by mesocoxae; middle part of anterior margin fused to mesosternal process; posterior 2/3 or more of metasternum convex; posterior margin biemarginate, slightly expanded posteriorly in middle, with relatively shallow median notch. Paracoxal sutures indistinct in dry- mounted specimens, well recognizable in transparent mounts; metasternum bordered on each side by narrow episternum. Setation of metasternum relatively long, moderately dense, suberect, except for asetose areas under mesocoxae. Metafurca with short and broad basal stalk and two long, slender, recurved and widely divergent furcal arms. Elytra (Figs 12, 19) entire, oval, convex, widest anterior to middle, base nearly straight or slightly concave, wider than base of pronotum; humeral callus moderately distinct, internally delimited by relatively short and shallow internal humeral impres- sion; each elytron bears pair of small circular basal foveae connected by U-shaped groove and located in depression between convex discal part and small basal lobe con- nected to mesonotum; in natural position foveae barely visible, covered by posterior margin of pronotum. Elytral punctation not ordered in rows, punctures relatively fine and sparse; elytral setation composed of moderately long, suberect to erect setae. Wing insertion and wing (Figs 12-13). Suralare large, subrectangular with rounded margins; axillary sclerites elongate, the uppermost sclerite large and S-shaped, two remaining sclerites smaller, elongate; subalare moderately large, elongate. Wings (Fig. 13) in all species fully developed, about twice as long as elytra. Wing venation highly reduced, limited to basal half of wing or less; costa highly reduced, very short; subcosta and radius short, distally fused together with anterior margin of wing; posterior venation composed of only two nearly parallel veins, which may represent mediana and cubitus (or fused cubitus + postcubitus). Distal half of wing without traces REVISION OF LOEBLITES 357 basisternum 10 hypomeron subcoxal fovea mesosternum ui \ .\__——— mesepisternum XS mesepimeron meso-metasternal fovea mesosternal process metasternum metepisternum gii - metafürca 1 1 paracoxal suture Fics 9-11 Loeblites sabahensis Franz: 9 — pronotum, ventral view, 10 — pronotum, left lateral view, 11 — mesosternum and metasternum, ventral view. Scale 0.5 mm. of veins, posterior margin bears row of long setae, anterior margin with very short setae, entire wing covered with extremely short, very dense setation. Legs. Procoxae relatively large, subconical, contiguous; mesocoxae slightly larger than procoxae, oval, separated by mesosternal process; metacoxae narrowly 358 P. JALOSZYNSKI separated, transverse, elongate, with well delimited internal posterior part adjacent to trochanters. Trochanters small and longer than wide, protrochanters fully separating procoxae from profemora; meso- and metatrochanters not separating meso- and metafemora from coxae. Femora long, clavate, with very slender proximal part (about half length of femur) and abruptly expanded distal part. Dorsal surface of clavate part of femora bears very narrow but distinct dorsal femoral groove and very small circular or oval pore near the widest place of femur. Tibiae relatively slender and long, straight or minimally recurved, pro- and especially mesotibiae in both sexes bear dense patch of golden setae along distal fourth or fifth of internal margin. Tarsi moderately long, relatively thick, tarsomeres I-IV subequal or only slightly decreasing in length, tarso- mere V elongate, nearly as long as II-IV together. Abdominal sternites. Six abdominal sternites visible (numbered I-VI in the present study), together shorter than metasternum. Sternite I relatively long, but only partly visible from under metacoxae, under coxae concave, with median longitudinal carina not reaching posterior margin of sternite. Sternites II-IV subequal in length, each about as long as half of I; sternite V about as long as 1.5 length of IV, sternite VI sub- triangular, with broadly rounded posterior margin, as long as V or slightly longer. Sutures between sternites arcuate or nearly straight; all sternites with variable puncta- tion and moderately dense, short and slightly suberect setation. Male genital segment (Figs 14-16) composed of tergite IX, sternite IX and tergite X. Tergite IX (Fig. 14) composed of two elongate lateral parts; sternite IX (Fig. 15) relatively large, subtriangular, with broadly emarginate anterior margin and rounded apex bearing several setae of various lengths, lateral margins convex in proxi- mal part and strongly concave in distal part; tergite X (Fig. 16) small, nearly penta- gonal, with anterior margin expanded in middle and posterior margin broadly emarginate. Aedeagus (Figs 20-22, 25-27, 30-32) with large and elongate median lobe widest near base and narrowing toward rounded apex; base with narrow median emargination; parameres long, symmetrical, without apical setae. Armature of internal sac relatively complicated; symmetrical (L. mastrigicornis and L. minor) or slightly to distinctly asymmetrical (L. sabahensis). Female genital segment (Figs 17-18) composed of pair of large, fused para- procts; pair of moderately broad valvifers fused with paraprocts, with dense setation along apical margin; pair of elongate, slender coxites with sparse and relatively long setae; and small proctiger (Fig. 18) with broadly emarginate proximal margin, with long, slender and pointed median projection in distal margin. Bursa copulatrix (Figs 24, 29, 35) highly diversified within the genus; well developed, darkly sclerotized and symmetrical in L. mastigicornis (Fig. 24), relatively lightly sclerotized and asym- metrical in L. sabahensis (Fig. 29), and L. minor lacks visible structures surrounding distal (i.e. genital) end of ductus spermathecae (Fig. 35). Spermatheca (Figs 23, 28, 34) capsular, well sclerotized, relatively dark, with apical excavation leading into internal tube or funnel; ductus spermathecae very thin, inserted in base of spermatheca, near short and thicker duct of very lightly sclerotized, in most cases barely recognizable accessory gland. REVISION OF LOEBLITES 359 anterior not process sal ies en < n. \\ iG Fr Ad suralare postscutellum a: - Pre osterior Rota, = SE ary subalare ae Ì . - 7 metepimeron Sa DI) costa subcosta radius __ mediana cubitus 13 —__—_m—rc Fics 12-13 Loeblites sabahensis Franz: 12 — mesonotum, metanotum, base of right elytron and wing, dorsal view, 13 — right wing, dorsal view. Prscl — prescutellum, prsc — prescutum, sc — scutum, scl — scutellum, psnt — postnotum. Scale 0.5 mm. KEY TO THE SPECIES OF LOEBLITES FRANZ 1 Pronotum densely covered with small, round granules . L. mastigicornis Franz - Rronotumswithoutseranules! saver. Fe errant PER 2 2 Elytral index: males 1.49-1.62, females 1.51-1.56 ....... L. sabahensis Franz - Elytral index: males 1.39-1.47, females 1.44-1.51 ........... L. minor sp. n. 360 P. JALOSZYNSKI Loeblites mastigicornis Franz Figs 20-24, 36 Loeblites mastigicornis Franz, 1986: 966, figs. 1, 2. Type material. Holotype male, white printed label “THAILAND: Chiang Mai, Doi Suthep, 1050 m, 5. XI. 1985, Burckhardt-Lôbl”, white handwritten label “Loeblites mastigicor- nis m.” and printed “det H. Franz”, red handwritten label “Holotypus” (MHNG). Paratypes: 2 males, 2 females, same data (MHNG). Redescription. Body large, slender, very convex, moderately dark brown, an- tennae and tibiae slightly lighter, palpi, femora and tarsi distinctly lighter than body; setation relatively sparse, moderately long, suberect, light brown. Male. Body length 2.77-2.82 mm (mean 2.8 mm). Head broader than long, widest at large, very convex and finely faceted eyes, length 0.45-0.47 mm (mean 0.46 mm), width 0.61-0.62 mm (mean 0.615 mm). Tempora about as long as eye length in dorsal view, rounded, strongly convergent posteriorly; vertex over twice as broad as long, regularly convex, with small and indistinctly delimited median tubercle adjacent to occipital constriction; supraantennal tubercles prominent, distinctly demarcated from frons but indistinctly delimited from vertex; frons relatively small, subtrape- zoidal, gradually lowering toward deep clypeal suture; clypeus very large, subtrape- zoidal, convex. Vertex posterior to supraantennal tubercles with small, round granules separated by distances about equal to granule diameters; frons with sparse and fine punctures, some with slightly raised margins; clypeus with coarse, small and very dense granules of irregular shape. Setation relatively sparse, setae thin, moderately long, suberect to erect. Antenna (Fig. 36) very slender, length 2.56-2.57 mm (mean 2.565 mm), minimally longer than 0.9 of body length, not thickened toward apex. Pronotum elongate, widest near anterior fourth, length 0.8-0.81 mm (mean 0.815 mm), maximum width 0.69-0.7 mm (mean 0.695 mm), width at base 0.47- 0.5 mm (mean 0.48 mm). Anterior and lateral margins rounded together, sides between middle and posterior third constricted, posterior margin nearly straight, mini- mally expanded posteriorly in middle; posterior collar demarcated from disc by trans- verse row of four large but very shallow, elongate pits. Discal part densely covered with small, round, very convex granules about twice as large as those on vertex, dis- tances between granules shorter than their diameters, posterior collar with very coarse and dense granules irregular in shape. Setation moderately sparse, anterior fourth of disc or less with setae as thin as those on head, posteriorly setae increasing in thickness, all setae moderately long, suberect. Elytra oval, widest at middle or just anterior to middle, length 1.52-1.54 mm (mean 1.53 mm), width 1.0-1.02 mm (mean 1.01 mm), EI 1.51-1.52. Humeral callus on each elytron moderately distinct; internal humeral impression broad and shallow; very narrow adsutural area in anterior !/,-!/3 slightly raised and demarcated by sharp edge; apices of elytra separately rounded. Punctation sparse and fine, punctures shallow and relatively indistinct; setation sparse, moderately long, suberect, setae slightly thinner than those on posterior part of pronotum. Hind wings well developed. Legs long and slender, all femora with stalk-like basal half and clavate distal half, dorsal femoral groove developed nearly from base of femur up to apex, circular pore very small, located in the highest point of dorsal margin of clavate part; tibiae slender, protibiae minimally recurved, meso- and metatibiae nearly straight; tarsi REVISION OF LOEBLITES 361 \ J paraproct 17 à Î > SF Lenta coxite Fics 14-18 Loeblites sabahensis Franz: 14 — tergite IX of male, dorsal view, 15 — sternite IX of male, ventral view, 16 — tergite X, dorsal view, 17 — ovipositor, dorsal view, 18 — proctiger, ventral view. Scale 0.1 mm. slender, tarsomeres I-IV only slightly reducing in length, tarsomere V as long as III-IV together or minimally longer. Aedeagus (Fig. 20-22) elongate, subtriangular, widest near base, narrowing toward rounded apex, length 0.66 mm. Base with deep median emargination or notch; parameres long, slender, exceeding apex of median lobe, broadened near apex, without visible setae; internal armature symmetrical, complicated, its most noticeable and darkly sclerotized part is basal complex composed of median capsule surrounded by two pairs of elongate structures, other parts of internal sac are relatively lightly sclerotized. 362 P. JALOSZYNSKI Female externally indistinguishable from male. Body length 2.74-2.82 mm, length of head 0.45-0.47 mm, width of head 0.62 mm, length of antenna 2.55-2.57 mm, length of pronotum 0.79-0.8 mm, maximum width of pronotum 0.67 mm, width of pronotum at base 0.47-0.5 mm, length of elytra 1.5-1.55 mm, width of elytra 1.0-1.02 mm, EI 1.5-1.52. Spermatheca as in Fig. 23, length 0.075 mm. Bursa copulatrix (Fig. 24) rela- tively large, length 0.24 mm, darkly sclerotized, symmetrical, in dorso-ventral view elongate and broadly constricted in distal third. Comments. Loeblites mastigicornis is unique in having pronotum with small, round, very convex granules densely covering disc. Loeblites sabahensis Franz Figs 1-19, 30-35, 37 Loeblites sabahensis Franz, 1992: 889, fig. 33. Type material. Holotype male, white printed label “SABAH: Poring Hot Springs, 500 m, 11. V. 1987, Burckhardt-L6bl”, white handwritten label “Loeblites sabahensis m.” and printed “det. H. Franz”, red handwritten label “Holotypus” (MHNG). Paratypes: 2 males, 1 fe- male, same data except for yellow handwritten identification label with printed “PARATYPUS” (2 in MHNG, 1 in NMW); 2 females, white printed label “SABAH: Kibongol V., 7 km N Tambunan, 700 m, 20. V. 1987, Burckhardt-Löbl” and identification label as above (MHNG); 1 female, white printed label “SABAH: Crocker Ra., 1270 m, km 60 rte Kota Kinabalu- Tambunan, 17. V. 87, Burckhardt-Löbl” and standard yellow identification label (MHNG); 1 female, white printed label “BORNEO Sabah Mt Kinabalu N.P. Por. H.S., area Kipungit Crk. 2, 490 m, 14. VIII. 1988, A Smetana (B112)”, and standard yellow identification label (NMW). Additional material studied. 5 males, 6 females, Borneo, Sabah, Batu Punggul resort, 24.-26. vi. 1988, Kodada & Ciampur leg. (PCPH, PCPJ); 1 male, Borneo, Sabah, Crocker Range, Gunung Emas, 1600 m, 31. v. 2001, Kodada & Ciampur leg. (PCPH). Redescription. Body moderately large, slender, very convex, moderately dark brown, antenna slightly lighter, palpi and legs distinctly lighter than body; overall setation moderately sparse and long, suberect, light brown. Male (Fig. 19). Body length 2.42-2.59 mm (mean 2.49 mm). Head (Figs. 1-4) very similar to that of L. mastigicornis, length 0.42 mm, width 0.55-0.57 mm (mean 0.56 mm), covered with sparse, small punctures with well delimited margins and sparse, moderately long, very thin setae. Antenna slightly less than 0.9 of body length, length 2.1-2.25 mm (mean 2.19 mm), as in Fig. 37. Pronotum (Figs 9-10) elongate, widest near anterior fourth, length 0.7-0.75 mm (mean 0.72 mm), maximum width 0.57-0.62 mm (mean 0.60 mm), width at base 0.41- 0.45 mm (mean 0.43 mm). Lateral and anterior margins rounded and separated by slightly indistinct angle; posterior margin arcuate, minimally expanded posteriorly in middle; sides strongly constricted between middle and posterior half; posterior collar demarcated from disc by transverse row of four large, deep, slightly elongate pits. Punctation sparse, composed of fine punctures with sharply delimited and slightly raised margins, sides of posterior collar with moderately coarse granulation; setation as that in L. mastigicornis. Elytra as in L. mastigicornis, length 1.3-1.42 mm (mean 1.35 mm), width 0.8- 0.95 mm (mean 0.87 mm), EI 1.49-1.62, punctation fine and sparse; setation moderately long, suberect, setae minimally increasing in thickness from base to middle of each elytron. Hind wings well developed. REVISION OF LOEBLITES 363 NRE GESTO io oa ee Fic. 19 Loeblites sabahensis Franz, dorsal habitus of male. Scale 0.5 mm. recurved. Legs as in L. mastigicornis, except for protibiae, which are nearly straight, not Aedeagus (Figs 30-33) elongate, widest near base, narrowing toward rounded apex, length 0.6 mm. Base with moderately deep median emargination; parameres 364 P. JALOSZYNSKI Fics 20-29 Loeblites mastigicornis Franz (20-24) and L. minor sp. n. (25-29): 20, 25 — aedeagus, dorsal view, 21, 26 — aedeagus, ventral view, 22, 27 — aedeagus, lateral view, 23, 28 — spermatheca, 24, 29 — bursa copulatrix, ventral view. Scale 0.2 mm for 20-22 and 25-27, 0.05 mm for 23-24, 28, 29. long, slender, exceeding apex of median lobe, broadened near apex, without visible setae; internal armature in most type specimens distinctly asymmetrical, in non-type males inner sac variable, from nearly symmetrical as in Fig. 33 to strongly asym- metrical. The most noticeable part of inner armature is median complex composed of dark central arcuate structure (capsule?) surrounded by relatively lightly pigmented lateral lobes bearing numerous tiny denticles in basal part. REVISION OF LOEBLITES 365 Female very similar to male but slightly longer. Body length 2.59-2.68 mm (mean 2.64 mm), length of head 0.45-0.46 mm (mean 0.455 mm), width of head 0.56- 0.59 mm (mean 0.57 mm), length of antenna 2.12-2.25 mm (mean 2.19 mm), length of pronotum 0.72-0.75 mm (mean 0.74 mm), maximum width of pronotum 0.57-0.65 mm (mean 0.61 mm), width of pronotum at base 0.42-0.45 mm (mean 0.43 mm), length of elytra 1.42-1.47 mm (mean 1.45 mm), width of elytra 0.91-0.97 mm (mean 0.94 mm), EI 1.51-1.56. Spermatheca as in Fig. 34, length 0.08 mm. Bursa copulatrix (Fig. 35) modera- tely darkly sclerotized, asymmetrical, subconical and slightly curved, length 0.12 mm. Comments. This species can be identified on the basis of lack of granulation on pronotum and moderately large body. Comparison of the elytral index, the shape of aedeagus or spermatheca and especially the bursa copulatrix is necessary to distinguish L. sabahensis from S. minor. The latter taxon has not been recognized by Franz and was included into the type series of L. sabahensis. Loeblites minor sp. n. Figs 25-29, 38 Type material. Holotype male, white printed label “Malaysia, Sabah, Batu Punggul Resort env., 24. VI.-1. VII. 1996, 11c, vegetation debris and forest floor litter accumulated around large trees near river”, red printed label “LOEBLITES minor m., det. P. Jatoszyfiski, 2004” (MHNG). Paratypes (all misidentified and bearing yellow identification labels with hand- written “Loeblites sabahensis m.” and printed “PARATYPUS”, labeled by H. Franz): 1 male, 1 female, white printed label “SABAH: Poring Hot Springs, 500 m, 7. V. 1987, Burckhardt-Löbl” (MHNG); 1 male, 2 females, white printed label “SABAH, # 15a, Poring Hot Springs, 500 m, 7. V. 1987, Burckhardt-Löbl” (2 in NMW, 1 in MHNG); all paratypes have been labeled during the present study with yellow, printed labels “LOEBLITES minor m., det. P. Jatoszynski, 2004”. Description. Body small, slender, very convex, moderately dark brown, an- tennae slightly lighter, palpi and legs distinctly lighter than body; setation moderately long and sparse, suberect. Male externally nearly identical to L. sabahensis, including setation and punc- tation, it differs only in slightly smaller body, shorter antennae and proportions of ely- tra. Body length 2.31-2.45 mm (mean 2.38 mm), length of head 0.39-0.41 mm (mean 0.4 mm), width of head 0.55-0.57 mm (mean 0.56 mm), antenna only slightly longer than 0.8 of body length, as in Fig. 38, length 1.95-2.05 mm (mean 1.99 mm); length of pronotum 0.67-0.72 mm (mean 0.69 mm), maximum width of pronotum 0.55-0.57 mm (mean 0.56 mm), width of pronotum at base 0.4-0.42 mm (mean 0.41 mm), length of elytra 1.25-1.32 mm (mean 1.29 mm), width of elytra 0.85-0.95 mm (mean 0.88 mm), EI 1.39-1 47. Aedeagus (Fig. 25-27) elongate, widest near base, narrowing toward rounded apex, length 0.49 mm. Base with moderately deep median emargination; parameres long, slender, exceeding apex of median lobe, broadened near apex, without setae; internal armature symmetrical, relatively darkly sclerotized, central complex composed of median capsule surrounded by subtriangular lateral structures and pair of elongate, nearly transverse sclerites in distal part of inner sac. Female externally differs from male only in higher elytral index. Body length 2.33-2.55 mm (mean 2.41 mm), length of head 0.41-0.45 mm (mean 0.43 mm), width of head 0.56-0.57 mm (mean 0.565 mm), length of antenna 1.9-2.0 mm (mean 366 P. JALOSZYNSKI 36 37 38 Fics 30-38 Loeblites sabahensis Franz (30-35, 37), L. mastigicornis Franz (36) and L. minor sp. n. (38): 30 — aedeagus of paratype, dorsal view, 31 — aedeagus, ventral view, 32 — aedeagus, lateral view, 33 — structures of inner sac, non-type male (variant), 34 — spermatheca, 35 — bursa copulatrix, ven- tral view, 36-38 — left antenna, dorsal view. Scale 0.2 mm for 30-32, 0.05 mm for 33-35, 0.5 mm for 36-38. 1.94 mm), length of pronotum 0.67-0.71 mm (mean 0.68 mm), maximum width of pronotum 0.55-0.56 mm (mean 0.55 mm), width of pronotum at base 0.41-0.42 mm (mean 0.41 mm), length of elytra 1.25-1.39 mm (mean 1.30 mm), width of elytra 0.87- 0.92 mm (mean 0.89 mm), EI 1.44-1.51. Spermatheca as in Fig. 28, length 0.062 mm. Bursa copulatrix (Fig. 29) missing or highly reduced to transparent, hymenous structures surrounding end of ductus sper- mathecae. Comments. This species is the smallest Loeblites, externally extremely similar to L. sabahensis. Examination of the aedeagus or spermatheca and bursa copulatrix is necessary for correct identification. REVISION OF LOEBLITES 367 Etymology. The specific Latin epithet “minor” refers to the smallest body amongst all species of the genus. DISCUSSION Within the Cyrtoscydmini, Loeblites shows the highest degree of similarity to Syndicus and Horaeomorphus. During the present study, the dorsal femoral groove was found in all members of this genus. This interesting character has been discovered very recently in Syndicus (Jatoszyñski, 2004); all Asiatic species of the Cyrtoscydmini known to the author (including all described species of Horaeomorphus, Borneosabahia Franz, Parastenichnus Franz, Protoscydmus Franz, Parastenichnaphes Franz, Siamites Franz, and many species of Euconnus Thomson, Microscydmus Saulcy & Croissandeau, Neuraphes Stevens, Scydmoraphes Reitter, and Stenichnus Thomson) have dorsal surface of femora without this structure. The general body shape of species belonging to Syndicus s. str. and Loeblites is very similar; on the other hand, the body shape (and some other characters) of Syndicus (Semisyndicus) is more similar to that of Horaeomorphus. The very long and thin antenna with well separated antennomere XI in Loeblites can be used as an unambiguous key character to distinguish this genus from Syndicus, which has the antennomere XI tightly inserted into the apex of the an- tennomere X, without separating interspace (a unique character within the family). Also, the single subapical tooth instead of two teeth and the sharp, finely and irregu- larly serrated cutting edge of mandibles found in Loeblites is not known among repre- sentatives of Syndicus and Horaeomorphus (however, details of mouthparts of Horaeomorphus are known only in H. sakishimanus Jatoszynski). An undescribed species of Horaeomorphus from Malaysia known to the author has the prostheca signi- ficantly shorter than that of H. sakishimanus, similar to the prostheca of L. sabahensis. Another unique feature of Loeblites is the long and slender median projection located in the posterior margin of the proctiger; this part of the female genital segment has broadly rounded posterior margin both in Syndicus and Horaeomorphus. The bursa copulatrix uniquely shaped in two out of three known species of Loeblites seems to be extremely diversified within the genus, and it has not been found in L. minor. This character is also variable within Horaeomorphus (Jatoszyñski, in preparation). Wing venation in Loeblites, thought similar to that found in the both subgenera of Syndicus, clearly differs in longer subcosta fused distally with radius (very short in Syndicus); wings in the remaining genera of the tribe have not been illustrated or described. Besides the general appearance and the presence of femoral grooves, members of Loeblites and Syndicus share the following features: the clypeus separated from the frons by a transverse groove; eyes with emarginated posterior or postero-ventral margin; antennal insertions located on latero-anterior margin of the head, widely separated; anterior tentorial foveae (overlooked in Jatoszyfiski, 2004, examined and confirmed during the present study in S. (s. str.) difficilis Jatoszyfiski and S. (Semi- syndicus) laei ranongianus Jatoszyñski) hidden in supraclypeal groove, hardly visible in dry-mounted specimens; occiput broad; mandible nearly planar, subtriangular; very similar maxilla (however, this structure seems to be little diversified within the tribe, Jatoszynski, unpublished observations); nearly identical design of palpi maxillare and 368 P. JALOSZYNSKI labiale; pronotum widest near anterior third, with narrow posterior collar separated by a row of ante-basal foveae; very similar details of mesosternum, metasternum, metan- otum and mesonotum, the latter structure with characteristic long posterior notal processes; each elytron with two basal foveae connected by an U-shaped groove and in natural position partly hidden under the posterior margin of the pronotum; similar wing venation; shape of legs; male and female genital segments; the aedeagus; and the spermatheca. Loeblites is more similar to Syndicus s. str. in the body shape (in Semi- syndicus the body shape is stouter and resembles that of Horaeomorphus), uniformly concave basisternal area of the mesosternum (in Semisyndicus with unique median “isle”, as illustrated in Jatoszynski, 2004, fig. 35D) and nearly straight base of elytra (strongly concave in Semisyndicus). In turn, Loeblites is more similar to Semisyndicus than to Syndicus s. str. in the shape of the posterior margin of the vertex, which bears a small median tubercle (not found in Syndicus s. str.); the aedeagus, which is espe- cially similar between all Loeblites and S. (Sem.) tenuicornis Jatoszynski (Jatoszynski, 2004, fig. 42A-C); in turn the copulatory organ of the latter species resembles that of Horaeomorphus. Loeblites also shares with Semisyndicus the shape of the sperma- theca, which has approximate insertions of the ductus spermathecae and the accessory gland, which are widely separated in Syndicus s. str. Interestingly, in representatives of Horaeomorphus, which have aedeagi similar to those of Loeblites, the spermatheca is distinctly different; it is also not similar to the spermatheca of Syndicus s. str. However, in one species of Horaeomorphus (H. nepalensis Franz) the posterior margin of the vertex bears a very small median tubercle like that found in Loeblites and Semi- syndicus, and the structures surrounding the opening of the ductus spermathecae inside the female genital segment are darkly sclerotized and relatively complicated (Jatoszynski, unpublished observations). Actual relationships between the three genera will be analyzed when important morphological details of Horaeomorphus have been described. The existing data strongly suggest a separate position of Loeblites, Syndicus and Horaeomorphus within the Cyrtoscydmini, a monophyly of this small group seems plausible. ACKNOWLEDGEMENTS I express my thanks to persons who arranged loans of the material used in my study: Dr Giulio Cuccodoro (MHNG), Dr Harald Schillhammer (NMW) and Mr Peter Hlavaè (KoSice, Slovakia). A significant part of this study was conducted during my stay in Japan; I’m greatly indebted to Dr Shühei Nomura (National Science Museum, Tokyo), who helped me in various ways during my work on the Oriental Scyd- maenidae. REFERENCES BORDONI, A. & CASTELLINI, G. 1973. Sulle specie Paleartiche del genere Mastigus Latreille con osservazioni su due specie dell’ Africa Australe (Coleoptera Scydmaenidae). Redia 54: 295-323. FRANZ, H. 1986. Ein neues Genus und ein neues Subgenus der Scydmaenidae aus Siidostasien. Revue suisse de Zoologie 93(4): 965-970. REVISION OF LOEBLITES 369 FRANZ, H. 1992. Monographie der Scydmaenidae (Coleoptera) von Sabah (NO-Borneo). Revue suisse de Zoologie 99(4): 859-953. JALOSZYNSKI, P. 2004. Revision of scydmaenid beetles of the genus Syndicus Motschulsky (Coleoptera, Scydmaenidae). National Science Museum Monographs 25, National Science Museum, Tokyo, 108 pp. NEWTON, A. F. & FRANZ, H. 1998. World catalog of the genera of Scydmaenidae (Coleoptera). Koleopterologische Rundschau 68: 137-165. NOMURA, S. 1991. Systematic study on the genus Batrisoplisus and its allied genera from Japan (Coleoptera, Pselaphidae). Esakia, Fukuoka 30: 1-462. O’KEEFE, S. T. & MONTEITH, G. 2000. Clidicus abbottensis O’ Keefe, a new species of Scyd- maenidae (Coleoptera: Staphylinoidea) from Australia with description of the larva. Memoirs of the Queensland Museum 46(1): 211-223. Alan. = Le i au de perio sd 2 de Ce REVUE SUISSE DE ZOOLOGIE 112 (2): 371-399; juin 2005 On Hadogenes angolensis Lourenco, 1999 syn. n. (Scorpiones, Liochelidae), with a redescription of H. taeniurus (Thorell, 1876) Lorenzo PRENDINI Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA. E-mail: lorenzo@amnh.org On Hadogenes angolensis Lourenco, 1999 syn. n. (Scorpiones, Liochelidae), with a redescription of H. taeniurus (Thorell, 1876). - The flat rock scorpion, Hadogenes taeniurus (Thorell, 1876), is redescribed, based on an examination of approximately 250 specimens from Angola and Namibia. Hadogenes angolensis Lourengo, 1999 is demonstrated to be a junior synonym. The distribution of H. taeniurus is mapped, and notes on its ecology and conservation status are provided. The synonymy of Hadogenes bifossulatus Roewer, 1943 with Hadogenes tityrus (Simon, 1888), rather than with H. taeniurus, is confirmed. Keywords: Scorpiones - Liochelidae - Hadogenes taeniurus - Hadogenes tityrus - synonymy - Angola - Namibia. INTRODUCTION Flat rock scorpions of the genus Hadogenes Kraepelin, 1894 represent an intriguing group of mostly large, extremely dorsoventrally compressed scorpions, all described species of which are obligate lithophiles, inhabiting the narrow cracks, crevices and spaces beneath exfoliations of weathered rock outcrops from South Africa to Tanzania. Besides dorsoventral compression, other ecomorphological adaptations facilitating existence in this specialized habitat include elongation of the metasoma and pedipalps, perhaps to aid with prey capture in confined spaces; greatly enlarged lateral ocelli relative to the median ocelli, to aid in anterior light perception; pronounced superciliary carinae to protect the median ocelli from abrasion; stout, spiniform setae on the ventral surfaces of the telotarsi and highly curved telotarsal ungues, to provide a vice-like grip on rock surfaces (Newlands, 1972a, 1972b, 1978; Newlands & Prendini, 1997; Prendini, 2001a). The tarsal adaptations of Hadogenes facilitate loco- motion on rock but hinder locomotion across other substrata. These scorpions are thus restricted to regions of rugged, mountainous topography and subject to allopatric speciation when mountain ranges become separated through erosion (Newlands, 1972a; Prendini, 2001b). With few exceptions, the distributional ranges of Hadogenes species are allopatric or parapatric as a result (Newlands, 1980; Prendini, 1995, 2001a). Manuscript accepted 16.11.2004 372 L. PRENDINI The taxonomy and phylogenetic position of Hadogenes have been the focus of several investigations in recent years (e.g., Newlands & Prendini, 1997; Lourenço, 1999; Prendini, 2000a, 2001a). The present contribution forms part of an ongoing revision of the taxonomy of the genus, and addresses the status of Hadogenes taeniu- rus (Thorell, 1876), a flat rock scorpion endemic to southern Angola and northern Namibia (Fig. 1). Hadogenes taeniurus was first described under the name /schnurus hahni Peters, 1862. However, this name was apparently forgotten after its description and not listed in any subsequent revisions or catalogs until its rediscovery by V. Fet. At the time of its rediscovery, the syntypes of /. hahni were examined (Prendini in Fet, 2000), determined to be conspecific with H. taeniurus, and synonymized accordingly. Although Peters’ (1862) name is the senior synonym, it has not been used since its description, whereas H. taeniurus has been widely accepted (Fet, 2000). After its description, H. taeniurus was synonymized, first with Hadogenes trichiurus (Gervais, 1843) by Kraepelin (1894) and then with Hadogenes troglodytes (Peters, 1861) by Kraepelin (1899), but its validity was upheld by all subsequent authors, including Kraepelin (1908, 1914). Hadogenes taeniurus remained poorly known — aside from a few citations, mostly providing new locality records or dis- cussing its diagnostic characters (Purcell, 1901; Kraepelin, 1908, 1914; Hewitt, 1918; Lawrence, 1928, 1955, 1959, 1961; Newlands, 1972a; Lamoral & Reynders, 1975) — until its redescription by Lamoral (1979). Lamoral (1979) examined most of the spe- cimens of H. taeniurus that were available at the time, described the trichobothria and hemispermatophore of the species for the first time, and restricted its known distri- bution to southern Angola and northern Namibia. Meanwhile, Newlands (1980), in an unpublished thesis, independently redescribed the species, mapped its known distri- bution in Angola and Namibia, and refined its diagnosis in a key to the species of Hadogenes. Newlands’ (1980) redescription and distribution map of H. taeniurus were never published. However, Newlands & Cantrell (1985) published Newlands’ (1980) key to the species of Hadogenes, as well as the chromosome number of H. taeniurus. Several authors previously noted the occurrence of H. taeniurus in Angola (Lawrence, 1959; Newlands, 1972a, 1980; Lamoral & Reynders, 1975; Lamoral, 1979). Three of these authors cited the Angolan specimens they examined (Lawrence, 1959; Newlands, 1972a, 1980; Lamoral, 1979). Indeed, the occurrence of H. taeniurus in that country was not questioned until a recent paper, in which a new species, Hadogenes angolensis Lourenço, 1999, was described. Lourenço (1999) justified this putative new species largely on a mistaken impression that the Angolan records of H. taeniurus, attributed solely to Lawrence (1959), were based on misidentifications (im- plying that the Angolan specimens represented another species, for which the name H. angolensis was proposed). Lourenco (1999) neglected to mention that the type locality of H. taeniurus is unknown (and could, in fact, be in Angola) and failed to provide any consistent diagnostic differences by means of which this putative new species might be distinguished from H. taeniurus. During the present study, the type specimens of H. an- golensis were compared and found to be conspecific with the type specimens of H. tae- niurus and I. hahni, and also with 247 non-type specimens of H. taeniurus from Angola and Namibia, including all the material examined by Lawrence (1959), Newlands ON HADOGENES ANGOLENSIS 373 NAMIBIA FIG. 1 Map showing the known distribution of Hadogenes taeniurus (Thorell, 1876) in Angola and Namibia (™), including the type localities for its synonyms: Zschnurus hahni Peters, 1862 (+) and H. angolensis Lourenço, 1999 (Je). (1972a, 1980) and Lamoral (1979). On the basis of this evidence, H. angolensis is here synonymized with H. taeniurus. The status of Hadogenes bifossulatus Roewer, 1943, long regarded as a dubious species (Lawrence, 1955; Newlands, 1972a, 1980; Lamoral & Reynders, 1975; Lamoral, 1979; Fet, 2000) was also investigated. Newlands (1972a) suggested that H. 374 L. PRENDINI bifossulatus might be synonymous with H. taeniurus. However, Newlands (1980) retracted his earlier suggestion, after examining the holotype of H. bifossulatus, and instead synonymized H. bifossulatus with Hadogenes tityrus (Simon, 1888). Newlands’ (1980) unpublished synonymy of H. bifossulatus was later published (and thereby validated) by Kovarik (1998), without explicit mention of the source, but the synonymy was not accepted by Fet (2000). During the present investigation, the holo- type of H. bifossulatus was re-examined and confirmed to be conspecific with H. tityrus, rather than with H. taeniurus. In light of this new evidence, the diagnosis and description of H. taeniurus are revised, its known distribution mapped on the basis of all available material, including a large number of new specimens (many from Angola) that have accumulated since the work of Lamoral (1979), and notes on its ecology and conservation status are provided. MATERIAL AND METHODS Many personally collected specimens of H. taeniurus were located at night using the ultraviolet (UV) light detection method (Honetschlager, 1965; Stahnke, 1972; Sissom et al., 1990). A portable UV lamp, comprising two mercury-vapour tubes at- tached to a chromium parabolic reflector and powered by a rechargeable 7 Amp/hr, 12 V battery, was used for this purpose. A few specimens were also found by inspecting rock crevices and exfoliations during the day. A portable Garmin™ GPS II Plus device was used for recording the geographical coordinates of collection localities in the field. All material examined, including type specimens, is deposited in the following collections: Albany Museum, Grahamstown, South Africa (AMGS); American Museum of Natural History, New York (AMNH), some bearing accession numbers from the Alexis Harington Collection (AH); California Academy of Sciences, San Francisco (CASC); Muséum d’histoire naturelle, Genève, Switzerland (MHNG); Musée Royal de |’ Afrique Centrale, Tervuren, Belgium (MRAC); National Collection of Arachnida, Plant Protection Research Institute, Pretoria, South Africa (NCA); Göteborgs Naturhistoriska Museet, Göteborg, Sweden (NMG); Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRM), bearing an accession number from the Julio Ferrer Collection (JF); National Museum of Namibia, Windhoek (NMNW); Natal Museum, Pietermaritzburg, South Africa (NMSA); South African Museum, Cape Town (SAMC), some bearing accession numbers from the John Visser Collection (JV); Natur-Museum Senckenberg, Frankfurt, Germany (SMF); Transvaal Museum, Pretoria, South Africa (TMSA); Zoologisches Museum, Humboldt-Universitat, Berlin, Germany (ZMHB); Zoologiska Institutionen, Lunds Universitet, Sweden (ZMLU); Zoologisches Institut und Zoologisches Museum, Universität Hamburg, Germany (ZMUH). Tissue samples of H. taeniurus have been stored (in the vapour phase of liquid nitrogen at -150°C) in the Ambrose Monell Collection for Molecular and Microbial Research (AMCC) at the AMNH. Photographs were taken in visible light as well as under long wave UV light using a Microptics™ ML-1000 digital photomicrography system. Measurements were taken using the ocular micrometer of a Nikon® SMZ-1500 stereomicroscope. Colour designation follows Smithe (1974, 1975, 1981), trichobothrial notation follows Vachon ON HADOGENES ANGOLENSIS 57/5) (1974), sternum terminology follows Soleglad & Fet (2003). Morphological termino- logy and mensuration follows previous papers (e.g., Newlands & Prendini, 1997; Prendini, 2000a, 2001a). A distribution map was produced using ArcView GIS Version 3.2 (Environ- mental Systems Research Institute, Redlands, California), by superimposing point locality records on coverages depicting the political boundaries, topography (500 m contour interval) and major sand systems in Angola and Namibia. A topographic contour coverage was created from the GTOPO30 raster grid coverage, obtained from the website of the U.S. Government Public Information Exchange Resource: http://edcdaac.usgs.gov/gtopo30/gtopo30.html. The coverage of sand systems was created by clipping and merging relevant polygons extracted from a coverage of the geology of Africa provided by the Department of Marine Geoscience, University of Cape Town, South Africa, with polygons extracted from a coverage of Namibian land- forms from the Namibian National Biodiversity Task Force (Barnard, 1998), downloaded from their website: http://www.dea.met.gov.na/programmes/biodiversity/ countrystudy.htm. In order to create a point locality geographical dataset for mapping the distrib- utional range of H. taeniurus, all records of sufficient accuracy were isolated from the material examined. Only a small proportion of the records were accompanied by geo- graphical coordinates or quarter-degree squares (QDS), usually entered by the collec- tor or subsequently added by the curator or collections manager. These were checked for accuracy and an attempt was made to trace coordinates for as many of the remain- ing records as possible, by reference to gazetteers, the official 1:250 000, 1:500 000 and 1:1000 000 topo-cadastral maps of Namibia published by the Government Printer, and the GEOnet Names Server (GNS): http://164.214.2.59/gns/html/cntry_files.html. Names of provinces, regions and magisterial districts of countries listed in the materi- al examined follow the most recent systems (post-1994). Spatial analyses were conducted, using ArcView, in order to ascertain whether the distribution of H. taeniurus is related to present environmental variables, to deter- mine the specific ecological correlates of its distributional range, and to calculate sta- tistics that could be used to define its conservation status. Coverages representing the topography, mean annual rainfall, biomes (as defined by Irish, 1994), vegetation types and protected areas in Namibia were used for these analyses. The GTOPO30 raster grid coverage was used for spatial analysis of topography. Polygon coverages representing the mean annual rainfall and vegetation types (as defined by Giess, 1971) of Namibia, were obtained from the website of the Namibian National Biodiversity Task Force (Barnard, 1998). The conservation status of H. taeniurus was assessed using a coverage of the protected area network in Namibia, also obtained from the Namibian National Biodiversity Task Force (Barnard, 1998). This was superimposed on the mapped dis- tributional range of H. taeniurus to determine whether any point locality records fall within the boundaries of protected areas. Number of known locality records, extent of the distributional range, occurrence inside and outside of protected areas, and pre- vailing land uses that might be construed as threats to the future survival of the species were then used as criteria for assigning H. taeniurus to one of the IUCN Red List Categories (IUCN, 2001). 376 L. PRENDINI SYSTEMATICS Hadogenes taeniurus (Thorell, 1876) Ischnurus hahni Peters, 1862: 27 (synonymized by Prendini in Fet, 2000: 389); Moritz & Fischer, 1980: 315. Ischnurus taeniurus Thorell, 1876: 254-258. Hadogenes trichiurus: Kraepelin, 1894: 115 (part). Hadogenes troglodytes: Kraepelin, 1899: 145 (part). Hadogenes taeniurus: Purcell, 1901: 206; Kraepelin, 1908: 267, 268 (part); Kraepelin, 1914: 117 (part); Hewitt, 1918: 163; Lawrence, 1928: 277, 278, tab.; Lawrence, 1955: 223, 252; Lawrence, 1959: 386; Lawrence, 1961: 154; Newlands, 1972a: 135 (part); Lamoral & Reynders, 1975: 540; Lamoral, 1979: 654-656, 657 (part), fig. 304, 305, 307-309; Newlands & Cantrell, 1985: 42, 44, tab. 2; Sissom, 1990: 125, fig. 3.24H; Kovarik, 1998: 133; Lourengo, 1999: 932, 934, 935, fig. 18, 21, 22; Fet, 2000: 389; Prendini, 2001a: 148, tab. 1; Prendini, 2001b: 137. Hadogenes sp. ?: Lawrence, 1927: 73. Hadogenes angolensis Lourenço, 1999: 932-936, fig. 1-17, 19, 20, 23, 25, tab. I (syn. n.); Kovarik, 2001: 83; Prendini, 2001a: 148, tab. 1; Prendini, 2001b: 136. Hadogenes taeniurus (?): Lourenço, 1999: 937, fig. 25. Type material (examined). Ischnurus taeniurus: holotype © [two legs broken, opistho- soma damaged] (NMG 121 [type no. 90]), ‘S. Africa’ [probably northern NAMIBIA (Purcell, 1901; Hewitt, 1918)], 28.x1.1864, C.J. Andersson. /schnurus hahni: 1 8,1 9 syntypes (ZMHB 2317/18), Neu-Barmen [Gross Barmen, 22°06°S 16°45’E], Otjimbingue [21°21’S 16°08’E], South West Africa [NAMIBIA: Otjozondjupa Region: Okahandja District], Hahn. Hadogenes angolensis: holotype dé [hemispermatophore dissected], paratype 2, 1 juv. d paratype (MHNG), ANGOLA: Huila District: Fazenda Bumbo, near Capangombe, 15°10’S 13°09’E, 21-26.vi.1954, W. Kisker & H. Barmann. Diagnosis. Hadogenes taeniurus appears most closely related to H. granulatus Purcell, 1901 and H. troglodytes, based on their large size (113-160 mm in d , 108-154 mm in 2) and similarity in habitus. All three species share the presence of postero- lateral oval depressions in sternite VII (Figs 10, 11), although these are most pro- nounced in H. taeniurus (Lawrence, 1928; Newlands, 1972a, 1980; Lamoral, 1979). They also share the presence of granulation on the telson of the adult male, which is more pronounced in H. granulatus (Purcell, 1901; Newlands, 1980). Hadogenes troglodytes differs from H. granulatus and H. taeniurus in the shape of metasomal segment I, which is higher than wide posteriorly, rather than wider than high posteriorly (Newlands, 1972a, 1980; Lamoral, 1979; Newlands & Cantrell, 1985). Hadogenes granulatus differs from H. taeniurus and H. troglodytes in two secondary sexual characters of the adult male. The lobe on the movable finger and corresponding notch in the fixed finger of the pedipalp chela are weakly developed to absent in H. granulatus, whereas the lobe and notch are more pronounced in H. taeniurus (Fig. 12) and H. troglodytes. The telson of H. granulatus is elongated and coarsely granular (Purcell, 1901; Newlands, 1980), compared with that of H. taeniurus and H. troglodytes, which is oval and finely granular (Fig. 21). Hadogenes taeniurus is readily separated from H. granulatus and H. troglodytes by the presence of a deep oval depression in the basal piece, between the pectines (Figs 8,9), which is absent in the other species. This character has not been noted previously. These species are further separated by the relatively shorter metasoma of the adult male H. taeniurus (usually 30-40% greater than the combined length of prosoma and meso- soma, compared with 50% and 45% greater in H. granulatus and H. troglodytes, respectively). ON HADOGENES ANGOLENSIS 377 Fics 2-5 Hadogenes taeniurus (Thorell, 1876), habitus of d and 2 (Farm Uisib 427 [AMNH]). 2. Dorsal aspect, d . 3. Ventral aspect, d. 4. Dorsal aspect, 2.5. Ventral aspect, 2. Scale bars = 10 mm. Description. The following description is based on the holotype © of I. taeniu- rus, the d and 9 syntypes of J. hahni, the holotype ¢ and paratype © of H. angolensis, and another d specimen in the AMNH [AH 4132]. Colour is described from a d and 2 recently collected at Farm Uisib 427 (AMNH). 378 L. PRENDINI Colour. Pale legs and tergites contrasting with darker chelicerae, carapace, pedi- palps, metasoma and telson (Figs 2-5). Posterior third of carapace, tergites, and ster- nites paler than anterior two-thirds of carapace, pedipalps, metasoma and telson. Telson often slightly paler than metasoma but darker than tergites. Chelicerae, pedipalp carinae, chela fingers, and ventral surfaces of leg femora sepia 119; carapace (anterior two-thirds), pedipalp chela manus, and intercarinal surfaces of patella and femur chestnut 32; carapace (posterior third), tergites and sternites olive-yellow 52; pectines and genital operculum straw yellow 36; legs (except ventral surfaces of femora) cinnamon 123A; metasomal segments I-V dusky brown 19; telson raw umber 123. Carapace. Three pairs of lateral ocelli, slightly smaller than median ocelli (Figs 6, 7). Median ocular tubercle with superciliary carinae well developed, protruding above ocelli, and interocular sulcus distinct. Anterior margin of carapace with median notch absent, triangular inset protruding slightly anteriorly. Anteromedian sulcus deep, suturiform, furcating anteriorly around triangular inset. Median longitudinal suture dis- tinct, continuous from anterior furcated sutures through ocular tubercle to posterior fur- cated sutures, which converge on ocular tubercle from posterior carapace margin. Posterior furcated sutures obsolete, discontinuous. Posteromedian and posteromarginal sulci distinct, but shallow. Paired median lateral and posterolateral sulci also distinct, shallow. Carapace almost entirely granular, except for surfaces behind lateral ocelli, and median lateral, posterolateral and posteromarginal sulci, which are smooth. Granulation almost uniformly fine, becoming coarse on antero-ocular and anterolateral surfaces. Chelicerae. Movable finger with distal internal tooth slightly smaller than distal external tooth, and opposable. Ventral aspect of fingers and manus with long, dense macrosetae. Pedipalps. Chela pentacarinate, with three distinct carinae; dorsal secondary and digital carinae obsolete (Figs 12, 13); external secondary carina strongly developed, costate granular (Figs 13, 14); ventroexternal carina strongly developed, crenulate, aligned parallel to longitudinal axis of chela, with distal edge disconnected from external movable finger condyle and directed toward a point between external and internal movable finger condyles, but closer to external condyle (Fig. 15); ventro- median carina obsolete, reduced to a vestigial granule proximally; ventrointernal carina also obsolete; internomedian and dorsointernal carinae weakly developed, each comprising a series of isolated spiniform granules; dorsomedian carina strongly devel- oped, composed of a continous row of spiniform granules; dorsal and ventrointernal in- tercarinal surfaces smooth, reticulate, becoming granulo-reticulate dorsally; dorsoint- ernal intercarinal surfaces with scattered spiniform granules, becoming finely granular on internal surface of fixed finger; external intercarinal surfaces coarsely granular. Chela with a rounded lobe on movable finger and corresponding notch in fixed finger. Dentate margins of chela fingers with double row of denticles, fused at the lobe/notch. Chela width 49.5% (45-54%) greater than chela height; length along ventroexternal carina 46% (43-49%) greater than width in d, and 39.5% (32-47%) greater in 2; length of movable finger 15.5% (10-21%) less than length along ventroexternal carina in gd, and 10.5% (3-18%) less in 2. Patella with seven carinae, six of them distinct (Figs 16-18); dorsoexternal carina obsolete; dorsointernal and ventrointernal carinae costate to costate granular; ON HADOGENES ANGOLENSIS = È Fics 6-11 Hadogenes taeniurus (Thorell, 1876), carapace, pectines and sternites of d and 2 (Farm Uisib 427 [AMNH]). 6. Carapace, 3.7. Carapace, 9 . 8. Pectines and basal piece, d . 9. Pectines and basal piece, ?. 10. Sternite VII, &. 11. Sternite VII, 2. Scale bars = 5 mm. 380 L. PRENDINI internomedian carina costate granular, composed of very large heavily sclerotized spiniform granules; externomedian and ventroexternal carinae granular; dorsoexternal and ventral intercarinal surfaces finely and uniformly granular, becoming granulo- reticulate on ventral surfaces; internal intercarinal surfaces smooth, except for a few scattered granules; anterior process strongly developed. Patella length 38.5% (32-46%) greater than width. Femur pentacarinate, with four distinct carinae; ventroexternal carina obsolete, reduced to a weak row of scattered granules; dorsoexternal and externomedian carinae granular, dorsointernal and ventrointernal carinae costate granular, composed of very large heavily sclerotized granules (Fig. 19); dorsoexternal and ventral intercarinal sur- faces finely and uniformly granular, becoming granulo-reticulate on ventral surfaces; internal intercarinal surfaces smooth, except for a few scattered spiniform granules. Femur length 62.5% (58-66%) greater than width (Table 1). Trichobothria. Neobothriotaxic major, type C (Figs 12-19), with the following segment totals (Table 1): femur 3 (1 d; 1 7; 1 e), patella 86-127 (2 d; 1 i; 28-49 v; 55- 75 e) and chela 90-126 (80-116 manus; 10 fixed finger, including 2 7). Total number of trichobothria per pedipalp, 179-256. Only femoral trichobothria, trichobothria in the d and i series of the patella, and trichobothria in the D, d, e and i series of the chela stable in number and distribution. External and ventral trichobothria of the chela and patella numerically and distributionally too variable for diagnostic purposes. Mesosoma. Tergites each with paired submedian depressions and obsolete me- dian carina. Pre-tergites of d and 2 smooth and shiny. Post-tergites of 4 uniformly finely granular, imparting a matt appearance to all surfaces, except for smooth subme- dian depressions; post-tergites I-VI of 9 smooth and shiny, but often with very fine and even granulation anteromedially, VII uniformly finely granular. Sternites smooth and shiny, each with paired longitudinal depressions internal to spiracles; VII additionally with a pair of deep posterolateral oval depressions (more prominent in d) and a pair of obsolete carinae, converging distally towards a shallow notch in distal apex (Figs 10, 11). Sternite VII 10% (3-17 %) wider than long in d, 15% (7-23%) wider than long in 9 (Table 1). Pectines. Basal piece between pectines with a deep oval depression (Figs 8, 9). Mesial margin of first proximal median lamella of each pecten angular, with pectinal teeth present along entire posterior margin in d; mesial margin of first proximal median lamella shallowly curved, with proximal fifth of posterior margin devoid of teeth in 2. Pectinal teeth: 18-24 (6), 14-20 (2). Sternum. Subpentagonal, type 2. Median longitudinal furrow deep and narrow along enire length. Genital operculum. Suboval, completely divided longitudinally, with genital papillae present (4). Subcordate, partially connected by a membrane in anterior two- thirds, with distinct distal lobes in posterior third, and with genital papillae absent (2). Legs. Femora each with paired granular carinae on ventral surface, becoming less developed on posterior legs. Basitarsi each with a few spiniform setae on prolateral and retrolateral margins, decreasing in number from anterior to posterior legs. Telotarsi each with two rows of three ventrosubmedian spiniform setae and a basal row of ventromedian spinules. Telotarsal laterodistal lobes truncated; median dorsal lobes ON HADOGENES ANGOLENSIS 381 extending to ungues. Telotarsal ungues short, distinctly curved, and equal in length. Retrolateral pedal spurs absent. Metasoma and telson. Metasomal segment I 13.5% (2-25 %) wider than high posteriorly (Table 1). Metasomal segments I-V progressively increasing in length, and decreasing in width, with segment V 36.5% (27-46%) narrower than segment I in à, 40% (32-43%) narrower in 2. Metasoma slender, width percentage of length for seg- ment I, 34% (29-39%) in 3, 45.5% (38-53%) in ®; for II, 15% (12-18%) in 5, 23.5% (22-25%) in 2; for III, 17% (15-19%) in 3, 25% (24-26%) in 9 ; for IV, 12.5% (10- 15 %) in 3, 19.5% (14-22%) in ®; and for V, 12.5% (11-14%) in 3, 17.5% (16-19%) in 2. Telson vesicle 16.5% (3-30%) wider than metasomal segment V in 4, 15% (4-26%) wider in 2; oval in shape, with flattened dorsal surface and rounded ventral surface (Figs 20, 21), height 36.5% (33-40%) of length. Aculeus short, 20% (14-26%) of vesicle length, and sharply curved. Total length of metasoma 45.5% (31-60%) greater than combined length of prosoma and mesosoma in 6, but 5% (0-10%) less than combined length of prosoma and mesosoma in 9. Eight carinae on segment I, six carinae on segments II-IV, and five carinae on segment V. Dorsosubmedian carinae of segment I becoming obsolete posteriorly, but distinct throughout length of segments II-V. Median lateral carinae fully developed on segment I, but absent from segments II-V. Segments I-IV with closely paired ventro- submedian carinae, fused into a single ventromedian carina on segment V. Ventrosubmedian and ventrolateral carinae costate on segment I, costate to costate granular on segments II-IV. Ventrolateral and ventromedian carinae of segment V composed of spiniform granules. Median lateral and dorsosubmedian carinae costate on segment I, dorsosubmedian carinae costate to costate granular (9) or granular (4) on segments II-V. Dorsosubmedian carinae of metasomal segments II, III and, to a lesser extent IV, each terminating posteriorly with a slightly enlarged, spiniform granule; dorsosubmedian carinae of other metasomal segments without spiniform granules posteriorly. Intercarinal surfaces smooth, except for lateral surfaces of segments II-V, finely granular in d. Telson smooth (@), or finely granular (3), and covered in long macrosetae. Hemispermatophore. Double hook near the base of the distal lamella; distal crest truncate (Figs 22, 23). Geographic variation. Specimens from higher elevations, which receive more rainfall, in the Khomas, Oshikoto and Otjozondjupa Regions of Namibia (e.g., Khomas Hochland, Otavi Highlands and Waterberg) are smaller and darker in colour, than specimens from lower elevations, which receive less rainfall, in Angola and the Erongo and Kunene Regions of Namibia (e.g., Damaraland and Kaokoveld). Ontogenetic variation. The presence of a lobe on the movable finger of the pedi- palp chela and a corresponding notch in the fixed finger (Figs 12, 13) are indicative of sexual maturity in most species of Hadogenes (Lawrence, 1966; Newlands & Prendini, 1997; Prendini, 2001a). The lobe and corresponding notch are absent from the fingers of the pedipalp chela in subadults and juveniles, developing in the final instar of species, such as H. taeniurus, in which these characters are present in the adults. In the specimens of Hadogenes examined for this study, sexual maturity was assessed by the presence of a lobe and notch in males and females, and by the presence L. 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PUIOSEIOMN :AI ewoseoW ‘TL ewoseoW I] eWOSsEyPW :] PWOSEIPMN TIA MUS :ELUOSOSA]N :dyedipaq 384 L. PRENDINI of fully developed paraxial organs in males or the gravid condition in females. The elongated metasoma (longer than the combined length of prosoma and metasoma), a secondary sexual characteristic only acquired in the final instar male (Lamoral, 1979; Newlands, 1980; Prendini, 2001a), is a further indication that male specimens are adult (Fig. 21, cf. female, Fig. 20). In all species of Hadogenes, juvenile males and females resemble each other, and adult females, very closely in general morphological features (besides the absence of a lobe and notch on the pedipalp chela fingers) until the final instar. The metasoma of the juvenile male is also shorter than the combined length of the prosoma and mesosoma. Sexual dimorphism. The characters of primary external sexual dimorphism are the undivided genital operculum of the female, which opens in a single flap (Fig. 9), whereas in the male, the operculum consists of two unconnected sclerites that open in- dependently and cover a pair of genital papillae (Fig. 8). Secondary sexual characters observed in adult males, compared with adult females and juveniles of both sexes, are as follows (Table 1): more slender pedipalp chela, with a less pronounced lobe on the movable finger and a less pronounced notch in the fixed finger (Fig. 12); more slender mesosoma (Fig. 2); metasoma elongated, longer than the combined length of the pro- soma and mesosoma (Fig. 21); increased granulation of the carapace (Fig. 6), tergites and metasoma; granular telson; greater number of pectinal teeth (Fig. 8). Chromosome number. Newlands & Cantrell (1985) recorded a chromosome number of 2n = 100. Remarks. Lourenco (1999: 932) prefaced his description of H. angolensis with a commentary on the known records of Hadogenes from Angola: ‘Sa présence en Angola est limitée a une seule citation de Lawrence (1959), qui signale Hadogenes taeniurus (Thorell) dans ce pays (Vila Arriaga près de Lungo: aujourd’hui Lubango, et Lucira). Newlands (1972), Lamoral (1979) et Lamoral & Reynders (1975) mentionnent à nou- veau Hadogenes taeniurus pour la partie sud du pays, sans pour autant préciser avoir examiné le matériel en question. La faune des scorpions d’ Angola a été jusqu’a présent trés peu connue. Les publications se limitent à celles de Monard (1929, 1937), suivies de Lawrence (1949, 1959, 1961) et Vachon (1950). Seul Lawrence (1959) mentionne le genre Hadogenes, mais il m’a été impossible de lo- caliser le matériel en question. L’étude de trois spécimens collectés dans la région du Plateau de l’Huila, près de Capangombe, permet la description d’une nouvelle espèce, qui présente des similitudes avec H. taeniurus. Ceci soulève la question d’une possible erreur d’identification de la part de Lawrence (1959), non vérifiée par Lamoral (1979). In this commentary, Lourenço (1999) claimed that Lawrence (1959) was solely responsible for the records of H. taeniurus from Angola, that the specimens on which these records were based, were not examined by subsequent authors (i.e., Newlands, 1972a; Lamoral & Reynders, 1975; Lamoral, 1979), and that Lawrence (1959) misidentified these specimens. The implication is that H. taeniurus does not occur in Angola and that these, as well as other Angolan Hadogenes specimens, represent an- other species, for which Lourenço (1999) devised the name H. angolensis. Lourenço’s (1999) claims are false. Newlands (1972a, 1980) listed the spe- cimens examined by Lawrence (1959), which reside in the TMSA, proving that Newlands examined them and concurred with Lawrence’s (1959) identification as H. taeniurus. The specimens referred to are: Lucira (TMSA 8941); Lungo near Vila ON HADOGENES ANGOLENSIS 385 Arriaga (TMSA 8919-8921). Lamoral (1979) did not examine these particular speci- mens, but examined others from Angola, which he also identified as H. taeniurus. These are: Assuncas [Assunçäo] (TMSA 10240); Benguela (NMSA 10002); Oncocua (NMNW 1602 [old 165]). All three authors evidently examined Hadogenes specimens from Angola and considered these specimens to be conspecific with H. taeniurus. Meanwhile, Lourenço (1999) provided no reason as to why H. taeniurus should not occur in Angola, neglected to mention that the type locality of H. taeniurus is unknown and could, in fact, be in Angola — although, as noted by Purcell (1901) and Hewitt (1918), it is more likely to be in the Damaraland region of Namibia — and failed to pro- vide any consistent diagnostic differences by means of which this putative new species might be distinguished from H. taeniurus. According to the brief diagnosis, H. angolensis can be separated from H. taeni- urus according to three differences (Lourengo, 1999): the ‘structure’ of sternite VII; the ‘structure’ of the hemispermatophore, particularly the distal lamella; the number of pectinal teeth (22/23 and 15/16 compared with 17/19 and 13/15 in the males and females of H. angolensis and H. taeniurus, respectively). The first two alleged differ- ences could not be discerned from the illustrations accompanying Lourengo’s (1999) description. Lourenco’s (1999) figures 16 and 17, purportedly illustrating the structural difference in hemispermatophores, are not even directly comparable with his figure 18, reproduced from Lamoral (1979), and there is no obvious difference between them, besides a slight curvature in the distal lamella of the hemispermatophore of H. an- golensis, which could be attributed to intraspecific variation or to an artifact of pre- servation. Lourenco’s (1999) figures 18 and 20 illustrate postero-lateral oval depressions in sternite VII of H. angolensis, a diagnostic character for H. taeniurus (Lawrence, 1928; Newlands, 1972a, 1980; Lamoral, 1979), as illustrated in Lourengo’s (1999) figures 21 and 22, also reproduced from Lamoral (1979). The third putative dif- ference in pectinal tooth count, based on a sample size of two males and one female, must be dismissed on the grounds that pectinal tooth counts are known to vary consi- derably both among and within populations of Hadogenes, as in many other scorpions (Purcell, 1899; Hewitt, 1918; Lamoral, 1979; Prendini, 2001c), and the alleged dif- ferences fall well within the range for H. taeniurus based on data presented by Lawrence (1928: 278) and new a survey of 23 males and 38 females (Table 2). No additional diagnostic differences were discerned in the present study, when the type specimens of H. angolensis were compared directly with those of H. taeniurus and J. hahni and also with 247 non-type specimens of A. taeniurus from Angola and TABLE 2. Pectinal tooth counts in a sample of Hadogenes taeniurus (Thorell, 1876), including the holotype, the syntypes of Zschnurus hahni Peters, 1862, and the holotype and paratype of Hadogenes angolensis Lourenço, 1999. (25) 2 (n= 38) left right left right Range 18-24 18-24 14-20 14-20 Mean 21 20 16 16 Median 21 20 16 16 Mode 19 20 16 16 386 L. PRENDINI Namibia, including all the material examined by Lawrence (1959), Newlands (1972a, 1980), and Lamoral (1979). Instead, the type specimens of H. angolensis displayed the diagnostic characters of H. taeniurus, including the abovementioned depressions in the basal piece and sternite VII, metasomal segment I width greater than height, and the relatively short metasoma of the adult male (23% greater than the combined length of prosoma and mesosoma). On the basis of this evidence, H. angolensis is hereby synonymized with H. taeniurus. Distribution. Hadogenes taeniurus is endemic to rocky and mountainous regions in southern Angola and northern Namibia. Its known distributional range is bordered approximately by the Huila Plateau to the north, the western limit of the Kalahari sand system to the east, the Namibian Central Highlands to the south, and the Skeleton Coast, Engo-Kunene and Baia dos Tigres-Curoca sand systems to the west (Fig. 1). In Angola, the species has been recorded from the Benguela, Cuanza Sul, Cunene, Huila and Namibé Districts. In Namibia, it has been recorded from the Erongo Region (Karibib and Omaruru Districts), Khomas Region (Windhoek District), Kunene Region (Khorixas, Opuwo and Outjo Districts), Oshikoto Region (Tsumeb District), and Otjozondjupa Region (Grootfontein, Okahandja and Otjiwarongo Districts). Although Newlands (1980) cited Uis [21°03’S 14°51’E] as the southernmost record, there are many records further south in the Erongo, Khomas and Otjozondjupa Regions, the southernmost being Farm Uitsig [on Uruganus 358, 22°49’S 16°17’E]. Kraepelin’s (1908) records of H. taeniurus from Kanya [Kanye] (Southern District, Botswana), Steinkopf and Kamaggas [Komaggas] (Northern Cape Province, South Africa), later listed by Lamoral & Reynders (1975), are erroneous (Kraepelin, 1914; Newlands, 1980). Kraepelin (1908) misidentified the specimens in question and later suggested (Kraepelin, 1914) that the specimens from Kamaggas might represent a new species. These specimens were re-examined during the present investigation, and found to be conspecific with Hadogenes gunningi Purcell, 1899 (Kanye, ZMHB 15110) and Hadogenes phyllodes Thorell, 1876 (Steinkopf and Komaggas, ZMHB 15111-15113). Kraepelin’s (1914) record from ‘Ababis, Hereroland’ [Farm Abbabis 3] (Hardap Region, Namibia) is also erroneous, although the specimen was examined at the ZMUH and confirmed to be conspecific with H. taeniurus. According to Newlands (1980), records from Kub [Farm Kub 657] (Hardap Region, Namibia) (AMGS), listed in Newlands (1972a), but supplied by Lawrence (in litt., 1969) were also based on misidentifications. This was also confirmed in the present study. The specimens in question (AMGS 8616, 8873, 8914) are conspecific with Hadogenes zumpti Newlands in Newlands & Cantrell, 1985. Lamoral’s (1979) records of H. taeniurus from the Hoanib River (NMNW 130), Portsmut (NMNW 222) and Gorob Mine (NMNW 517) are also based on misidentifi- cations. These specimens are conspecific with H. tityrus. The known locality records of H. taeniurus fall within the following range of altitudes (percentage of locality records indicated in parentheses): 300-600 m (5%), 600-900 m (27%), 900-1200 m (31%), 1200-1500 m (31%), 1500-1800 m (6%). The distributional range falls mostly within the Savanna biome (Irish, 1994), where locality records occur in the Mopane Savanna (66%), Thornbush Savanna (9%), ON HADOGENES ANGOLENSIS 387 Fics 12-15. Hadogenes taeniurus (Thorell, 1876), carinae, trichobothria and macrosetae on the dextral pedipalpal segments of d and 9 (Farm Uisib 427 [AMNH]). 12. Chela, dorsal aspect, d . 13. Chela, dorsal aspect, 9 . 14. Chela, external aspect, 9 . 15. Chela, ventrointernal aspect, 2. Scale bar = 5 mm. 388 L. PRENDINI Mountain Savanna and Karstveld (7%), Forest Savanna and Woodland (3%), and Highland Savanna (3%) vegetation zones (Giess, 1971). However, a significant pro- portion of records fall within the Desert and Nama Karoo biomes (Irish, 1994), in the following vegetation zones (Giess, 1971): Semi-desert and Savanna Transition (10%), Northern Namib (2%). This species inhabits a moderately to very arid region, receiving less than 650 mm annual rainfall, most of which falls in the summer (December to May). Known locality records fall within the following mean annual rainfall ranges: 0-50 mm (3%), 50-100 mm (4%), 100-150 mm (5%), 150-200 mm (13%), 200-250 mm (13%), 250- 300 mm (21%), 300-350 mm (14%), 350-400 mm (13%), 400-450 mm (4%), 500- 550 mm (3%), 550-600 mm (2%), 600-650 mm (2%). Ecology. Hadogenes taeniurus is an obligate lithophile, which inhabits the narrow cracks, crevices and spaces beneath exfoliations of weathered granite rocks, but can also be found under large flat rocks resting on bedrock (Prendini, 2001b). The distributional range of H. taeniurus is allopatric with that of its closest relatives, H. granulatus and H. troglodytes, which occur on the eastern side of the Kalahari sand system in Botswana, Malawi, Mozambique, Zambia, Zimbabwe and South Africa, and parapatric with that of H. tityrus, which occurs in drier habitats further south and west in Namibia and South Africa (Prendini, 1995, 2001a). Hadogenes taeniurus has been collected in sympatry with A. tityrus, 5 km SE of Blue Drum, Kaokoveld (AMNH), but the two species were not syntopic. Hadogenes taeniurus inhabited large crevices in granite outcrops, whereas the much smaller A. tityrus inhabited narrow fissures in schist outcrops. Hadogenes taeniurus has also been collected in sympatry with other scorpion species, including the bothriurids, Lisposoma elegans Lawrence, 1928 and L. jose- hermana Lamoral, 1979; the buthids, Hottentotta conspersus (Thorell, 1876), Parabuthus brevimanus (Thorell, 1876), P. granulatus (Ehrenberg, 1831), P. kraepeli- ni Werner, 1902, P. villosus (Peters, 1862), Uroplectes otjimbinguensis (Karsch, 1879), and U. planimanus (Karsch, 1879); the scorpionids, Opistophthalmus brevicauda Lawrence, 1928, O. carinatus (Peters, 1861), O. cavimanus Lawrence, 1923, O. chrysites Lawrence, 1967, O. lamorali Prendini, 2000, O. gibbericauda Lamoral, 1979, and O. wahlbergii (Thorell, 1876). Conservation status. Hadogenes taeniurus is largely unaffected by the dual threats of habitat destruction and collection for the exotic pet trade that threaten many other species of Hadogenes (Prendini, 2001a). This species is presently known from ca. 100 localities, falling within 80 QDS, in a region of low agricultural potential (Barnard, 1998) where, besides mining, which occurs at sporadic sites across its dis- tributional range (e.g., Uis), there are few other threats to its survival. Furthermore, the species has been recorded from at least two protected areas (Iona Reserve in Angola; Waterberg Plateau Park in Namibia) and may occur in others. For example, records of H. taeniurus from localities close to the western boundary of Etosha National Park (e.g., Hobatere, Kowares), suggest that it probably occurs in rocky habitats in the south and west of the park. Hadogenes taeniurus is therefore assigned the Least Concern IUCN Red List Category. ON HADOGENES ANGOLENSIS 389 Fics 16-19 Hadogenes taeniurus (Thorell, 1876), carinae, trichobothria and macrosetae on the dextral pedi- palpal segments of © (Farm Uisib 427 [AMNH]). 16. Patella, dorsal aspect. 17. Patella, external aspect. 18. Patella, ventral aspect. 19. Femur, dorsal aspect. Scale bar = 5 mm. 390 L. PRENDINI Material examined (type material, see above). No data: J. Visser, 1 2 (SAMC C4286 [JV 3284]), 2 juv. d (SAMC C4264 [JV 3119], C4265 [JV 3120]). ANGOLA: Benguela District: be- tween Cainbanbo [Caibambo, 13°01’S 14°00’E] and Cubal [13°02’S 14°15’E], 2.iii.1972, J. Visser, 1 juv. d (TMSA 10373), 3.111.1972, J. Visser, 1 subad. d (TMSA 10374); Hanha [13°21’S 14°24’E], 13-18.v.1925, Lang & Boulton, Vernay Angola Expedition, 1 6 3 2 3 sub- ad. d 2 subad. 2 2 juv. d (AMNH [27278]); Lobito Bay [12°20’S 13°34’E], vi.1932, K.H. Barnard, 1 d (SAMC B7343); near Baia Farta, Benguela [12°36’S 13°13’E], 8.ix.1970, J. Visser, 1 juv. d 1 juv. 2 (NMSA 10002). Cuanza Sul District: Gabela, 25 km W [10°51’S 14°15’E], 26.v.1974, W.D. Haacke, 2 subad. 2 (TMSA 10987, 10988), 1 juv. d (TMSA 10986). Cunene District: Cahama, 2 mi NW [16°17’S 14°19°E], 22.11.1971, W.D. Haacke, 1 juv. 2 (TMSA 14332); Oncocua, 23 km W [16°39°S 13°13’E], 6.ix.1969, C.G. Coetzee, 1 juv. 2 (NMNW 1602 [old 165]), 6-7.1x.1969, C.G. Coetzee, 1 4 1 subad. d (NMNW 165); Ruacana [17°24’S 14°12’E], 3.xii.1972, Crawford Cabral, 1 subad. d 1 subad. 2 1 juv. 2 (MRAC 166.889). Huila District: Chibemba, 5 km S [15°46’S 14°45’E], 24.111.1971, W.D. Haacke, 1 4 (TMSA 10234), 1 2 (TMSA 10235); Chibemba, 11 km S [15°47’S 14°45’E], 2.v.1974, WD. Haacke, 1 4 (TMSA 10933), 1 juv. 2 (TMSA 10934); Hungueria [15°19’S 13°32’E], 1970, Crawford Cabral, 1 juv. 6 (MRAC 207.424); Otchinjau, 15 km W towards Oncocua [16°22’S 13°56’E], 10.iv.1971, W.D. Haacke, 1 9 (TMSA 10425). Namibé District: Assunçäo [14°52’S 13°06’E], 24.11.1971, W.D. Haacke, 1 2 (TMSA 10236); Assungào, 5 km E [14°52’S 13°07’E], 26..1971, W.D. Haacke, 1 juv. 2 (TMSA 10240); Lucira [13°52’S 12°32’E], 13.1x.1956, G. Rudebeck, 1 subad. 9 (TMSA 8941 [ex NMSA]); Lungo near Vila Arriaga [14°21’S 13°15’E], 10.ix.1957, G. Rudebeck, 1 9 (TMSA 8919), 1 subad. 2 (TMSA 8921), 1 juv. d [not 2 cf. Lawrence (1959: 386)] (TMSA 8920); Saiona River, 25 km NW Cainde [15°21’S 12°31’E], 16.iv.1971, W.D. Haacke, 1 2 (TMSA 10422), 1 juv. 2 (TMSA 10423); Vila Arriaga, 5 mi E [14°46’S 13°22’E], 21.v.1958, ES. Ross & RE. Leech, 1000 m, 1 2 (CASC). Iona Reserve: Iona, 7 km towards Oncocua [16°56’S 12°37’E], 8.iv.1971, W.D. Haacke, 1 2 (TMSA 10424). NAMIBIA: Erongo Region: Karibib District: Farm Ameib 60 [21°49’S 15°37’E], 12.11.1972, C.G. Coetzee & M.-L. Penrith, 2 juv. 2 (NMNW 333); Dawisaub [Farm Davetsaub 29, 22°27’S 16°11°E],J. Visser, 1 juv. & (SAMC C4179 [JV 3169]), 1-16.iv.1984, J. Visser, 1 juv. 2 (SAMC C4164 [JV 3102]); Farm Dobbelsberg 99 [21°53’S 16°01’E], 13.1.1981, A. Harington, sympatric with P. brevimanus, 1 juv. 6 (AMNH [AH 1792]); Farm Johann Albrechtshöhe 44 [21°56’S 16°05’E], 9.ii1.1970, H. Mittendorf, 1 juv. d (NMNW 160); Farm Springbokfontein 21 [21°37’S 15°27’E], Erongo Mountains, 25.xii.1988, A. Harington, 1 2 2 juv. 9 (NMNW 2038), in cracks on granite hillside, sympatric with P. kraepelini and O. carinatus, 1 9 (AMNH [AH 3671)); Otjimbingue [21°21°S 16°08’E], Wilhelmstal [21°55’S 16°18’E], 1-16.iv.1984, J. Visser, 2 juv. 3 (SAMC C4170 [JV 3147], C4187 [JV 3148]), 1 juv. 2 (SAMC C4171); Otjimbingue to Windhoek [21°55’S 16°18’E], 1-16.iv.1984, J. Visser, 1 juv. 9 (SAMC C4268 [JV 3150]). Omaruru District: ‘Damaraland’, ix.1911, 1 subad. d (ZMUH); Khorixas, 84 km S [20°53’S 15°00’E], 22.1.1986, J. Visser, 1 & (SAMC C4215 [JV 4155]); Ozondati, 16 km N towards Okonyenye Mountains [20°52’S 15°21’E], 1.iv.1979, W.D. Haacke, 1 2 (TMSA 11569); Uis mine, Uis river [21°11’S 14°52’E], 12.ix.1962, B. Grobbelaar, 1 subad. d (NMNW 167); Uis tin mine [21°13’S 14°51’E], Uis, 30.11.1969, J.J. Nel, 1 6 (TMSA 9416); Uis, 5 km W [21°15’S 14°50°E], 15.iv.1980, A. Harington, in rock cracks, syntopic with U. planimanus, 1 juv. 3 (AMNH [AH 1416]); Uis, at Kamanjab turnoff [21°14’S 14°51’E], 18 .iv.1980, A. Harington, in rock crack at summit of extremely rocky hill, 1 2 (AMNH [AH 1356]). Khomas Region: Windhoek District: Farm Uitsig [on Uruganus 358, 22°49’S 16°17’E], Wilhelmstal [21°55’S 16°18’E], 15.11.1963, C. de Wet, 1 6 (NMNW 61); Goreangab Dam [22°32’S 17°02’E], v.1970, H. Strauss, 1 juv. 2 (NMNW 252). Kunene Region: Khorixas District: Farm Engelbrecht 272 [19°56°S 14°42°E], 14.x11.1988, A. Harington, rocky area, sympatric with H. conspersus, P. bre- vimanus, P. villosus, O. carinatus and O. wahlbergii, 1 ¢ (AMNH [AH 4132]); Hobatere Lodge, Farm Marenphil 641, 19°19.89°S 14°22.45°E, 20.1.1998, L. Prendini & E. Scott, 1050 m, granitic hills near camp, in crevices of rock outcrops at night, syntopic with L. elegans, P. krae- pelini, P. villosus, O. carinatus, and O. gibbericauda, 3 ? (AMNH); Hobatere Lodge [19°19°S 14°22’E], 10.viii.1990, S. Braine, 1 2 (NMNW 1475), 11.x.1993, B. Brell, 1 juv. (NMNW 2037), 11.1997, S. Braine, 1 d (NMNW 2046), 2000, S. Braine, 1 d (NMNW 2048); Hobatere [19°15°S 14°23’E], 12.x.1993, M. Griffin & M.A. de Kock, 1 subad. d (NMNW 1556); ON HADOGENES ANGOLENSIS 391 Kamanjab, 68 km W [19°49°S 14°11’E], J. Visser, 1 juv. 2 (SAMC C4208 [JV 4019]); Khorixas, 2 km E [20°22’S 14°59°E], 19.iv.1980, A. Harington, sympatric with O. wahlbergii, 1 subad. 2 (AMNH [AH 1399]); Khorixas, 5 km S [20°24’S 14°58’E], 13.xii.1988, A. Harington, sympatric with H. conspersus, P. brevimanus, P. granulatus, P. kraepelini, O. cari- natus and O. wahlbergii, 1 ¢ (AMNH [AH 4066]); Makukous Spring, 6 km N [19°33’S 13°53’E], 27.iv.1976, W.D. Haacke, 1 juv. & (TMSA 11226); Onguati, 44.3 km N [19°34’S 14°29’E], 18.vi.1993, A. Bauer, 1 2 (NMNW 1529); Welwitschia (Khorixas) [20°22’S 14°58’E], 1.1963, F. Gaerdes, 1 juv. 4 (NMSA 9049). Opuwo District: Anabib (Orupembe), about 100 mi W Ohopoho (Opuwo) [18°11’S 12°31’E], 6.vi.1951, P. & G. Brinck, G. & I. Rudebeck, on dry hillside, 1 juv. & [not 9 cf. Lawrence (1955: 223)] (ZMLU L51/4285); base of Vanzyl’s Pass, 17°39.827’S 12°40.852’E, 11.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 635 m, UV detection, in crevices in schist slope in arid savanna, syntopic with H. conspersus, U. otjimbinguensis, U. planimanus and O. brevicauda, 1 ? (AMNH); Beesvlakte [19°02’S 14°14’E], 15.vi.1992, M. Griffin & M.A. de Kock, 1 d 1 juv. 2 (NMNW 2036); Blue Drum, 5 km SE on S bank of Ondondujengo River, 17°48.894’S 12°25.626’E, 15.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 841 m, UV detection, in crevices in granite outcrops near riverbed, arid savanna, syntopic with H. conspersus, P. brevimanus, P. granulatus, P. villosus, U. otjimbinguensis, U. planimanus, O. brevicauda, O. cavimanus and O. chrysites, H. tityrus collected on same slopes from crevices in schist, 1 d [measured] 1 2 (AMNH), 1 sub- ad. d 1 subad. 2 (NMNW 2665), 1 subad. d (AMCC 126098); Caimaeis [19°20’S 14°00’E], i-iv.1926, S.A. Museum expedition, 1 juv. d 1 juv. 9 [rehydrated] (SAMC B6071); Epupu falls on Kunene River [17°01’S 13°15’E], 26.iv.1970, W.D. Haacke, 1 subad. 9 (TMSA 9799); Epupa falls, Kunene River, 8 km SE [17°03’S 13°14’E], 25.iv.1970, W.D. Haacke, 1 juv. d (TMSA 9798); Farm Kowares 276 [19°03’S 14°21’E], i-iv.1926, S.A. Museum expedition, 1 9 [rehydrated] (SAMC B6967), 1 juv. & [rehydrated] (SAMC B6974); Hartmann Valley, S of Angola border [17°34’S 12°17’E], iii.1995, C.R. Owen, 2 2 (NHRM [JF 80]); Hartmann’s Valley, ca. 39 km N of Orange Drum, 17°26.396’S 12°15.868’E, 14.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 699 m, UV detection, in crevices on granite inselberg on gravel plain, syntopic with A. conspersus, 2 3 [measured] (AMNH); Kaoko Otavi [18°18’S 13°39’E], i-iv.1926, S.A. Museum expedition, 2 juv. & [rehydrated] (SAMC B6801); Kaokoland [17°37’S 12°12’E], 15.x.1988, M. Griffin & H. Kleynhans, between stones, rocky hillside, 1 juv. 9 (NMNW 1069); Kaokoveld, 1.ix.1955, E. van Koen, 1 2 1 juv. d (SMF 39267); Kaokoveld, 17°36.842’S 12°52.107’E, 9.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 1339 m, arid savanna, found walking on rock in the late afternoon, 1 juv. 2 (AMNH); Kunene River [17°26°S 14°02’E], iii.1923, R.F. Lawrence, 1 juv. d 1 juv. 2 (SAMC B5420, damaged), 1 juv. 2 (SAMC B5428, damaged); Kunene River, 37 km E of Epupa [17°01’S 12°55’E], 10-11.iv.1991, M. Griffin, 2 juv. d (NMNW 2049); Ohopoho [Opuwo], 40 km N [17°43’S 13°53’E], 12.11.1975, L. Schulze, 1 juv. d (TMSA 10969); Okokatuwo [17°27°S 12°32’E], 10.v.1991, E. Griffin, between rock slabs, 3 juv. 4 1 juv. 2 (NMNW 1449); Ongongo Community Campsite, 6 km N of Warmquelle, 19°08.427°S 13°49.173’E, 20.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 730 m, UV detection, in crevices in calcrete on rocky slope near freshwater spring, arid savanna, syntopic with A. conspersus, P. brevimanus, U. otjimbinguensis, U. planimanus, O. brevicauda and O. cavimanus, 1 ? [measured] (AMNH), 1 juv. 2 (NMNW 2666); Opura [Opuwa?, 18°04’S 13°51’E], vii-ix.1982, J. Coetzer, 1 2 (NCA 84/83); Oruhona [17°41’S 13°36’E], 28.iv.1970, W.D. Haacke, 2 2 (TMSA 9807, 9809), 1 sub- ad. d (TMSA 9808); Orumana [18°15’S 13°54’E], 111.1973, Mr Vermaak, 1 juv. 2 (NMNW 437); Otjihepa Mountains [17°16’S 12°39’E], 11.vi.1992, M. Griffin & M.A. de Kock, 3 & (NMNW 2052); Otjijangasemo [Otjijandjasemo, 17°26’S 13°16’E], 27.iv.1970, W.D. Haacke, 1 Juv. 2 (TMSA 9800); Otjitanda, 5 km E, 17°38.820’S 13°51.814’E, 9.i.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 1338 m, UV detection, in crevices in granitic hillside in fairly dense broadleaf savanna, syntopic with P. brevimanus, P. granulatus, P. kraepelini, P. vil- losus, U. otjimbinguensis, U. planimanus and O. carinatus, 1 d [measured] 4 2 (AMNH), 1 subad. d 2 subad. 9 (NMNW 2667), 3 juv. & 1 juv. 2 (AMCC 126102); Otjitundua [18°39’S 14°14’E], i-iv.1926, S.A. Museum expedition, 1 juv. d (SAMC B6973, damaged, without meta- soma); Ruacana [17°26’S 14°21’E], 21.iv.1970, W.D. Haacke, 1 juv. ¢ (TMSA 9839); Ruacana, 2 km NNE, 17°26.709’S 14°21.396’E, 6.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. 392 L. PRENDINI Martins, 1138 m, UV detection, in crevices in calcrete on rocky slope (edge of Kunene River val- ley), arid savanna, syntopic with A. conspersus, P. brevimanus and O. carinatus, 1 2 (AMNH), 1 juv. 4 (AMCC 126101), 1 juv. d (NMNW 2673); Ruacana, 5 km NNE, 17°25.869’S 14°21.522’E, 6.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 1109 m, UV detec- tion, in crevices in sandstone at base of rocky slope (edge of Kunene River valley), arid savan- na, syntopic with H. conspersus, P. brevimanus, U. planimanus and O. carinatus, 1 2 (AMNH), 2 subad. 2 (NMNW 2668), 1 juv. d 1 juv. 2 (AMCC 126099); Sanitatas [18°17’S 12°40’E], 30.iv.1970, W.D. Haacke, in crack of layered sandstone-slate, 1 subad. d (TMSA 9811); Swartboois Drift, on Kunene River [17°22’S 13°52’E], 23.iv.1970, W.D. Haacke, 1 2 (TMSA 9790), 1 juv. 2 (TMSA 9789), 24.iv.1970, W.D. Haacke, 1 2 (TMSA 9793), 1 subad. 6 (TMSA 9792), 1 subad. 2 (TMSA 9791), 1 juv. 4 (TMSA 9794); Swartbooisdrif, 2 km E, 17°20.693’S 13°51.427’E, 7.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 732 m, UV detec- tion, in crevices in sandstone on S bank of Kunene River, arid savanna, syntopic with H. cons- persus, P. brevimanus, U. planimanus, O. carinatus and O. gibbericauda, 1 £ 1 subad. 4 (AMNH), 1 ad. [pedipalp chela] 1 juv. d 1 juv. ? (AMCC 126100), 4 juv. 2 (NMNW 2669); Warmquelle [19°01’S 13°49’E], J. Visser, 1 juv. d (SAMC C4210 [JV 4092]); Zebra Mountain North [17°08’S 13°27’E], 31.1.1972, G.J. Menge, 1 juv. & 1 juv. [fragment] (TMSA 10442). Outjo District: Farm Glücksburg 152 [20°15’S 16°06’E], 23.iv.1980, A. Harington, sympatric with H. conspersus, P. brevimanus, U. planimanus, O. carinatus and O. wahlbergii, 1 ? (AMNH [AH 3578]); Farm Hoas 273 [19°55’S 14°45’E], 1971, J.R. Labuschagne, 1 9 (TMSA 10132), 1 juv. [pedipalp] (TMSA 10152), 18.1.1972, J.R. Labuschagne, 1 subad. d (TMSA 10269); Farm Kamanjab 104, ca. 2 km towards Ruacana [19°37’S 14°50°E], 21.iv.1980, A. Harington, in gran- ite rock cracks, 1 2 (AMNH [AH 1317]); Farm Kamanjab 190 [19°34’S 14°52’E], 21.iv.1980, A. Harington, in granite rock cracks near town, 1 2 (AMNH [AH 1387]), 1 subad. 2 (AMNH [AH 1386]); Farm Kaross 237 [19°21°S 14°31’E], 7.x.1986, E. Griffin, in rocks, 1 juv. 6 (NMNW 1209), 10.11.1987, E. Griffin, between slabs of stone, 1 juv. 2 (NMNW 1022); Farm Palafontein 158 [20°15’S 16°11’E], iv.1980, A. Harington, 2 subad. 2 (AMNH [AH 3586, 3587]); Farm Weerlig 11 [20°03’S 15°01’E], 21.iv.1980, A. & L. Harington, in granitic rock cracks, sympatric with O. carinatus and O. wahlbergii, 2 2 (AMNH [AH 1333, 1334]); Kamanjab [19°38’S 14°50’E], J. Visser, 1 juv. & (SAMC C4206 [JV 4007]), 1 juv. 2 (SAMC C4207 [JV 4006]), i-iv.1926, S.A. Museum expedition, 1 subad. 9 [rehydrated] (SAMC B6089), 21.iv.1980, A. Harington, 1 juv. & 1 juv. 2 (AMNH [AH 4260]); Kamanjab, 3 km N [19°37°S 14°48°E], 5.iv.1976, B. Lamoral & L. Ferguson, in vertical crevices of rocky outcrops, at night, 1 juv. 2 (NMSA 10747); Kamanjab, 30 km W [19°36’S 14°33’E], J. Visser, 1 subad. 2 (SAMC C4211 [JV 4093]); Kamanjab, 34 km W [19°36’S 14°10’E], J. Visser, 1 juv. 9 (SAMC C4205 [JV 4002]), 23.1.1986, J. Visser, 1 2 (SAMC C4299 [JV 4167]), 1 2 (SAMC C4305 [JV 4166]). Oshikoto Region: Tsumeb District: Farm Varianto on Elandshoek 771, 19°22.773’S 17°44 45€ E, 4.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 1500 m, UV detection, in crevices in dolomite at base of rocky hill in dense broadleaf savanna, fairly humid with dense litter layer, syntopic with L. josehermana, U. otjimbinguensis, U. planimanus and O. carinatus, 1 2 (AMNH), 1 subad. d 1 juv. d (NMNW 2670); Farm Uithoek 770, 9 mi S Tsumeb [19°20’S 17°39’E], 6.iv.1970, W.D. Haacke, 1 2 (TMSA 9786); Tsumeb [19°13’S 17°43’E], iv.1984, J. Visser, 1 2 (SAMC C4282 [JV 3227]). Otjozondjupa Region: Grootfontein District: Farm Uisib 427, 15 km NW of Otavi, 19°33.132’S 17°14.124’E, 2.1.2004, L. Prendini, E. Scott, T. & C. Bird, Q. & N. Martins, 1293 m, UV detection, in crevices in dolomite at base of rocky hill in dense broadleaf savanna, fairly humid with dense litter layer, syntopic with L. josehermana, U. otjimbinguensis, U. planimanus and O. carinatus, 7 à [1 measured] 14 2 2 subad. & 5 subad. 2 (AMNH), 4 subad. 9 7 juv. d 5 juv. 2 (NMNW 2671), 1 juv. d 1 juv. 2 (AMCC 126103); Tsumeb, 40 km S [19°34’S 17°30’E], J. Visser, 2 juv. 9 (SAMC C4165 [JV 3116], C4166 [JV 3117]). Okahandja District: Gross Barmen [22°06°S 16°45°E], xii.1981, H. Otto, 1 subad. 9 (AMNH [AH 2234]); Okahandja [21°59’S 16°55’E], 1955, F. Gaerdes, 1 juv. d 1 juv. 2 (SMF 39323), v.1960, F. Gaerdes, 1 juv. d [not 2 cf. Lamoral (1979: 657)] (NMSA 7318). Otjiwarongo District: Waterberg [20°25’S 17°15’E], 5.iv.1970, P.J. Buys & P.G. Olivier, 1 juv. 6 (NMNW 168); Waterberg [20°31’S 17°14’E], 5.iv.1970, W.D. Haacke, 1 4 [fragmented, dehydrated] 3 juv. © 1 juv. [exuvium] (TMSA 9835). Waterberg Plateau Park: Main camp, 20°30.803°S 17°14.753’E, 1.1.2004, L. Prendini, E. Scott, ON HADOGENES ANGOLENSIS 393 Fics 20-21 Hadogenes taeniurus (Thorell, 1876), metasoma and telson of d and ® (Farm Uisib 427 [AMNH]). 20. Lateral aspect, 9 . 21. Lateral aspect, d . Scale bar = 10 mm. Q. & N. Martins, 1440 m, UV detection, in crevices in sandstone on eastern slope of Waterberg, fairly dense mixed savanna, sympatric with L. josehermana, U. otjimbinguensis, U. planimanus and O. carinatus, 3 2 [2 measured] (AMNH), 1 subad. & 1 subad. £ 1 juv. 2 (NMNW 2672). Dubious records: NAMIBIA: Erongo Region: Swakopmund District: Hentiesbaai [22°07°S 14°17’E], iv.1970, J. Viljoen, 1 2 (NMNW 74). Hardap Region: Maltahöhe District: Ababis [Farm Abbabis 3, near Solitaire, 23°57’S 16°03’E], 10.x.1910, G.E. Müller, 1 subad. 2 (ZMUH). Karas Region: Bethanie District: Helmeringhausen, just south [25°55’S 16°50’E], J. Visser, 1 juv. 9 (SAMC C4209 [JV 4025]). Hadogenes tityrus (Simon, 1888) Ischnurus tityrus Simon, 1888: 383-384. Hadogenes tityrus: Kraepelin, 1894: 118; Kraepelin, 1896: 136; Kraepelin, 1899: 145-146; Kraepelin, 1901: 272; Kraepelin, 1908: 268; Kraepelin, 1914: 117; Hewitt, 1918: 163; Hewitt, 1925: 270; Werner, 1936: 189; Roewer, 1943: 232; Lawrence, 1955: 252; Lawrence, 1961: 154; Lawrence, 1962: 221; Lawrence, 1966: 5-7, fig. 3c; Newlands, 1972a: 135; Vachon, 1974, fig. 95, 109, 125-127; Lamoral & Reynders, 1975: 540; Lamoral, 1979: 657-661, fig. 302, 303, 310-317; Newlands & Cantrell, 1985: 37-43, fig. 1-3, tab. 1, 2; Lourengo, 1989: 166, pl. 4, fig. 9-11; Newlands & Prendini, 1997: 80, 81; 394 L. PRENDINI Kovarik, 1998: 133; Fet, 2000: 389, 390; Prendini, 2000b: 110, tab. 1; Prendini, 2001a: 148, tab. 1; Prendini, 2001b: 137. Hadogenes bifossulatus Roewer, 1943: 222-234, fig. 10, 10a-d, pl. 5 (synonymized by Kovarik, 1998: 133); Lawrence, 1955: 254; Newlands, 1972a: 134, 135; Lamoral & Reynders, 1975: 538; Lamoral, 1979: 661; Fet, 2000: 387. Hadogenes taeniurus: Lamoral, 1979: 657 (NMNW 130, 222, 517). Hadogenes tityurus: Biicherl, 1964: 61. Type material (examined). Ischnurus tityrus: holotype £ (not 4) (MNHN RS 0378), ‘Kalahari’, South West Africa [NAMIBIA]. Hadogenes bifossulatus: holotype subad. d (not 2) (SMF 6739/146), Waterberg [Otjozondjupa Region: Otjiwarongo District], South West Africa [NAMIBIA]. Remarks. Hadogenes bifossulatus has long been regarded as a dubious species on account of the inadequate original description (Lawrence, 1955; Newlands, 1972a, 1980; Lamoral & Reynders, 1975; Lamoral, 1979; Fet, 2000). Newlands (1972a) sug- gested that H. bifossulatus might be synonymous with H. taeniurus, perhaps because the type locality of H. bifossulatus occurs in an area from which H. taeniurus had pre- viously been recorded, and different Hadogenes species are seldom sympatric. However, after examining the holotype of H. bifossulatus, Newlands (1980) syno- nymized the species with H. tityrus. Newlands’ (1980) unpublished synonymy of A. bi- fossulatus was later published (and thereby validated) by Kovarik (1998), without explicit mention of the source, but Fet (2000) did not accept the synonymy. During the present investigation, the holotype of H. bifossulatus was re-examined and confirmed to be conspecific with H. tityrus, rather than with H. taeniurus. Although not adult, the holotype presents many of the diagnostic characters of H. tityrus: small size; pale colouration; greatly enlarged lateral ocelli; long, slender pedipalp chela with short fingers, and with dorsal and ventral margins of manus subparallel; less than 140 tri- chobothria per pedipalp; very short metasoma, total length 25% less than combined length of prosoma and mesosoma; distinctive shape of the metasomal segments (e.g., metasomal segment I width greater than height, dorsosubmedian carinae of segment III only terminating posteriorly with enlarged, spiniform granules); lateral margins of sternite VII strongly convex. The type locality of H. bifossulatus nevertheless remains questionable, because the nearest records of H. tityrus are much further south and west, in habitats considerably drier than the Waterberg, as is typical for this species. It seems unlikely, although perhaps not impossible, that H. tityrus occurs in the Waterberg. Material examined (type material, see above). NAMIBIA: Erongo Region: Omaruru District: Brandberg [21°14’S 14°30’E], B.H. Lamoral, 1977, 1 2 (TMSA 18350); Brandberg, Basswald Rinne [21°10’S 14°38’E], 17-18.iv.1980, A. Harington, in cracks in red rocks, 2 9 2 subad. 2 (AMNH [AH 1369-1371]); Brandberg, Goaseb gorge [21°14’S 14°35’E], 20.xii.1988, A. Harington, on foothills, under granite rocks, 1 2 (AMNH [AH 4001]); Numas Valley, 21°06’S 14°23’E, 13.viii.1995, M. & E. Griffin, between slabs of stone, 1 d (NMNW 1863); Numaskloof, Brandberg, 21°07.48°S 14°25.54’E, 17.1.1998, L. Prendini & E. Scott, 470 m, rocky granite slopes at base of mountain, UV detection, 1 d (AMNH); Brandberg West mine [21°00°S 14°09’E], 27.iii.1964, F. Motonane, 1 © 1 subad. d (NMNW 90); Brandberg West, 7 km from turnoff towards Uis [21°06’S 14°17’E], 28.1.1981, A. Harington, 1 2 1 juv. d (AMNH [AH 2221]); 5 km SE of Brandberg West-Cape Cross-Uis T-junction [21°05’S 14°16’E], 22.xii.1988, A. Harington, granite exfoliations of low boulders, no hill per se, 1 9 (AMNH [AH 3757]). Swakopmund District: Namib-Naukluft Park: Gorob Mine [23°33’S 15°25°E], 15.11.1974, J. Tebje, 1 6 (NMNW 517) [misidentified as H. taeniurus by Lamoral (1979: 657)]; Rössing, Crusher Dust Area [22°28’S 15°02’E], 28.viii-25.ix.1984, J. Irish & ON HADOGENES ANGOLENSIS 395 Fics 22-23 Hadogenes taeniurus (Thorell, 1876), hemispermatophore (holotype d of H. angolensis Lourenço, 1999, MHNG). 22. Ental aspect. 23. Ectal aspect. Scale bar = 1 mm. H. Rust, preservative pitfall traps, 1 2 (NMNW 867). Khomas Region: Windhoek District: Farm Portsmut [33 23°06’S 16°26’E], 7.11.1969, P.G. Olivier, 1 juv. d (NMNW 222) [misiden- tified as H. taeniurus by Lamoral (1979: 657)]. Kunene Region: Opuwo District: Skeleton Coast Park: Hoanib River, 8 mi E of dunes [19°23’S 13°06’E], 11.xi.1965, C. Brits, 1 2 (NM- NW 130) [misidentified as H. taeniurus by Lamoral (1979: 657)]. ACKNOWLEDGEMENTS Financial support towards the 1998 and 2003/2004 expeditions to Namibia, during which specimens of H. taeniurus were personally collected, was provided by the Department of Pharmacology, University of Stellenbosch (kindly arranged by Gerbus Miiller) and the Stavros S. Niarchos Foundation, respectively. I thank the fol- lowing people and institutions for providing permits to collect scorpions in Namibia and/or assisting with the permit application process during the respective years: Colin Craig and Holger Kolberg, Ministry of Environment and Tourism (MET), Namibia; Eryn Griffin and Tharina Bird (NMNW). I thank the following people for participating on the expeditions and for personally collecting some of the specimens: Elizabeth Scott; Chris and Tharina Bird; Quinton and Nicole Martins. I thank the following people for assisting with the loan of types and additional specimens of Hadogenes and/or allowing access to the collections under their care during my visits: Sarah Gess (AMGS); Charles Griswold and Darrell Ubick (CASC); Charles Lienhard and Lionel Monod (MHNG); Rudy Jocqué (MRAC); Torbjôrn Kronestedt and Julio Ferrer (NHRM); Ted von Proschwitz (NMG); Debbie Jennings (NMSA); Tharina Bird (NMNW); Ansie Dippenaar-Schoeman and Annette van den Berg (NCA); Margie Cochrane (SAMC); Peter Jiger and Julia Altmann (SMF); Klaas Manamela and Barbara Dombrowsky (TMSA); Jason Dunlop (ZMHB); Roy Danielsson (ZMLU); Hieronymus Dastych (ZMUH). This is the fifth paper that includes material from the 396 L. PRENDINI Alexis Harington scorpion collection and I reiterate my thanks to those involved in its transferral to the AMNH, listed by name in previous papers. 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The American Museum of Natural History, New York, 37 pp. SOLEGLAD, M. E. & FET, V. 2003. The scorpion sternum: Structure and phylogeny (Scorpiones: Orthosterni). Euscorpius 5: 1-34. STAHNKE, H. L. 1972. UV light, a useful field tool. BioScience 22: 604-607. THORELL, T. 1876. Etudes scorpiologiques. Atti della Società Italiana di Scienze Naturali 19: 75-272: VACHON, M. 1950. Subsidios para o estudo da biologia na Lunda. Remarques sur les scorpions de l’ Angola (Première note). Publicacöes Culturais da Companhia de Diamantes de Angola (Separata): 5-18. VACHON, M. 1973 [1974]. Étude 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 (3) 140: 857-958. WERNER, F. 1936. Neu-Eingänge von Skorpionen im Zoologischen Museum in Hamburg (pp. 171-193). Festschrift zum 60. Geburtstage von Professor Dr. Embrik Strand, vol. 2. Riga. ON HADOGENES ANGOLENSIS 399 Note added in proof While this paper was in press, Acosta & Fet (2005) reinstated Hadogenes hahni (Peters, 1862) as the valid name for the species referred to in this paper as Hadogenes taeniurus (Thorell, 1876). As noted on p. 372 of the present paper, Prendini (in Fet, 2000: 389) examined the syn- types of Ischnurus hahni Peters, 1862, determined them to be conspecific with H. taeniurus, and proposed that /. hahni and H. taeniurus were synonymous. Fet (2000) gave precedence to Thorell’s (1876) binomen, H. taeniurus, because Peters’ (1862) name, /. hahni, had not been cited since the original description, and H. taeniurus was the only name used for the species since 1876. However, as pointed out by Acosta & Fet (2005), Fet’s (2000) action violates the Principle of Priority (ICZN, 1999). The nearest provision is the Reversal of Precedence (Art. 23.9), according to which priority may be reversed when long unused names are involved, but only if two conditions are met. In this particular case, the first condition (Art. 23.9.1.1) is fulfilled. The senior synonym was not used as a valid name for the species after 1899. However, the second condition (Art. 23.9.1.2) is not. In the 50 years immediately preceding Fet’s (2000) action, there are only nine works citing H. taeniurus as the valid name for the species, although 25 are required. Recognition that H. hahni is the valid name for this species implies that Hadogenes angolensis Lourenço, 1999 = Hadogenes hahni (Peters, 1862), syn. n. ACOSTA, L. E. & FET, V. 2005. Nomenclatural notes in Scorpiones (Arachnida). Zootaxa 934: 1-12. Available from: http://www.mapress.com/zootaxa/. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE (ICZN). 1999. International Code of Zoological Nomenclature. 4fh edition. The International Trust for Zoological Nomen- clature, clo The Natural History Museum, London, 306 pp. (i = MI LAN wo Ta î eal Ora te Beati Inca kn AN ‘A ANR sift tag Fe cot Seo N = ay Prot croi uses de LE , { SM pie 4} Sah REVUE SUISSE DE ZOOLOGIE 112 (2): 401-408; juin 2005 Astyanax pampa (Characiformes, Characidae), a new species from the southernmost boundary of the Brazilian subregion, Argentina Jorge R. CASCIOTTA, Adriana E. ALMIRON & Maria de las Mercedes AZPELICUETA Divisiön Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo, Paseo del Bosque, 1900 La Plata, Argentina. E-mail: jrcas@fcnym.unlp.edu.ar Astyanax pampa (Characiformes, Characidae), a new species from the southernmost boundary of Brazilian subregion, Argentina. - Astyanax pampa sp.n., the new species of characid described herein, is known from arroyo Las Mostazas, an Atlantic Ocean drainage, and the lower rio Colorado basin. Astyanax pampa sp. n. is distinguished from all other Astyanax species by the possession of 17-20 branched anal-fin rays, one maxillary tooth unicuspidate to tricuspidate, short anal-fin base (24.2-30.3% of SL), and long caudal peduncle (10.5-12.7% of SL). Keywords: Freshwater fishes - characids - Buenos Aires - systematics. INTRODUCTION The streams of the Atlantic Ocean drainage and the lower rio Colorado represent the southern limit of the Brazilian subregion. In that area, the diversity of freshwater fishes is low, about 15 species (Casciotta et al., 1999) and supossedly, the fish species are well known. Thus, collecting trips in the streams of Atlantic Ocean drainages and rio Colorado are not frequent. In the last years, some trips in the area allow us to confirm the presence of several freshwater species including a new species of Astyanax. Astyanax pampa is included in this genus following Eigenmann (1921, 1927) by the presence of two series of premaxillary teeth, maxilla with few or no teeth, dentary with strong teeth in front and minute conical ones on the sides, lateral line complete, caudal fin naked, shallow body depth, and a complete series of scales in the predorsal area. The aim of this paper is to describe Astyanax pampa sp. n. which has the southernmost distribution of the genus. MATERIAL AND METHODS The specimens were cleared and counterstained (C&S) following Taylor & Van Dyke (1985). Measurements were taken as straight line distances in mm, using digital calliper; all measurements are expressed as percentages of SL or indicated length. Standard length is taken from tip of snout to hypural joint; head length includes opercular flap; peduncle length is taken from insertion of last anal-fin ray to hypural Manuscript accepted 09.12.2004 402 J. R. CASCIOTTA ET AL. joint. Measurements of holotype and paratypes (Table 1) include two cleared and stained specimens. Values of holotype are indicated by an asterisk. Vertebrae count includes four vertebrae of Weber apparatus and the CP1+U1 as one element. Scales below lateral line are counted from that line to pelvic-fin origin. Perforated scales of lateral line were counted from first scale posterior to opercle to last scale on caudal peduncle. Institutional abbreviations are as listed in Leviton et al. (1985), excluded AI (Asociaciön Ictiolégica, La Plata, Argentina), and CI-FML (Fundacion Miguel Lillo, Tucumän, Argentina). Comparative material examined (SL in mm). Astyanax abramis (Jenyns, 1842): MLP 9427, 2 ex., 102.0-113.0, Argentina, Misiones, rio Parana. Astyanax asuncio- nensis Géry, 1972: MLP 8660, 5 ex., 43.6-61.4, Argentina, Santiago del Estero, Banado de Anatuya. Astyanax eigenmanniorum (Cope, 1894): ANSP 21627-28, 2 paratypes, 42.5-49 4, Brazil, Rio Grande do Sul. AI 167,5 ex., 30.3-54.6, Brazil, Rio Grande do Sul, Rio Jacui basin, Arroio do Conde. MLP 9160, 6 ex., 36.8-80.2, Argentina, Buenos Aires, man-made ponds in Los Talas. Astyanax cf. fasciatus (Cuvier, 1819): MLP 8668, 4 ex., 61.0-67.7, Argentina, Santiago del Estero, Bañado de Figueroa. MLP 8798, 17 ex., 28.8-39.6, Argentina, Formosa, highway from Formosa to Clorinda. Astyanax ita Almirön et al., 2002: MLP 9599, holotype, 64.0, Argentina, Misiones, rio Iguazu basin, arroyo Tateto. Astyanax latens Mirande et al., 2004: CI-FML 3400, holotype, male, 44.3, Argentina, Salta, rio Bermejo basin, arroyo El Oculto. Astyanax leonidas Azpelicueta et al., 2002: MLP 9580, holotype, male, 45.6, Argentina, Misiones, rio Parana basin, headwaters of arroyo Urugua-i. Astyanax ojiara Azpelicueta & Garcia, 2000: MLP 9470, holotype, male, 50.5, Argentina, Misiones, rio Uruguay basin, arroyo Benitez, headwaters of arroyo Yaboti-Mini. Astyanax paris Azpelicueta et al., 2002: MLP 9584, holotype, 75.6, Argentina, Misiones, rio Uruguay basin, Arroyo Fortaleza. Astyanax pynandi Casciotta et al., 2003: MACN-ict 8543, holotype, 52.0, Argentina, Corrientes, Esteros del Iberä, Laguna Iberä, Lobo-Cuä. Astyanax saguazu Casciotta et al., 2003: MLP 9603, holo- type, 63.0, Argentina, Misiones, Uruguay basin, arroyo Once Vueltas. Astyanax steno- halinus Messner, 1962: AI 132, 2 ex. C&S, 42.6-55.7, Argentina, Buenos Aires, Rio de la Plata basin, Punta Indio, unnamed stream. Astyanax troya Azpelicueta et al., 2002: MACN-ict 8310, holotype, 73.8, Argentina, Misiones, rio Parana basin, arroyo Cuñapirü Chico. Astyanax tupi Azpelicueta et al., 2003: MACN-Ict 8646, holotype, 70.1, Argentina, Misiones, arroyo Cuñapirü in Balneario of Aristöbulo del Valle. RESULTS Astyanax pampa sp. n. Figs 1-6, Table 1 Holotype. MACN-ict 8651, 63.7 mm SL, female, Argentina, province of Buenos Aires, arroyo Las Mostazas (38°5°S, 61°27’W), coll. J. Casciotta and A. Almirön, October 1997. Paratypes. AI 134, 8 ex., 38.3-73.0 mm SL, collected with the holotype. MHNG 2644.22, 4 ex., 37.7-65.0 mm SL, collected with the holotype. AI 135, 2 ex. C&S, 43.8-46.0 mm SL, collected with the holotype. AI 166, 2 ex. C&S, 27.5-31 mm SL, Argentina, province of Buenos Aires, rio Colorado, coll. M. Azpelicueta, A. Almirön and J. Casciotta, December 1994. Diagnosis. Astyanax pampa sp. n. is distinguished from all other Astyanax species by the possession of 17-20 branched anal-fin rays, one maxillary tooth uni- NEW ASTYANAX FROM BUENOS AIRES PROVINCE 403 FIG. 1 Astyanax pampa sp. n., holotype, female, MACN-ict 8651, 63.7 mm SL, Argentina, province of Buenos Aires, arroyo Las Mostazas. cuspidate to tricuspidate, short anal-fin base (24.2-30.3% of SL), and long caudal peduncle (10.5-12.7% of SL). Description. Morphometrics of holotype and 14 paratypes are presented in Table 1. Maximum body depth located inmediately anterior to dorsal-fin origin (Fig. 1). Dorsal profile of body convex from snout to dorsal-fin origin, with concavity over supraoccipital area; almost straight from dorsal-fin origin to adipose fin, then slanted ventrally to caudal peduncle, concave along caudal peduncle to base of caudal- fin rays. Ventral profile of body similar to dorsal profile, more curved at pelvic-fin origin. Dorsal-fin origin nearer tip of snout than base of caudal-fin rays. Adipose fin located anterior to vertical through base of posteriormost anal-fin rays. Pelvic-fin origin situated anterior to vertical through dorsal fin-origin. Anal-fin origin placed posterior to vertical through last dorsal-fin ray insertion. Tip of pectoral fin surpassing pelvic-fin origin in males, even in small specimens. Tip of pelvic fin reaching anal-fin origin in males. Head length moderate, mouth horizontal and terminal; snout smaller than eye. Lower jaw slightly longer than upper jaw. Premaxilla with slender ascending process (Fig. 2); alveolar process bearing two series of teeth. Outer row with 3 (1 ex.), 4 (12* ex.), 5 (2 ex.), tricuspidate or tetracuspidate teeth. Inner series with 5 teeth; symphysial tooth tetracuspidate, remaining teeth tricuspidate to pentacuspidate, with central cusp larger. Maxilla long; laminar process with one small unicuspidate or tricuspidate tooth (Fig. 2). Dentary with 3 (11* ex.) or 4 (4 ex.) large teeth penta- cuspidate followed by a median-sized tooth tricuspidate or pentacuspidate, and 3-4 smaller teeth unicuspidate or tricuspidate (Fig. 3). 404 J. R. CASCIOTTA ET AL. TABLE 1. Morphometric data of the holotype and 14 paratypes of Astyanax pampa sp. n. Standard length expressed in mm. Holotype Range Mean SD Standard length 63.7 37.7-73.0 % of standard length Predorsal distance 32.1 49.5-55.0 53.1 1.29 Prepelvic distance 50.5 46.1-52.8 49.6 1272. Preanal distance 68.6 65.1-69.5 67.5 1.24 Body depth 39.6 36.9-42.2 38.7 1637] Dorsal-fin base roll 15271553 14.1 0.61 Anal-fin base ZI, 24.2-30.3 27.8 1.69 Pectoral-fin length 25.0 22.3-26.0 24.5 1.16 Pelvic-fin length 19.3 18.4-21.6 20.0 1.01 Distance between pectoral and pelvic-fin origins 25" 20.5-25.6 229) 1.44 Distance between pelvic and anal-fin origins 21.0 18.5-22.2 20.1 1.10 Head length 27.8 26.5-30.3 28.6 Lil Caudal peduncle depth 13.7, 12.8-14.2 135 0.46 Caudal peduncle length 12.7 10.5-12.7 11.4 0.80 % of head length Snout length 25.4 21.9-26.5 24.0 1229 Eye 322 30.9-42.3 37.1 3.03 Interorbital distance 32.8 31.3-34.8 3851 1.09 Postorbital length 40.1 40.1-46.9 43.4 1.90 Maxillary length 27 25.0-29.9 27.6 1.53 Eye scarcely larger than interorbital. Third infraorbital not contacting latero- sensory canal of preopercle. Dorsal fin with 111,9 rays; first dorsal-fin ray very small, easily visible in some specimens. Distal margin of dorsal fin slightly straight. Anal fin with iii-v, 17 (1* ex.), 18 (4 ex.), 19 (4 ex.), 20 (6 ex.) rays. Males with anal-fin distal margin straight and females with anteriormost rays somewhat lengthened to form a small lobe. Caudal fin with 1,17,i principal rays, upper lobe scarcely shorter and narrower than lower lobe. Pectoral-fin with i,11-14 rays; 11 (1 ex.), 12 (8 ex.), 13 (5* ex.), 14 (1 ex.); margin rounded. Pelvic-fin with 1,7 rays and margin slightly rounded. Hooks on anal and pelvic fins of males; one pair on each segment, on posterior branch of ray. Hooks extended on first to eighth branched anal-fin rays, curved anteriorly. Pelvic fin with large hooks on second to seventh branched rays. Scales cycloid. Lateral series with 35 (7* ex.), 36 (4 ex.), 37 (4 ex.) perforated scales. Scales between dorsal-fin origin and lateral line 6-7, 6 (14* ex.), 7 (1 ex.); scales between lateral line and pelvic-fin origin 5-6, 5 (10 ex.), 6 (5* ex.). Scales around caudal peduncle 16-17. Nine to 13 scales forming an irregular row between tip of supraoccipital spine and dorsal-fin origin. Nine to 12 scales situated along anal-fin base, covering base of all unbranched and first 8 to 12 branched anal-fin rays. Few scales on caudal-fin base. — NEW ASTYANAX FROM BUENOS AIRES PROVINCE 405 FIG. 2 Astyanax pampa sp. n., 46.0 mm SL, right upper jaw in lateral view. Scale bar = 1 mm. In 4 cleared and stained specimens, first branchial arch with 19 to 21 gill rakers (2 on hypobranchial, 11-12 on ceratobranchial, 1 on cartilage, and 7-8 on epibranchial). Caudal fin with 9-10 dorsal and 8-10 ventral procurrent rays. Thirty three to 35 verte- brae. Ten to eleven dorsal-fin pterygiophores between neural spines of vertebrae 9-10 and 18-19; 19 to 20 anal-fin pterygiophores between hemal spines of vertebrae 18-19 and 27-28. Eleven or twelve pairs of ribs. Five or six supraneurals. Color in alcohol: Background light yellowish, dorsal region of body and head sligthly darker. Lateral band extended from posterior margin of opercle to caudal peduncle, forming a caudal spot; this band continuing onto middle caudal-fin rays. One vertical triangular humeral spot with its vertex directed ventrally, located posterior to vertical through pectoral-fin insertion. A second humeral spot very faint, posterior to vertical through middle pectoral fin. Dark chromatophores concentrated on scale mar- gins of uppermost portion of body. Small faint chromatophores present on premaxilla and around nares. Mouth surrounded by chromatophores. All fins hyalines. Anal and caudal fins with dark chromatophores concentrated on distal margins (Fig. 2). Etymology. The specific epithet pampa, a noun in apposition, is a Quichua word that named one of the groups of aborigines that lived in the plain regions of Buenos Aires province. 406 J. R. CASCIOTTA ET AL. Fics 3-4 Right lower jaw in medial view. 3, Astyanax pampa sp. n., 46.0 mm SL; 4, Astyanax eigenman- niorum, 47.8 mm SL. Scale bar = 1 mm. Distribution. Astyanax pampa sp. n. is known from arroyo Las Mostazas and the lower rio Colorado basin, both environments running through the south of Buenos Aires province into the Atlantic Ocean. Habitat. The arroyo Las Mostazas is a short stream with variable depth ranging from 0.60 to 1 m. The water was turbid because of rainfall, with slow current and muddy bottom (Fig. 5). The lower rio Colorado basin presently has a net of irrigation cannals, 0.30 to 1 m deep with the caudal varying between 0.20 to 3 m3 /seg”!; the current velocity can reach 1.5 km/h! (Almirön et al., 1997). DISCUSSION Astyanax pampa Sp. n. comes from arroyo Las Mostazas and the lower rio Colorado basin. Both environments are at the southernmost border of the zoogeo- graphic Brazilian subregion in the cis-Andean basins and they are probably isolated from the remaining Brazilian drainages at least since middle Miocene (Casciotta et al., 1999). Astyanax pampa (cited as A. eigenmanniorum by Almirön et al., 1997; Casciotta et al., 1999) is the only species of the genus inhabiting the streams and rivers of southern Buenos Aires province. The species A. abramis, A. asuncionensis, A. eigenmanniorum, A. fasciatus, and A. stenohalinus do not surpass the Rio de la Plata NEW ASTYANAX FROM BUENOS AIRES PROVINCE 407 FIG. 5 Habitat of Astyanax pampa sp. n., at arroyo Las Mostazas, type locality. basin, about 37° S. There is only one and old record of Astyanax fasciatus (in Eigenmann, 1921 following Günther, 1880) that surpasses the geographical limit of the species mentioned above because it reaches the rfo Negro basin (about 41° S) but it has never been found again in that river. Astyanax pampa sp. n., Cheirodon interruptus (Jenyns, 1842), Bryconamericus iheringii (Boulenger, 1887), Oligosarcus jenynsii (Günther, 1864), and Gymno- characinus bergii Steindachner, 1903, are the species of characiforms with the south- ernmost distribution in South America. Astyanax pampa Sp. n. is similar to A. eigenmanniorum in shape, color pattern, and number of perforated scales of lateral line system. Astyanax pampa sp. n. and A. eigenmanniorum differ in having lower number of branched anal-fin rays (17-20 vs. 22-24), shorter anal-fin base (24.2-30.3 vs. 30.3-32.6 % SL), longer caudal peduncle (10.5-12.7 vs. 8.5-9.4 % SL). Furthermore, A. pampa sp. n. is distinguished from A. eigenmanniorum by the lower number of smaller dentary teeth (2-5 vs. 6-8), the distal tip of maxilla not reaching vs. surpassing the anterior margin of the eye, and depth of dentary higher vs. lower (see both dentaries in Figs 3, 4). Astyanax pampa sp. n. has similar number of branched anal-fin rays that A. brachypterygium Bertaco & Malabarba, 2001, A. cremnobates Bertaco & Malabarba, 2001, A. intermedius Eigenmann 1908, A. ita Almirön et al., 2002, A. lati- ceps (Cope,1894), A. leonidas Azpelicueta et al., 2002, A. paranae Eigenmann, 1914, A. scabripinnis (Jenyns, 1842), and A. obscurus (Hensel, 1870). Astyanax pampa sp. n. is distinguished from A. brachypterygium and A. cremnobates by the longer anal- fin base (24.2-30.3 vs. A. brachypterygium 17.0-23.3 and A. cremnobates 18.4-24.1% of SL). Astyanax pampa sp. n. has 17-20 branched anal-fin rays whereas A. intermedius has 21-26 rays. Astyanax pampa sp. n. bears one unicuspidate to tricuspidate maxillary tooth and A. ita has one heptacuspidate maxillary tooth; in addition, A. laticeps has three or four maxillary teeth. 408 J. R. CASCIOTTA ET AL. The caudal-fin hooks of males in A. leonidas differentiate it from A. pampa sp. n. The body depth of A. paranae (less than 33% of HL) separates A. pampa sp. n. with deeper body (36.9-42.2% of SL). The width of the interorbital (38.5% of HL) and the predorsal length (46.2% of SL) of the holotype of A. scabripinnis (following Melo, 2001) distinguish that species from A. pampa sp. n. (31.3-34.8% of HL and 49.5-55.0% of SL respectively). The numerous dark dots present on body, opercle, and all fins and the body depth (33.3% SL) of A. obscurus differenciatte it from A. pampa sp.n. (36.9- 42.2% SL). ACKNOWLEDGEMENTS The authors thank C. Tremouilles (Museo de La Plata, UNLP) for help with the figures, L. Malabarba (Museu de Ciéncias e Tecnologfa, PUCRS) for gift of A. eigen- manniorum topotypes, S. Körber for translation of the original description of A. obscurus, L. Soibelson for assistance in field trips, Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET) PID 3895/92, Facultad de Ciencias Naturales y Museo (UNLP), and Comisiön de Investigaciones Cientificas de la Provincia de Buenos Aires (CIC) for financial support. REFERENCES ALMIRON, A. E., AZPELICUETA, M. DE LAS M., CASCIOTTA, J. R. & LOPEZ CAZORLA, A. 1997. Icthyogeographic boundary between the Brazilian and Austral subregions in South America, Argentina. Biogeographica 73: 23-30. CASCIOTTA, J., ALMIRON, A., CIONE, A. & AZPELICUETA, M. DE LAS M. 1999. Brazilian freshwater fish assemblages from southern Pampean area, Argentina. Biogeographica 75: 67-78. DE MELO, F. A. G. 2001. Revisäo taxonömica das espécies do género Astyanax Baird e Girard, 1854, (Teleostei: Characiformes: Characidae) da regiäo da Serra dos Orgaos. Arquivos do Museo Nacional de Rio de Janeiro 59: 1-46. EIGENMANN, C. H. 1921. The American Characidae. Memoirs of the Museum of Comparative Zoology 43: 209-310. EIGENMANN, C. H. 1927. The American Characidae. Memoirs of the Museum of Comparative Zoology 43: 311-428. GUNTHER, A. 1880. A contribution to the knowledge of the fish fauna of the Rio de la Plata. Annals Magazine of Natural History, ser. 5, 6: 7-13. Leviton, A. E., GIBBS, Jr., R. H., HEAL, E. & Dawson, C. E. 1985. Standars in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802-832. TAYLOR, W. R. & VAN DyKE, G. C. 1985. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119. REVUE SUISSE DE ZOOLOGIE 112 (2): 409-420; juin 2005 Morphology and systematic status of Coluber karelini mintonorum Mertens, 1969 (Reptilia: Squamata: Colubrinae) Beat SCHATTI! & Andrea STUTZ? ! Apartado postal 383, San Miguel de Allende, Gto. 37700, Republica Mexicana. 2 Talgutstrasse 25, CH-8400 Winterthur, Schweiz. Morphology and systematic status of Coluber karelini mintonorum Mertens, 1969 (Reptilia: Squamata: Colubrinae). - Mintons’ racer is a valid species of the racer genus Platyceps Blyth. It is, so far, only reported from southern Afghanistan and Baluchistan (Iran, Pakistan). Morpho- logically, P. mintonorum (Mertens) differs from all potentially sympatric Platyceps spp. Baluchi records of P. cf. rhodorachis (Jan) are in need of further studies. Keywords: Platyceps mintonorum - species status - morphology - Afghanistan - Baluchistan - Platyceps spp. - sympatry. INTRODUCTION From the systematic point of view, species of the genus Platyceps Blyth, 1860 (see Schatti & Utiger, 2001) are among the most difficult and intriguing taxa within Old World racers. This is particularly true for P. rhodorachis (Jan, 1863) and the type species, P. ventromaculatus (Gray, 1834), reported from northern India to the Near East (e.g., Smith, 1943; Minton, 1966). They are commonly considered to be closely related to P. karelini (Brandt, 1838) from the eastern Caspian region to Pakistan (e.g., Terentjev & Chernov, 1949; Khan, 1997). The taxonomic history of Sindo-Arabian Platyceps spp. is tricky, resulting in considerable confusion about species concepts as well as mixed-up morphological and distribution data. Leviton (1959), for instance, noted that the ranges of P. karelini, P. rhodorachis, and P. ventromaculatus ‘from southwestern Asia [...] overlap most extensively’, and “they exhibit the same ranges of morphological variation, the same color pattern variations, and are found in similar environmental situations”, concluding that “their recognition is based upon characters of rather nebulous taxonomic value, and their present partition is not entirely satisfactory.” Minton (1966) could not identify various racers from Chagai District in Baluchistan, Pakistan, “with Afghan specimens reported by Leviton and Anderson (1961) or with Iranian specimens of rhodorachis reported by Steven C. Anderson (1963).” He considered that these Baluchi snakes “may represent an undescribed form, perhaps related to rhodorachis” (Minton, 1966: 172). A few years later, Coluber kare- lini mintonorum Mertens, 1969, described on the basis of 17 specimens from Chagai Manuscript accepted 22.12.2004 410 B. SCHATTI & A. STUTZ and Kharan District in southwest Pakistan, was named after Sherman A. Minton, Jr., and his wife Madge. Referring to the status of the “Variegated Sand Racer”, Minton (1966) noted that “its relationships to other racers of the Middle East and North Africa will not be clear until monographic treatment of the group is undertaken.” Although Khan (1997) started “a new approach” on “the Coluber karelini-rhodorachis-ventromaculatus species complex”, and despite the description of new obscure taxa (Khan & Khan, 2000), the remarks of Leviton (1959) and Minton (1966) regarding the systematics of Saharo-Sindian Platyceps spp. hold, with some reservation, to the present day. P. rhodorachis auct., reported from the Himalayas to the western Sahara, is a species complex, and Middle East populations of P. ventromaculatus auct. may be conspecific with P. rogersi (Anderson, 1893) (Schätti & McCarthy, 2004; Schätti, 2005). This is the first in a series of papers dedicated to Sindian taxa of the Platyceps rhodorachis-ventromaculatus group of species. The objective of this investigation is to analyse the intraspecific variation and distribution of Mintons’ racer, assess the sys- tematic status of this taxon, and elaborate its morphological distinction from presum- ably closely related species. MATERIAL AND METHODS The study is based on 33 specimens including 14 of the 16 paratypes and Minton’s (1966) “Variegated Sand” racers except SAM 695 and UMMZ 123436. An additional individual, CAS 120543 from “10 km N Darweshan” (Afghanistan), is in very poor condition, and no data were ascertained. The material was borrowed from the American Museum of Natural History, New York (AMNH), California Academy of Sciences, San Francisco (CAS), Field Museum of Natural History, Chicago (FMNH), Muséum d’histoire naturelle, Genève (MHNG), and the Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main (SMF). Further acronyms used in the text are BMNH (The Natural History Museum, London, formerly British Museum of Natural History), NMW (Naturhistorisches Museum, Wien), SAM (collection of Sherman A. Minton), and UMMZ (University of Michigan, Museum of Zoology, Ann Arbor). Morphological terms and cephalic measurements are explained in Schatti (1988) and Schatti & McCarthy (2004). Numbers in parenthesis indicate intraspecific variation. Cephalic measurements were ascertained with a calliper. The head length is in a straight line from the anterior tip of the rostral to the posterior end of the median suture of the parietals. The head width equals the distance between the lateral edge of the supra- oculars at the middle of the eye. The frontal length is along the midline from the anterior tip to the posterior border, and its width equals the maximum distance between the lateral projections. The internasal, prefrontal, and parietal length were measured along their respective median suture. The distance from the nostril to the eye is the shortest span from the posterior border of the external nose opening to the orbit. Scale formulae give the number of longitudinal dorsal scale rows (dsr) at the 20th ventral, midbody, and five ventrals in front of the vent. The reduction pattern is expressed in terms of ventrals and as a percentage of their total number (%ven), based on the average of the right and left side. Maxillary teeth were examined on the right COLUBER KARELINI MINTONORUM 411 bone. The length of the hemipenis in situ and the insertion of the Musculus retractor penis magnus are expressed in absolute numbers of subcaudals and as a percentage thereof (%sub). Scientific names of the taxa discussed in this paper are usually given in full on- ly at their first appearance in the text. Coordinates are from Mertens (1969) and, in the case of Afghanistan, the GEONET database (http://earth-info.nima.mil). RESULTS Platyceps mintonorum (Mertens, 1969) comb. n. — Mintons’ racer Coluber karelini Brandt, 1838 [partim]. - Leviton, 1959: 454, Tb. IV (“Chah-i-Angir”, CAS 84630-33); Leviton & Anderson, 1961: 275 (same material [Chah-i-Anjir]) 1). Coluber sp. - Minton, 1966: 122 [172], Pl. 24.2 [Nushki] (vicinity of Nushki, “2 miles northwest of Ahmad Wal”, and “Chagai” [Nushki]: AMNH 87460, 88464, 96222, FMNH 140277- 78, SAM 695, and UMMZ 123436 [last two not examined]). Coluber karelini mintonorum Mertens, 1969: [3, 10] 56, Fig. 17 [holotype], Tb. [unnumbered]. - “Zangi-Nawar” [29°26’N 65°47’E], Chagai District, Baluchistan, Pakistan [SMF 62942, dé, not examined] (paratypes from Dalbandin, Kharan, Nok Kundi, Nushki, “between Nushki and Dalbandin”, and Surdehgari [not located]: SMF 62923, 62931-37, 62939, 62943 [not examined], 62944-48, 62949 [not examined]). Coluber karelini [partim]. - Gasperetti, 1988: 219, Fig. 29 [dorsal and lateral view of head, dorsal section of body scales: “Chah-i-Angir”, Nushki: CAS 84632, 101593] (see Dis- cussion); Clark, 1990: 33 [35], Tb. [unnumbered] (vicinity of Darweshan, “40 Km. SE Kandahar”: CAS 120541, 120715-17). Coluber sp. - Latifi, 1991: [67] 104 [footnote 3] (see Discussion). Coluber ventromaculatus Gray, 1834 [partim]. - Disi, 1993: 111 (“Pakistan, Belouchistan”: “MHNG 1359.12” [MHNG 2443.12]). Coluber karelini [partim]. - Khan, 1997: 56 (see Discussion). Coluber rhodorachis (Jan, 1863) [partim]. - Khan, 1997: 56 (see Discussion). Coluber karelini mintonorum. - Latifi, 2000: 263 (“Sistan and Baluchistan’). Material examined (an asterisk denotes specimens used for the calculation of head indices; see also Material and Methods). AFGHANISTAN: CAS 84630-33 (“Chah-i-Angir” [Chah- i-Anjir], 31°41’N 64°20’E, 8,3 2 ©), 120541 (10 km N Darweshan, ca. 31°00’N 64°10’E, ©), 120715 (“10-20 km NE Darweshan”, 9), 120716-17 (“40 km SE Kandahar” [31°36°N 65°42’E], 6 9). PAKISTAN (Baluchistan: Chagai and Kharan districts): *AMNH 87460 (“near Nushki”, ca. 29°33’N 66°01’E, 9), *88464 (2 km NW Ahmadwal, ca. 29°25’N 65°56’E, d), 96222 (“2 miles east of Nushki”, 2); *CAS 101498, *101592-93 (Nushki, dd, 9): *FMNH 140277-78 (Nushki, 2 2); *MHNG 2443.11-12 and *2629.98 (“Baluchistan”, 6, 9 2); *SMF 62923 (Dalbandin, 28°53’N 64°26’E, 2 paratype), *62931-32 (“6 Meilen nördlich Nushki”, 9 2 paratypes), 62933 (Kharan, 28°34’N 65°25’E, 2 paratype), 62934 (Nushki, 9 paratype), 62935 (“between Nushki and Dalbandin”, 8 paratype), 62936-37 (Nushki, d $ paratypes), *62939 (Nok Kundi, 28°47’N 62°44’E, d paratype), *62944-48 (Dalbandin, 2 dd,3 989 paratypes). 1) Leviton & Anderson (1961) discussed seven specimens from Chah-i-Anjir (Dasht-i-Margo Desert) reported earlier as Coluber karelini (Leviton, 1959). This material was re-examined by Sergjus A. Chernov (St. Petersburg) who referred CAS 84634-36 (see Appendix) to Platyceps rhodorachis ladacensis (Anderson, 1871) and CAS 84630-33 to P. karelini (as Coluber auct.). However, scale data, the description of the dorsal colour pattern, and accession numbers were mixed up in Leviton & Anderson (1961) as already noted by Mertens (1969: 56). 412 B. SCHATTI & A. STUTZ MORPHOLOGY Snout in dorsal view comparatively pointed, rostral usually projecting “and strongly concave on under side” (Minton, 1966; “auf der Unterseite stark ausgehöhlt”, Mertens, 1969), 1.25-1.56 times broader than high 2). Internasals shorter than pre- frontals, nearly equal length in FMNH 140278, SMF 62923, 62944, and the holotype (fide Mertens, 1969). SMF 62949 (not examined) with two pairs of internasals (“in zwei hinteinanderstehenden Paaren ausgebildet [...], wodurch das Rostrale hinten einen spitzen Fortsatz aufweist”, Mertens, 1969; see footnote 2). Frontal 1.23-1.39 times longer than broad, 1.23-1.70 times longer than internasals and prefrontals, 0.87-1.07 times as long as parietals. Posterior border of the latter somewhat indented towards the median suture. Head 1.90-2.25 times longer than broad. Nostril-eye distance 0.79-0.96 times length of internasals and prefrontals. Loreal rectangular or pentagonal, usually as long as high and situated on second and third supralabial (third in SMF 62931, 62948); coalesced with nasal in SMF 62933. Preocular single, paired or, sometimes, with an incomplete but distinct suture (e.g., in CAS 120541, on right side of CAS 101498, and left in SMF 62945) 3); in contact with frontal except on left side of SMF 62944. Anterior subocular sometimes quite large (e.g., CAS 84631), slightly larger than posterior subocular in AMNH 87460; 2 anterior suboculars in SMF 62933 (first larger), 62942 (left, holotype), and 62949 (fide Mertens, 1969). Normally 9 supralabials (last 3 larger), 8 on right side of holotype (Mertens, 1969), 10 in AMNH 87460 (left side); fifth, fifth and sixth, sixth, or none in contact with eye. Posterior subocular usually present and slightly larger than anterior (see above); absent in CAS 84630, FMNH 140277, SMF 62933 as well as on one side in CAS 120715-16, MHNG 2443.11, 2629.98, SMF 62937, and 62939; 2 posterior sub- oculars on one side in SMF 62935, 62939, and 62948, anterior small. Two postoculars (see footnote 3) and anterior temporals; 2 or 3 (4) scales in second row; anterior temporals comparatively large in SMF 62937, small in CAS 84632, and lower scale generally larger (upper particularly small in SMF 62933). Usually 10 (9-11) sublabials, the 4 anterior in contact with first inframaxillary, sixth (fifth) largest. Anterior chin shields shorter and usually broader than posterior pair; the latter anteriorly separated by 2 rows of scales and usually 4 to 5 (3) posteriorly. Gulars in 4 (3-5) oblique rows between the posterior inframaxillaries and the first ventral. Ventrals 221-240 (Sd 225-231, 22 221-240); anal scute divided; 110-127 (4 d 110-122, 2 2 113-127) paired subcaudals; sum of ventrals and subcaudals 336- 360 (336-350 and 337-360, respectively). Up to 242 ventrals and 128 subcaudals according to Clark (1990) and Latifi (1991), respectively ®. 2) Mertens (1969: 58) noted that the shape of the rostral is a variable feature [“Andererseits gibt es unter mintonorum, vor allem unter den älteren weiblichen Tieren, vereinzelt Individuen, bei denen die Schnauze vorne erheblich abgerundet ist (z. B. bei dem groBen 2 SMF 62949 mit der anomalen Rostralbeschilderung)”]. 3) Minton (1966) and Mertens (1969) included the anterior and posterior subocular(s) in their number of preoculars and postoculars, respectively. 4) Ventral data in Minton (1966), Mertens (1969), and Clark (1990) include preventrals. COLUBER KARELINI MINTONORUM 413 Dorsals with paired apical pits, in 19-19-13 rows except SMF 62944 which has 21 longitudinal dsr from ventrals 47 (right) and 49 (left) to 105 and 101, respectively (increase and decrease involving row 9); this female reduces the number of dsr at ventral 142 (rows 8+9, 61%ven), 147 (3+4, 63%ven), and 187 (6+7, 81%ven). In males, the first and second reduction occur between ventrals 126 and 136 (55-60%ven) and 130-137 (57-60%ven), respectively; the third (last) reduction is situated at the level of ventrals 149-169 (67-74%ven). In females (except SMF 62944), the values are 121- 139 (55-59%ven), 127-145 (56-61 %ven), and 144-171 (64-72%ven), respectively. The sequence of the first and second reduction is variable but, in most cases, the anterior is paravertebral (rows 7+8 in males except CAS 101593 [8+9, left], 7-9 in females); rows 3+4 (2+3 on left side in CAS 84632, 4+5 on left in CAS 101593) are involved in the lateral reduction. The third is paravertebral, or vertebral (i.e., rows 7+8) in four females (CAS 84632, 120717, MHNG 2629.98, SMF 62931). Without specification, Minton (1966) indicated an increase to 21 dsr on the anterior trunk in two specimens and a reduction to 11 dsr in front of the vent in one. Longest specimens 1110 + 360 mm (d, SMF 62942, holotype) and 1170 mm snout-vent length (incomplete tail 287 mm, 2, SMF 62949) according to Mertens (1969: Tb.). Smallest individual (AMNH 96222) 295 + 102 mm. Tail/body length ratio in holotype (3) 0.32, 0.33-0.36 for remaining d d, and 0.32-0.37 in £ 2. To conclude from these data, the snout-vent length of 1190 mm for the holotype as stated in the original diagnosis (Mertens, 1969) is most probably not correct. Dorsal ground colour light tan to pale grey, “top of head often dull reddish, with shields narrowly edged with cream” (Minton, 1966). A dark, light-edged spot between the frontal and parietal and along the parietal suture in the holotype. Circumocular scales light (yellowish) except for a dark blotch below the eye. Temporals yellowish, with irregular darker marks. Dorsal scales dark-edged, forming cross-bars or two series of juxtaposed and slightly transverse paravertebral bars. Dark flecks may also occur along the lower flank. SMF 62943 (not examined) with a more distinct pattern on the anterior portion of the trunk (Mertens, 1969). Venter and underside of tail “immaculate pearly gray to white” (Minton, 1966) or yellowish (e.g., CAS 101592-93). Maxillary with 14-16 teeth, anterior series subisodont, diastema distinct, posterior two teeth enlarged, last offset laterad. Palatine teeth 9. Hemipenis subcylin- drical, sulcus spermaticus simple; spinose throughout, apical area calyculate, apex (in situ) at subcaudals 8-11 (7-10%sub); insertion of Musculus retractor penis magnus at subcaudals 26-28 (21-25%sub). DISTRIBUTION AND ECOLOGY Given the published records and collecting sites of the examined specimens, Mintons’ racer seems to be restricted to southern Afghanistan, southeastern Iran (Sistan-va-Baluchistan), and Baluchistan province (Chagai and Kharan districts) in Pakistan (Fig. 1). “Surdehgari”, the collecting site of a paratype (SMF 62943), could not be located. The specimens reported by Minton (1966) “have all been collected in the desert basin near Nushki.” Mertens (1969) recorded the type series of Coluber karelini mintonorum from the sandy districts of Chagai and Kharan. Material from southeastern B. SCHATTI & A. STUTZ 414 ps mintonorum MHNG 2443.12 (3) from Baluchistan Province, Pakistan. - Platyce PLATE 1 COLUBER KARELINI MINTONORUM 415 Pakistan Fic. 1. Known distribution of Platyceps mintonorum. The localities (@) are based on the examined material and Zangi Nawar, the type locality (%). Triangles (A) mark three collecting sites of P. cf. rhodorachis (see Discussion) from the ““Gandrani Caves” (SAM 679), Band Murad Khan (SMF 62928), and Kharan (SMF 62938). Open symbols denote two approximate locations (“between Nushki and Dalbandin” and “Gandrani Caves”). Iran “in the collection of the State Razi Institute near Teheran” (Minton, 1966) is from “Sistan and Baluchistan province (Zabol, Zahedan)” (Latifi, 1991). Minton’s (1966) Nushki series was obtained “in relatively flat sandy country with sparse vegetation. All were in the open, three being taken between sunrise and midmorning and two between sunset and darkness. They are very quick, alert snakes and savage biters. One captive fed on sand lizards (Acanthodactylus).” According to Clark (1990), Mintons’ racer is “Very common at the Darweshan and Kandahar sites” where it was “found both in non-sandy and sandy biotopes, both firm and loose sands” between 800 and 1’060 m. 416 B. SCHATTI & A. STUTZ One specimen from the vicinity of Kandahar [Qandahar] “was found lying in the open so bloated with a large gecko (Teratoscincus) that it had just consumed that it was incapable of movement” (Clark, 1990: 28, 33). Latifi (1991) reported this species to feed on “mice in captivity.” DISCUSSION Based on head shape, viz., a pointed (“zugespitzt”) snout, and the presence of a posterior subocular or, sometimes, a complete row of scales separating the eye from the supralabials, Mertens (1969) considered Mintons’ racer a subspecies of Platyceps kare- lini (Brandt). This opinion was reached in spite of considerable (“erhebliche”) differences in ventral and subcaudal scale counts as well as a marked fading of the head and dorsal colour pattern vis-a-vis Karelin’s racer. Morphologically, Platyceps mintonorum and P. karelini are easily separable. Apart from differences in body length (less than 750 mm snout-vent length in the latter), dorsal colour pattern (transverse black blotches in karelini), and hemipenis features (length, insertion of retractor muscle; in prep.), these species differ quanti- tatively in a number of head measurements and scale characters including the dorsal scale reduction pattern of males, and they are most distinct in ventral and subcaudal counts and the sum thereof (Tbs 1-2). TABLE 1. Head measurements and cephalic scales in Platyceps mintonorum and P. karelini. Rostral length/height, frontal length/width, length of frontal/internasals and prefrontals, condi- tion of preocular (1: single, 2: paired), number of supralabials in contact with eye, and posterior subocular (0: absent, 1: present). Unusual condition in parenthesis; rare states in brackets. Data for P. karelini are based on 19 specimens from Afghanistan, Iran, and Pakistan (see Appendix). character / rostral frontal frontal /inter- preocular supralabial posterior taxon length length nas. + prefr. cont. eye subocular P. mintonorum 1.25-1.56 1.23-1.39 1.23-1.70 1 or 2 0-2 1 (0) P. karelini 1.50-1.76 1.36-1.53 1.18-1.45 1 [2] 1 [0] 1 TABLE 2. Body scale characters in Platyceps mintonorum and P. karelini (see Tb. 1 and Discussion). Ventrals, subcaudals, and sum of ventrals and subcaudals in dd and 2 9, as well as position of second reduction (from 17 to 15 dsr) in d d ‚expressed in terms of ventrals and as a percentage of their total number (%ven). See Morphology for ventral and subcaudal data of P. mintonorum reported by Clark (1990) and Latifi (1990). character / ventrals subcaudals sum second taxon dé 2g dé 2g dé QUE reduction (dd) P. mintonorum 225-231 221-240 110-122 113-127 336-350 337-360 130-137 (57-60) P. karelini 193-207 202-212 90-105 86-106 290-311 292-314 119-130 (60-64) Higher body scale counts for Platyceps karelini than reported here are given, for instance, for a female from the Kavir Desert in Iran with 216 ventrals (Nilson & Andrén, 1981) and an unsexed specimen from Mastung, Pakistan, with 110 subcaudals (Khan & Ahmed, 1987). Without further evidence (“nach Literatur-Angaben”), Mertens (1969) noted up to 220 ventrals and 117 subcaudals for P. karelini. These data exceed by far authentic values ard, therefore, seem highly unlikely. Khan’s (1997) total counts (282-317) for this species were probably added up from the extremes of ventrals (192-207) and subcaudals (90-110). — COLUBER KARELINI MINTONORUM 417 Ultimately, alleged parapatric ranges led Mertens (1969) to refer the new taxon to Platyceps karelini (“Die neue Form ist [...] nichts anderes als ein Vikariant von kare- lini, der mit dieser Natter kaum [sic] sympatrisch auftritt”) 5). However, sympatry of Mintons’ racer and P. karelini is documented, for instance, near Darzi Chah, roughly 65 km west of Nushki (SMF 64629, Mertens, 1969), probably in Afghanistan (see Appendix). There, these taxa occur together, among other places, at Chah-i-Anjir (CAS 84634-35, see footnote 1, Appendix, and below). Mertens (1969) thought that at least some of the specimens reported by Boulenger (1889) as “Zamenis karelini” from the border region in Afghanistan (“aus dem afghanischen Grenzgebiet”) probably belonged to Mintons’ racer. However, three specimens from the “Helmund River” and “between Tirphul and Kilki” (Herat province) collected by the Afghan Boundary Commission and deposited in the BMNH collection with 200-209 ventrals and 86-106 subcaudals (Boulenger, 1893: 402) are not Platyceps mintonorum (unpubl. data). Minton (1966) correctly stated that “Although I originally identified these Nushki snakes as karelini on the basis of circumocular lepidosis, the two are quite different in pattern and in ventral and subcaudal counts.” That author emphasised that “the regular presence of a third preocular, high ventral count, and body pattern” as well as a “more pointed” snout distinguish the “Variegated Sand Racer” (Platyceps mintonorum) from P. karelini (Brandt), P. rhodorachis (Jan), and P. ventromaculatus (Gray) (see footnotes 2-3 and 5). Minton (1966) concluded that the “nearest relative appears to be rhodorachis.” Latifi (1991), most likely following this quotation, con- sidered Mintons’ racer to be “a valid taxon but probably not a subspecies of karelini. It may be closer to rhodorachis.” Khan (1997) believed that two Platyceps mintonorum figured in Gasperetti (1988: Fig. 29), 1.e., CAS 84632 from Chah-i-Anjir (Afghanistan, see footnote 1) and CAS 101593 from Nushki, were P. karelini, and that “specimens from this region have neither dorsal pattern nor orbito-labial and temporal stripes as vivid as observed in C.[oluber] karelini from Quetta-Peshin, Baluchistan.” According to Khan (1997), Mintons’ racer was “erected [...] on similar C. rhodorachis from the Baluchistan high- land.” This erroneous conclusion resulted from enormous confusion as to the system- atics of certain Sindian Platyceps spp. Minton (1966) and Mertens (1969) stated that, apart from head shape and the presence or absence of the posterior subocular, Platyceps mintonorum and P. rhodo- rachis auct. differ in their habitus (rhodorachis is more slender). However, the speci- fic allocation of some female Baluchi racers referred to P. rhodorachis and their 5) From a comparison with Platyceps rhodorachis and P. ventromaculatus (as Coluber spp.) “ziehe ich den SchluB, daB mintonorum, ein Vikariant von karelini, am besten als Subspecies von dieser Natter aufzufassen ist. Die kleine Liicke zwischen der Plusvariante bei karelini (220) und der Minusvariante bei mintonorum (227) wird zweifellos durch weitere Funde tiber- briickt werden. Bei der Zahl der Subcaudalia ist das bereits der Fall [...]. Auf den Zeichnungsunterschied ist kein großer Wert zu legen” (Mertens, 1969: 59). “In der äußeren Erscheinungsform herrscht zwischen der neuen Form und karelini, abgesehen von der Zeichnung, eine recht große Übereinstimmung. Der spitze Kopf ist bis zu einem gewissen Grade auch karelini eigen [...]. Er ist stärker zugespitzt und vor allem die Schnauze stärker vorspringend als bei rhodorachis, ventrimaculatus [sic] oder [Hemorrhois] ravergieri, aber keinesfalls stärker als bei den meisten karelini” (Mertens, 1969: 58). 418 B. SCHATTI & A. STUTZ distinction from P. mintonorum require some comments ©). This is the case with SAM 679 from the “Kud River [approx. 26°17’N 66°13’E] near the Las Bela-Kalat border” (“Gandrani Caves”), SMF 62938 from Kharan, and SMF 62928 (subadult) from Band Murad Khan (25°06’N 67°00°E) on the Hab River (Fig. 1). SAM 679 (not examined) shows “a pattern more suggestive of this form [i.e., Platyceps mintonorum] than of coastal rhodorachis, and circumocular scales as in rhodorachis” (Minton, 1966: 123). The latter feature, 1.e., the lack of a posterior subocular, does not allow a positive identification of that species and a distinction from P. mintonorum (see Morphology). SMF 62928 (237 ventrals, 124 subcaudals) and 62938 (236 and 115, respectively; see footnote 4) possess, as SAM 679, a single anterior subocular and two supralabials (fifth and sixth) entering the eye (i.e., no posterior subocular). The dorsal colour pattern of SMF 62938 is surprisingly similar (“Erstaunlich ähnlich”) to a female Platyceps mintonorum (SMF 62933) obtained at the same locality (Kharan) but the specimen lacks a posterior subocular and the rostral is not projecting (“zeichnet sich [...] außer durch die bezeichnende Subocular-Pholidose durch ein kleines, nicht vorspringendes Rostrale aus”, Mertens, 1969: 59). However, that author emphasised that, occasionally, the snout of Mintons’ racer is considerably rounded, particularly in large females (see footnote 2). SMF 62938 has the loreal and nasal fused on the left side. SMF 62928 is remarkable for somewhat elevated head indices vis-a-vis Platyceps mintonorum (i.e., frontal 1.73 times longer than internasals and prefrontals, head length/width 2.34, nostril-eye distance 1.0), the shape of the posterior border of the parietals (straight instead of indented), a tail/body ratio of 0.38 (snout-vent and tail length 365 + 140 mm, respectively), and dsr reduction pattern, viz., the second fusion involving the fourth and fifth (instead of third and fourth) rows. For the time being, it cannot be ruled out that SMF 62928 and/or 62938 belong to Platyceps mintonorum, or that this species intergrades with P. rhodorachis auct. By all means, their systematic status requires further studies. ACKNOWLEDGEMENTS Thanks to Robert C. Drewes and Jens V. Vindum (San Francisco), Konrad Klemmer (Frankfurt/M.), Harold K. Voris (Chicago), and Richard A. Zweifel (New York) for approving the loans of specimens. 6) Khan & Khan’s (2000) description of three new racer taxa from Pakistan begs a thorough response (in prep.). In any case, Coluber ventromaculatus bengalensis Khan & Khan, 2000 and C. ventromaculatus khanorum Barabanov, 2002 (nomen novum) are objective junior synonyms of C. ventromaculatus Gray, 1834. The description of C. ventromaculatus indusai Khan & Khan, 2000 violates rules of the code (e.g., Art. 16.4, ICZN, 1999; Hallermann et al., 2001). COLUBER KARELINI MINTONORUM 419 REFERENCES ANDERSON, J. 1871. A list of the reptilian accession to the Indian Museum, Calcutta, from 1865 to 1870, with a description of some new species. Journal of the Asiatic Society of Bengal [2] 40 (1): 12-39. ANDERSON, J. 1893. On a new species of Zamenis and a new species of Bufo from Egypt. The Annals and Magazine of Natural History [6] 12: 439-440. ANDERSON, S. C. 1963. Amphibians and reptiles from Iran. 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ScHATTI, B. & MCCARTHY, C. 2004. Saharo-Arabian racers of the Platyceps rhodorachis complex — description of a new species (Reptilia: Squamata: Colubrinae). Revue suisse de Zoologie 111 (4): 691-705. SCHATTI, B. & UTIGER, U. 2001. Hemerophis, a new genus for Zamenis socotrae Giinther, and a contribution to the phylogeny of Old World racers, whip snakes, and related genera (Reptilia: Squamata: Colubrinae). Revue suisse de Zoologie 108 (4): 919-948. SMITH, M. A. 1943. The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese sub-region. Reptilia and Amphibia. Vol. III. Serpentes. London, Taylor & Francis, XI + 583 pp. TERENTJEV, P. V. & CHERNOV, S. A. 1949. Opredelitel presmykajuszezikhsja i zemnovodnykh [Determination key to reptiles and amphibians] (31 enlarged ed.). Moscow & Leningrad, Sovjet Science Press, 340 pp. APPENDIX. Comparative sample of Platyceps karelini. AFGHANISTAN: BMNH 73.1.7.10 (“Kila-i-Fath, Seistan”, juv.), 82.3.20.2 (Kandahar, 3), 86.9.21.102 (Tirpul, 9), 86.9.21.103 (“Kilki”, 9); CAS 84634-36 (Chah-i-Anjir, 2, juv. dd), 103785 (Herat — Islam Qala, ©), 120540 (vicinity of Tashkurghan, 3), 120714 (45 km E Herat, 2 ); SMF 64629 (Darzi Chah, juv. 6). IRAN: FMNH 141604 (Pahlavi Dezh, 9); NMW 25446.3 (“Vor Neh”, juv.), 25446.4 (“Zirkuch”, 9), 25446.5 (“Gulu Chakak”, 2), 25446.6 (“Tscha Sam”, ). PAKISTAN: AMNH 96219 (“near Pishin”, 6); SMF 62924 (Khuzdar, 3), 62940 (Quetta, 2). REVUE SUISSE DE ZOOLOGIE 112 (2): 421-438; juin 2005 Oribatids from Brunei IV (Acari: Oribatida). (Acarologica Genavensia CVI) Sandor MAHUNKA Department of Zoology, Hungarian Natural History Museum and Systematic Zoology Research Group of the Hungarian Academy of Sciences, Baross utca 13, H-1088 Budapest, Hungary. E-mail: mahunka@zoo.zoo.nhmus.hu Oribatids from Brunei IV (Acari: Oribatida) (Acarologica Genavensia CVI). - Fifteen oribatid species are recorded from Brunei; five are new to science and for four of them a new genus (Nasobelba gen. n.) is established in the family Suctobelbidae. The following new combinations are proposed: Nasobelba inenodabilis (Hammer, 1980) comb. n. (= Suctobelbella ineno- dabilis Hammer, 1980) and Nasobelba transitoria (Balogh & Mahunka, 1974) comb. n. (= Suctobelba transitoria Balogh & Mahunka, 1974). A list of the hitherto published oribatid species of Brunei is given in addition. Keywords: Acari - Oribatida - taxonomy - new genus - new species - Brunei. INTRODUCTION The oribatid fauna of Brunei is exceptionally rich and therefore its study extremely rewarding. In this part of the island of Borneo some restricted parts of the mountain vegetation are still well preserved and also large patches of the Towland rain forests are intact. For this reason Dr B. Hauser, then Head of the Arthropoda Department of the Muséum d’histoire naturelle, Geneva, undertook alone a second collecting trip in 1996 to Brunei (with stops in Singapore and Hong Kong) with a focus on soil fauna. I had the opportunity to study a small part of this material and again I found many interesting species. So far, I have published three papers discussing the oribatids living in Brunei (Mahunka 1995, 1997, 2001). Besides giving new distributional data of some rare and hardly known species (e.g. Gehypochthonius xarifae Strenzke, 1963, Epilohmannoides esulcatus Ohkubo, 1979, Notogalumna praetiosa Sellnick, 1959), I there described 37 new species. In my last paper it struck me that the suctobelbid fauna of this territory is exceptionally rich. Therefore, in my new investigations, I concentrated upon this family. Five species are again new for science and four also represent a new genus Manuscript accepted 02.09.2004 422 S. MAHUNKA (Nasobelba gen. n.). The fifth species is a peculiar new species of Parasuctobelba Hammer, 1977. I describe the new taxa in this paper, completing them with some known species and add new distributional data. In describing the new species, I use morphological terms applied in earlier work (e.g., Mahunka & Mahunka-Papp, 2001). A first list of the hitherto published 64 species of beetle mites from Brunei concludes this study. LIST OF IDENTIFIED SPECIES For localities see list at the end of this paper (p. 436). Phthiracaridae Perty, 1841 Notophthiracarus hauseri Mahunka, 1995 Locality: SBH-96/12: 10 specimens. Distribution: Brunei (hitherto known from the type locality only): second record for Brunei. Temburongiidae Mahunka, 1990 Temburongia patoi Mahunka, 1990 Localities: SBH-96/11: 3 specimens; SBH-96/12: 4 specimens. Distribution: Sarawak, Brunei. Microtegeidae Balogh, 1961 Suctotegeus tumescitus Mahunka, 1987 Locality: SBH-96/12: 5 specimens. Distribution: Sabah, Brunei: second record for Brunei. Microzetidae Grandjean, 1936 Teraja sungai Mahunka, 1997 Locality: SBH-96/9: 2 specimens. Distribution: Brunei. Damaeolidae Grandjean, 1965 Fosseremus laciniatus (Berlese, 1905) Locality: SBH-96/11: 2 specimens. Distribution: Cosmopolitan (?); second record for Brunei. Carabodidae C. L. Koch, 1837 Hardybodes penicillatus Mahunka, 1995 Locality: SBH-96/12: 3 specimens. Distribution: Brunei. Dampfiellidae Balogh, 1961 Dampfiella zellwegeri Mahunka, 1997 Locality: SBH-96/11: 3 specimens. Distribution: Brunei (hitherto known from two localities only): second record for Brunei. Rhynchoribatidae Balogh, 1961 Oxyamerus hauserorum Mahunka, 1987 Localities: SBH-96/11: 3 specimens; SBH-96/12: 2 specimens. Distribution: Sabah, Brunei: first record for Brunei. Suctoribates foliatus Mahunka, 1997 Localities: SBH-96/11: 2 specimens; SBH-96/12: 5 specimens; SBH-96/15a: 3 spe- cimens. Distribution: Brunei (hitherto known from two localities only): second record for Brunei. Suctobelbidae Jacot, 1938 Condylobelba bruneiensis Mahunka, 2001 Locality: SBH-96/12: 3 specimens. Distribution: Brunei (hitherto known from six localities): second record for Brunei. Nasobelba agathis gen. n., sp. n. Locality: SBH-96/12. Nasobelba coronata gen. n., sp. n. Localities: SBH-96/11, SBH-96/15a. ORIBATIDS FROM BRUNEI IV 423 Nasobelba hauseri gen. n., sp. n. Locality: SBH-96/11. Nasobelba undosa gen. n., sp. n. Localities: SBH-96/9, SBH-96/11. Parasuctobelba quinquecostata sp. n. Locality: SBH-96/15a. DESCRIPTIONS OF NEW TAXA Nasobelba gen. n. Diagnosis: Family Suctobelbidae. Rostral apex nasiform, following a rostral elevation, with large rostral and smaller accessory teeth. Tectopedial field reduced, paratectopedial field absent. Lamellar knob present, prebothridial rib mostly absent, or reduced. Bothridial lobe present, interbothridial field often reduced. Sensillus with a conspicuously long peduncle. Two pairs of notogastral condyles fusing toone pair of large apophyses in the sejugal region. Polygonal pattern on notogaster and ventral plate. Sternal apodemes more or less reduced, epimeral fields not touching medially. Epimeral setal formula: 3 — 1 — 3 — 3. Anogenital setal formula: 6 — 1 — 2 — 3. Setae ad; in paraanal position. Type species: Nasobelba agathis sp. n. Remarks: Hammer (1980) already drew the attention to the difficult generic placement of Suctobelbella inenodabilis from Java. She also properly placed Sucto- belba transitoria Balogh & Mahunka, 1974 in the same species group. I relegate both species to this new genus: Nasobelba inenodabilis (Hammer, 1980) comb. n. and Nasobelba transitoria (Balogh & Mahunka, 1974) comb. n. The new genus is well characterised by the somewhat sclerotised anteromedian part of the prodorsum, the form of the sensillus, the peculiar pattern on the notogaster and ventral plate, and the position of the adanal setae. It seems to be related to the genus Fenestrobelba Balogh, 1970, however, the new genus is distinguishable from Fenestrobelba by te presence of its tectopedial field and lamellar knob, by the form of sensillus, the pattern on the surfaces and by the large rostral tooth, which is missing in Fenestrobelba (Balogh, 1970; Balogh & Balogh, 1992). Derivatio nominis: Named after the form of the rostral apex. Nasobelba agathis sp. n. Figs 1-6 Material examined: Brunei: Holotype: SBH-96/12, 2 paratypes from the same sample. Holotype and 1 paratype: MHNGI, 1 paratype: (1664-PO-02) HNHM?. Diagnosis: Rostral apex nasiform, well protruding, rostral tooth large. Anterior part of prodorsum with small tubercles arranged in longitudinal rows. Head of the sen- sillus symmetric. Notogastral condyles partly separated from each other. Median part of the notogaster and the ventral plate with polygonal pattern. Nine pairs of long and simple notogastral setae, setae h, much shorter than h;. Measurements: Length of body: 245-267 pm, width of body: 110-114 um. ! MHNG = deposited in the Muséum d’histoire naturelle, Geneva. 2 HNHM = deposited in the Hungarian Natural History Museum, Budapest, with the identi- fication number of the specimens in the Collection of Arachnida. 424 S. MAHUNKA Fics 1-3 Nasobelba agathis gen. n., sp. n. — 1, body in dorsal view; 2, body in ventral view; 3, body in lateral view. ORIBATIDS FROM BRUNEI IV 425 Fics 4-6 Nasobelba agathis gen. n., sp. n. — 4, basal part of prodorsum; 5, rostrum in lateral view; 6, genital region. Prodorsum: Rostral apex strongly protruding, nasiform. Rostral teeth larger than the 2 equally long accessory teeth. Rostral elevation well separated, surface with small tubercles posteriorly, arranged in longitudinal column. Tectopedial fields comparatively small, located posteriorly, near to bothridium (Fig. 1). Paratectopedial field reduced, only a small part visible anteriorly, therefore anterior part of prodorsum nearly smooth. Lamellar knob normally developed, interbothridial field well developed basally, along the bothridial lobe (Fig. 4). Rostral setae geniculate, lamellar and inter- lamellar setae short, simple. Sensillus very long, its head symmetrical, widely lan- ceolate. Lateral part of podosoma: Lateroprodorsal pattern absent. Rostral elevation rising from the prodorsal surface, well tuberculate or granulate (Fig. 5). Exobothridial region (Fig. 3) well sclerotised. Notogaster: Lateral notogastral condyles fused with the median ones, but their original form observable (Fig. 1). Typical polygonal pattern seen on the middle part of the notogaster. Among the fields observable are also larger ones. Nine pairs of long and simple notogastral setae, c narrower than la and ms. Setae h, much shorter than hp. Setae p, and p, minute. Ventral parts (Fig. 2): Epimeres located far from each other, sternal apodeme absent, there is a wide sternal field in this region. Epimeral setal formula: 3 — 1 — 3 — 3. All setae well developed, comparatively long. Some tubercles present along the posterior epimeral borders. Ventral plate with well developed polygonal pattern reaching over to the anal plates posteriorly, but absent laterally. Anogenital setal 426 S. MAHUNKA formula: 6 — 1 — 2 — 3. Anterior and posterior genital setae are longer than those in the middle part of the genital plates (Fig. 6). Aggenital setae located laterally, much farther from each other than the adanal setae. Setae ad; in paraanal position, all setae in this region long. Lyrifissure iad conspicuously long. Legs: Trochanter III and IV and all femora granulate. Remarks: See Nasobelba undosa sp. n. Derivatio nominis: Named after the Agathis (Araucariaceae) forest in Brunei. The name is a noun in apposition. Nasobelba coronata sp. n. Figs 7-11 Material examined: Brunei: Holotype: SBH-96/11, 11 paratypes from the same sample and 2 paratypes: SBH-96/15a. Holotype and 8 paratypes deposited in MHNG, 2 paratypes in HNHM (1665-PO-02). Diagnosis: Rostral apex nasiform, fused with a plate bearing small median teeth, these teeth reaching over the apex. Rostral teeth very large. Anterior part of prodorsum with a few small irregularly arranged tubercles. Head of the sensillus asym- metric. Notogastral condyles completely fused. Median part of notogaster and the ven- tral plate with irregular pattern. Nine pairs of notogastral setae, h,; and h, equal in length. Anal plates granulate medially. Measurements: Length of body: 198-211 wm, width of body: 104-112 pm. Prodorsum: Rostral apex seems to be very wide, true nasiform apex hardly observable. The apices of the anteromedian teeth, bearing the rostral plate and the rostral teeth, form a wide, crown-shaped anterior margin (Fig. 11). Rostral elevation weakly developed, only with some, 2-3, small tubercles. Outer margin of the tecto- pedial field conspicuously long (Fig. 7), a weak paratectopedial field observable ante- riorly. Lamellar knob well developed, located basally between the bothridia. Rostral setae geniculate, lamellar setae very long, robust. Interbothridial setae short, directed backwards. Sensillus very long, its head asymmetrical. Lateral part of podosoma: Lateroprodorsal pattern conspicous, polygonal (Fig. 10). Exobothridial region well sclerotised. Notogaster: Notogastral condyles fused, seem to be one pair condyles with two apices each. Notogastral pattern small, located medially, formed of irregular lines (Fig. 7). Nine pairs of notogastral setae, their ratio different from the previous species. Setae h, slightly shorter than h,, but longer than h3. Setae p, and p> short, but not minute. Ventral parts: Basically similar to the preceding species but the epimeral setae longer (Fig. 8), setae /b particularly long. Setae Ja and 4b arising very near to each other. Along the posterior epimeral border a line of small tubercles present, a pile of secretions observable behind them. Ventral plate with a region of interconnecting lines just posterior to genital plates. Anogenital setal formula: 6 — 1 — 2 — 3. The position of setae also similar to that of the preceding species. Surface of anal plates granulate, the granules form an elliptical pattern (Fig. 9). Legs: Similar to the preceding species. Remarks: See Nasobelba undosa sp. n. Derivatio nominis: Named after the shape of the structure in the rostral region. 427 ORIBATIDS FROM BRUNEI IV Fics 7-11 Nasobelba coronata gen. n., sp. n. — 7, body in dorsal view; 8. body in ventral view; 9, anal plates; 10, podosoma in lateral view; 11, rostral part of prodorsum. 428 S. MAHUNKA Nasobelba hauseri sp. n. Figs 12-16 Material examined: Brunei: Holotype: SBH-96/11. 2 paratypes from the same sample. Holotype and 1 paratype deposited in MHNG, 1 paratype in HNHM (1666-PO-02). Diagnosis: Rostral apex wide, rostral and 2 pairs of accessory teeth nearly equal. Surface of prodorsum without tubercles. Head of the sensillus symmetric, narrow. Notogastral condyles fused. Whole surface of notogaster and ventral plate with a dense polygonal pattern. Nine pairs of very long notogastral setae, h; shorter than hp. Anal plates smooth. Measurements: Length of body: 232-237 pm, width of body: 118-122 um. Prodorsum: Rostral apex nasiform. Rostral teeth large, accessory teeth of equal length (Fig. 14). Rostral elevation separated, its surface smooth. Tectopedial fields comparatively small, located posteriorly near to the bothridium (Fig. 12). Para- tectopedial field reduced, only a small part visible anteriorly. Lamellar knob weakly developed, its anterior border opened or absent and seemingly consisting of two parts. Interbothridial field weakly developed, its posterior margins and the same part of the basal lobe finely granulate. Rostral setae geniculate, lamellar setae long, interlamellar setae short, simple. Sensillus very long, its head symmetrically lanceolate, narrow. Lateral part of podosoma: Lateroprodorsal pattern absent. Rostral elevation ris- ing from the prodorsal surface, smooth. Exobothridial region well sclerotised (Fig. 16). Notogaster: Lateral notogastral condyles fused with the median ones, both apices rounded anteriorly. Notogastral pattern covering nearly the entire surface, polygonal, consisting of mostly small fields. Nine pairs of notogastral setae, some of them very long, their distal ends mostly curved (Fig. 12). Setae c much shorter than la and ms. Setae h, not shorter than hy. Setae pj and p, comparatively long. Ventral parts (Figs 13): Similar to the preceding species, epimeres located far from each other, sternal apodeme absent. Epimeral setal formula: 3 — 1 — 3 — 3. AII setae well developed, comparatively long. The posterior epimeral borders with some tubercles (Fig. 15). Ventral plate with well developed polygonal pattern reaching to the anal plates. Anogenital setal formula: 6 — 1 — 2 — 3. Anterior and posterior genital setae longer than those in the middle part of the genital plates. Aggenital setae located laterally, much farther from each other than the adanal setae. Setae ad, in paraanal position, all setae in this region long. Lyrifissure iad long. Remarks: See Nasobelba undosa sp. n. Derivatio nominis: I dedicate the new species to my friend, Dr B. Hauser (Muséum d’histoire naturelle, Geneva), the collector of this very interesting material. Nasobelba undosa sp. n. Figs 17-20 Material examined: Brunei: Holotype: SBH-96/9, 1 paratype SBH-69/11. Holotype deposited in MHNG, paratype in HNHM (1667-PO-02). Diagnosis: Rostral apex nasiform, well protruding, rostral tooth large. Anterior part of prodorsum with irregular small tubercles. Peduncle of sensillus very long, curved, head of the sensillus symmetric. Surface of notogaster and the ventral plate with an irregular polygonal pattern. Nine pairs of long and flagellate notogastral setae, p, and p> short, simple. Measurements: Length of body: 242-247 um, width of body: 133-141 wm. ORIBATIDS FROM BRUNEI IV 429 D VX SALE RE Fics 12-16 Nasobelba hauseri gen. n., sp. n.— 12, body in dorsal view; 13, body in ventral view; 14, rostrum in lateral view; 15, genital region; 16, podosoma in lateral view. 430 S. MAHUNKA Fics 17-20 Nasobelba undosa gen. n., sp. n.— 17, body in dorsal view; 18, body in ventral view; 19, genital region; 20, basal part of prodorsum. ORIBATIDS FROM BRUNEI IV 431 Prodorsum: Rostral apex strongly protruding, nasiform. Rostral teeth large, accessory teeth much narrower. Rostral parts with some irregular small tubercles. Tectopedial fields comparatively large, their inner margin observable (Fg. 17). Para- tectopedial field reduced, only a small part visible anteriorly. Basal part of the pro- dorsum distinctly covered by fine granules. Lamellar knob normally developed, some tubercles or short crests present along it (Fig. 20). Interbothridial field weakly developed basally, along the bothridial lobe. Rostral setae geniculate, lamellar and interlamellar setae normal or short, simple. Sensillus with very long peduncle, its head symmetrically lanceolate, narrow. Lateral part of podosoma: Lateroprodorsal pattern absent. Rostral elevation rising from the prodorsal surface. Exobothridial region well sclerotised. Notogaster: Lateral notogastral condyles fused with the median ones, their ori- ginal form still observable (Fig. 17). Typical polygonal pattern reaching to the sejugal margin anteriorly. There are long small fields medially, their borders mostly curved. Nine pairs of long, flagellate notogastral setae, c, shorter than la and ms. Setae h; longer than hy. Setae p, and p> also short. Ventral parts (Fig. 18): Similar to the other species. Epimeres located far from each other, sternal apodeme absent. AIl epimeral setae well developed, comparatively long. Surface of the posterior epimeres covered by fine granules. Along the posterior epimeral borders some small tubercles present, continuing in lines (Fig. 19). Ventral plate with well developed polygonal pattern reaching the anal plates posteriorly. Lateral surface also covered by minute granules. Anogenital setal formula: 6 — 1 — 2 — 3. Anterior and posterior genital setae are longer than those in the middle part of the genital plates. Aggenital setae located much more medially than in the preceding species. Anal setae located in the posterior part of the anal plates. Setae ad, in paraanal position, all setae in this region long. Lyrifissure iad conspicuously long. Derivatio nominis: Named after the form of the notogastral setae. Remarks: The genus Nasobelba gen. n. comprises 6 species, all from the Oriental Region. The four new species are well characterizable and distinguished from the previously known species by the form of the sensillus and the notogastral setae of different lengths, and the composition of the notogastral and ventral pattern or the shape of the prodorsal structures. All six species can be distinguished by using the following key: 1 Rostral apex crown-shaped, with small teeth medially. Notogastral sur- faceswithrirte gularpattern esente ae: coronata Sp. n. = Rostral apex nasiform, without small teeth. Notogastral surface with polygonalipattemssstonesio ate a bet Sete re ig oy 2 D Prodorsal surface with small tubercles medially. Setae h, much shorter (hanicornyd tette has. Bio dae: o Bei An. RR IN ET 3 = Prodorsal surface smooth. Setae h| and c, nearly equal in length ......... + 3 Prodorsal tubercles arranged in a longitudinal column. Notogastral setae rather shorter, /p shorter than the distance between them ....... agathis sp. n. = Prodorsal tubercles irregular. Notogastral setae rather longer, /p longer than the distance between them iaia Aeg SUI win transitoria (Balogh & Mahunka, 1974) comb. n. 432 S. MAHUNKA 4 Lamellar knob opened anteriorly. Posterior part of the interbothridial field and bothridial lobe finely granulated .................. hauseri sp. n. - Lamellar knob normal, closed anteriorly. Posterior part of the inter- bothridial field and bothridial lobe smooth, or the entire basal surface pranulatedt u. na OA LIE RI AN ERRE 5 3 Notogastralsetaesllagellater. M0 2.2093 HE I IE undosa sp. n. - Notogastral setae normal ............ inenodabilis (Hammer, 1980) comb. n. Parasuctobelba quinquecostata sp. n. Figs 21-23 Material examined: Brunei: Holotype: SBH-96/15a, 6 paratypes from the same sample. Holotype and 4 paratypes deposited in MHNG, 2 paratypes in MHNG, 2 paratypes in HNHM (1668-PO-02). Diagnosis: Rostrum wide, conical, without teeth or incisures. Lamellar knob long. Sensillus clavate. Bothridium large and wide. Two pairs of sejugal condyles, their median part connected with each other. Five thick, longitudinal crests present, median one longer than the lateral crest. Ten pairs of short notogastral setae. Posterior border of the epimeral region with digitiform extensions and tubercles. Anogenital setal formula: 5-1-2-3. Measurements: Length of body: 159-170 pm, width of body: 91-100 um. Prodorsum: Rostrum conical, its margin nearly smooth, neither teeth, incisures nor lobes present. There are short transversal crests with 3 sharply pointed tubercles each behind the rostrum (Fig. 22). They are connected laterally, so this border and a longitudinal lateral strong crest form a pair of fields, appearing to be paratectopedial fields. True tectopedial field absent. Lamellar knob elongated. Interbothridial field with a well sclerotised basal part. Rostral setae geniculate, lamellar setae simple, inter- lamellar ones short, curved. Sensillus club-shaped. Lateral part of podosoma: Well developed polygonate lateroprodorsal pattern present. Exobothridial region well sclerotised, partly granulated. Above the acetabula a strong lath running anteriorly from acetabula IV (Fig. 23). Notogaster: Two pairs of sharply pointed, interconnected notogastral condyles present. Two pairs of shorter lateral costulae and one unpaired, long median costula present, they are thick and rise well out from the surface (Fig. 21). The lateral ones with tubercles, some tubercles also existing along the anterolateral margin. Median costula also connected with the lateral ones and bearing a cordyform cavity anteriorly. 10 pairs of short, nearly spiniform notogastral setae present, two posterior ones observable only in lateral view (Fig. 23). Ventral parts: Sternal apodeme absent, epimeres not touching medially, a wide sternal field exist between them. Posterior border of the epimeral region with a pecu- liar structure bearing some digitiform expansions and behind them some large tubercles (Fig. 22). Some large alveoli observable laterally. Epimeral setal formula: 3 — 1 —3 —3, all epimeral setae simple and short. Anogenital setal formula: 5 — 1 — 2 — 3. Aggenital setae located near to the lateral margin of the ventral plate, far from each other. Adanal setae equal in length, setae ad, in paraanal position. Lyrifissures iad long. Remarks: On the basis of the shape of the prodorsum and the peculiar noto- gastral structure, the new species without doubt belongs to the genus Parasuctobelba ORIBATIDS FROM BRUNEI IV 433 Fics 21-23 Parasuctobelba quinquecostata sp. n. — 21, body in dorsal view; 22, body in ventral view: 23, body in lateral view. 434 S. MAHUNKA Hammer, 1977. The new species is distinguishable from the known species by the five long costulae, which are absent in the other species. Derivatio nominis: Named after the structure of the notogaster. CHECK-LIST OF THE HEREWITH PUBLISHED ORIBATID SPECIES OF BRUNEI, WITH COMPLETE LIST OF LOCALITIES This check-list contains the bibliographic references and all data concerning the collecting localities. To make it more effective for ecological and biogeographical pur- poses, the complete locality-list gives the possibility to decode all published localities. Eniochthoniidae Grandjean, 1947 1. Eniochthonius sumatranus (Mahunka, 1989) — Mahunka, 1997: Bru-88/41. Gehypochthoniidae Strenzke, 1963 2. Gehypochthonius xarifae Strenzke, 1963 — Mahunka, 1997: Bru-88/41. Parhypochthoniidae Grandjean, 1932 3. Parhypochthonius asiaticus Mahunka, 1997 — Mahunka, 1997: Bru-88/41. Brachychthoniidae Thor, 1934 4. Sellnickochthonius muara Mahunka, 1995 — Mahunka, 1995: Bru-88/12; Bru-88/24. 5. Sellnickochthonius planus (Chinone, 1974) — Mahunka, 1995: Bru-88/24. Phthiracaridae Perty, 1841 6. Hoplophthiracarus (Plonaphacarus) aculeatus Mahunka, 1995 — Mahunka, 1995: Bru-88/38; Mahunka, 1997: Bru-88/46. 7. Notophthiracarus hauseri Mahunka, 1995 — Mahunka, 1995: Bru-88/21; Mahunka, 2005: SBH-96/12. Synichotritiidae Walker, 1965 8. Sabahtritia lienhardi Mahunka, 1995 — Mahunka, 1995: Bru-88/29; Mahunka, 2001: Bru-88/32. Temburongiidae Mahunka, 1990 9. Temburongia patoi Mahunka, 1990 — Mahunka, 1990 and Mahunka, 1995: Bru-88/21, Bru-88/38, Mahunka, 1997: Bru-88/41, Bru-88/46; Mahunka, 2001: Bru-88/29, Bru-88/38; Mahunka, 2005: SBH-96/11, SBH-96/12. Epilohmanniidae Oudemans, 1923 10. Epilohmannia nortoni Mahunka, 1997 — Mahunka, 1997: Bru-88/29, Bru-88/41, Bru-88/46. 11. Epilohmannoides esulcatus Ohkubo, 1979 — Mahunka, 1997: Bru-88/29, Bru-88/41, Bru-88/46. Lohmanniidae Berlese, 1916 12. Papillacarus lienhardi Mahunka, 1997 — Mahunka, 1997: Bru-88/29. Hermanniellidae Grandjean, 1934 13. Bruneiella sultan Mahunka, 1997 — Mahunka, 1997: Bru-88/41, Bru-88/46. Microtegeidae Balogh, 1961 14. Microtegeus sabahnus Mahunka, 1987 — Mahunka, 1997: Bru-88/21, Bru-88/41. 15. Suctotegeus tumescitus Mahunka, 1987 — Mahunka, 1997: Bru-88/41, Mahunka, 2005: SBH-96/12. Eremaeozetidae Balogh, 1972 16. Eremaeozetes maculosus Mahunka, 1995 — Mahunka, 1995: Bru-88/24. Microzetidae Grandjean, 1936 17. Anakingia borneensis Mahunka, 1997 — Mahunka, 1997: Bru-88/41. 18. Teraja asymmetrica Mahunka, 2001 — Mahunka, 2001: Bru-88/38. 19. Teraja sungai Mahunka, 1997 — Mahunka, 1997: Bru-88/41, Bru-88/46; Mahunka, 2001: Bru-88/35; Mahunka, 2005: SBH-96/9. 20. Teraja tuberculata (Mahunka, 1987) — Mahunka, 2001: Bru-88/35. 21. Teraja wongi Mahunka, 1995 — Mahunka, 1995: Bru-88/32; Mahunka, 1997: Bru-88/29; Mahunka, 2001: Bru-88/32. ORIBATIDS FROM BRUNEI IV 435 Damaeolidae Grandjean, 1965 22. Fosseremus laciniatus (Berlese, 1905) — Mahunka, 2001: Bru-88/12; Mahunka, 2005: SBH-96/11. Eremobelbidae Balogh, 1961 23. Eremobelba porcella Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/24. Peloppiidae Balogh, 1943 24. Austroceratoppia serapi Mahunka, 1996 — Mahunka, 2001: Bru-88/29. Carabodidae C. L. Koch, 1837 25. Congocepheus orientalis Mahunka, 1987 — Mahunka, 1997: Bru-88/41. 26. Gymnobodes semengok Mahunka, 1996 — Mahunka, 2001: Bru-88/35. 27. Hardybodes flabellatus Mahunka, 1995 — Mahunka, 1995: Bru-88/21, Bru-88/32, Bru-88/38; Mahunka, 1997: Bru-88/41. 28. Hardybodes penicillatus Mahunka, 1995 — Mahunka, 1995: Bru-88/35; Mahunka, 2001: Bru-88/24; Mahunka, 2005: SBH-96/12. 29. Pasocepheus eremaeozetoides Mahunka, 1995 — Mahunka, 1995: Bru-88/32. Tectocepheidae Grandjean, 1954 30. Tegeozetes tunicatus Berlese, 1913 — Mahunka, 2001: Bru-88/32, Bru-88/38. Otocepheidae Balogh, 1961 31. Dolicheremaeus andulauensis Mahunka, 1997 — Mahunka, 1997: Bru-88/41, Bru-88/46; Mahunka, 2001: Bru-88/24. 32. Dolicheremaeus bruneiensis Aoki, 1967 — Aoki, 1967: Brunei City, 21.1.1962. 33. Dolicheremaeus furcillatus Mahunka, 1997 — Mahunka, 1997: Bru-88/21, Bru-88/46. 34. Dolicheremaeus wallacei Mahunka, 1997 — Mahunka, 1997: Bru-88/29, Bru-88/46. 35. Otocepheus durian Mahunka, 1997 — Mahunka, 1997: Bru-88/46. Dampfiellidae Balogh, 1961 36. Dampfiella zellwegeri Mahunka, 1997 — Mahunka, 1997: Bru-88/21, Bru-88/46; Mahunka, 2005: SBH-96/11. Luxtoniidae Mahunka, 2001 37. Luxtonia hauseri Mahunka, 2001 — Mahunka, 2001: Bru-88/35. Oppiidae Grandjean, 1951 38. Arcoppia teraja Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/32. 39. Corynoppia andulau Mahunka, 2001 — Mahunka, 2001: Bru-88/21. 40. Graptoppia sundensis (Hammer, 1980) — Mahunka, 2001: Bru-88/21, Bru-88/35. 41. Karenella bruneiana Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/46. 42. Oppiella nova (Oudemans, 1902) — Mahunka, 2001: Bru-88/24, Bru-88/32, Bru-88/35. 43. Ptiloppia lienhardi Mahunka, 2001— Mahunka, 2001: Bru-88/21, Bru-88/32, Bru-88/35. 44. Pulchroppia burckhardti Mahunka, 1987 — Mahunka, 2001: Bru-88/35. 45. Senectoppia kerangas Mahunka, 2001 — Mahunka, 2001: Bru-88/32. Rhynchoribatidae Balogh, 1961 46. Oxyamerus hauserorum Mahunka, 1987 — Mahunka, 2005: SBH-96/11, SBH-96/12. 47. Suctoribates foliatus Mahunka, 1997 — Mahunka, 1997: Bru-88/29, Bru-88/41; Mahunka, 2005: SBH-96/11, SBH-96/12, SBH-96/15a. Suctobelbidae Jacot, 1938 48. Bruneibelba separata Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/35. 49. Bruneibelba tuberosa Mahunka, 2001 — Mahunka, 2001: Bru-88/35. 50. Coartobelba pauper Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/29, Bru-88/41, Bru-88/46. 51. Condylobelba agathis Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/29, Bru-88/35, Bru-88/46. 52. Condylobelba bruneiensis Mahunka, 2001 — Mahunka, 2001: Bru-88/21, Bru-88/24, Bru-88/29, Bru-88/35, Bru-88/41, Bru-88/46; Mahunka, 2005: SBH-96/12. 436 S. MAHUNKA 53. Condylobelba sculpturata Mahunka, 2001 — Mahunka, 2001: Bru-88/35. 54. Nasobelba agathis Mahunka, 2005 — Mahunka, 2005: SBH-96/12. 55. Nasobelba coronata Mahunka, 2005 — Mahunka, 2005: SBH-96/11, SBH-96/15a. 56. Nasobelba hauseri Mahunka, 2005 — Mahunka, 2005: SBH-96/11. 57. Nasobelba undosa Mahunka, 2005 — Mahunka, 2005: SBH-96/9, SBH-96/11. 58. Parasuctobelba quinquecostata Mahunka, 2005 — Mahunka, 2005: SBH-96/15a. 59. Suctobelbella subcomplexa (Balogh & Mahunka, 1968) — Mahunka, 2001: Bru-88/21, Bru-88/41. 60. Suctobelbella variosetosa (Hammer, 1961) — Mahunka, 2001: Bru-88/35, Bru-88/38, Bru-88/46. Haplozetidae Grandjean, 1936 61. Bolkiah hauseri Mahunka, 1997 — Mahunka, 1997: Bru-88/41; Bru-88/46. 62. Borneozetes lanceolatus Mahunka, 1997 — Mahunka, 1997: Bru-88/29. Oribatellidae Jacot, 1925 63. Lamellobates orientalis Csiszar, 1961 — Mahunka, 2001: Bru-88/35. Galumnidae Jacot, 1925 64. Notogalumna praetiosa Sellnick, 1959 — Mahunka, 2001: Bru-88/24. LIST OF LOCALITIES FOR ALL PUBLISHED RECORDS (EXPEDITIONS OF 1988 AND 1996) 1988 Bru-88/12: Brunei (Brunei-Muara District): près du pont sur le ruisseau “Sungai Lubang Barus” ~ sur la route venant de Tutong, à 33 km de Bandar Seri Begawan, prélèvement de sol dans les angles formés par les contreforts de deux grands arbres proches des habitations, env. 20 m; 16.X1.1988; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). Bru-88/21: Brunei (Belait District): “Andulau Forest (“Mixed dipterocarp forest”), K-7 (“Kompartment 7”), prélèvement de sol dans les angles formés par les contreforts de grands arbres, 50 m; 19.X1.1988; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). Bru-88/24: Brunei (Brunei-Muara District): “Berakas Forest Reserve” au nord de Bandar Seri Begawan sur la route, à 19,5 km de Muara (= à 102,5 km de Kuala Belait), forêt “Kerangas” (= “Tropical heath forest”), prélèvement de sol au pied de Casuarina nobilis Whitmore (Casuarinaceae), 30 m; 20.XI.1988; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). Bru-88/29: Brunei (Belait District): Sungai Liang, “Arboretum Forest Reserve”, forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contre- forts de deux arbres appelés “Nyatho”, 90 m; 21.X1.1988; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). Bru-88/32: Brunei (Belait District): “Labi Hills Forest Reserve”, “Teraja”, à 42 km au sud de Sungai Liang (= 12 km au Sud de Labi), environs de “Rumah Panjang” (= Longhouse du Kampong Teraja), forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contreforts d’un très grand arbre, 40 m; 22.XI.1988; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). Bru-88/34: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secon- daire qui bifurque, à 32 km de Kuala Belait, vers le sud, forêt “Kerangas” (=“Tropical heath forest”) formée presque exclusivement par Agathis dammara (Lambert) L. G. Rich. (Araucariaceae), sur et sous écorces, 10 m; 23.X1.1988; leg. B. Hauser. Bru-88/35: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secon- daire qui bifurque, à 32 km de Kuala Belait, vers le sud, forêt “Kerangas” (=“Tropical heath forest”) formée presque exclusivement par Agathis dammara (Lambert) L. G. Rich. (Araucariaceae), prélèvement de sol au pied de Agathis dammara, 10 m; 23.XI. 1988; leg. B. Hauser — (extraction par appareil Berlese à Hong Kong). Bru-88/38: Brunei (Temburong District): “Peradayan Forest Reserve” (= “Bukit Patoi”), à 14,5 km de Bangar (= 2,5 km de Labu), forêt primaire (“Mixed dipterocarp forest”), prélè- vement de sol dans les angles formés par les contreforts de grands arbres morts, 80 m; 24.X1.1988; leg. B. Hauser — (extraction par appareil Berlese à Hong Kong). ORIBATIDS FROM BRUNEI IV 437 Bru-88/41: Brunei (Belait District): Sungai Liang, “Arboretum Forest Reserve”, forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contre- forts d’arbres appelés “Kempas” (= Koompassia malaccensis Maing. & Benth. [Fabaceae]), 20 m; 25.XI.1988; leg. B. Hauser — (extraction par appareil Berlese à Hong Kong). Bru-88/46: Brunei (Belait District): “Andulau Forest Reserve”, à 3,5 km au sud de Sungai Liang (= à 39,5 km de Labi), forêt primaire (“Mixed dipterocarp forest”), K-8 (“Kompartment 8”), prélèvement de sol dans les angles formés par les contreforts d’un grand arbre, 70 m; 26.X1.1988; leg. B. Hauser — (extraction par appareil Berlese à Hong Kong). 1996 SBH-96/9: Brunei (Belait Dstrict): Sungai Liang, “Arboretum Forest Reserve”, forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contre- forts de Alstonia angustiloba Miq. (Symplocaceae), 20 m; 29.XI.1996; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). SBH-96/11: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secondaire qui bifurque, à 32 km de Kuala Belait, vers le S; forêt “Kerangas” (= “Tropical heath forest”) formée presque exclusivement par Agathis dammara (Lambert) L. G. Rich. (Araucariaceae), “Genetic Resources Protection Area”, prélèvement de sol au pied d’un gros A. dammara, 10 m; 30.XI.1996; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). SBH-96/12: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secondaire qui bifurque, à 32 km de Kuala Belait, vers le S; forêt “Kerangas” (= “Tropical heath forest”) formée presque exclusivement par Agathis dammara (Lambert) L. G. Rich. (Araucariaceae) “Genetic Resources Protection Area”, prélèvement de sol au pied de A. dammara, 10 m; 30.X1.1996; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). SBH-96/15a: Brunei (Belait District): “Labi Hills Forest Reserve”, “Teraja”, à 42 km au S de Sungai Liang (= 12 km au Sud de Labi), environs de “Rumah Panjang” (= Longhouse du Kampong Teraja), forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contreforts de deux très grands arbres, 20 m; 2.XI1.1996; leg. B. Hauser — (extraction par appareil Berlese à Bandar Seri Begawan, Brunei). ACKNOWLEDGEMENTS First and foremost I wish to thank Dr B. Hauser, the collector of this very rich material. Our hearty thanks are due also to the curator of the collection, Dr P. Schwendinger, and the Director of the Museum, Dr V. Mahnert, for the opportunity to study this collection in Geneva. For corrections to the manuscript I extend my sincere thanks to Dr M. Luxton (National Museum of Wales, Cardiff). Special thanks are due to Dr B. Hauser for his editorial work while preparing the manuscript, for the preparation of the faunistic list of Brunei and all the technical assistance with the Geneva collection. For the linguistic revision I wish to thank Dr L. Zombori (Budapest). This project was partly sponsered by the Hungarian Scientific Research Fund (OTKA 38319 and 45889). REFERENCES AOKI, J. 1967. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata) II. Subfamily Tetracondylinae. Bulletin of the National Science Museum, Tokyo 10: 297-359. BALOGH, J. 1970. New oribatids (Acari) from Ceylon. The scientific results of the Hungarian soil zoological expeditions. Opuscula Zoologica Budapest 10: 33-67. 438 S. MAHUNKA BALOGH, J. & BALOGH, P. 1992. The oribatid mites genera of the world. Hungarian Natural History Museum, Budapest, I: 263 pp., II: 375 pp. HAMMER, M. 1980. Investigations on the oribatid fauna of Java. Biologiske Skrifter. Kongelige Danske Videnskabernes Selskab 22(9): 1-79 + pls. 1-47. MAHUNKA, S. 1990. A survey of the superfamily Euphthiracaroidea Jacot, 1930 (Acari: Oribatida). Folia Entomologica Hungarica 51: 37-80. MAHUNKA, S. 1995. Oribatids from Brunei I (Acari: Oribatida). New and interesting mites from the Geneva Museum LXXV. Revue suisse de Zoologie 102: 913-942. MAHUNKA, S. 1997. Oribatids from Brunei II (Acari: Oribatida). (Acarologica Genavensia LXXXII). Revue suisse de Zoologie 104: 661-700. MAHUNKA, S. 2001. Oribatids from Brunei III (Acari: Oribatida). (Acarologica Genavensia XCT). Revue suisse de Zoologie 108: 317-349. MAHUNKA, S. 2005. Oribatids from Brunei IV (Acari: Oribatida). (Acarologica Genavensia CVI). Revue suisse de Zoologie 112: 421-438. MAHUNKA, S. & MAHUNKA-PAPP, L. 2001. Oribatids from Switzerland V. (Acari: Oribatida: Suctobelbidae 2). (Acarologica Genavensia XCVII). Revue suisse de Zoologie 108: 355-385. REVUE SUISSE DE ZOOLOGIE 112 (2): 439-455; juin 2005 Trojanella serbica gen. n., sp. n., a remarkable new troglobitic travunioid (Opiliones, Laniatores, Travunioidea) Ivo M. KARAMAN Department of Biology and Ecology, Faculty of Sciences, Trg D. Obradoviéa 2, 21000 Novi Sad, Serbia and Montenegro. E-mail: karaman@ib.ns.ac.yu Trojanella serbica gen. n., sp. n., a remarkable new troglobitic travuni- oid (Opiliones, Laniatores, Travunioidea). - A new species of travunioid from Serbia, Mt. Stara Planina, is described and a new genus is established. The new species exhibits some characters and combination of characters not known from any other species. It cannot be placed at this time in any of the described families of the superfamily Travunioidea. Some unknown details of male morphology in Abasola hofferi (Travuniidae) are presented. Keywords: Travunioidea - Mt. Stara Planina - troglobite - Travuniidae - Abasola hofferi - penis structure. INTRODUCTION Thanks to intensive biospeleological research done by the Belgrade Institute for Protection of Nature team under Dragan Pavicevic’s leadership, an unusual new species of troglobitic opilionid has been discovered. Specimens of the new species were found in a pothole in Mt. Stara Planina in south-east Serbia, near the Bulgarian border. The specimens were collected from the 60 m deep pothole, between stones of the breakdown covering its bottom. The new species belongs to the suborder Laniatores and the superfamily Travunioidea which is widespread in temperate zones of both hemispheres. In the southern hemisphere this superfamily has a typical Gondwanan distribution, whereas in the northern hemisphere it is represented by a small number of genera and species found locally in some parts of Europe, North America, Japan, and Korea. In all areas, representatives of this superfamily mostly occur as rare relict elements, often as troglo- bionts, which indicates the great age of the group. Characteristics of the new species and the combination of characteristics make the species unique in the group. So it was not difficult to establish that it belongs to a new genus. On the basis of the criteria of significance of certain characters and of the current classification of the Travunioidea into families and subfamilies, the position of this new genus is rather disputable. By its sternum structure, form of spiracle and the type of claws structure, this genus seems closely related to the family Triaenonychidae, which representatives are predominantly distributed in the southern hemisphere (in the Manuscript accepted 01.12.2004 440 I. M. KARAMAN northern hemisphere, given the current composition, this family is represented by 12 genera and 23 species in North America, Japan and Korea). On the other hand by the ovipositor morphology and midgut anatomy this new genus is closely related to the north hemispheric families Cladonychidae and Travuniidae. Systematic of the superfamily Travunioidea is somewhat confusing with five families and eight subfamilies, in some cases uncritically erected. A comprehensive revision of the whole superfamily Travunioidea is needed. Recent confusing syste- matic of the superfamily, the extremely specific penis structure and combination of relevant characters of Trojanella serbica gen. sp. n., made me decide to put it in Travunioidea incertae sedis. Male specimens of Abasola hofferi Silhavy, 1936 (family Travuniidae) were first collected in the cave Pokljuka Gornja (type locality) in Montenegro (Silhavy, 1936). The material was used for comparison and for learning more about the morpho- logy of this little known species. Details of penis structure in A. hofferi question the validity of the current composition of the Travuniidae. RESULTS TRAVUNIOIDEA incertae sedis Trojanella gen. n. The genus of small long legged Travunioidea is defined by morphological characters of the genitalia. Penis with elongated truncus and movable articulated, flattened, bifurcate glans. Truncus of the penis terminally widened; musculature settled in the terminally widened part of the truncus and in the glans. Glans wide and flattened, terminally bifurcated, laterally with 2 strong teeth-like protrusions. Ovipositor with 4 lobes terminally; dorsal and ventral one as well as the ovipositor body covered with sparce denticles: lateral lobes smooth, each bearing ventral and dorsal rows of few spines. Spiracle not concealed, semicircular in shape. Sternum narrow, basally widened with setae; subapically trapezoidally expanded. Claws II and IV with a pair of elongated, flattened branches parallel to widely flattened medial prong. Shape of these claws similar to peltonychium in Travuniidae. Juveniles with claws III and IV of peltonychium type. Type species. Trojanella serbica sp. n. Etymology: Trojan — in south Slavic mythology the demon or the god of night and darkness. According to legends, most often lives in ruins of towns and fortresses. The name is diminutive of femine gender. Trojanella serbica sp. n. Figs 1-4, SA, 6-7 Material examined: From an unnamed pothole, at the locality of Vladikina Ploëa in the Visoëica River gorge, near the village of Rsovci, on Mt. Stara Planina, Serbia (Serbia & Montenegro), 900 m asl (UTM - FN48), 2.10.2002. leg. I.Karaman: 1 & holotype, 5 d paratypes, 8 2 paratypes and 2 juv. (+1 adult specimen used for molecular analyses); ibid., 30.05.2002, leg. S. Ognjenovié: 2 juv.; ibid., 2.10.2002-2.06. 2003, from traps, leg. S. Ognjen- ovic: 3 d paratypes and 3 9 paratypes. 441 TROJANELLA SERBICA "MOA [RIOT :(wur Eg’ | ‘odAqered af eu) u ‘ds € "u ‘98 D21q19$ DJJOUDIOUL © | “OWA = NA se \ WPS ET TN N) SETT \ 442 I. M. KARAMAN Holotype 3 (Inv.No 1324),7 5,10 © paratypes and 4 juv. are deposited in the author’s collection at the Department of Biology and Ecology — Novi Sad (Serbia & Montenegro), 1 4 and 1 ® paratypes in the Muséum d’histoire naturelle, Geneva, Switzerland. DESCRIPTION Male: Blind troglobitic species. Male holotype 1.7 mm (male paratypes 1.58- 1.83 mm) long, measured from frontal edge of dorsal scutum. Body uniformly yellowish amber in colour. Dorsum densely granular with sparse rows of setae and pronounced lateral grooves in its median area (Figs 2A-B). Grooves starting with two lateral creases on both sides of carapace region on same level as ocular tubercle and above ozopores. Creases joined in a depression at posterior end of dorsal scutum. A pair of grooves anteriolateral, near and parallel to frontal edge of dorsal scutum. Conical ocular tubercle distant from anterior edge of dorsal scutum for almost twice its length. Coxal lobes II anteriomesally with strong conical apophyses (male secondary sexual character) ventrally directed (Fig. 2C), sparsely setose. Coxal lobes III anterio- mesally with a pair of small tubercles (on both sides of the sternal subapical expansion). Sternum as in Fig. 2C, basally widened, with two pairs of setae (short and long ones); sternum slightly trapezoidally expanded at level of junction between coxae I and III. Chelicerae (Figs 3A-B). Basal segment gradually widened distally, dorsal surface terminally with two tubercles bearing setae, ventrally with few setae; second segment anteromesally with two spinelike tubercles bearing subapical setae, anteriorly with several tubercles bearing apical setae and with concentration of terminal setae. Pedipalps (Figs 3C-D) in a form of catching basket, armed with strong elongate spinelike tubercles, each bearing a strong elongate seta subapically; coxae ventrally with a low tubercle bearing small setae and a strong spinelike tubercle proximally; trochanter ventrally with 1 strong spinelike tubercle, dorsally with 1 tubercle bearing small setae; femur strong, ventrally with a row of 6 spinelike tubercles (proximal 3 stronger than others), medio-proximally with 2 spinelike tubercles, dorsally with 2 low spinelike tubercles; patella medially with 2 spinelike tubercles, laterally with 1 spine- like tubercle; tarsus with 4 strong spinelike tubercles on both sides (laterally and medially), terminally with 2 strong setae. Claw elongated. All articles bearing sparce hairs and setae, on dorsal sides usually placed on low tubercles. Sparce hairs on tarsal segment in longitudinal rows. Legs elongated, segments cylindrical; coxae ventrally with a row of tubercles bearing setae subapically; each femur basally with a false articulation (Fig. 7B). Calcanei I-IV elongated. Ratio of calcaneus/astragalus of metatarsi I-IV: 0.78/0.91/1.08/1.56. Tarsal formula: I-11; I-24; III-3; IV-3. Tarsal claws of first and second legs simple (Fig. 4A), first claw somewhat more sickle-shaped than second one; claws of third and fourth legs (Figs 4C-D) with lateral branches almost parallel to longer median prong, median prong and basal parts of branches dorso-ventrally flattened. TROJANELLA SERBICA Se) Bn eh ee vos SK .2 255857 eS oO ein ee o Sas CESSO ee ER COMTE ORE Mowe Es ETES S BES S S = = Sos à Bal oO Le E eos 055052550 fumetti n > > ar en © Gea . SRH: e Sn = Sea O S cQESS < CO fe) D ES EE Sour © Im VA er is 5.828 SEDE sd "Mora Jouur ‘dyed -Ipod ‘q ‘MIA Jono ‘dyedipod ‘9 :MOIA J9]NO ‘BIII -I[949 “gq :MOIA JOUUI ‘RI99I 949 ‘V :(W OLI ‘odAjesred ogewi) "u ‘ds “u ‘u98 919 -198 pyjouploaL € OM 445 TROJANELLA SERBICA urge AI MEO ‘Œ :2[IU9A -nf‘u ‘ds “u ‘u93 D91q19$ ‘I ‘AI MET. ‘I ‘III Melo ‘I MEI CV (uu gf ‘ad -eied jew) "u ‘ds “u ‘uas p21q -195 pjjouploA] LAC 446 I. M. KARAMAN FIG. 5 Trojanella serbica gen. n., sp. n. (holotype): A, penis, ventral view. Abasola hofferi (male, 1.23 mm): B, penis (contracted), lateral view. A. hofferi (male, 1.43 mm): C, terminal part of penis, lateral view; D, penis terminal part, ventral view. Scale line: A, B = 5007; C, D = 1007. Measurements of legs (in mm): Tr Fe Pa Ti Mt Ta Di Leg I 0.31 2.16 0.5 1293 1.82 2.38 9.1 Leg II 0.36 3.44 0.7 3.22 3.19 4.9 15.81 Leg III 0.42 2.58 0.56 1.96 2.66 2.24 10.42 Leg IV 0.39 322 0.56 24 3.16 322 12.95 re TROJANELLA SERBICA 447 Penis (Fig. SA; Figs 6A-D). Truncus elongated with wide basis, from the basal narrowing gradually widened toward the apex; musculature concentrated in distal part of truncus and in glans. Glans wide and flattened, terminally bifurcated, laterally with a pair of divergent tooth-like protrusions, ventrally with 2 pairs of short spines. Seminal opening situated in between branches of glans bifurcation, close to its ventral side (Fig. 6C, see arrow). Glans musculature composed of a median trapezoidal mus- cle extending through entire length of glans to apical parts of bifurcation (on inner side of each branch), and by smaller muscle bands situated on outer sides of bifurcation branches. Inner structure of glans complex, with 2 groups of cells (possibly glandular structures) lateral to trapezoidal muscle. This group of cells extended from terminal parts of truncus to level of lateral teeth-like protrusions. If these structures really represent glands, their canals could open on the tips of the bifurcation branches. Unfortunately the tiny membranous structures at the tip of the bifurcations as seen at high magnification under LM and SEM (Fig. 6D) does not allow me to reach a conclusion about the presence of glandular openings on them.* Female: Similar to male in somatic characters but differs in lacking the medial conical apophyses on coxal lobes II. Genital operculum wider than in male (Fig. 2D). Body length (from frontal edge of dorsum) 1.54-1.91 mm. Ovipositor cylindrical, with 4 terminal lobes (Fig. 7A); ovipositor body and outer sides of dorsal and ventral lobes covered with sparse denticles; lateral lobes smooth, on the ventral side bearing 3 small spines, on the dorsal side 4-5 small spines; eight spherical receptacles in radial arrangement. Intestinal complex. First pair of intestinal diverticula ramified in ramus anterior and ramus coxalis. Second pair of intestinal diverticula ramified in ramus transversalis and ramus longitudinalis. Third pair of intestinal diverticula with two rami - ramus medianus and ramus lateralis. Ramus medianus remarkably shorter than ramus lateralis; ramus lateralis in distal half rectangularly ventrally oriented. First half of the ramus lateralis on the distal end (at the place where it’s rectangularly changing orien- tation to the ventral side) is heavily widened. TRAVUNIIDAE Absolon & Kratochvil, 1932 Abasola hofferi Silhavy, 1936 Figs 5B-D, 8,9 Material examined: From Pokljuka Gornja Cave (type locality), near the village of KneZlaz, Krivoëije, Montenegro (Serbia & Montenegro), 1.07.1997 leg. I. Karaman: 1 d, 1 juv.; ibid., 9.07.1999, leg. S. Ognjenovié: 1 ¢; Jama Duboka pothole, ibid., 15.09.2003, leg. S. Ognjenovic: 1 juv. Specimens are deposited in the author’s collection at the Department of Biology and Ecology — Novi Sad (Serbia & Montenegro). Dorsal scutum anteriorly with visible remnants of 2 tubercles (Fig. 8A). The anterior one, more pronounced and coarsely ornamented, is situated on the frontal edge of the dorsal scutum and probably represents the remnant of some larger protrusion. The other one is noticeably smaller and located more posteriorly and thus is pre- sumably the remnant of an ocular tubercle. * Histological studies of the inner structure of the glans are undertaken by Prof. J. Martens and will be published in due time. 448 I. M. KARAMAN 3 5 D SO n een © Gey = SOIR SIE IDE ESS S . yo SISERR o .= e E Sas Ame > ” 089 - RMC UE V0O0£tas® . 3 nem mi TS > DES ada >> sE S_LE.O ave ES 29 pad > = eo Egr fto =a So. DE Es T 3 SLA a) SS {SI ofS fog NO Ss a SS Bars ds So © Paws 449 TROJANELLA SERBICA "smole AQ payed -Ipur UONE|nonIe 9S[EJ YM INUIJ ‘q :(Wu O/T ‘odAqered sfeun) "u ‘ds “u ‘uas 0919 -198 ‘J, ‘10JISOdIAO ‘V ‘(um 851 ‘odAqered ayeuray) ‘u ‘ds “u ‘u98 po -1Q19$ DJJ2UDIO4I Ll | 450 Wirt "MIA jeısye] ‘ouues ‘a :MOIA [PTU9A ‘(P91981JuU09) stuod jo jied [BUTI] ‘7 MOIA penuoa ‘Apoq Jo ed 1orojue ‘4 MOIA [BSIOP ‘LunJN9S [ESIOP JO wed our ‘y ‘(WU ET | ‘Opew) 110/foy pjospqy 8 Old TROJANELLA SERBICA 451 REGLE D ons > 85805 SCESO O ON a DL Saar a SAS Soa © © Sion 2£432 452 I. M. KARAMAN Penis structure (Figs 5B-D, 8C-D); truncus stout, basally and terminally narrowed, continuous with glans without clear transition; subterminally with two pairs of lateral membranous wing-like lamina, basal pair bigger and dorsally directed (Fig. SC); ventral side of glans with a small subterminal stylus; 2 pairs of small spines subterminally, bigger one ventrolaterally above wing-like lamina, smaller one ventrally on both sides of stylus; a muscle in basal 2/3 of truncus. DISCUSSION With some details of their structures (primarily of the penis), the species Trojanella serbica gen. n., sp. n. and Abasola hofferi additionally complicate the already rather confusing classification of Travunioidea. It is difficult to single out any differentiating character used so far that could clearly define a particular family or sub- family of Travunioidea. Claw structure seems not so useful in family and subfamily classification of the Travunioidea, as it was summarised by Hunt & Hickman (1993) for Triaenonychidae. Too much attention was paid to this character and has sometimes caused absurd results. An example are species Paranonychus brunneus (Banks, 1911) from the north-west of North America and Mutsunonychus fuscus Suzuki, 1976 from . the north-east of Japan, which are obviously congeneric. Based solely on claw struc- ture, they were classified into two different subfamilies of the Triaenonychidae: Paranonychinae Briggs, 1971 and Kaolinonychinae Suzuki, 1975, respectively (Briggs, 1971; Suzuki, 1976). Dorsoventrally flattened claws II and IV (with central prong and side branches in the same plane) in 7rojanella serbica gen. n., sp. n., although of the triaenonychid type, seem closer to the peltonychium type of claws present in A. hofferi (i.e. Dinaric travunioids) than to the claws of Triaenonychidae from the southern hemisphere (typical triaenonychid type). Claw structure cannot be a defining character for the family Travuniidae either, because the obvious great differences in penis structures of Dinaric Travuniidae and the genus Peltonychia do not confirm their monophyletic origin. It can be assumed that the closeness by this character is a consequence of independent occurrence of the same, probably convergent neotenic characteristic. The current composition of the Travuniidae is also largely a result of a, in my opinion flawed, perception of Europe as a region with more or less homogeneous fauna. The genus Trojanella gen. n., Dinaric travuniids and Balkan Cyphophthalmi (as representatives of the old relictual fauna) suggest that a part of the Balkan fauna must have had a long and significant period of autonomous and independent development, in an isolated region (which surely must have had its own dynamics). The sternum structure seems still useful in the classification of Travunioidea. Based on this character Trojanella gen. n. is close to Triaenonychidae. However, among Dinaric travuniids there are remarkable differences, from a wedge-shaped ster- num in Dinaria vjetrenicae Hadzi, 1932 (Hadzi, 1932: fig. 6) to a wider sternum type with subterminal expansion in A. hofferi, and an almost Triaenobuninae type sternum in Abasola borisi Hadzi, 1973 (Hadzi, 1973: fig.15c). The spiracle structure may be a character of importance for classifying Travunioidea. There are significant differences in spiracle structures among species discussed in this paper. Trojanella serbica gen. n., sp. n. has unconcealed spiracles, TROJANELLA SERBICA 453 semicircular in shape, which are present also in south hemisphere Triaenonychinae. In Abasola hofferi the spiracles are small, rounded, exposed on a tubercle-like protrusion and laterally directed. This type of spiracle (exposed) is also present in some north hemisphere Travunioidea (Dinaric travuniids; Paranonychinae Briggs, 1971; Kaolinonychinae Suzuki, 1975; Nippononychinae Suzuki, 1975 and Yuria pulcra Suzuki, 1964). The concealed type of spiracles is present in south hemispheric Triaenobuninae and some north hemispheric Travunioidea, as well. The penis structure and musculature are no doubt very important in the classi- fication of this group. Yet, it seems that they have not been used consistently when defining certain taxonomic categories. One of the main differentiating characters of the family Cladonychiidae (Martens, 1978, 1986) is the basal position of penis muscula- ture. However, according to HadZi’s description and drawings (Hadzi, 1935), the type species of the family, Cladonychium corii HadZi, 1935 [synonymized by Briggs (1969) with Erebomaster acanthina (Crosby & Bishop, 1924)] has the truncus with a fully developed muscle. Despite this significant deviation, the current composition of the family Cladonychidae, based on the penis structure, maybe is not questioned”. This discrepancy may be assigned to something which appears as a general tendency with- in the Travunioidea (and wider, within Opiliones) towards simplification of the terminal part of the penis (both structurally and functionally, causing reduction of the muscula- ture) and prolongation of the truncus, followed by glans reduction (with subsequent displacement i.e. concentration of the remaining musculature in the widest, basal part of the truncus). This tendency (and gradual transition), ranging from the complex penis structure in Australian Triaenonychidae to the simple penis in species belonging to the genus Peltonychia, can manifest itself in various stages of reduction and changes in closely related taxa, which is considered to be the case with Cladonychidae, as well. It is also possible that cases of convergent similarity with respect to the penis structure may occur among phylogenetically remote taxa. Trojanella serbica gen. sp. n. deviates from this tendency because it is obvious that the terminal part of the penis (glans) is functionally very complex and highly apomorphic. The specific structure of the glans may be derived from the complex structure of the terminal part of the penis in Australian and South African triaenonychid taxa shown by Martens (1986) by complete reduction of the dorsal and dorsolateral plates and by merging the stylus with the ventral plate (i.e. “Sensillentràger”). With respect to the penis structure, this species represents an unique and isolated phylogenetic line in the superfamily. It is interesting that the great heterogeneity of penis structures in Travunioidea is almost invariably followed by the presence of subterminal, laterally diverging struc- tures (obviously functionally very important) of different structural types, from dorso- lateral plates in south hemispheric genera to diverging thorns of the glans in the species of the genus Peltonichya. In Trojanella gen. n. they are present in the form of diverg- ing teeth on the glans. Penis structure in A. hofferi may also be derived from the complex types of Australian and South African triaenonychids. The basal pair of membranous wing-like lateral lamina may represent dorsolateral plates of the terminal part of the truncus, But the ovipositor structure in Cladonychidae also shows remarkable differences. A compre- hensive revision of this family is needed. 454 I. M. KARAMAN which most triaenonychids have. The part of the penis distally to these structures may be the glans, i.e. the ventral plate and the stylus that have become a glans. The struc- ture of the penis in D. vjetrenicae may be derived from the penis structure of A. hof- feri, the glans of which is greatly reduced and the dorsolateral plates strongly developed and with ventral orientation. The structure of the intestinal complex could give useful data for phylogenetic consideration, however there are available data only for a few species (Dumitrescu 1974, 1976). Dumitrescu (1976) noticed differences in midgut anatomy between southern hemispheric and northern hemispheric Travunioidea. After him southern hemispheric Travunioidea posses the ramus exterior of the third pair of intestinal diverticula in comparison with species on the northern hemisphere. In 7rojanella ser- bica gen. n., sp. n. absence of the ramus exterior is evident, but a widened median part of ramus lateralis could be interpreted as shortened and reduced “true ramus lateralis”. In such interpretation the part distally of that widening could represent ramus exterior. In that case the midgut anatomy of 7. serbica gen. n., sp. n., could represent a transi- tion form between “southern” and “northern” type of the intestinal complex. To make final conclusions about taxonomic value and usefulness of the intestinal structure in solving problems in Travunioidea, details of this structure in higher number of species are needed. Unfortunately I was not able to investigate completely the intestinal structure of A. hofferi. Significant differences in the penis structure of Abasola hofferi and Dinaria vjetrenicae suggest that a diverse travuniid fauna (at least two genera and obviously more than two species) exists in one relatively small Dinaric region (south-eastern Herzegovina, southern Dalmatia and south-western Montenegro). So far six species in three genera — Travunia, Abasola and Dinaria — have been described, at times extremely superficially and uncritically (without any clear differentiating characteris- tics given for the genera). In at least two cases the descriptions are based on juvenile specimens. The confusing situation is additionally complicated by imprecise location data for some caves where certain species were recorded, and by unknown destiny of some specimens. In order to get a clearer picture of this undoubtedly distinct phyletic line of Travunioidea, it is necessary to obtain detailed information on the morphology of all the species described so far. Unfortunately this will be difficult in some cases. The terrain where the Durovié peéina cave [type locality of Abasola troglodytes (Roewer,1915)] is situated has been devastated and made part of an airport complex (R. Ozimec in pers. comm.), while the Dejanova peéina cave (type locality of Abasola borisi Hadzi, 1973) is now submerged in an artificial lake. Once the Dinaric travuniids have been revised, the species Dinaria vjetrenicae and Abasola hofferi may no longer be placed in to the genera where they currently are. Laniatorid Paralola buresi Kratochvil,1951 inhabits the same regions as T. ser- bica gen. n., sp. n.(western part of Mt Stara Planina), and Lola insularis Kratochvil, 1937 (both in the Phalangodoidea-Phalangodidae) is found in the south Dinaric region, like Dinaric travuniids. After comparing characteristics of the species Trojanella ser- bica gen. n., sp. n. and Abasola hofferi (as representatives of Dinaric travuniids) it is clear that both are phylogenetically very distant from each other. It would be inte- TROJANELLA SERBICA 455 resting to research further into interrelations and phylogenetical distance of the genera Paralola and Lola (Phalangodidae) for the purpose of comparison. ACKNOWLEDGEMENTS I thank biospeleologist Sinisa Ognjenovié who has collected the first specimens of Trojanella serbica gen. sp. n.. Phil Sirvid, Nobuo Tsurusaki, Graham Milledge, Jürgen Gruber and Mark Harvey have provided the necessary literature. Moméilo Popovié has made the nice drawing of 7. serbica gen. n., sp. n. Misa Bokorov produced the SEMs. Roman Ozimec shared information on the type locality of Abasola troglodytes. Jürgen Gruber, Peter Schwendinger, Darrell Ubick and Jochen Martens critically commented on an earlier version of this manuscript. I am grateful to them all. REFERENCES BRIGGS, T.S. 1969. A new Holarctic family of laniatorid phalangids. The Pan-Pacific Entomo- logist 45: 35-50. BRIGGS, T.S. 1971. The harvestmen of family Triaenonychidae in North America (Opiliones). Occasional Papers of the California Academy of Sciences 90: 1-43. DUMITRESCU, D. 1974. Contribution à l’étude de l’appareil digestif (intestin moyen) des Opilions (Arachnida). Travaux du Muséum d'Histoire Naturelle “Grigore Antipa” 14: 95-107. DUMITRESCU, D. 1976. Recherches morphologiques sur l’appareil digestif (intestin moyen) des Gonyleptomorphi (Arachnida, Opilionida). Travaux du Muséum d'Histoire Naturelle “Grigore Antipa” 17: 17-30. HADZ1, J. 1932. Prilog poznavanju peéinske faune Vjetrenice (in serbian cirilic). Glas Srpske Kraljevske akademije 151: 102-157. Hanzi, J. 1935. Ein eigentümlicher neuer Höhlen-Opilionid aus Nord-Amerika, Cladonychium corii g. n. sp. n. Biologia Generalis 11: 49-72. Hapzı, J. 1973. Novi taksoni suhih juZin (Opilionidea) v Jugoslaviji. Razprave, Classis IV, Historia Naturalis et Medicina, SAZU, Ljubljana, 16: 1-120. HUNT, G.S & HICKMAN, J.L. 1993. A revision of the Genus Lomanella Pocock and its implifi- cations for family level classification in the Travunioidea (Arachnida: Opiliones: Triaenonychidae). Records of the Australian Museum 45: 81-119. MARTENS J. 1978. Spinnentiere, Arachnida, Weberknechte, Opiliones. Die Tierwelt Deutschlands, Gustav Fischer, Jena, 64:1-449. MARTENS J. 1986. Die Grossgliederung der Opiliones und die Evolution der Ordnung (Arach- nida)(pp. 289-310). Jn: BARRIENTOS, J. A.(ed.). Actas 10 Congreso International de Aracnologia, Jaca/España, Septiembre 1986, vol. 1. Instituto Pirenaico de Ecologia & Grupo de Aracnologia, Barcelona, X + 428 pp. SUZUKI, S. 1976. Two triaenonychid harvestmen from the northeast, Japan (Triaenonychidae, Opiliones, Arachnida). Journal of Science of the Hiroshima University B, 26: 177-185. SILHAVY, V. 1936. Novy jeskynni sekaë z Jugoslavie, Abasola hofferi n.sp. Sbornik entomo- logického oddeleni Narodniho Musea v Praze, 14: 208-213. PU Anny = Be dI 9 salvia; dir Ay wet) Lau Gli È Ja m eu Tp - ] A. 454, ét, Sade ent REVUE SUISSE DE ZOOLOGIE 112 (2): 457-509; juin 2005 First records of Pauropoda (Millotauropodidae; Pauropodidae) from Gabon with the description of 16 new species (Pauropoda and Symphyla of the Geneva Museum XIV) Ulf SCHELLER Higgeboholm, Häggesled, S-53194 Järpäs, Sweden. First records of Pauropoda (Millotauropodidae; Pauropodidae) from Gabon with description of 16 new species (Pauropoda and Symphyla of the Geneva Museum XIV). - Two collections of Pauropoda (Myriapoda) from Gabon were studied. Twenty-three species were identified, 16 of them are new to science and are described here: Allopauropus gabonicus sp. n., A. akonesis sp. n., A. barrai sp. n., A. ipassaensis sp. n., A. singesensis sp. n., A. cleofanus sp. n., A. cylindricus sp. n., A. suppeditatus sp. n., A. isodacintrai sp. n., A. stenygros sp. n., A. phakoides sp. n., A. bovistellus sp. n., A. lambdoides sp. n., Cauvetauropus pistillifer sp. n., Hemipau- ropus elongatus sp. n., H. bilobatus sp. n. A key to the species of the sub- genus Perissopauropus in Allopauropus is presented. Most species found in tropical West Africa have not been collected elsewhere, indicating a high degree of endemism. Species occurring outside West Africa more often have ranges including the islands of the Indian Ocean and/or south Asia rather than North or South Africa. The wide range element is poor in species. Keywords: Myriapoda - taxonomy - soil fauna - Africa - biogeography. INTRODUCTION The Pauropoda of tropical West Africa have been studied by Remy, several papers in 1948-1962, and by Scheller, papers in 1975 and 1995. Their studies were based on material from Senegal, Gambia, Guinea, Sierra Leone, the Ivory Coast, Cameroon, Congo, and Angola but even if many species have been reported, the taxo- nomic and distributional knowledge of the West African pauropods is still very incom- plete. One gap is partly filled by this study of the first collections from Gabon. Prof. P. A. Remy (7) was one of the participants of the C.N.R.S. expedition to Gabon in 1962. Unfortunately he died right in the midst of the collection work there, but he and his colleagues, Dr G. Bernardi (+), Paris, Prof. B. Condé (+), Nancy, and Prof. P. P. Grassé (7), Paris, had the opportunity to collect in the Ogooué-Ivindo District in north-eastern Gabon, particularly in the neighbourhood of the Biological Station of Makokou. Their material has been studied here together with material collected by Dr J. A. Barra, Strasbourg, who studied the soil fauna SW of Makokou, of the Plateau Forestier d’Ipassa and of the Île aux Singes in the Ivindo River, 10 km Manuscript accepted 16.08.2004 458 U. SCHELLER downstream Makokou. The two collections, 342 specimens in all, included 23 species. Sixteen species are new to science are described below. The material, preserved in alcohol, is deposited in the collections of the Department of Arthropods and Entomology I, Natural History Museum of Geneva. ABBREVIATIONS AND MEASUREMENTS Abbreviations: ad. ..., subad. ... and juv. ... = an adult, a subadult or a juvenile specimen with the number of pairs of legs indicated. Measurements: length of the body in mm and range of variation in adult paratypes given in brackets. Indication of absolute lengths are always applicated with um. Otherwise the text refer to relative lengths. In the section Systematics the names of the collectors are given by surname only. SYSTEMATICS Order HEXAMEROCERATA MILLOTAUROPODIDAE Genus Millotauropus Remy, 1950 1. Millotauropus angustiramosus Remy Figs 1-3 Millotauropus angustiramosus Remy, 1955a: 117-118, figs 1-5. Material examined. Plateau Forestier d’Ipassa, primary forest, soil, 1 ad.(d), 1 subad. 10(9), 7.VI.1966 (loc. IPA5/E7); ibidem, 1 juv. 8(sex?), 11.VI.1966 (loc. IPA6/CI 9); ibidem, at base of plant, 1 ad.(2),2 subad.9(d), 1 subad. 8( 2), 27.VI.1966 (loc. IPA9/AN2); ibidem, at base of fern, 1 subad.10(d), 27.VI.1966 (loc. IPA9/AN3): ibidem, at base of fern, 1 subad. 9(2), 27.V1.1966 (loc. IPA9/AN4); ibidem, at base of fern, 1 ad.(?), 27.VI.1966 (loc. IPA9/ANS) (all leg. Barra). - Mbeza, secondary forest, near trail at old plantation, 1 ad.(9), 20.11.1962 (loc. 5, leg. Condé). — Mayiga, left side of road to Booué, at trail near stream, 1 sub- ad. 10(d), 22.11.1962 (loc. 7, leg. Condé). - Makokou, dell with source, near the water intake, 1 subad. 8(£), 18.VII.1962 (loc. 48, leg. Condé). Total number. 13 specimens. General distribution. The species is known from Angola only and was described by Remy (1955a) from the Lunda District. Later two more Angolan specimens were reported from two other districts, Cuanza-Norte and Cabinda (Scheller, 1975). Taxonomic remarks. Remy had 3 subad. 10 specimens only when he erected the species and therefore his description is unusually brief. Though the specimens reported above are not in the best condition, it has been possible to emend the description in the following respects. Antennae. Chaetotaxy of segments 4-6: 7+p / 6+R / p+p’+p”+R’+f. Distal part of flagella tapering, pointed. Their relative lengths (base segments included): F = 100, F’ = 256(-259). Antennal branch R on 5° segment cylindrical, 1.4 times as long as wide, its flagellum F 3.0 times longer than R. Distal branch R’ subcylindrical, 1.3 times as long as its greatest diameter. Relative lengths of setae of segment 6 (F = 100): p = (196-)200(-207), p’ = (75-)78(-81), p” = (45-)47(-50). Forked organ f as long as basal PAUROPODA FROM GABON 459 Fics 1-3 Millotauropus angustiramosus Remy, genital papillae. 1, subad. 9, right, anterior view; 2. subad 10, right, lateral view; 3, ad. 11, anterior view. segment of F’ and with 3 distally furcate end-branches. Antennal branches almost glabrous, but R’, basal segment of F’ and basal part of p with short but distinct oblique pubescence. Genital papillae (Figs 1-3). Base with convex sides, papilla straight, subcylin- drical, somewhat tapering, 3.9 times as long its greatest diameter; seta 0.4 of the length of organ; pubescence usually very short, most distally distinct, dense, oblique. Seta on coxa of leg 2 as other coxal setae of anterior legs. Papillae in subad. 10 conical, with convex sides, 3.3 times as long as their greatest diameter; seta 0.3 of the length of organ, in subad. 9 subcylindrical, distal part strongly tapering and with a very short distal seta; pubescence minute, dense. Pygidium. Remy (1955a) said that the st were very thin and according to his fig. 1 they were glabrous too. In the material from Gabon, and in the Angolan spe- cimens studied by me, they are proportionately shorter and with a few hairs. A few characters of the pygidial sternum are deviating too. The setae bj in the Gabon spe- cimens are tapering, not subcylindrical, and they are longer than their interdistance, not 0.8 of that length. Moreover are the branches of the anal plate tapering in the distal half, not cylindrical. The latter character was found also in the Angolan specimens. Order TETRAMEROCERATA PAUROPODIDAE Genus Allopauropus Silvestri, 1902 Subgenus Allopauropus s. str. 460 U. SCHELLER 2. Allopauropus (A.) bicornis Remy Allopauropus bicornis Remy, 1948a: 568-569, fig. 1. Material examined. Plateau Forestier d’Ipassa, primary forest, litter, lad. 9(2) 23.VI.1966 (loc. IPA8/IVIS, leg. Barra). - Mbeza, secondary forest, near trail at old plantation, 1 juv. 6, 20.11.1962 (loc. 5, leg. Condé). Total number. 2 specimens. General distribution. Known from tropical Africa only: Kenya (Remy, 1948a) and Angola (Scheller, 1975). 3. Allopauropus (A.) dundoensis Remy Allopauropus dundoensis Remy, 1955a: 121-122, fig. 3. Material examined. Mbeza, secondary forest, near trail at old plantation, 1 subad. 8(2), 20.11.1962 (loc. 5, leg. Condé). Total number. | specimen. General distribution. A. (A.) dundoensis has been collected on both sides of the Atlantic, in Amazonia (Scheller, 1994, 1997) and in Angola (Remy, 1955a; Scheller, 1975). 4. Allopauropus (A.) sphaeruliger Remy Allopauropus sphaeruliger Remy, 1948b: 116-117, fig. 1. Material examined. Plateau Forestier d’Ipassa, primary forest, under piece of wood, 1 ad. 99), 11.VI.1966 (loc. IPA6/CIS, leg. Barra); ibidem, soil, 1 juv. 3, 23.VI.1966 (loc. IPA8/AVCT3, leg. Barra). Total number. 2 specimens. General distribution. A. (A.) sphaeruliger is widely distributed in tropical Africa and has been collected in south and east Asia as well. Previous African records are from Gambia (Remy, 1958a), the Ivory Coast (Remy, 1948b, 1952a, 1953), Angola (Scheller, 1975), Madagascar (Remy, 1956b; Remy & Rollet, 1960), Réunion (Remy, 1956b), Mauritius (Remy, 1959b). 5. Allopauropus (A.) gabonicus sp. n. Figs 4-13 Type material. Holotype: ad. 9(9), GABON, Plateau Forestier d’Ipassa, primary forest, soil, 20.V.1966 (loc. I PA3/4, leg. Barra). Other material. Plateau Forestier d’Ipassa, primary forest, soil, 1 juv. 6 (moulting), 7.VI.1966 (loc. IPA5/E8); ibidem, 1 juv. 3, 23.VI.1966 (loc. IPA8/AMC2); ibidem, at base of plant, 1 juv. 3, 27.VI.1966 (loc. IPA9/AN4, leg. Barra). Total number. 4 specimens. Diagnosis. The new species is close to Allopauropus (A.) aculeatus Remy and A. (A.) stilifer Remy, from Congo (Remy, 1954) and Angola (Remy, 1955a), res- pectively. These three species together form a homogenous group characterized by great similarities in the shape of the anal plate and in the general chaetotaxy of the pygidial tergum. The anal plate is narrowest anteriorly, linguiform and with a single stiletto-shaped appendage, and the setae of the pygidial tergum are of very similar shape and length. A. gabonicus sp. n. is distinguished from both A. aculeatus and A. stilifer in the pubescence of the pygidial setae a3 (strong sparse hairs in A. gabonicus sp. n., very short or glabrous in A. aculeatus and A. stilifer) and the shape of the sternal antennal branch (subcylindrical with indistinct anterior truncation in A. - PAUROPODA FROM GABON 461 n re E (Si = - - - - - - - - - - - - - - u € Fics 4-13 Allopauropus (A.) gabonicus sp. n., holotype, ad. 9(9). 4, head, median and right part, tergal view; 5, lateral group of setae and posterior margin of right temporal organ; 6, left antenna, ter- gal view; 7, collum segment, median and left part, sternal view; 8, tergite VI, posterior part; 9, seta on trochanter of leg 9; 10, tarsus of leg 9; 11, pygidium, median and left part, sternal view; 12, anal plate, lateral view; 13, stylus, lateral view. Scale a: Figs 4-5, 7-8, 10; b: Figs 9, 11; c: Figs 6, 12-13. 462 U. SCHELLER gabonicus Sp. n., truncated posteriorly too in A. stilifer and proportionately short and wide in A. aculeatus). Moreover is the antennal globulus different (large spherical in A. gabonicus sp. n., very small in A. aculeatus and longish in A. stilifer). Etymology. A latinized adjective of the name Gabon. DESCRIPTION Length. 1.02 mm. Head (Figs 4, 5). Tergal setae blunt, striate, median ones rather long, clavate; some lateral ones long, (sub)cylindrical. Relative lengths of setae, 15° row: a, = 10, a; =11; 220 row: aj = 13, a, = 20, a, = 17; 3 row: aj = 10, a, = 11; 4" row: a, = II, a, = 27, a3 = 25, ag = 15; lateral group: /; = 21,1, = 20, /3 = 15. Ratio aj/aj — a, is in 1st, 3rd and 4th rows 1.1, in 2"4 row 0.6. No posterior aperture in temporal organs. Other parts of these organs not studied. Head cuticle glabrous. Antennae (Fig. 6). Segment 4 with 6 setae, these cylindrical, distally tapering, striate. Relative lengths of setae: p = 100, p’ = 84, p° = 62, 97 = 18772323109 Tergal seta p 1.1 times as long as tergal branch r. The latter somewhat fusiform, 5.1 times as long as its greatest diameter and 1.3 times as long as sternal branch s, the latter 2.9 times as long as its greatest diameter and with its anterodistal corner somewhat truncate. Seta g somewhat thinner than p”, cylindrical, blunt, striate, 0.8 of the length of s. Relative lengths of flagella (basal segments included) and basal segments: Fj = 100, bs, = 8; Fy = 42, bs, = 7; F3 = 73, bs3 = 8. F> thinnest. F| 2.5 times as long as f, F; and F; 1.6 and 2.5 times as long as s respectively. Distal calyces of F| and F3 hemi- spherical, those of F smaller and conical. Globulus g 1.1 times as long as wide; 8-9 bracts of unequal lengths; width of g 1.1 times as wide as greatest diameter of r. Antennae glabrous. Trunk. Setae of collum segment (Fig. 7) simple, subcylindrical, blunt, striate, sublateral ones 1.7 times as long as submedian ones; sternite process low, blunt, glabrous; appendages short, subhemispherical with flattened caps, glabrous. Setae on anterior tergites as submedian setae on head; on posterior tergites longer, most posteriorly tapering and shortly pubescent. 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 8) 0.7 of their inter- distance and 0.9 of the length of pygidial setae a. Bothriotricha. Relative lengths: T, = 100, T> = 98, T; = ?, T, = 136,75 = 216. All with very thin axes, those of 73 thickest; pubescence minute. Legs. Setae on coxa and trochanter (Fig. 9) of leg 9 furcate, branches curved, subequal in length; primary one thickest, tapering, pointed, striate, secondary one cylindrical, blunt, indistinctly striate. Corresponding setae on more anterior legs not studied. Tarsus of leg 9 (Fig. 10) tapering, 5.4 times as long as its greatest diameter, very slender, distal part subcylindrical. Proximal seta tapering, pointed, with very short oblique pubescence, 1/4 of the length of tarsus, 2.5 times as long as distal seta, the latter somewhat clavate, striate. Cuticle of tarsus almost glabrous. Pygidium (Fig. 11). Tergum. Posterior margin rounded. Relative lengths of setae: a, = 100, a, = 104, az = 202, st = 23. a, and a, tapering, glabrous, the former straight, pointed, converging, the latter curved inwards and diverging. a3 and st PAUROPODA FROM GABON 463 (Fig. 13) curved inwards, the former tapering, pointed, diverging and with sparse, oblique, long pubescence, the latter clavate, converging, almost glabrous. Distance a, — a; 0.6 of the length of a,, distance aj — a; 4.3 times as long as distance a, — az; dis- tance st — st 2.3 times as long as st and 0.8 of distance a, — ay. Sternum. Posterior margin between bj with broadly V-shaped indentation. Relative lengths of setae (a; = 100): bj = 131, b, = 45, b3 = 64. bj and b3 subcylin- drical, the former with very faint pubescence most distally, the latter somewhat striate; b> tapering, pointed, glabrous. b; 1.4 times as long as their interdistance; by about as long as distance dj - b>; b3 0.5 of distance b3 — b3. Anal plate (Fig. 12) directed downwards, about as broad as long, narrowest an- teriorly, lateral margins almost straight, posterolateral corners rounded and posterior margin straight; stiletto-shaped appendage, as long as the plate, protruding backwards from middle of posterior margin. 6. Allopauropus (A.) akonesis sp. n. Figs 14-22 Type material. Holotype: ad. 9( 2), GABON, 11 km W Makokou, at road to Booué, forest near Ntsibelong, 19.11.1962 (loc. 4, leg. Condé). Paratypes: same data as for holotype, 7 ad. 9(2),3 subad. 8(2), 1 juv. 6. Other material. Same data as for holotype, 2 juv. 5, 11 km W Makokou, at road to Booué, forest near Ntsibelong, right bank of the Ivindo River, 2 ad. 9(@), 19.11.1962 (loc. 3B, leg. Condé). — Mayiga, Endoumé, old plantation near the village, soil, 6 ad. 9(9), 2 subad. 8( 2), 3 juv. 6, 1 juv. 5, 1 juv. 3, 21.11.1962 (loc. 7, leg. Condé); ibidem, forest near the village, 1 ad. 9(2), 1 juv. 6, 1 juv. 5, 21.11.1962 (loc. 7, leg. Condé & Remy); Mayiga, forest near road to Booué, at trail, 2 subad. 8(£), 22.11.1962 (loc. 8, leg. Condé & Remy). — Loualouah, right bank of the Ivindo River, 1 ad. 9(@), 1 subad. 8(@), 8.III.1962 (loc. 3bis, leg. Grassé); ibidem, right bank of the Ivindo River, under piece of wood, 4 ad. 9(@), 1 juv. 6, 1 juv. 5, 1 juv. 3, 10.111.1962 (loc. 3, leg. Grassé); ibidem, 1 ad. 9(£), under piece of wood, 8-10.III.1962 (loc. 23, leg. Condé & Remy). — Edoungavion, 1 ad. 99), 19.11.1962 (loc. 4, leg. Condé & Remy) and 2 ad. 9(2), 2.111.1962 (loc. 15bis, leg. Condé); ibidem, at beginning of trail to Alarmintang, near stream, under piece of wood, 1 ad. 9( 9), 2.III.1962 (loc. 15, leg. Condé). — Belinga, at trail along the drinking-water pipe, under piece of wood, 2 ad. 9(2),2 subad. 8(2),1 juv.5,16.II 1962 (loc. 33bis, leg. Condé); Belinga, 1 ad. 9(2), 3 subad. 8(9), 4 juv. 6, 4 juv. 5, 17.111.1962 (loc. 35, leg. Condé & Bernardi); Belinga, under small stone, 1 ad. 9(9), 1 juv. 6, 22.VII.1 962 (loc. 55, leg. Condé). — Mvadhi, end of trail to Dubost forest, under stones in laterite, 2 subad. 8(2), 9.IX.1962 (loc. 101, leg. Condé); ibidem, miners crossroad, under stones, 2 ad. 9(2), 24.IX.1962 (loc. 107, leg. Condé). Total number. 69 specimens. Diagnosis. The new species is close to Allopauropus (A.) stilifer and A. (A.) aculeatus, the former described from Angola (Remy, 1955a) and the latter from Belgian Congo (Remy, 1954) later also reported from Angola (Remy, 1955a). Together with the preceding species, A. (A.) gabonicus sp. n., these three species form a homogenous group characterised by great similarities in the shape of the anal plate and in the general chaetotaxy of the pygidial tergum. The anal plate is narrowest anteriorly, linguiform and with a single stiletto-shaped posterior appendage, and the setae of the pygidial tergum are of similar shape and length. A. (A.) akonesis sp. n. is distinguished from A. (A.) stilifer by the shape of the sternal antennal branch [only anterodistal corner truncated in A. (A.) akonesis sp. n., both anterior corners truncated in A. (A.) stilifer], the st [straight and distinctly fusiform in A. (A.) akonensis sp. n., thinner, curved inwards and clavate in A. (A.) stilifer], the shape of the posterior margin of the pygi- 464 U. SCHELLER dial sternum [median indentation deep with steep lateral margins in A. (A.) akonesis sp. n., Shallow and with rounded inner margins in A. (A.) stilifer]. There are dissimilarities in the shape of the anal plate too [appendage with two small knobs at its base and appendage itself probably perforated and with convex margins in A. (A.) akonesis sp. n., straight margin and neither knobs nor perforation in A. (A.) stilifer]. Many charac- ters separate A. (A.) akonesis sp. n. from A. (A.) aculeatus, e.g. the shape of the tergal antennal branch [very slender, 6.0-6.5 times as long as its greatest diameter in A. (A.) akonesis sp. n., 3 times as long as its greatest diameter in A. (A.) aculeatus Remy], the globulus g [distinct conical stalk in A. (A.) akonesis sp. n., short-stalked in A. (A.) aculeatus]. Moreover, the appendage of the anal plate is at least as long as the plate in A.(A.) akonesis sp. n., about 0.3 of the length of the plate in A. (A.) aculeatus) and the b; are proportionately much longer in A. (A.) akonesis sp. n. than in A. (A.) aculeatus. Etymology. From Greek akonesis = forming a sharpening (referring to the pointed appendage of the anal plate). DESCRIPTION Length. (1.20-)1.39(-1.52) mm. Head (Fig. 14). Tergal setae blunt, striate, median ones rather long, somewhat clavate; some lateral ones long, (sub)cylindrical. Relative lengths of setae, 1°! row: a, = 10; a, —12,, 294 row: a; = 12; a, = (16-)18, a3 = 1516); row ta Na 9(-10); 4" row: aj = 9(-10), a, = 19(-21), az = (19-)22, a, = 14(-15); lateral group: J, = 18(-19), 1, = 15(-18), 13 = 22(-24). The ratio aj/aj — aj is in 18t row (1.4-)1.5, 224 row 0.7, 3™ row (1.1-)1.3 and 4" row 1.5. Length of temporal organs (0.7-)0.8 of their shortest interdistance; neither pistil nor posterior aperture present. Head cuticle glabrous. Antennae (Fig. 15). Segment 4 with 6 thin, cylindrical, distally tapering, striate setae. Relative lengths of setae: p = 100, p’ = 84(-88), p” = (47-)48(-52), p” = 10(-15), r = (31-)32(-33), u = (10-)12. Tergal seta p as long as tergal branch f. The latter somewhat fusiform, 6.0(-6.5) times as long as its greatest diameter and (1.3-)1.5 times as long as sternal branch s, the latter (2.3-)2.4(-2.7) times as long as its greatest diameter and with truncate anterodistal corner. Seta g somewhat thinner than p”, cylin- drical, blunt, striate, (0.7-)0.8 of the length of s. Relative lengths of flagella (basal seg- ments included) and basal segments: Fj = 100, bs, = 5; Fy = 33(-47), bs, = 4; F3 = 73(-88), bs3 = 5(-6). Flagella similar in thickness; Fj (3.1-)3.5 times as long as ¢, F, and F3 (1.7-)1.8(-2.3) and (3.6-)3.8(-3.9) times as long as s respectively. Distal calyces conical, with rounded tip. Globulus g (1.3-)1.4 times as long as wide; = 10 thin bracts present; width of g (1.3-)1.4 times as long as wide and 0.9 of greatest diameter of r. Antennae glabrous. Trunk. Setae of collum segment (Fig. 16) furcate, subcylindrical, somewhat tapering, striate, secondary branch rudimentary, blunt, glabrous. Sublateral setae 1.5 (- 1.6) times as long as submedian ones; sternite process with broad base, anterior part narrow, incised distally; appendages short, wide, with flattened caps. Process and appendages with faint pubescence. Setae on anterior tergites as submedian setae on head; on posterior tergites longer, most posteriorly tapering and almost glabrous. 4+4 setae on tergite I, 6+6 on PAUROPODA FROM GABON 465 17 DIE 16 Fics 14-22 Allopauropus (A.) akonesis sp. n., holotype, ad. 9(2). 14, head, median and right part, tergal view; 15, right antenna, sternal view; 16, collum segment, median and left part, sternal view; 17, tergite VI, right posteromedian part; 18, setae on coxa of leg 9; 19, seta on trochanter of leg 9; 20, tarsus of leg 9; 21, pygidium, median and left part, sternal view; 22, anal plate, lateral view. Scale a: Figs 16-20; b: Figs 14-15; c: Figs 21-22. I-IV, 6+4 on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 17) (0.5-)0.7 of their interdistance and 1.1(-1.2) times as long as pygidial setae a). Bothriotricha. Relative lengths: T, = 100, 7, = (98-)104, 73 = (106-)112(-114), T, = 2(123), T = ?(189-198). All possessing very thin axes with short oblique pubes- cence. 466 U. SCHELLER Legs. Setae on coxa (Fig. 18) and trochanter (Fig. 19) of leg 9 furcate, branches similar, cylindrical, blunt, striate, subequal in length on coxa, secondary branch longest on trochanter. Corresponding setae on more anterior legs with rudi- mentary secondary branches. Tarsus of leg 9 (Fig. 20) tapering, (5.6-)5.9(-6.3) times as long as its greatest diameter, in middle and distal parts very slender, distal 2/3 subcylindrical. Proximal seta tapering, pointed, with very short oblique pubescence, (0.3-)0.4 of the length of tarsus, (3.3-)3.6 times as long as distal seta, the latter is somewhat clavate, with short oblique pubescence. Cuticle of tarsus almost glabrous. Pygidium (Fig. 21). Tergum. Posterior margin rounded but with shallow inden- tations outside st. Relative lengths of setae: a, = 100, a; = (119-)128, a; = (204-) 255, st = 20(-23). aj, a, and az long, thin, tapering, glabrous, somewhat diverging, a, curved outwards, a, and a; curved inwards; st straight, bladder-shaped, tapering dis- tally, converging, faintly pubescent. Distance a, — a, about as long as aj, distance a, — a, about 6 times longer than distance a, — az; distance st — st (2.7-)2.9 times as long as st and (0.5-)0.6 of distance a, — a}. Sternum. Posterior margin between bj with median indentation having steep sides; a median, broadly triangular lobe below anal plate. Relative lengths of setae (a, = 100): bj = 135(-155), by = (57-)60(-65), b3 = (74-)89. bj and b3 of similar shape, thin, tapering, blunt, very short pubescence or striate only most distally; b> tapering, pointed, glabrous. b,1.4(-1.5) times as long as their interdistance; b, (0.8-)0.9 of dis- tance bj — bo; b; 0.5 of distance b3 — b3. Anal plate (Figs 21, 22) directed obliquely upwards, narrowest anteriorly, broader than long, lateral margins convex, posterolateral corners rounded and posterior margin straight. A stiletto-shaped appendage protruding backwards from posterior margin. Appendage longer than plate and with two small knobs at base and probably a perforation in the middle of basal part. 7. Allopauropus (A.) barrai sp. n. Figs 23-31 Type material. Holotype: subad. 8(9), GABON, Plateau Forestier d’Ipassa, primary forest, from fruit on soil, 20.V.1966 (loc. IPA3/Fruit B, leg. Barra). Paratypes: GABON, Plateau Forestier d’Ipassa, primary forest, 3 subad. 8(1 2 , 2 sex?), 27.VI.1966 (IPA9/AN6, leg. Barra). Other material. Plateau Forestier d’Ipassa, primary forest, in soil, 1 juv. 5, 7.VI.1966 (loc. IPAS/E13, leg. Barra); ibidem, base of fern, 1 juv. 6, 1 juv. 5, 27.VI.1966 (loc. IPA9/AN4, leg. Barra). — Ntsibelong, 1 subad. 8(£), 19.11.1962 (loc. 3, leg. Condé). — Mayiga, Endoumé, 1 Juv. 6, 12.11.1962 (loc. 7, leg. Condé). Total number. 9 specimens. Diagnosis. A. (A.) barrai sp. n. belongs to a homogenous group of species described from the Ivory Coast by Remy (1948b): A. (A.) liticen Remy, A. (A.) buci- nator Remy and A. (A.) vouauxi Remy. These four species have great similarities in the antennae and the bothriotricha, and in the pygidium with its anal plate. The new species is distinguished from Remy’s three species by the shape of the 73 [with swelling in the middle in A. (A.) barrai sp. n., axis thin without swelling in the others] and the pos- terior part of the pygidial tergum [broadly triangular in A. (A.) barrai sp. n., rounded with median indentation in the others]. Etymology. Dedicated to the collector, Dr J.A. Barra. PAUROPODA FROM GABON 467 Fics 23-31 Allopauropus (A.) barrai sp. n., holotype, subad. 8(9). 23, right temporal organ with interior pistil, lateral view; 24, right antenna, sternal view; 25, collum segment, median and left part, sternal view; 26, 7]; 27, 73; 28, seta on trochanter of leg 8; 29, tarsus of leg 8; 30, pygidium, sternal view; 31, anal plate, lateral view. Scale a: Figs 26-27; b: Figs 23-25, 28-29; c: Figs 30-31. DESCRIPTION Length. (0.95-)0.97 mm. Head. Tergal setae not available for study. Temporal organs (Fig. 23) short, in lateral view triangular and faintly pubescent. No posterior aperture but an inner ovoid pistil in posterior half. 468 U. SCHELLER Antennae (Fig. 24). Segment 4 with 5 setae and rudiment of a 6fh one, i. e. p””. Setae cylindrical, blunt, annulate. Relative lengths of setae: p = 100, p’ =p” = (43-)45, r = 50(-52), u = (6-)7. Tergal seta p 0.9 of length of tergal branch r. The latter some- what fusiform, 2.8(-3.0) times as long as its greatest diameter and 1.4 times as long as sternal branch s, this (1.7-)1.9 times as long as its greatest diameter and with distinctly truncate anterodistal corner. Seta g somewhat thinner than p, cylindrical, blunt, striate, 1.2 times as long as s. Relative lengths of flagella (basal segments included) and basal segments: F = 100, bs, = 6; Fy = 27(-28), bs, = (2-)3; F3 = (77-)79, bs; = 5. Fy thinnest. F, 3.5(-3.6) times as long as r, F and F3 1.6 and (4.5-)4.6 times as long as s, respectively. Distal calyces of F, and F3 hemispherical, those of F smaller and coni- cal. Flagella axes cylindrical below calyx. Globulus g as long as wide, with ~ 9 thin bracts; width of g 0.6(-0.7) of greatest diameter of t. Antennae faintly pubescent. Trunk. Setae of collum segment (Fig. 25) simple, subcylindrical, blunt, striate, sublateral ones 2.9 times as long as submedian ones; sternite process with small base and narrow anterior lengthening with apical incision; appendages low, with wide caps. Appendages faintly pubescent, process distinctly pubescent anteriorly, caps glabrous. Setae on tergites not available for study. Bothriotricha (Figs 26, 27). Relative lengths: 7; = 100, T = 104, 73 = 112, Ty = 178. All with simple straight axes. T, and 7, possessing thin axes, most proximally glabrous, more outwards with thin pubescence hairs, these at first short and simple, then longer and branched. Proximal half of 73 thin but at the middle a distinct clavate thickening, axis outside it very thin; pubescence on proximal half and on thickening consisting of short simple oblique hairs, on distal half as on 7} and 7). 75 with minute pubescence of very simple oblique hairs. Legs. Tibia of legs 2-8 short, annulate. Setae on coxa and trochanter (Fig. 28) of leg 8 furcate, branches thin, cylindrical, striate, primary branch almost twice longer than secondary branch. More anteriorly setae on trochanter similar but those on coxae simple without rudiment of secondary branch. Tarsus of leg 8 (Fig. 29) 2.8 times as long as its greatest diameter, tapering. Setae with short oblique pubescence, proximal one cylindrical and pointed, distal one somewhat clavate. Proximal seta 0.3 of length of tarsus, about as long as distal seta. Cuticle of tarsus faintly pubescent. Pygidium (Fig. 30). Tergum. Posterior part between setae a3 obtusely triangular. Relative lengths of setae: a, = 100, a, = (91-)97, a3 = 153(-179), st = 20(-25). ay, ay and a; long, thin, tapering, glabrous except most distally, curved inwards; st bladder- shaped, with faint pubescence, converging. Distance a, — a, 0.8 of length of aj, dis- tance a, — a, about 4 times as long as distance a, — a3; distance st — st 2.1(-2.2) times as long as st and 0.5 of distance a, — ay. Sternum. Posterior margin with broadly V-shaped indentation between b, and small triangular lobe with rounded tip below anal plate. Relative lengths of setae (a, = 100): bj = 87(-97), b> = (41-)50, b3 = 22(-23). Setae striate, b, and b, thin, tapering; b3 subcylindrical. bj1.1 times as long as their interdistance; by 1.2(-1.3) times as long as distance bj — b>; b; 0.2 of distance b3 — b3. Anal plate (Figs 30, 31) directed obliquely upwards, about as broad as long, narrowest anteriorly, lateral margins convex and posterior margin almost straight, PAUROPODA FROM GABON 469 posterolateral corners right-angled. Two submedian appendages projecting backwards from sternal side of posterior margin, these 0.6 of length of plate, cylindrical, blunt, somewhat curved inwards. Plate and appendages glabrous. 8. Allopauropus (A.) ipassaensis sp. n. Figs 32-44 Type material. Holotype: ad. 9(9 ), GABON, Plateau Forestier d’Ipassa, primary forest, soil, 23.VI.1966 (loc. IPA8/AVCTS, leg. Barra). Total number. 1 specimen. Diagnosis. Among several similar species in the subgenus the new species can be distinguished by the following combination of characters: the tergal setae of the head are cylindrical, the temporal organs have a distinct inner pistil, the antennal branches r and s are proportionately long, the distinctly stalked antennal globulus g has few bracts, the appendage of the collum segment has a deep anterior incision, and the bothriotricha T, and 7, have long branched hairs on distal half. Etymology. A latinized adjective of the name Ipassa. DESCRIPTION Length. 0.72 mm. Head (Fig. 32). Tergal setae cylindrical, thin, blunt, striate, of medium lengths or long. Relative lengths of setae, 15t row: a, = 10, a, = 11; 24 row: a, = 12, a; = 13, az = 11; 35 row: a, = 8, a, = 10; 4 row: a, = a4 = 9, a, = 17, az = 18; lateral group: 1, = 13,1,=13 = 10. Ratio ay/aı - aj in 1°! row 1.5, in 229 row 0.9, in 34 row 0.6 and in 4th row 0.8. Temporal organs 1.1 times as long as their shortest interdistance; in pos- terior half a claviform pistil (Figs 33, 34) in a depression of the cuticle; pistil 3 times longer than its greatest diameter. No small posterior aperture present. Head cuticle glabrous. Antennae (Fig. 35). Segment 4 with 6 setae, 5 of them cylindrical, distally ta- pering, striate, one rudimentary. Relative lengths of setae: p = 100, p’ = 86, p = 37, p”=9,r=34,u= 1. Tergal seta p as long as tergal branch r. The latter very slender, somewhat tapering proximally, 5.8 times as long as its greatest diameter and 1.3 times as long as sternal branch s, the latter 2.9 times as long as its greatest diameter and with its anterodistal corner somewhat truncate. Seta q cylindrical, blunt, striate, as thick as p’, almost 0.8 of length of s. Lengths of flagella (basal segments included) and basal segments: F = ?, bs; = 6; F> = 48, bs, = 6; F3 = 50, bs; = 6 um. F, thinnest; F, and F 1.8 and 1.9 times as long as s respectively. Distal calyces of F, and F; propor- tionately small and conical. Axis of flagella not widened below calyx. Globulus g 1.4 times as long as wide; 8-9 bracts; width of g 0.8 of greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment (Fig. 36) furcate, cylindrical, blunt, striate, with rudimentary secondary branch, sublateral setae 1.5 times as long as submedian setae; sternite process narrow anteriorly, with deep anterior incision; appendages low and with flattened caps; appendages and anterior part of process with short but distinct pubescence. Setae on anterior tergites as submedian setae on head; on posterior tergites longer, and shortly pubescent. 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V and 4+2 470 U. SCHELLER 42 a — pee at à —— Fics 32-44 Allopauropus (A.) ipassaensis sp. n., holotype, ad. 9(9). 32, head, median and right part, tergal view; 33, pistil, sternal view; 34, pistil, lateral view; 35, left antenna, sternal view; 36, collum segment, median and left part, sternal view; 37, tergite VI, right posteromedian part; 38, 7; 39, T3; 40, seta on coxa of leg 9; 41, seta on trochanter of leg 9; 42, tarsus of leg 9; 43, pygidium, median and left part, sternal view; 44, anal plate, lateral view. Scale a: Figs 32, 37-42; b: Figs 33-34, 36; c: Figs 35, 43-44. PAUROPODA FROM GABON 471 on VI. Posterior setae on tergite VI (Fig. 37) 1.3 times as long as their interdistance and as long as pygidial setae a]. Bothriotricha (Figs 38, 39). Their relative lengths: 7 = 100, 7, = 103, 73 = 111, T, = 145, T; = 211. All with very thin axes, those of 73 thickest. Pubescence hairs short simple, oblique everywhere except on distal half of 7, and 7,, hairs there increasing in length outwards and becoming branched. Legs. Setae on coxa (Fig. 40) and trochanter (Fig. 41) of leg 9 furcate, branches cylindrical, blunt, shortly pubescent; branches equal in length on coxal seta, primary branch twice longer than secondary branch on seta of trochanter. More anterior setae with rudimentary secondary branch. Tarsus of leg 9 (Fig. 42) tapering, 4.6 times as long as its greatest diameter. Proximal seta long, tapering, pointed, with short oblique pubescence; its length 0.5 of length of tarsus and 3.6 times as long as cylindrical, blunt, striate, distal seta. Cuticle of tarsus somewhat pubescent. Pygidium (Fig. 43). Tergum. Posterior margin evenly rounded. Relative lengths of setae: aj = 100, a, = 93, a; = 107, st = 18. a, and a, tapering, glabrous, the former also directed upwards, straight, pointed, the latter curved inwards. a3 as a> but longer and directed obliquely outwards. st somewhat fusiform, a little curved inwards, converging. Distance a, — a, 0.4 of length of aj, distance a, — a, = 5 times longer than distance a; — az; distance st — st 2.4 times as long as st and as long as distance a, — aj. Sternum. Posterior margin between bj with deep, broadly V-shaped indentation and rounded lobe below anal plate. Relative lengths of setae (aj = 100): bj = 112, b, = 38 and 40, b; = 33. Setae thin, b, and b, tapering and pointed, the former almost glabrous, the latter glabrous; b3 cylindrical, blunt, striate. b,1.7 times as long as their interdistance; b> 0.7-0.8 of distance bj — b>; b3 0.4 of distance b; — b3. Anal plate (Figs 43, 44) directed obliquely upwards, about as broad as long, narrowest anteriorly, lateral margins convex, posterolateral corners rounded and pos- terior margin straight with minute median indentation; two cylindrical, blunt, striate, somewhat diverging appendages protruding backwards from sternal side of posterior margin, length of appendages 0.6 of length of plate. 9. Allopauropus (A.) singesensis sp. n. Figs 45-52 Type material. Holotype: ad. 9(9), GABON, Île aux Singes, in the Ivindo River, 10 km downstream Makokou, primary forest, at base of fern, 11.VII.1966 (loc. IS3/AN4 B, leg. Barra). Paratype: same data as for holotype, 1 ad. 9(9), 4.VII.1966 (loc. IS2/7, leg. Barra). Total number. 2 specimens. Diagnosis. A. (A.) singesensis sp. n. may be closest to A. (A.) inornatus (Hansen) from Paraguay and A. (A.) siamensis (Hansen) from Thailand (Hansen, 1902). They have similarities both in antennae and pygidium. Good distinctive characters in relation to A. (A.) inornatus (Hansen) are the shape of the tergal setae on the head [thin and cylindrical in A. (A.) singesensis sp. n., thicker and clavate in A. (A.) inornatus], the shape of the st [subcylindrical in A. (A.) singesensis sp. n., distinctly clavate in A. (A.) inornatus] and the shape of the anal plate [longer than broad, with sharp posterolateral corners and posteriorly directed appendages in A. (A.) singesensis sp. n., as long as broad, with rounded posterolateral corners and diverging appendages 472 U. SCHELLER Fics 45-52 Allopauropus (A.) singesensis sp. n., holotype, ad. 9(2). 45, head, median and right part, tergal view; 46, left antenna, sternal view: 47, collum segment, median and left part, sternal view; 48, tergite VI, right posteromedian part; 49, setae on coxa of leg 9; 50, seta on trochanter of leg 9; 51, tarsus of leg 9; 52, pygidium, sternal view. Scale a: Fig 48; b: Fig 45, 49-51; c: Figs 46-47, 522 PAUROPODA FROM GABON 473 in A. (A.) inornatus]. The new species is distinguished from A. (A.) siamensis by the length of the distal seta on the tarsus of leg 9 [0.5 of length of tarsus in A. (A.) singe- sensis sp. n., 0.1-0.2 in A. (A.) siamensis], the length of the pygidial setae [long in A. (A.) singesensis sp. n., short in A. (A.) siamensis] and the shape of the st [cylindrical in A. (A.) singesensis sp. n., thick and clavate in A. (A.) siamensis]. Etymology. A latinized adjective of the name Singes. DESCRIPTION Length. (0.75-)0.78 mm. Head (Fig. 45). Tergal setae of medium lengths or long, thin, cylindrical, blunt, glabrous. Relative lengths of setae (holotype only), 15 row: aj = 10, a) = 11; 24 row: acilia a - =11; 3drow: a), =5,a, = 8; 4 row: a) =5,a; a 19, a, = 9; lateral group not studied. Ratio aj/aj — a, in 15 row 0.9, in 299 row 1.0, in 37d row 1.2 and in 4!h row 0.7. Temporal organs 1.3 times as long as their shortest interdistance; in posterior half a small claviform pistil in a depression of the cuticle. No posterior aperture. Head cuticle glabrous. Antennae (Fig. 46). Segment 4 with 5 setae and rudiment of a 6fh one, i. e. p””. Setae thin, cylindrical, blunt, striate-annulate. Relative lengths of setae: p = 100, p’ = 68(76), p = 35(38), r = 27(29), u= 8. Tergal seta p as long as tergal branch r. The latter somewhat fusiform, 5.1(5.6) times as long as its greatest diameter and 1.1(1.2) times as long as sternal branch s; this 3.1(3.4) times as long as its greatest diameter and with its anterodistal corner distinctly truncate. Seta q thinner than p, cylindrical, striate, 0.6 of length of s. Relative lengths of flagella (basal segments included) and basal seg- ments: F; = 100, bs, = 6; Fy = (73)74, bs, = 5; F3 = (74)75, bs3 = 6. F, thinnest. Fy (2.6)2.9 times as long as r, F' and F 2.2 and 2.2(2.3) times as long as s respectively. Distal calyces conical. Flagella axes very little widened below calyx. Globulus g 1.3(1.5) times as long as wide with = 9 thin bracts; width of g 0.8 of greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment (Fig. 47) furcate, with thin rudiment of secondary branch; main branch cylindrical, on sublateral setae striate, on submedian setae annulate. Sublateral setae 1.4 times as long as submedian ones; sternite process low, anterior lengthening with minute apical incision; appendages with distal half cylindrical and proximal half conical, caps flat. Appendage and processes glabrous. Setae on anterior tergites glabrous, somewhat thickening distally, on posterior tergites tapering distally. 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 48) 1.2 times as long as their interdistance and as long as pygidial setae a). Bothriotricha. Relative lengths: T, = 100, T> = (65)67, T3 = 52(60), T4 = 61(62), Ts = (84)93. All with simple straight axes, 73 thickest. Anterior bothriotricha glabrous, posterior pairs with faint pubescence only most distally. Legs. Setae on coxa (Fig. 49) and trochanter (Fig. 50) of leg 9 furcate, branches (sub)cylindrical, with sparse pubescence; branches subequal in length on coxal seta, secondary branch longest on seta of trochanter. Secondary branch rudimentary on more anterior setae. 474 U. SCHELLER Tarsus of leg 9 (Fig. 51) tapering, 4.3(5.0) times as long as its greatest diameter. Setae thin, with minute pubescence, proximal seta pointed, distal one cylindrical. Proximal seta 0.5 of length of tarsus and 4.6 times as long as distal seta. Cuticle of tarsus glabrous. Pygidium (Fig. 52). Tergum. Posterior part between setae a3 evenly rounded. Relative lengths of setae: a; = 100, a; = (71)80, az = 104(112), st = 16. a}, a; and az thin, tapering, curved inwards, glabrous, a; converging, a; diverging, st subcylindrical and converging, straight, with minute pubescence. Distance a, — a; (0.5-)0.6 of length of aj, distance aj — a; 3.5(4.0) times as long as distance a, — az; distance st — st 4.9(5.0) times as long as st and 0.8 of distance a, — aj. Tergum glabrous. Sternum. Posterior margin with broad and shallow indentation between bj. Relative lengths of setae (aj = 100): bj = (123)141, by = (50)54, b3 = (39)43. Setae thin, dj and b,, tapering, with short oblique pubescence most distally, b3 cylindrical, glabrous. dj (1.4)1.5 times as long interdistance; b, as long as distance dj — b>; b3 0.4 of distance b3 — b3. Sternum glabrous. Anal plate glabrous, 1.1 times as long as broad, narrowest anteriorly, lateral margins convex and posterior margin somewhat convex, posterolateral corners distinct. Two submedian appendages projecting backwards from sternal side of posterior margin. These 0.5 of length of plate, straight, cylindrical, blunt, striate. 10. Allopauropus (A.) cleofanus sp. n. Figs 53-60 Type material. Holotype: ad. 9(9), GABON, Mayiga, Endoumé, old plantation near the village, in soil, 21.11.1962 (loc. 7, leg. Condé). Total number. 1 specimen. Diagnosis. The subequal length of the flagellae F, and F; and the pygidial ster- num with setae b3 on distinct diverging lobes are characters which place A. (A.) cleo- fanus sp. n. close to A. (A.) bicornis Remy from Kenya (Remy, 1948a) and Angola (Remy, 1955a). There are also similarities in the general shape of the anal plate, but it has good distinctive characters too [narrowest anteriorly and with pointed appendages in A. (A.) cleofanus sp. n., broadest anteriorly and with cylindrical appendages in A. (A.) bicornis]. Etymology. From Anglo-Saxon cleofan = cleave, split (posterior half of anal plate). DESCRIPTION Length. 0.98 mm. Head (Fig. 53). Tergal setae subcylindrical, blunt, striate, some sublateral ones long. Relative lengths of setae, 1° row: a, = 10, a, =11; 294 row: a) = a3 = 12,4, = 15; 374 row: aj = 10, a, = 12; 4th row: aj = 12, ay = 15, az = 16, ay = 12; lateral group: l, = l3 = 15, 1, = 12. Ratio aj/a, — aj in 1 row 1.2, 24 row 1.0, 319 row 2.6 and 4fh row 1.3. Temporal organs 2.4 times as long as their shortest interdistance. Small pos- terior aperture outside posterior margin at level of /,. Head cuticle glabrous. Antennae (Fig. 54). Segment 4 with 4 thin, cylindrical, striate setae. Relative lengths of setae: p = 100, p’ = 83, p” = 52,r = 24. Neither p”’ nor u present. Tergal seta p 0.8 of length of tergal branch r. The latter somewhat fusiform, 3.6 times as long as its greatest diameter and 1.2 times as long as sternal branch s; this 3.2 times as long PAUROPODA FROM GABON 475 1, u U 1 i! I li il E t {] } ] Ss À Q 1 f in # ll ll r -- YU et u 1} 4 ‘ il / N . \7 (à p 0 "> PG, I7V\ 4 CANON 4 ( N Gp” Q \ \\ N x IS ' \ | 5 | 58 D Fics 53-60 Allopauropus (A.) cleofanus sp. n., holotype, ad. 9(9). 53, head, median and right part, tergal view; 54, left antenna, tergal view; 55, tergite VI, right posteromedian part; 56, seta on trochanter of leg 9; 57, seta on coxa of leg 8; 58, tarsus of leg 9; 59, pygidium, median and left part, sternal view; 60, anal plate, lateral view. Scale a: Figs 53, 56-58; b: Fig 55; c: Figs 54, 59-60. 476 U. SCHELLER as its greatest diameter and with its anterodistal corner truncate. Seta g as p”, 0.7 of length of s. Relative lengths of flagella (basal segments included) and basal segments: F, = 100, bs, = 8; Fy = 82, bs, = 8; F3= 81, bs3 = 8. F 2.3 times as long as t, F> and F3 2.2 times as long as s. Distal calyces small, helmet-shaped. Globulus g 1.6 times as long as wide, almost 0.4 of length of s, = 10 bracts; width of g 0.9 of greatest diameter of t. Antennae glabrous. Trunk. Collum segment not studied. Setae on anterior tergites as submedian se- tae on head; on posterior tergites longer, most posteriorly tapering and glabrous. 4+4 setae on tergite I, 6+6 on I-IV, 6+4 on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 55) as long as their interdistance and 1.2 times as long as pygidial setae a. Bothriotricha. Relative lengths: 7} = 100, T> = 159, 73 = 165, T, = ?, T, = 271. All with very thin axes, those of 73 thickest; pubescence minute. Legs. Setae on coxa and trochanter of leg 9 (Fig. 56) furcate, shortly pubescent, branches equal in length, main branch curved, secondary branch straight. Corres- ponding setae on more anterior legs (Fig. 57) furcate, main branch foliform, with short pubescence, and secondary branch rudimentary, blunt, glabrous. Tarsus of leg 9 (Fig. 58) tapering, slender, 5.0 times as long as its greatest diameter. Proximal seta tapering, pointed, with minute pubescence, 0.4 of the length of tarsus, 3.7 times as long as clavate, striate, distal seta. Cuticle of tarsus glabrous. Pygidium (Fig. 59). Tergum. Posterior margin evenly rounded. Relative lengths of setae: a, = 100, a; = 133, a3 = 247, st = 58. Setae glabrous, a}, a; and a; diverging, a, also cylindrical, curved outwards, a; tapering, somewhat curved inwards, a; cylin- drical, somewhat curved inwards; st cylindrical, somewhat curved inwards and converging. Distance a, — a; 1.6 times as long as aj, distance aj — a, twice longer than distance a, — az; distance st — st 2.7 times as long as st and as long as distance a, — ay. Sternum. Posterior margin between b, with deep and broad indentation with flat bottom, setae b, protruding from outer part of distinct, somewhat diverging lobes. Relative lengths of setae (a; = 10): bj = 52, by = 13, b3 = 12. b, and D; subcylindrical, the former somewhat widened most distally and with short pubescence on distal half, the latter glabrous; b, tapering, glabrous. b,1.9 times as long as their interdistance; b> 0.9 of distance bj — by; b3 0.7 of distance b3 — b3. Anal plate (Fig. 60) directed obliquely upwards, appendages obliquely down- wards, plate 1.3 times longer than broad, narrowest anteriorly, lateral margins convex, deep posterior incision reaching to middle of plate, posterior corners strongly tapering. One long appendage, 0.5 of length of plate protruding from distal part of each of the two posterior lobes. Subgenus Decapauropus Remy, 1957 11. Allopauropus (D.) bouini Remy, 1955 Allopauropus Bouini Remy, 1955a: 129-130, fig. 8. Material. Île aux Singes, in the Ivindo River, 10 km downstream Makokou, primary for- est, in soil, depth 0-5 cm, 12 ad. 9(9), 4.VII.1966 (loc. IS2/6, leg. Barra). Total number. 12 specimens. General distribution. Earlier reported from Angola (Remy, 1955a; Scheller, 1975), Borneo (Scheller, 2001), Florida (Remy, 1958b) and Canada (Scheller, 1984). The species is rare and has a very wide but discontinuous distribution. PAUROPODA FROM GABON 477 12. Allopauropus (D.) proximus Remy, 1948 Allopauropus proximus 1948a : 572-573, fig. 4. Material. Île aux Singes, in the Ivindo River, 10 km downstream Makokou, primary forest, in dead wood, 1 ad. 9(9), 25.V.1966 (loc. IS1/2, leg. Barra). — Ntsibelong, right side of the Ivindo River, under wood, 1 ad. 9( 9), 4.VII.1966 (loc. IS1/2, leg. Barra); ibidem, right bank of the Ivindo River, under bark, 40 ad. 9(9), 6 juv. 6, 19.11.1962 (loc. 3, leg. Grassé). — Loualouah, right side of the Ivindo River, under bark, 5 ad. 99), 10.11.1962 (loc. 3, leg. Condé); ibidem, under bark on trunk and in litter, 6 ad. 9(9), 8-10.1I1.1962 (leg. Remy); ibidem, right bank of the Ivindo River, under bark, 2 ad. 9(9), 1 juv. 6, 8.III.1962 (loc. 3bis, leg. Grassé). — Mayiga, Endoumé, old plantation near village, in soil, 3 ad. 9(2), 21.11.1962 (loc. 7, leg. Remy); Mayiga, forest near road to Boué, 2 ad. 9(2), 22.11.1962 (loc. 8, leg. Remy). — Edoungavion, at road to Boué, near Ntsibelong, 6 ad. 9(2), 1 juv. 6, 19.11.1962 (loc. 4, leg. Bernardi); ibidem, beginning of trail to Alarmintang, near small stream, under bark on soil, 4 ad. 9(£), 1 juv. 6, 2.111.1962 (loc. 15, leg. Remy). — Mbeza, secondary forest, near trail at old plan- tation, 16 ad. 9(2), 20.11.1962 (loc. 5, leg. Condé & Remy); ibidem, 9 ad. 9(2), 1 juv. 6, 20.11.1962 (loc. 6, leg. Condé & Remy). — Makokou, small valley, in dell with source, near the water intake, 2 ad. 9(2), 1 juv. 6, 18.11.1962 (loc. 48, leg. Foulé); ibidem, Loaloa, 11 ad. 99), 1 subad. 8( 2), 18.11.1962 (loc. 2, leg. Condé); ibidem, near the quay of Somifer, under brick and bark, 2 ad. 9(9), 24.11.1962 (loc. 11, leg. Remy). — Plateau Forestier d’Ipassa, primary forest, soil, 1 juv. 6, 20.V.1966 (loc. IPA3/B1, leg. Barra); ibidem, primary forest, soil, 8 ad. 99), 20.V.1966 (loc. IPA3/4, leg. Barra); ibidem, primary forest, soil, 6 ad. 9(9), 1.VI.1966 (loc. IPA/4, leg. Barra); ibidem, primary forest, burned soil, 6 ad. 99), 11.VI.1966 (loc. IPA/6C11, leg. Barra); ibidem, primary forest, burned soil, 2 ad. 9(2), 11.VI.1966 (loc. IPA/6C13, leg. Barra); ibidem, primary forest, under wood, 1 ad. 9( 9), 1 stad.?, 11.VI.1966 (loc. IPA/6C15, leg. Barra); ibidem, primary forest, litter, 1 juv. 6, 11.VII.1966 (loc. IPA/6CI1, leg. Barra); ibidem, primary forest, at foot of Palisota, 1 ad. 9(9), 11.VI.1966 (loc. IPA/6C111, leg. Barra); ibidem, primary forest, base of plant, 3 ad. 9(2), 27.VI.1966 (loc. IPA/9AN6, leg. Barra); ibidem, primary forest, at foot of fern, 11 ad. 99), 3 juv. 6, 27.VI.1966 (loc. IPA/9AN7, leg. Barra). Total number. 166 specimens. General distribution. Earlier reported from Angola (Remy, 1955a; Scheller, 1975). The species is widely and discontinuously distributed in the tropics and sub- tropics of America, Africa and south Asia. 13. Allopauropus (D.) cylindricus sp. n. Figs 61-71 Type material. Holotype: ad. 9(9), GABON, Plateau Forestier d’Ipassa, primary forest, soil, 20.V.1966 (loc. IPA3/4, leg. Barra). Paratypes: same data as for holotype, 2 ad. 9( 9). Other material. Plateau Forestier d’Ipassa, primary forest, soil, 1 juv. 6, 20.V.1966 (loc. IPA3/4, leg. Barra). - Edoungavion, at road to Booué, near Ntsibelong, 1 ad. 9(9), 19.11.1962 (loc. 4, leg. Bernardi). — Mayiga, left side of road to Booué, at trail near small stream, 1 ad. 9(£), 22.11.1962 (loc. 7, leg. Condé & Remy). — Makokou, Loaloa, 1 ad. 9(2), 18.11.1962 (loc. 2, leg. Condé). Total number. 7 specimens. Diagnosis. Some species in Allopauropus are characterized by a complicated ramification of the bothriotrica combined with a linguiform anal plate with two posterior appendages. Among them the new species is closest to A. (D.) machadoi Remy from Angola (Remy, 1955a). The 73 are almost identical, but the two species are easily distinguished by, e.g. the length of the antennal flagella [F 0.7 of Fj in A. (D.) cylindricus sp. n., F3 only a little shorter than F, in A. (D.) machadoi], the length of the setae on the tarsus of the last pair of legs [proximal seta 0.3-0.4 of the distal one in A. (D.) cylindricus sp. n., 0.75 in A. (D.) machadoi], the shape of the styli and the anal plate [styli cylindrical and lateral margins of anal plate concave anteriorly in A. (D.) 478 U. SCHELLER cylindricus sp. n., somewhat clavate styli and anal plate with convex lateral margins in A. (D.) machadoi]. There are also similiraties with A. (D.) arbusculosus Remy & Bello from Madagascar (Remy & Bello, 1960) as to the antennae, bothriotricha and pygidium. Etymology. From Latin cylindrus = roller (referring to the appendages of the collum segment). DESCRIPTION Length. (0.51-)0.52(-0.56) mm. Head (Fig. 61). Tergal setae of medium length, subcylindrical, blunt; most of them annulate, posterolateral ones striate. Relative lengths of setae, 15 row: aj = 10, a, = (13-)14; 204 row: a, = (9-)10(-11), a) = (14-)16, a3 = 14(-15); 374 row: aj = (10-)11, a, = (13-)14(-18); 4th row: aj = (10-)12, a, = 20(-24), az = 14, ag = 20(-24); lateral group (one paratype only) (Fig. 62): l, = 7, /, = 15,/3 = 24. Ratio ay/aj — a, in 1st and 4th rows (0.5-)0.6, in 294 and 314 rows 0.5. Temporal organs (1.1-)1.3(-1.4) times as long as their shortest interdistance. Neither pistil nor posterior aperture close to the posterior margin of temporal organ. Head cuticle glabrous. Antennae (Fig. 63). Segment 4 with setae p, p’ andr; p” and p” not ascertained. Setae cylindrical, p and p’ annulate, r very thin and striate. Relative lengths of setae: p = 100, p’ = (52-)59, r = (65-)88. Tergal seta p 1.7(2.1) times as long as tergal branch t. The latter subcylindrical, 1.8(-2.0) times as long as its greatest diameter and (as long as —)1.1 times as long as sternal branch s, this 1.6(-1.7) times as long as its greatest di- ameter and with its anterodistal corner distinctly truncate. Seta g somewhat thinner than p’, cylindrical, blunt, annulate, as long as (-1.4 times as long as) length of s. Relative lengths of flagella (basal segments included) and basal segments: F, = 100, bs, = 4(-5); Fy = 27(-33), bs, = 2; F3 = 71(-75), bs3 = 4. F (5.7-)6.4 times as long as t, F> and F; (1.7-)1.8 and (4.6-)4.7 times as long as s, respectively. Distal calyces with very small caps and distal part of flagella axes strongly fusiform, the one of F} almost as large as the whole sternal branch s. Globulus g 1.2(-1.3) times as long as wide and its width 0.7(-0.8) of greatest diameter of t; 9(-10) bracts present. Antennae glabrous. Trunk. Setae of collum segment (Fig. 64) simple, subcylindrical, blunt, striate, sublateral one annulate distally. Sublateral setae (2.6-)3.0(-3.3) times as long as sub- median setae; sternite process narrow, anterior half cylindrical, blunt, without incision; appendages short, very wide, cylindrical, with flat caps. Both process and appendages with very short pubescence. Setae on tergites as submedian setae on head; 4+4 setae on tergite I, 6+6 on II- IV,? on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 65) 0.3 of their interdis- tance and 0.5 of the length of pygidial setae a. Bothriotricha (Figs 66, 67). Relative lengths: 7, = 100, 7; = (104-)111(-113), T3 = 112(-119), Ty = (134-)139, T; = (156-)167. All but 7; branched, axes and branches thin. 7,, 7, and 7, similar to each others, with straight main axis and somewhat bow- shaped oblique simple branches; length of these branches reaching 0.2 of length of bothriotrix; 7, with (5-)7 branches, a little more on T> and 7,. T; bifurcate, one of the branches thin and straight, the other with thicker axis and curved. In two paratypes curved branch with a small end-swelling. Pubescence oblique-erect, hairs simple except on inner side of curved branch of T;. PAUROPODA FROM GABON 479 Fics 61-71 Allopauropus (A.) cylindricus sp. n., 61, 63-71 holotype, ad. 99), 62 paratype ad. 9(9). 61, head, median and right part, tergal view; 62, lateral group of setae; 63, left antenna, sternal view; 64, collum segment, median and left part, sternal view; 65, tergite VI, posteromedian part; 66, T;; 67, T3; 68, seta on coxa of leg 9; 69, tarsus of leg 9; 70, pygidium, sternal view; 71, anal plate, lateral view. Scale a: 64, 66-69; b: Figs 61-63, 65, 70-71. 480 U. SCHELLER Legs. Setae on coxa (Fig. 68) and trochanter of all legs simple, cylindrical, striate-annulate, blunt. Tarsus (Fig. 69) of leg 9 tapering, 3.6 times as long as its greatest diameter. Proximal seta short, curved, cylindrical, striate, blunt, 0.1(-0.2) of length of tarsus. Distal seta long cylindrical, annulate, blunt, (2.6-)3.8 times as long as proximal seta. Cuticle of tarsus glabrous. Pygidium (Fig. 70). Tergum. Posterior margin rounded. Relative lengths of setae: aj] = 100, a, = (62-)75(-77), az = (122-)125(-141), st = 63(-90). Setae thin, cylin- drical; a, straight, somewhat diverging, annulate; a, and a3 curved inwards and down- wards, annulate distally, diverging. st curved inwards, converging. Distance a, — aj (as long as —) 1.1 times as long as length of a}, distance a, — a; 1.5 times as long as dis- tance a, — a3; distance st — st (2.5-)3.2(-3.3) times as long as st and 1.8(-2.0) times as long as distance a, — aj. Sternum. Posterior margin between b, with broad indentation. Relative lengths of setae (a)=100): bj = (344-)425, b> = (95-)118. Setae tapering, striate-annulate, b, pointed, diverging, curved inwards. b,1.4(-1.5) times as long as interdistance; b, (0.5-)0.6(-0.7) of distance bj — bp. Anal plate (Figs 70, 71) linguiform, narrowest anteriorly, lateral margins (straight —) somewhat indented; plate (1.2-)1.4 times as long as broad, directed back- wards. Two posterior appendages protruding from posterosternal margin; appendages thin, straight, cylindrical, blunt, (0.7-)0.8 of length of plate. Plate and appendages glabrous (the latter with minute pubescence). 14. Allopauropus (D.) suppeditatus sp. n. Figs 72-80 Type material. Holotype: ad. 9( 2 ), GABON, Plateau Forestier d’Ipassa, primary forest, in soil, 17.VI.1966 (loc. IPA7/VM4, leg. Barra). Paratypes: GABON, Plateau Forestier d’Ipassa, primary forest, in soil, 1 ad. 9(2), 17.VI.1966 (loc. IPA7/VM2, leg. Barra); ibidem, 1 subad. 8. (2), 17.V1.1966 (loc. IPA7/VMS, leg. Barra). Other material. Edoungavion, at road to Booué, near Ntsibelong, 1 ad. 9(2), 19.11.1962 (loc. 4, leg. Bernardi). Total number. 4 specimens. Diagnosis. A. (D.) suppeditatus Sp. n. has several characters in common with A. (D.) tenuis Remy (1948b), a species widely distributed in the tropics. There are simi- larities in the general shape of the head, antennae, bothriotricha T,, 7 and 7, and in the chaetotaxy of the pygidium. A. (D.) suppeditatus sp. n. is distinguished from A. (D.) tenuis by the shape of the distal part of the antennal flagellum F, [axes only insignificantly widened in A. (D.) suppeditatus sp. n., almost ovoid in A. (D.) tenuis], the shape of the antennal globulus g [with narrow stalk in A. (D.) suppeditatus sp. n., with wide stalk in A. (D.) tenuis] and the shape of the 73 [simple and without end- swelling in A. (D.) suppeditatus sp. n., branched and with end-swelling in A. (D.) tenuis]. A. (D.) suppeditatus sp. n. has also similarities with A. (D.) cylindricus sp. n. described above. Distinctive characters are the shape of the 7; [axis simple in A. (D.) suppeditatus sp. n., bifurcate in A. (D.) cylindricus sp. n.], the length of the distal seta on the tarsus of leg 9 [twice longer than proximal seta in A. (D.) suppeditatus sp. n., 2.6-3.8 times as long as proximal seta in A. (D.) cylindricus sp. n.] and the length of the st [as long as a, in A. (D.) suppeditatus sp. n., shorter than a, in A. (D.) cylindricus sp.n.]. PAUROPODA FROM GABON 481 Fics 72-80 Allopauropus (D.) suppeditatus sp. n., holotype, ad. 9(9). 72, head, median and right part, tergal view; 73, right antenna, tergal view; 74, collum segment, median and left part, sternal view; 75, T;; 76, T3; 77, seta on coxa of leg 9; 78, seta on trochanter of leg 9; 79, tarsus of leg 9; 80, pygidium, sternal view. Scale a: Figs 74-79; b: Figs 72-73, 80. 482 U. SCHELLER Etymology. From Latin suppedito = have in abundance (referring to the branch- es on some bothriotricha). DESCRIPTION Length. 0.56(-0.72) mm. Head (holotype only) (Fig. 72). Tergal setae of medium length, blunt, striate, those in 15° row somewhat clavate, the others cylindrical. Relative lengths of setae, 15t row: a] = 10, ay = 11; 204 row: a, = 10, ay = 17, a3 = 9; 3" row: a, = 9, a, = 16; 4th row: a; = 7, a, = 10, az = 13, a, = 19; lateral group: /; = 31, I, = 24, 1; = 16. Ratio a,/a, — a; in 18 row 1.1, in 2"4 row 0.5, in 39 row 0.6 and in 4" row 0.4. Temporal organs large, about twice longer than their shortest interdistance. No pistil; posterior aperture not ascertained. Head cuticle glabrous. Antennae (Fig. 73). Segment 4 with 4 cylindrical, striate-annulate, blunt, setae, r thinnest. Relative lengths of setae: p = 100, p’ = 39(41), p’ = 25(24), r = (62)65. Tergal seta p 1.7(1.8) times as long as tergal branch r. The latter fusiform, 2.0(2.2) times as long as its greatest diameter and 1.1(1.2) times as long as sternal branch s, this 1.3(-1.5) times as long as its greatest diameter and with its anterodistal corner distinct- ly truncate. Seta g somewhat thinner than p, cylindrical, striate, blunt, 1.3 times as long as length of s. Relative lengths of flagella (basal segments included) and basal segments: F; = 100, bs, = 5(6); Fy = (26)27, bs, = 2; F3 = 65(68), bs; = 5.F; (6.0)7.1 times as long as f, F; and F; 1.5(1.7) and 3.7(4.3) times as long as s respectively. Distal calyces with small caps and distal part of flagella axes strongly widened in F;. Globulus g 1.2 times as long as wide; 13(14) bracts present; width of g 0.8(0.9) of greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment (Fig. 74) simple, cylindrical, blunt, annulate. Sublateral setae (3.5)4.3 times as long as submedian setae; sternite process triangular, anteriorly pointed, without incision; appendages obliquely cylindrical, with flat caps. Minute pubescence on anterior part of process only. Setae on tergites as posteromedian setae on head. 4+4 setae on tergite I, 6+6 on II-IV, ? on V and 4+2 on VI. Bothriotricha (Figs 75, 76). Relative lengths (holotype only): T; = 100, 7 = 102, 73 = 120, 74 = 111, 7; = 176. All with very thin axes and all but 73 and 75 branched. 7), 7, and 7, similar to each other, with straight main axis provided with several simple, bow-shaped, oblique branches, their length reaching up to 0.2 of length of bothriotrix; 7, with 10 branches, 7, and 7, similar to 7,. 73 and 75 with simple straight axis, very thin in distal half of the former and in distal 3/5 of the latter. Pubescence on 7, T>, 74 and on most proximal parts of 73 and 7; consisting of short straight oblique hairs, on distal half of 73 long, erect, whorled and partly branched hairs, on most distal part of 75 short erect hairs which at least partly may be branched. Legs. Setae on coxa (Fig. 77) and trochanter (Fig. 78) of leg 9 cylindrical, blunt, striate; seta on coxa simple, seta on trochanter furcate with thin secondary branch 0.5 of length of primary branch. More anterior setae simple, straight, blunt, without rudi- ments of secondary branches. Tarsus of leg 9 (Fig. 79) strongly tapering, 2.5(-3.0) times as long as greatest diameter. Setae on tarsus somewhat curved, cylindrical, blunt, striate; proximal seta 0.1 of length of tarsus and 0.5 of length of distal seta. Tarsus glabrous. PAUROPODA FROM GABON 483 Pygidium (Fig. 80). Tergum. Posterior margin rounded. Relative lengths of setae: a, = 100, a,= (65)70, az = 110(121), st = 94(113). Setae thin, cylindrical, striate; a, straight, somewhat diverging; a, and a3 curved inwards, somewhat diverging, st curved inwards and converging. Distance a, — a, (as long as —) 1.1 times as long as length of a,, distance a, — a, about 2.5 times as long as distance a, — az; distance st — st 1.5 times as long as st and 1.6 times as long as distance a; — aj. Tergum glabrous. Sternum. Posterior margin between bj with shallow indentation. Relative lengths of setae (aj = 10): bj = 35(42), b> = 12. bj and bp cylindrical, somewhat tapering, pointed, striate, the latter strongly diverging and curved inwards. bj 1.2(1.3) times as long as interdistance; b, 0.8(0.9) of distance dj — by. Sternum glabrous. Anal plate as long as broad, linguiform, narrowest anteriorly, lateral margins somewhat convex, posterior margin convex, with two posteriorly directed appendages, these thin, straight, cylindrical, striate, blunt, 0.5 of length of plate. Plate glabrous. 15. Allopauropus (D.) isodacintrai sp. n. Figs 81-89 Type material. Holotype: ad. 9(9), GABON, Edoungavion, at road to Booué, near Ntsibelong, 19.11.1962 (loc. 4, leg. Bernardi). Paratypes: same data as holotype, 2 ad. 9(d ,2). Other material. Mayiga, Endoumé, forest near the village, 1 ad. 9(9), 12.11.1962 (loc. 7, leg. Condé & Remy). Total number. 4 specimens. Diagnosis. A. (D.) isodacintrai sp. n. is a very close relative of A. (D.) dacintrai Scheller from Sierra Leone (Scheller, 1995). The two species are very alike as to the general chaetotaxy of the head, antennae and legs. The shape of the bothriotricha is also similar and the pygidium with the anal plate shows several similarities. The two species are well distinguished by the following characters: the shape of the axis of the distal part of the flagellae F, and F; [only somewhat widened in A. (D.) isodacintrai sp. n., very thick in A. (D.) dacintrai], the shape of the process of the collum segment [no incision anteriorly in A. (D.) isodacintrai sp. n., incised in A. (D.) dacintrai], the size of the end-swelling of bothriotrix 73 [0.1 of length of bothriotrix in A. (D.) iso- dacintrai sp. n., twice that length in A. (D.) dacintrai], the shape of the genital papillae [distal half curved inwards by a transverse fold in A. (D.) isodacintrai sp. n., papillae straight with even inner side in A. (D.) dacintrai], the st [long and striate in A. (D.) iso- dacintrai sp. n., short and glabrous in A. (D.) dacintrai] and the number of appendages of the anal plate [4 in A.(D.) isodacintrai sp. n., 2 in A. (D.) dacintrai]. Etymology. From Greek isos = like [referring to the resemblance with Allopauropus (D.) dacintrai]. DESCRIPTION Length. (0.38-)0.48 mm. Head (Fig. 81). Submedian and sublateral setae of medium length, lateral ones fairly long. Relative lengths of setae, 18 row: aj = 10, a) = 9(-10); 294 row: a, = 9, a; = 19, a3 = (16-)17; 314 row: aj = (8-)9, a, = 7(-8); 4 row: aj = 7(-8), a, = 13(-14), a; = 18, ay = 17; lateral group: /, = 30, I, = 20(-23), /3 = 19(-20). Ratio aj/aj — a, in 15t and 4th rows 0.9, in 29d row 0.4 and in 3 row 0.8. Temporal organs about as long as shortest interdistance. No pistil; probably posterior aperture at level of /j and /;. Head cuticle glabrous. 484 U. SCHELLER Fics 81-89 Allopauropus isodacintrai sp. n., 81-84, 86-89 holotype, ad. 9(2), 85, paratype, ad. 9(9). 81, head, median and right part, tergal view; 82, right antenna, sternal view: 83, collum segment, median and left part, sternal view; 84, T3; 85, genital papillae, anterior view; 86, seta on coxa of leg 9; 87, seta on trochanter of leg 9; 88, tarsus of leg 9; 89, pygidium, sternal view. Scale a: Figs 83, 85-88; b: Figs 81, 84: c: Figs 82, 89. PAUROPODA FROM GABON 485 Antennae (Fig. 82). Segment 4 with 4 cylindrical, striate, blunt setae, r very thin. Relative lengths of setae: p = 100, p’ = (52-)54, p” = (25-)27, r = 31. Tergal seta p 1.4 times as long as tergal branch 1. The latter fusiform, (2.0-)2.4 times as long as greatest diameter and (0.9-)1.1 times as long as sternal branch s, this 1.6(-1.8) times as long as its greatest diameter and with its anterodistal corner distinctly truncate. Seta q subcylindrical, striate, as long as length of s. Relative lengths of flagella (basal seg- ments included) and basal segments: F; = 100, bs, = (7-)8; Fy = 32, bs, = 2(-3); F3 = 71(-86), bs3 = 8(-9). Fy 4.5(-5.5) times as long as 7, Fy and F3 (1.5-)1.6 and 3.6(-4.2) times as long as s, respectively. Distal calyces with small caps and distal part of flagella axes only somewhat widened. Globulus g large, 1.3 times as long as wide; 16 bracts present; capsule bottom flat; width of g 1.1 times as wide as greatest diameter of 1. Antennae glabrous. Trunk. Setae of collum segment (Fig. 83) simple, striate, blunt, submedian setae somewhat clavate, sublateral setae cylindrical. Sublateral setae 2.4 times as long as submedian setae; sternite process triangular, pointed anteriorly without incision; appendages subcylindrical with flat caps. Short pubescence on appendages and anterior part of process. Bothriotricha (Fig. 84). Relative lengths of bothriotricha: T, = 100, 7, = 101(- 105), 73 = 81(-109), 7, = ?, 7; = (?98-)118(-169). All with straight, simple, very thin axes, 7], T> and T, exceedingly thin. 73 with thickest axis which most distally provided with a swelling, 4 times longer than wide and reaching 0.1 of length of bothriotrix. Pubescence of bothriotricha composed of simple, straight, short, oblique hairs, these strongest on distal half of 73 below the swelling; the latter with very short erect pubescence. Genital papillae (paratype) (Fig. 85). Subconical, distal half curved inwards, glabrous, 1.8 times as long as their greatest diameter; subapical seta 0.4 of the length of papilla. Coxal seta of leg 2 of the same shape as other coxal setae on anterior legs. Legs. Setae cylindrical blunt striate, on coxa of leg 9 (Fig. 86) simple, on trochanter (Fig. 87) furcate with the branches subequal in length. Corresponding setae more anteriorly simple, without rudiments of secondary branches. Coxal setae some- what clavate and shorter than setae on trochanter. Tarsus of leg 9 (Fig. 88) tapering, (3.5-)4 times as long as greatest diameter. Setae on tarsus cylindrical, striate, tapering distally; proximal seta short, curved, 0.1 of length of tarsus; distal seta longer, (1.5-)2.0 times longer than proximal seta. Cuticle of tarsus almost glabrous. Pygidium (Fig. 89). Tergum. Posterior margin broadly triangular behind st. Relative lengths of setae: a, = 100, a,= (80-)85, a3 = 140(-154), st = 54(-70). Setae subcylindrical, curved inwards, distal half tapering, st converging, striate, aj, a> and az almost glabrous. Distance aj — a, 0.8(-0.9) of length of a}, distance a; — a, about twice longer than distance a, — az; distance st — st (1.9-)2.0 times as long as st and (1.1-)1.3 times as long as distance a, — a}. Sternum. Posterior margin between b, with shallow indentation. Relative lengths of setae (a,=100): bj = 308(-350), b> = 61(-76). bj cylindrical, striate, blunt; by tapering, curved inwards, pointed, distal half striate. bj (1.4-)1.5 times as long as interdistance; b, 0.5 of distance bj — bp. Anal plate with concave lateral margins, posterior !/, divided into two short branches by a V-shaped incision; branches subcylindrical, distally squarely truncate. 486 U. SCHELLER Plate with 4 diverging appendages: two large terminal ones, as long as plate, clavate; and two short sternal ones, cylindrical, 0.2 of length of large appendages. Plate and appendages glabrous. 16. Allopauropus (D.) stenygros sp. n. Figs 90-98 Type material. Holotype: ad. 9(9), GABON, Île aux Singes, in the Ivindo River, 10 km downstream Makokou, primary forest, in soil, 4.VI.1966 (loc. IS2/5, leg. Barra). Paratypes: same data as holotype, 3 ad. 9( 2), (leg. Barra); ibidem, 1 ad. 94), 25.V.1966 (loc. IS1/1, leg. Barra); ibidem, 2 ad. 9( 9), 4.VI.1966 (loc. IS2/6, leg. Barra). Total number. 7 specimens. Diagnosis. Allopauropus (D.) stenygros sp. n. belongs to a group of species, which occurs in the Oriental region and tropical West Africa, and which possesses linguiform anal plates with two short posterior appendages and also similarities in the shape of the antennae. They are all incompletely known but good distinctive characters for A. (D.) stenygros sp. n. have been recognized in the following organs: in relation to À. (D.) viarti Remy (Remy, 1961) from Pondichéry (sternal antennal branch, anterior bothriotricha, genital papillae, pygidial setae aj and a); to A. (D.) nemoralis Remy (Remy, 1956b) from Madagascar (setae on leg 9, pygidial setae a, and a, appendages of anal plate); to A. (D.) lupiger Remy (Remy, 1959b) from Mauritius (bothriotricha, setae of pygidial tergum, anal plate); to A. (D.) socius Remy (Remy, 1948b) from the Ivory Coast (antennal branches, globulus g, posterior part of anal plate); and to A. (D.) minutissimus Remy (Remy, 1948b) from the Ivory Coast (tergal antennal branch, anterior bothriotricha, posterior part of anal plate). Etymology. From Greek stenygros = narrow (referring to the anterior part of anal plate). DESCRIPTION Length. (0.42-)0.43(-0.55) mm. Head. Not studied. Antennae (Fig. 90). Segment 4 with 4 blunt setae, p, p’ and p” clavate, annu- late, r cylindrical. Relative lengths of setae: p = 100, p’ = (43-)47, p” = (38-)40(-43), r = (55-)59(-64). Tergal seta p (1.8-)2.1 times as long as tergal branch r. The latter fusiform, 1.5(-1.7) times as long as its greatest diameter and (1.5-)1.7 times as long as sternal branch s, this 1.4(-1.5) times as long as its greatest diameter and with its an- terodistal corner truncate. Seta g as seta p, (1.5-)1.7 times as long as s. Relative lengths of flagella (basal segments included) and basal segments: Fj = 100, bs; = 7; F = (35-)38(-40), bs, = 3(-4): F3 = (81-)84(-88), bs; = 7. F, (5.4-)5.6 times as long as f, F, and F3 1.7 and 3.8(-4.2) times as long as s, respectively. Distal calyces small, helmet-shaped. Globulus g 1.2 times as long as wide; 6-7 bracts present, width of g (0.5-)0.7 of greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment (Fig. 91) simple, cylindrical, blunt, striate, sub- lateral one (3.5-)3.6(-3.7) times as long as submedian one; narrow anterior part of sternite process without apical incision; appendages short, with flattened caps; both process and appendages glabrous. Setae on tergites not studied. PAUROPODA FROM GABON 487 Fics 90-98 Allopauropus stenygros sp. n., 90-93, 95-98 holotype, ad. 9(2), 94, paratype, ad. 9(9). 90, left antenna, sternal view; 91, collum segment, median and left part, sternal view; 92, 7}; 93, 73: 94, genital papillae and seta on coxa of leg 2, anterior view; 95, seta on trochanter of leg 9; 96, tarsus of leg 9; 97, pygidium, sternal view; 98, anal plate, lateral view. Scale a: Figs 92-94; b: Figs 91, 95-98; c: Fig. 90. 488 U. SCHELLER Bothriotricha (Figs 92, 93). Relative lengths: T; = 100, T; = (105-)110, 73 = (105-)104(-111), 74 = 95(-109), 75 = 115(-119). All except 7; with simple axes, the lat- ter bifurcate with branching in the middle. One branch short and straight, the other curved and 0.5 of length of bothriotrix. Pubescence consisting of short simple hairs, oblique on 75, of increasing length distally and there with partly branched hairs on T, = I: Genital papillae. (Fig. 94, paratype). Twice wider than long, widest near base, conical, pointed distally, glabrous, apical seta long, 0.9 of length of papilla. Seta on coxa of leg 2 simple, somewhat clavate, annulate. Legs. Setae on coxa and trochanter (Fig. 95) of leg 9 furcate, branches thin, cylindrical, blunt, densely striate, secondary branch much shorter than primary branch. Corresponding setae on more anterior legs simple, probably without rudiment of secondary branch. Tarsus of leg 9 (Fig. 96) tapering, (2.4-)2.5 times as long as its greatest dia- meter. Setae cylindrical, somewhat curved, striate; proximal seta 0.1(-0.2) of length of tarsus, 0.4 of length of distal seta. Cuticle of tarsus glabrous. Pygidium (Fig. 97). Tergum. Posterior margin evenly rounded. Relative lengths of setae: aj = 10, a; = 4(-5), az = (12-)14(-15), st = 7(-8). a, straight, a), az and st curved inwards, a, and st cylindrical converging, striate, a; somewhat clavate, az glabrous, tapering, diverging. Distance a, — a, (1.7-)2.2 times as long as aj, distance aj — ay 3 times longer than distance a, — az; distance st — st 2.3(-2.5) times as long as st and 1.5 times as long as distance aj — aj. Tergum glabrous. Sternum. Posterior margin between b, with broad shallow bow-shaped inden- tation. Relative lengths of setae (a, = 10): bj = (48-)52, by = (20-)23(-24). Setae striate, bj tapering, b> cylindrical. bj 1.2(-1.3) times as long as interdistance; b 0.5 of distance b, — b,. Sternum glabrous. Anal plate (Figs 97, 98) directed obliquely upwards, 1.8 times as long as greatest breadth, broadest in the middle, lateral sides distinctly concave in anterior part, convex in the middle, posteriorly lengthened and sharpened to a point; two diverging clavate glabrous appendages protruding backwards from sternal side of distal part of plate, their length 0.3 of length of plate. 17. Allopauropus (D.) phakoides sp. n. Figs 99-109 Type material. Holotype: ad. 9(2), GABON, Île aux Singes, in the Ivindo River, 10 km downstream Makokou, primary forest, in soil, 4.VII.1966 (loc. IS2/3, leg. Barra). Paratypes: GABON, Belinga, at trail along the drinking-water pipe, under piece of wood, 1 ad. 9(9), 16.III. 1962 (loc. 33, leg. Condé); ibidem, 2 ad. 9(9), 22.VII.1962 (loc. 55, leg. Condé). — Ntsibelong, right bank of the Ivindo River, under bark, 2 ad. 9 ,9), 19.1.1962 (loc. 3, leg. Grassé). — Edoungavion, | subad. 9( 9), 26.11.1962 (loc. 12, leg. Condé & Remy). Total number. 7 specimens. Diagnosis. Allopauropus (D.) phakoides sp. n. is characterized by the following combination of characters: setae thin, antennal branches subequal in length, antennal globulus g with many bracts, bothriotricha with thin straight axes, pubescence on 73 longest and partly branched, pygidial setae a, short and clavate, anal plate lens-shaped with two posterior striate appendages. The species has affinities to A. (D.) eburnensis and A. (D.) vesperalis both described by Remy from the Ivory Coast (Remy, 1957b) PAUROPODA FROM GABON 489 Fics 99-109 Allopauropus (D.) phakoides sp. n., 99-103, 105-109 holotype, ad. 9(2), 104, paratype, ad. 9(d). 99, head, median and right part, tergal view; 100, right antenna, tergal view; 101, collum segment, median and left part, sternal view; 102, tergite VI, posterior part; 103, 73; 104, genital papillae and seta on coxa of leg 2, anterior view; 105, seta on coxa of leg 9; 106, seta on trochanter of leg 9; 107, tarsus of leg 9; 108, pygidium, tergal view; 109, anal plate, lateral view. Scale a: Figs 101, 103, 105-107; b: Figs 99, 102, 104, 108-109; c: Fig. 100. 490 U. SCHELLER but are easily distinguished from them by the shape of the pygidial setae aj and the anal plate. Etymology. From Greek phakos = lentil and lens of the eye (referring to the shape of the anal plate). DESCRIPTION Length. (0.54-)0.72 mm. Head (Fig. 99). Tergal setae of medium length or fairly long, thin, cylindrical, striate-annulate. Relative lengths of setae, 15 row: a, = 10, a; = (10-)12; 2nd row: ay= 9(-10), a, = 2(18-20), a3 = 11(-14); 314 row: a, = (6-)10, ay = (11-)14; 4 row: a, = (6-)8, dy = 12(-15), a3 = a, = (11-)14; lateral group: I, = (28-)29(-35), /, = (19-)20 (-22), 13 = 2(22-23). Ratio a,/a, — a, in 18 row 1.0(-1.2), 24 row 0.6(-0.8), 3™ row (0.7-)0.9, 4h row (0.6-)0.7. Temporal organs large, (2.1-)3.1 times as long as their shortest interdistance; small pistil in posterior part and small aperture at posterior mar- gin at level of /,. Head cuticle glabrous. Antennae (Fig. 100). Segment 4 with setae p, p’ p” and r; p’” not ascertained. Setae cylindrical, striate-annulate. Relative lengths of setae: p = 100, p’ = (36-)37(-43), p” = 30(-42), r = (52-)63. Tergal seta p 1.4(-1.7) times as long as tergal branch r. The latter somewhat fusiform, 2.7(-3.2) times as long as its greatest diameter and (1.1-)1.3 times as long as sternal branch s, this (1.6-)1.7(-1.9) times as long as its greatest diameter and with its anterodistal corner distinctly truncate. Seta g cylindrical, blunt, annulate, (as long as —) 1.3 times as long as length of s. Relative lengths of flagella (basal segments included) and basal segments: F; = 100, bs, = 5(-6); F = 27(-37), bs, = 2(-3); F3 = (74-)84(-87), bs3 = 5(-6). Fy (4.5-)4.8(5.1) times as long as f, F, and F3 (1.7-)1.9(-2.1) and (4.6-)5.3 times as long as s, respectively. Distal calyces with small caps and distal part of flagella axes only somewhat widened. Globulus g 1.5(-1.6) times as long as wide and its width (0.8-)0.9 of greatest diameter of r; = 12 bracts present. Antennae glabrous. Trunk. Setae of collum segment (Fig. 101) simple, subcylindrical, blunt, striate- annulate. Sublateral seta (1.8-)1.9 times as long as submedian seta; sternite process narrow anteriorly and without apical incision; appendages short, wide, cylindrical, caps flat. Both process and appendages glabrous. Setae on tergites as submedian setae on head; 4+4 setae on tergite I, 6+6 on II- IV, 6+? on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 102) (0.3-)0.4(-0.5) of their interdistance and (2.5-)2.8(-3.2) times as long as pygidial setae ay. Bothriotricha (Fig. 103). Relative lengths: T,; = 100, T, = 137(-150), 73 = (101- )103(-114), 74 = 102(-121), 75 = 172(-182). All with straight and very thin axis. Pubescence consisting of short, simple, oblique hairs on T,, T>, 74 and Ts but hairs of increasing length distally and there partly branched hairs on 73. Genital papillae (Fig. 104, paratype). Conical, narrowing in distal 2/3, 2.1 times as long as their greatest diameter, glabrous; distal seta long, thin, almost 0.6 of length of papilla. Legs. Setae on coxa and trochanter of leg 9 cylindrical, striate, blunt; coxal seta (Fig. 105) simple, seta on trochanter (Fig. 106) furcate with secondary branch thin and 0.6 of length of primary branch. Corresponding setae on more anterior legs, (including PAUROPODA FROM GABON 491 setae on coxa of leg 2 in male), simple, cylindrical, without rudiments of secondary branch. Tarsus of leg 9 (Fig. 107) tapering, 3.1(-3.5) times as long as its greatest diameter. Setae striate-annulate, proximal seta short, tapering, curved, (0.2-)0.3 of length of tarsus; distal seta cylindrical, striate, about as long as proximal seta. Cuticle of tarsus glabrous. Pygidium (Fig. 108). Tergum. Posterior margin rounded. Relative lengths of setae: aj = 10, a, = (20-)22(-25), az = 28(-33), st = (22-)27. Setae thin, striate, converging, a, short, straight, clavate, a), az and st somewhat curved inwards, a; pointed. Distance a, — a, (3.0-)4.5 times as long as aj, distance a, — a, about as long as distance a, — az; distance st — st 2.2(-3.1) times as long as sf and (as long as —) 1.1 times as long as distance a, — a). Tergum glabrous. Sternum. Posterior margin between b, almost straight apart from small lobe with shallow median incision just below anal plate. Relative lengths of setae (a,=10): bj = 125(-134), b> = 48(-57). Setae tapering, striate, b diverging, somewhat curved inwards. bj (1.3-)1.4(-1.5) times as long as interdistance; b, as long as (— 1.2 times as long as) distance dj — bp. Anal plate (Figs 108, 109) narrowest anteriorly, subcircular, lens-shaped, (1.0-)1.1 times as long as greatest breadth, directed backwards-upwards. Two posterior appendages protruding backwards from its posterosternal part; they are thin, straight, cylindrical, striate, 0.6 of length of plate. Plate and sternum glabrous. 18. Allopauropus (D.) bovistellus sp. n. Figs 110-118 Type material. Holotype: ad. 9(6 ), GABON, Plateau Forestier d’Ipassa, primary forest, in soil, 1.VI.1966 (loc. IPA4/AVC3, leg. Barra). Total number. 1 specimen. Diagnosis. A. (D.) bovistellus sp. n. is unique in the odd shape of its antennal globuli, the shape of the genital papillae and in the short ring-shaped tibiae of legs 2-8 and the stalk of the apical organ of the tarsi. Etymology. From Greek bovista = puffball (referring to the shape of antennal globulus). DESCRIPTION Length. 0.77 mm. Head. Only partly available for study. Tergal setae cylindrical, blunt, striate. Relative lengths of setae, 15t row: a, = a, = 10; 229 row: aj = 9, a, = 15, az = 7; 359 row: a, and a, = ?; 4th row: a, = 7, ay = 15, a; = 12, ag = 20; lateral group: J; = 1, = 20, /, = 19. Temporal organs have neither pistil nor posterior aperture. Antennae (Fig. 110). Segment 4 with at least 3 short annulate setae, probably p’, p” and r. Their length 5, 2 and 3 um respectively. Tergal branch almost cylindrical, 1.5 times as long as its greatest diameter and as long as sternal branch s, this 1.1 times as long as its greatest diameter and with its anterodistal corner somewhat truncate. Seta annulate, 1.1 times as long as length of s. Relative lengths of flagella (basal segments included) and basal segments: F, = 100, bs, = 9; Fy = 23, bs) = 4; F3 = 70, bs3 =7. Fy 5.4 times as long as t, F, and F; 1.3 and 3.8 times as long as s, respectively. Distal calyces with very small caps and distal part of flagella axes with fusiform widening in 492 U. SCHELLER 113 20 pm DR “TS Fics 110-118 Allopauropus (D.) bovistellus sp. n., holotype, ad. 9(¢). 110, left antenna, sternal view; 111, col- lum segment, median and left part, sternal view; 112, 73; 113, genital papillae and basal part of leg 2, anterior view; 114, seta on coxa of leg 9; 115, seta on trochanter of leg 9; 116, tarsus of leg 9; 117, tibia and tarsus of leg 8 with claw and empodium; 118, pygidium, sternal view. Scale a: Figs 114-117; b: Figs 110-113, 118. F, and F;, in the shape of an inverted cone in F,. Globulus g 1.3 times as long as wide, in shape of a puffball with proximal half almost cylindrical; = 17 very thin bracts; capsule fungiform; width of g 1.2 times as long as greatest diameter of t. Antennae glabrous except for basal segments of flagella, which have distinct pubescence arranged in whorls. PAUROPODA FROM GABON 493 Trunk. Setae of collum segment (Fig. 111) simple, cylindrical, blunt, striate- annulate. Sublateral seta 2.6 times as long as submedian seta; sternite process narrow, without anterior incision; appendages short, wide, with flat caps. Process with minute pubescence. Setae on tergites not studied. Bothriotricha. Axes very thin except in 7; (Fig. 112). The latter thin proximal- ly but widens to a longish club, probably with distal thin lengthening. Genital papillae (Fig. 113). Small, conical, pointed, protruding from large bases, these as long as papillae; seta 0.3 of length of papilla. “Coxal” seta of leg 2 inserted on lateral part of base of papilla. Seta cylindrical, with distinct end-swelling, striate. Legs. Setae on coxa and trochanter of all legs simple, blunt, striate-annulate. Seta on coxa (Fig. 114) and trochanter (Fig. 115) of leg 9 cylindrical, annulate; coxal seta longest. Tarsus of leg 9 (Fig. 116) cylindrical, tapering in distal third, 2.8 times as long as its greatest diameter. Setae straight, short, proximal seta pointed, 0.1 of length of tarsus and 0.5 of length of cylindrical distal seta. Cuticle of tarsus glabrous. Tibiae of legs 2-8 short, annulate (Fig. 117). Claw and empodium on distinct stalk, in leg 8 0.3 of length of tarsus. Pygidium (Fig. 118). Tergum. Posterior margin between a, almost straight. Relative lengths of setae: a, = 10,a,= ?, a3 = 11, st= ?. aj and a; straight, cylindrical, almost glabrous, the latter diverging. a, lacking and st not identified, may be very short. Distance a, — a, twice longer than aj, distance aj — a, considerably shorter than distance a> — az; distance st — st 0.3 of distance a, — a). Sternum. Posterior margin between D, straight, with small median V-shaped incision. Relative lengths of setae (aj = 10): bj = ?, b> = 11. bp cylindrical, blunt, striate, almost as long as distance bj — b;. Anal plate narrowest anteriorly, about as broad as long, posterior margin some- what indented in the middle, posterolateral corners rounded; two diverging appendages protruding backwards from submedian part of posterior margin; appendages lanceo- late, curved outwards, diverging, about as long as plate. Sternum glabrous. Subgenus Perissopauropus Scheller, 1997 Key to the species of Perissopauropus 1 Stalk of antennal globulus g thick, subcylindrical; setae bj of pygidial stemumilanceolate?. Ue ra Ey IN ET AND RN as PAR tridens Scheller - Stalk of antennal globulus g thin, conical; setae bj of pygidial sternum thinstaperino RRSSAA AE Url DIRSI RE Re i 2 2 Antennal setae u well developed, cylindrical ............. bounourei Remy - Antennal'setacizziudimentaryCONICALERE tt TNA II NE ce 3 3 Pygidial setae a, and a, thin, tapering, pointed; pubescence on pygidial st long, decreasing in length outwards ............... amphikomos Scheller - Pygidial setae a, and a, thick, cylindrical, blunt; pubescence on pygi- dial st distinct but not long, and all of the same length ...... lambdoides sp. n. 494 U. SCHELLER 19. Allopauropus (Perissopauropus) lambdoides sp. n. Figs 119-130 Type material. Holotype: ad. 9(9), GABON, Plateau Forestier d’Ipassa, primary forest, at base of plant, 27.VI.1966 (loc. IPA9/AN6, leg. Barra). Paratypes: GABON, Edoungavion, under bark on soil, 2 ad. 9( 9), 21.11.1962 (loc. 5, leg. Condé). — Mbeza, secondary forest, near trail at old plantation, 1 subad. 8(9), 1 juv. 6, 20.11.1962 (loc. 5, leg. Condé & Remy); Other material. Plateau Forestier d’Ipassa, primary forest, at base of plant, 1 ad. 9(@), 1 subad. 8(2), 1 juv. 6, 27.VI.1966 (loc. IPA9/AN2, leg. Barra); ibidem, at base of fern, 1 juv. 6, 27.VI.1966 (loc. IPA9/AN3, leg. Barra). - Mbeza, secondary forest, near trail at old plantation, 1 subad. 8( 2), 20.11.1962 (loc. 5, leg. Condé & Remy). — Edoungavion, at road to Booué, 1 ad. 9(3), 19.11.1962 (loc. 4, leg. Bernardi); ibidem, under bark, 1 subad. 8(£), 1 juv. 6, 20.11.1962 (loc. 12, leg. Condé & Remy); ibidem, at foot of big tree, under bark on soil, 1 ad. 9(9), 2.111.1962 (loc. 12bis, leg. Condé). — Mvadhi, end of trail to Dubost forest, under stones in laterite, 1 subad. 8(2), 1 juv. 6, 9.IX.1962 (loc. 101, leg. Condé). — Mayiga, Endoumé, right side of road to Booué, at trail near small stream, 2 ad. 9(£), 12.11.1962 (loc. 7, leg. Condé & Remy); ibidem, 1 juv. 5, 22.11.1962 (loc. 8, leg. Condé & Remy). — Belinga, under moss-covered stone, 1 ad. AG), 17.111.1962 (loc. 35, leg. Condé & Bernardi). Total number. 20 specimens. Diagnosis. Three species have been described in the subgenus Perissopau- ropus: A. (P.) bounourei Remy from Cameroon (Remy, 1955b), A. (P.) tridens Scheller from Angola (Scheller, 1975) and A. (P.) amphikomos Scheller from Central Amazonia (Scheller, 1997). They form, together with the species described below, a homogenous group which has many characters in common. The new species is distinguished from A. (P.) bounourei by the shape of the antennal seta u (short and conical in A. (P.) lambdoides sp. n., longer and cylindrical in A. (P.) bounourei), the pygidial seta a; (thick, straight, blunt in A. (P.) lambdoides sp. n., thin, curved inwards, tapering, pointed in A. (P.) bounourei) and by the shape of the anal plate (branches slender and tergal appendages curved inwards in A. (P.) lamb- doides Sp. n., branches short and tergal appendages straight in A. (P.) bounourei). A.(P.) lambdoides sp. n. is distinguished from A. (P.) tridens by the shape of the antennal globulus g [stalk thin in A. (P.) lambdoides sp. n., thick in A. (P.) tridens), the shape of the antennal setae and the pubescence of the antennae (setae pointed, antennal stalk, branches and globulus g almost glabrous in A. (P.) lambdoides sp. n., setae blunt, with distinct pubescence in A. (P.) tridens], by the pubescence on the bothriotrix Ts [sparse and depressed hairs in A. (P.) lambdoides sp. n., dense and hairs oblique in A. (P.) tridens], by the shape of the st [curved inwards and converging in A. (P.) lamb- doides sp. n., straight, directed backwards in A. (P.) tridens], by the pubescence on the pygidial setae a3 and st [fairly dense pubescence of short hairs in A. (P.) lambdoides sp. n., a few long hairs in A. (P.) tridens], by the shape of the pygidial setae bj [curved, tapering, with distinct pubescence on one side only in A. (P.) lambdoides sp. n., straight, lanceolate, with very dense short pubescence on all sides in A. (P.) tridens] and by the shape of the appendages of the anal plate [distinctly tapering in A. (P.) lamb- doides sp. n., subcylindrical and blunt in A. (P.) tridens]. A. (P.) lambdoides sp. n. is distinguished from A. (P.) amphikomos by: the length proportion p/p’ [1.2 in A. (P.) lambdoides sp. n., 1.5 in A. (P.) amphikomos], the occurrence of seta p’” [rudimentary knob in A. (P.) lambdoides sp. n., not visible in A. (P.) amphikomos], the shape of the antennal globulus g [distinct stalk in A. (P.) lamb- doides sp. n., very short stalk in A. (P.) amphikomos], the shape of the seta r on the 4t antennal segment [pointed in A. (P.) lambdoides sp.n., blunt and distal part annulate in PAUROPODA FROM GABON 495 Q K In. à 2 0 m i ie ik 2 2 a u " \ a R / \ i N b \ IN rer ey C ——— ——m@ Fics 119-124 Allopauropus (Perissopauropus) lambdoides sp. n., holotype, ad. 9(2). 119, head, median and right part, tergal view; 120, right antenna, sternal view; 121, collum segment, median and left part, sternal view; 122, tergite VI, posterior part; 123, Tj; 124, 73. Scale a: Figs 121, 123-124; b: Fig. 122; c: Figs 119-120. A. (P.) amphikomos], the shape of the setae on the pygidial tergum [a, and a, straight, cylindrical, blunt in A. (P.) lambdoides sp. n., curved inwards, tapering in A. (P.) amphikomos], the st [curved, pubescence hairs of about the same length in A. (P.) lamb- 496 U. SCHELLER doides sp. n., almost straight, pubescence hairs decrease in length outwards in A. (P.) amphikomos], and the length of the submedian appendages of the anal plate [<0.5 of length of plate in A. (P.) lambdoides sp. n., >0.5 of length of plate in A. (P.) amphikomos]. Etymology. From the Greek letter lambda (referring to the shape of the anal plate). DESCRIPTION Length. (0.77-)0.97(-1.02) mm. Head (Fig. 119). Tergal and lateral setae long, tapering, all terminated by a thin straight hair and covered with a very distinct pubescence of somewhat curved oblique hairs. Relative lengths of setae, 15 row: a, = 10, a, = (10-)11; 2nd row: a, = (10-)12, a, = (13-)15, az = (12-)13; 3rd row: aj = (12-)15, a, = (10-)12(-13); 4th row: aj = (13-)19, a, = (15-)19, az = (18-)20(-25), ag = (12-)15(-16); lateral group: / = 15(-16), I, = (10-)12, 13 = (15-)19. Ratio aj/aj — aj in 18 row 1.4, 24 row (0.5-)0.6, 314 row 1.3 and 4th row (0.9-)1.2(-1.3). Length of temporal organs 0.9 of their shortest inter- distance. No interior pistil present but a clavate vesicular appendage projecting back- wards from a point somewhat anterior to posterior margin, length of appendage 0.4 of length of temporal organ. Posterior aperture not ascertained. Head cuticle glabrous, temporal organs with vesicle possessing sparse minute pubescence. Antennae (Fig. 120). Segment 4 with 5 distal setae, these tapering, pointed, from base and outwards pubescent-striate-annulate, r thinnest. Relative lengths of se- tae: p = 100, p’ = 83(-84), p” = (37-)53, r = (22-)23, u = (3-)5. Tergal seta p 0.9(-1.1) times as long as tergal branch r. The latter slender, cylindrical, distally obliquely trun- cate, (4.0-)4.4(-4.9) times as long as its greatest diameter and as long as sternal branch s, this subcylindrical, 3.0(-4.1) times as long as its greatest diameter and with its anterodistal and posterodistal corners equally truncate. Seta g similar to p”, 0.5(-0.6) of length of s. Relative lengths of flagella (basal segments included) and basal seg- ments: Fj = 100, bs, = (8-)10; F, =74(-78), bs, = (7-)9; F3 = 69(-71), bs3 =7. F 2.3 times as long as ¢, F and F3 1.6(-1.7) and 1.5(-1.7) times as long as s, respectively. Distal calyces of F, and F3 very small; distal part of flagella axes with a distinct sub- globular swelling just below calyx. Globulus g pyriform, (1.6-)1.7(-1.8) times as long as wide; 12 bracts present; width of g (0.7-)0.8 of greatest diameter of +. Antennae glabrous except for bracts of globuli, these with a very distinct pubescence, and the base segments of the flagella, which have a minute pubescence. Trunk. Setae of collum segment (Fig. 121) furcate, main branch tapering, point- ed, with oblique pubescence; secondary branch rudimentary, cylindrical, blunt, glabrous. Sternite process broad, pointed anteriorly. Appendages wide, with flat caps. Sparse short pubescence on anterior part of process and appendages. Setae on tergites as on tergal side of head, somewhat increasing in length pos- teriorly. 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V, 4+2 on VI. Posterior setae on tergite VI (Fig. 122) 1.2 times as long as interdistance and somewhat longer than length of pygidial setae a). Bothriotricha (Figs 123, 124). Relative lengths: T, = 100, 7, = 107(-113), T3 = 117(-137), T4 = (169-)197, 7; = (206-)210(-253). All with simple straight axes; PAUROPODA FROM GABON 497 Fics 125-130 Allopauropus (Perissopauropus) lambdoides sp. n., 126-130, holotype, ad. 9(9), 125, paratype, ad. 94). 125, genital papillae and seta on coxa of leg 2, anterior view; 126, seta on coxa of leg 9; 127, seta on trochanter of leg 9; 128, tarsus of leg 9; 129, pygidium, sternal view; 130, anal plate, lateral view. Scale a: Fig. 128; b: Figs 125-127; c: Figs 129-130. pubescence hairs strong, straight, simple, strongest on 73 and Ts, erect on distal half of T, and T,, otherwise oblique. Genital papillae (Fig. 125, paratype). Conical, 2.4 times as long as their greatest diameter, glabrous, distinctly narrowing and curved inwards in distal half; seta 0.5 of length of papilla. 498 U. SCHELLER Legs. Setae on coxa (Fig. 126) and trochanter (Fig. 127) of leg 9 simple, sub- cylindrical, cleft distally, with distinct oblique pubescence, seta on trochanter 1.4 times as long as coxal seta and also more slender than seta on coxa. More anterior setae simple, not cleft distally and without rudiments of secondary branches except on coxal seta on leg 2 in males, this with a cylindrical, blunt, glabrous, rudimentary, secondary branch. Tarsus of leg 9 (Fig. 128) slender, straight, tapering, 5.6(-6.8) times as long as greatest diameter. Setae on tarsus very dissimilar: proximal seta straight, tapering, pointed, with long distal hair and a few long depressed more proximal hairs, seta (0.3-)0.4 of length of tarsus and 3.9(-4.7) times as long as distal seta; the latter short, 0.1 of length of tarsus, fork-shaped as a trident with straight spinous prongs, the middle one longest. Pubescence coarse, arranged in rows lengthways, hairs on tergal side long and somewhat increasing in length toward proximal end, there glabrous between proximal seta and the upper end of tarsus. Pubescence on lateral and sternal sides shorter and denser but distinct. Pygidium (Fig. 129). Tergum. Posterior margin straight, with evenly rounded corners. Relative lengths of setae: aj = 100, a, = (51-)71, az = (104-)147, st = (43-)59. Setae of two types: a, and a, thick, straight, cylindrical, blunt, glabrous; a; and st tapering, pointed, filiform distally, curved inwards and with distinct oblique pu- bescence, strongest on st. Distance a, — a; (0.5-)0.8 of length of a}, distance a, — a; (1.4-)2.5 times as long as distance a, — az; distance st — st 1.2(-1.3) times as long as st and about as long as distance a, — aj. Cuticle of tergum sparsely covered with long spinous pubescence most posteriorly. Sternum. Posterior margin between bj straight apart from shallow indentation just below anal plate. Relative lengths of setae (a,=100): bj = (82-)94(-130), by = (46-)62(-66). bj tapering, glabrous on one side, pubescence increasing in length dis- tally; b> curved inwards, diverging, with long distal hair. bj 0.9 of (— as long as) inter- distance, by 1.2(-1.3) times as long as distance bj — bp. Anal plate (Figs 129, 130) Y-shaped, with its prongs 1.7 times as long as their common base; distal part of each prong terminated by a tapering blunt extension, 0.7 of length of prong; on sternal side of each extension a spine very similar to st but smaller. Plate and appendages with distinct oblique pubescence; cuticle of sternum with long sparse pubescence. Stage subad. 8. Setae dj and d> tapering, pointed, with oblique pubescence; d, 1.4 times as long as distance dj — dj and twice longer than d). Genus Cauvetauropus Remy, 1952b Subgenus Nesopauropus Scheller, 1997 20. Cauvetauropus (N.) pistillifer sp. n. Figs 131-139 Type material. Holotype: ad. 9( 2 ), GABON, Plateau Forestier d’Ipassa, primary forest, in soil, 20.V.1966 (loc. IPA3/b4, leg. Barra). Total number. 1 specimen. Diagnosis. C.(N.) pistillifer sp. n. may be closest to C. (N.) subtilis Scheller from Sri Lanka (Scheller, 1970). They are very alike as to the general shape of the antennae and the bothriotricha, but can be easily distinguished by: the shape of the PAUROPODA FROM GABON 499 136 137 Fics 131-139 Cauvetauropus (N.) pistillifer sp. n., holotype, ad. 9(9). 131, posterior part of temporal organ with aperture and lateral group of setae; 132, right antenna, sternal view; 133, collum segment, median and left part, sternal view; 134, tergite VI, posterior part; 135, 73; 136, seta on coxa of leg 9; 137, seta on trochanter of leg 9; 138, tarsus of leg 9; 139, pygidium, sternal view. Scale a: Fig. 133; b: Figs 131, 132, 134-138; c: Fig. 139. 500 U. SCHELLER tergal antennal branch [3.3 times as long as its greatest width in C. (N.) pistillifer sp. n., 2.1 in C. (N.) subtilis], the shape of the setae on coxa and trochanter of leg 9 [simple in C.(N.) pistillifer sp.n., furcate in C. (N.) subtilis], the shape of the setae on the tarsus of leg 9 [proximal seta pointed, 0.4 of the length of the tarsus in C. (N.) pistillifer sp. n., blunt, 0.5 of the length of the tarsus in C. (N.) subtilis], the shape of the setae a, of the pygidial tergum [short, bladder-shaped in C. (N.) pistillifer sp. n., long, cylindrical in C. (N.) subtilis], and the shape of the anal plate [subcircular with long appendages in C. (N.) pistillifer sp. n., subrectangular with short appendages in C. (N.) subtilis Scheller]. Etymology. From Latin pistillum = club-shaped pounder and ferre = carry (referring to the setae a, of pygidial tergum). DESCRIPTION Length. 0.85 mm. Head. Only partly available for study. Tergal setae cylindrical, blunt, striate. Relative lengths of setae, 15 row: a, = a, = 10; 229 row: a, = 9, a, = 15, az = 7; 3 row: aj and ay = ?; Ath row: a; = 7, a, = 15, a3 = 12, aq = 20; lateral group: I, = 1; = 20, I, = 19. No pistil; posterior aperture (Fig. 131) close to posterior margin of tem- poral organ at level of /; and /;. Antennae (Fig. 132). Segment 4 with 4 thin, cylindrical, striate-annulate setae, p’ and r very thin. Relative lengths of setae: p = 100, p’ = 35, p” = 27, r = 61. Tergal seta p 1.3 times as long as tergal branch r. The latter subcylindrical, 3.3 times as long as its greatest diameter and 1.3 times as long as sternal branch s, this1.4 times as long as its greatest diameter and with its anterodistal corner distinctly truncate. Seta q similar to p, 1.4 times as long as length of s. Relative lengths of flagella (basal segments included) and basal segments: F = 100, bs, = 4; F, = 28, bs, = 3; F3 = 81, bs; = 4. F 5.2 times as long as r, Fy and F3 1.9 and 5.7 times as long as s, respecti- vely. Distal calyces with very small caps and distal part of flagella axes strongly widened, almost ovoid and with many discs. Globulus g 1.3 times as long as wide; 17 bracts present; width of g as long as greatest diameter of r. Antennae glabrous. Trunk. Setae of collum segment (Fig. 133) simple, cylindrical, blunt, striate. Sublateral seta 2.2 times as long as submedian seta; sternite process narrow, triangular, without anterior incision; appendages short, wide, caps flat. Process with minute pu- bescence in anterior half. Setae on tergites cylindrical, blunt, striate. 4+2 setae on tergite VI (Fig. 134); posterior setae there 0.4 of interdistance and 3.7 times as long as pygidial setae a]. Bothriotricha (Fig. 135). Axes very thin and simple. T, and 7, not studied. Lengths of other trichobothria: 73 = 98, 74 = 70 and 7; = 135 um. Pubescence on all except 73 very short, erect distally; 73 with short simple oblique hairs in proximal half, these longer towards distal part, there also arranged in whorls and branched. Legs. Setae on coxa and trochanter of all legs simple, blunt, striate. Seta on coxa of leg 9 (Fig. 136) somewhat clavate and 0.7 of length of cylindrical seta on trochanter (Fig. 137). Tarsus of leg 9 (Fig. 138) tapering, 3.1 times as long as its greatest diameter. Setae pointed, proximal seta long, somewhat curved, with short oblique pubescence, 0.4 of length of tarsus and 1.5 times as long as distal striate seta. Cuticle of tarsus glabrous. PAUROPODA FROM GABON 501 Pygidium (Fig. 139). Tergum. Posterior margin rounded, with shallow indenta- tion between st. Relative lengths of setae: a, = 10, a, = 23, a3 = 40, st = 33. a, short, subglobular, a, somewhat clavate, a3 and st subcylindrical, blunt; a, and a3 somewhat curved inwards, st converging. Distance a, — a, 5.7 times as long as aj, distance a, — a, 2.5 times as long as distance a, — az; distance st — st 2.5 times as long as st and 1.5 times as long as distance a, — a,. Tergum glabrous. Sternum. Posterior margin between b; with broad shallow indentation. Relative lengths of setae (a, = 10): bj = 190, b, = 62. dj and b; tapering distally, pointed, striate, the latter curved inwards. b,1.4 times as long as interdistance; b, about as long as distance b, - bo. Anal plate narrowest anteriorly, short, 1.3 times as broad as long, linguiform, lateral margins convex, posterior margin with shallow sternal incision; each posterior lobe with a cylindrical, blunt, shortly striate and posteriorly directed appendage, 0.9 of length of plate. Sternum with anal plate glabrous. Genus Hemipauropus Silvestri, 1902. Subgenus Hemipauropus s. str. 21. Hemipauropus (H.) elongatus sp. n. Figs 140-151 Type material. Holotype: ad. 9(d ), GABON, Plateau Forestier d’Ipassa, primary forest, in soil, 7.VI.1966 (loc. IPAS/ES, leg. Barra). Paratype: ibidem, 1 ad. 9(¢), 7.VI.1966 (loc. IPAS/E16, leg. Barra). Other material. Plateau Forestier d’Ipassa, primary forest, in soil, 1 juv. 6, 7.VI.1966 (loc. IPAS/E17, leg. Barra). - Edoungavion, under bark on soil, 1 ad. 94), 21.11.1962 (loc. 12, leg. Condé & Remy). Total number. 3 specimens. Diagnosis. Hemipauropus (H.) elongatus Sp. n. is well delimited from the other species of the genus by the very long and slender tarsi of the last pair of legs and the very thin setae a3 on the pygidial tergum. It may be closest to H. (H.) angolanus Remy from Angola (Remy, 1955a) and H. (H.) obrei Remy from Mauritius (Remy, 1959b). Good distinctive characters in relation to H. (H.) angolanus are the length of the antennal setae p and p’ [p>p’ in A. (H.) elongatus sp. n., p [long and subcylindrical in A. (H.) elongatus sp. n., short and conical in 7. (H.) angolanus] and the shape of the setae a; of the pygidial tergum [very thin distally in 7. (H.) elongatus sp. n., tapering and blunt in 7. (H.) angolanus Remy]. The new species is distinguished from H. (H.) obrei as follows: the antennal globulus g is short-stalked and 1.4-1.5 times as long as its greatest diameter in H. (H.) elongatus sp. n., long-stalked and 2.5 times as long as its greatest diameter in H. (H.) obrei; the apical organ of the antennal flagella F, and F> are cylindrical in A. (H.) elongatus sp. n., subovoid in H. (H.) obrei and the anal plate is more slender, 4.5 times as broad as long in H. (H.) elongatus sp. n., 2.5 times in A. (H.) obrei. Hemipauropus (H.) elongatus sp. n. has also similarities with H. (H.) leonensis Scheller from Sierra Leone (Scheller, 1995) but that species has a much shorter and U. SCHELLER 502 el PAUROPODA FROM GABON 503 proportionally wider stalk of the antennal globulus g, proportionally shorter seta g and tarsi, and the seta a3 on the pygidial tergum is thicker and has a distal hair. Etymology. From Latin elongatus = prolonged (referring to the tarsi). DESCRIPTION Length. 0.62(-0.69) mm. Head, holotype only (Fig. 140). Tergal setae of 15 row and a, in 24 row of medium length, other tergal and lateral setae fairly long; main part of them leaf-shaped, glabrous or with very minute pubescence; az in 24 and 4 rows thin, striate, the former cylindrical, blunt, and the latter tapering, pointed. Relative lengths of setae, 15 row: ay = 10, ay = 11; 204 row: a, = 13, a, = 18, az = 16; 374 row: a, = 16, a, = 17; 4th row: aj = 16, a, = 15, a3 = 21, aq = ?; lateral group: J, = 16, I, = 20, I; = 40. Ratio a,/a, — aj in 15 row 1.1, 2"4 row 0.6, 3"4 row 1.6 and 4" row 0.8. Length of temporal organs about as long as shortest interdistance. No pistil but very small aperture at posterior margin of temporal organs (Fig. 141). Head cuticle glabrous. Antennae (Fig. 142). Segment 4 with 5 cylindrical, blunt, striate setae, p some- what widened distally. Relative lengths of setae: p = 100, p’= 85(90), p” = 35(40), r = 35(39), u = 5. Tergal seta p 0.9 of length of tergal branch r. The latter fusiform, 2.9(3.1) times as long as greatest diameter and (0.8)0.9 of length of sternal branch s, this 2.3(2.4) times as long as greatest diameter and with anterodistal corner distinctly trun- cate. Seta q thinner than p and p’, cylindrical, blunt, striate, (0.7)0.8 of length of s. Relative lengths of flagella (basal segments included) and basal segments: Fj = 100, bs, = (14)16; F, = (54)60, bs, = (11)13; F3 = (94)95, bs; = (13)14. F and F3 thinner than F}. F 2.7(3.0) times as long as t, Fy and F3 (1.4)1.5 and 2.4 times as long as s, respectively. Distal organs of Fj and F, subcylindrical, 2.5 times as long as greatest diameter and consisting of slightly curved and flattened bracts around a sessile ovoid capsule; distal organ of F almost twice longer and also thicker than that of F,; those of F, small and subhemispherical, without surrounding bracts. Globulus g 1.5 times as long as wide, with thin stalk; 10 bracts present; width of g 0.8 of greatest diameter of t. Antennae glabrous except for basal segments of flagella, these with a minute pubescence. Trunk. Setae of collum segment (Fig. 143) furcate, main branch leaf-shaped, broadest in the middle, with minute pubescence; secondary branch rudimentary, cylin- drical; these setae 4 times longer than their greatest diameter; sublateral seta 1.7 times as long as submedian seta; sternite process short, broad, triangular, not cleft distally, glabrous; appendages cylindrical with wide and flat caps, the latter minutely pubescent. Fics 140-151 Hemipauropus (H.) elongatus sp. n., holotype, ad. 9(d). 140, head, median and right part, tergal view; 141, part of temporal organ and lateral group seta /,; 142, right antenna, sternal view; 143, collum segment, median and left part, sternal view; 144, tergite VI, posterior part; 145, T3; 146, genital papillae and seta on coxa of leg 2, anterior view; 147, seta on coxa of leg 9; 148, seta on trochanter of leg 9; 149, tarsus of leg 9; 150, pygidium, sternal view; 151, anal plate, lateral view. Scale a: Fig. 143; b: Figs 140-141, 144-149; c: Fig. 142; d: Figs 150-151. 504 U. SCHELLER Setae on tergite I as submedian setae on head, narrower more posteriorly. Anterior tergites transversely divided, I and II distinctly so, III incompletely so; 4+4 setae on I, 6+6 on II-V and a single row of 4 setae on VI (Fig. 144). Reticular cuticle pattern very weak, visible on tergites III-VI only. Bothriotricha (Fig. 145). Relative lengths (holotype only): 7} = 100, 7, = 78, T3=97,Ty=?,T5= 174. All with very thin axes and minute pubescence; axes of 73 thickest. Genital papillae (Fig. 146). Basal segments short but well developed, with nar- row collar. Papillae twice longer than greatest diameter, slowly tapering distally, inner side almost straight, outer side strongly curved in distal third. Distal seta very short and thin. Setae on coxa of leg 2 not deviating from other coxal setae of anterior legs. Legs. Setae on coxa (Fig. 147) and trochanter (Fig. 148) of leg 9 furcate; primary branch leaf-shaped, with subparallell lateral margins, 3.7 times as long as greatest width; secondary branch clavate, much shorter and thinner, about 0.6 of length of main branch. Both branches minutely pubescent. Corresponding setae more anteriorly with rudimentary secondary branches. Tarsus of leg 9 (Fig. 149) long, tapering, very slender, (6.7)8.0 times as long as greatest diameter. Both setae tapering, pointed, almost glabrous; proximal seta 0.1(0.2) of length of tarsus and 1.3(1.9) times as long as distal seta. Cuticle of tarsus almost glabrous. Pygidium (Fig. 150). Tergum. Posterior margin with rounded median bulge car- rying small but distinct posteromedian extension. Relative lengths of setae: a, = 100, dy = 110, az = (189)194, st = ?. Setae glabrous, aj and a; lanceolate, broadest in proximal third, az thin, tapering, pointed, a; and a; curved inwards. sf very small or lacking. Distance aj — a; 1.3(1.9) times as long as a, , distance a, — a, twice longer than distance a; — az; distance st — st 0.8 of distance a, — ay. Sternum. Margin between bj with a broad median pointed triangular lobe pro- jecting backwards below anal plate. Relative lengths of setae (aj = 100): bj = 210(260). Neither b, nor b3 present. bj proportionately short, 0.7 of interdistance, tapering, glabrous, somewhat curved inwards. Anal plate (Figs 150, 151) glabrous, with broad base with two thin, almost straight posterolateral spines and a posteromedian, unusually thin, forked appendage, 3 times longer than breadth of plate at base. 22. Hemipauropus (H.) bilobatus sp. n. Figs 152-160 Type material. Holotype: ad. 9(9), GABON, Edoungavion, beginning of trail to Alarmintang, near small stream, under wood, 2.111.1962 (loc. 15, leg. Condé). Paratypes: same data as for holotype, 2 ad. 96 ©). Other material. Edoungavion, under bark on soil, 3 ad. 9(29 ,1sex ?), 2.II.1962 (loc. 15bis, leg. Condé & Remy). Total number. 6 specimens. Diagnosis. The large posterolateral lobes on the pygidial sternum have not been found in other species of the genus. Etymology. From Latin bis = two and lobus = rounded projection (referring to the posterior part of the pygidial sternum). PAUROPODA FROM GABON 505 Fics 152-160 Hemipauropus (H.) bilobatus sp. n., 152-155, 157-160, holotype, ad. 9(2), 156 paratype, ad. 9(d). 152, head, median and right part, tergal view; 153, left antenna, sternal view; 154, collum segment, median and left part, sternal view; 155, tergite VI, posterior part; 156, genital papillae and seta on coxa of leg 2, anterior view; 157, seta on trochanter of leg 9; 158, tarsus of leg 9; 159, pygidium, tergal view; 160, anal plate, lateral view. Scale a: Figs 152, 154-158; b: Figs 153, 159-160. 506 U. SCHELLER DESCRIPTION Length. (0.98-)1.20(-1.21) mm. Head, holotype only (Fig. 152). Tergal setae fairly long; main part of them leaf- shaped, with short dense pubescence; a3 in 2nd and 4th rows thin, pointed, striate. Relative lengths of setae, 1°! row: a; = 10, a, = 12-13; 2nd row: a, = 13, a, = 18-19, az =18; 314 row: a] = 13, a, = 14; 4" row: a, = 15,a, = 13,a3 = ?, ay = ?; lateral group: I, = 22, 1, = 30, 13 = 21. Ratio ay/aj — aj in 15 row 1.2, 299 row 0.5, 3'd row 1.6 and 4th row 1.0. Length of temporal organs 0.8 of shortest interdistance. No pistil but very small aperture at posterior margin of temporal organ. Head cuticle glabrous. Antennae (Fig. 153). Segment 4 with 3 striate setae, p and p’ cylindrical, blunt, r subcylindrical, tapering, pointed, p” a rudimentary knob only. Relative lengths of setae: p = 100, p’ = (85-)87, r = (46-)51. Tergal seta p (0.8-)0.9 of length of tergal branch r. The latter fusiform, 3.9(-4.6) times as long as greatest diameter and 0.7(-0.8) of length of sternal branch s, this (3.5-)4.2 times as long as greatest diameter and with anterodistal corner distinctly truncate. Seta g thinner than p and p’, subcylindrical, tapering, striate, 0.5(-0.6) of length of s. Relative lengths of flagella (basal segments included) and basal segments: F; = 100, bs, = 10(-11); F = (52-)54(-55), bs, = 10; F; = (96-)112, bs; = (12-)14. F| 2.4 times as long as 7, F, and F3 0.9 and (1.8-)1.9 times as long as s, respectively. Distal organs of F; and F> similar, subcylindrical, somewhat campanulate, 2.1 times as long as greatest diameter and consisting of thin, slightly curved bracts around a sessile ovoid capsule; distal organ of F3 small and subhemi- spherical, without surrounding bracts. Globulus g 1.6(-1.8) times as long as wide with conical stalk; = 10 bracts present; width of g 0.6 of greatest diameter of t. Antennae glabrous. Trunk. Setae of collum segment (Fig. 154) furcate, main branch leaf-shaped, broadest in the middle, with minute pubescence; secondary branch rudimentary, cylin- drical; these setae 3.8-5.8(-5.9) times as long as greatest diameter; sublateral seta 1.3(-1.4) times as long as submedian seta; sternite process short, broad, triangular; not cleft distally, anterior part distinctly pubescent; appendages widest proximally; caps flat, wide, glabrous; base segments with minute pubescence. Setae on tergite I as submedian setae on head, more posteriorly narrower. 4+4 setae on I, 6+6 on I-IV, 4+2 on V and a single posterior row of 4 setae on VI. Submedian setae on VI (Fig. 155) lanceolate, broadest in proximal part, almost glabrous, 0.3 of interdistance. Reticular cuticle pattern very weak. Bothriotricha. Axes very thin, pubescence very short, T,, 7, and 7, thinnest. Relative lengths: T, = 100, 7, = = 100,73 = (141-)168, 7, = 150(-153), Ts = (237-)288. Genital papillae (Fig. 156). Conical, twice longer than greatest diameter; distal seta short, thin, 0.2 of length of papilla. Setae on coxa of leg 2 not deviating from other anterior coxal setae. Legs. Setae on coxa and trochanter of leg 9 (Fig. 157) furcate; primary branch leaf-shaped, with subparallell lateral margins, seta on trochanter 5.4 times as long as greatest width; seta on coxa proportionately broader; secondary branch shorter and thinner than primary branch, somewhat clavate, about 0.7 of length of main branch; distinct oblique pubescence on both branches. Corresponding setae on more anterior legs with rudimentary secondary branches. Tarsus of leg 9 (Fig. 158) slender, tapering, PAUROPODA FROM GABON 507 5.4(-5.7) times as long as greatest diameter. Setae minutely pubescent, proximal seta thin, tapering, pointed; distal seta short, clavate. Proximal seta 0.2 of length of tarsus and (3.6-)4.1 times as long as distal seta. Cuticle of tarsus with faint pubescence. Pygidium (Fig. 159). Tergum. Posterior part of pygidium broadly triangular, margin between st with two small submedian lobes separated by a rounded indentation. Relative lengths of setae: aj = 10, a; = (14-)15, a3 = 23(-27), st = 2(-3). Setae almost glabrous, a, and a, lanceolate, broadest in proximal third, somewhat curved inwards; az tapering, strongly curved inwards. st short, lanceolate. Distance a, — a; (1.1-)1.2 times as long as a, distance a, — a; (2.3-)2.8 times as long as distance a, — a3; distance st — st (1.2-)1.3 times as long as distance a, — a]. Sternum. Margin between b, with one large median and two small submedian posterior bulges. Lateral margins just outside bj with large lobes projecting posteriorly; lobes blunt, with convex sides. Relative lengths of setae (a, = 10): bj = (26-)29. Neither b, nor b3 present. bj almost straight, tapering, minutely pubescent, (0.8-)0.9 of interdistance. Anal plate (Figs 159, 160) with somewhat coarse surface, broad base with two short thin almost straight diverging posterolateral spines and a long and broad forked posteromedian appendage, 2.2 times as long as breadth of plate at base. Genus Polypauropus Remy, 1932 23. Polypauropus afrioccidentalis Scheller Polypauropus afrioccidentalis Scheller, 1995: 41-43, figs 177-188. Material examined. GABON, Plateau Forestier d’Ipassa, primary forest, in soil, ad. 9(9), 17.VI.1966 (loc. IPA7/VMA, leg. Barra). Total number. 1 specimen. Taxonomic remarks. Compared with the type material from Sierra Leone, the specimen from Gabon has proportionately long pygidial setae a, and bj, somewhat larger setae t, and the posterior margin of the small process between the anal plate setae has two glabrous posterior lobes instead of an undulated margin with some pubescence hairs. General distribution. Previously known only from the locus typicus in Sierra Leone (Scheller, 1995). REMARKS ON THE WEST AFRICAN PAUROPODA FAUNA Many species were previously known from tropical West Africa, from Senegal in the north to Angola in the south. Remy has reported the main part by the describing of material from the Ivory Coast (1948b, 1952a, 1953, 1957b), Congo (1954, 1956a, 1962), Angola (1955a), Cameroon (1955b), Senegal (1957c), Gambia (1958a) and Guinea (1959a). The rest of the species have been described by Scheller, who has studied material from Angola (1975) and Sierra Leone (1995). From these papers, and with addition of the species reported here, a list of 111 identified species can be put together, about 15% of the world-fauna known up to now. Most of them, 91species or 82% of the known West African fauna, have not been collected elsewhere, indicating both a high diversity and a high degree of endemism. Those 20 species found outside 508 U. SCHELLER tropical West Africa have mostly ranges which include the islands of the Indian Ocean and/or south Asia, but seldom North Africa and never South Africa. A few species have extremely large distribution ranges (Allopauropus sphaeruliger Remy, A. bouini Remy and A. proximus Remy) and may belong to an old subcosmopolitan element, and some of the new species show similarities with species now living far away. For example Allopauropus singesensis sp. n. has relations to species in Paraguay and Thailand, A. stenygros Sp. n. to species in the Oriental region, A. lambdoides sp. n. to species not only in tropical Africa but also in Brazil, Cauvetauropus pistillifer sp. n. seems to be related to congeners in Sri Lanka and Hemipauropus elongatus sp. n. not only to African species but also to a species on Mauritius. ACKNOWLEDGEMENTS The author is indebted to the collectors, with special attention to the late Professor B. Condé, Nancy, and Dr Jean-A. Barra, Obernai, for sending me the material which formed the basis of this paper. REFERENCES HANSEN, H. J. 1902. On the genera and species of the order Pauropoda. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 1901: 323-424, pls. 1-6. REMY, P. A. 1932. Un pauropode de Banyuls-sur-Mer type d’une famille nouvelle: Polypauropus duboscqi nov. gen., nov. sp. Archives de Zoologie expérimentale et générale 74: 287- 303. REMY, P. A. 1948a. Pauropodes de l’ Afrique orientale britannique. Proceedings of the Zoological Society of London 118: 568-574. REMY, P. A. 1948b. Pauropodes de la Côte d’Ivoire, Afrique occidentale française. Mémoires du Muséum national d'Histoire naturelle Paris (n.S.) 27: 115-151. Remy, P. A. 1950. Les Millotauropus, types d’un nouveau groupe de Pauropodes. Comptes rendus des séances de l’Académie des Sciences Paris 230: 472-473. REMY, P. A. 1952a. Pauropodes récoltés par M. Maurice Vuillaume en Côte d’Ivoire. Bulletin de la Société entomologique de France 57(4): 49-53. Remy, P. A. 1952b. Contribution a la faune endogée du Sahara. Pauropodes. Bulletin de la Société zoologique de France Paris 77: 51-61. REMY, P. A. 1953. Pauropodes de Côte d’Ivoire (nouvelles récoltes de M. Maurice Vuillaume). Bulletin de la Société Zoologique de France 78(1): 28-33. Remy, P. A. 1954. Pauropodes d’Afrique tropicale. Jn: SCHOUTEDEN, H. (ed.). Miscellane Zoologica. Annales du Musée Royal du Congo Belge, Bruxelles, Tervuren C- Zoologie 1: 249-256. REMY, P. A. 1955a. Pauropodes de |’ Angola. Publicagoes culturais da Companhia de Diamantes de Angola, Lisboa 24: 115-134. Remy, P. A. 1955b. “Allopauropus bounourei” Pauropode nouveau du Cameroon. Archives d’Anatomie, d’Histologie et d’Embryologie 37(2, 4/8): 131-134. REMY, P. A. 1956a. Contribution à l'étude de la microfaune endogée de 1’ Afrique tropicale: Palpigrades et Pauropodes. II. Pauropodes du Congo Belge. Revue de Zoologie et de Botanique africaines 53: 327-335. REMY, P. A. 1956b. Pauropodes de Madagascar. Mémoires de l’Institut scientifique de Madagascar (A)10: 101-229. Remy, P. A. 1956b. Pauropodes de l’île de la Réunion. 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SCHELLER, U. 1994. Pauropoda of a secondary forest near the Taruma Mirim River, Amazonas, Brazil (Myriapoda, Pauropoda, Pauropodidae). Amazoniana 13: 65-130. SCHELLER, U. 1995. First record of Pauropoda (Myriapoda) from Sierra Leone. Entomologica scandinavica, Supplementum 47: 1-53. SCHELLER, U. 1997. Pauropoda from upland and inundation forests in Central Amazonia, Brazil (Myriapoda, Pauropoda: Millotauropodidae, Pauropodidae). Amazoniana 14: 223-300. SCHELLER, U. 2001. Pauropoda (Myriapoda) from Sabah (East Malaysia) (Pauropoda and Symphyla of the Geneva Museum XI.) Revue suisse de Zoologie 108: 949-986. SILVESTRI, F. 1902. Ordo Pauropoda. Jn: BERLESE, A. (ed.). Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Portici. Li Im ca LS EEN dae ALe lez i! ‘7 " 9 INSEE (Vi opie b D aad i BE LE (aie > i : inn aa à lil ANT de le if el Ji it, CAS TER | u : ) vad y : E = REVUE SUISSE DE ZOOLOGIE 112 (2): 511-517; juin 2005 Two new taxa of Galagete (Lepidoptera, Autostichidae) from the Galapagos Islands, Ecuador Patrick SCHMITZ & Bernard LANDRY ©) Muséum d’histoire naturelle, C. P. 6434, CH-1211 Geneva 6, Switzerland. E-mail: patrick.schmitz@mhn.ville-ge.ch / bernard.landry @mhn.ville-ge.ch Two new taxa of Galagete (Lepidoptera, Autostichidae) from the Galapagos Islands, Ecuador. - A new species and a new subspecies of the genus Galagete (Lepidoptera, Autostichidae) from the Galapagos Islands are described and illustrated. Galagete griseonana sp. n., known only from males, is endemic to the island of Santa Cruz. Galagete pecki flavofasciata ssp. n., known from both sexes, occurs on the islands of Santa Cruz and Santiago, where it is also believed to be endemic. Keywords: Micro moths - Autostichidae - new species - new subspecies - endemic - Galapagos Islands. INTRODUCTION Autostichidae are gelechioid moths characterised by the gnathos being an articulated band with an unarticulated mesial hook, and by the presence of spiniform setae in a band across the abdominal terga. The caterpillars feed on dead or decaying plant or animal tissue (Hodges, 1998). With a total of 13 species (one of which is not described), the genus Galagete Landry represents the first documented case of an extensive radiation of endemic Lepidoptera in the Galapagos Islands comparable in size to the radiation of the Darwin’s finches with some species confined to one single island (Landry, 2002). The second largest radiation in Galapagos Lepidoptera found so far contains only three species (genus Utetheisa Hübner, Arctiidae). For the purpose of enabling recognition and documentation of the elements of the Galagete radiation, and because we were able to obtain sufficient material in early 2004, we are describing below a new species of Galagete and a new subspecies of Galagete pecki Landry. MATERIAL AND METHODS The 51 specimens forming the basis of this study were collected by ourselves mostly during one expedition to the Galapagos in March-April 2004. Three of the specimens were collected by the second author in 1989 and 1992. The moths were collected at light, either with an ultra-violet light or a mercury vapour light suspended next to a white sheet. (1) Corresponding author Manuscript accepted 01.11.2004 512 P. SCHMITZ & B. LANDRY In listing the label data of the holotypes, the information is copied as found on the labels with slashes to express changes of lines, and abbreviations spelled out in square brackets, except “m” for “meters.” As regard the lists of paratypes, the speci- mens' data are listed first in alphabetical order of island collected and then in order of dates collected, the information is recorded without indications of line changes, the abbreviations, except for distances, "GPS" (= Global Positioning System), and cardi- nal points, are spelled out only once at first encounter, collecting localities are report- ed without accented letters, dates are standardised, and collectors' information is stan- dardised and placed in parentheses. For each taxon’s holotype the data label is printed in black on white card stock while the holotype label is hand-written in black ink on red card stock. Other than the above mentioned, the following acronyms are used: BMNH for The Natural History Museum (London, England), CNC for Canadian National Collection of Insects, Arachnids, and Nematodes (Ottawa, Ontario, Canada), CDRS for Charles Darwin Research Station (Santa Cruz Island, Galapagos), MHNG for "Muséum d'histoire naturelle de Genève" (Geneva, Switzerland), and USNM for National Museum of Natural History (Washington, D. C., USA). Genitalia were dissected after the abdomen had macerated in a cold 20% KOH solution overnight. The dissected parts were kept in lactic acid stained with orange G for description purposes. They were subsequently stained with chlorazol black and mounted on slides in Euparal. The forewing lengths were measured with a reticule on a stereomicroscope. The illustrations of the moths and genitalia were made with the AutoMontage® system using a JVC® video camera mounted on a Leica MZ APO stereomicroscope or a Zeiss Axioskop compound microscope. Illustrations of all Galagete species can be viewed on the web site of the MHNG at www.geneva- city.ch/musinfo/mhng/. RESULTS Our expedition to the Galapagos in 2004 brought only one new island record for previously described Galagete species: G. turritella Landry is also present on San Cristobal Island. Contrary to what is recorded in Landry (2002, p. 848) the holotype of G. consimilis Landry is not dissected and in the diagnosis for G. consimilis (same page of Landry, 2002), the 314 line should read “G. darwini” instead of “G. consimilis”. One of the paratype females of G. seymourensis Landry was wrongly associated with slide number BL 1344; the correct slide number is BL 1343. DESCRIPTIONS Galagete griseonana sp. n. Figs 1, 3-5 Galagete sp.; Landry, 2002: 819, 820, fig. 15. Holotype d, [1] “ECU[ADOR], Galapagos, Santa Cruz/ C[harles] D[arwin] R[esearch] S[tation]/ base of El Barranco/ GPS: S 00°44.305’ W 90°18.105’/ 18.11.2004, u[ltra] v[iolet] I[ight]/ leg/it]. B[ernard]. Landry, P[atrick]. Schmitz”. [2] *HOLOTYPE/ Galagete/ griseonana/ Schmitz & Landry”. Specimen in perfect condition deposited in the MHNG. Paratypes, Ecuador: 30 & , from the island of Santa Cruz, Galapagos Islands, collected at UVL by B. Landry and P. Schmitz, unless specified otherwise. 1 4 (dissected, slide MHNG 2703), CDRS, arid zone, 3.11.1989, M[ercury] V[apour] L[amp] (B. Landry); 2 é, with same TWO NEW TAXA OF GALAGETE 513 data as holotype; 7 & (one dissected, slide BL 1596), CDRS wall of Invert[ebrate]s. Lab[ora- tory]., GPS: S 00°44.478’ W 90°18.132’, 19.111.2004; 20 3 (one dissected, slide BL 1595), same locality but 6.iv.2004. Deposited in the BMNH, CNC, CDRS, MHNG, and USNM. Diagnosis. Among the species of Galagete, G. griseonana can be distinguished by its very small size (wingspan: 5.7-7.1 mm) and by its forewing appearing uniform- ly grey-brown. This combination of size and colour is unique. Some specimens of Galagete darwini Landry (wingspan: 7.0-9.0 mm) and Galagete pecki Landry (wingspan: 7.5-8.7 mm) are only slightly larger, but their background colour is brown or beige respectively with distinct markings (see Landry, 2002, figs 7, 8, and 13). Galagete cinerea Landry is also a grey-brown species, but it is much larger (wingspan: 8.5-11.2 mm) and has darker markings (see Landry, 2002, fig. 14). Description. MALE (n=31) (fig. 1). Head grey-brown with whitish beige scales around eye. Haustellum and maxillary palpus whitish beige. Labial palpus grey-brown on first segment; second segment whitish beige medially and dorsally, laterally grey- brown except for apical whitish beige ring laterally and ventrally; third segment grey- brown ventrally, whitish beige dorsally. Antennal scape with few whitish beige scales apically; flagellum grey-brown. Thorax dorsally grey-brown except for shining whitish grey-brown metascutellum. Foreleg coxa grey-brown at base, whitish beige apically; femur, tibia, and tarsomeres mostly grey-brown with whitish beige at apex of tibia and apex of tarsomere I and V. Midleg femur whitish beige with grey-brown on dorsal edge apically; tibia pale grey-brown with whitish beige apically and on spurs; tarsomeres I-V mostly whitish beige with grey-brown at base of each segment. Hindleg whitish beige. Male wingspan: 5.7-7.1 mm (Holotype: 6.8 mm). Forewing grey-brown, with sometimes slightly darker grey-brown markings as small spots submedially and in cubital fold and postmedially at the end of cell; fringe grey-brown. Hindwing pale greyish brown, fringe pale whitish beige. Abdomen whitish beige, without modified scales. Male genitalia (n=3) (figs 3-5). Basal half of uncus only slightly angled from second half; second half not produced dorsally, apical margin only slightly concave; arms slightly laterally compressed, triangular, short, apically rounded; dorsal crests broadly rounded, poorly demarcated. Median hook of gnathos rather short and thick, very slightly upturned and pointed apically. Dorsal connection of tegumen wide; pedunculi short and broad. Lateral arms of transtilla moderately long, broad, rounded, median surface with fan-shaped scales towards base and setae mostly apically, edges setose; median arm as long as or slightly longer than lateral arms, very narrow for whole length. Valva of medium length and width, dorsal margin angled ventrally very gently before apex; ventral margin angled dorsally from about 1/3 and with subbasal notch; apex broadly rounded; costa melanised from base to about 2/3; sacculus rather wide, of medium length, apically flattened and blunt. Juxta symmetrical, somewhat heart shaped, with rather shallow and broadly rounded median notch. Vinculum bulbous, short and rounded, not projecting dorsoapically. Aedeagus narrow, slightly arched, larger at base with very short coecum penis; apical 1/3 open ventrally, dorsal wall slightly bent to right, narrowly rounded; vesica with abundant spicules, without cornuti. FEMALE. Unknown. 514 P. SCHMITZ & B. LANDRY Etymology. From the Latin griseus, grey, and nanus, a dwarf, referring to the colour of the forewing and the size of this species. Biology. Unknown although moths are attracted to light in the arid zone between February and April. Distribution. Currently known only from the Galapagos island of Santa Cruz; presumed to be endemic to the archipelago and possibly to Santa Cruz. Remarks. This species had been recognised as new by Landry (2002) but was not described then because only one damaged specimen was available. Based on our collecting experience, the available sample being made of males only may reflect a sexually-linked seasonality pattern, or that the females are flightless, or that they are not attracted to light, or that their flying period each night is later than that of the males. It remains to be tested. Galagete pecki flavofasciata ssp. n. Fig. 2 Galagete sp.; Landry, 2002: 866, fig. 36 E, F. Holotype d, [1] “ECU[ADOR], Galapagos, Santa Cruz/ C[harles] D[arwin] R[esearch] S[tation], wall of Invert[ebrate]s. Lab[oratory]./ GPS: elev[ation]. 11 m, S 00°44./478° W 90°18.132’ 6.1v.2004/ ufltra] v[iolet] l[ight]/ leg/it]. B[ernard]. Landry, P[atrick]. Schmitz”. [2] ’HOLOTYPE/ Galagete/ pecki flavofasciata/ Schmitz & Landry”. Specimen complete except for small notch at apex of left forewing and few segments missing on left flagellum. Deposited in the MHNG. Paratypes, Ecuador: 9 dg, 10 2 from the Galapagos Islands, collected at UVL by B. Landry and P. Schmitz, unless specified otherwise. Santa Cruz: 1 d,3 ®, transition zone, recently cut road, GPS: S 00°42.528° W 90°18.849’, 12.11.2004; 1 d, 1 © , low agriculture zone, GPS: S 00°42.132° W 90°19.156°, 13.11.2004; 6 d (one dissected, slide BL1600), 2 2 (one dissected, slide BL 1601), CDRS base of El Barranco, GPS: S 00°44.305° W 90°18.105’, 18.11.2004; 1 &, 1 2, CDRS wall of Inverts. Lab., GPS: elev[ation]. 11 m, S 00°44.478’ W 90°18.132’, 19.iii.2004; 1 2 (dissected, slide BL 1306), Finca Vilema, 2 km W [of] Bella Vista, 1.iv.1992, M[ercury] V[apour] L[amp] (B. Landry); 1 9, with same data as holotype. Santiago: 1 2, Aguacate, 520 m elev., 7.iv.1992, M[ercury] V[apour] L[amp] (B. Landry). Deposited in the BMNH, CNC, CDRS, MHNG, and USNM. Diagnosis. Among the species of Galagete, G. pecki flavofasciata can be easi- ly distinguished by its yellowish-orange (or whitish beige) forewing markings. Only Galagete pecki pecki, G. levequei Landry, and Galagete cristobalensis Landry are similar in wing markings (see Landry, 2002, figs 10-13), but the background colour of their forewings is white to beige or pale greyish brown. Description. MALE (n=10) (fig. 2). Head whitish beige to yellowish-orange with few thin dark brown scales along posterior eye margin. Haustellum and maxillary palpus whitish beige. Labial palpus greyish brown, shining on first segment; whitish beige on second segment sometimes with brown scales laterally at base; third segment whitish beige except for pair of usually connected blackish brown spots on second half laterally and ventrally and with small blackish brown spot ventrally at base. Antennal scape and flagellum blackish brown. Thorax dorsally blackish brown at base to 1/5-1/3, less so on tegula, whitish beige to yellowish orange in middle including most of tegu- la and lateroposteriorly, dark greyish brown at apex medially; metascutellum greyish brown, shining. Foreleg coxa whitish beige; femur blackish brown with whitish beige at apex; tibia blackish brown with whitish beige at apex, base, and small patch medially; tarsomere I blackish brown with whitish beige at apex; tarsomeres II-IV TWO NEW TAXA OF GALAGETE 515 Fics 1-2 Holotypes of Galagete species. 1. Galagete griseonana sp. n.; 2. G. pecki flavofasciata ssp. n. blackish brown. Midleg femur and tibia as in foreleg, tibial spurs whitish beige; tarsomere I, II and V blackish brown with whitish beige at apex; tarsomere III and IV blackish brown. Hindleg femur whitish beige; tibia greyish brown with whitish beige apically, spurs whitish beige; tarsomere I, II, and V with whitish beige at apex; tarsomere III and IV blackish brown. Wingspan: 6.0-6.9 mm (Holotype: 6.5 mm). Forewing blackish brown with whitish beige to yellowish-orange as small basal spot, subbasal band of variable width sometimes separated by line of blackish brown scales in cubital fold, two small spots of variable shape, sometimes connected, one on coastal margin between 2/5 and 1/2 and one slightly larger on inner margin at 1/2, and a trans- verse band of variable width postmedially from costa to below cell or to inner margin; fringe dark greyish brown. Hindwing and fringe greyish brown. Abdomen greyish brown, shining, without modified scales, but whitish beige on valvae. Male genitalia (n=1) (fig. 30 A-D in Landry, 2002) as in Galagete pecki pecki. FEMALE. (n=10). Colour as in males but often with more extensive whitish beige to yellowish orange markings. Antenna slightly thinner than those of males. Forewing length: 6.4-8.2 mm. Female genitalia (n=2) (fig. 36 E, F in Landry, 2002) as Galagete pecki pecki (fig. 32 A, B in Landry, 2002). Other dissected specimen differs from that illustrated on fig. 36 E, F (Landry, 2002) in having more broadly rounded lobes of sternum VIII, more broadly emarginated apical margin of tergum VIII, and more distinct scobination on membrane posterad ostium. Both dissected specimens differ from G. p. pecki in having a more simple signum, without smaller spines at base of lateral spines, and no scobination on corpus bursae. Etymology. From the Latin flavus, yellow, and fascia, band, referring to the colour and pattern of the forewings. Biology. Unknown except for the fact that adults come to light and fly from March to April in habitats between near sea level to 520 m in elevation. Distribution. Presently known only from the islands of Santa Cruz and Santiago, the subspecies is presumed to be endemic to the Galapagos. 516 P. SCHMITZ & B. LANDRY Fics 3-5 Male genitalia of Galagete griseonana sp. n. 3. Whole genitalia without aedeagus (slide BL 1595); 4. Aedeagus in dorsal view (slide BL 1595); 5. Aedeagus in lateral view (slide BL 1596). Remarks. The absence of noticeable differences in the genitalia was the first reason we decided to recognise this species as part of Galagete pecki. However, the male (6.0-6.9 mm) and female (6.4-8.2 mm) wingspans of Galagete pecki flavofasciata are smaller than those of the respective sexes (7.5-8.7 and 8.2-8.3 mm) of G. pecki pecki, which is known only from Isabela Island. These differences in wing size and forewing coloration in addition to their allopatry were the reasons we decided to describe the G. pecki specimens collected on the islands of Santa Cruz and Santiago as a distinct subspecies (G. pecki flavofasciata). We were also comforted in this decision when we found that a specimen of G. pecki flavofasciata had a 2.7 % sequence diver- TWO NEW TAXA OF GALAGETE 517 gence in a 1.4-kb fragment of the Cytochrome Oxidase I (COI) gene with G. p. pecki (P. Schmitz, unpublished). This amount of intraspecific divergence in this gene is within the range of results obtained in other Lepidoptera studies (Landry er al., 1999). In comparison, the smallest amount of interspecific divergence in the same fragment of COI between Galagete species that are easily recognised on the basis of genitalic characters is 4.9 % (P. Schmitz, unpublished). Hopefully these subspecies of Galagete pecki will not be united by anthropogenic factors that would alter their evolutionary paths. ACKNOWLEDGEMENTS We are very thankful to the authorities of the Galapagos National Park and those of the Charles Darwin Research Station for allowing fieldwork and for logistical support in 1989, 1992, and 2004. We are especially grateful to Lazaro Roque-Albelo and family for their hospitality and logistical help in 2004. The comments of L. Roque- Albelo and Ole Karsholt on our manuscript were also appreciated. This work was carried out with the financial support of a “Bourse Augustin Lombard,” the MHNG, and the University of Geneva to P. Schmitz, and the MHNG to B. Landry. REFERENCES HODGES, R. W. 1998. The Gelechioidea (pp. 131-158). Jn: KRISTENSEN, N. P. (ed.). Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol. 1: Evolution, Systematics, and Bio- geography. Walter de Gruyter, Berlin & New York, x + 491 pp. LANDRY, B. 2002. Galagete, a new genus of Autostichidae representing the first case of an extensive radiation of endemic Lepidoptera in the Galapagos Islands. Revue suisse de Zoologie 109: 813-868. LANDRY, B., POWELL, J. A. & SPERLING, F. A. H. 1999. Systematics of the Argyrotaenia francis- cana (Lepidoptera: Tortricidae) species group: Evidence from mitochondrial DNA. Annals of the Entomological Society of America 92: 40-46. REVUE SUISSE DE ZOOLOGIE 112 (2): 519-539; juin 2005 The Western Palaearctic species of Stenomicra Coquillett (Diptera, Periscelididae, Stenomicrinae), with description of a new species of the subgenus Podocera Czerny Bernhard MERZ! & Jindfich ROHACEK? 1 Muséum d'histoire naturelle, c.p. 6434, CH-1211 Genève 6, Switzerland. E-mail: bernhard.merz@mhn.ville-ge.ch 2 Department of Entomology, Silesian Museum, Masarykova 35, CZ-746 46 Opava, Czech Republic. E-mail: rohacek@szmo.cz The Western Palaearctic species of Stenomicra Coquillett (Diptera, Periscelididae, Stenomicrinae), with description of a new species of the subgenus Podocera Czerny. - Stenomicra (Podocera) soniae sp. n. is described from Central and Southeast Europe, and compared with morpho- logically similar species. The type material of Stenomicra (Podocera) deli- cata (Collin, 1944) is revised and the species redescribed. Both European species of Podocera Czerny are illustrated and their biology and distribu- tion reviewed with a number of new records. A key to the four Western Palaearctic species of Stenomicra is presented. Keywords: Periscelididae - Stenomicra (Podocera) - new species - Western Palaearctic. INTRODUCTION The systematic position of Stenomicra Coquillett was for a long time subject to controversial opinions since Coquillett (1900) originally placed it in the Drosophilidae. The genus was later included in the Anthomyzidae (Collin, 1944; Sabrosky, 1965), Asteiidae (Malloch, 1927), Aulacigastridae (Hennig, 1971; Sabrosky, 1975; Cogan, 1976; Irwin, 1982; Teskey, 1987) or treated as separate family, Stenomicridae (Rohäëek, 1983, 1987, 1997; Papp, 1984, 2001; Khoo & Sabrosky, 1989; Andersson, 1991, Tschirnhaus, 1992; Bächli, 1997, 1998; Chandler, 1998; Rohacek & Bartäk, 2001; Merz, 2002; Drake, 2004). McAlpine (1978) provisionally referred the genus to the Periscelididae. This proposition is now widely accepted, often with subfamilial sta- tus given to the group (Grimaldi & Mathis, 1993; Mathis & Papp, 1998; Tschirnhaus, 1999; Freidberg & Mathis, 2002) and is therefore adopted in the present paper. Sabrosky (1975) proposed that Stenomicra may be divided into two well-distin- guished subgenera, Stenomicra s. str. and Podocera Czerny, 1929, and this division was followed by subsequent workers (Papp, 1984; Freidberg & Mathis, 2002). Chandler (1998) suggested, however, that Podocera Czerny is preoccupied by Podo- Manuscript accepted 08.11.2004 520 B. MERZ & J. ROHACEK cera Latreille, 1817. The latter spelling (Podocera, Latreille, 1817) must be conside- red an incorrect subsequent spelling of Podocerus Leach, 1814, to which Latreille ex- plicitly refers. For this reason the name Podocera Latreille does not fall within the ru- les of the Code and Podocera Czerny is the valid genus-group name (ICZN, Articles 3373tand#3379) Very little is known about the biology of Stenomicra. According to Williams (1939), larvae of a Hawaiian species were found in water-holding leaf bases of several monocotyledones. Adults of S. delicata (Collin, 1944) were collected on vegetation in an old empty pond (Collin, 1944) and subsequently recorded to be closely associated with tussocks of large Carex species (Howe & Howe, 2001; Howe et al., 2001; Drake 2004). Stenomicra cogani Irwin, 1982, was originally found in vegetation beside a small lake (Irwin, 1982), later collected in bogs and marshes from large stands of Carex paniculata (Howe & Howe, 2001), Typha (Merz, 2002), large Carex spp. [C. panicu- lata, C. elata, C. rostrata, C. acutiformis (Drake, 2004)], and on large Carex in a wa- ter pond in Northern Italy (Merz, unpublished observation). Freidberg & Mathis (2002) described S. jordanensis from specimens swept on Cyperus. These observations suggest that most Stenomicra may be associated with grass-like monocotyledones in wetland habitats and may be either phytophagous or saprophagous. Stenomicra has an almost world-wide distribution, with about 20 extant species described from all biogeographical regions (Freidberg & Mathis, 2002) and two fossil species from Dominican Amber (Grimaldi & Mathis, 1993). However, according to Freidberg & Mathis (2002) over 100 species from the Neotropical Region await description. About 10 undescribed species are deposited in the collection of the “Muséum d’histoire naturelle, Genève”, mainly from the Oriental and Australasian Regions. Evidently, these tiny flies are still very incompletely studied. Collin (1944) described the first species, S. delicata, occuring in the Western Palaearctic Region. Later, Irwin (1982) added a species from Wales, and Freidberg & Mathis (2002) a species from Israel. Andersson (1991) mentioned two probably undescribed species from Sweden but he did not describe them. During a recent field trip near Geneva three specimens of a Stenomicra (Podocera) were collected by beating tussocks of Carex. Their identification showed that previous records of S. delicata from Switzerland (Bächli, 1997, 1998), Czech Republic (Rohaéek, 1995a, 1997; Rohälek & Bartäk, 2001) and Slovakia (Papp, 1978; Rohäcek, 1983, 1986, 1987, 1995b, 1997) were misidentifications and refer to a hither- to undescribed species. To clear up the morphological limits of the two Western Palaearctic species of the subgenus Podocera a key and detailed descriptions of both species are presented below. MATERIAL AND METHODS The material which was studied in this paper is deposited in the following collections: BUB Fakultät für Biologie, Universität Bielefeld, Germany HNHM Hungarian Natural History Museum, Budapest, Hungary MHNG Muséum d’histoire naturelle, Genève, Switzerland SMO TAU UMO CGB CMB WESTERN PALAEARCTIC STENOMICRA (PODOCERA) S2i Silesian Museum, Opava, Czech Republic Entomological collection of the Tel-Aviv University, Tel-Aviv, Israel Hope Entomological Collections, University Museum, Oxford, Great Britain private collection G. Bächli, Dietikon, Switzerland private collection M. Bartak, Praha, Czech Republic In the “Material” section the Swiss cantons are indicated according to the offi- cial abbreviations: GE = Genève, JU = Jura, TI = Ticino, VS = Valais, ZH = Zürich. Terminology of morphological structures follow the first chapters in Papp & Darvas (2000) and Freidberg & Mathis (2002). Structures of antennae are according to Stuckenberg (1999). Abdominal sternites and tergites are abbreviated as “S” and “T” respectively in the descriptions. KEY TO WESTERN PALAEARCTIC SPECIES OF STENOMICRA COQUILLETT, 1900 1 Costal section between apices of veins R2+3 and R4+5 at least 3 times as long as section between R4+5 and M1+2 (Fig. 29); anterior fronto-or- bital seta much smaller and shorter than posterior fronto-orbital seta, usuallysrechnate TER subgenus Stenomicra s. str. 2 Costal section between apices of veins R2+3 and R4+5 shorter or about as long as section between R4+5 and M1+2 (Figs 4, 17); both fronto-or- bital setae subequal in length and strength, anterior seta usually inclina- LORS SAS AAD) EE RE RE SIE TSI O subgenus Podocera 3 Posterior crossvein (DM-Cu) absent; body almost entirely yellow (HIDE) CARRO ge al EE ha A dele nee a ee S. cogani Irwin, 1982 Both crossveins present; body yellow with longitudinal black vittae on scutum (laterally of line of dorsocentral setae), pleurae (anepisternum and katepisternum, Fig. 29), scutellum (laterally at base) and abdomen (paired spots on T 3 and 4, see Freidberg & Mathis, 2002, Fig. 2) D NOR CISA E AE ae nae ABORTO S. jordanensis Freidberg & Mathis, 2002 Wing (Fig. 4) without alula, CuA2 developed as a fold (cell cup partly closed); base with broad milky white crossband; pterostigma brown, distally uniformly hyaline; no gap between last two dorsocentral setae (Fig. 25); all femora and tibiae uniformly yellow (Fig. 24); basal two abdominal tergites ivory yellow, remaining tergites orange-yellow to pale brown (T 3 and 4); surstylus bilobate (Figs 8, 12); two bare, asym- metric pregonites present (Figs 10, 11); only left postgonite developed (ERRATO A S. delicata (Collin, 1944) Wing with a small alula, without trace of CuA2 (cell cup open); surface uniformly hyaline, but base of wing with very narrow milky white cross- band and both crossveins bordered with milky frame (Fig. 17); last two dorsocentral setae separated by a distinct gap (Fig. 27); hind femur and hind tibia each with a black ring (Fig. 26); abdomen more or less uni- formly brown; surstylus simple (Figs 20, 23); only right pregonite pre- sent, covered with small tubercles (Figs 19, 22); two rather large post- gonitestdeveloped (Rigs 2D)h Vari ak NERO ER I RE oe S. soniae sp. n. S22 B. MERZ & J. ROHACEK TAXONOMIC PART Stenomicra Coquillett Stenomicra Coquillett, 1900: 262. Type species: Stenomicra angustata Coquillett, by original designation. Subgenus Podocera Czerny Podocera Czerny, 1929: 93. Type species: Podocera ramifera Czerny, by original designation. Synonymy and subgeneric status with Stenomicra by Hendel, 1931: 10. Neoscaptomyza Séguy, 1938: 347. Type species: Neoscaptomyza bicolor Séguy, by original designation. Synonymy with Stenomicra (Podocera) by Sabrosky, 1975: 664. Diadelops Collin, 1944: 265. Type species: Diadelops delicata Collin, by original designation. Synonymy with Stenomicra s. lat. by Sturtevant, 1954: 560. Synonymy with Podocera by Irwin, 1982: 235. Stenomicra (Podocera) delicata (Collin) Figs 1-12, 24-25 Diadelops delicata Collin, 1944: 266. Stenomicra delicata: Sabrosky, 1965: 211; Howe & Howe, 2001:48; Howe et al., 2001: 148; Drake, 2004: 3. Stenomicra (Podocera) delicata: Irwin, 1982: 235; Papp, 1984: 62; Tschirnhaus, 1999: 171. Stenomicra (Diadelops) delicata: Chandler, 1998: 142. TYPE MATERIAL Lectotype d (here designated), Great Britain: “S. L. [= Sussex Lodge], Nmkt. [= Newmarket], dry pond, 11.7.43 [= 11.VII.1943]”, “VC-Type 645, Diadelops, deli- cata Collin 2” (UMO). Paralectotypes: 1 2 , same data; 1 , same data, but 5.VII.1942; 19, same data, but 1.VII.1942; 16, same data, but 19.VII.1942 (all UMO). This species was described based on 45 8,522 all from the same locality (Collin, 1944; Pont, 1995). Of these, the above listed 23 d and 39 2 could be exa- mined. The male which is herewith selected as lectotype is pinned on a minuten pin, in good condition with few setae on the head missing and with the minuten pin obscuring some setulae on the scutum. The other male was dissected. Its terminalia agree with Figs 1-12. ADDITIONAL MATERIAL (5 specimens) Germany: 18, NW [= Nordrhein-Westfalen], Bielefeld, Hohberge, 30.VI. 1992, F. Piichel (BUB). 19, SH [= Schleswig-Holstein], Kreis Plön, Trentmoor, between Preetz and Plön, 23.1V.1993, Malaise trap, Kassebeer (BUB). Switzerland: 13,2? 9, GE, Jussy, Prés de Villette, 475 m, 18.V.2004, B. Merz (MHNG). DIAGNOSIS This is a very small, slender species (wing length less than 2 mm) which can be distinguished from the other, more robust Western Palaearctic species of Podocera (S. soniae) by the absence of an alula, but the presence of a faint CuA2 (cell cup therefore partly closed); the broad white crossband at wing base with brown pterostigma, remaining wing surface uniformly hyaline; the grey thorax including scutellum but ventral half of pleura yellow (Figs 24, 25); all femora and tibiae entirely yellow; the basal two abdominal tergites ivory yellow, the remaining tergites orange-yellow to pale brown, and very different male terminalia. Further differences are given in Tab. 1. 523 WESTERN PALAEARCTIC STENOMICRA (PODOCERA) (17 314) Jopusys (TT 'S14) yuosaıd sıLyaunuÄse 7 (ZZ 314) osopn8ni ‘yuosord JysLı ATUO (€Z “Stq) opduns Apeorde UMOIQ AJULIOJIUN SULI NOLI UNA VII) JO pay [eseq pue Into} Jo pry) jesıde (ed AI9A :]oyoo]e ur suauroods) J981q 0) UMOIG (LI 819) oueay ayyM-Ay]TU Aq paropioq (LI 31H) UISASSOIS JO 9981) OU (LI 314) pueqsso1s oyYyM-AY[TU MOLTeU AIOA (LI 'S14) jounstp ynq ‘]ews (LI Std) 1:17 (91 ‘3t4) des peoiq Aq payeredes ‘Sur 98J9$ [BIJU290S10p OM] JS] A213 Aquuozrun MOIPA Apeorde ‘Aas (FI (314) YSIY ue) JOpim (FI 14) Juosqu (6 314) moys (OT 314) [Jews *Jusssıd yoy Aquo (TI ‘OI S31H) 22eq og “Juasaıd SLmawwÄse 7 (ZI 314) aegogıq Apesıde umomg aed 0} MOJ[2A-33ue.Jo S9)I819) JOLIOSOd ‘MOT[OA-AIOAT S9ITI9] OM] [USE MO][9A Aquuoziun UM ( (313) Sura Jo 3s91 se outre y Aquuoziun (4 (Sl) UI9ASSOI9 X9AUO9 “JUTRT (p ‘SI4) PUEGSSOIO ONYM-ANTIUI prog WIM Juosqe (p 3a) TE (£ 319) des è Aq pariedos jou ‘193u0] 19s [ENUS90SIOP Ise] Aquo MOT[9 Apenuoa ‘4018 Ajfesıop A213 Aquuopiun (7 314) spia uey) 10YSIY (7 (814) yuasaıd awopodereyq :5 onuoG]sod ‘5 opuodald 5 sn]jsIns 7D INOJO9 :V eıgn pue INUI] pury INO]O9 :] Joyfey JO QOUN :M (ND-WC ‘IN-H) SUISASSOI : MA I[99 feue JO UISJEU [UJSIP : M aseq :M einge :M ORI YIPIM : YI] :M BIOS [UHU99IOSIOP iL wunura]sidoue INO[O9 :L Wn][omos INOTOS :L suol :H Indro90 uo 989$ 1e][990]sod [fews :H "u ‘ds apiuos ‘¢ (UII0D) 2102HP 'S IMIVICYI ‘SUIM = M ‘XCIOUL = L S99] = 7] ‘PeoH = H ‘(ofeur) erjPIU9N = H ‘U9wOpgy = y :suONeIAsIqqy "u ‘ds 202008 *S pue (ppel “UNJOD) VIDIYap PAINUOUIIS Jo ASojoydiou Jo uosuedwo) ‘| ATAVI 524 B. MERZ & J. ROHACEK Fics 1-6 Stenomicra (Podocera) delicata (Collin). 1, head laterally; 2, head dorsofrontally; 3, thorax dor- sally; 4, wing; 5, abdomen dorsally; 6, postabdomen + Sth sternum ventrally (genitalia omitted). Abbreviations: ads = additional sclerite; S = Sternite; T = Tergite. (Del. J. Rohacek) WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 525 DESCRIPTION OF MALE Wing length. 1.60 mm. Body length: 1.3 mm (1.25-1.5 mm according to Collin, 1944). Head (Figs 1-2). Yellow, but thinly ash-grey microtrichose on posterior part of frons and partly on occiput posterior of compound eye; compound eye with short, but distinct interfacetal setulae; form as in S. soniae; frons bare, parallel-sided, about 1.35 times as high as wide; ocellar triangle slightly raised, in the middle of frons; face as in S. soniae, the pair of small sensillae medially more closely together; antenna as in S. so- niae; arista large, plumose, rays much longer than diameter of postpedicel, dorsally with 5, ventrally with 3 long rays; mouthparts as in S. soniae. Chaetotaxy: colour of major setae and setulae yellow to brown depending on viewing angle and illumination; 2 subequal fronto-orbital setae in anterior half of frons, anterior fronto-orbital seta procli- nate and inclinate, posterior fronto-orbital seta reclinate; 1 long, lateroclinate medial vertical seta; anteromedially with 1 slightly shorter, proclinate pseudopostocellar seta; dorsally of pseudopostocellar seta with 1 very short, upright postocellar seta on occiput; no ocellar seta; 1 porrect, lateroclinate and slightly dorsoclinate pseudovibrissa; ventrally of them with 1 short, dorsoclinate seta and with 2 long and 3 short ventro- clinate genal setae. Thorax (Fig. 3). Ground colour yellow, but scutum, scutellum, subscutellum and dorsal half of pleura to a line of middle of anepisternum and base of halter thinly ash grey microtrichose. Chaetotaxy: colour of setae and setulae as on head; 5 minute acro- stichal setulae in 1 row in anterior half of scutum; 1 strong posterior dorsocentral seta and 7-8 much shorter setulae anteriad in 1 line, without gap between last two setae; 1 short postpronotal seta; 2 stronger subequal notopleural setae; 1 postalar seta; 2 minute setulae dorsally of notopleuron anterior of suture; 1-2 minute postsutural intraalar setulae; scutellum with 1 apical scutellar seta and 1 minute dorsolateral setula; 1 kate- pisternal seta. Wing (Fig. 4). Rather elongate, maximal length:width ratio 3:1; alula absent; vein CuA2 present as fold, convex, below better developed, other venation as in S. soniae; ratios on C: section 2/3 (distance between R1 and R2+3/R3+3 and R4+5) about 18; section 3/4 (distance between R2+3 and R4+5/R4+5 and M) about 0.85. Last section on M (DM-Cu to tip) about 3 times as long as distance between crossveins. Halter white; squamae yellow, small. Legs. Entirely yellow, only last tarsal segment of fore and mid leg blackish, and apical third of last hind tarsomere slightly brownish; fore femur with 2 posteroventral brown setae; mid tibia with one brown, ventral apical seta. Abdomen (Figs 5-6). Tergites 1+2 ivory-yellowish, remaining tergites of yellow ground colour, but with badly delimited brown patches, T3 and T4 darkest, T5 and T6 distinctly paler. T3-T6 less transverse and narrower than those of S. soniae (Fig. 5). T6 well developed, relatively long (twice as long as in S. soniae). Sternites large, pale yellow; S1 short, transverse and wider than S2; S2-S5 becoming wider posteriorly, subequal in length or S5 slightly shorter. Abdominal spiracles strongly reduced, hardly visible. Postabdominal sclerites (S6-S8, see Fig. 6) fused to asymmetrical complex, markedly longer than in S. soniae, but similarly shaped. Strip-shaped marginal part belonging to S6 and stripe-like ventral projection of right anterior corner of S8 meeting 526 B. MERZ & J. ROHACEK (Fig. 6) on ventral side. Additional sclerite (Fig. 6) on right side behind S6-S8 complex larger and darker brown than that of S. soniae, somewhat resembling a head of mattock. Male genitalia (Figs 7-12). Epandrium (Figs 7, 8) orange yellow, distinctly higher than long, shortened ventrally and less broad than that of S. soniae but with similar setosity including 3 anteroventral setae on each side. Anal fissure broad but cerci smaller and closer each to other than those of S. soniae. Surstylus (Figs 8, 12) normally so strongly bent inside (inclined) that almost invisible in lateral view (and therefore omitted in Fig. 7; in Fig. 8 the surstylus is somewhat unnaturally straigh- tened); in widest extension view (Fig. 12) more slender than in S. soniae with dilated and densely micropubescent proximal third, slender middle part and widened distal third provided with 2 distinct teeth. Bacilliform sclerite formed by two separate and setose sclerites (Figs 7, 8), more slender than in S. soniae. Hypandrium (Figs 7, 10, 11) similarly formed as in S. soniae but both its lateral parts provided with robust anteroventral appendages (? pregonites) being different on left (having apical and ven- tral preapical tooth, Fig. 10) and right (simply apically pointed, Fig. 11) sides. Aedeagal complex asymmetrical, with only left postgonite developed (Fig. 10) though small and weakly sclerotized (right postgonite absent in contrast to large one in S. soniae). Basiphallus (Fig. 9) more robust than in S. soniae, with short and blunt epiphallus and with richly microsetulose ventrolateral lobes. Distiphallus similarly armed as that of S. soniae, but apical processes (including preapical sclerite) smaller and differently shaped (Fig. 9). Phallapodeme (Fig. 9) more robust than in S. soniae, with strongly bent ventral process. Ejaculatory apodeme (Fig. 7) also thicker than in S. soniae with more robust distal part and with smaller dark proximal part. DESCRIPTION OF FEMALE Wing length. 1.75-1.95 mm. Body length: 1.5 mm. Morphology. As male, but differing in the following points: Head more exten- sively darkened, with only anterior margin of frons yellow; face medially grey micro- trichose; most of occiput and postgena grey microtrichose. Abdominal tergites 3-7 yellow, laterally indistinctly darkened; 2 large, black spermathecae translucent on level of segment 7. Terminalia not dissected. BIOLOGICAL OBSERVATIONS The specimens from Jussy were collected by holding an insect net under large tussocks of Carex, which were beaten and shaken with a wooden stick (= “tussocking”, see Drake, 2004). The flies were sitting on dead leaves and did not fly away when approched by an aspirator. In the tubes they were crawling into a piece of paper and did not come out even when shaking the tube. One specimen was observed walking anteriorly, posteriorly and laterally with the same speed, with the head always directed upwards in the net. A similar behaviour was observed by Williams (1939) and Sabrosky (1975). The wings remained closed during these movements. Despite two hours of careful searching at the same place no additional specimens were collected. In England S. delicata is closely associated with tussocks of large species of Carex (eg. C. paniculata, C. acutiformis) in both inland and coastal wetlands and may be obtained WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 527 Fics 7-12 Stenomicra (Podocera) delicata (Collin), male genitalia (specimen from Germany, Bielefeld). 7, genitalia laterally (surstylus omitted); 8, external genitalia caudally (left surstylus omitted); 9, ae- deagal complex laterally (ejaculatory apodeme and postgonite omitted); 10, left side of hypan- drial complex with postgonite laterally; 11, right side of hypandrial complex laterally; 12, left sur- stylus ventrocaudally (widest extension). Abbreviations: bp = basiphallus; bs = bacilliform scle- rite; ce = cercus; dp = distiphallus; eja = ejaculatory apodeme; ep = epandrium; hy = hypandrium; pg = postgonite; pha = phallapodeme; prg = pregonite; sst = surstylus. (Del. J. Rohacek) only by beating or suction sampling method (Drake, 2004; P. J. Chandler, J. W. Ismay & B. Schulten, personal communication, 2004). DISTRIBUTION Described from Great Britain and later found in Germany (Tschirnhaus, 1999) and Switzerland (new record). 528 B. MERZ & J. ROHACEK Stenomicra (Podocera) soniae sp. n. Figs 13-23, 26-27 Stenomicra delicata: Papp, 1978: 198; Bachli, 1997: 34, 1998: 289; Rohaéek, 1997: 79; Rohaéek & Bartäk, 2001: 378. (Misidentifications) Stenomicra (Podocera) delicata: Rohaéek, 1983: 133, 1986: 146, 1987: 255, 1995a: 173, 1995b: 150. (Misidentifications) Stenomicra sp. n.: Tschirnhaus, 1992: 458. Stenomicra spec.: Buck, 1996: 371. MATERIAL (74 specimens) Holotype d, Czech Republic: “CS, Kunice-garden, Malaise trap, Bartäk, 49.56N/14.40E, 430 m, 8-9.VI.1985” (SMO). The holotype is laterally pinned on a cardpoint and is in very good condition. Paratypes: Bulgaria: 1 2 , Sliven, 13 km N, 42.49N/26.16E, 800 m, damp val- ley, 21.VII.1987, Bartak (CMB). Czech Republic: 19 , same data as holotype, but 13- 19.VII.1985 (MHNG); 16, Prostejov-Kunstät, car net, 5.VIII.1992, Bartàk (SMO); 12, Duchcov, 50.36.30N/13.43.30E, 220 m, floodplain forest, MT [= Malaise trap], 18.V.-15.V1.1998, Bartak (CMB); 14 , same data, but 25.vi.-23.vii.1998, Bartäk leg. (SMO); 1 6, Podyji NP, Braitava, 48°52’N, 15°49’E, 31.V.-2.VI.2002, mixed wood, car-net, Bartik (CMB); 19, Bohemia, Hradec Krälove, 5.VII.1997, Zeman leg. (Museum of East Bohemia, Hradec Krälové). Germany: | 2, BW [=Baden-Wiirttem- berg], Niefern nr. Pforzheim, 13.-22.VI.1992, Malaise-trap, Schmid-Egger (BUB); 32 2,BW, Ravensburg, Bad Waldsee, 16.VI.-15.VII.1995, Eklektor, M. Buck (BUB); 14, Hessen, Darmstadt, 18.VI.-2.VII.1989, Eklektor, M. Buck (BUB); 16, NW [= Nordrhein-Westfalen], Bielefeld, Universität, 3.-10.VII.1986, pan trap, M. von Tschirnhaus (BUB); 99 9, NW, Bielefeld, Werther Str., 23.-30.VI.1987, pan traps, M. von Tschirnhaus (BUB, MHNG); 2222, NW, Bad Lippspringe, 18.VI.1989, B. Robert (BUB, MHNG); 39 2, NW, Köln, Dünnwald, 27.VI.-4.VII.1989, Malaise trap, J. Wehlitz (BUB); 22 2, NW, Köln-Pöll, 30.V.-6.VI.1989, Malaise trap, J. Wehlitz (MHNG); 19, same, 27.VI.-4.VII.1989 (BUB); 19, same, 11.-18.VII.1989 (BUB); 12, same, 1.-8.VIII.1989 (BUB); 1¢, NW, SE Krefeld, Nature Reserve “Die Spey”, 26.V.-2.VI.1990, Malaise trap, Birnbrich er al. (BUB); 1 2 , same, 2.-9.VI.1990 (BUB); 16, same, 9.-16.VI.1990 (BUB); 19 , same, 16.-24.VI.1990 (BUB); 1 4 , same, 24.VI- 1.VII.1990 (MHNG); 16, same, 8.-15.VII.1990 (MHNG); 16, RP [= Rheinland- Pfalz], Eifel-Gebirge, Kr. Daun, Gönnersdorf, 29.VI.-6.VII.1991, Malaise trap, Cölln & Pompé (BUB); 1 2, SH [= Schleswig-Holstein], Kr. Plön, Sieversdorf, Postsee, 22.- 29.V.1983, yellow pan trap, M. von Tschirnhaus (BUB). Romania: 19, Carasova, 6 km N, 45.16N/21.53E, flowering meadow, 3.VII.1987, Bartàk (SMO). Slovak Republic: 1 2 , Slovakia or., Slovensky kras, Zädiel (at light) 15.VI.1981, J. Rohäëek (SMO); 19, same, but 16.VI.1981 (SMO); 19, Slovakia or., Nova Sedlica env. (distr. Humenné), sweeping over boggy meadow, 7.VII.1993, J. Rohaéek (SMO); 1 9 , Bärtfa [= Bardejov, NE Slovakia], Csergö h. [= Cergov Mts.], 5.VII.1969, Mihälyi (HNHM). Switzerland: 19, GE, Geneva, in a glass pavilion, 20.VII.1982, J. Steffen (MHNG); 12, JU, Delémont, 2-6.VIII.1974, G. Bächli (CGB); 12, TI, Bolle di Magadino, 17- 20.VI.1995, FZ [= Fangzelt], Merz & Bächli (HNHM); 29 ©, TI, Gordola, 580-660 m, 1997, M. Moretti (CGB); 12, VS, Leuk-Pfynwald, 614100/290070, 600m, 7.VI.2001, Merz & Landry (MHNG); 12, ZH, Embrach-Haumiili, 400 m, 10.VII.1997, B. Merz (MHNG); 1 2 , ZH, Dietikon, 15-19.VII.1989, G. Bachli (CGB); 19, ZH, Dietikon, 26.VI.-21.VII.1997, C [= Canopy], G. Bächli (CGB). WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 529 T1+2 Fics 13-18 Stenomicra (Podocera) soniae sp. n. 13, head laterally; 14, head dorsofrontally; 15, antenna laterally (medial side); 16, thorax dorsally; 17, wing; 18, abdomen dorsally. Abbreviations: sen = sensilla; S = Sternite; T = Tergite. (Del. J. Rohacek) ETYMOLOGY Named after Mrs Sonia Guyot who supports the scientific work of the senior au- thor in many ways. DIAGNOSIS This species is unmistakable among Western Palaearctic Stenomicra by the fol- lowing combination of characters: body mainly ash grey (Figs 26, 27); wing with 530 B. MERZ & J. ROHACEK small, but distinct alula, cell cup open; hyaline with very narrow milky-white cross- band at base and with both crossveins bordered by a narrow milky-white frame; legs yellow, but mid and hind femora and corresponding tibiae each with black ring; abdo- men uniformly brown. DESCRIPTION OF MALE Wing length. 1.75-2.05 mm. Body length: 1.4-1.5 mm. Head (Figs 13-15). Ground colour ash-grey, but a narrow stripe along posterior margin of compound eyes, gena and face translucent yellow; antennae and proboscis yellow; compound eye with short, but distinct interfacetal setulae; in lateral view elon- gate oval, oblique, slightly kidney shaped with posterior margin concave; gena at most one eighth as high as compound eye; frons bare, slightly wider than high, parallel- sided, ocellar triangle slightly raised in the middle of frons; fronto-orbital plate raised; face concave, raised ventrally at vibrissal angle; medially with carina; mediodorsally with a pair of sensillae (very shortened spine-like setae - see Figs 13, 14); antenna (Fig. 15): scape very small, ring-like, with one black dorsal setula; pedicel large, as long as postpedicel, cape-like, apicodorsally with one outstanding black seta, basally with a ring of 5-6 black setulae; postpedicel dorsoapically with a brush of soft, pale setulae; arista black, Drosophila-like with long rays which are much longer than width of post- pedicel; in addition to apical fork dorsally with 5-6 and ventrally with 2-3 rays; mouth- parts: palpus minute, about as long as wide, almost invisible; proboscis not projecting in front of vibrissal angle, with fleshy labellae. Chaetotaxy: colour of major setae pa- ler or darker brown depending on viewing angle and illumination; 2 subequal fronto- orbital setae in anterior half of frons, anterior fronto-orbital seta proclinate and incli- nate, posterior fronto-orbital seta reclinate; 1 long, lateroclinate medial vertical seta; anteromedially with 1 slightly shorter, proclinate pseudopostocellar seta; postocellar seta on occiput dorsally of pseudopostocellar absent; no ocellar seta; 1 porrect, latero- clinate and slightly dorsoclinate pseudovibrissa; ventrally with 1 short, dorsoclinate seta, and with 2 long and 3 short ventrally directed genal setae along mouth margin. Thorax (Fig. 16). Ash-grey microtrichose, but tip of scutellum and ventral part of pleura between coxae yellow. Chaetotaxy: colour of setae and setulae as on head; 5- 7 minute acrostichal setulae in 1-2 irregular rows in anterior half of scutum; one row of 5-7 dorsocentral setae, the penultimate much longer than the anterior setulae and se- parated from the longer and stronger prescutellar dorsocentral seta by a distinct gap; 1 short postpronotal seta; 2 long, subequal notopleural setae; 1 postalar seta; 1-2 minute presutural and 1-2 minute postsutural intraalar setulae; scutellum with 1 apical scutellar seta and with 1-2 small dorsolateral setulae; 1 katepisternal seta, other pleural sclerites bare. Wing (Fig. 17). Less elongate than in S. delicata, length/width ratio of 2.7; alu- la small, but distinct; C reaching M; BM-Cu absent; no trace of CuA2, cell cup entire- ly open; crossveins R-M and DM-Cu present; distance between crossveins 1.5-2 times the length of DM-Cu; R2+3 very long and converging towards M near tip; ratios on C: section 2/3 (distance between R1 and R2+3/R2+3 and R4+5) over 10; section 3 and 4 (distance between R2+3 and R4+5/R4+5 and M) of about same length; last section of M about 2.5 times as long as penultimate section between crossveins; Sc weak, WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 531 reaching C as fold where the latter has an indistinct break; surface of wing hyaline, but with a milky-white, narrow stripe from node of R2+3 and R4+5 to vein CuA1, and with both crossveins surrounded by milky-white frame. Halter with yellow stem and black knob (but knob usually yellow in specimens in alcohol), squamae small, dark brown, fringe blackish. Legs. Mainly yellow, but apical third of hind femur and basal third of hind tibia each with a distinct, broad black ring; mid femur and mid tibia with indistinct, small brown infuscation around knee; last tarsal segment of front and mid leg blackish, and apical third of last hind tarsomere slightly brownish; fore femur posteroventrally with 2 brown, long setae; mid tibia with one apicoventral brown seta; tibiae without dorsal preapical seta. Abdomen (Fig. 18). Tergites completely brown, Tl medially and T6 somewhat paler. T3-T6 distinctly wider and more transverse than those of S. delicata, bent on lat- eral sides of abdomen; T6 particularly shortened (Fig. 18). Preabdominal sternites S2- SS large, orange brown. S1 short and transverse, with only lateral parts dark pigment- ed and looking like divided in 2 small sclerites. S2 longest (distinctly longer than any of other sternites) but narrowest; S3-S5 becoming slightly wider and shorter posteriorly, S5 widest and most transverse. Abdominal spiracles near margins of tergites, strongly reduced. Postabdominal sclerites (S6-S8) forming a fused complex shorter than that of S. delicata. S6 reduced to transverse anterior marginal stripe of this complex; part belonging to S7 also small, subtriangular on left side. S8 slightly asym- metrical but its right anterior corner far projecting as strip-like process on ventral side almost to meet S6. There is a small, pointed additional sclerite in membrane on right side behind the latter process. Male genitalia (Figs 19-23). Epandrium relatively long, longer than high and very broad (see Figs 19-20), uniformly shortly setose and with 3 setae in anteroventral comers. Anal fissure broadly semicircular. Cerci well developed, widely separate, each with 1 long apical seta and number of short setae. Surstylus (Figs 19, 20, 23) strongly inclined, in widest extension view (Fig. 23) with simply rounded apex, broader and densely microtrichose in proximal two-thirds of lateral side and with setae mainly concentrated on medial side. Bacilliform sclerite bipartite, not fused medially; each its part movably connected with surstylus and distinctly setose (Fig. 20), anteriorly reaching to posterior margin of hypandrium as also is an Y-shaped slender sclerite attached by its arms to cerci (Fig. 20). Hypandrium (Figs 19, 22) asymmetrically frame-shaped, but posteromedially disconnected; its lateral parts expanded and spar- sely setulose, anteromedial part slender and connected with ventral process of phallapodeme; left side of hypandrium anteroventrally provided with a large, suboval, densely tuberculate appendage (? pregonite, see Figs 19, 22) while right side is simple but with more setulae (Fig. 22). Aedeagal complex distinctly asymmetrical (see Figs 21, 22), particularly as regards unevenly developed postgonites, left being small and suboblong (in lateral view), right one very large but pale pigmented, lobe-shaped, with concave inner side and convex outer side. Basiphallus short, inversely V-shaped in caudal view (Fig. 22), with distinct posterodorsal process (epiphallus) and ventrolateral lobes rounded and distinctly microsetulose (on left side — Fig. 21). Distiphallus (Fig. 21) relatively small, ventrally largely membranous, dorsally weakly sclerotized and 532 B. MERZ & J. ROHACEK À ı IN: VAM ALF RZZ Say A die yf Naa Fics 19-23 Stenomicra (Podocera) soniae sp. n., male genitalia (specimen from Germany, Krefeld). 19, genitalia laterally; 22, internal genitalia caudally; 20, external genitalia caudally; 21, aedeagal complex laterally (ejaculatory apodeme omitted); 23, left surstylus ventrolaterally (widest extension). Abbreviations: bp = basiphallus; bs = bacilliform sclerite; ce = cercus; dp = distiphal- lus; eja = ejaculatory apodeme; ep = epandrium; hy = hypandrium; pg = postgonite; pha = phal- lapodeme; prg = pregonite; sst = surstylus. (Del. J. Rohaéek) WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 533 micropubescent, with densely spinose apex including a slender, left-curved and point- ed sclerite and a small terminal, finely spinulose lobe. Phallapodeme short (Fig. 21), with simple anterior rod and distinct ventral process connected with hypandrium. Ejaculatory apodeme (Fig. 19) larger than phallapodeme, with darker and more robust proximal part and slender anterior rod with expanded but hyaline apex. DESCRIPTION OF FEMALE Wing length. 2.00-2.40 mm. Body length: 1.6-2.0 mm. Morphology. As male, but differing in the following points: Head more exten- sively darkened, with only gena and antennal grooves yellow, but facial carina grey microtrichose; abdominal tergites uniformly brown, posterior margin of tergites 3 and 4 narrowly yellow, posterior tergites slightly darker; sternites orange-brown. Spermathecae not visible externally without dissection. Terminalia not studied. BIOLOGICAL OBSERVATIONS Almost nothing is known about the biology of S. soniae. Tschirnhaus (1992) suggested that larvae may develop in axils of Angelica silvestris filled with water. Most specimens were collected with various traps, and only 3 specimens were swept with a net (Tab. 2). It is remarkable that males were almost exclusively collected with Malaise traps, whereas females could be obtained with all methods. The apparent rarity of these flies may be explained by the difficulties in collecting them with the sweeping of vegetation, the most frequently used collecting method. The discovery of a number of specimens collected “in the air” with car nets, window traps and at light is an indication that S. soniae is rather vagile, unlike S. delicata (see above). The ecological tolerance of S. soniae seems to be much less specific than of the other 3 species of the genus in the Western Palaearctic region that are all associated with a wet environment and the presence of large Cyperaceae (Carex, Cyperus, see introduction). Only 4 males and 13 females were collected in swampy areas, whereas the other specimens were found in various dry to wet open grassland and in forests (Tab. 3). Even garden vegetation and towns offer suitable habitats for this species with at least 1 male and 8 females caught in these places. This observation gives evidences that S. soniae is much more abundant than the material in museums would suggest. Adults were found from May to August. TABLE 2. Collecting methods for Stenomicra soniae sp. n. Abbreviations for countries: CH = Switzerland; CZ = Czech Republic; GE = Germany; SK = Slovakia. Method males females origin depository Malaise trap 9 13 CZ, GE BUB, SMO, CMB pan trap 1 10 GE BUB eclector 1 3 GE BUB window trap 3 CH MHNG, CGB light 2 SK SMO car net 1 1 CZ SMO, CMB beer trap 1 CH CGB banana-yeast trap l CH CGB sweeping 3 CH, SK MHNG, SMO no information 25 534 B. MERZ & J. ROHACEK TABLE 3. Habitats for Stenomicra soniae sp. n. Information was taken from unpublished Excel file data from M. von Tschirnhaus (received together with specimens), field book notes of the first author, and label data. Habitat males females 8 garden, town 1 meadows 1 pastures 1 shrubs 4 forests (open, deciduous) 1 unknown 4 N Boaov DISTRIBUTION Widely distributed in Central and Southeastern Europe from Northern Germany to Bulgaria. AFFINITIES The following remarks are based on descriptions and illustrations, but not on examination of specimens, except for S. fascipennis Malloch (Material studied: 26 dg, 222, Japan, Mt. Enaki, Yokohama City, Honshu, 15.VIIL.2002, M. Sueyoshi, MHNG). The new species belongs undoubtly to the subgenus Podocera Czerny by the subgeneric characters given in the key above, which is partly taken from Sabrosky (1975) and Freidberg & Mathis (2002). Within the 12 valid species of Podocera 5 species have characteristic milky-white crossveins R-M and DM-Cu as in S. soniae: S. fascipennis Malloch, 1927, from the Eastern Palaearctic and Oriental Regions, S. fae- niata Hennig, 1956, from the Neotropics, and the Afrotropical species S. biconspicua Sabrosky, 1975, S. trimaculata Sabrosky, 1975, and S. uniconspicua Sabrosky, 1975. S. fascipennis differs from all other species, including S. soniae, by the presence of a fourth whitish crossband near the distal margin of the wing and by the extremely short ultimate section of CuA1 which is much shorter than DM-Cu (in S. soniae at least 1.35 times as long, see Sueyoshi & Mathis, 2004, Fig. 2f). The 3 Afrotropical species all have a more extensive milky-whitish spot over DM-Cu which reaches dorsally at least R4+5, and all species have the ventral half of the anepisternum yellow, as well as entirely yellow femora (Sabrosky, 1975). In S. soniae the milky-white area is confined to a small surface around DM-Cu without reaching R4+5, the anepisternum is entirely grey, and the mid and hind femora have both a subapical brown to black ring. According to Hennig (1956), S. taeniata seems to have a rather similar wing pattern as S. soniae, but the wing base is more broadly whitish (only very narrowly in S. soniae), and the prescutellar dorsocentral seta is not separated by a gap from the penultimate seta (in S. soniae with a distinct gap). All dorsocentral setae except for the prescutellar dorsocentral seta are short in S. taeniata, whereas the last two dorsocentral setae are distinctly longer than the other dorsocentral setae in S. soniae. Our knowledge of the genus is still preliminary. Probably less than 10% of the species that are stored in entomological collections are named. It is therefore premature to propose phylogenetic relationships. Moreover, no safely identified species outside the Western Palaearctic region were available for study. However, based on the comparison with other species of Stenomicra it is reasonable to assume that S. soniae WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 535 Fics 24-29. Habitus photos of Western Palaearctic Stenomicra. 24, S. delicata (Collin), lateral view; 25, same, dorsal view; 26, S. soniae sp. n., lateral view; 27, same, dorsal view; 28, S. cogani Irwin, dorsal view; 29, S. jordanensis Freidberg & Mathis, lateral view. (Photos by A. Nobile, San Francisco). 536 B. MERZ & J. ROHACEK may be more closely related to the 3 Afrotropical species based on wing pattern (Sabrosky, 1975, Figs 5-7) and general description than to Stenomicra (Podocera) from other biogeographical regions. Subgenus Stenomicra Coquillett Stenomicra Coquillett, 1900: 262. Stenomicra (s. str.) cogani Irwin Fig. 28 Stenomicra cogani Irwin, 1982: 235. Material studied: Germany: 19, Nordrhein-Westfalen, Bielefeld-Nord, 2.VII. 1999, M. v. Tschirnhaus (BUB); 16, Schleswig-Holstein, Kr. Plön, Lauker See, Kühren, 6.VI.992, M. v. Tschirnhaus (BUB). Italy: 106 6, 322, Mantova pr. Marmirolo, Bosco d. Fontana, 50 m, 45.12N/10.45E, 25.V.2001, Merz & Mason (MHNG); Spain (new record): 14,39 2, Andalusia, Alcala de los Gazules, 48km NNW Tarifa, road no 440, 6.1V.1993, M. v. Tschirnhaus (BUB). In 1982, Irwin described and illustrated this species, including its external male terminalia, but no information was given about hypandrium and the aedeagal complex. The species is unique in the Western Palaearctic Region by its entirely yellow colour and the absence of the DM-Cu-crossvein (Fig. 28). Stenomicra (s. str.) jordanensis Freidberg & Mathis Fig. 29 Stenomicra jordanensis Freidberg & Mathis, 2002: 48. Material studied: 18,22 9 , Israel: Park HaYarden, 14.1V.1999, A. Freidberg (TAU) (Paratypes). An excellent description was provided by Freidberg & Mathis (2002). The ter- minalia of this species, however, were not studied. S. jordanensis can be distinguished from the other 3 Western Palaearctic species by the presence of vittae on the thorax (Fig. 29). ACKNOWLEDGEMENTS It is our pleasure to thank J. Ismay and B. Schulten (Oxford, England) for demonstrating the technique to collect small flies from grass tussocks and for un- published data about the biology and distribution of S. delicata. We sincerely thank A. Freidberg (Tel Aviv, Israel) for loan of specimens and his help during the various stages of preparation of this paper, and to P. J. Chandler (Melksham, England) for literature and information about British species of Stenomicra. We are indepted to A. Pont and J. Hogan (Oxford, England) for loan of types of S. delicata and to G. Bächli (Zürich, Switzerland), M. Bartäk (Praha, Czech Republic), L. Papp (Budapest, Hungary), M. Sueysohi (Washington D. C., USA) and M. von Tschirnhaus (Bielefeld, Germany) for loan or donation of specimens. A. Nobile (California Academy of Sciences, San Francisco, USA) produced the photos of the colour plate, F. Marteau (Geneva) helped us with the prepartion of the plates, Ch. Lienhard (Geneva) gave precious advice on nomenclatorial problems, and W. N. Mathis (Washington D. C., USA) kindly reviewed WESTERN PALAEARCTIC STENOMICRA (PODOCERA) 537 the manuscript. Our thank is extended to the “Service des forêts, de la protection de la nature et du paysage” (G. Dändliker, Geneva) for the delivery of a collecting permit for Natural Reserves in the canton Geneva. REFERENCES ANDERSSON, H. 1991. Dvärgflugor, Stenomicridae, en för Norden ny flugfamily (Diptera). Entomologisk Tidskrift 112: 149-151. BACHLI, G. 1997. Die Arten der Tanypezidae, Dryomyzidae, Periscelididae, Acartophthalmidae, Aulacigastridae und Stenomicridae in der Schweiz (Diptera). Mitteilungen der Entomologischen Gesellschaft Basel 47 (1): 29-34. BACHLI, G. 1998. 84. Stenomicridae (p. 289). In: MERZ, B., BACHLI, G., HAENNI, J.-P. & GONSETH, Y. (eds). Diptera - Checklist. Fauna Helvetica 1: 1-369. Buck, M. 1996. 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Minier- und Halmfliegen (Agromyzidae, Chloropidae) und 52 wei- tere Familien (Diptera) aus MALAISE-Fallen in Kiesgruben und einem Vorstadtgarten in Köln. Decheniana — Beihefte 31: 445-497 & 3 plates. TSCHIRNHAUS, M. VON, 1999. Periscelididae (pp. 170-171). In: SCHUMANN, H., BAHRMANN, R. & STARK, A. (eds). Checkliste der Dipteren Deutschlands. Entomofauna Germanica 2. Studia Dipterologica, Supplement 2: 1-354. WILLIAMS, F. X. 1939. Biological studies in Hawaiian Water-Loving Insects. Part III. Diptera or Flies. B. Asteiidae, Syrphidae and Dolichopodidae. Proceedings of the Hawaiian Ento- mological Society 10: 281-315. REVUE SUISSE DE ZOOLOGIE 112 (2): 541-542; juin 2005 Synonymic note on the monobasic genus Ophryomedon Wasmann, 1916 (Coleoptera, Staphylinidae, Paederinae)* Guillaume de ROUGEMONT Parfondeval, F-76660 Londinières, France. Synonymic note on the monobasic genus Ophryomedon Wasmann, 1916 (Coleoptera, Staphylinidae, Paederinae). - Ophryomedon marginatus Naomi, 1995 is designated a junior synonym of Ophryomedon crenatus Wasmann, 1916. Keywords: Staphylinidae - Paederinae - Ophryomedon crenatus - margi- natus - synonymy. Wasmann (1916) described the myrmecophilous beetle Ophryomedon crenatus from Malacca, Malaysia. Naomi (1995) described Ophryomedon marginatus from Japan (Honshu and Kyushu) and, relying on Wasmann’s description, gave as diagnos- tic differences a longer third segment of the labial palpi, different colour pattern, and broader abdomen of the Japanese specimens. While sorting unidentified Staphylinidae in the collection of the Geneva Natural History Museum I found eight specimens belonging to the genus Ophryomedon labelled as follows: 4 exx.: JAPON, Nara, Nara, 27-31.VII.1980, Cl. Besuchet (coll. Geneva Mus., 1 ex. coll. Rougemont); 1 ex.: INDIA, U.P., Kumaon, Bhim Tal 1450-1550 m., 5.X.1979, I. Löbl (coll. Geneva Mus.); 1 ex.: INDIA, Assam, Manas, 200 m., 22.X.1978, Besuchet — Löbl (coll. Rougemont). Naomi (1995) has provided a good description and illustrations of O. margi- natus Naomi, with which the new material agrees in all respects. The colouration and extent of the dark elytral band are somewhat variable, and in one specimen from Nara the whole body is slightly infuscate, so that the elytral marking almost disappears from lack of contrast with the ground colour. Even without examining the type of O. cre- natus Wasmann, it can be confidently asserted that all specimens examined belong to a single, evidently widely distributed species. The new synonymy established is thus: Ophryomedon crenatus Wasmann 1916: 202. Ophryomedon marginatus Naomi, 1995: 162. Syn. n. * 45th contribution to the knowledge of Staphylinidae Manuscript accepted 03.09.2004 542 G. DE ROUGEMONT REFERENCES NAOMI S.-I. 1995. Descriptions of two new species of the subfamily Paederinae (Coleoptera, Staphylinidae) from Japan. Japanese Journal of systematic Entomology 1 (2): 161-168. WASMANN, E. 1916. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien Aus- gefuehrt im Auftrage der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen, V: Termitophile und myrmecophile Coleopteren. Gesammelt von Herm Prof. Dr. v. Buttel-Reepen in den Jahren 1911-1912 (207. Beitrag). Zoologische Jahrbiicher, Abteilung fiir Systematik, Geographie und Biologie der Tiere, Jena, 39: 169-210. REVUE SUISSE DE ZOOLOGIE 112 (2): 543-558; juin 2005 Ten years trends in the oligochaete and chironomid fauna of Lake Neuchâtel (Switzerland) Brigitte LODS-CROZET* & Olivier REYMOND Laboratoire du Service des Eaux, Sols et Assainissement, Boveresses 155, CH-1066 Epalinges, Switzerland. E-mail: brigitte.lods-crozet@sesa.vd.ch Ten years trends in the oligochaete and chironomid fauna of Lake Neuchâtel (Switzerland). - Quantitative surveys of benthic macroinverte- brates (oligochaetes and chironomids) were conducted during 1992, 1997, 2000 and 2002 at a depth of 40 m to monitor the biological quality of sedi- ments in Lake Neuchatel. Recent declines in frequency of occurrence and abundance of oligochaete species characteristic of oligotrophic conditions (Stylodrilus heringianus, Embolocephalus velutinus) contrasted with the improvement of water quality metrics. Total phosphorus concentrations in lake waters decreased from 63 mg m’? in 1980 to 10 mg m’? in 2002. Since 1992, significant reductions of total zoobenthic biomass have been recorded and the chironomid community structure reflects typical oligo- mesotrophic conditions in the upper sediment layer. The population of the oligochaete species Potamothrix vejdovskyi is in clear expansion and indi- cates that this new species for the lake (1986) has found good conditions for a successful colonization. These divergent responses within the zoobenthic community are discussed according to three main hypotheses: implications of toxic pollutants like heavy metals and organic micropollutants in the sediment, impact of algae on oxygen conditions on the bottom layer and different biological responses of oligochaetes to competition for food and space. Keywords: Zoobenthos - biomonitoring - profundal zone - sediment contamination - organic deposition - micropollutant. INTRODUCTION In marine and lacustrine ecosystems, it is generally accepted that pelagic phyto- detrital organic material, derived from autochtonous and allochthonous inputs, drives profundal zoobenthos production (Brinkhurst, 1974; Graf, 1989). The major part of autochtonous input arises from phytoplankton primary production which is controlled largely by nutrient elements (mainly P and N) coming in the system (Hàkanson & Jansson, 1983). Thus, profundal macroinvertebrate growth is supported directly by nutrients in fresh or decaying algae or indirectly via microbial production (Johnson & Wiederholm, 1992). * Corresponding author Manuscript accepted 24.11.2004 544 B. LODS-CROZET & O. REYMOND The deep bottom fauna of lakes is dominated by oligochaete worms and chiro- nomid larvae. Additional invertebrates are molluscs, crustaceans, watermites, flat- worms and minor insect taxa which usually occur in rather limited numbers (Wiederholm, 1980). The two main taxa have different biological traits: chironomid larvae have a surface deposit-feeding strategy, which show many diets (algae and freshly deposited detritus, sediment bacteria and invertebrates), whereas oligochaete are subsurface deposit-feeders (mainly bacterial food and organic matter). Their repro- ductive behavior is also distinctive: oligochaetes reproduce within the sediment whereas chironomids have aerial stages and oviposit on the water surface (Johnson & Wiederholm, 1992; Goedkoop & Johnson, 1993). Therefore, they do not respond equally to environmental changes. Under conditions of recovery, oligochaetes seem to maintain their community structure for a longer time than chironomids (Wiederholm, 1980; Lang, 1998). In Lake Neuchâtel, despite decreasing phosphorus concentration since the 1980s, densities of oligochaete species intolerant of oxygen depletion and organic deposition tended to decrease after 1992 (Lang 1999, 2001). To examine this trend, the same biomonitoring survey as in 1992, 1997 and 2000 was performed in 2002 in order to monitor the zoobenthic assemblages and further investigations were carried out on potential toxic contaminants of lake sediments. The present study focuses also on the change of the zoobenthic communities during the last ten years. STUDY AREA, SITES AND METHODS STUDY AREA Lake Neuchâtel, located at the southern foot of the Jura Mountains, is one of the largest Swiss lakes with a surface area of 215 km?. Its orientation is parallel to the major wind directions, resulting in a good oxygen supply even to the deepest waters. Oxygen concentrations one meter above bottom sediment have always been above 7 mg L:! from 1982 to 2002 (Pokorni-Aebi, 2002). Six major tributaries discharge into both ends of the lake (Fig 1). The main characteristics of the lake are given in Table 1. In 2002, physico-chemical and phytoplanktonic features classified this lake as mesotrophic (Häkanson, 1980). SAMPLING SITES In 2002, 30 sites were sampled along the southeasthern shore of Lake Neuchâtel (Fig. 1), between the villages of Portalban and Font. This biomonitoring transect is located along the largest fringing wetland in Switzerland (The “Grand Carigaie” reserve), which is an almost 40 km-long continuous marshland belt. The sites, distri- buted evenly (500 m apart), were sampled six times in May, before the mass emergence of chironomids. At each site, six cores (16 cm2, 30 cm long) were taken with a gravity corer lowered from the water surface. Similar biomonitoring surveys were previously undertaken in 1992, 1997 and 2000 respectively by Lang & Reymond (1993) and Lang (1999, 2001). In 1984, 16 sam- pling sites were distributed along four transects, regularly spaced (4 km apart) in the same area. Along each transect, samples were collected at 40, 60, 90 and 120 m deep. OLIGOCHAETES AND CHIRONOMIDS 545 Switzerland Neuchâtel 5 km ot = Wetland: Grande-Carigaie reserve Fic. 1 Lake Neuchätel showing inflows, outflow and sampling sites divided into 3 zones. TABLE 1. Main characteristics of Lake Neuchätel (Lang, 1996; Pokorni-Aebi, 2002) Depth (m) Mean 64 Max 153 Lake surface area (km?) 215 Volume (km?) 13.8 Altitude of lake surface (m a.s.l.) 429 Area of drainage basin (km?) 2672 Average altitude of the basin 780 No. of total circulation per year 2 Theoretical residence time (year) 8.2 Total phosphorus (mg m-3)4 1980 63.0 2002 10.0 Secchi disc transparency (m) 2002 2.5-9.0 Mean algal biomass (0-20 m)(g m) 2002 1.26 Chlorophyll-a (0-20 m)(mg m->) 2002 2.2-6.5 4 Mean value for the spring circulation Three zones on the 2002 transect were differenciated along a gradient of organic sedimentation (Lang, 2001). 546 B. LODS-CROZET & O. REYMOND CORE INVESTIGATIONS From 1997, each sediment core was described in the laboratory according to thickness of three main vertical layers: 1) the thin brownish surface layer of oxydized sediment, ii) the intermediate reduced black layer, which can be considered as a descriptor of organic deposition and matches with the 1940s to 1950s (Steinmann er al., 2003) and iii) the compact grey or brown clay found underneath. After measure- ment of the thickness of the reduced black layer to the nearest 0.5 cm, each core sediment was sieved (mesh size 0.2 mm) and retained material preserved in 5% formalin. Collected macrofauna consisted mainly of lumbriculids, tubificids and chiro- nomids which were sorted out and counted under a low magnification (6x) binocular microscope. Oligochaetes and chironomids were then mounted (Reymond, 1994) and identified to species or groups of species according to Brinkhurst (1971), and Wiederholm (1983, 1986). Some species of lumbriculids and tubificids are known to be sensitive to oxygen depletion and organic deposition (Jonasson, 1969; Milbrink, 1978, 1980; Lang, 1990), and can be considered as bioindicators of oligotrophic conditions. The bioindicative value of species or species groups for chironomid larvae was based on Saether (1979). In 1984 and 1992, chironomids were counted separetely in each core, but they were identified only as a whole sample based on 64 and 171 cores, respectively. In 1997, 2000 and 2002, chironomids were counted and identified separetely in each core. Chironomid material prior to 2002 was re-examined and identified to species or groups of species. CHEMICAL ANALYSES In 2003, three composite samples were collected along the same transect including the upper 5 cm of sediment which represent approximately twenty years of sedimentation (Steinmann er al., 2003). Chemical analyses were then performed on heavy metals (Cd, Cu, Cr, Ni, Pb and Zn) using Inductively Coupled Plasma — Atomic Emission Spectrometry (ICP-AES). PCBs were determined by GC/MS after puri- fication. Seven congeners were included (No. 28, 52, 101, 118, 138, 153, 180). PAHs (PolyAromatic Hydrocarbons) were determined by HPLC coupled with a fluorescence detector. They include Fluoranthene, Benzo(b)Fluoranthene, Benzo(k)Fluoranthene, Benzo(a)Pyrene, Benzo(g.h.i)Perylene, Indeno(1,2,3-cd)Pyrene. STATISTICAL ANALYSES Frequency of occurrence was calculated as the number of samples containing a particular species divided by the total number of samples. Relative abundance was the total number of individuals of a particular species collected in all samples in one par- ticular year divided by the total number of oligochaetes or chironomids. Differences between years or zones for sediment parameters were tested with a one-way ANOVA. Kruskall-Wallis rank tests were used to test differences in zoobenthic abundance and biomass. All analyses and graphical displays were undertaken using SPSS software (10.1) for Windows. OLIGOCHAETES AND CHIRONOMIDS 547 RESULTS SEDIMENT CONDITION AND CONTAMINATION The thickness of the reduced black layer of soft sediment decreased significant- ly in 2002 along a eastern-western gradient (zone 1 to 3) (F = 30.80, P < 0.0001). Similar trends occurred in 1997 and 2000 (F = 7.99 and 38.24 respectively, P < 0.0001) (Fig. 2). Micropollutant analyses of heavy metals, PCBs and PAHs in bottom sediments showed similar results in the three zones defined above (Table 2). Few values for PCBs and PAHs were above the quantitation limits of the methods. Fluorantene had the most elevated concentration (0.274 mg Kg” in zone 3). EM 205 E CE X U 154 E = FA O 10- \ = 2 LISE LU | ce | X | | O 5 < — a JE 07 T T | Pre 1997 2000 2002 YEAR FIG. 2 Gradient of organic deposition, expressed as the thickness of the dark reduced layer between 1997 and 2002 and between zones. The median, percentiles (25th and 75h) and 1.5 interquartile range are indicated. TABLE 2. Chemical characteristics of superficial sediments in the three zones of Lake Neuchâtel. All values are expressed as mg Kg on a dry sediment weight basis. REF: reference values Zone Cd Er Cu Ni Pb Zn >RCB 26 HAP 1 0.220 123 132655126 10.7 26.8 << OEE!) < 1.092 2 0.264 12.4 1341 12.6 SL 29 4 < 0.105 <1.111 3 0.265 14.1 142 146 1255 332 < 0.105 < 1.159 216 HAP REF 0.21 30! 30! 60! 0.12 1.02 l : reference values (Vernet & Viel, 1984); 2 : reference values (Osol, 1998) B. LODS-CROZET & O. REYMOND 548 SUOHIPUO9 9IYdOHOSI[O JO 10JR91pUI : O {exe} OÙ] JO J9uasaıd : + 081 081 SLI [LT 081 081 SLI IZI #9 59109 JO ON * * * * * * ‘dds snsunjcuny * * * * # * ‘13 uinsojnoaqnu unpipadkjog * * (U9SIOIN) SIMOJYI Sadipua10191N * * ‘dds sadipua10191q * * x * s* ‘dds snwuououry201dt4 10007 LSI 0 IT + CET IF6€ OCFLI 00 681 v9 * * (P9PS19N97) PU19D109 DYUITLIS 10007 86 OF + 60€ OT + 06 x FSLI ICI + L97 £'8c vrl 0% * ik ‘dds (4) snippjo04d 10007 wel cL ¥ 169 LE F9ET OCT + PITT 901 + 871 6 CS LIT VIL * * 19 suaosajou DHI0SdOHIN © 1000 > 909 CT + O9I SI + LS TIFOY OLI FSLE IERI SCI VL * ‘18 DINJIAS1U Du2d0pP]2D4104 O 2060 sro SFIC LF VI 67 IT L9 F EI CE et ['O k * (990) zuuvuyof vidojadossvp 0 YHVAINONOAIHD o 5 * * * (oqn19) sHDq1Dq SAPYIKAOUNUDS J * * * * * k (YOZVINN 9 Aysaoploa, ) S1SU91ADPJOUI X1AYJOWDIO J * xk (paeppog) 1(q4amos DANIYIUDAY * (JOIN) Xofıgnı xafiqny 9000 OCT £9 + 085 TLFSLS OLFITS PL + S68 14 £ 89 079 L9 OOI [ * (usspeystW) SISUAIUOULUIDY XIAYJOUDIOT (sojiuoAnf) sny11pouwurT 10007 S8SS8IT ESIFS6EI 8FO£6 6LF099 6ÿ + 887 OSL vv9 Téÿ 8c 90} [ * (IILLIOA) D021Punfosd snpı4pouunT + (oparede]9) astouffoy snpapouwT 10007 1907 ScIFISCI SIL + 8IIL Do F969 LY + 6ST CLO TIS 6th we 0 (OQUIH) 1Kysaoploa x14Y10WDIOA EIEO IT I IT + VOI 97 F EST TFOI CCF 701 6 €1 ooc Vél SEI oT * (uasiq) xosaf puodsonds © 000 Tor OT +79 91 F9L di + C8 de + p9I V8 IRE AZIO CGI * (oqnio) snunnjoa punodsonds © 1000 7 €S'SS CFL CFL 6 + SC S + 86€ Ul Tl OV EIT ore [ * oparede]) snupidunoy SNPAPO]MS © 1000 €85 OT * 76 Og F SOC LIF i + LSI 8 CI GUEST S'OT * JOYOSIOI] paummdups Djavyrig 0 VLAVHOOSITO d [| 007 0007 L661 T661 TOT 0007 L661 ss «HHT 861 8I6I WON) Souepunqy (%) souarimo90 Jo Aouonbe1{ -[OJeyonaN ONE] OU) Ur poI99][09 euNry PILIOUOIUO pue ajavyoos!fo JO (AS F) sourpunqe pue soue1nd90 Jo Aouonboiy ‘oousSa1q ‘£ ATAY,L OLIGOCHAETES AND CHIRONOMIDS 549 EVOLUTION OF ZOOBENTHIC COMMUNITIES In 2002, zoobenthos (Table 3) consisted mainly of lumbriculid and tubificid worms (12 species) and chironomid larvae (8 taxa). The main oligochaete taxa were present at each biomonitoring survey. Branchuira sowerbyi, Potamothrix moldaviensis and Potamothrix vejdovskyi were recent for the lake and the latter is a new colonist since 1986 (Lang & Reymond, 1993). For two of the four oligochaete species bioindicators of oligotrophic conditions (Stylodrilus heringianus and Embolocephalus velutinus), frequency of occurence and relative abundance tended to decrease through years with a strong shift after 1992 (Fig. 3A), whereas for Bichaeata sanguinea and Spirosperma ferox, no characteristic trends were seen (Fig. 3A and 3B, Table 3). Concerning the non-oligotrophic type worms, Potamothrix vejdovskyi showed a significant pattern, expanding spatially (present in 67% of the sites in 2002) and numerically (its population was five times higher in 2002 than in 1992) in no more than ten years (Kruskall-Wallis rank test, k = 76.13, p < 0.05) (Table 3 and Fig 3B). Populations of Potamothrix hammoniensis and Tubifex tubifex were started to decrease (Kruskall-Wallis rank test, k = 18.10, p < 0.05). In contrast, Limnodrilus species showed an inverse trend (Kruskall-Wallis rank test, k = 101.28, p < 0.05). The chironomid community was dominated by three taxa bioindicators of oligo- trophic conditions, Macropelopia fehlmanni, Paracladopelma nigritula gr. and Micropsectra notescens gr. which occurred in 42-78% of the samples (Fig 3C), depending on years. The presence of Micropsectra contracta (notescens gr.) was confirmed by pupal examination. Procladius, a rather euryecious and predatory genus was the fourth dominant taxon in terms of frequency of occurrence and abundance. Total mean zoobenthic biomass (oligochaete and chironomid larvae) decreased significantly between 1992 and 2002 (Kruskall-Wallis rank test, k = 50.67, p < 0.05) and also between the three zones (Fig 4, Table 4) following the same east-west gradient of organic deposition. Mean total abundance of worms (Fig 5) increased significantly through years (Kruskall-Wallis rank test, k = 58.72, p < 0.05); Potamothrix vejdovskyi and Limno- drilus were mainly responsible for this change. The major augmentation in oligo- chaetes occurred between 1997 and 2000. Chironomid assemblages showed an up-and- down pattern in their total abundance, mainly reflecting variations in Micropsectra density (Kruskall-Wallis rank test, k = 52.10, p < 0.05). DISCUSSION DIVERGENT TRENDS AT THE ZOOBENTHIC COMMUNITY LEVEL The zoobenthic communities reflected different trends with regard to the time- scale and the type of organism considered. Despite a phase of eutrophication during the 1970s, oligochaete assemblages retained their native composition since the beginning of the xxth century (Monard, 1919). Three new species of oligochaetes, from the Ponto-Caspian area, appeared in the lake during the last century (Potamothrix molda- viensis, P. vejdovskyi and Branchiura sowerbyi (Brinkhurst & Jamieson, 1971; Lang, 1984; Lang & Reymond, 1993). In 1918, Embolocephalus velutinus was the most & O. REYMOND 550 (%) JONVGNNAV SAILV1S4y | 2002 (%) JONVGNNAV JAILV 144 OLIGOCHAETES AND CHIRONOMIDS 551 MH Macr feh NC | Mi Micr not 757 N Para nig RELATIVE ABUNDANCE (%) 1992 1997 2000 2002 YEAR FIG. 3 Relative abundance of species (+ CL) between 1992 and 2002. A: 3 oligochaete species indica- tors of oligotrophic conditions (Bich san: Bichaeta sanguinea; Embo vel: Embolocephalus velutinus; Styl her: Stylodrilus heringianus); B: more tolerant oligochaete species to oxygen depletion and organic deposition: Pota vej: Potamothrix vejdovskyi and Spir fer: Spirosperma ferox; C: chironomid taxa (Macr feh: Macropelopia fehlmanni; Micr not: Micropsectra notescens gr.; Para nig: Paracladopelma nigritula gr.). frequent species followed by Potamothrix hammoniensis, Stylodrilus heringianus and Tubifex tubifex (Monard, 1919). Since 1984, oligochaete fauna has been dominated by the more tolerant taxa Potamothrix hammoniensis and Tubifex tubifex in terms of frequency and abundance. After 1992, a strong reduction in density of two species intolerant to oxygen depletion and/or organic deposition (Brinkhurst, 1964; Milbrink, 1978, 1983) was noted (Stylodrilus heringianus and Embolocephalus velutinus). The dominating position of Tubifex tubifex at both ends of the trophic scale would seem to be rather controversial (Milbrink, 1978, 1980). As is now widely recognized, at low organic content, 7. tubifex is generally a dominating species together with Stylodrilus and Embolocephalus species and abundances remains low (Milbrink, 1978; Milbrink et al., 2002). This was the case in the beginning of the XX" century in Lake Neuchâtel. The persistance in 2002, of a low density of Stylodrilus heringianus and Embolo- cephalus velutinus, combined with a reduction Potamothrix hammoniensis and Tubifex tubifex (frequency and abundance) indicated that the oligochaete species structure does not provide a coherent signature of the lake sediment quality. In contrast to oligochaetes, the bioindicative value of chironomid communities suggested good conditions at the water-sediment interface since 1992 with, in addition, 552 B. LODS-CROZET & O. REYMOND 30.07 20.05 MEAN ZOOBENTHIC BIOMASS | I 1992 1997 2000 2002 YEAR Fic. 4 Mean zoobenthic biomass (oligochaete worms and chironomid larvae) (FW, g m?) between 1992 and 2002 and between zones. The median, percentiles (25th and 75th) and 1.5 interquartile range are indicated. TABLE 4. Summary of the analysis of variance (one-way ANOVA) inter-zones for two variables. Variable ANOVA inter-zone F P Black reduced layer 1997 7299 < 0.001 2000 38.24 < 0.001 2002 30.80 < 0.001 Zoobenthic mean biomass 1992 5233 0.006 1997 10.52 < 0.001 2000 15.49 < 0.001 2002 4.46 0.013 the presence and the settlement of Paracladopelma nigritula gr., a species group characteristic of oligotrophic lakes (Saether, 1979). This taxon was also present in Lake Geneva at a depth of 40 m in 1995 (Lods-Crozet, unpubl. data) and in Lake Annecy (Verneaux & Verneaux, 2002). Chironomid composition and richness (11 taxa) during the last ten years is analogous to that of other lakes of the Alpine region, for the same depth: Lake Constance (Reiss, 1968), Lake Geneva (Lods-Crozet & Lachavanne, 1994) and Lake Starnberger (Gerstmeier, 1989). Furthermore, the inter-annual up and down pattern (Fig. 5) could be explained by the fact that, if profundal communities are food limited, then year-to-year fluctuations in autochtonous algal production may ulti- OLIGOCHAETES AND CHIRONOMIDS 333 a : 40007 oe ui 30007 fom O Z 5 pot Z 2000- 5 a 2 . i 1000- ae a me cy a ia ibm eo br Sei Sada 1992 1997 2000 2002 YEAR Fic. 5 Total abundance (No. m2) of oligochaete worms (circles) and chironomid larvae (triangles) (+ CL) between 1992 and 2002. mately affect profundal zoobenthos communities resulting in time-delayed population responses (Johnson & Wiederholm, 1992). In addition, two species of molluscs (Pisidium conventus, P. personatum), typ- ical of deep lakes were found. These two species were also present in the deep zone of the oligotrophic Lake Annecy, France (Mouthon, 2002) and were rare or absent in meso-eutrophic Swiss lakes (P. Stucki, pers. comm.) THE EFFECTS OF ABIOTIC AND BIOTIC FACTORS Chemical and phytoplanktonic metrics in the pelagic waters of lake Neuchatel indicate that mean total phosphorus concentrations in the water column have markedly decreased from 38 mg m in 1984 to 18 mg m in 1992 and 10.0 mg m in 2002. Consequently, mean algal biomass has also decreased (Pokorni-Aebi, 1997, 2002). In 554 B. LODS-CROZET & O. REYMOND contrast, during a similar phase of recovery from eutrophication in Lake Geneva, good relationships were found between pelagic chemical parameters and zoobenthic communities, with a significant increase of oligochaete species bioindicators of oligo- trophic conditions (Lang, 1998). In order to understand these divergent trends between zoobenthic groups we have explored three main hypotheses. One of them is contamination of sediment by micropollutants (heavy metals, PCBs and PAHs) and their potential impact on zooben- thic communities. Anthropogenic contaminants were detected along the 15 kilometres- long transect, but no spatial differences were noted. Heavy metals concentrations were relatively low compared to those measured in Lake Geneva in 1983 (Mondain-Monval et al., 1983) and in 2003 at 70 m-depth in the same lake (Lods-Crozet, unpublished data). Compared to background concentrations observed prior to industrialisation (Vernet & Viel, 1984) mean values were close and even lower than background levels (Table 4). PCBs in Lake Neuchâtel have similar values as at 70 m-depth in Lake Geneva (Lods-Crozet, 2003, unpublished data). Their long-term persistence in the upper layer of bottom sediments, despite their banning in industrial processes since 1986, could explain the residual contamination. PAHS levels are higher than those measured at 70 m depth in Lake Geneva: 0.193 mg Kg"! (Lods-Crozet, 2003, unpublished data). The Swiss reference is a zero value for these synthetic and per- sistant contaminants (PCBs and PAHs) but reasonable limits could be chosen on the basis of threshold values determined for agricultural valorisation of wastewater sludges (Osol, 1998) (Table 2). Similar levels of sediment contamination by heavy metals and PAHs were found in Lake St-Frangois, Canada (Pinel-Alloul et al., 1996) and attempts to correlate contaminants and macroinvertebrate metrics were unsuccessful. It should be stressed that sediment toxicity assessment is not an easy task (Luoma & Carter, 1993) and sediment quality criteria essentially take into account the capacity of benthic organisms to tolerate the different contaminant concentrations. Substantial doubt exists about pol- lutants concentrations that pose significant ecological dangers because the bioavailable fraction of toxicants in sediments is rarely measured (Pardos er al., 2004). The use of sediment quality assessment metrics, developped by Smith et al. (1996) permitted the following considerations. The Probable Effect Level (PEL) on benthic organisms, showed that the concentrations of metallic and organic (PAHs) contaminants detected were well below those estimated by the PEL (Smith er al., 1996). On the other hand, in laboratory experiments, Nalepa (1991) pointed out that sublethal effects were apparent at lower concentrations than those reported from Lake Ontario sediments. For instance, Stylodrilus heringianus ceased feeding after 58 days at PCB concentrations over 0.05 mg Kg! (PCB congeners not known). Concentrations at this level were found in Lake Neuchâtel in 2003. In addition, tubicifids feed continuously in a conveyor-belt fashion by ingesting particles in bulk at depth and defecating on the sediment surface. Such bioturbation has been reported to significantly increase the flux of metals and organic contaminants from sediment into the water column (Reynoldson, 1987; Reible et al., 1996). While results of laboratory experiments may not be appli- cable to the field situation, these results do demonstrate the possibility of low-level, long-term chronic impacts of pollutants on oligochaete populations (Nalepa, 1991). OLIGOCHAETES AND CHIRONOMIDS 555 Another hypothesis, proposed by Lang (1999) was the changes in the phyto- plankton composition between 1993 and 1996, with the dominance of large diatoms (Butty er al., 2003). These have a high sinking rate and as a result, sedimented intact on the bottom surface. Algal signatures at the sediment level were also observed by Steinmann et al. (2003) using the analysis of suspended material from the lake epilimnion. Settlement of these algae is known to increase oxygen uptake at the water- sediment interface (Johnson et al., 1989; Lang, 1999). Consequently, some oligochaete species, sensitive to oxygen depletion, were able to be disturbed in their growth and reproductive behavior. The last hypothesis concerned competition mechanisms. Few studies have actually demonstrated increases or decreases in population growth rate by interaction between oligochaete species (Milbrink, 1993; Timm, 1996). The rapid expansion of Ponto-Caspian Potamothrix vejdovskyi, which may compete for habitat and food with other oligochaete species like Stylodrilus heringianus and Embolocephalus velutinus, should not to be neglected. Furthermore, increase of lake water temperature during the last fifteen decades (Lazzarotto et al., 2002; Pokorni-Aebi, 2002) may have enhance growth and reproduction of this opportunistic species and strengthened up its expansion over more stenothermic native species. Our study confirms the potential multiple controls of the macroinvertebrate community in anthropogenically stressed ecosystems by both simple and combined effects of chemical, toxicological, biological and ecological factors. Further impro- vement of lake biomonitoring programs must address assessment of the structure of predominant and profundal zoobenthic community (oligochaetes, chironomids and bivalves molluscs). We consider that Lake Neuchâtel, in terms of biological quality of sediments is in phase of recovery from eutrophication, by using zoobenthic biomass, chironomid community structure and Pisidium assemblages as descriptors. The decline of characteristic oligochaete species indicators of oligotrophic conditions from 1992 should be monitored further in order to better understand the causes of this apparent degradation. ACKNOWLEDGEMENTS We thank Raymond Ducret for his technical assistance in the field, Claude Lang for providing data and helpful comments and Pascal Stucki for the Pisidium iden- tifications. The authors are grateful to Sandra Knispel and Emmanuel Castella for valuable comments on an early draft of the manuscript and to two anonymous reviewers for their constructive remarks. The authors also thank John E. Brittain for correcting the English in a subsequent version of the manuscript. REFERENCES BRINKHURST, R. O. 1964. Observations on the biology of lake-dwelling Tubificidae. Archiv für Hydrobiologie 60: 385-418. BRINKHURST, R. O. 1971. A guide for the identification of British aquatic Oligochaeta. Freshwater Biological Association, Scientific Publication 22, The Ferry House, 55 pp. BRINKHURST, R. O. 1974. The benthos of lakes. The Macmillan Press, London, 190 pp. 556 B. LODS-CROZET & O. REYMOND BRINKHURST, R. O. & JAMIESON, B. G. M. 1971. Aquatic Oligochaeta of the world. Oliver & Boyd, Edinburgh, 860 pp. BUTTY, L., JEANRENAUD, D. & POKORNI-AEBI, B. 2003. Suivi de l’environnement neuchâtelois en 2002. Bulletin de la Société Neuchâteloise des Sciences Naturelles 126: 169-184. GERSTMEIER, R. 1989. Phenology and bathymetric distribution of the profundal chironomid fau- na in Starnberger See (F.R. Germany). Hydrobiologia 184: 29-42. GOEDKOOP, W. & JOHNSON, R. K. 1993. Modelling the importance of sediment bacterial carbon for profundal macroinvertebrates along a lake nutrient gradient. Netherlands Journal of Aquatic Ecology 26: 477-483. GRAF, G. 1989. Benthic-pelagic coupling in a deep-sea benthic community. Nature 341: 437-439. HAKANSON, L. 1980. 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POKORNI-AEBI, B. 2002. Suivi de la qualité des eaux du lac de Neuchatel (Suisse). Bulletin de la Société Neuchâteloise des Sciences Naturelles 125: 135-143. REIBLE, D. D., Popov, V., VALSARAJ, L.J., THIBODEAUX, F., LIN, F., DIKSHIT, M., TODARO, M.A. & FLEEGER, J.W. 1996. Contaminant fluxes from sediment due to tubificid oligochaete bioturbation. Water Research 30: 704-714. REISS, F. 1968. Okologische und systematische Untersuchungen an Chironomiden (Diptera) des Bodensees. Archiv fiir Hydrobiologie 64: 176-323. REYMOND, O. 1994. Préparations microscopiques permanentes d’oligochètes: une méthode simple. Bulletin de la Société vaudoise des Sciences naturelles 83: 1-3. REYNOLDSON, T. B. 1987. The role of environmental factors in the ecology of Tubificid Oligo- chaetes - an experimental study. Holarctic Ecology 10: 241-248. SAETHER, O. A. 1979. Chironomid communities as water quality indicators. Holarctic Ecology 2: 65-74. SMITH, S. L., MACDONALD, D. D., KEENLEYSIDE, K. A., INGERSOLL, C. G. & FIELD, L. J. 1996. A preliminary evaluation of sediment quality assessment values for freshwater ecosystems. Journal of Great Lakes Research 22: 624-638. STEINMANN, P., ADATTE, T. & LAMBERT, P. 2003. Recent changes in sedimentary organic matter from Lake Neuchâtel (Switzerland) as traced by Rock-Eval pyrolysis. Eclogae Geologicae Helvetiae 96: 109-116. TIMM, T. 1996. Tubifex tubifex (Müller, 1774) (Oligochaeta, Tubificidae) in the profundal of Estonian lakes. Internationale Revue der gesamten Hydrobiologie 81: 589-596. VERNEAUX, V. & VERNEAUX, J. 2002. Assessing lake functioning using the macrobenthic community with special reference to Chironomidae (Diptera). A subalpine lake (Lake Annecy) as an example. Archiv fiir Hydrobiologie 154: 61-78. VERNET, J. P. & VIEL, M. 1984. Métaux lourds dans les sédiments lacustres (pp. 227-238). Rapport de la Commission internationale pour la protection des eaux du Léman contre la pollution, Le Léman, synthèse 1957-1982, Lausanne. 558 B. LODS-CROZET & O. REYMOND WIEDERHOLM, T. 1980. Use of benthos in lake monitoring. Journal of the Water Pollution Control Federation 52: 537-547. WIEDERHOLM, T. 1983. Chironomidae of the Holarctic region. Keys and Diagnoses. Part 1. Larvae. Entomologica Scandinavica Suppl. 19: 1-449. WIEDERHOLM, T. 1986. Chironomidae of the Holarctic region. Keys and Diagnoses. Part 2. Pupae. Entomologica Scandinavica Suppl. 28: 1-482. REVUE SUISSE DE ZOOLOGIE 112 (2): 559-572; juin 2005 Three new species of Ancistrus Kner (Teleostei: Siluriformes: Loricariidae) from the upper Tapajés and Tocantins rivers Sonia FISCH-MULLER!, Alexandre R. CARDOSO, José F. P. da SILVA? & Vinicius A. BERTACO? 1 Muséum d’histoire naturelle, PO. Box 6434, CH-1211 Geneva 6, Switzerland. E-mail: sonia.muller@mhn.ville-ge.ch 2 Laboratério de Ictiologia, Museu de Ciéncias e Tecnologia, Pontiffcia Universidade Catélica do Rio Grande do Sul. Av. Ipiranga, 6681, 90619-900 Porto Alegre, RS, Brazil. E-mail: alexrc@pucrs.br, pezzi@pucrs.br — ubertaco@pucrs.br Three new species of Ancistrus Kner (Teleostei: Siluriformes: Lori- cariidae) from the upper Tapajés and Tocantins rivers. - Three new species of Ancistrus are described: Ancistrus tombador sp. n. from the upper rio Tapajôs basin, and Ancistrus reisi sp. n. and Ancistrus jataiensis sp. n. from the upper rio Tocantins basin. The three species differ from their congeners by the absence of an adipose fin. Loss of the adipose fin was only rarely observed in Ancistrini. In the three new species it is replaced by a series of small unpaired platelets forming a low crest. Ancistrus tombador further differs from all congeners by a unique combination of characters: naked margin of snout large and tentacles usually absent in both sexes; body very narrow (cleithral width 27.5-31.2% SL); long caudal peduncle partic- ularly depressed (depth 8.3-9.2% SL). Ancistrus reisi is distinguished from A. jataiensis by measurements, including: predorsal length (respectively: 43.8-46.4% SL versus 47.5-49.3% SL); occipital depth (14.9-17.0% SL versus 17.0-19.5% SL); and caudal peduncle length (27.7-30.9% SL, versus 24.6-27.1% SL). Keywords: Ancistrini - new species - Amazon River drainage - Brazil - catfishes - taxonomy. INTRODUCTION With 53 valid species plus numerous undescribed forms (Fisch-Muller, 1999, 2003), Ancistrus Kner, 1854 is the type genus and the most speciose genus of the Ancistrini Kner, 1853. This latter taxon was recently moved from subfamily to tribe rank in subfamily Hypostominae by Armbruster (2004). Ancistrus exhibits a wide dis- tribution in the Neotropical Region with highest species diversity observed in the Amazon system. It was found to be monophyletic on the basis of osteological (Schaefer, 1987), molecular (Montoya-Burgos et al., 1997, 1998), and biochemical (Fisch-Muller, 1999) data. Manuscript accepted 19.01.2005 560 S. FISCH-MULLER ET AL. During a field expedition of the Laboratory of Ichthyology of PUCRS (MCP) to Central Brazil in January 2002, with the exploration of upper rio Tapajés basin, a very distinct new species of Ancistrus was discovered. This new species is peculiar in the absence of the adipose fin, which is replaced by median platelets forming a low crest, a character never described for Ancistrus until present. In the same year, the MCP received some fishes from the upper Tocantins basin collected by the Centro de Biologia Aquâtica (UCG) for identification, and surprisingly two additional and new species of Ancistrus with a similar loss of the adipose fin were recognized. The objec- tive of this paper is to describe these recently discovered species, while contributing to the advancement of the “All Catfish Species Inventory”. MATERIAL AND METHODS Measurements and counts follow Fisch-Muller et al. (2001). Morphometric characters other than standard length (SL) are expressed as percentages of SL, except for sub-units of the head, which are expressed as percentages of head length (HL). Specimens are deposited in the Museu de Ciéncias e Tecnologia, Pontificia Universidade Catölica do Rio Grande do Sul, Porto Alegre (MCP) and in the Muséum d’histoire naturelle, Geneva (MHNG). Vertebrae counts, including Weberian and ural complexes, were done on radiographs of type specimens, and the plates replacing the adipose fin were examined on cleared and stained specimens (c&s) prepared according to the method of Taylor & Van Dyke (1985). Comparative material includes the pri- mary type specimens of all but six described Ancistrus species. DESCRIPTIONS OF NEW SPECIES Ancistrus tombador sp. n. Fig: 1-7Tablest Holotype. MCP 33000, male, 62.8 mm SL; Brazil: Mato Grosso: Porto dos Gauchos: igarapé Ribeirào Preto, on the road MT-338, about 26 km SE from Porto dos Gaüchos (11°39°27”S, 57°12°7”W), a tributary of rio Arinos, upper rio Tapajos basin; coll. R. E. Reis, L. R. Malabarba, E. H. Pereira, V. A. Bertaco, and A. R. Cardoso, 19 Jan 2002. Paratypes. Brazil: Mato Grosso: rio Arinos basin, rio Tapajos drainage. MCP 33001, 11 (10 measured; 1 c&s), 24.9-57.6 mm SL; MHNG 2647.55, 5, 35.1-55.6 mm SL; collected with the holotype. MCP 33002, 2 (1 measured), 13.7-52.9 mm SL, stream on the road MT-338, about 39 km N from Tapurah (12°1’49”S, 56°33’59”W), Tapurah, same collectors, 19 Jan 2002. MCP 33004, 1, 52.0 mm SL; stream in the road between the ferryboat of rio Arinos and Tapurah, about 6 km E from Tapurah, same collectors, 18 Jan 2002. DIAGNOSIS. Ancistrus tombador is easily distinguished from all congeners — except A. reisi and A. jataiensis by the absence of an adipose fin. A series of 3 to 5 median, unpaired platelets forming a low crest is present in place of the adipose fin. These platelets are followed by at least two dermal plates (more often three) of the dorso-lateral series that join dorsally, while in A. reisi and jataiensis they are nearly immediately followed by the plate-like procurrent caudal rays. Ancistrus tombador further differs from all described Ancistrus species by the unique combination of the following characters: naked margin of snout large and tentacles usually absent in both sexes, body very narrow (cleithral width 27.5-31.2% SL, mean 29.4), and long caudal peduncle particularly depressed (depth 8.3-9.2% SL, mean 8.7). NEW ANCISTRUS FROM THE AMAZON DRAINAGE Fic. 1 Ancistrus tombador sp. n., holotype, MCP 33000, male, 62.8 mm SL; Brazil: Mato Grosso: igarapé Ribeirào Preto, upper Tapajos basin. DESCRIPTION. Morphometrics and meristics presented in Table 1. Possibly small-sized species; body very narrow at origin of dorsal fin, narrowing progressively and regularly to caudal peduncle end; head and body strongly depressed, postoccipital elevation weak when present, caudal peduncle very low. 562 S. FISCH-MULLER ET AL. Snout rounded with large naked margin in both sexes. Tentacles absent in both sexes except in one specimen (male, 52.9 mm SL) with single small tentacle on dorso- lateral extremity of naked area. Eye medium-sized, dorsal margin of orbit elevated, interorbital area slightly concave. Exposed part of opercle roughly triangular with posterior part elongated; dermal plates of postopercular area never numerous, generally large and contiguous with pterotic-supracleithrum, leaving large naked area around opercle. Evertible cheek odontodes short, not reaching posterior margin of opercle, with fleshy base variable in length and thickness. Oral disk roughly circular, lips covered with minute papillae. Lower lip large but not reaching pectoral girdle, its border with very small papillae. Maxillary barbel short, about as long as buccal papilla. Mandibular tooth row short, premaxillary tooth row slightly wider; teeth numerous, bifid, with main cusp large; lateral cusp minute, pointed, never reaching more than one third mesial cusp length. Supraoccipital plate usually well delimited from surrounding plates, particular- ly from central plate of first predorsal row. Curved to triangular nuchal plate always present before dorsal-fin spinelet. Five series of lateral plates; mid-dorsal, and mid- ventral series ending at level of small median platelets replacing the adipose fin. Last plate in median series usually of similar size and shape that penultimate plate. Odontodes present on body plates except along dorsal-fin base and on widely extended area below anal fin; odontodes generally short, only very slightly longer on ventral margin of opercle and on pectoral-fin spine of males. Abdomen entirely devoid of plates. Presence of small preanal platelet (first anal-fin pterygiophore) covered with odontodes. Dorsal-fin origin slightly anterior to pelvic-fin origin; dorsal fin short, when laid back its tip not reaching series of median platelets replacing adipose fin. Adipose fin absent, replaced by series of 3 to 5 small median platelets between lateral scutes, forming slightly raised crest. These platelets separated from usual median platelets preceeding caudal fin (procurrent caudal rays) by 2 or 3 plates of dorso-lateral series that join dorsally. Pectoral spine reaching anterior third of ventral spine. Anal fin short. Caudal fin short, slightly concave. Fin-ray formulae: D 1,7; P 1,6; V 1,5; A 14; Ci, 13, e) ORCH GG EX): Vertebrae: 28 (holotype and one paratype) COLORATION IN ALCOHOL. Head brownish; body dorsally and laterally brownish with four or five weak ill-defined paler bands; inferior part of caudal peduncle paler. Insconspicuous small round spots on naked margin of snout only, or up to dorsal-fin origin. These spots generally whitish and sprinkled with dark chromatophores; dark chromatophores sometimes numerous on whitish area, thus resulting in dark spots in- stead of ligth spots (more often in small specimens). Lips yellowish. Ventral surface of head and abdomen pale brown to yellowish. Fin rays generally brown spotted whitish to orange; membranes hyaline, unpigmented at least in their middle. Spots sometimes forming bands on dorsal and caudal fins, with tips yellowish to orange-colored (Fig. 1). DISTRIBUTION. 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BOITO GOL IGES LI sc SU] ajounpod yepneD SIT L'CT9:0T 9 607 CNG “SCEVOE EI TE 610, SIETE SU] [eurWOpgy SIT IITETET OST SEC SIVTTET El VEC STE SV. ICONE aC UISUO] IOVIOYI OTT GIETOEC 97657 LIVORNO = tall OSG UG IGO AE SV Suo] ourds ITAJOd V9C SIEGES Se SiC CLEC BUS SISTER CRT SIL CAVO LS CHE IEESESC Suv] ourds [e.10)99d CCC DEGNA bee Lic OT El Sic? Vie O€CS6l LI Ot? Su] seq UY [ESI0( 6 CC AALST CVC € SOL TT OL YET Iye= OT SCT LL SET Su] surds pesto] 68€ GOV-G9IE I Gov. Pe Gikeniiaze Bl SLs SiS Ce OIL EOI. eee ORE sus] peoH (00/43 SES, Do ESE Sanger ts. fail Sis OG GNSS LEG NEI EST UIPIM [ey We) 81 S6IOLI 9 él OO Oil ek 6591 COTONE I 69 ‘pdop [endi9o0 (14 COCA G/L? OC Gr Ov VASE el LS (AA ESA GAI Wavy, Yous] [ESIOPIIT HLONAT ANVANVLS YO SLNIINAA 6 St 07S-S0r 9 OFS PISTES ID ICE ETES 109 697 STEVE LI 879 (wur) ySua] PIEPueIS ueoW oduey u H ueoW oduey u H urn 9SUPY u H S1SU91DJDL snajsiouy 18194 SNAJSIIUV 4OPDQUIO] SNASTOUY ‘u ‘ds sisuamol snysiouy pue "u ‘ds 18194 snajsiouy “u ‘ds soppquior snajsiouy Jo sodAyesed pue (H) sodAjojoy ou} Jo vyep Snsuow pur sLNOWOYdIO| “| ATAVL 564 S. FISCH-MULLER ET AL. FIG. 2 Collection localities of Ancistrus tombador sp. n. (squares) from upper Tapajös basin, Ancistrus reisi sp. n. (circle) and Ancistrus jataiensis sp. n. (star) from upper Tocantins basin. Symbols may represent more than one locality. ETYMOLOGY. The specific name tombador (a noun in apposition) refers to the Serra do Tombador, where the type locality is situated, in Mato Grosso. Ancistrus reisi sp. n. Fig. 3; Table 1 Holotype. MCP 34818, male, 60.8 mm SL; Brazil: Goiäs: Mambaf: cörrego das Dores, tributary of the rio Vermelho (14°29’S, 46°6’W), rio Tocantins basin; coll. Centro de Biologia Aquatica - UCG, 15 Dec 2002. Paratypes. Brazil: Goiäs: Mambai: rio Tocantins basin. MCP 33924, 4 (2 measured), 19.8-49.8 mm SL, collected with the holotype. MCP 33922, 5 (4 measured; 1 c&s), 35.6-54.5 mm SL; MHNG 2652.91, 4 (3 measured), 22.4-37.8 mm SL; same locality and collector as holotype, 9 Sep 2002. MCP 33925, 7 (2 measured), 15.9-47.9 mm SL; ribeiräo das Araras, trib- utary of rio Vermelho (14°29°S, 46°6’W); same collector as holotype, 14 Dec 2002. DIAGNOSIS. Ancistrus reisi differs from all congeners except A. tombador and A. jataiensis by the absence of an adipose fin. A series of 2 to 4 median unpaired platelets forming a low crest replaces the adipose fin. These platelets are nearly imme- diately followed by the plate-like procurrent caudal rays, while in A. tombador they are followed by at least two lateral plates that join dorsally before the procurrent caudal rays. Ancistrus reisi further differs from A. tombador by several morphometric and meristic characters, by the coloration pattern, and by the snout bearing tentacles (see under diagnosis of A. tombador). It is distinguished from A. jataiensis by the following morphometric characters: predorsal length (43.8-46.4% SL, mean 44.9, NEW ANCISTRUS FROM THE AMAZON DRAINAGE 565 FIG. 3 Ancistrus reisi sp. n., holotype, MCP 34818, male, 60.8 mm SL. Brazil: Goiäs: cörrego das Dores, upper Tocantins basin. versus 47.5-49.3% SL, mean 48.1), occipital depth (14.9-17.0% SL, mean 16.0, versus 17.0-19.5% SL, mean 18.1), and caudal peduncle length (27.7-30.9% SL, mean 28.7, versus 24.6-27.1% SL, mean 26.3). DESCRIPTION. Morphometrics and meristics presented in Table 1. Small species; body large, narrowing regularly to caudal peduncle end; head strongly depressed, postoccipital elevation weak when present. 566 S. FISCH-MULLER ET AL. Snout rounded with naked margin generally ill-delimited by posterior dermal plates; these plates not regular in size and shape, and rarely joined together. Naked area in males large but never reaching nostrils (narrower in females), with fleshy tentacles (up to 20, in holotype). Tentacles restricted to dorso-lateral portion of naked area in small males (from 36.5 mm SL); also present on anterior portion of snout in larger males (from 54.3 mm SL); present in one row bordering snout as well as dorsally in largest specimen (holotype). Tentacles not branched. Eye small, dorsal margin of orbit not elevated, interorbital area slightly convex. Exposed part of opercle variable in shape but never very long; dermal platelets of postopercular area variable in shape, some large and contiguous with pterotic-supra- cleithrum, smaller anteriorly. Evertible cheek odontodes long respective to size of specimens, posterior one reaching often far beyond posterior margin of opercle, with fleshy base sometimes long and thick. Oral disk enlarged, widened laterally; lips covered with minute papillae. Lower lip large but not reaching pectoral girdle, its border formed by unequal small flaps and smooth. Maxillary barbel short. Mandibular tooth row wide, premaxillary tooth row equal in length or slightly shorter; teeth numerous, bifid, with main cusp large and long; lateral cusp minute, pointed, never reaching more than one third mesial cusp length. Supraoccipital plate usually well delimited from surrounding plates, particu- larly from central plate (or two plates) of first predorsal row. Odontodes very short on head; central part of supraoccipital slightly granular. Exposed part of nuchal plate minute or totally covered with skin and dorsal-fin spinelet often reduced. Five series of lateral plates; mid-dorsal and mid-ventral series ending at level of median platelets replacing adipose fin. Last plate of median series sometimes smaller than penultimate plate. Odontodes present on body plates except along dorsal-fin base and on widely extended area below anal fin; odontodes slightly longer on ventral margin of opercle and on pectoral-fin spine of males. Abdomen entirely devoid of plates. First anal-fin pterygiophore totally covered by skin. Dorsal-fin origin slightly anterior to pelvic-fin origin; dorsal fin short, when laid back its tip not reaching median platelets replacing adipose fin. Adipose fin absent, replaced by 2 to 4 small median platelets between lateral scutes, forming slightly raised crest. These platelets immediately followed by smaller and unraised median platelets (procurrent caudal rays) preceeding caudal fin. Pectoral spine reaching anterior third of ventral spine. Anal fin short. Caudal fin short, slightly concave. Fin-ray formulae: D 1,7; Pi,6; V 1,5; A 1,4; C i, 13, i (4 specimens, including holotype) or more generally 1,14,1. Vertebrae: 28 (holotype and one paratype) COLORATION IN ALCOHOL. Dorsal surface brown, with undefined brown-reddish or brown-yellowish areas: small and rounded lighter spots often present on snout but never very contrasted (including holotype; Fig. 2). Ventral surface lighter; lips yello- wish; belly yellowish (rarely) to light brown; when pigmented, chromatophores pre- sent on its entire surface including central part, and unspotted. Fin rays generally brown, finely spotted with whitish to orange, membranes hyaline, unpigmented at least in their middle. Except in largest specimens, spots usually forming narrow bands on caudal fin; caudal fin with lower and upper tips often yellowish to orange. NEW ANCISTRUS FROM THE AMAZON DRAINAGE 567 DISTRIBUTION. Ancistrus reisi was collected in small rivers of the upper Tocantins drainage in Tocantins State (Fig. 2). ETYMOLOGY. The new species is named after Roberto Reis for his active contri- bution to the knowledge of the Neotropical ichthyofauna. Ancistrus jataiensis sp. n. Fig. 4; Table 1 Holotype. MCP 35244, male, 54.0 mm SL; Brazil: Goiäs: Mambaï: cörrego Jataf, tributary of the rio Vermelho (14°29’S, 46°6’W), rio Tocantins basin; coll. Centro de Biologia Aquâtica - UCG, 8 Sep 2002. Paratypes. MCP 33921,3 (1 c& s), 41 .6-50.6 mm SL; MHNG 2652.92, 1,44.2 mm SL; collected with the holotype. MCP 33923, 1, 40.5 mm SL; same locality and collector as holotype, 15 Dec 2002. DIAGNOSIS. Ancistrus jataiensis differs from all congeners except A. tombador and A. reisi by the absence of an adipose fin. A series of 2 to 4 median unpaired platelets forming a low crest replaces the adipose fin. These platelets are nearly imme- diately followed by the plate-like procurrent caudal rays, while in A. tombador they are followed by at least two lateral plates that join dorsally before the procurrent cau- dal rays. Ancistrus jataiensis further differs from A. tombador by several morphomet- ric and meristic characters, by the coloration pattern, and by the snout bearing tenta- cles (see under diagnosis of A. tombador). It is distinguished from A. reisi by the fol- lowing morphometric characters: predorsal length (47.5-49.3% SL, mean 48.1, versus 43.8-46.4% SL, mean 44.9), occipital depth (17.0-19.5% SL, mean 18.1, versus 14.9- 17.0% SL, mean 16.0), and caudal peduncle length (24.6-27.1% SL, mean 26.3, versus 27.7-30.9% SL, mean 28.7). DESCRIPTION. Morphometrics and meristics presented in Table 1. Small species; body thickset, predorsally long and large, with short caudal peduncle; head moderately depressed, weak postoccipital elevation. Snout rounded with naked margin generally ill-delimited by posterior dermal plates; these plates not regular in size and shape, and rarely joined together. Naked area relatively large in males (narrower in females), with fleshy tentacles (up to 27, in holotype). Tentacles present at dorso-lateral portion of naked area and on anterior portion of snout in males from 41.6 mm SL; in one row bordering snout as well as dorsally in largest specimens (from 50.6 mm SL). Some tentacles branched in larger males (including holotype). Eye very small, dorsal margin of orbit not elevated; interorbital large, particu- larly in large specimens, and slightly convex. Exposed part of opercle of variable shape but never very long; dermal platelets of postopercular area variable in shape, generally large and contiguous with pterotic-supracleithrum. Evertible cheek odontodes strong and numerous, long respective to size of specimens, posterior one reaching far beyond posterior end of opercle in largest specimens (including holotype), with fleshy base sometimes long, thick and branched. Oral disk enlarged, widened laterally; lips covered with minute papillae. Lower lip reaching or almost reaching anterior margin of pectoral girdle, its border formed by unequal small flaps, and smooth. Maxillary barbel short. Mandibular tooth row wide, premaxillary tooth row equal in length or slightly shorter; teeth numerous, bifid, with 568 S. FISCH-MULLER ET AL. Fic. 4 Ancistrus jataiensis sp. n., holotype, MCP 35244, male, 54.0 mm SL. Brazil: Goiäs: cörrego Jataî, upper Tocantins basin. main cusp large and long; lateral cusp minute, pointed, never reaching more than one third mesial cusp length. Supraoccipital plate usually hardly delimited from surrounding head plates; well delimited from central plate (or two plates) of first predorsal row. Odontodes very short on head; central part of supraoccipital slightly granular. Exposed part of nuchal plate minute or totally covered with skin and dorsal-fin spinelet often reduced. Five series of lateral plates; mid-dorsal and mid-ventral series ending at level of median platelets replacing adipose fin. Last plate of median series rarely smaller than penul- NEW ANCISTRUS FROM THE AMAZON DRAINAGE 569 timate plate. Odontodes present on body plates except along dorsal-fin base and on widely extended area below anal fin; odontodes slightly longer on ventral margin of opercle and on pectoral-fin spine of males. Abdomen entirely devoid of plates. First anal-fin pterygiophore totally covered by skin. Dorsal-fin origin slightly anterior to pelvic-fin origin; dorsal fin short, when laid back its tip not reaching median platelets replacing adipose fin. Adipose fin absent, replaced by 2 to 4 small median platelets between lateral scutes, forming slightly raised crest. These platelets immediately followed by smaller and unraised median platelets (procurrent caudal rays) preceeding caudal fin. Pectoral spine reaching anterior third of ventral spine. Anal fin short. Caudal fin short, slightly concave. Fin-ray formulae: D 1,7; P16; V15; Ai4; Ci,l4,1. Vertebrae: 28 (holotype and one paratype) COLORATION IN ALCOHOL. Dorsal surface light reddish-brown to dark brown (including holotype; Fig. 4); small and rounded lighter spots present on snout or on entire head (holotype), rarely very contrasted. Ventral surface slightly lighter; lips yellowish; belly light brown to dark brown, with chromatophores present on its entire surface including central part, and usually covered with light spots of variable shape. Fins brownish similar to dorsal surface, spotted with whitish to orange, membranes variably pigmented; spots usually forming bands on caudal fin, with upper and lower tips often yellowish to orange. DISTRIBUTION. Ancistrus jataiensis was found only at the type locality, a small tributary of the rio Vermelho, upper Tocantins basin (Fig. 2). ETYMOLOGY. The specific epithet jataiensis (an adjective) is derived from Jataf, the name of the only river where the species was found, in Goiäs state. DISCUSSION The absence of an adipose fin is known for several loricariids but it is quite rare for Ancistrini, where it is characteristic of the few species of the genera Acanthicus Spix & Agassiz, 1829, Leptoancistrus Meek & Hildebrand, 1916, Lipopterichthys Norman, 1935, and of two species of Lithoxus (L. pallidimaculatus Boeseman, 1982; L. surinamensis Boeseman, 1982). The adipose fin is also absent occasionally in Chaetostoma anomalum Regan, 1903 and in C. venezuelae (Schultz, 1944). The loss of the adipose fin was never observed before in Ancistrus. All described species of the genus have an adipose fin formed by a raised and curved spine and by a membrane, that often extends beyond the end of the spine. This adipose fin is preceded by one or more small median unpaired bony plates, named pre-adipose scutes in Schaefer (1987). In the three Ancistrus species described here, a series of unpaired median platelets replacing the adipose fin forms a slightly raised crest. Replacement of the adipose fin by such plates was described for Lipopterichthys, and in Hypostomini tribe for species of Corymbophanes Eigenmann, 1909 and Hemipsilichthys vestigipinnis Pereira & Reis, 1992. Ancistrus is characterized by a naked anterior snout margin with fleshy tentacles that develop during growth. The width of the naked margin, as well as the distribution, 570 S. FISCH-MULLER ET AL. number, and size of tentacles vary between sexes and, to a lesser extent, between species (Muller, 1990; Sabaj et al., 1999). In all species but A. bolivianus (Steindachner, 1915), A. aguaboensis Fisch-Muller, Mazzoni & Weber, 2001, A. minu- tus Fisch-Muller Mazzoni & Weber, 2001, and an undescribed Peruvian species, the naked area is very narrow along the entire snout margin in females while it is much wider at least laterally on the snout in both small and large males, allowing for sex identification. Ancistrus tombador is thus one additional described Ancistrus species for which sex cannot be easily determined on the basis of this character. The possibi- lity that specimens described here are all immatures, and that this species grows to a much larger size than observed cannot be excluded. However, in comparison to all examined material, it is evident that it is characterized by the poor development of these fleshy protuberances. The genus name Xenocara Regan, 1904, a synonym of Ancistrus which is still sometimes found in the aquarium trade, was used by some authors to include species without tentacles, as proposed by Eigenmann (1905, 1910). However, we have never observed any Ancistrus species [including Ancistrus latifrons (Giinther, 1869), type species of Xenocara] completely without tentacles. Ancistrus reisi and A. jataiensis from the upper Tocantins basin are quite similar to each other in appearance, as they are also to A. aguaboensis and A. minutus. The latter two are very likely endemic from the upper rio Tocantins basin and were collected at the Serra da Mesa dam. However, while they were found to be cryptic, A. reisi and A. jataiensis can be distinguished by several morphometric characters. The body of A. jataiensis is more elongated anteriorly and shortened at the caudal peduncle; it is deeper and larger than in A. reisi, it shows a larger interorbital and inter- nostril, a smaller eye relative to head length, and has shorter paired and unpaired fins except for the anal fin. In addition to the absence of the adipose fin, the two new species each differ from A. aguaboensis and A. minutus by several measurements, counts, and the color pattern. Other known Ancistrus species from the upper Tocantins basin are Ancistrus cryptophthalmus Reis, 1989, a blind species living in caves of the rio Angélica-Bezerra system, and at least one undescribed epigean species found in the same area (Fisch-Muller er al., 2001). Reports on the ichthyofaunal composition of the Tocantins and Tapajos rivers are rare (e.g. Santos ef al., 1984; Merona, 1987; Miranda & Mazzoni, 2003) and the ichthyofauna of the upper reaches of these basins is still poorly known. However the Tocantins river basin was pointed out as an area of endemism for several Neotropical freshwater fish groups by different authors (e.g. Vari, 1988; Menezes & Lucena, 1998; Lima & Moreira, 2003). In its upper part especially, it also appears to represent an area of high endemism for the Ancistrini as shown by the presence of three recently disco- vered Hemiancistrus species (Cardoso & Lucinda, 2003), and by at least six distinct and apparently endemic species of Ancistrus including those described here. ACKNOWLEDGEMENTS The Central Brazil Expedition was supported by the project “Conhecimento, conservaçäo e utilizaçäo racional da diversidade da fauna de peixes do Brasil” (PRONEX/CNPgq, proc. 661058/1997-2), coordinated by Naércio A. Menezes. We also thank Francisco L. T. Garro (UCG), who first collected the specimens of Ancistrus reisi NEW ANCISTRUS FROM THE AMAZON DRAINAGE 571 and À. jataiensis during research on the upper streams of the Tocantins basin. We are particularly grateful to Heraldo Britski, Volker Mahnert, Roberto Reis, and Claude Weber for their useful comments on the manuscript, to Bernard Landry and Andreas Schmitz for reading the proof, and to Florence Marteau for appreciation of the figures. REFERENCES ARMBRUSTER, J. W. 2004. Phylogenetic relationship of the suckermouth armoured catfishes (Loricariidae) with emphasis on the Hypostominae and the Ancistrinae. Zoological Journal of the Linnean Society 141: 1-80. CARDOSO, A. R. & LUCINDA, P. H. F. 2003. Three new species of Hemiancistrus (Teleostei: Siluriformes: Loricariidae) from the rio Tocantins basin with comments on the genus. Ichthyological Exploration of Freshwaters 14(1): 73-84. EIGENMANN, C. H. 1905. The mailed catfishes of South America (Loricariidae). Science 21: 792- 795: EIGENMANN, C. H. 1910. Catalogue of the fresh-water fishes of tropical and South temperate America (pp. 375-511). In: SCOTT, W. B. (ed.). 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Phylogenetic relations- hips of the Loricariidae (Siluriformes) based on mitochondrial rRNA gene sequences (pp. 363-374). In: MALABARBA, L. R., REIS, R. E., VARI, R. P., LUCENA, Z. M. S. & Lucena, C. A. S. (eds). Phylogeny and Classification of Neotropical Fishes. Edipucrs, Porto Alegre, 603 pp. MULLER, S. 1990. Etude méristique et morphométrique d’Ancistrus piriformis Muller et Ancistrus pirareta Muller (Pisces, Siluriformes, Loricariidae). Revue suisse de Zoologie 97: 153-168. ST? S. FISCH-MULLER ET AL. SABAJ, M. H., ARMBRUSTER, J. W. & PAGE, L. M. 1999. Spawning in Ancistrus (Siluriformes: Loricariidae) with comments on the evolution of snout tentacles as a novel reproductive strategy: larval mimicry. Ichthyological Exploration of Freshwaters 10: 217-229. SANTOS, G. M. Dos, JÉGU, M. & MERONA, B. DE 1984. Catalogo de peixes comerciais do baixo rio Tocantins. Eletronorte/CNPq/INPA, Manaus, 83 pp. SCHAEFER, S. A. 1987. Osteology of Hypostomus plecostomus (Linnaeus), with a phylogenetic analysis of the loricariid subfamilies (Pisces: Siluroidei). Contributions in Science 394: iii-iv, 1-31. TAYLOR, W. R. & VAN DYKE, G. C. 1985. Revised procedures for staining and clearing small fis- hes and other vertebrates for bone and cartilage study. Cybium 9 (2): 107-119. VARI, R. P. 1988. The Curimatidae, a lowland neotropical fish family (Pisces: Characiformes): Distribution, endemism, and phylogenetic biogeography (pp. 343-377). In: VANZOLINI, P. E. & HEYER, W. R. (eds). Proceedings of a workshop on Neotropical distribution pat- terns. Academia brasileira de Ciéncias, Rio de Janeiro, 488 pp. REVUE SUISSE DE ZOOLOGIE Tome 112 — Fascicule 2 HONORARY MEMBERSHIP TO PROF. S.C. STEARNS .................... Hou, Zhong-E & Li, Shugiang. Gammarus species from River Jumahe, China (Crustacea, Amphipoda, Gammaridae) ................... Zicsi, Andrés & CUENDET, Gerard. Über eine neue Octodrilus-Art aus Erankreichi(@ligochaeta: BEumbrieidae) 2 men ee, LIENHARD, Charles. Description of a new beetle-like psocid (Insecta: Pso- coptera: Protroctopsocidae) from Turkey showing an unusual sexual MOD AIS TES A Na LE, JALOSZYNSKI, Pawel. Revision of the Oriental Genus Loeblites Franz (ColeopterasSeydmaenidae)e a EE RE RE n PRENDINI, Lorenzo. On Hadogenes angolensis Lourenço, 1999 syn. n. (Scorpiones, Liochelidae), with a redescription of H. taeniurus (dihore ll IS TO) ER og Coph RS pue Pere APR M CASCIOTTA, Jorge R., ALMIRON, Adriana E. & AZPELICUETA, Marfa de las Mercedes. Astyanax pampa (Characiformes, Characidae), a new species from the southernmost boundary of the Brazilian subregion, ANO SMART E ERICE e Re le SCHATTI, Beat & STUTZ, Andrea. Morphology and systematic status of Coluber karelini mintonorum Mertens, 1969 (Reptilia: Squamata: Colonne) sé romans EN era MAHUNKA, Sandor. Oribatids from Brunei IV (Acari: Oribatida). (Acaro- losica GenavensialeN Wainer trea et scent EN KARAMAN, Ivo M. Trojanella serbica gen. n., sp. n., a remarkable new troglobitic travunioid (Opiliones, Laniatores, Travunioidea) . ....... SCHELLER, Ulf. First records of Pauropoda (Millotauropodidae; Pauropo- didae) from Gabon with the description of 16 new species (Pauropoda and Symphyla of the Geneva Museum XIV) .................... SCHMITZ, Patrick & LANDRY, Bernard. Two new taxa of Galagete (Lepidoptera, Autostichidae) from the Galapagos Islands, Ecuador . . . MERZ, Bernhard & ROHACEK, Jindrich. The Western Palaearctic species of Stenomicra Coquillett (Diptera, Periscelididae, Stenomicrinae), with description of a new species of the subgenus Podocera Czerny ..... ROUGEMONT, Guillaume DE. Synonymic note on the monobasic genus Ophryomedon Wasmann, 1916 (Coleoptera, Staphylinidae, Paederi- MAS E RA EEA yk IE alli ie) no a LODS-CROZET, Brigitte & REYMOND, Olivier. Ten years trends in the oligo- chaete and chironomid fauna of Lake Neuchatel (Switzerland) ...... FISCH-MULLER, Sonia, CARDOSO, Alexandre R., SILVA, José F. P. DA & BERTACO, Vinicius A. Three new species of Ancistrus Kner (Teleostei: Siluriformes: Loricariidae) from the upper Tapajés and Tocantins TV ETS Er RO PA SOE Ren RR ERI Pages 311-312 313-327 329-331 333-349 351-369 371-399 401-408 409-420 421-438 439-455 457-509 511-517 519-539 541-542 543-558 559-572 REVUE SUISSE DE ZOOLOGIE Volume 112 — Number 2 HONORARY MEMBERSHIP TO PROF. S.C. STEARNS .................... Hou, Zhong-E & Li, Shugiang. Gammarus species from River Jumahe, China (Crustacea, Amphipoda, Gammaridae) ................... Zicsi, Andras & CUENDET, Gerard. On a new Octodrilus species from Érance (OligochaetatEumbhrcidie) CEE RE LIENHARD, Charles. Description of a new beetle-like psocid (Insecta: Pso- coptera: Protroctopsocidae) from Turkey showing an unusual sexual MOD ALS TN HE OT OLI NE CR RE EE JALOSZYNSKI, Pawel. Revision of the Oriental Genus Loeblites Franz (@oleoptera, Scydmaenidac) LI E PRENDINI, Lorenzo. On Hadogenes angolensis Lourenco, 1999 syn. n. (Scorpiones, Liochelidae), with a redescription of H. taeniurus (Hotel RS): RE are RS Le TEE CASCIOTTA, Jorge R., ALMIRÖN, Adriana E. & AZPELICUETA, Maria de las Mercedes. Astyanax pampa (Characiformes, Characidae), a new species from the southernmost boundary of the Brazilian subregion, INTÉENTINAR U TI ee DDR SCHATTI, Beat & STUTZ, Andrea. Morphology and systematic status of Coluber karelini mintonorum Mertens, 1969 (Reptilia: Squamata: (COMBINATE RATIO OI MAHUNKA, Sandor. Oribatids from Brunei IV (Acari: Oribatida). (Acaro- logiealGenayensiaeNDr LIL ISEE KARAMAN, Ivo M. Trojanella serbica gen. n., sp. n., a remarkable new troglobitic travunioid (Opiliones, Laniatores, Travunioidea) . ....... SCHELLER, Ulf. First records of Pauropoda (Millotauropodidae; Pauropo- didae) from Gabon with the description of 16 new species (Pauropoda and Symphyla of the Geneva Museum XIV) .................... ScHMITZ, Patrick & LANDRY, Bernard. Two new taxa of Galagete (Lepidoptera, Autostichidae) from the Galapagos Islands, Ecuador . . . MERZ, Bernhard & ROHACEK, Jindrich. The Western Palaearctic species of Stenomicra Coquillett (Diptera, Periscelididae, Stenomicrinae), with description of a new species of the subgenus Podocera Czerny ..... ROUGEMONT, Guillaume DE. Synonymic note on the monobasic genus Ophryomedon Wasmann, 1916 (Coleoptera, Staphylinidae, Paederi- DIARI ri N ot A LoDS-CROZET, Brigitte & REYMOND, Olivier. Ten years trends in the oligo- chaete and chironomid fauna of Lake Neuchatel (Switzerland) ...... FISCH-MULLER, Sonia, CARDOSO, Alexandre R., SILVA, José F. P. DA & BERTACO, Vinicius A. Three new species of Ancistrus Kner (Teleostei: Siluriformes: Loricariidae) from the upper Tapajés and Tocantins TANCES i OR RR EA LE Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 311-312 313-327 329-331 333-349 351-369 371-399 401-408 409-420 421-438 439-455 457-509 511-517 519-539 541-542 543-558 559-572 PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N©s 1-17 (1908-1926) ........ série Fr. (prix des fascicules sur demande) REMUEIDEPALÉOBIOLOGIE Ss 22) EN Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris) ....... par fascicule Fr. THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION, WITH KEYS FOR DETERMINATION ISIN OSERG S45 ID AMONT RE OE O PR EE Em te Fr. CLASSIFICATION OF THE DIPLOPODA RATESEIOFEMANG2 Sil Dis 99 auc a En e Fr. LES OISEAUX NICHEURS DU CANTON DE GENÈVE P. GÉROUDET, C. GUEX & M. MAIRE Sep AnomDIEUSESICATteS CUM SUTES SLOSS ce TEO ees iets Fr. CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ A LA CONNAISSANCE DES ECHINODERMES ACTUELS MATANGOUX Gap MOSS PSM ie TI ANNE RAR SE spats Meets ac EM Seele ee Fr. RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) MAEINER J NVUESD ec KSMAREDA 02,0. 1998 ran NET Er Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: SHELLS AND RADULAS V@BEINETSIAWUESTI:SAK VIARED A992) iia RTE RER SERRE crete Er: O. SCHMIDT SPONGE CATALOGUE R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 p., 1992...................... „Er: ATLAS DE REPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE AS KEIDER, VITAELLENI GV MAHNERT,48P., 1993. 20.......000 00.02. Fr. THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST NAIEINER US Os MOOS reas eee es Mo en Werk Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE Ut Ce CNIS POSTO LE OO Fr. 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) (Instrumenta Biodiversitatis I), I. LOBL, xii + 190 p., 1997 ...................... Fr. CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis II), 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, SIP LIDO E E ee TAR SIONE ICON IDIOTI DION Ico e, NIETO Fr. PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY (Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS, ALMA SD 2002 RAP de doi deo api e rai so, ino aaa e E Fr. REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) (Instrumenta Biodiversitatis VI), G. SABELLA, CH. BUCKLE, V. BRACHAT CACSBESUCHETAVIC2 83 (Daj AO ERE RO TSI Fr. 285.— 35.— 35.— 30.— 30.— 45.— 60.— 70.— 180.— 100.— avan sc | (‘dia ITS À a sap sata ta RES un T N ot pan Gy: othe ete Ed Volume 112 - Number 2 - 2005 Revue suisse de Zoologie: Instructions to Authors The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific results based on the collections of the Muséum d'histoire naturelle, Geneva. Submission of a manuscript implies that it has been approved by all named authors, that it reports their unpublished work and that it is not being considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of colour plates and large manuscripts. All papers are refereed by experts. In order to facilitate publication and avoid delays authors should follow the Instructions to Authors and refer to a current number of R.S.Z. for acceptable style and format. Papers may be written in French, German, Italian and English. Authors not writing in their native language should pay particular attention to the linguistic quality of the text. Manuscripts must be typed or printed (high quality printing, if possible by a laser-printer), on one side only and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins must be at least 25 mm wide. Authors must submit three paper copies (print-outs), including tables and figures, in final fully corrected form, and are expected to retain another copy. Original artwork should only be submitted with the revised version of the accepted manuscript. We encourage authors to submit the revised final text on a disk (3,5”), or on a CD-R, using MS-WORD or a similar software. The text should be in roman (standard) type face throughout, except for genus and species names which should be formatted in italics (bold italics in taxa headings) and authors’ names in the list of references (not in other parts of the text!), which should be formatted in SMALL CAPITALS. LARGE CAPITALS may be used for main chapter headings and SMALL CAPITALS for subordinate headings. Footnotes and cross-references to specific pages should be avoided. Papers should conform to the following general layout: Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, full name(s) and surname(s) of author(s), and full address(es) including e-mail address(es) if possible. Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should summarise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed by up to 10 keywords, separated by hyphens, which are suitable for indexing. Some of the terms used in the title may be omitted from the list of keywords in favour of significant terms not mentioned in the title. Introduction. A short introduction to the background and the reasons for the work. Material and methods. Sufficient experimental details must be given to enable other workers to repeat the work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen- clature must be strictly followed. Cite the authors of species on their first mention. Results. These should be concise and should not include methods or discussion. Text, tables and figures should not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp. n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by synonyms, material examined and distribution, description and comments. All material examined should be listed in similar, compact and easily intelligible format; the information should be in the same language as the text. Sex symbols should be used rather than “male” and “female” (text file: $= 6, £= 9). Discussion. This should not be excessive and should not repeat results nor contain new information, but should emphasize the significance and relevance of the results reported. References. The author-date system (name-year system) must be used for the citation of references in the text, e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown et al. (1995) or (Brown et al., 1995; White et al., 1996) should be used. In the text authors’ names have to be written in standard type face. However, in the list of references they should be formatted in SMALL CAPITALS (see below). The list of references must include all publications cited in the text and only these. References must be listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated for each reference. The title of the paper and the name of the journal must be given in full in the following style: PENARD, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. PENARD, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61. MERTENS, R. & WERMUTH, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: WENZEL, R. L. & TIPTON, V. J. (eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit 122 x 180 mm, each table on a separate sheet and numbered consecutively. Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain from mixing drawings and half tones. If possible, originals of figures (ink drawings, photographs, slides) should be submitted (together with the revised version of the accepted manuscript). Original drawings will not be returned automatically. The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. 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