5,51 sh HE IN DI DE Hi 4 DR) 2 RAA , ves 1,45. re ee IR DER Hu yt CAL No. a LA: # 1,59 PARA LA) Pa DT HU , RIAD MEE) CHEN a tra SUR AL 1 ry ig’ 1943, BELA areata DALIA ry LALA AZIO di di Int, [AA } at) Dt) n è dr CA in, HE , OCR DI “à ; MENO OAT DI gl; UV D PARU 1 (AC AA EU # wy in? i ' A aitu LIT RAT #, Ù FX Au ' rà M LEY MAR an HAE ER NUE Un) tot AUS mn NA { nn Ay DUMAS, yy nd a NA MONS ni sr CRUE NA hi MR ORLA tat CRON EAN BR ah KERN sun wd 3 HOCH Meat its a N (HAE ar OU M A He à (NA Y ae ON MU fi year HAE Wrote | DP A an wy ey oy DAI (A ACT NA HAE faz ise ONE x mira una a SUIS 4 3%, TARDE n rac ATA 244 MIRI » AR À ls Woe ON i 1 4 Tome 99 Fascicule 3 1992 REVUE SUISSE ZOOLOGIE ANNALES DELA SOCIÉTÉ SUISSE DE ZOOLOGIE © ET DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE GENEVE IMPRIMERIE SITO SEPTEMBRE 1992 ISSN 0035-418X REVUE SUISSE DE ZOOLOGIE TOME 99 — FASCICULE 3 Publication subventionnée par la Société helvétique des Sciences naturelles et la Société suisse de Zoologie VOLKER MAHNERT Directeur du Muséum d'Histoire naturelle de Genève FRANÇOIS BAUD Conservateur au Muséum d'Histoire naturelle de Genève DANIEL BURCKHARDT Chargé de recherche au Muséum d'Histoire naturelle de Genève Comité de lecture Le président de la Société Suisse de Zoologie Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des vertébrés — Muséum de Genève Le président du comité: Claude BESUCHET — Systématique des Insectes — Muséum de Genève Patrick GUÉRIN — Physiologie et éthologie des arthropodes — Institut de Zoologie, Neuchâtel Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchatel Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchâtel Olivier RIEPPEL — Morphologie, Paléontologie — Paläontologisches Institut, Zürich Paul SCHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f. Zoologie, Basel Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel Beat TSCHANZ — Ethologie des Vertebres — Ethologische Station Hasli, Bern Claude VAUCHER — Systématique des Invertébrés — Muséum de Genève La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie, systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 PRIX DE L'ABONNEMENT DES 1972: SUISSE ER225 UNION POSTALE Fr. 230. — (en francs suisses) Les demandes d'abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'Histoire naturelle, Genève Tome 99 | Fascicule 3 1a + 7 1992 REVUE SUISSE ZOOLOGIE ANNALES DELA SOCIÉTÉ SUISSE DE ZOOLOGIE ET DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE GENEVE IMPRIMERIE SITO SEPTEMBRE 1992 REVUE SUISSE DE ZOOLOGIE TOME 99 — FASCICULE 3 Publication subventionnée par la Société helvétique des Sciences naturelles et la Société suisse de Zoologie VOLKER MAHNERT Directeur du Muséum d'Histoire naturelle de Genève FRANÇOIS BAUD Conservateur au Muséum d'Histoire naturelle de Genève DANIEL BURCKHARDT Chargé de recherche au Muséum d'Histoire naturelle de Genève Comité de lecture Le président de la Société Suisse de Zoologie Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des vertébrés — Muséum de Genève Le président du comité: Claude BESUCHET — Systématique des Insectes — Muséum de Genève Patrick GUÉRIN — Physiologie et éthologie des arthropodes — Institut de Zoologie, Neuchâtel Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchâtel Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchâtel Olivier RIEPPEL — Morphologie, Paléontologie — Paläontologisches Institut, Zürich Paul SCHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f. Zoologie, Basel Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel Beat TSCHANZ — Ethologie des Vertébrés — Ethologische Station Hasli, Bern Claude VAUCHER — Systématique des Invertébrés — Muséum de Genève La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie, systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 PRIX DE L'ABONNEMENT DES 1972: SUISSE ER 225 UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d'abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'Histoire naturelle, Genève | | | | | Revue suisse Zool. | Tome 99 | Fasc. 3 p. 471-627 | Genève, septembre 1992 | The Scaphidiidae (Coleoptera) of the Nepal Himalaya’ by Ivan LÖBL? With 191 figures ABSTRACT The Scaphidiidae are remarkably diverse in the Himalaya. Its more than 200 species represent approximately 15% of the total known world fauna. Most of the species are found in the wetter eastern part of the Central Himalaya and most of its presumably endemic taxa are found at altitudes ranging from 2000 m to 3500 m. Collections made in the West Himalaya are not representative for the regional fauna, and almost no data are available from the East Himalaya. The following taxa are new to science: Ascaphium ochripes sp.n., Episcaphium unicolor sp. n., Scaphidium gurung sp.n., S. holzschuhi sp.n., S. melanogaster sp.n., S. nepalense sp.n., S. thakali sp.n., Pseudobironium bicolor sp. n., P. ineptum sp.n., P. rufitarse sp.n., Baeocera cribrata sp.n., B. crinita sp.n., B. errabunda sp.n., B. laminula sp.n., B. martensi sp.n., B. mustangensis sp.n., B. reducta sp.n., B. schawalleri sp.n., B. sordidoides sp.n., B. thoracica sp.n., B. tuberculosa sp.n., Scaphi- soma adjacens sp.n. S. alacre sp.n., S. aurorae sp.n., S. baloo sp.n., S. bhareko sp.n., S. clavigerum sp.n., S. coalitum sp.n., S. fatuum sp.n., S. fratellum sp.n., S. fulcratum sp.n., S. inquietum Sp.n., S. interjectum sp.n., S. invalidum sp.n., S. jado sp.n., S. kanchi sp.n., S. nepalense sp.n., S. nima sp.n., S. pinnigerum sp.n., S. praesigne sp.n., S. sikkimense sp.n., S. simplicipenis sp.n., S. varians sp.n., Baeotoxidium yeti sp.n., Scaphobaeocera zdenae sp.n., Toxidium spectabile sp.n., Xotidium gen.n. uniforme sp.n, and Bironium nepalense sp.n. New synonymies are: Yparicum Achard, 1920 with Cyparium Erichson, 1845, Scaphidium grande var. inimpressum Pic, 1920 and S. grande var. subannulatum Pic, 1915 with Scaphidium grande Gestro, 1879, and Baeocera pseudolenta Löbl, 1979 with Baeocera lenta (Löbl, 1971). Ascaphium minor Pic, 1956 is raised to species level. Baeocera montanella is a new name proposed to replace B. montanum Löbl, 1971 (nec ! Some specimens used in this study are the results of the Himalaya Expeditions of J. Martens, No. 177. — For No. 176 see: Senckenbergiana biol., 72 (4/6) 1992. — J.M. sponsored by Deutscher Akademischer Austauschdienst and Deutsche Forschungsgemeinschaft. 2 Muséum d'Histoire naturelle, Case postale 434, CH-1211 Genéve 6. 472 IVAN LÖBL Baeocera montana (Pic, 1955)). New combinations are: Scaphidium vernicatum (Pic, 1956) from Scaphium, Scaphidium dureli (Achard, 1922) and S. coomani (Pic, 1925) from Scaphidiolum, Cyparium yunnanum (Achard, 1920) from Yparicum, Xotidium pygmaeum (Löbl, 1971), X. montanum (Löbl, 1971) and X. notatum (Löbl, 1977) from Toxidium. Diagnostic characters are given for all species of Scaphidium, Hemiscaphium, Cyparium, and Pseudobironium occurring in the area studied. Lectotypes are designated for Scaphidium arrowi Achard, S. coomani (Pic), S. harmandi Achard, S. semilimbatum Pic, S. baconi Pic, S. assamense Pic, Hemiscaphium brunneopictum Achard, Pseudobironium almoranum Achard, P. castaneum Pic, Scaphisoma minutissimum Champion, S. cribri- penne Champion and S. bedeli Achard. The genera and species occurring in the Himalaya, in the Meghalaya and in Pakistan are keyed out, as well as all Asian species of Ascaphium, Episcaphium, Scaphoxium, Toxidium and Xotidium. Distributional data are presented for all Nepalese species, and additional data are given for numerous Indian species. INTRODUCTION Mycophagy is a common feature within the Coleoptera. About half of the beetle families include mycophages species and in several families all species are possibly fungivores (LAWRENCE 1989). This trophic guild is particularly diverse in the tropics. In the Dumoga-Bone National Park in north Sulawesi 23.4% of all sampled Coleoptera were fungivores, while only 17.4% phytophages (HAMMOND 1990). Obviously all Scaphidiidae feed on fungi (NEWTON 1984), and they are decidedly more diverse and abundant than sometimes assumed (PAULIAN 1949; 1988). In samples of forest litter arthropods I have made in different parts of Asia (e.g., in Sri Lanka, India, Thailand, Indonesia, Malaysia) the scaphidiids represent 2 to 6% of the total number of beetles. Surprisingly, this is true also for collections I have made in the mountains of north India in which temperate elements (e.g., Carabidae: Trechus, Staphylinidae: Micropeplus, Quedius and Tachinus, Latridiidae: Corticariini, Cryptophagidae: Cryptophagus, Atomaria) are well represented. Most of the previous north Indian collections came from the Darjeeling district of West Bengal, and from various localities in Kumaon and Garhwal. Nepal which became accessible on a regular basis for field research since 1949, remained until recently a “terra incognita” located between these two areas with better documented scaphidiid fauna. This circumstance, and the encouragements of my fried Ales Smetana, with whom I undertook three trips to Nepal Himalaya, were significant for realising this paper. It is hoped that it increases substantially the knowledge about the scaphidiids of the Central Himalaya. As far as the more western portion of the Himalaya is concerned, recent collections from Himachal Pradesh and from north Pakistan, i.e., from the Hindu Kush and Kohistan ranges, provide both fairly signicant data. Unfortunately, almost nothing is known about the scaphidiids (and about other cryptic forest litter Arthropoda as well) from areas east of the Darjeeling district, i.e., from Bhutan and from Arunachal Pradesh. Hopefully, field research will become possible in these areas before the human activity significantly alters the natural ecosystem. SCAPHIDIIDAE OF NEPAL 473 In the beginning was field work... The descent from Phulcoki to Godawari. The author with samples of sieved forest litter in and on his rucksack (above). Camp in Induwa Khola Valley. Samples are placed in the “Winkler-Moczarski” devices (below). IVAN LÖBL 474 TS FIG. 1. Himalayan region. Circles indicate areas from which signicant collections of Scaphidiidae have been made. SCAPHIDIIDAE OF NEPAL 475 TECHNIQUES MATERIAL This study is based on adults exclusively. Most of the material studied was collected by sifting forest litter and extracted from the samples by means of the “Winkler- Moszarski” devices (see BESUCHET & al. 1987). This method was used not only by Ales Smetana and me, but also by H. Franz, J. Martens & W. Schawaller, and S. Vit. A smaller portion of the scaphidiids was taken from fungi on dead logs, under bark or under decaying pieces of wood on the forest floor. Several specimens, mainly Scaphidium, were swept from vegetation or found in Malaise traps. The flight interception traps have been used only once in Nepal by A. Smetana during his 1985 trip. It is possible that this method may provide additional information, especially on Cyparium species, which were found in large number in flight interception traps in South and North America by S. & J. Peck. In Thailand, P. Schwendinger used pit- fall traps for collecting arachnids and other forest litter arthropods. They proved to be successful for sampling scaphidiids, although I am not aware of any scaphidiids obtained by this method in the Himalaya. I have tried to accumulate material from all known sources. The only more important Himalayan collection I know of that was not examined, is that housed in the Forestry Institute in Dehra Dun. It appears not to be accessible for revision. The Himalayan material available amounted to about 8000 specimens, half of them from Nepal. I have published several papers on the Himalayan scaphidiids (LOBL 1970; 1977; 1984; 1986a; 1986b; 1986c; 1987a; 1988; 1990a), in which a number of taxa have been described. Information on several other species occuring in the Himalaya have been given in those relating to the scaphidiids of other areas (e.g. LOBL 1981; 1990b). The diagnostic characters of all these taxa are not repeated in the present study, except for characters used in the keys. All Himalayan members of Scaphidium, Hemiscaphium, Cyparium, and Pseudobironium were inadequately described, and were not at all, or only briefly, treated in my previous papers. Therefore, the diagnostic specific characters in these genera are discussed in greater detail. The acronyms of institutions which loaned material or in which material is housed are listed below. BMNH The Natural History Museum, London CNCC Canadian National Collection of Insects, Ottawa IZK Institute of Systematic and Experimental Zoology, Krakow MCSN Museo Civico di Storia Naturale, Genova FIG. 2. Political division of Napel. Only districts in which treated material was collected are numbered. District bondaries dotted, Zonal bondaries in simple brocken lines, bondaries of Development Centres in double brocken lines. a: Eastern, b: Central, c: Western, D: Far Western, e: Extreme Western Development Centre: 1. Bara; 2. Chitawan; 3. Dhading; 4. Dhankuta; 5. Dolakha; 6. Gorkha; 7. Ilam; 8; Jumla; 9. Kathmandu; 10. Kaski; 11. Lamjung; 12. Manang; 13. Mustang; 14. Myagdi; 15. Nuwakot; 16. Panchthar; 17. Parbat; 18. Parsa; 19. Patan; 20. Rasuwa; 21. Sankhua Sabha; 22. Sindhupalcok; 23. Solukhumbu; 24. Taplejung; 25. Terhathum. Synonymies and alternative transcriptions of district names: Chitawan = Chitwan; Patan = Lalitpur; Sankhua Sabha = Sankhuwasawa, Sankhuwasabha. 476 IVAN LÖBL Idle nf # BOS FIGS 3 TO 6. Prothoracal cavity with “pocket” covered by spinose structures; 3 and 4. Baeocera sp., 5 and 6. Baeocera pubiventris Löbl. SCAPHIDIIDAE OF NEPAL 477 FIGS 7 AND 8. Spinose structures of prothoracal cavity in Baeocera, detail. MHNG Museum d’Histoire naturelle, Geneva MNHN Muséum National d'Histoire naturelle, Paris NMB Naturhistorisches Museum, Basel NMP National Museum, Prague NSNT National Science Museum, Tokyo SMNS Staatliches Museum für Naturkunde, Stuttgart ZSI Zoological Survey of India These acronyms, the names of private collections, and the names of collectors are given in paranthesis, when referring to the material studied. AREA Most of the material treated in the present study was collected within the political boundaries of Nepal. Additional material comes from Himachal Pradesh, and in much lesser extent from Sikkim or other areas. All identified taxa are keyed as are a few additional unidentified ones, known to occur in the Himalayan region. The Himalaya comprises the ranges between the rivers Indus and Brahmaputra. The Karakorum and Kohistan - Baluchistan ranges in the west, and the Arakan - Chin - Yoma fold belt and the Mogok belt in the east are of the same orogenesis (MOLNAR & TAPONNIER 1977, MASCLE & al. 1990). While a reasonable set of data is available from some portions of the western ranges, hardly anything is known about the Scaphidiidae of the two eastern fold belts. Therefore, the few species described to date from Burma are not treated in this 478 IVAN LÖBL paper. For practical reasons, the Scaphidiidae from the Garo and Khasi Hills in Meghalaya, and those from Pakistan, are included in the keys, as well in the discussion on the biogeography. Most of the collected material has been found in areas shown in the Fig. 1. DISTRIBUTIONAL RECORDS Nepal occupies a central position within the Himalaya. Since 1980 is it divided into five macro-regions: Eastern, Central, Western, Far Western, and Extreme Western Development Regions (MAJUPURIA & MAJUPURIA 1983). Each of them is sub-divided into two or three Zones (listed sometimes as Provinces) which include a various number of Districts (Fig. 2). This administrative division is respected in the present study, although it does not reflect biogeography. Indeed, it is often extremely difficult to locate data from locality labels which record the provenience of specimen as “E.Nepal” or “C.Nepal” in combination with a name of a small village, as frequently seen in different collections. An attempt has been made to arrange the records in east-west or west-east direction. The names of the political units are given once only in the respective material section. The data under “type material” do not appear under “material examined”. To avoid misleading information, only one kind of transcription of geographic names is used throughout, regardless of the original spelling on locality labels. Thus all specimens from “Phulcauki” or “Lethe” are quoted as from Phulcoki or Lete. The difficulties in listing Nepalese material are due to inadequate maps, different ways of transcription and misinterpretation of local information. However, the localities of the numerous expeditions made by J. Martens & al. have been shown on maps in MARTENS (1987) and in SCHAWALLER (1991). The collecting sites of A. Smetana and me are shown on maps in SMETANA (1988). For more information see the above papers. All data are based on material I have examined. DIAGNOSTIC CHARACTERS The length of the specimens is measured from the middle of the apical pronotal margin to the inner apical angle of the elytra. Width is measured at the largest point. Size and colour pattern of teneral specimens was not taken in account. The relative length of antennomeres is measured at the same magnification (see LOBL 1974; 1990b). In Scaphisoma, the level of inner and outer apical angles of elytra is compared in dorsal view. The indicated size of the metepisternum refers to the free visible portion. Abdominal sterna are counted from the beginning of the first visible free segment, i.e., 3rd sternum. Mouthparts and genitalia are mounted in Canadian balsam. For more details see LOBL (1990b). The presence or absence of the microsculptured patches on the 6th abdominal tergum is not examined for the diagnostic purpose. It is covered by the elytra and the wings, and not easy to see. The characters of the mouthparts are not used below genus level and potential species characters in chaetotaxy have not been investigated. The prothorax includes a pair of internal cavities (or pockets) covered by dense spinose or setose structures (Figs 3 to 8) in several genera. NEWTON (1984) was the first to record the presence of this unusual feature in scaphidiids, and thought that they may function as mycangia or spore-retaining structures. These cavities are usually distinct when viewed under a stereoscopic microscope, when the prothorax is separated from the mesothorax. The fine structure as seen in scanning electron micrographs shows significant differences among the species of same the genus (1.e., Baeocera, Figs 3 to 8). I have not SCAPHIDIIDAE OF NEPAL 479 seen these cavities in any of the examined species of the Scaphium-Scaphidium groups, neither in Cyparium, Scaphisoma, Caryoscapha, Afroscaphium, Toxidium, Scaphischema, Scaphicoma, Sapitia, and Baeoceridium. They are present in all the specimens of Pseudobironium, Baeocera, Scaphobaeocera, Scaphoxium, Baeotoxidium, Xotidium, Brachynopus, Brachynoposoma and Zinda examined, and are particularly well developed in Pseudobironium. It is likely that the internal thoracic cavities evolved independly in different groups, as these genera do not form a holophyletic group. The anapleural suture separating the metasternum from the mesepisternum is always distinct in Scaphidiidae. Most members of the group exhibit a subparallel ridge or stria joined to the anapleural suture more or less close to the mesocoxal cavity (but never reaching its margin), delimiting a small area believed to be the mesepimeron by previous authors. A.F. Newton, Jr. (pers. comm) suggested that this area cannot be the mesepimeron, because of the absence of a corresponding internal suture, and because it never extends to the coxal cavity. The examination of the internal surface of the mesothorax I have made in several scaphidiids confirms his observation. However, the relative length of the visible ridge (or stria) is an useful species character in some groups, especially in.Scaphisoma. It is named “mesepimeral ridge” in this paper. Primitively, the parameres of the aedeagus are facing the ventral side of the body. In some groups the aedeagus has rotated 90 degrees (e.g., in Scaphium, Baeocera, Scaphobaeocera), with dorsal side facing the right side of the body. In this study the ventral/dorsal/lateral sides are referred to as their plesiomorphic position. Similarly, the “right” and “left” sides of the aedeagus are referred to as primitive within the abdomen which is inverted in the figures. The ventro-apical wall of the basal bulb of the median lobe is strongly sclerotized, being the site of the attachment of the parameres. This area is often more or less protruding, or even forms an apophysis longer than the basal bulb in some species of the Scaphisoma unicolor group. In this study this structure is referred as the ventral process. NATURAL HISTORY AND ECOLOGY The occurence of sporophores is inpredictible and often ephemeral. Thus, there is a tendency toward polyphagy in fungivores. In addition, polyphagy in mycophagous insects may be explained by its low cost (HANSKI 1989). The scaphidiids, which are putative primary fungivores, are generally found on living fungi (e.g. KLIMASZEVSKI & PECK 1987), although exceptions may occur. Large numbers of Scaphium castanipes Kirby have been attracted to traps made from an assortiment of fungi in the Yukon Territory (Smetana, pers.comm.), and the polyphagy in scaphidiids has been reported several times, e.g., ASHE 1984, HAMMOND & LAWRENCE 1989, LESCHEN 1988, LESCHEN & al. 1990, NEWTON 1984, Nuss 1975. Obviously, most species feed either on fruiting bodies of slime moulds (Baeocera, Scaphobaeocera), or on hyphae of Polyporaceae and Agaricales. Gilled fungi are source for few groups only, e.g., Scaphium castanipes, Cyparium (ASHE 1984, NEWTON 1984). The only biological data on Himalayan scaphidiids are given in NEWTON & STEVENSON, 1990. Details of the life history of any scaphidiid species have yet to be established. Most species seem to lay a single egg (pers. observation). Information on morphology of immature stages is summarized in NEWTON 1991, and additional pertinent information is 480 IVAN LÖBL given in ASHE 1984, KOMPANTSEV & POTOTSAKAYA 1987, LAWRENCE & NEWTON 1980, and LESCHEN 1988. Three larval instars are recorded for Caryoscapha americanum. The larval development in Baeocera is very rapid, taking only a few days (NEWTON 1991), pupae in Scaphisoma and Caryoscapha were observed after 11 days, and adults emerged 17 to 41 days after collecting date (LESCHEN 1988). Scaphidiids occur from spring to winter in temperate climate (LÖBL & STEPHAN 1993; pers. observation). Usually, Himalayan records are from samples made during the spring and in autumn. This tends to reflect the sampling activity rather than the phenology: heavy snowfall at higher elevation and drought at lower altitudes make collecting difficult, if not impossible in winter. Collecting mycophagous beetles such as scaphidiids is harder during the summer monsoon rainfall. The altidutinal records indicate highest diversity of the Himalayan scaphidiids between 1000 and 2500m, but several species were found above 3000m (e.g., Scaphidium biundulatum, Baeocera microptera, B. crinita, B. thoracica, B. errabunda, Scaphisoma inguietum, S. aurorae, S. jado, S. clavigerum, S. sikkimense, S. bhareko), and one (Baeocera microptera) is found at 4000 m. Since Myxomycetes have been recorded at 4800 m in Khumbu Himal (POELT 1967) the Baeocera may occur above 4000 m. In my oppinion, the relative paucity of data from lower elevations (Terrai, Siwaliks, Dum- valleys), is due to greater interest in the study of higher altitude habitats with, presumably, endemic species, an unfavourable climate repelling researchers, the short period in year favourable for sampling, and difficulties in locating suitable sampling sites. Scaphidiids often aggregate and several species may cohabit on a single fungus growth (LOBL 1986b). Unlike many other fungivorous beetles (e.g., Ciidae, Pterogeniidae, Tenebrionidae) they have not developed any secondary sexual characters which are correlated with food or mating competition. SYSTEMATICS TAMANINI (1969) subdivided the scaphidiids in two groups of family level, the Scaphidiidae and the Scaphisomidae. This has not been followed by any subsequent worker. To the contrary, KASULE (1966) reduced the scaphidiids to a subfamily of the Staphylinidae, and several modern authors (e.g., KASULE 1968, LAWRENCE 1982, HAMMOND & LAWRENCE 1989) treated them as a subfamily. LAWRENCE & NEWTON (1982) considered the scaphidiids as a possible member of the oxyteline group. NAOMI (1985) retained the family rank for scaphidiids, and considered them be the sister group of the scydmaenids with which they constitute the sister group of his Oxyporidae. For practical reasons, I prefer to treat the group as a family, in view of the still insufficient knowledge of the phylogeny of the Staphylinoidea. The classification and suprageneric names proposed by ACHARD (1924a) are not used here, as in any of my previous papers. Geographic variation has been little examined and clines are unknown within the scaphidiids. Only three species have been found to be polytypic. Two of them, Baeocera mussardi and Scaphisoma assimile, are represented each by a subspecies in the studied area. The polytypy in these species is based on minor discontinuous morphological characters. Their aedeagi do not exhibit any distinguishing characters compared with the nominate subspecies (e.g., Fig. 149). For more detail see LOBL (1963; 1979). The taxonomic treatement in these cases reflects rather typological than biological thinking. 16 SCAPHIDIIDAE OF NEPAL 481 KEY TO THE GENERA OF SCAPHIDIIDAE OF CONTINENTAL ASIA (genera not recorded from the Himalaya in parenthesis) Segments of antennal club symmetrical. Scutellum large, visible. Robust species. Rrocoxalicavitiesiclosed a picalliyà tt a SRE Ter 2 Segments VII to XI always, XI often, asymmetrical. Scutellum not visible, or only small apical portion point exposed. Usually small, graceful species. Procoxal cavities OPEMAPICAllyX 4 REEL NER ER ee 7 Eyemoßnotehede. NENNE RD O 3 Eye motched rn SR AA LINDNER I RN 4 Occipital region visible. Basal segment of metatarsus short. Elytron with longitu- dinal striae. Prosternum long. Tibiae without rows of spines .......... Scaphium Kirby Head retracted into thorax to eye margin. Basal segment of metatarsus long. Elytron without longitudinal striae. Prosternum narrow. Pro- and mesotibia with rows of SPIES RS IE REN SE Cyparium Erichson Base of elytron impressed to receive extended basal angle of pronotum. Centre of metasternum pubescent in male. Head retracted into pronotum ........................... 6 Base of elytron not impressed, basal pronotal angle not prolonged apically. Meta- sternum glabrous in both sexes. Head with occipital region visible . Sa RER) Elytral disc with rows of longitudinal, deep, punctate striae . Ace ibe wis Elytraldisesvaithoußstriaer. ae Blasi ani Lewis Mesosternalikeelibifidibasally® Re eg an en Scaphidium Olivier Mesosternalikeelisimple RE Eee RR II Hemiscaphium Achard Dhirdiantennomere Shorts triangular) Ars ne ne RE A TN 8 ihixdfantennomere clongate;subcylindrical) Re O 11 Body appearing glabrous, setae usually restricted to punctures ......................... 9 BOGVECONSPICUOUSIY PUDESCENE Errno PA NE MR RE EAN CORANO ARRE tone 10 Apical segment of maxillary palpus slender, tapering, Without any stria . Se ER RER ER dirti iL ee RAEE nine Leach Apical segment of maxillary palpus flattened, triangular, with striate outer mar- CREER E PLS SR N Be an Caryoscapha Ganglbauer MÉSepimerabridoe distinct 2.2. Cr ne (Sapitia Achard) Mesepimeralridee absente na. een Mystrix Champion Basal pronotal angle almost always extended to or beyond level of anapleural suture. Mesepimeral ridge distinct. Antennal insertion situated near clypeal suture ...........12 Basal pronotal angle never extended apically and not reaching level of anapleural suture. Mesepimeral ridge usually obsolete. Antennal insertion situated more or less taskromkelypealisuturesma. ne RE NN N I SNO 14 Meso2 and metacoxae! distant her m ne Baeocera Erichson Meso-and metacoxae,stronelyzapproximateh nn. a ER I En 13 Elytron with parasutural stria, discal surface usually distinctly microsculptured SIERRA ANA E PET STO INA ERI I COSO ORI RCA SORTE. Scaphobaeocera Csiki Elytron without parasutural stria, and disc not microsculptured ... Baeotoxidium Löbl Body narrow. Meso- and metacoxae approximate ....................................... 15 Body wide Meso-andimetacoxae distante man. LT 18 Third antennal segment curved. Hypomeron explanate basally, with margin distinctly CONVEX Re rococo LL TTI I cree ME OR Da Lawns ars Scaphoxium Löbl Third antennal segment straight. Hypomeron not explanate basally, with margin AMOS STALLONE Simca ARR Jonnie II PALO Pe RITO III RE 16 Meso- and metatarsus notably longer than respective tibia Scaphicoma Motschulsky 482 IVAN LÖBL — Meso- and metatarsus about as long as respective tibia ................................. 17 17 Elytron with basal stria entire, joined to sutural and lateral striae. Labial palpus 2- i SOSMEM CO rey ttt TI O dud eens RI Xotidium gen. n. — Elytron with basal stria usually ne laterally, sutural stria shortened. Labial palpus 3-segmented .. RATE AUS .. Toxidium Le Conte 18 Legs and antennae conspicuously Jong. “Antennal segments m to XI very thin. Antennal insertion situated near upper margin of eye .................... Bironium Csiki — Legs an antennae moderately long. Antennal segments III to XI robust. Antennal insertion situatedinearClypeal suture Wi. re I Eee 19 19 Antennomere XI asymmetrical, moderately longer than preceding segment, segments VIlttoxSelonsate MARNE RES ER EEE Pseudobironium Pic — Antennomere XI symmetrical, conspicuously long, longer than two preceding seg- ments combined, segments VII to X short ....................... (Amaloceromorpha Pic) Scaphium Kirby Scaphium Kirby, 1837; type species: Scaphium castanipes Kirby, 1837, by monotypy. This genus includes four species: the Nearctic S. castanipes Kirby, the Asian S. quadraticolle Solsky, S. immaculatum (Olivier) from Central Europe and Mediterranean area, and S. ferrugineum Reitter of presumed South African origin (REITTER 1880). Scaphium vernicatum Pic from China is hereby transferred to Scaphidium, comb. nov. Scaphium quadraticolle Solsky from ex “Soviet Central Asia” has been recently recorded from northern Pakistan (LOBL 1986b). A new record is “Naltartal, Umg. Jagot, 1- 4.VII.1974 F. Gartner”,one specimen found by the Austrian Karakorum Expedition (Coll. H. Franz). It is possible that the range of this species extends to the Himalaya. Ascaphium Lewis Ascaphim Lewis, 1893; type species: Ascaphium sulcipenne Lewis, 1893, by present designation. Members of Ascaphium have conspicuously striate elytra as those of Scaphium, but they differ from the latter by the elongate basal segment of the metatarsus. The genus includes five species and one “variety” which is here elevated to species rank. An additional species occurs in Nepal. The range of Ascaphium extendes from Japan, China and North Vietnam to Central Himalaya. KEY TO THE SPECIES OF Ascaphium 1 Elytron with sutural, marginal, and six discal striae ending almost at same level near (EI NA ES RER RE tetto TT PTT soso 2 — Elytron with four inner discal striae ending far from apical margin, outer stria or StH ae Teduced....222.22 Henne denen een Goo Eee I I © 5 2 First and third discal striae joined near elytral apex, beyond second stria ........ Beh ee see ste ee ORO le elaine nate LED NES tonkinense Achard =) Elytron' with discal striae not joined apically... ne 3 3 Row of sub-basal pronotal punctures sparse, more or less interrupted, not impressed in middle. Body and femora black, tibiae, tarsi and antennae bright ochreous Ri nie AUR SANG ee ochripes sp.n. SCAPHIDIIDAE OF NEPAL 483 — Pronotum with impressed dense row of sub-basal punctures. Femora and tibiae uniformly dark, or apical portion of tibiae somewhat paler than femora ............... 4 4 Second and third elytral striae starting at base and joined with basal stria ............... Mei ui IR e I ssülcipennejlewis — Second and third elytral striae starting beyond base and not joined with basal stria MPP ANA RA IS DIA RE ANI IS RSI CRT SULCILSCARIC 5 Pronotal disc very finely punctate, punctures on pronotal centre much smaller than tROSSIONKITON SEHR A RAR minor Pic (status nov.) — Pronotal disc and frons about equally, coarsely punctate ................................. 6 6 Length 3.2 - 3.5 mm. Antennal club dark with distinctly paler apical segment. Mesosternal keel low, with straight horizontal margin in lateral view .. apicale Lewis — Length 3.8 - 4.5 mm. Antennal club unicolorously dark or with apical portion of last sement somewhat paler. Mesosternal keel high, with sinuate margin in lateral WIG LARE SITES ERE VA NERI SR AR ER QAR BERMAN: OR A HERR tibiale Lewis Ascaphium ochripes sp.n. Holotype, female: Nepal, Sankhua Sabha distr., Chichila, 2200m, 24.IV.1984 (Löbl & Smetana) (MHNG). Diagnosis. Large-size species with ochreous tibiae and tarsi (Fig. 9). Subbasal row of pronotum sparse. Elytral disc with Ist and 2nd striae not joined with basal stria, 3rd joined with basal stria, Ist and 2nd striae joined apically. Metasternal grooves long, approximate. Description. Length 6 mm. Body, head and femora black. Apex of abdomen, tibiae, tarsi, antennae and palpi bright ochreous. Frons at narrowest point 0.31 mm, with fine and sparse punctation becoming denser and coarser apically, vertex with punctures larger than intervals. Antennae long, with segments VII to X about 2.5x as long as wide, and XI almost 3x as long as wide. Pronotum at base 2.6 mm wide, with lateral margin oblique in basal half, arcuate in apical half (dorsal view); sub-basal row of punctures irregular, sparse, in central portion more or less interrupted, laterally impressed. Elytron with straight basal stria and six deep discal striae (in addition to sutural and marginal striae): Ist and 2nd starting just beyond basal stria and not joined with latter, joined together apically; 3rd joined with basal stria, 4th starting laterally of basal stria, 3rd and 4th weakly incurved and somewhat longer than Ist or 2nd; 5th and 6th starting almost at same level behind humeral callosity, not or barelly incurved apically and almost as long as Ath. All striae distinctly punctate, extended to narrow, irregularly puncate apical area of elytron. Intervals convex, very finely punctate. Apical area not flattened, with fine and shallow punctures. Mesosternal keel gradually lower toward apical margin, with ridge horizontal in apical half (lateral view). Metasternal median grooves long, approximate, moderately convergent, each delimited by elongate apical impression. Protibia straight, mesotibia weakly sinuate, metatibia weakly incurved. Remarks. This species may be readily distinguished by its colour pattern in combination with the sparse row of the sub-basal punctures of the pronotum. It shares long lateral elytral striae and the large size of body with A. sulcipenne, A. sinense, and A. tonkinense. The remaining species, A. apicale, A. tibiale, A. minor, and undescribed species from Burma and Taiwan represented in the MHNG, have reduced lateral striae and shortened four discal striae. Ascaphium minor was described as a variety of A. sinense (Pic, 1956) although it differs conspicuously from the latter in many characters, including in the much smaller size. 484 IVAN LÖBL Episcaphium Lewis Episcaphium Lewis, 1893; type species: Episcaphium semirufum Lewis, 1893; by monotypy. This is a small genus confined to Asia. The type species occurs in Japan, the second member, E. saucineum (Motschulsky) in Sri Lanka, and E. unicolor sp.n., described below, in Eastern Nepal. KEY TO THE SPECIES OF Episcaphium 1 Elytron pale, with dark transverse fasciae in centre and along apical margin. Median process of mesosternum not margined, mesosternal transverse keel-line absent ........ RARA SIOE ANNE ARE RE ae Ne TER saucineum (Motschulsky) =" Elytron uniformly reddish-brown. .........2.2...2002220000 MN INT Eee 2 Head, pronotum and elytra unicolored. Middle portion of Ist ventrite not micro- SOUT OLIN te an Te ENIT E unicolor sp.n. — Head black, thorax black or reddish, elytra and abdomen reddish or ochreous. Entire hustabdominal ventrite microsculptured 4.02 nenne semirufum Lewis N Episcaphium unicolor sp.n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley, Lamobagar Gola, 1000 - 1400 m, 27.V. - 3.V1.1980 (Holzschuh) (MHNG). Paratype, male: Nepal, Taplejung distr., SE Yamputhin to Yamputhin, 2000 - 1650 m, mainly Alnus forest, 26. and 30.1V.1988 (Martens & Schawaller) (SMNS). Diagnosis. Body and legs uniformly reddish-brown. Metasternum with two small apico-median tubercles. Middle portion of first ventrite with obsolete micro- sculpture. Description. Length 3.2 - 3.6 mm. Body and legs reddish-brown, antennomeres I to VI ochreous, following segments dark brown to blackish. Punctation on frons finer than that on vertex. Pronotum with lateral margins weakly sinuate in dorsal view, disc almost evenly arcuate in lateral view, not impressed near base; sub-basal row formed by coarse punctures. Elytron with four or five discal rows of moderately large punctures, intervals finer and irregularly punctate. Apical area flattened, bearing shallow large punctures situated very closely to each other. Median mesosternal process with raised, keel-like margins extended laterally and forming transverse keels joined with apico-lateral margins of mesosternum. Metasternum entirely very finely punctate, with two small elongate medio-apical tubercles; surface flat between and laterally tubercles, impressions absent. Abdominal segments sparsely and very finely punctate, with microsculpture consisting of punctures, except middle portion of Ist ventrite lacking microsculpture. Meso- and metatibia incurved. Segments 1 to 3 of male protarsi distinctly enlarged. Aedeagus 1.12 - 1.19 mm long. Median lobe with curved apical portion and robust ventral process. Internal sac bearing pair of bifid, slender sclerites situated at base of thin laminar structure. Latter narrowed and incurved apically. Membranes of internal sac extremely finely denticulate apically, with setose structures in centre, and with extremely fine scale-like proximal structures. Remarks. Episcaphium unicolor shares the margined mesosternal process and the transverse mesosternal keel with E. semirufum. It may be distinguished from the latter species by the colour pattern, the vertex with punctures coarser than those on the frons, the SCAPHIDIIDAE OF NEPAL 485 finer punctures of the sub-basal pronotal row and of the elytral rows, and by the obsolete microsculpture on the middle portion of the 1st abdominal ventrite. Scaphidium Olivier Scaphidium Olivier, 1790; type species: Scaphidium quadrimaculatum Olivier, 1790, designated by Leach 1815. Isoscaphidium Achard, 1922 (sg); not validated by designation of type species. Hyposcaphium Achard, 1922 (sg); not validated by designation of type species. Pachyscaphidium Achard, 1922 (sg); type species: Scaphidium arrowi Achard, 1920, by monotypy. Parascaphium Achard, 1923; type species: Scaphidium optabile Lewis, 1893, by monotypy. Scaphidium with over 200 species is distributed over all major regions but absent from oceanic islands and from areas with cool climate. ACHARD (1922a) defined four subgenera, all based on overall similarities. He failed to place most of the species validly described to that date in any of these subgenera. They were used in papers dealing with the Japanese species (SHIROZU & Morimoto 1963, LOBL 1967) but ignored by most of the other authors (i.e., CHAMPION 1927, MıwA & MITONO 1943, LOBL 1975b and 1976a). Actually, roughly 90% of the species of Scaphidium have not been assigned to any subgenus. In absence of defined species groups based on autapomorphies the subgeneric names are not used for the 20 Himalayan species treated below. Three of these species, S. biundulatum Champion, S. gurung sp.n. and S. nepalense sp.n., all with obviously restricted ranges at higher altitudes, are possibly relatives of S. quadrimaculatum Olivier. Scaphidium arrowi Achard from “British Bootang” belongs likely to the same group. This species and S. dureli Achard have not been found in any modern collection, and I have not located their type localities. The remaining species were collected at lower altitude and are or seem to be widely distributed. Scaphidium incrassatum Achard from Burma is represented in the MHNG by a specimen labelled “Assam”. The species is not included in the key but may be easily recognised by colour pattern of the elytra (Fig. 12). Scaphidium clatratum Achard and S. ocellatum Achard from “Assam” are not included in this study. KEY TO THE HIMALAYA SPECIES OF Scaphidium Me LCOnOtUMm and elytra unicolored tI 2. SR onotumtand/orielytra)bieolored. une. ce ee euere ae 8 PRE orsalisurtace onbodyiochreouse ren er ee MO ee 3 — Body blackish-brown to black, elytra and pronotum sometimes with bluish or viola- COONS AIS eS oe settee ROSCA Cn ee E trom ERIC ENTER Eee NE 4 Blhorassochreousiventrallyan ee. cinnamomeum Champion ora dackeventrallyi se e RS A0 melanogaster sp.n. 4 Meso- and metafemur black with reddish transverse fascia. Species 6 to 7 mm long Ne RE TA tna grande Gestro ze emoraunormlysdark4Smaller SDECIES PANIER SERRE EAN re I 5 Elytra and pronotum violaceous or bluish, remaining surface of body black ............. CURE CGR EC OCH OR CBC AO COC ORA ARIA ER RS à ear RE A cyanellum Oberthü 486 IVAN LÖBL N RES ee biseriatum Champion — Elytral disc without trace of rows of coarse punctures ................................... 7/ 7 Lenght 6 mm. Lateral pronotal margin sinuate in dorsal view. Male profemur not keelede sr ot e ei CLI NE dureli (Achard) — Length 4.5 mm. Lateral pronotal margin not sinuate in dorsal view. Male profemur keelediventrally 2.22 2.2. REST LIT thakali sp.n. 8 Elytron black or blackish-brown, with single reddish sub-basal fascia ................. 9 =) Colour pattern: ofelytron different Sr TR 10 9755 Male profemurswithiventralitoothen.n nee CE sylhetense Achard — Male profemur with several ventral tubercles ............................. holzschuhi sp.n. 10 Elytron black or blackish, with two ochreous or yellowish fasciae or spots ......... 11 =) Colour pattem of elytron different "7.2... CE TE ES 16 Me Bronotumiuniformlyiblack ...........22. a ioe cose ees ee eee RCE OREM 12 =’ Pronotum black with reddish pattern... det «see eee EEE EOE 15 12 Elytron with wide fasciae or spots covering more than half of total surface ......... 13 — Elytron with narrow fasciae or spots covering less than third of total surface ....... 14 13 Margins of anterior elytral fascia strongly dentate. Pronotal punctures fine, smaller thansintervalser nenne en ee ent cos nue biundulatum Champion — Margins of anterior elytral spot weakly dentate. Pronotal punctures fairly coarse, as lansevasrormlarsen than intervialS acest RR arrowi Achard 14> Apical half of male:protibia‘evenly thick o. nepalense sp.n. — Inner margin of male protibia angulate in apical half, tibia narrowed towards apex ..... CH ARPS SMA e ARR a ONC io Le EURE AR Aen cacao gurung sp.n. 15 Elytron with anterior fascia strongly narrowed in middle, posterior fascia not reachingeapiealimargin a... ee RT harmandi Achard — Elytron with anterior fascia almost evenly wide, posterior fascia extended to apical TMM AN SUM a RO ed cies LE RITI RT coomani (Pic) 16 Elytron yellowish, with base, sutural, lateral and apical margins black rubritarse Pic — Elytron ochreous, with black or brown spots and/or fasciae ............................ 17 WiElytralidiscrentirely very finely punctate RR e sce ese eee semilimbatum Pic =) | Elytral disc With TOWS Of coarse punctures ...... 22202. Me ENS EEE 18 18 Elytral disc with two long and two short rows of coarse punctures ......... baconi Pic — Elytral disc with one long and one short row of coarse punctures ...................... 19 19 Pronotal spots not reaching basal margin. Elytron with wide central and apical fasciae, and with small humeral spot .......................... septemnotatum Champion — Pronotal spots reaching basal margin. Elytron with humeral spot absent ................. Scaphidium cinnamomeum Champion Scaphidium cinnamomeum CHAMPION, 1927: 270. Diagnosis. Length 3.2 - 3.4 mm. Head and body ochreous. Femora and antennomeres I to VI as body, tibiae darker, antennomeres VII - XI dark brown to blackish. Frons narrow. Pronotum strongly vaulted above level of elytra and strongly inclined apically, sub-basal row of punctures broadly interrupted in middle, lateral margins distinctly sinuate, discal punctation very fine. Elytron vaulted, with impressed sutural area; discal punctation fine with exception of two longitudinal rows of coarse punctures and several additional coarse punctures situated laterally. Male profemur incurved, ventrally flattened, with anterior and posterior ventral edge distinct, basal half bearing ventral row of erected setae; subapical tubercle minute. Meso- and metafemur slender in both sexes. SCAPHIDIIDAE OF NEPAL 487 Male protibia slender, somewhat incurved and weakly thickened apically, meso- and metatibia distinctly incurved and thickened apically. Aedeagus (Figs 45,63) with median lobe weakly narrowed apically in dorsal view, ventral wall suddenly tapering apically. Parameres each sinuate in apical half, with rather strongly enlarged apex. Internal sac complex (Fig.83). Type material. Holotype, female, labelled “W. Almora Divn. Kumaon U.P. nov. 1918, H.G.C.” (BMNH). Materia examined, 5 specimens: India, Himachal Pradesh, 10 km NW Sarahan, 1700 m, 7.X.1988 (Vit) (MHNG). Distribution. North India, Uttar Pradesh: Kumaon and Himachal Pradesh. Remarks. This species was found under oak bark, on stump with fungi, and in decaying plant debris. Scaphidium melanogaster sp.n. Holotype, male: India, Himachal Pradesh, Kulu Valley, Vashisht Baths N Manali,1990m, 13.X.1988 (Vit) (MHNG). Paratypes, 4: India, Himachal Pradesh, 10km NW Sarahan, 7.X.1988 (Vit) (MHNG) 2 females; Uttar Prades, Mussoorie, Rabit Farm, 1300 m, 10.VII.1989 (Riedel) (SMNS) 1 female; Nepal, Sankhua Sabha distr., above Sheduwa, 2550 m, 30.11.1982 (A. & Z.Smetana) (MHNG) 1 female. Diagnosis. Small-sized species with ochreous dorsal surface and black ventral side of thorax. Elytral punctation very fine. Male profemur and protibia without conspicuous sexual characters. Description. Length 3.3 mm. Head, pronotum, hypomera, elytra, and abdomen uniformly ochreous. Ventral surface of thorax, hypomera excepted, femora, tibiae, and antennal club black. Tarsi dark reddish-brown, antennal segments I to VI ochreous. Frons rather narrow. Pronotum vaulted above level of elytra, rather strongly inclined anteriad, with lateral margins distinctly sinuate; sub-basal row of punctures interrupted in middle, punctures relatively small, joined by striae; discal punctation almost obsolete. Elytron vaulted, with sutural area flat and impressed; punctures along sutural stria relatively small, about as large as basal punctures; discal punctation very fine, somewhat more distinct than that of pronotum; in addition, disc with two longitudinal rows of coarse punctures; apical area of disc weakly flattened, with enlarged but extremely shallow punctures. Pygidium and propygidium evenly very finely punctate, both bearing microsculpture well visible at 50x magnification. Male with metasternal pubescence short and sparse. Profemur weakly thickened, without conspicuous sexual characters. Meso- and metafemur slender. Protibia somewhat incurved, hardly stouter apically. Meso- and metatibia distinctly incurved, slender. Aedeagus with median lobe similar to that of S. cinnamomeum, but with lateral margins of distal portion somewhat concave. Parameres (Fig. 66) each gradually wider toward middle third, from there narrowed, in apical portion again wider; inner margin of parameres concave between middle and apical portion. Internal sac complex (Fig. 82). Remarks. This species is similar to S. cinnamomeum with which it shares the interrupted sub-basal row of pronotal punctures and the reduced metasternal pubescence in male. It may be readily distinguished by the black ventral surface of the thorax and by the black femora and tibiae. Besides, it differs from S. cinnamomeum by the less vaulted pronotum, by the elytral and abdominal punctation (the pygidium and propygidium are notably more coarsely punctate in S. cinnamomeum), and by the sexual characters in male. The holotype was found in decaying plant debris, the two paratypes were taken together with S. cinnamomeum from under oak bark. 488 IVAN LÖBL Scaphidium grande Gestro "Scaphidium grande GESTRO, 1879: 50; ACHARD 1920a:56; 1920b:125; 1920c:211; CHAMPION 1927:268; LÒBL 1990b:511. Scaphidium grande var. subannulatum Pic, 1915c:3. - syn.nov. Scaphidium grande var. inimpressum Pic, 1920:189. - Sa nov. Scaphidiolum grande; ACHARD 1924b:91. Diagnostic characters. Length 6 - 7 mm. Head and body black. Profemur and all tibiae black. Meso- and metafemur black, each with wide reddish fascia. Tarsus and antennomeres I to VI dark brown to blackish, club black. Frons moderately wide. Pronotum vaulted, with centre above level of elytra, discal punctation very dense. Elytron rather flat, with raised sutural area, without row of coarse punctures, discal punctation irregular, more or less fine. Male with ventral side of profemur and inner side of protibia tuberculate. All femora slender. Protibia weakly incurved, almost evenly stout, with minute apical denticle. Meso- and metatibia somewhat incurved. Metasternal pubescence dense and long. Aedeagus (Fig. 39) with median lobe wide, parameres (Fig. 55) weakly enlarged and incurved apically, internal sac comlex (Fig.71). Type material. Holotype male, from “Sarawak” (MCSN), examined. Material examined, 6 specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley bellow Num, 1050 m, 21. and 22.1V.1984 (Löbl & Smetana) (MHNG). Distribution. North India, Nepal, Burma, Thailand, Laos, Malaysia, Vietnam, Indonesia, Taiwan. This is a rather variable species, especially in size and elytral punctation. The Himalayan specimens are smaller than those I have seen from southeast Asia. The infrasubspecific forms inimpressum Pic, 1920 and subannulatum Pic, 1915 are based on characters without taxonomic significance. The specimens from Eastern Nepal were found under wood debris in a subtropical forest. Scaphidium cyanellum Oberthiir Scaphidium cyanellum OBERTHÜR, 1884: 5; CHAMPION 1927: 271. Diagnostic characters. Length 3.4 - 3.8 mm. Head dorsally, pronotum and elytra very dark violet or bluish, with metallic shine. Remaining surface of body, femora, tibiae, and antennal club black. Antennomeres I to VI and tarsi reddish-brown to black. Frons rather narrow. Pronotum vaulted above level of elytra, moderately inclined anteriad, with distinctly sinuate lateral margins; discal punctation dense and rather coarse, sub-basal row of coarse punctures not interrupted. Elytron vaulted, with irregular discal punctation and without trace of discal rows of coarse punctures, large discal punctures usually smaller than large punctures of pronotal disc; sutural area raised. Pygidium and propygidium finely punctate, both with more or less distinct microsculpture. Male with long oblique metasternal pubescence; profemur thickened, ventral side flattened, anterior margin subangulate ventrally. Metafemur somewhat stouter than mesofemur. Male protibia straight beyond weakly incurved basal portion, gradually stouter toward shallow subapical notch, then narrowed and with oblique inner margin. Meso- and metatibia straight or hardly arcuate, gradually thickened apically. Aedeagus (Fig. 42) with apical portion of median lobe evenly wide in dorsal view, in ventral view tapering toward apex. Parameres each somewhat incurved apically. Internal sac complex (Fig. 74). Type material. Holotype, female, from “Ind.bor.’(MNHN), examined. SCAPHIDIIDAE OF NEPAL 489 Material examined, 9 specimens: Nepal, Parbat distr.,Kali Gandaki Khola, Kopchepani, 1600 m, 18.V1.1986 (Holzschuh) (MHNG) 1; Kopchepani-Ghasa, 1600-2000 m, 19.V1.1986 (Probst) (MHNG) 1; Patan distr., Godawari, 1500 m, 22-25.V1.1983 (Brancucci) (NMB) 1; Godawari, 1500-1600 m, 6.VII.1986 (Holzschuh) (MHNG) 1; Sankhua Sabha distr., Arun Valley, Chichila-Mure, 1900 m, 24.V.1980 (Holzschuh) (MHNG) 1; Chichila, 1950 m, 29.V.1983 (Brancucci) (NMB) 1; Arun Valley, Num, 1550 m, 5-6.V1.1983 (Brancucci) (NMB) 1; Thernathum distr., Phulvari-Waku, 1200-1600 m, 9.VI.1985 (Brancucci) (NMB,MHNG) 2. Distribution: India “North India”, Meghalaya, Nepal. Scaphidium biseriatum Champion Scaphidium biseriatum CHAMPION, 1927: 268. Diagnostic characters. Length 3.8 - 4.5 mm. Body, femora and tibiae black or blackish-brown, apex of abdomen paler. Antennomeres I to VI and tarsi reddish, antennal club brown. Frons narrow. Pronotum vaulted above level of elytra, strongly inclined apically, with lateral margins sinuate; discal punctation rather fine, sub-basal row of punctures not interrupted at middle. Elytron moderately convex, with 2 or 3 rows of coarse punctures in addition to irregular, fine discal punctation; sutural area impressed. Male with long oblique metasternal pubescence. Male profemur moderately thickened, . margined and with one row of tubercles ventrally. Metafemur barely thickened. Male protibia somewhat incurved, becoming stouter from base to middle, evenly thick in apical half, with small apical tooth on ventral side. Meso- and metatibia somewhat arcuate. Aedeagus (Fig. 51) with median lobe wide dorsally, ventral side tapering. Parameres (Fig. 58) each incurved, somewhat enlarged at apex. Internal sac complex (Fig.76). Type material. Holotype male from “Sunderdhung V., W.Almora divn., 8-1200 feet” (BMNH), examined. Material examined, 23 specimens: Nepal, Sankhua Sabha distr., Arun Valley, Lamobagar Gola, 1000-1400 m, 27.V.-2.V1.1980 (Holzschuh) (NMB) 1; bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 19; Taplejung distr., confluence of Kabeli Khola and Tada Khola, 1000 m, 23-25.IV.1988 (Martens & Schawaller) (SMNS) 1; India, Uttar Pradesh, Garhwal, Gangani, 1250 m,13-20.VI.1981 (Brancucci) (NMB) 1; West Bengal, Darjeeling distr., Tindharia near Kurseong, 7.V.1980 (Hisamatsu) (MHNG) 1. Distribution. North India, Nepal. Scaphidium dureli (Achard) comb. n. Scaphidiolum dureli ACHARD, 1922c:37. Diagnostic characters. Length 6 mm. Head, pronotum and most of ventral surface black. Elytra and hypomera blackish, with weak reddish shine. Femora and tibiae black, tarsi dark reddish brown.Antennomeres I to VI dark reddish brown, VII to X black, XI ochreous. Eyes large. Frons moderately wide. Pronotum moderately vaulted above level of elytra, strongly inclined apically, with distinctly sinuate lateral margins, sub-basal row of punctures regular, not interrupted in middle, consisting of coarse punctures, discal punctation fine and dense. Elytron without any discal row of coarse punctures; punctation obsolete on humeral area, on most surface similar to that of pronotum. Femora slender, in male without particular sexual chgaracters. Male protibia strongly arcuate, slender except for moderately thickened apex. Mesotibia in male weakly arcuate, metatibia almost straight. Aedeagus with wide median lobe gradually narrowed toward tip. Apical portion of parameres weakly widened. Internal sac complex. 490 IVAN LÖBL Type material. One male syntype from “British Bootan Padong L.Durel 1913” (MHNG), examined. An additional male bearing same locatity data but not Achard’s identification ‘label (MNHN) is possibly a syntype also. Remarks. I have not been able to locate the type data. The species may be readily distinguished from S. grande by the uniformly flack femora and by the shape of the male profemur and protibia. Scaphidium thakali sp.n. Holotype, male: Nepal, Mustang distr., Lete, 2550 m, 2.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 4: Nepal, Mustang distr., Lete, 24.IX.1971 (Franz) (Coll.H.Franz, MHNG) 2 females; Myagdi distr., Gasa-Tatopani, Kali Gandaki Valley, 2000-2500 m, 20.VI.1986 (Holzschuh) (NMB) 1 female; Chitre, Ghar Khola, 2400 m, 26-31.V.1984 (Holzschuh) (MHNG) 1 male. Diagnosis. Medium-sized species with unicolored black body. Elytral punctation irregular. Male profemur with anterior margin keeled, male protibia thickened in apical half and notched near apex. Description. Length 4.5 mm. Body black, head and legs very dark reddish- brown. Antennomeres I to VI ochreous, VII to X black, XI dark brown. Pronotum vaulted above level of elytra, strongly inclined apically, with lateral margins oblique in basal half (dorsal view), discal punctation dense and rather coarse, sub-basal row of coarse punctures deeply impressed, not interrupted at middle. Elytron with moderately and evenly rounded lateral margin, dorsally moderately convex; area between sutural margin and deeply impressed sutural stria distinctly raised; basal punctures not elongate, becoming gradually larger laterad; discal punctation irregular, without any longitudinal row of coarse punctures, most punctures much finer than those on pronotum; apical portion of elytron somewhat flattened and with punctation coarser than that on centre. Propygidium and pygidium very finely punctate, with microsculpture distinct on propygidium and on base of pygidium, obsolete on most of pygidium. Male with metasternal pubescence long, dense, erected. Profemur thickened, with anterior margin expanded, forming small rounded keel. Metafemur somewhat stouter than mesofemur. Basal half of protibia incurved, apical half thickened and with straight outer margin, subapical notch shallow. Ventral margin of protibia weakly convex above and straight below notch. Meso- and metatibiae almost straight, barely sinuate in dorsal view. Aedeagus (Fig.52) weakly expanded apically, with ventral wall abruptly narrowed. Parameres (Fig. 61) each sinuate, narrowed subapically. Internal sac complex (Fig. 78). Remarks. Scaphidium thakali exhibits secondary sexual characters very similar to those in S. harmandi, except for the slender protibia. However, these two species may be readily distinguished by their distinct colour patter. S. harmandi differs also in the higher vaulted pronotum and in the shape of the aedeagus. Scaphidium sylhetense Achard Scaphidium sylhetense ACHARD, 1920e: 263; Champion 1927:269. Diagnostic characters. Length 4.0 - 4.6 mm. Head dark reddish-brown to blackish. Thorax and most of elytra black, lateral area of pronotum sometimes reddish. Elytron with reddish sub-basal fascia (Fig.15). Abdomen reddish or ochreous with dark pygidium and dark genital segments. Femora and tibiae black, tarsi dark brown. Antennomeres I to VI dark brown or reddish, club black. Frons narrow. Pronotum vaulted SCAPHIDIIDAE OF NEPAL 491 above level of elytra, strongly inclined apically, with notably sinuate lateral margins; sub- basal row not interrupted, discal punctation rather coarse. Elytron moderately convex, with discal punctation similar to that of pronotum, except for very finely punctate base and sub- basal fascia; disc without any row of coarse punctures. Metasternal pubescence in male long, dense, decumbent. Male profemur thickened, flattened ventrally, margined along anterior edge, beyond narrowed and with concave margin beyond small subapical tooth. Metafemur hardly thickened. Male protibia weakly incurved, very weakly thickened apically, with row of more or less distinct tubercles on inner side. Meso- and metatibiae somewhat incurved. Aedeagus (Fig. 53) with apical portion of median lobe slender. Parameres (Fig. 59) evenly arcuate, not enlarged apically. Internal sac with small sclerites (Fig.79). Type material. The male lectotype from Sylhet (BMNH) was designated by Champion (1927), not examined. Material examined (males only), 13 specimens: India, West Bengal, Darjeeling distr., Trindharia near Kurseong, 7.V.1980 (Hisamatsu) (MHNG) 1; Sukuna near Matigara, 7.V.1980 (Hisamatsu) (MHNG) 1; Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 1; Hedangna - Num, 800 m, 16.VI.1983 (Brancucci) (NMB, MHNG) 3; Arun Valley, Lamobagar Gola, 1000-1400 m, 25.V.-3.VI.1980 (Holzschuh) (MHNG) 5; same but 1400 m, 8-14.V1.1983 (Brancucci) (NMB) 1; Myandi distr., Tatopani, 1100-1400 m, 27- 28.V1.1986 (Holzschuh) (MHNG) 1. Distribution. Bangla Desh, North India, Nepal. Remarks. CHAMPION (1927) noted that the “anterior femora are toothed beyond the middle beneath”. This, in combination with characters given by Achard, does define the species. For further details see remarks under S. holzschuhi. The specimens from eastern Nepal were found on and under logs and wood debris; some were swept from vegetation. Scaphidium holzschuhi sp. n. Holotype, male: Nepal, Sankhua Sabha distr. Arun Valley, Lamobagar, 1400 m, 8-14.V1.1983 (Holzschuh) (MHNG). Paratypes, 4 males: Sankhua Sabha distr., Arun Valley, Hedangna - Num, 800 m, 16.VI.1983 (Brancucci) (NMB,MHNG) 2; Hedangna - Navagaon, 5.IV.1980 (Holzschuh) (MHNG) 1; bottom Arun Valley bellow Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 1. Diagnosis. Medium-sized species with distinct colour pattern. Elytral punctation fine or very fine. Male profemur with minute tubercles. Inner margin of male protibia strongly sinuate. Description. Length 4.0 - 4.5 mm. Head dark reddish-brown. Thorax black except for small reddish sub-basal area on each side of pronotum. Elytron black with ochreous sub-basal fascia. Abdomen ochreous with dark pygidium and dark genital segments. Legs dark brown. Antennomeres I to VI ochreous, club black. Frons narrow. Pronotum weakly vaulted above level of elytra, strongly inclined apically, with distinctly sinuate lateral margins; discal punctation dense, rather fine, basal punctation very fine, sub-basal row not interrupted at middle. Elytron moderately convex, very finely punctate on base and on ochreous fascia, beyond latter more or less finely punctate; punctures along sutural stria coarser than largest discal punctures; basal punctures not elongate; interval between sutural margin and sutural stria raised. Pygidium and propygidium very finely punctured and microsculptured. Male with metasternal pubescence long, dense, oblique. Profemur distinctly thickened, flattened ventrally, with anterior ventral margin irregular, bearing several minute tubercles. Metafemur only somewhat thicker than mesofemur. 402 IVAN LÖBL Protibia incurved in basal third, with outer margin somewhat, inner margin strongly sinuate (lateral view); stouter towards apical third, from there narrowed toward apex, at ‘apex only somewhat thicker than at base. Meso- and metatibia very weakly arcuate, gradually weakly thickened apically. Aedeagus (Fig. 54) with narrow apical portion of median lobe. Parameres and internal sac (Fig. 78) very similar to those in S. sylhetense. Remarks. This species may be distinguished from S. sylhetense by the shape of the male profemur and protibia. The pronotal punctation seems to be somewhat finer in S. holzschuhi than in S. sylhetense, but it does not provide reliable distinguishing character. Consequently, only males may be positively identified. Scaphidium biundulatum Champion Scaphidium biundulatum CHAMPION, 1927:269. Diagnostic characters. Length 5.5 - 6.0 mm. Head, body including apex of abdomen, antennae and legs black. Elytron with two large ochreous fasciae (Fig. 10). Frons wide. Pronotum not vaulted above level of elytra, moderately inclined apically, with fine discal punctation; sub-basal row not interrupted at middle. Elytron with fine discal punctation, without any row of coarse punctures, impunctate on pale fasciae. Male profemur thickened, its central portion explanate anteriorly; metafemur thickened. Male protibia incurved in basal half, straight in apical half, gradually stouter from basal fifth toward apical third, then narrowed toward apex; inner margin angulate, inner surface glabrous apically. Meso- and metatibia arcuate. Tarsi somewhat longer in male than in female. Aedeagus (Fig.48) with wide median lobe. Parameres not enlarged apically. Internal sac complex (Fig. 68). Type material. Holotype, male from Tonglu (Darjeeling district near Nepal frontier), BMNH, examined. Material examined, 10 specimens: Nepal, Panchtar distr., Dhorpar Kharka, 2700 m, Rdododendron-Lithocarpus forest, 13-16.1V.1986 (Martens & Schawaller) (SMNS,MHNG) 3; Taplejung distr., ascent to pasture Lassetham from Omje Khola, 2400-3150 m, 6.V.1988 (Martens & Schawaller) (SMNS,MHNG) 2; Sankhua Sabha distr., above Pahakhola, 2600-2800 m, Quercus semecarpifolia-Rhododendron, 31.V.-3.VI.1988 (Martens & Schawaller) (SMNS, MHNG) 3; India, Sikkim, Bakkim-Choka, 2670-3050 m, nr. Yuksam, 25.IX.1983 (Sakai) (M.Sakai, MHNG) 2. Distribution. Eastern Nepal, western Darjeeling distr. and western Sikkim. Remarks. CHAMPION (1927) stated that S. biundulatum resembles closely to S. fryi Achard. I have examined one syntype of the latter species. It is notably smaller and has the colour pattern of the elytra significantly different (Fig. 11) from that in S. biundulatum. Scaphidium arrowi Achard Scaphidium arrowi ACHARD, 1920e: 264. Diagnostic characters. Length7 mm. Head, body, legs and antennae black. Each elytron with two ochreous spots. Anterior spot large, extending from sutural stria to lateral stria and from basal row of punctures upto mid-length of elytron, with deeply emargined apical margin in male, somewhat irregular, oblique apical margin in female. Apical spot smaller, wider than long, suboval, separated from sutural stria and frora apical elytral margin by narrow dark area. Frons wide. Eyes relatively small. Pronotum not vaulted above leve! of elytra, strongly inclined apically, with strongly SCAPHIDIIDAE OF NEPAL È 493 sinuate lateral margins; sub-basal row of punctures not interrupted in middle; discal punctation dense and coarse, most punctures about as large as or larger than intervals. Elytron without any discal row of coarse punctures; spots impunctate, black areas inbetween densely and coarsely punctate, punctures larger than those of pronotum and about as large as intervals. Metasternal pubescence in male short, decumbent. Male femora not thickened, longer than those in female. Profemur flattened and margined ventrally, with ventral margin sinuate in frontal view. Male protibia thickened toward apical seventh, then narrowed, with inner margin oblique, not notched; basal half of protibia weakly curved, apical half straight. Mesotibia longer than in female, arcuate. Mesotarsus conspicuously long, with ventral pubescence long. Aedeagus with median lobe wide, parameres evenly slender, setose, bearing minute tubercles on apical portion of inner side, and with internal sac complex. Type material. Lectotype, female, labelled “Bhotan Padong” (NMP), two paralectotypes, females, labelled “British Bootan Padong L.Durel 1913” (NMP,MHNG), and one male paralectotype labelled “Assam” (BMNH), by present designation. Remarks. This species shares many characters with S. quadrimaculatum and its allies but exhibits setose and apically tuberculate parameres. The male paralectotype is in poor state. It has been pinned and its hind legs are missing. Scaphidium gurung sp. n. Holotype, male: Nepal, Parbat distr., Ghoropani pass, 2700 m, S slope, 9.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 8: as holotype but 6.X., 2 males; nr. Ghoropani pass, 2700-3100 m, 5-9.X.1983 (Löbl & Smetana) (MHNG) 1 female; Bhantanti-Ghoropani, 2500-2800 m, 10.V.1984 (Holzschuh) (MHNG) 1 male, 2 females; Chitre, 2400 m, 26-31.V.1984 (Holzschuh) (MHNG) 1 male; Mustang distr., Kopchepani, 1500-1700 m, 15.V.1984 (Bhakta) (NMB) 1 female. Diagnosis. Large species with black body and distinct colour pattern of elytra, latter finely punctate. Male profemur thickened, margined anteriorly. Male protibia with angulate inner margin. Description. Length 5.0 - 5.5 mm. Head, body, antennae and legs black. Elytron with narrow whitish or yellowish sub-basal fascia and with two subapical spots of same colour (Fig. 14), spots sometimes joined. Frons wide. Pronotum similar to that of S.biundulatum, discal punctation fine, basal punctation very fine, sub-basal row not interrupted at middle. Elytron with fine discal punctures similar to those of pronotum, without any row of coarse discal punctures, with humeral area very finely punctate, basal punctures not elongate; pale spots and fasciae impunctate. Pygidium and propygidium very finely punctate, distinctly microsculptured. Male with metasternal pubescence long and dense. Profemur thickened, ventrally flattened, margined anteriorly. Metafemur moderately thickened, widest beyond middle. Protibia incurved, evenly thin in basal fourth, then gradually stouter, from widest point toward apex obliquelly narrowed, with angulate inner margin. Meso- and metatibiae incurved. Aedeagus (Fig.49) with short, wide, apically narrowed distal portion. Parameres (Fig.56) each incurved, not enlarged apically. Internal sac complex (Fig.70). Remarks. This species may be readily distinguished from S. biundulatum by the smaller size and by the colour pattern. 494 IVAN LÖBL Scaphidium nepalense sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, 2550 m, 29.IV.1984 (Löbl & Smetana) (MHNG). Paratypes, 75: as holotype, 6 males, 13 females; same but 2500m, 30.IV., 5 males, 1 female; same but 2600 m, 13.X.1983, 1 male; Phulcoki, 20. and 22.1V.1982 (A. & Z. Smetana) (CNC, MHNG) 1 male, 4 females; Kathmandu distr., Siwapuri, 24.11.1982 (Rougemont) (MHNG) 2 males, 1 female; Siwapuri, 2500 and 2540 m, 8.X.1981 (Sakai) (Coll. Sakai) 2 females; Siwapuri, 2400- 2500 m, 29.IV.-1.V.1985 (Smetana) (MHNG, CNC) 25 specimens; Sindhupalcok distr., Gul Bhanjyang, 2600 m, 6.IV.1981 (Löbl & Smetana) (MHNG) 3 males, 3 females; Kutumsang, 2200- 2400 m, 6.IV.1981 (Löbl & Smetana) (MHNG) 1 male; Malemchi, 2800 m, 14-18.IV.1981 (Löbl & Smetana) (MHNG) 2 males, 3 females. Diagnosis. Large species with black body and distinct colour pattern of elytra. Elytra finely and irregularly punctate. Male profemur ventrally flattened, margined anteriorly. Apical half of male protibia evenly wide. Description. Length 4.5 - 5.0 mm. Very similar to S. gurung, body in average smaller, elytron (Fig.13) with yellowish colour pattern. Punctation on pronotal disc notably coarser than that in S. gurung. Elytral punctation usually finer than that of pronotum, punctures along sutural stria about as coarse as discal punctures; outer basal punctures somewhat elongate. Male with metasternal pubescence long and dense. Profemur moderately thickened, ventrally flattened and glabrous, margined anteriorly. Metafemur weakly thickened beyond middle. Protibia somewhat incurved near base, in apical half weakly and evenly thickened. Meso- and metatibia moderately incurved. Aedeagus (Fig.50) with ventral wall of median lobe gradually tapering toward apex, parameres (Fig.57) each arcuate, apically moderately enlarged, internal sac complex (Fig.69). Remarks. This species differs notably from S. gurung and S. biundulatum in the shape of the male protibia. The females of S. nepalense may be distinguished by the yellowish elytral pattern. These three species are possibly vicarious. Most specimens were found in decaying vegetational debris along large oak logs and under oak bark, on logs with fungi. Scaphidium harmandi Achard Scaphidium harmandi ACHARD, 1920f: 125. Diagnostic characters. Length 4.5 - 5.0 mm. Head reddish. Pronotal coloration variable, generally most of surface black and lateral margin with reddish spot or fascia, sometimes median area reddish. Elytron black, with two reddish transverse fasciae (Fig.16). Abdomen black, apical segments sometimes more or less reddish. Legs blackish or dark reddish-brown, antennomeres Ito VI ochreous, antennal club black. Pronotal disc not vaulted above level of elytra, with lateral margins not sinuate, oblique near base, discal punctation coarse, sub-basal row of punctures not interrupted. Elytron moderately convex, discal punctation fine, rather irregular, without row of coarser punctures; sutural area flat, not impressed. Male metasternum with long dense pubescence. Male profemur widened, with keeled middle of anterior margin. Meso- and metafemur slender. Male protibia with incurved basal third, then with straight outer margin, sinuate inner margin; tibia thickened beyond middle, with apical fourth slenderer than middle, but stouter than base, with small glabrous area near apical margin. Aedeagus (Fig.40) with median lobe narrowed beyond level of ventral process, moderately enlarged apically. Parameres each incurved, narrowed subapically. Internal sac complex (Fig.72). SCAPHIDIIDAE OF NEPAL 495 Type material. Lectotype female, labelled “Sikkim Harmand 1886” (MNHN), by present designation. Material examined, 13 specimens. Nepal, Myandi distr., Kopchepani, 1600 m, 18.VI.1986 (Holzschuh) (MHNG) 1; Kathmandu distr., Siwapuri, 1700-2100 m, 25.V1.1988 (Martens & Schawaller) (MHNG) 1; Sankhua Sabha distr., Chichila, 2200 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 3; forest NE Kuwapani, 2250 m, 24.1V.1984 (Smetana & Löbl) (MHNG) 1; same but 2600 m, 15.IV.1982 (A. & Z.Smetana) (MHNG) 1; Induwa Khola Valley, 2100 m, 17.1V.1984 (Löbl & Smetana) (MHNG) 1; Basantapur, 2100, 30.V.-2.VI.1985 (Brancucci) (NMB) 1; Taplejung distr., Yamputhin cultural land, 1650-1800 m, 26.IV.-1.V.1988 (Martens & Schawaller) (SMNS, MHNG) 3; between Hellok and lower Gunsa Khola, 2000-1620 m, 18.V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. Nepal, Sikkim (or Darjeeling district). Remarks. Scaphidium sinense Pic has similar colour pattern (Fig.17) as S. harmandi. Scaphidium semilimbatum Pic Scaphidium semilimbatum Pic, 1917: 3. Diagnostic characters. Length 4.5 mm. Head reddish-brown. Pronotum reddish-brown with black base and two small black median spots; hypomeron reddish except for narrow black area near procoxal cavity. Most of elytron ochreous; basal, lateral and apical margins black, humerus black, disc with black transverse fascia (Fig.18). Most of ventral surface reddish, mesepimeron, mesepisternum and centre of metasternum black, mediobasal portion of 1st abdominal ventrite and metasternum darkened. Pygidium and propygidium reddish, base of pygidium black. Legs dark reddish. Antennomeres I to VI reddish, club black. Frons rather wide. Pronotum vaulted above level of elytra, with moderately sinuate lateral margins. Elytron convex, with rather fine punctation similar to that of pronotum, without discal row of coarser punctures. Male profemur stout, with keeled anterior margin in basal half. Male protibia moderately arcuate in basal half, with apical half of outer margin straight, inner margin sinuate between basal third and apex, and with shallow subapical notch. Meso- and metatibia each long, former weakly incurved, latter straight. Aedeagus with apical portion of median lobe enlarged near tip. Parameres each sinuate, almost equally wide in apical half. Internal sac complex. Type material. Lectotype, male, labelled “Kurseong (R.P.Astruc)”(MNHN), by present designation. Material examined, 1 specimen: Nepal, Sankhua Sabha distr., bottom Arun Vallew below Num, 1050 m, 21.1V.1984 (Löbl & Smetana) (MHNG). Distribution. East Nepal and India, West Bengal: Darjeeling district. Remarks. Kurseong is a village in the Darjeeling district quoted erronously by Pic (1917) as a Chinese locality. The single specimen of S. semilimbatum kept in the MNHN bears Pic’s handwritten identification label. There is no doubt that this is the specimen he had described. Scaphidium coomani (Pic), comb. n. Scaphidiolum coomani Pıc, 1925:323. Diagnostic characters. Length 3.5 - 3.7 mm. Head with frons reddish or ochreous and vertex black. Body black, pronotum with two large lateral ochreous spots extended over upper portion of hypomera. Elytron with large sub-basal and narrower 496 IVAN LÖBL apical ochreous fasciae (Fig.20). Apex of abdomen blackish or dark brown. Femora black, tibiae dark brown to blackish, tarsi and antennomeres I to VI reddish-brown, VII to X and basal half of XI black, apical half of segment XI paler. Frons very narrow. Pronotum notably vaulted above level of elytra, with sinuate lateral margins and very fine discal punctation. Elytra convex, with fine discal punctation, without discal row of coarser punctures. Male profemur and protibia without conspicuous features. Male meso- and metatibia longer than in female, protarsus enlarged. Aedeagus (Fig.43) with median lobe gradually narrowed apically. Parameres (Fig.64) each distinctly incurved, enlarged apically. Internal sac complex (Fig.75). Type material. Lectotype, female, labelled “LACTHO Tonkin de Cooman” (MNHN), by present designation. Material examined, 6 specimens: Nepal, Sankhua Sabha distr., Arun Valley, Lamobagar Gola, 1000-1400 m, 27.V.-3.VI.1980 (Holzschuh) (MHNG, NMB) 4; India, Meghalaya, Garo Hills, 2500 ft. VII.-VIII.1917 (Kemp) (MHNG,ZSI) 2. Distribution. Vietnam, North India, Nepal. Scaphidium baconi Pic Scaphidium baconi Pic, 1915c: 43. Scaphidium assamense Pic, 1915c: 43. Scaphidium baconi; ACHARD, 1922d: 263; LOBL, 1990b: 511. Diagnostic characters. Length 4.5 - 5.2 mm. Head ochreous. Pronotum yellowish or ochreous, with two black discal spots and two small, more or less darkened subapical spots. Elytron yellowish or ochreous, with five black spots: larger humeral, latero- central and apical, and two smaller near sutural stria (Fig.23); sutural area, epipleuron and pseudopleron black. Abdomen yellowish, pygidium sometimes with more or less dark spot. Upper portion of hypomeron yellowish, lower portion black. Ventral surface of thorax black except for yellowish lateral portion of metasternum. Femora yellowish with black base, profemur with black spot on ventral side. Tibiae and tarsi dark reddish-brown. Antennal segments I to VI reddish-brown, club black. Frons narrow. Pronotum somewhat vaulted above level of elytra, strongly inclined apically, with hardly sinuate lateral margins and very fine discal punctation. Elytral disc rather flat, with two long and two short rows of coarse punctures; discal punctation fine near base, gradually coarser from middle toward apex. Male profemur weakly thickened, its anterior ventral edge margined and bearing a row of minute tubercles, and with few minute subapical tubercles along posterior ventral edge. Male protibia slender, sinuate, weakly thickened apically. Meso- and metatibia arcuate, evenly slender. Aedeagus (Fig.47) with median lobe abruptly narrowed subapically. Parameres each incurved. Internal sac complex (Fig.81). Type material. Lectotype of S. baconi, female, labelled “Ind.bor. Bacon” (MNHN), by present designation; lectotype of S. assamense, female, labelled “Margherita Assam IV & V.1889” (MNHN), by present designation. Material examined, 3 specimens: Nepal, Sankhua Sabha distr., Arun Valley, Mure, 2000-1300 m, 9.VI.1983 (Brancucci) (NMB) 1; Taplejung distr., SE Yamputhin to Yamputhin, 2000-1650 m, and Yamputhin cultural land, 1650 - 1800 m, 26.IV. - 1.V.1988 (Martens & Schawaller) (SMNS,MHNG) 2. Distribution. Vietnam, India, Nepal. Scaphidium rubritarse Pic Scaphidium rubritarse Pıc, 1915b: 36. SCAPHIDIIDAE OF NEPAL 497 Diagnostic characters. Length 4.5 mm. Head, most of pronotum, hypomera, and abdomen ochreous. Basal portion of pronotum and scutellum black. Elytron yellowish, with black base between suture and humerus, black lateral and apical margins, and black epipleuron and pseudopleuron (Fig.19). Ventral surface of thorax, femora, tibiae and antennal club black. Tarsi dark brown, antennomeres I to VI ochreous. Frons rather wide. Pronotum vaulted above level of elytra, strongly inclined apically, with lateral margins oblique near base, arcuate in apical half (dorsal view); discal punctation very fine, sub-basal row not interrupted at middle. Elytron weakly vaulted, with flat sutural area and very fine punctation, latter obsolete on yellowish surface of disc. Male profemur thickened, ventrally flattened, with convexly arcuate ventral margin. Meso- and metafemur slender. Protibia in basal half moderately incurved, stouter toward apex, with shallow subapical notch; inner margin concave between base and apical third, weakly oblique in apical third. Meso- and metatibia moderately incurved. Aedeagus (Fig.41) with apical portion of median lobe almost evenly wide, apical margin of ventral wall broadly rounded. Parameres incurved (Fig.62), evenly wide. Internal sac complex (Fig.73). Type material. Not located. According to the description from Java. Possibly lost from the Pic’s collection which is housed in the MNHN. Material examined, 1 specimen: Nepal, Panchtar distr., betw. Gitang Khola Valley and Dhopar Kharka, cultural land, mixed forest, 1750-2100 m, 13.1V.1988 (Martens & Schawaller) (MHNG). Distribution. Java, Nepal. Remarks. The identity of the Nepalese specimen is based on its conspicuous colour pattern which corresponds with the description. The type material of S. rubritarse has not been traced in the MNHN. Scaphidium septemnotatum Champion Scaphidium septemnotatum CHAMPION, 1927: 270. Diagnostic characters. Length 3.0 - 3.3 mm. Head and body ochreous, pronotum with two black spots, elytron with black humeral, median and apical spots, pygidium with single large black spot (Fig.21). Legs and antennomeres I to VI ochreous, club black. Frons narrow. Pronotum vaulted abclve level of elytra, strongly inclined apically, with distinctly sinuate lateral margins and very fine discal punctation. Elytron convex, with one long and one short row of coarse discal punctures, punctation fine on discal centre, coarser on apical third. Male profemur weakly incurved, pubescent ventrally. All tibiae weakly incurved, evenly slender. Aedeagus (Fig.44) with median lobe tapering beyond middle. Parameres (Fig.65) apically incurved and enlarged. Internal sac complex (Fig.80). Type material. Holotype, male, from “Gopaldhara, Nepal-Sikkim Frontier H.Stevens 10.IX.18” (BMNH), examined. Material examined, 5 specimens: Nepal, Parsa distr., nr. Birganj Lothar, 5- 19.IX.1967 (Canadian Nepal Exp.) (CNC, MHNG). Distribution. India: Darjeeling distr., Nepal. Scaphidium species Similar to S. septemnotatum but with less vaulted pronotum, the pronotal spots lying basally and the elytron without humeral spot (Fig.22). It possibly represents an undescribed species. 498 IVAN LÖBL Material examined, 1 female:. Nepal, Parsa distr., nr. Birganj Lothar, 450 ft, 5.IX.1967 (Canadian Nepal Exp.) (CNC). Hemiscaphium Achard Hemiscaphium Achard, 1922; type species: Scaphidium striatipenne Gestro, 1879, by original designation. ACHARD (1922b) assigned to this genus 12 species, all confined to south east Asia. Their descriptions are incomplete and useless not only for a phylogenetic analysis but even for identification. Further species of this genus were described in Scaphidium. Only one species is known to occur in the Himalayan region. Hemiscaphium brunneopictum Achard Hemiscaphium brunneopictum ACHARD, 1922b: 35. Diagnostic characters. Length 3.3 - 3.7 mm. Head, most of body and legs ochreous. Pronotum with two dark brown or black elongate median spots and with minute spot on central part of each lateral margin. Elytron with base, apex, pseudopleuron and epipleuron dark brown to black, and with dark transverse fascia on centre of disc (Fig.24). Center of pygidium usually darkened. Antennomeres I to VI ochreous, VII to X black, XI more or less pale. Pronotum with lateral margins weakly convex or oblique; discal punctation very fine, sub-basal row of punctures lying in impressed stria, not interrupted at middle. Elytron with two long discal rows of coarse punctures, accompanied by one short inner and two short outer rows of coarse punctures; discal punctation very fine on basal half, coarse on apical third. Male femur slender, not modified. Male protibia weakly incurved, almost evenly slender, protarsi not lobed, with segments 1 to 3 somewhat wider than in female. Aedeagus (Fig.46) slender, median lobe weakly narrowed apically, with rounded apex. Parameres (Fig.67) straight, hardly enlarged at apex. Internal sac moderately complex (Fig.84). Type material. Lectotype, female, labelled “Karen Mts Birma” (NMP), by present designation. Material examined, 10 specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 3; Arun R., 800 m, Hedanga- Num, 16.VI.1983 (Brancucci) (NMB) 1; India, West Bengal, Darjeeling distr., Sukna, 200 m, 7.X.1978 (Besuchet & Löbl) (MHNG) 1; same but Sevoke (MHNG) 1; Meghalaya, Garo Hills, above Tura, 3500-3900 ft., 15.VII.-30.VIII.1917 (Kemp) (ZSI, MHNG) 3; “Ober-Assam” (Hartert) (ZSM) 1. Distribution. Burma, India, Nepal. Cyparium Erichson Cyparium Erichson, 1845; type species: Cyparium palliatum Erichson, 1845, by monotypy. Yparicum ACHARD, 1920b; type species: Yparicum yunnanum Achard, 1920, by monotypy. - syn. nov. Achard distinguished Yparicum from Cyparium by the presence of a row of denticles on the outer margin of the protibia in the former genus. Obviously, he overlooked that the outer margin of the protibiae bears a row of denticles in all species of Cyparium. SCAPHIDIIDAE OF NEPAL 499 This genus is poorly represented in the Himalayan collections. Five species occur in North India, but none was yet found in Nepal. KEY TO THE HIMALAYAN SPECIES OF Cyparium 1 Body black, elytron with yellowish humeral spot and yellowish apical fascia, not IMICROSCUIDIUTE A ne en A EE en Nr es montanum Achard — Colour pattern different, elytron with or without microsculpture ........................ 2 2 Elytron yellowish or ochreous, with dark brown or black pattern along discal rows of punctures, along sutural and along lateral margins ...................... bowringi Achard — Elytron more or less dark reddish, without distinct colour pattern ...................... 3 5222 Metastemnum entirely: very finely punetaten....................en.0... plagipenne Achard — Metasternum with lateral portion much coarser punctate than centre ................... 4 case Bodyanot.iridescent,Lensth2.8: mm... meer enee nee khasianum Löbl BodyzinideseentwEarsenspeciesinn nn ee species indet. Cyparium montanum Achard Cyparium montanum ACHARD, 1922c: 41; CHAMPION, 1927: 272; LOBL, 1984: 60. Diagnostic characters. Length 3.1 - 3.5 mm. Body not iridescent, black, elytron with small yellowish humeral spot and larger yellowish apical fascia. Antennomeres I to VI ochreous, antennal club dark brown or black. Elytron with four rows of discal punctures, surface laterad of outer row coarsely and irregularly punctate. Elytral microsculpture absent, microsculpture on visible tergites conspicuous, consisting of punctures. Metasternum much coarser punctate laterally than on centre. Aedeagus (Fig.85) with long, slender apical portion of median lobe, internal sac simple, parameres slender, sinuate. Type material. Lectotype from Simla, (India, Himachal Pradesh) (BMNH), designated in CHAMPION (1927), not examined. Material examined, Il specimens. India, Himachal Pradesh, Barog forest 4 km SW Solan, 1500 m, 8.X.1988 (Vit) 2; Solan, N end, 1300 m, 9.X.1988 (Vit) 2; 10 km NW Sarahar, 1700 m, 7.X.1988 (Vit) 2; Kulu Valley, Naggar, 1850 m, 16.X.1988 (Vit) 4; Kulu Valley, Vashisht, 1900 m, 13.X.1988 (Vit) 1 (all MHNG). Distribution. India (Himachal Pradesh, Uttar Pradesh, West Bengal), Bhutan. Remarks. This species may be readily distinguished by its conspicuous colour pattern. The specimens from Himachal Pradesh and from Uttar Pradesh were found on decaying plant debris and on decaying fungi. Cyparium bowringi Achard Cyparium bowringi ACHARD, 1922c: 42; LOBL, 1979: 84; 1984: 59; 1991: 126. Diagnostic characters. Length 3.5 - 3.9 mm. Body iridescent, dark brown to black. Elytron yellowish or ochreous, with dark brown to black pattern along sutural and lateral margins, and on discal rows of punctures. Antennomeres I to VI yellowish, club dark brown. Elytral disc with four rows of coarse punctures starting basally or sub-basally and with two short lateral rows starting behind a small, irrerularly punctate area. Elytral microsculpture distinct. Visible tergites with microsculpture 500 IVAN LÖBL consisting of striae. Metasternum entirely very finely punctate. Aedeagus (Fig.86) with rather short, narrow apical portion of median lobe. Internal sac simple, bearing long, basally incurved duct, without sclerites. Parameres moderately long, straight in lateral view. Type material. Lectotype, female, from Java, designated in Löbl (1979). Material examined, 11 specimens. India, Himachal Pradesh, 10 km NW Sarahan, 1700 m, 7.X.1988 (Vit) (MHNG). Distribution. India (Himachal Pradesh, Meghalaya, Tamil Nadu), Indonesia (Java). Remarks. The species may be readily distinguished by the elytral coloration (see LOBL 1991). Cyparium plagipenne Achard Cyparium plagipenne Achard, 1922c: 41; CHAMPION, 1927: 271; LOBL, 1984: 60. Diagnostic characters. Length 3.5 - 3.7 mm. Body not iridescent. Head and thorax dark brown to blackish. Elytra reddish, laterally and apically paler than on centre. Abdomen darker than elytra, paler than thorax. Antennae brown, club more or less darkened. Elytron microsculptured, with four discal rows of coarse punctures and with a lateral group of coarse punctures forming one or two short rows. Microsculpture on visible tergites consisting of transverse striae. Entire metasternum very finely punctate and distinctly microsculptured. Aedeagus (Fig.87) with apical portion od median lobe short, rather robust. Internal sac with setose and papilar structures and bearing sclerites. Parameres each sinuate in lateral view. Type material. Lectotype, male, labelled “India Orient” designated by Champion 1927, examined. Material examined. In addition to the lectotype, two specimens from “India Orient” (NMP) and from “W.Almora divn. Kumaon U.P. Oct. 1917 H.G.C.”, both recorded in Champion 1927 as C. plagipenne (BMNH). Distribution. North India. Remarks. The Kumaon specimen is a female differing from the lectotype and from the second specimen labelled “India Orient” by the brighter reddish elytra and darker antennal club. As C. plagipenne is well defined by the aedeagal characters only, additional material is needed to confirm the specific assignment of the Kumaon specimen. Cyparium species A female specimen from “Sikkim: Gopaldhara, Rungbong Vall. H.Stevens” (BMNH) identified by CHAMPION (1927) as C. plagipenne differs by the iridescent body and by the metasternum laterally more coarsely punctate than on the centre. It possibly represents an undescribed species. Pseudobironium Pic Pseudobironium Pic, 1920; type species: Pseudobironium subovatum Pic, 1920, by monotypy. Morphoscapha ACHARD, 1920; type species: Morphoscapha grossum Achard, 1920, by original designation. SCAPHIDIIDAE OF NEPAL 501 Most of the 22 members of the genus are distributed in subtropical and tropical Asia. Two species, P. lewisi Achard and P. ussuricum Löbl, occur in temperate eastern Asia. Pseudobironium globosum Löbl from New Caledonia is of uncertain relationship and the sole species occuring southeast of the Wallace Line. The possibly sister group of Pseudobironium, Pseudobironiella Löbl, is restricted to Madagascar. As in the three previous genera, many species of Pseudobironium are inadequately described and/or their male sexual characters are unknown. At the moment it is difficult to define species groups although several species share possible synapomorphies absent from other members of the genus: P. sparsepunctatum (Pic) and P. languei (Achard) have a short and stout maxillary palpus; P. almoranum Champion and P. ussuricum Löbl are characterized by short und stout antennal segments VII to XI; P. vitalisi (Achard), P. carinense (Achard), P. castaneum Pic and P. rufitarse sp.n. share wide parameres (the parameres are slender in P. bicolor sp.n., P. feai Pic, P. fasciatum Löbl, P. impressipenne Löbl, P. ineptum sp.n., P. lewisi Achard, P. plagifer Löbl, P. sinicum Pic, and P. subglabrum (Löbl); P. vitalisi and P. castaneum have the lateral wall of the median lobe tuberculate. Pseudobironium globosum Löbl, with reduced tibial spurs, is distinguished by the aedeagus with a particularly elongate flagellum. KEY TO THE NEPALESE SPECIES OF Pseudobironium 1 Pronotum and elytra with distinct colour pattern ............................. bicolor sp.n. — Body uniformly black or very dark reddish-brown ....................................... 2 2 First abdominal ventrite not microsculptured laterally ...................... ineptum Sp.n. — Entire abdomen, including first visible ventrite, microsculptured ....................... 3 3 Antennomeres VII to XI stout, XI usually not SoS 2x as longe as wide . és ae " almoranum Champion — Antennomeres VI to XI slender, XI 2. 5 to 4x : as long as wide EN Rent See 4 4 Body, femora and tibiae dark reddish-brown or reddish-black, not deep black. AntennomereMhmuchishortenthan "77 Are ee castaneum Pic — Body, apex of abdomen excepted, femora and tibiae deep black. Antennomere VI as IT ES Re IE OO ANI Ne ARTO rufitarse Sp.n. Pseudobironium bicolor sp.n. Holotype, male, Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG). Paratypes, 6: Nepal, as holotype, 3 females; India, Meghalaya, Khasi Hills, between Mawsynram and Balat, 16 km from Mawsynram, 1000 m, 27.X.1978 (Besuchet & Löbl) (MHNG) 1 female; Meghalaya, Garo Hills, above Tura, 3500-3900ft., 15.VIL.-30-VIIL.1917 (Kemp) (ZSD) 1 female; Thailand, Chiang Mai prov., Doi Pui, 1250 m, 14.III.1982 (Rougemont) (MHNG) 1 female. Diagnosis. Medium-sized species with distinct colour pattern on pronotum and on elytra (Fig.25). Median and apico-lateral portion of metasternum microsculptured. Abdomen with microsculpture consisting of striae. Parameres of aedeagus enlarged apically; internal sac bearing central and lateral sclerites. Description. Length 2.8 mm. Body and legs ochreous or reddish-brown. Pronotum with dark brown to black median spot enlarged anteriad (sometimes T-shaped), and with darkened basal margin. Elytron darkened along margins, and with dark basal and central spots. Apical segment of maxillary palpus 4x as long as wide, tapering. Antennae 502 IVAN LÖBL long, segment II about 1.7x longer than III and somewhat longer than VI, segments VII to _ XI slender, XI 3.5 to 4x as long as wide, apically granulate. Pronotal and elytral punctation fine, on lateral areas finer than that on centre. Elytral disc evenly inclined apically, with puctation notably dense apically, sutural area flat. Pygidium very finely punctate. Median and apico-lateral portion of metasternum microsculptured, median portion of metasternum rather finely punctate, lateral portion of metasternum extremely finely punctate. Ventrites with microsculpture consisting of transverse striae. First visible ventrite with very fine punctation, hardly visible at 100x magnification. Meso- and metatibia somewhat incurved. Segments 1 to 3 of male protarsus notably widened, narrower than apex of protibia. Aedeagus (Fig.88) 0.86 mm long. Internal sac with lateral sclerites accompanying central tubes and with membranes bearing short denticulate structures. Parameres each enlarged apically. Remarks. Pseudobironium bicolor may be readily distinguished by its colour pattern. This species has been recorded from India and Thailand (LOBL 1982; 1990b). Pseudobironium almoranum Champion Pseudobironium almoranum CHAMPION, 1927: 273; LOBL 1969: 324; 1982: 160; 1986b: 343. Diagnostic characters. Body black, apical segment of maxillary palpus slender, tapering. Antennae short, segments VII to XI short, XI usually less than 2x as long as wide. Pronotal and elytral punctation fine. Lateral portion of metasternum very finely punctate, without microsculpture. Ventrites with distinct microsculpture consisting of punctures. Aedeagus (Fig.89) 0.76 - 0.86 mm long. Internal sac with simple flagellum extended to level of weakly sclerotized valves, bearing setose apical structures. Parameres slender, almost straight. Type material. Lectotype, female, and 2 paralectotypes, females, labelled “Kumaon W.Almora India H.G.C.” (BMNH), by present designation. Material examined, 21 specimens: Nepal, Mustang distr., Kali Gandaki Khola, Kalopani, 2400 m, 17-19.V.1984 (Holzschuh) (MHNG) 1; Patan distr., Phulcoki, 1700 m, 10.V.1981 (Löbl) (MHNG) 3; Sindhupalcok distr., Pokhare NE Barahbise, 2700 m, 7.V.1981 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., Chichila - Mure, 1900 m, 24.V.1980 (Wittmer) (NMB) 1; Chitre, 2200-2400 m, 28-29.V.1985 (Holzschuh) (MHNG) 1; Gorza, 2100 m, 5-6.VI.1985 (Brancucci) (NMB) 1; above Pahakhola, 2600-2800 m, 31.V. - 3,VI.1988 (Martens & Schawaller) (SMNS, MHNG) 4; Arun Valley, between Mure and Hurure, 2050-2150 m, 9-17.VI.1988 (Martens & Schawaller) (SMNG, MHNG) 4; Chichila, 1900 m, 18-20.VI.1988 (Martens & Schawaller) (SMNS, MHNG) 2; Taplejung distr., SE Yamputhin, 2000-1650 m, 20. and 30.IV.1988 (Martens & Schawaller) (SMNS) 2; Yamputhin, 1650-1800 m, 26.IV. - 1.V.1988 (Martens & Schawaller) (MHNG) 1. Distribution. North India: Himachal Pradesh, Uttar Pradesh, Nepal. Pseudobironium rufitarse sp.n. Holotype, male: Nepal, Taplejung distr., Yamputhin, open forest, 1650-1800 m, 26.IV.- 1.V.1988 (Martens & Schawaller) (SMNS). Paratypes: as holotype, 2 females (SMNS, MHNG). Diagnosis. Body black. Elytron with subapical hump. Metasternum without microsculpture. Abdominal microsculture consisting of short transverse arcs. Aedeagus SCAPHIDIIDAE OF NEPAL 503 with parameres robust, sinuate; apex of median lobe incurved and pointed, internal sac bearing extremely fine setose structures and very slender median tube. Description. Length 3.4 mm. Body black, apical abdominal segments reddish. Antennomeres I to VI ochreous, club brown. Femora and tibiae black, tarsi ochreous. Apical segment of maxillary palpus 3.5 - 4x as long as wide, tapering. Antenna long, with slender segments III to XI; III somewhat shorter than II, segment VI about 1.6x longer than III, as long as XI, latter 2.5 - 3x as long as wide. Pronotal punctation very fine. Elytron with small subapical hump. Sutural area flat anteriorly, somewhat raised apically. Elytral punctation near base very fine, similar to that of pronotum, becoming gradually coarser apically. Pygidium very finely punctate. Metasternum without microsculpture. Middle portion of metasternum densely and coarsely punctate, except on impunctate middle line. Lateral portion of metasternum very finely punctate. All tibiae somewhat incurved. Segments I to III of male protarsus conspicuously enlarged, basal segment wider than apex of tibia. Segments I to III of male mesotarsus somewhat widened. Aedeagus (Figs 92-94) 1.32 - 1.33 mm long. Median lobe gradually narrowed apically, with incurved and pointed tip. Parameres robust, sinuate in dorsal view, almost straight and gradually narrowed in lateral view. Internal sac with long thin median tube and with membranes bearing apical setose structures. Remarks. This species may be readily distinguished by the pale tarsi cons- trasting with the dark tibiae and body. It is well characterized also by the shape of the median lobe and parameres. Pseudobironium ineptum sp. n. Holotype, male: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG). Diagnosis. Body reddish. Elytron without subapical hump. Metasternum and lateral portion of first ventrite lacking microsculpture. Abdominal microsculpture consisting of punctures. Aedeagus with almost straight, slender parameres and short, incurved tip of median lobe. Internal sac with long flagellum and a pair of incurved central valves. Description. Length 2.7 mm. Body, femora and tibiae reddish-brown, apex of abdomen and tarsi paler. Antennae ochreous. Apical segment of maxillary palpus about 2x as long as wide, tapering. Antennae long, segments III to XI slender, III almost as long as II, VI about 1.5x longer than III, shorter than XI, latter almost 3x as long as wide. Pronotal punctation fine. Elytral disc regularly inclined toward narrow, flattened apical area, with punctation coarser than that on pronotum; sutural area flat. Metasternum without microsculpture, middle portion densely and rather coarsely punctate, except for impunctate median line. Lateral portion of metasternum very finely punctate. First abdominal ventrite very finely punctate, laterad without, in middle with, microsculpture. Abdominal microsculpture consisting of punctures, almost obsolete on pygidium, distinct on propygidium. Meso- and metatibia weakly incurved. Segments I to III of male protarsus strongly widened, narrower than apex of tibia. Segments I to III of male mesotarsus somewhat enlarged. Aedeagus (Figs 90,91) 1.0 mm long. Median lobe irregularly narrowed towards apex, with short incurved tip. Parameres slender, each moderately incurved near base, beyond almost straight, evenly wide between expanded base and apex. Internal sac with very long, almost straigh flagellum, two central valves formed by filamentous laminae, and with short stick-like sclerites. 504 IVAN LÖBL Remarks. This species may be distinguished from other members of the genus _of similar size and colour by the shape of the parameres in combination with the metasternum lacking visible microsculpture. Pseudobironium castaneum Pic Pseudobironium castaneum Pıc, 1923: 17. Diagnostic characters. Large-sized species with dark reddish to blackish body. Apical segment of maxillary palpus slender and tapering. Antenna long, segments VII to XI slender, XI 3.5 - 4x as long as wide. Punctation on pronotal disc fine, that on elytral disc rather coarse. Lateral portions of metasternum and of Ist abdominal ventrite very finely punctate, former without microsculpture, latter with microsculpture consisting of punctures. Aedeagus (Figs 95,96) 1.34 - 1.40 mm long. Median lobe wide, with lateral tubercles, abruptly narrowed at apex, with bilobed ventral process. Membranes of internal sac lamellar, flagellum accompanied by stick-like sclerites. Parameres robust, each enlarged sub-basally and forming a lobe extended to median line; apical portion of parameres moderately enlarged in dorsal view. Type material. Lectotype, male, labelled “Lac Tho Tonkin” (MNHN), by present designation. Material examined, 4specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley bellow Num, 1050 m, 22.1V.1988 (Löbl & Smetana) (MHNG). Distribution. Vietnam, Nepal. Remarks. This species is very similar to P. carinense (Achard) which differs by the absence of tubercles on the lateral wall of the median lobe. The Nepalese specimens of P. castaneum are somewhat darker than the specimens from Vietnam I have seen. Baeocera Erichson Baeocera Erichson, 1845; type species: Baeocera falsata Achard, 1920, by subsequent designation (Opinion 1221) ICZN 1982. Sciatropes Blackburn, 1903; type species: Sciatropes latens Blackburn, 1903, by monotypy. Cyparella Achard, 1924; type species: Scaphisoma rufoguttatum Fairmaire, 1898, by monotypy. Eubaeocera Comell, 1967; type species: Baeocera abdominalis Casey, 1900, by original designation. Amaloceroschema Löbl, 1967 (sg); type species: Baeocera (Amaloceroschema) freudei Löbl, 1967, by original designation. This speciose genus is well represented in the Himalaya. Forty species have been distinguished and named but seven additional species known in female sex only were recognised in the collections studied. These are not included in the present study. For practical reasons the four species known only from Meghalaya (B. senilis, B. pseudincisa, B. hygrophila, and B. montrosetibialis) are included in the key provided below. Baeocera montana (Pic, 1955) was transferred to Baeocera from Scaphisoma in LößL (1987b: 845). Hence, Baeocera montana Löbl, 1971 from India is a secondary junior homonym. I replace the name Baeocera montana Löbl by Baeocera montanella nom. nov. Among the 30 species found in Nepal Himalaya 11 are new and appear to be endemic to certain areas of the eastern half of the Nepal territory. This is a considerably high percentage of endemism when compared to that in the Western Himalaya where only two species out of 15 are possibly endemic. In the Darjeeling district only three species out of 14 appear to be endemic. SCAPHIDIIDAE OF NEPAL 505 Although all Himalayan Baeocera were found in moist forest litter, many show little macrohabitat preferences. Baeocera ventralis is common in the evergreen dipterocarp forest and in the subtropical deciduous forest, but occurs also in the temperate, mixed broad-leaved and oak forest up to about 2000 m altitude. Other species exhibit similar considerable range in vertical distribution, often exceeding 1000 m (e.g. B. hamifera, B. khasiana, B. mussardi roberti, B. signata, B. sordidoides, B. vilis, B. wittmeri). Baeocera serendibensis which occurs at 400 m altitude in the Corbett National Park in Kumaon was found up to 2750 m in Nepal, and the range of B. microptera extends from 1500 m in Himachal Pradesh to the alpine scrub zone at 4000 m in Khumbu Himal. The Himalayan Baeocera belong to several species groups, defined mainly by the male genital characters, but only some of them are based on convincing synapomorphies (e.g. curtula, excelsa, insolita, lenta, ventralis, monstrosa, and satana groups): 1. The brevicornis group is characterized by the symmetrical, slender median lobe with apical portion tapering and moderately incurved apically; the ostium is situated apico-eudorsally and is covered by two overlapping membranous valves; the internal sac is armed with a slender, incurved elongate sclerite which is sometimes accompanied by a second slender sclerite. The parameres are symmetrical, slender and simple. Two subgroups can be recognised within this group: 1. The brevicornis subgroup, characterized by the body moderately convex both dorsally and ventrally. Baeocera brevicornis and B. serendibensis distributed throughout the Indian subcontinent, four additional East Asian species, B. problematica Löbl from Kenya and several unnamed Afrotropical species belong here. 2. The nobilis subgroup is characterized by the strongly convex body. Baeocera sordidoides from Nepal, B. sordida Löbl from Japan and the Mediterranean B. nobilis Reitter are assigned to this subgroup. 2. The monospecific /aminula group. It shares the essential characters with the nobilis subgroup, but the internal sac is covered by very fine setae and spines, and is armed by three distal sclerites. 3. The monospecific inermis group shares the aedeagal characters with the former two groups, but the internal sac lacks any sclerotized structures. The metacoxae are approximate. 4. The pilifera group is characterized by the median lobe similar to that in the former groups but with the distal portion short. The internal sac is armed by a single, slender and proximally incurved sclerite. The parameres are symmetrical, enlarged and pubescent. The short basal angles of the pronotum which do not cover the mesepimeral ridge is another possible synapomorphy of the group to which Baeocera pilifera from the Darjeeling district, and B. schwendingeri Löbl and B. pyricola Löbl from Thailand are assigned. 5. The monospecific but polytypic mussardi group is characterized by the symme- trical aedeagus with the median lobe consisting of moderately large basal and long distal portions. The ostium is situated dorsally near apex of the median lobe and the valves are contiguous medially. The internal sac is vesicular and weakly sclerotized, with minute basal sclerites. The parameres are strongly enlarged in sagital plane. The tarsi are consicuously short. 6. The monospecific khasiana group is characterized by the symmetrical aedeagus similar to that of the species of the brevicornis group, but with the distal portion of the median lobe relatively short. The internal is complex sac, bearing overlapping flat sclerites, and membranes covered by setose and spinose structures. 7. The lenta group is characterized by the symmetrical median lobe and parameres, similar to those of the species of the brevicornis group, although they are usually wider. The internal sac is armed with two or three unevenly long, curved, flattened and partly 506 IVAN LÖBL overlapping sclerites, which are joined proximally and form a more or less complex „structure; accessory basal sclerites are often present. The ejaculatory duct is thin and not sclerotized, enters in the proximal end of the sclerites and reappears at the distal end of the longer dorsal sclerite; the distal portion of the ejaculatory duct is very long, extrudes from the ostium and usually formes several rings on the external surface of the median lobe; the proximal portion of the ejaculatory duct enters often at an enlarged or vesicular portion before reaching the complex of sclerites. In the /enta group the metepisternum is often obsolete with suture indicated by a row of punctures. Most of the Asian and several Melanesian species are assigned to this group, in addition to B. pacifica Löbl from the Caroline Is., B. australica Löbl from Australia and B. umtalica Löbl from Zimbabwe. It includes following Himalayan species: B. cribrata, B. crinita, B. inculta, B. lenta, B. longicornis, B. manasensis, B. martensi, B. microps, B. microptera, B. pigra, B. pseudincisa, B. pseudovilis, B. puncticollis, B. reducta, B. signata, B. vesiculata, B. vilis, and B. wittmeri. Baeocera martensi, B. pigra, B. wittmeri, and B. vidua Löbl from Thailand have in common the internal sac bearing short sclerites accompanied by rows of sclerotized, apically pointed scales, situated at the left side of the ejaculatory duct. These three species may be separated in a distinct subgroup (subgroup pigra). Another subgroup seems to be formed by B. vilis, B. pseudovilis and B. inculta which have the dorsal sclerite of the internal sac enlarged and apically not delimited from membranous structures. 8. The ceylonensis group shares most characters with the /enta group, but the aedeagus differs in having the internal sac bearing a proximal tuft of sclerotized spines. In addition, the species of the group have notched parameres, characteristic elytral punctation, and a distinct metepisternum. The group includes five Oriental species : B. ceylonensis (Löbl), B. dilutior Lobl, B. franzi (Löbl), B. semiglobosa (Löbl) and B. ventralis (Löbl). The latter is widely distributed and occurs also in the Himalaya. 9. The monospecific senilis group is characterized by the symmetrical aedeagus with tapering median lobe split apically in ventral and dorsal lobe (bilobed type). The ostium is situated apically. The internal sac is armed by a simple stick-like sclerite. The parameres are straight and slender. 10. The monospecific excelsa group shares the bilobed median lobe with the senilis group, but the internal sac has a long, flat, spiral sclerite, and the parameres are enlarged. 11. The insolita group is defined by the trilobed distal portion of the median lobe. The basal bulbous portion of the median lobe is very large, with the feebly sclerotized apico-dorsal portion overlapping the base of the unpaired, large ventral lobe, and paired narrow dorsal lobes. The internal sac is complex, bearing numerous overlapping scales, spines and denticles. Large sclerites are absent. The parameres are slender, simple, and symmetrical. The group includes B. insolita Löbl from Thailand and B. schawalleri from Nepal. 12. The monospecific bengalensis group is defined by the very large and somewhat asymmetrical bulbous basal portion of the median lobe which ovelaps apically the ostium and the base of the symmetrical, short and flat distal portion. The armature of the internal sac consists of large and strongly sclerotized denticles, stick-like pieces and pointed scales. The parameres are symmetrical, slender and simple. The sister group of the bengalensis group is possibly the macrops group which differs notably in having the distal portion of the median lobe asymmetrical. 13. The curtula group includes species having a large symmetrical median lobe with a long distal portion, the dorsal wall of which is uneven, formed by two valves. The ostium is situated latero-apically. The internal sac is provided with a long, more or less SCAPHIDIIDAE OF NEPAL 507 strongly sclerotized flagellum and is armed by one or two flagellar guide-sclerites. Baeocera kapfereri Reitter from North Africa and six Asiatic species: B. curtula Achard, B. freyi Löbl, B. hammondi Löbl, B. callida Löbl, B. hamifera Löbl, and B. mustangensis sp.n. are assigned here. The latter three occur in the Himalayan region. Several New World species (deflexa and congener groups) are close to this group. 14. The monstrosa group is characterized by the asymmetrical aedeagus with the basal bulbous portion of median lobe very large and more or less overlapping the ostium, and by the small, strongly sclerotized and irregularly curved distal portion. The internal sac is extremely complex, with overlapping large sclerotized teeth and membranes covered by scales, papillae and spines. The flagellum is absent. The parameres are asymmetrical, one is lobed, the other one bears usually an apophyse. Baeocera monstrosa (Löbl), B. inaequicornis Champion and B. gilloghyi (Löbl) occuring in the Himalaya, B. paradoxa (Löbl) from Sri Lanka, several additinal Oriental species, and the Japanese B. nakanei Lobl belong to this group. 15. The satana group includes species with the distal portion of the median lobe split dorsally, forming two asymmetrical, strongly sclerotized pieces. The left piece is pointed and short, the right piece is long and usually lobed. The ostium (not seen) is apparently situated between these pieces. The internal sac lacks flagellum and is very complex, armed by sclerotized teeth-like structures and by membranous spines-like structures and papillae. The parameres are asymmetrical, more or less enlarged but simple. In the male the metatibiae are modified and the colour pattern differs from that in the female. In addition to B. satana Nakane from Japan, five Himalayan species belong here: B. dentipes, B. errabunda, B. monstrosetibialis, B. thoracica, and B. tuberculosa. KEY TO THE HIMALAYAN SPECIES OF Baeocera (including Meghalayan species) 1 Lateral portion of metasternum very finely punctate, with punctures barely distinct at [O0xemapnificatloni: a area ee ee een anale san ae cou re 2 — Lateral portion of metasternum coarsely punctate, with punctures distinct at 24x MAENIEICAHON. eee gage Sone Se sn seen cn ns eds ee VE 15 2 Sutural stria extended along elytral base to form basal stria joined with lateral stria ..3 aaepebasal stria. not joined: with lateral:Stria 7 cite... 9 3 Propygidium and pygidium very finely punctate. Body very dark reddish-brown to AUX cod sn ne eis Bene PERS Re ce 4 = Propygidium and pygidium with coarse punctures .....4................2....e.. ces 6 4 Distal portion of median lobe bearing ventral bidentate lobe; apex of paramere EHlars ed Mm RE aan ante mustangensis sp. n. — Distal portion of median lobe without any ventral protuberance; apical half of paramere equaliy.wider: Soc are kun Sots tots lene a x. 5 5 Flagellum of internal sac slender, joined with proximal end of sclerotized complex; flagellar guide-sclerite flat, curved at apex, not hook-shapes ............... callida Löbl — Flagellum of internal sac wide and moderately long, joined with central part of sclerotized complex; flagellar guide-sclerite robust, hook-shaped ....... hamifera Löbl PR Barameres:simplercre Mares ee os RG a bs oe heist dsl dere: 7 D ONE para mere Will APOPNY SIS) 21.55. 45cm areas nen 8 7 Parameres almost straight, truncate at apex ......................,....... gilloghyi (Löbl) Parameres irregularly curved, hook-shaped at apex .................... pubescence Löbl 508 24 IVAN LÖBL Right paramere with basal apophysis, left paramere enlarged and lobed apically ....... 30800 do SOEUR DLO TORS COS SRE EUR cn ERO dote pali i ER nn np inaequicornis Champion Right paramere simple, left paramere with large apophysis starting from centre ........ ne > predoni nto] e et ER net abo 02 MS ANR monstrosa (Löbl) Mesocoxaliline arcuate; finelyapunctate re rec ee ce khasiana Löbl Mesoxocal line parallel or subparallel to coxal foramen, coarsely punctate ......... 10 Length 1.4 mm. Body uniformly pale reddish-brown .................. bengalensis Löbl Parsenspecies Colour pattern differenti tI 11 Male metatibia with one or more denticles on inner side near base. Elytra more or less'dark, never black: nor ea ee TT 12 Male metatibia not dentate. Body entirely black in female, black with ochreous prothorax and metepisternum in male... RE 14 Inner side of male metatibia bearing numerous minute denticles .......................... RIONI TI RITI monstrosetibialis Löbl Inner side of male metatibia bearing one to three sub-basal teeth ..................... 13 Parameres slender and sinuate, apically not or weakly narrowed ....... thoracica sp. n. Parameres incurved, left paramere with middle portion wider than that of right Paramere,narrowediapicallys ar. see ee Ce ELLE dentipes Löbl Male with basal and apical pronotal margins black. Median lobe strongly enlarged WINE CRONISTA IA acco tuberculosa sp. n. Pronotum entirely ochreous in male. Median lobe not enlarged apically ................. Se Seed SR aon tC OAL DERE READ Sue SE ri nei eli in Tao errabunda sp. n. Hypomeron distinctly punctate 2. 16 Hypomeron:iMpunctate ...-...... criterio tt RT 19 Sutural stria short, evanescent before reaching level of elytral base .................. 17 Sutural stria long, curved along pronotal lobe and extended laterally along elytral DASÉ A I E po 00000000 18 Entire hypomeron coarsely punctate; mesepisternum distinctly punctate. Lateral portion of pronotum much more coarsely punctate than pronotal centre cribrata sp.n. Only small area of hypomeron punctate; mesepisternum impunctate. Pronotum EVENIYADUNCLALCHE nni puncticollis Löbl Metepistermumpundistinct rn RS RE ER microptera Löbl Metepisternum large, separated from metasternum by deep suture ....... excelsa Löbl Sutural stria reduced, not visible on anterior part of elytron ............................ 20 Sutural'stria longer, reaching’elytral’base............. ORTI eee 22 Antennae short, segment VIII not or somewhat longer than wide. Small species 1.0 - MMAIONSE RE RE Re crinita sp. n. Antennae longer, segment VIII elongate. Larger species ............................... 21 Sutural stria visible only on apical part of elytron .......................... reducta sp. n. Sutural stria longer, distinct between apex and basal 1/3 to 1/5 of sutural length ........ SIR IE AIA AE RR IL no hygrophila Löbl Sutural striainot extended alongelytral’baser............... 222 ose ee eee eee eee 23 Sutural stria curved at base and more or less extended along elytral base ............ 24 Length 0.9 - 1.0 mm. Antenna short, segment VIII about as long as wide ................ REMI RAMPE SIE VRR RSS LI ana II PR PC A M microps Löbl Length 1.75 - 2.25 mm. Antennae long, segment VIII slender, much longer than wide. Metatarsus conspicuously short, only somewhat longer than half of metatibia N N ET ET N e AIA mussardi (Löbl) Punctation of elytron and pronotum very fine, or elytron with few somewhat larger PUACIUTES HE RER SM EEE SSIS 25 32 33 38 SCAPHIDIIDAE OF NEPAL 509 Punctation of elytron notably coarser than that of pronotum, often irregular, or coarsely punctate area restricted to anterior half of elytron ............................. 28 Apex of median lobe exceeding somewhat level of mid-length of parameres ........... 25000000 DD RIA ER AR RE wid Mian QE, AR PASSAT blot oe inermis Löbl Apex of median lobe reaching almost level of apices of parameres ................... 26 Internal sac of aedeagus with simple, straight long median sclerite ........ senilis Löbl Internalksacioraedeasus’different. 2.2.2.2... me ee 27 Internal sac with two slender sclerites, one almost straight and flat, second curved, narrowedtandipointed/apieally’ u... res sordidoides sp. n. Internal sac with asymmmetrical apical sclerotized lamina accompanied by two slendemelonsate SClemlest en nee eee dre laminula sp. n. Elytron with basal stria extended to middle of basal margin or to humeral area, not No medkwathbla(Cralls(fa st Rene Res RAR ne A MR nr Tr Ne 29 Basal stria of elytron extended laterally and joined with lateral stria .................. 34 Most of basal half of elytron coarsely punctate, remaining elytral surface impunctate or very finely punctate, with punctures about as tiny as those on pronotum ......... 30 ÉMtralpunctationtdiffe rente MR IR ee 31 Parameres curved, with emarginate middie of inner margin, not narrowed apically .... 66000 0088 BER NG ANS II Dents, AI LA sie tnmneee ventralis (Löbl) Parameres straight, dentate, narrowed in apical half .................... manasensis Löbl Metepisternum wide, separated from metasternum by distinct, finely punctate suture 00000 EEE EEE ran a RITO AA schawalleri sp. n. Metepisternum not visible or very narrow, separated from metasternum by row of CO ATSC MUM CLUES mace IA RELITTO 32 Lateral margin of elytron straight. Elytral punctation relatively fine. Small species, ltl oul o8) mm Alone use rer E crane ete Cid EE signata Löbl Lateral margin of elytron rounded, or straight in middle portion only ................. 33 Parameres of aedeagus not or moderately narrowed apically. Membranes of internal sactcoyered.byspointedseales m am inne EE RAA EE lenta (Löbl) Parameres of aedeagus much narrower in apical half than in basal half, dentate in middle. Membranes of internal sac with minute, extremely short spines ................. PN SRF be ir. oc ane RI ATI ITER pseudincisa Löbl Lateral portion of metasternum with apical row of coarse punctures and a few coarse punctures near anterior and lateral margins. Parameres pubescent ........ pilifera Löbl Entire or most of lateral portion of metasternum covered by coarse punctures. Barameres5ssmootie.. re... er N IR an kenn hasse ee 35 Pronotal punctation rather coarse, distinct at 24x magnification ........ martensi Sp. n. Pronotal punctation very fine, not or barely visible at 24x magnification ............ 36 Metepisternum distinct, separated from metasternum by deep, punctate suture ..... 37 Metepisternum often indistinct, separated from metasternum by row of coarse PURCLUTE Sa ORI ya SP een IERI seele RR i Nee 38 Antenna short, segment VII almost 2x longer than segment VII. Elytron rather finely punctate, with very finely punctate base. Base of Ist ventrite rugose ..................... où EC N D CA Ste ES inset eae eae brevicornis (Löbl) Antenna long, segment VII moderately longer than segment VIII. Elytron, base excepted, coarsely punctate. Base of Ist ventrite with elongate punctures ............... i a a EE ve DE IST RI MS Ae A serendibensis (Löbl) Apical third to half of elytron very finely punctate, basal half of elytron coarsely punctate. First ventrite without any coarse punctures except those margining base 39 510 IVAN LÖBL — Elytron entirely coarsely punctate, or punctation becoming gradually finer toward apex and almost evanescent on latero-apical area. First ventrite usually coarsely punctate beyond basal row of punctures ..............2.....2.2.n ne 40 39 Parameres straight, evenly wide between apex and enlarged base .... pseudovilis Löbl — Parameres sinuate, enlarged in middle and apically ......................... inculta Löbl 40 Internal sac of aedeagus with large vesica, entered by ejaculatory duct before reaching complex Of Sclerites ........0........ dus ET 41 — Internal sac of aedeagus without basal vesica ........................................... 42 41 Parameres straight, parallel to each other .................................. vesiculata Löbl — Parameres somewhat curved, usually diverging apically ............. longicornis (Löbl) 42 Apex of median lobe of aedeagus situated far beyond level of mid-length of parameres. Elytron with rather large smooth latero-apical area ............... vilis Löbl — Apex of median lobe of aedeagus situated at about level of mid-length of parameres. Elytron with narrow smooth latero-apical area, or entirely coarsely punctate ....... 43 43 Parameres conspicuously narrow, parallel to each other, somewhat uneven ............ DS RC SENSEO OA a Sale I EN EN et ee wittmeri Löbl 4 Parameres rather Wide: Curved ........... 0000 nes TE pigra (Löbl) NEW RECORDS Baeocera serendibensis (Löbl) Material examined, 199 specimens: Nepal, Mustang distr., Kali Gandaki Valley, Mishi N Ghasa, 21.IX.1971 (Franz) (Coll. H. Franz) 2; Kali Gandaki Valley, between Lete and Ghasa, 25.IX.1971 (Franz) (Coll. H. Franz) 1; Kaski distr., Khorkore near Pokhara, ravine, 26.IX.1977 (Franz) (Coll. H. Franz) 1; Tandarakot, trail Pokhara - Ghoropani, ca 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz, MHNG) 6; Parbat distr., Ghoropani Pass, 2750 m, 5.X.1983 (Löbl & Smetana) 1; Gorkha distr., Buri Gandaki, Labubesi-Gorlabesi, 900-1000 m, 29.VII.1983 (Martens & Schawaller) (SMNS) 1; Darondi Kola between Sangu and Gorkha, 1200 m, 14.VIII.1983 (Martens & Schawaller) 1; Rasuwa distr., trail from Fulang Temple via Dinguari Khola, 150 m above bottom of valley, 11.X.1971 (Franz) (Coll. H.Franz) 1; Kathmandu distr., Nagarjun forest, 1650 m, 2.1V.1981 (Löbl & Smetana) (MHNG) 1; Nagarjun, Jamacok, 1400-1600 m, 18.VIIL.1983 (Martens & Schawaller) (SMNS, MHNG) 6; Gokarna forest, 1400 m, 3.VIII.1970 and 3.X.1971 (Franz) (Coll. H. Franz) 4; Gokarna forest, 1.IX.1981 and 20.X.1983 (Löbl & Smetana) (MHNG) 12; Patan distr., 2 km S Godawari, 1700 m, 20.X.1983 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., below Sheduwa, 2550 m, 30.11.1982 (A. & Z. Smetana) (MHNG) 1; bottom Arun Valley below Num, 1050 - 1100 m, 21.-22.1V.1984 (Löbl & Smetana) (MHNG) 30; Arun Valley near Num, 1500 m, 29.11.1982 and 1500-1600 m, 10.IV.1982 (A. & Z. Smetana) (MHNG) 7; Ilam distr., between Ilam and Mai Pokhari, 1400-1600 m, 8.IV.1988 (Martens & Schawaller) (SMNS) 1; Taplejung distr., confluence of Kabali Khola and Tada Khola, 1000 - 1050 m, 23.-25.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 52; ascent to Khebang from Tada Khola, 1500 m, 25.1V.1988 (Martens & Schawaller) (SMNS) 1; Yamputhin, 1650-1800 m, 26.1V.1988 (Marterns & Schawaller) (SMNS) 1; Yamputhin, 1650-1800 m, 26.IV.1988 (Marterns & Schawaller) (SMNS) 1; Yamputhin, 1650 -1800 m, 226.IV.1988 (Marterns & Schawaller) (SMNS) 3; India, Himachal Pradesh, Nahan, 930 m, 3.X.1988 (Vit) 1; 10 km NW Sarahan, N. Nahan, 1700 m, 7.X.1988, (Vit), 40; Barog forest 4 km SW Solan, 1500 m, 8.X.1988 (Vit) 19; Kulu Valley, Vashisht Baths N. Manali, 1900 m, 13.X.1988 (Vit) 1; Kulu Valley, Naggar, 1850 m, 16.X.88 (Vit) 2; 12 km E Mandi, 750 m, 25.X.1988 (Vit) 3 (all Indian specimens in MHNG). Distribution. Pakistan, India (Himachal Pradesh, Uttar Pradesh: Garhwal and Kumaon, West Bengal: Darjeeling distr., Assam, Meghalaya, Kerala), Sri Lanka, Western, Central and Eastern Nepal, Thailand. SCAPHIDIIDAE OF NEPAL 511 Baeocera brevicornis (L6bl) Material examined, 2 specimens: Nepal, Parsa distr., Terai, Amlekganj, 8.X.1972 (Franz) (Coll. H. Franz, MHNG). Distribution. Sri Lanka, India (Kerala), Nepal. Baeocera mussardi roberti Löbl Material examined, 96 specimens: India, Uttar Pradesh, Mussoorie, Rabit Farm, 1300m, 10.VII.1989 (Riedel) (SMNS) 1; Nepal, Kathmandu distr., Rani Ban SE Sanogau, 1500-1600 m, 25.1V.1988 (Brachat) (MHNG) 1; Patan distr., Phulcoki SE Godawari, ca 1800 m, 22.IV.1988 (Brachat) (MHNG) 2; Dhading distr., Ankhu Kola Valley, Ankhu Sangu, 650 m, 24.-25.VII.1983 (Martens & Schawaller) (SMNS) 1; Sankhua Sabha distr., Arun Valley near Num, 1500-1600 m, 29.III. and 10.1V.1982 (A. & Z. Smetana) (MHNG) 27; bottom Arun Valley below Num, 1050-1100 m, 21.-22.1V.1984 (Löbl & Smetana) (MHNG) 56; bottom Arun Valley between Hedangna and Num, 950-1000 m, 6.-9.VI.1988 (Martens & Schawaller) (SMNS) 1; below Sheduwa, 2550 m, 30.11.1982 (A. & Z. Smetana) (MHNG) 1; Induwa Khola, 1750 m, 14.IV.1984 (Löbl & Smetana) (MHNG) 1; Dhankuta distr., Arun Valley, Lamobagar Gola, 1000-1400 m, 27.V.-3.V1.1980 (Holzschuh) (NMB) 1; Taplejung distr., confluence of Kabeli Khola and Tada Khola, 1000-1050 m, 23.-25.1V.1988 (Martens & Schawaller) (SMNS) 1; Yamputin cultural land, 1600-1800 m, 26.IV.- 1.V.1988 (Martens & Schawaller) (SMNS) 1; Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) 2. Distribution. India (Tamil Nadu, Kerala, West Bengal: Darjeeling distr., Uttar Pradesh: Garhwal), Bhutan, Eastern and Central Nepal. Remarks. The size of the body of B. mussardi roberti is rather variable. It is in several specimens similar to that in mussardi s.str. which occurs in Sri Lanka and in Thailand. The conspicuously coarse punctation of the pronotum and the punctate mesepimera in roberti provide good diagnostic characters. Most specimens of roberti were found in subtropical and in mixed broad-leaved forests. Baeocera lenta (Löbl) Material examined, 4 specimens: Nepal, Sankhua Sabha distr., Arun Valley bottom between Hedangna and Num, 950-1000 m, 6.-8.VI.1988 (Martens & Schawaller) (SMNS, MHNG), 3; Arun Valley bottom below Num, 1050 m, 20.1V.1984 (Löbl & Smetana) (MHNG) 1. Distribution. Sri Lanka, India (Tamil Nadu, Kerala, Meghalaya), Eastern Nepal. Remarks. This is a common species in Sri Lanka and in Southern India. The populations from Kerala and Tamil Nadu were described as B. pseudolenta Löbl, based on the seemingly significant characters in the shape of the basal portion of the sclerites of the internal sac. However, reexamination of the aedeagi, using better optics, showed that the previously observed differences are due partly to the different degree of the sclerotization of the aedeagus, and partly to the infraspecific variability. Hence, Baeocera pseudolenta LoBL, 1979 is junior synonym of B. lenta (LÔBL, 1971) - syn. nov. Baeocera longicornis (Löbl) Material examined, 15 specimens: Nepal, Kaski distr., Pokhara Lake, hill facing temple near Pinta, 20.1X.1978 (Franz) (Coll. H. Franz) 1; Pande settle, via Pokhara, ca 2000 m, SIL IVAN LÖBL 29.IX.1971 (Franz) (Coll. H. Franz) 1; Gorkha distr., Darondi Khola betw. Doreni and Motar, 900-750 m, 13.VIII.1983 (Martens & Schawaller) 1; Kathmandu distr., Gokarna forest, 1.IV.1981 and ~ 10.IX.1983 (Löbl & Smetana) 4; Sankhua Sabha distr., botton Arun Valley, below Num, 1050 m, 21.IV.1984 (Löbl & Smetana) 2; Taplejung distr., confluence of Kabeli Khola and Tada Khola, 1000- 1050 m, 23-25.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 4; Yamputhin, 1650 - 1800 m, 26.IV.1988 (Martens & Schawaller) 1. Distribution. Sri Lanka, India (West Bengal: Darjeeling distr., Assam, Meghalaya, Uttar Pradesh: Kumaon and Garhwal), Western, Central and Eastern Nepal, Thailand. Remarks. Mostof the specimens previously recorded from Northern India as B. vesiculata belong to this species. Baeocera vesiculata Löbl Material examined, 7 specimens: Nepal, Myagdi distr., Myagdi Khola, Muri, 2100-2300 m, III.1970 (Martens) (SMNS) 1; Kathmandu distr., Nagarjun forest, 1650 m, 2.IV.1981 (Löbl & Smetana) 1; Patan distr., Godawari, 6000', 7.-13.VIII.1967 (Canadian Nepal Exp.) (CNC) 1; 2 km S Godawari, 1700 m, 20.X.1983 (Löbl & Smetana) (MHNG) 1, Sindhupalcok distr., above Chaubas, 2600 m, 5.IV.1981 (Löbl & Smetana) (MHNG) 1; Burlang Bhanjyang, 2600 m, 5.IV.1981 (Löbl & Smetana) (MHNG) 2. Distribution. India (Kerala, Meghalaya), Western and Central Nepal. Remarks. The tentatively associated females are not listed. The species may be distinguished from B. longicornis only by the shape of the parameres. Baeocera pigra (Löbl) Material examined, 1specimen: Nepal, Kathmandu distr., Gokarna forest, 1400 m, 3.X.1971 (Franz) (Coll. H. Franz). Distribution. Sri Lanka, India (Kerala, Tamil Nadu, Meghalaya, Assam, Uttar Pradesh: Kumaon), Central Nepal, Thailand. Baeocera wittmeri Löbl Material examined, 30 specimens: India, Uttar Pradesh, Mussoorie, Rabit Farm, 1300 m, 10.VII.1989 (Riedel) (SMNS) 1;Nepal, Kaski distr., Tandarakot, trail Pokhara - Ghoropani, ca 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz) 1; Mustang distr., Kali Ghandaki Valley, between Lete and Ghasa, 25.IX.1971 (Franz) (Coll. H. Franz) 1; Kathmandu distr., Gokarna forest, 1.IV.81 and 10.IX.1983 (Löbl & Smetana) (MHNG) 2; Nagarjun forest near Kathmandu, 1650 m, 2.1V.1981 (Löbl & Smetana) (MHNG) 1; Patan distr., Godawari, 1600 m, 31.11.1984 (Löbl) (MHNG) 1; Phulcoki, near Dalikhel, ca 1900 m, 21.IX.1977 (Franz) (MHNG) 1; Sindhupalcok distr., above Chaubas, 2500 m, 4.1V.1981 (Löbl & Smetana) (MHNG) 2; Burlang Bhanjyang, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 1, Solukhumbu distr., ravin near Shutje S Lughla, 7.X.1975 (Franz) (MHNG) 1; Sankhua Sabha distr., forest S Mangsingma, 2200 and 2300 m, 11. and 13.1V.1984 (Löbl & Smetana) (MHNG) 6; NE Kuwapani, 2250 m, 24.1V.1984 (Löbl & Smetana)l; Arun Valley , Chichila, 1900-2000 m, 18.-20.VI.1988 (Martens & Schawaller) (SMNS) 1; Chichila, 2300 m, 26.11.1982 (A. & Z. Smetana) 1; bottom Arun Valley below Num, 1050 and 1100 m, 20 .and 21.1V.1984 (Löbl & Smetana) (MHNG) 2; Arun Valley, between Mure and Hurure, 2050-2100 m, 17.VI.1988 (Martens & Schawaller) (SMNS, MHNG) 4; below Sheduwa, 2100-2550 m, 9.1V.1982 (A. & Z. Smetana) (MHNG) 4. SCAPHIDIIDAE OF NEPAL 513 Distribution. India (West Bengal: Darjeeling district, Uttar Prades: Kumaon, Garhwal), Western, Central and Eastern Nepal. Baeocera vilis Löbl Material examined, 68 specimens: Nepal, Myagdi distr., Kali Gandaki Valley, Dana, 20.IX.1971 (Franz) (Coll. H. Franz) 1; Mustang distr., Kali Gandaki Valley, between Ghasa and Lete IX.1971 (Franz) (MHNG) 1; Parbat distr., above Shika near Ghoropani, 26.IX.1971 (Franz) (MHNG) 1; Kathmandu distr., Gokarna forest 1400 m, 31.I11.-1.IV.1981 and 20.X.1983 (Löbl & Smetana) (MHNG) 21; Gokarna forest, 3.X.1971 (Franz) (Coll. H. Franz) 1; Bajalu Park, Kath- mandu, 1400 m, 17.1II.1980 (Martens & Ausobsky) (SMNS) 1; Rani Beni SE Sanogau, 1500-1600 m, 25.IV.1988 (Brachat) (MHNG) 4; Nagarjung, Jamacok, 1400-1600 m, 18.VIII.1983 (Martens & Schawaller) (SMNS) 12; Nagarjung, 1650 m, 2.1V.1981 (Löbl & Smetana) (MHNG) 5; Patan distr., Godawari, 1600 m, 31.11.1984 (Löbl) (MHNG) 1; above Godawari, 1700 m, 19.11.1980 (Martens & Ausobsky) (SMNS, MHNG) 2; Sindhupalcok distr., Burlang Bhanjyang, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi Khola bellow Malemchi, 2000 m, 15.IV.1981 (Löbl & Smetana) (MHNG) 1; Zorum River between Durmtali and Korthali, Ting San La, near Barahbise, ca 1900 m, VII.1980 (Franz) (Coll. H. Franz) 1; Sankhua Sabha distr., below Sheduwa, 2550 m, 30.11.1982 (A. & Z. Smetana) (MHNG) 1; Induwa Khola Valley, 2000 m, 16.1V.1984 (Löbl & Smetana) (MHNG) 1; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 2; 2 km E Mangsingma, 1900 m, 19.1V.1984 (Löbl & Smetana) 1; Ilam distr., Gitang Khola Valley, 1750 m, 11.-13.1V.1988 (Martens & Schawaller) (SMNS) 2; Panchtar distr., Paniporua, 2300 m, 16.-20.1V.1980 (Martens & Schawaller) (SMNS) 2; Taplejung distr., ascent to Khebang from Tada Khola, 1500 m, 25.IV.1988 (Martens & Schawaller) (SMNS, MHNG) 3; above Yamputhin, left bank of Kabeli Khola, 1800- 2000 m, 27.-29.1V.1988 (Martens & Schawaller (SMNS) 2. Distribution. India (West Bengal: Darjeeling distr.; Sikkim, Uttar Pradesh: Kumaon), Bhutan, Wertern, Central and Eastern Nepal. Remarks. Females of B. vilis may be readily associated with the males by their elytral punctation, notably by the latero-apical, very finely punctate area extended conspicuously anteriad. Baeocera pseudovilis Löbl Material examined, 10 specimens: Nepal, Kaski distr., Tandarakot between Pokhara and Ghoropani Pass, 1000 m, 18.IX.1971 (Franz) (Coll.H.Franz) 1; Parbat distr., above Shika near Ghoropani, 25.IX.1971 (Franz) (MHNG) 1; Ilam distr., Citang Khola Valley, 1750 m, 11.-13.IV.1988 (Marterns & Schawaller) (SMNS) 1; Taplejung distr., Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (MHNG) 1; India, Himachal Pradesh, Khadjiari E Dalhousie, 1950 m, 21.X.1988 (Vit) (MHNG) 4; 10 km NW Sarahan, NW Hahan, 1700 m, 7.X.1988 (Vit) (MHNG) 1; Barog forest SW Solan, 1500 m, 8.X.1988 (Vit) (MHNG) 1. Distribution. India (Himachal Pradesh, Uttar Pradesh: Kumaon, West Bengal: Darjeeling distr.), Western and Eastern Nepal. Baeocera signata Lobl Material examined, 22 specimens: Nepal, Manang distr., forest W Bagarchap, 2200 m, 241X.1983 (Löbl & Smetana) 1; Latha Manang W Bagarchap, 2350 m, 22.IX.1983 (Löbl & Smetana) 1; Parbat distr., Goropani Pass, 2700-2850 m, 5-6.X.1983 (Löbl & Smetana) 5; Kathmandu distr., Gokarna forest, 1400 m, 1.IV.1981 (Löbl & Smetana) 3; Sindhupalcok distr., Chaubas, 2600 m, 514 IVAN LÖBL 5.IV.1981 (Löbl & Smetana) 1; Malemchi, 2800 m, 14.1V.1981 (Löbl & Smetana) 1; Malemchi Khola below Malemchi, 2100 m, 15.1V.1981 (Löbl & Smetana) 1; above Shermathang, 2900 m, — 26.IV.1981 (Löbl & Smetana) 1; Pokhare NE Barahbise, 2700 m, 2.V.1981 (Löbl & Smetana) 3; Sankhua Sabha distr., forest S Mangsingma, 2200 m, 19.1V.1984 (Löbl & Smetana) 4; 2 km E Mangsingma, 1900 m, 19.1V.1984 (Löbl & Smetana) 1 (all MHNG). Distribution. India (West Bengal: Darjeeling distr.), Western, Central and Eastern Nepal. Baeocera microptera Löbl Material examined, 93 specimens: Nepal, Mustang distr., between Ghasa and Lete, 25.1X.1971, (Franz) (Coll.H.Franz, MHNG) 6; Mishi N Ghasa, Kali Gandaki Valley, 21.1V.1971 (Franz) (Coll. H. Franz) 1; Manang distr., Latha Manang W Bagarchap, 2400 m, 23.IX.1983 (Löbl & Smetana) (MHNG) 3; Gorka distr., Buri Gandaki, Nyak, 2270-2450 m, 1.VIII.1983 (Martens & Schawaller) (SMNS, MHNG) 5; Solukhumbu distr., Dugdima near Lughla via Blasda Pass, ca 4000 m, 4.X.1975 (Franz) (MHNG) 1; S Lughla, between Shutje and Bhum, ca 2200 m, 7.X.1975 (Franz) (Coll. H. Franz) 1; ravine near Shutje, S. Lughla, 7.X.1975 (Franz) (Coll. H. Franz, MHNG) 2; Sankhua Sabha distr., Arun Valley, betw. Mure and Hurure, 2050-2150 m, 17.VI.1988 (Martens & Schawaller) (SMNS) 1; pass NE Mangmaya, 2300 m, 6.1V.1984 (Löbl & Smetana) (MHNG) 1; Ilam distr., Mai Pokhari, 2100-2200 m, 25.-27.III.1980 (Martens & Ausobsky) (SMNS) 2; India, Uttar Pradesh, Joshimat, Pulna, 4.VIII.1989 (Riedel) (SMNS) 5; Himachal Pradesh, Baroq forest SW Solan, 1500 m, 8.X.1988 (Vit) (MHNG) 4; Kulu Valley, Chijoga S Manali, 1900 m, 12.X.1988 (Vit) (MHNG) 2; Kulu Valley, Vashisht Baths N Manali, 1900 m, 13.X.1988 (Vit) (MHNG) 5; Kulu Valley, Naggar 1850 m, 16.X.88 (Vit) (MHNG) 43; Dalhousie, 1950 m, 20.X.1988 (Vit) (MHNG) 7; Katalope Sanct. E Dalhousie, 2400 m, 22.X.1988 (Vit) (MHNG) 4; Jutogh, 10 km W Simla, 2000 m, 29.X.1988 (Vit) (MHNG) 2. Distribution. Pakistan, India (Himachal Pradesh, Uttar Pradesh: Kumaon and Garhwal), Western and Eastern Nepal. Remarks. This species has reduced wings and cannot fly, which is surprising in the light of its wide distribution. It occurs in conniferous and broad-leaved forests. The specimen found at Dugdima at 4000 m altitude represents the highest record for the genus. Baeocera ventralis (Löbl) Material examined, 152 specimens: Nepal, Kaski distr., Pokhara City, 820 m, 15- 18.V1.1976 (Wittmer & Baroni Urbani) (NMB) 1; Khorkore near Pokhara, 26.IX.1977 (Franz) (Coll. H. Franz) 1; Tandarakot, trail Pokhara - Ghoropani, ca 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz) 1; hill above Bennas near Pokhara, 20.IX.1978 (Franz) (Coll. H. Franz) 5; Gorkha distr., Arughat Suteo, 600-700 m, 27.VII.1983 (Martens & Schawaller) (SMNS) 2; Buri Gandaki, Suteo - Labubesi, 700 - 1000 m, 29.VII.1983 (Martens & Schawaller) (SMNS) 3; Darondi Khola between Doreni and Motar, 900-750 m, 13.VIII.1983 (Martens & Schawaller) (SMNS) 3; near Bimal Nagar, Terai, 500 m, 28.IX.1977 (Franz) (Coll. H. Franz) 4; Dhading distr., Ankhu Khola Valley, Ankhu Sangu, 650 m, 24-25.VII.1983 (Martens & Schawaller) (SMNS) 1; Buri Gandaki facing Pangshing, 1750 m, 31.VII.1983 (Martens & Schawaller) 1; Chitawan distr., Royal Chitawan National Park, 9.X.1980 (Franz) (Coll. H. Franz) 10; Parsa dist., Terai, Amlekganj 7-10.X.1972 (Franz) (Coll.H.Franz) 2; near Amlekganj, Khingar, Siwalik, 660 m, 18.X.1977 (Deharveng) (MHNG) 26; Kathmandu distr., Nagarjung, Jamacok, 1400-1600 m, 18.VIII.1983 (Martens & Schawaller) (SMNS) 1; Patan distr., Caukel Dara, near Bajrajogini, 1600-1700 m, 23.1V.1988 (Brachat) (MHNG) 2; Phulcoki near Dalikhel, ca 1900 m, 21.1V.1977 (Franz) (H. Franz) 4; Sindhupalcok distr., 3 km N Bahunepati, 900 m, 28.IV.1981 (Löbl & Smetana) (MHNG) 1; above Barahbise, 1550 m, 6.VIIL.1970 (Franz) (Coll. H. Franz) 1; Sankhua Sabha distr., Lamobagar Gac, 1400 m, 28-30.V.1980 (Wittmer) (NMB) 1; SCAPHIDIIDAE OF NEPAL 515 bottom Arun Valley below Num, 1100 m, 21.1V.1984 (Löbl & Smetana) (MHNG) 1; Ilam distr., Sanishare, 5 km N, feet of Siwalik Mts, 270-300 m, 3-5.1V.1988 (Martens & Schawaller) (SMNS) 7 ; Taplejung distr., Kabeli Khola below Limbudin, 900 m, 1.IX.1983 (Martens & Daams) (SMNS) 2; confluence of Kabeli Khola and Taka Khola, 1000-1050 m, 23-25.IV.1988 (Martens & Schawaller ) (SMNS, MHNG) 72. Distribution. Pakistan, India (Himachal Pradesh, Uttar Pradesh: Garhwal and Kumaon, West Bengal: Darjeeling district, Assam, Meghalaya), Western, Central and Eastern Nepal, Bhutan, Thailand. Baeocera khasiana Löbl Material examined, 8 specimens: Nepal, Ilam distr., Mai Pokhari, 2100-2200 m, 9- 10.1V.1988 (Martens & Schawaller) (SMNS) 1; Panchthar distr., Paniporua, 2300 m, 16.-20.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 2; Taplejung distr., Omje Kharka NW Yamputhin, 2300- 2500 m, 1-6.V.1988 (Martens & Schawaller) (SMNS) i; Sankhua Sabha distr., above Tashigaon, 3100 m, 7.IV.1982 (A. & Z. Smetana) (MHNG) 2; Sindhupalcok distr., Malemchi, 2900 m, 14.1V.1981 (Löbl & Smetana) (MHNG) 1, Patan distr., Phulcoki,, 2300 m, 10.V.1981 (LöBl) (MHNG) 1. Distribution. India (Meghalaya), Eastern and Central Nepal, Thailand. Remarks. Only females were found in Thailand. Their identity has to be confir- med. Baeocera excelsa Löbl Material examined, 1 specimen: Nepal, Sankhua Sabha distr.,Arun Valley, between Mure and Hurure, 2050-2150 m, 9-17. VI.1988 (Martens & Schawaller) (SMNS). Distribution. India (Uttar Pradesh: Kumaon), Eastern Nepal. Baeocera callida Löbl Material examined, 66 specimens: Nepal, Sankhua Sabha distr., Arun Valley bottom, below Num, 1050-1100 m, 20. and 21.IV.1984 (Löbl & Smetana) (MHNG) 24; below Sheduwa, 2550 m, 30.11.1982 (A. & Z., Smetana) (MHNG) 2; Taplejung distr., confluence of Kabeli Khola and Tada Khola, 1000-1050 m, 23-25.IV.1988 (Martens & Schawaller) (SMNS) 4; Kathmandu distr., Gokarna forest, 1400 m, 1.1V.1981 (Löbl & Smetana) (MHNG) 1; Kaski distr., Tandarakot between Pokhara and Ghoropani Pass, ca 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz) 1; India, Uttar Pradesh, Mussoorie, Rabit Farm, 1300 m, 10.VII.1989 (Riedel) (SMNS) 1; Himachal Pradesh, Jutogh 10 km W Simla, 2000 m, 29.X.1988 (Vit) (MHNG) 7; Kulu Valley, Chijoga S Manali, 1900 m, 12.X.1988 (Vit) (MHNG) 1; Kulu Valley, Naggar, 1850 m, 16.X.1988 (Vit) (MHNG) 7; Kulu Valley, Vashisht Baths N Manali, 1900 m, 13.X.1988 (Vit) (MHNG) 1, Barog forest 4 km SW Solan, 1500 m, 8.X.1988 (Vit) (MHNG) 17. Distribution. Eastern, Central and Western Nepal, India (Uttar Pradesh: Kumaon and Garhwal, Himachal Pradesh), Pakistan. Remarks. The records from Pakistan and Garhwal are based on females and are therefore not very reliable (LOBL, 1986b). However, the new findings support previous identification. Baeocera monstrosa (Löbl) Material examined, 3 specimens: India, Himachal Pradesh, 12 km E Mandi, 750 m, 25.X.1988 (Vit) (MHNG). 516 IVAN LÖBL Distribution. Sri Lanka, India (Kerala, Tamil Nadu, Uttar Pradesh: Kumaon, _ Himachal Pradesh). Baeocera hamifera Löbl Material examined, 34 specimens: Nepal, Mustang distr., 2 km N Kalopani, 2500 m,. 1.X.1983 (Löbl & Smetana) (MHNG) 3; Lete, 2550 m, 2.X.1983 (Löbl & Smetana) (MHNG) 2; N Lete, 24.IX.1971 (Franz) (Coll. H. Franz) 1; Tangsang near Tukche, Takola, ca 3000 m, 23. IX.1971 (Franz) (Coll. H. Franz) 1; Manang distr., Marsyandi, Thanjok-Chame 2250 m, 17.1V.1980 (Martens & Ausobsky) (SMNS) 1; Kathmandu distr., Nagarjung, Jamacok, 1400-1600 m, 18.VIII.1983 (Martens & Schawaller) (SMNS) 1; Patan distr., Phulcoki, 2300, 2500 and 2700 m, 10.V.1981, 16.X.1983, and 28-29.1V.1984 (Löbl & Smetana) (MHNG) 6; Sindhupalcok distr., Malemchi Khola below Malenchi, 2100 m, 15.IV.1981 (Löbl & Smetana) (MHNG) 1; Malemchi, 2800 m, Malemchi, 2800 m, 14.IV.1981 (Löbl & Smetana) (MHNG) 1; Pokhare NE Barahbise, 2700 and 2800 m, 2. and 7.V.1981 (Löbl & Smetana) (MHNG) 4; Sankhua Sabha distr., below Sheduwa, 2100-2500 m, 9.IV.1982 (A. & Z. Smetana) (MHNG) 3; Induwa Khola Valley, 2100m, 17.IV. 1984 (Löbl & Smetana) (MHNG) 1; Panchthar distr., between Paniporua and Hinwa Khola Valley, 2300 -1850 m, 20.1V.1988 (Martens & Schawaller) (SMNS) 2; Taplejung distr., Worebung Pass, 2000 m, 21.1V.1988 (Martens & Schawaller) (SMNS) 4; Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (SMNS) 1; upper Tamur Valley, resthut side of the valley, 2450 m, 19.V.1988 (Martens & Schawaller) (SMNS) 2. Distribution. India (West Bengal: Darjeeling distr.), Western, Central and Eastern Nepal. Baeocera pubiventris Löbl Material examined, 5 specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.IV.1984 (Löbl & Smetana) (MHNG) 2; India, Himachal Pradesh, 12 km E Mandi, 25.X.88 (Vit) (MHNG) 3. Distribution. Thailand, Eastern Nepal, India (Himachal Pradesh). NEW SPECIES Baeocera sordidoides sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, 2700 m, 16.X.83 (Löbl & Smetana) (MHNG). Paratypes, 19: as holotype but 15.X.1983, 1 male; Patan distr., Phulcoki, 2500 m, 28.- 29.IV.1984, 1 male, 2 females and at 2600 m, 14.X.1983 (Löbl & Smetana) 1 female (MHNG); 2 km S Godawari, 1700 m, 19.X.1983 (Löbl & Smetana) (MHNG) 1 female; Kathmandu distr. Shewapuri, 2100 - 2300 m, 25.VI.1988 (Martens & Schawaller) (SMNS) 1 male; same but 2400 m, 3.IV.1985 (Smetana) (MHNG) 1 male Sindhupalcok distr., Chaubas, 2600 m, 5.IV.1981 (Löbl & Smetana) (MHNG) 1 male, 2 females; Malemchi, 2800 m, 14.1V.1981 (Löbl & Smetana) 1 female; Manang distr., forest W. Bagarchap, 2200 m, 21:IX.1983 (Löbl & Smetana) (MHNG) 1 male, 1 female; Mustang distr., Taksang, Tukche, IX.-X.1971 (Franz) (Coll. H. Franz) 1 male; Parbat distr., Ghoropani Pass, N slope, 2750 m, 5.X.1983 (Löbl & Smetana) (MHNG) 1 male, 1 female; India, Uttar pradesh, Joshimat, Pulna, 2300 m, 4.VIII.1989 (Riedel) (SMNS) 1 male, 1 female; Mussoorie, Rabit Farm, 1300 m, 10.VII.1989 (Riedel) (SMNS) 1 female. Diagnosis. Medium-sized species with slender aedeagus and symmetrical median lobe, similar to that of species of brevicornis group; internal sac with two elongate SCAPHIDIIDAE OF NEPAL 517 curved sclerites; median lobe bilobed between ventral process and base of apical portion. Body, metasternum excepted, very finely punctate. Metacoxae approximate. Description. Length 1.40 - 1.55 mm, width 0.88 - 1.0 mm. Strongly convex, especially ventrally, body, femora and tibiae more or less dark brown, antennae and tarsi paler. Eyes large. Antennae moderately long, relative length of segments as follows: III 11, IV 12, V 16, VI 14, VII 18, VII 12, IX 19, X 17, XI 21 (holotype). Segments III to VI evenly wide, V about 4 x as longe as wide; VII 3x as long as wide; VIII about 2.5x as long as wide; IX to XI each much wider than VII, XI almost 2 x as long as wide. Pronotum with rounded lateral margins except near base; lateral keels not visible in dorsal view; punctation very fine, usually barely visible at magnification 50x. Tip of scutellum exposed. Elytra rather narrowed apically; lateral keel not visible in dorsal view and lateral margin feebly rounded; sutural area flat, very finely punctate; discal punctation sparse and very fine, punctures very shallow, but much larger than those on pronotum, often barely visible at magnification 24x; sutural stria deep, complete, extended along basal margin, forming basal stria joined with lateral stria. Wings fully developed. Pygidium extremely finely punctate and microsculptured. Hypomeron and mesepisternum impunctate. Mesepimeral ridge about twice as long as interval between its end and mesocoxa. Mesosternum with convex centre; middle impunctate area rather large, limited laterally by finely and very densely punctate areas. Lateral portion of metasternum smooth near metacoxa, elsewhere more or less coarsely punctate, punctures often elongate. Mesocoxal area 0.04 - 0.05 mm long, with coarse marginal punctures. Metepisternum 0.04 - 0.06 mm wide, not or barely narrowed anteriad, with deep, straight, punctate suture. Metacoxae relatively approximate. First ventrite with mediobasal hump; without microsculpture and very finely punctate; basal punctures coarse, not elongate. Protibiae straight, meso- and metatibiae somewhat curved. Segments 1 to 3 of protarsi moderately enlarged in male. Aedeagus (Figs 97,98) 0.48 - 0.52 mm long. Median lobe with long, tapering apical portion and bisinuate strongly sclerotized ventral wall beyond level of ventral process. Parameres uniformly wide, hardly curved. Internal sac with strongly curved basal sclerite. Remarks. This new species resembles B. sordida Löbl from Japan. Both species share most of the significant characters, including the relatively approximate metacoxae, which distinguish them from the membres of the brevicornis group. The aedeagi of B. sordida and B. sordidoides are similar, that of B. sordidoides differs in the median lobe bearing a bilobed process situated between the ventral process and the base of the narrowed apical portion. Baeocera inermis Löbl and B. laminula sp.n. described below are also difficult to separate from B. sordida/sordidoides by their external characters. Only the male genitalia provide positive diagnostic characters. Most specimens of B. sordidoides were found in oak and mixed broad-leaved forests in Central and Western Nepal. Baeocera laminula sp.n. Holotype, male: Nepal, Manang distr.,forest W Bagarchap, 2200 m, 21.IX.83 (Löbl & Smetana) (MHNG). Paratypes, 4: as holotype 1 male, 1 female; Parbat distr., Ghoropani Pass., N slope, 2750 m, 5.X.1983 (Löbl & Smetana) 1 male; Patan distr., Phulcoki, 2600 m, 20.IV.1982 (A. & Z. Smetana) 1 male (all MHNG). Diagnosis. Medium-sized species with symmetrical, rather wide aedeagus. Parameres almost straight in dorsal view, sinuate in lateral view. Internal sac with large basal vesicular portion bearing very fine spines, and distal portion with asymmetrical and 518 IVAN LÖBL flat median sclerite and two slender lateral sclerites. Body, metasternum excepted, very _ finely punctate. Metacoxae approximate. Description. Length 1.50 - 1.65 mm, width 0.96 - 1.07 mm. Body dark reddish-brown to blackish. Most external characters as in B.sordidoides, antennae longer, with ratio of segments as follows: IH 15, IV 15, V 17, VI 15, VII 18, VII 15, IX 20, X 19, XI 22 (holotype). Elytral punctation still finer than in B. sordidoides, anterior portion of sutural stria accompanied by several distinct punctures. Smooth latero-apical area of metasternum larger, metepisternal suture deeper and wider, with coarser punctures. Aedeagus (Figs 99, 100) 0.55 - 0.60 mm. Median lobe with fairly short apical portion. Internal sac with apical, irregularly margined lamina joined with two slender sclerites. Remarks. This species may be distinguished from B. sordida/sordidoides and from B. inermis by the shape of the median lobe and by the internal sac. Several females from other localities than those of the males, or slightly differing by their external characters, are not certainly conspecific with the males of B. laminula and therefore are not included in the type series. Baeocera reducta sp. n. Holotype, male: Nepal, Kaski distr., above Dumbus, 2100 m, 8-10.V.1980 (Martens & Ausobsky) (SMNS). Paratypes, 4 females: as holotype (SMNS, MNHG). Diagnosis. Micropterous member of the lenta group, with very short sutural striae of elytra, impunctate hypomeron, entire elytral surface coarsely punctate, parameres moderately sinuate and wide distal sclerite of internal sac of aedeagus. Description. Length 1.15 - 1.30 mm, width 0.78 - 087 mm. Body strongly convex, pale reddish-brown, antennae and tarsi yellowish. Eyes moderately large. Antennae long, relative length of segments as follows: III 12, IV 13, V 15, VI 12, VII 16, VII 14, IX 19, X 18, XI 20 (holotype). Segments III to VI slender, V somewhat wider than IV or VI; VII about 3x as long as wide, VIII 3.5 x as long as wide. Pronotum with rounded lateral margins; lateral keels not visible in dorsal view; punctation very fine, well delimited, visible at 24x magnification. Scutellum covered by pronotal lobe. Elytra rather strongly narrowed apically, with rounded lateral margins; lateral keel not visible in dorsal view; sutural area not or barely raised; sutural stria very shallow, distinct only in apical portion, evanescent on anterior half of sutural length. Entire punctation coarse and dense (including on humeral area), punctures near apex somewhat smaller and closer than those on centre. Wings strongly reduced. Pygidium very finely punctate. Hypomeron impunc- tate. Mesepisternum with several more or less distinct punctures. Entire metasternum coarsely punctate; punctures large, less dense laterally than on centre. Mesocoxal area very narrow. Mesepisternum not distinct. First ventrite coarsely punctate except on small smooth lateral area. Basal punctures not elongate. Pro- and mesotibiae straight, metatibiae somewhat curved. Segments 1 to 3 of protarsi somewhat enlarged in male. Aedeagus (Figs 101,102) 0.36 mm long. Median lobe with relatively large basal bulb and very short apical portion. Parameres very weakly sinuate. Remarks. This species shares with B. hygrophila Löbl most of the external and aedeagal characters. It may be distinguished from B. hygrophila in the much paler coloration of the body, the shorter sutural striae of the elytra, the reduced wings, the punctate mesepisternum and in the shape of the sclerites of the internal sac of the aedeagus. All specimens of B. reducta were found in a moist Sarauja napaulensis forest in western Nepal. SCAPHIDIIDAE OF NEPAL 519 Baeocera crinita sp. n. Holotype, male: Nepal, Sankhua Sabha distr., range S Mangsingma, 2800 m, 7.IV.1984, (Löbl & Smetana) (MHNG). Paratypes, 26: Sankhua Sabha distr., as holotype 1 female; “Bakan” W of Tashigaon, 3200m, 5.1V.1982 (A. & Z. Smetana) (MHNG) 1 female; Thudam, 3550-3650 m, 25.-27.V.1988 (Martens & Schawaller) (SMNS, MHNG) 4 males, 5 females; above Pahakhola, 2600-2800 m, 3.VI.1988 (Martens & Schawaller) (SMNS, MHNG) 1 male, 14 females. Diagnosis. Small-sized micropterous member of the /enta group characterized by very short antennae with wide segments VII to XI, very short sutural striae of elytra and distinctly microscultured Ist ventrite. Description. Length 1.0 - 1.1 mm, width 0.60 - 0.73 mm. Body moderately convex, dark reddish-brown. Antennae, tarsi and apex of abdomen paler. Eyes small. Antennae very short, relative length of segments as follows: III 8, IV 6, V 8, VI 6, VII 8, VIII 6, IX 9, X 10, XI 12 (holotype). Segments III and V somewhat wider than IV, VI notably wider than V; VII to XI each conspicuously wide, not or somewhat longer than wide. Dorsal punctation dense and fine, distinct at 24x magnification, that of pronotum somewhat finer than that of elytra. Pronotal and elytral keels not visible in dorsal view. Scutellum covered by pronotal lobe. Elytra rather strongly narrowed apically; sutural area usually somewhat raised, except anteriorly; sutural stria unsually short, visible only along apical half of sutural margin. Wings almost completely atrophied. Hypomeron and metepisternum impunctate. Mesepimeral ridge about 2x longer than interval between its end and mesocoxa. Middle portion of metasternum convex, entirely densely and rather finely punctate. Punctation of lateral portion of metasternum coarser and less dense than that of centre. Mesocoxal area very narrow. Metepisternum flat, 0.02 - 0.04 mm wide, with suture indicated by row of coarse punctures. Abdomen microsculptured. Pygidium extremely finely punctate. First ventrite with middle portion densely and rather finely punctate, laterally almost impunctate, basal punctures coarse. Tibiae straight. Segments 1 to 3 of male pro- and mesotarsi somewhat enlarged. Aedeagus (Figs 103-105) 0.29 - 034 mm long. Median lobe slender, its ventro-apical portion tapering. Parameres weakly sinuate. Remarks. This species seems to be related to B. microps Löbl which has a similar shape of the body, reduces eyes and wings, short antennae with wide segments VII to XI, and a similar aedeagus. It may be distinguished by the shorter sutural striae of the elytra, the impunctate lateral portion of the Ist ventrite, and by the darker and somewhat larger body. Baeocera crinita and B. microps are possibly vicarious and are restricted to small areas in Eastern Nepal and in Western Bengal, respectively. Baeocera microps has been found only in evergreen oak forest. The vertical range of B. crinita extends from the oak to the Betula/Rhododendron zone. Baeocera cribrata sp. n. Holotype male: Nepal, Panthar distr., Paniporua, 2300 m, 16 - 20.IV. 1888 (Martens & Schawaller) (SMNS). Paratypes, 7: as holotype (SMNS, MHNG) 1 male, 1 female; Nepal, Sankhua Sabha distr., Chi- chila above Ahale, 2300 m, 26.III.1982 (A. & Z. Smetana) (MHNG) 1 male; NE Kuwapani, 2250 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 1 male; forest S Mangsingma, 2200 m, 11-13.IV.1984 (Löbl & Smetana) (MHNG) 1 female; Nuwakot distr., Dinguari Kola above Trisuli Bazar, IX.-X.1971 (Franz) (Coll. H. Franz) 2 males. 520 IVAN LÖBL Diagnosis. Brachypterous member of the /enta group with conspicuously | coarsely punctate pronotum and coarsely punctate mesepisternum and hypomeron. Sutural stria of elytron very short. Aedeagus with relatively long median lobe and long sclerites of internal sac. Description. Length 1.15 - 1.30 mm, width 0.78 - 0.90 mm. Body reddish- brown, antennae and tarsi paler. Eyes small. Antennae of average length, with slender segments III to VI; relative length of segments in holotype as follows: III 13, IV 11, V 14; VI 13, VII 16, VIII 14, IX 17, X 16, XI 18. Segment VII about 3x as long as wide, VIII about 3.5x as long as wide. Pronotum with rounded lateral margins, lateral keels not visible in dorsal view; entire punctation very dense and coarse, that of centre distinct at 12x magnification , near lateral margin still coarser, with punctures larger than intervals. Scutellum barely exposed or completely covered by pronotal lobe. Elytra strongly convex, apically narrowed; lateral keel not visible in dorsal view; sutural stria very short, distinct only in apical half of sutural length ; sutural area flat; whole discal surface densely and very coarsely punctate, punctures about as large as those of lateral portion of pronotum. Wings reduced, not functional. Pygidium very finely punctate. Entire hypomeron coarsely punctate. Mesepimeral ridge about 2x longer than interval between its end and mesocoxa. Mesepisternum with numerous coarse punctures. Metasternum with smooth, convex centre; punctation around smooth area and near metacoxa very dense and coarse, elsewhere less dense but still coarser. Mesocoxal area very narrow. Metepisternum indistinct. Entire first ventrite without visible microsculpture (100x), densely and coarsely punctate. Basal punctures not elongate. Pro- and mesotibia straight, metatibia somewhat curved. Segments 1 to 3 of male protarsus somewhat enlarged. Aedeagus (Figs 106-108) 0.37 - 0.39 mm long. Median lobe with long basal bulb and very short apical portion. Parameres hardly sinuate. Remarks. Baeocera cribrata is possibly closely related to B. puncticollis with which it shares a similar aedeagus, the strongly shortened sutural striae and the punctate hypomeron. It may be easily separated from the latter by the much more coarsely punctate pronotum and by the punctate mesepisterna. The species is brachypterous. It occurs in mixed broad-leaved forest in Central and Eastern Nepal. Baeocera martensi sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley below Num, 1050 m, 22.IV.1984 (Löbl & Smetana) (MHNG). Paratypes, 20: Sankhua Sabha distr., as holotype (MHNG) 5 males, 6 females; Arun Valley bottom betw. Hedanga and Num, 950-1000 m, 6-8.V1.1988 (Martens & Schawaller) (SMNS, MHNG) 3 males, 6 females. Diagnosis. Member of the /enta group with distal portion of ejaculatory duct accompagnied by a bunch of moderately sclerotized denticles; parameres weekly sinuate; pronotum distinctly punctate; whole elytral disc coarsely punctate. Description. Length 1.0 - 1.2 mm, width 0.64 - 0.78 mm. Body more or less dark reddish brown, tarsi and antennae paler. Eyes moderately large. Antennae with narrow segments III to VI, similar to those in related species. Pronotum with rounded margins, lateral keels not visible in dorsal view, punctation rather fine, barely visible at 12x magnification, distinct at magnification 24x. Scutellum covered by pronotal lobe. Elytra narrowed apically, with sutural area raised in most specimens; sutural stria complete, extended along basal margin and joined with lateral margin; lateral keel not SCAPHIDIIDAE OF NEPAL 521 visible in dorsal view; punctation entirely coarse and dense, apical area finer punctate than remaining surface; most punctures distinctly larger than intervals between them. Wings apparently fully developed. Pygidium very finely punctate. Ventral surface similar as in related species. Hypomeron and mesepisternum impunctate; metasternum and Ist ventrite coarsely punctate, small smooth central area of metasternum excepted; mesocoxal area very narrow, visible portion of metepisternum narrow but distinct. Segments 1 to 3 of male protarsus moderately enlarged. Aedeagus (Figs 109 to 111) 0.29 - 0.33 mm long. Median lobe with large basal bulb and short apical portion. Parameres very weakly sinuate. Internal sac with a bunch of denticles. Remarks. This species is closely related with B. pigra, B. vidua and B. wittmeri; all share similar shape of the aedeagus, and the conspicuous sclerotized denticles along the ejaculatory duct, situated apically of the sclerotized pieces of the internal sac. These four species may be distinguished by the shape of the parameres, which are straight and evenly very narrow in B. vidua, curved and distally narrowed in B. pigra, and conspicuously long in B. wittmeri. Baeocera martensi may be readily distinguished from these three species by the much coarser pronotal punctation. All specimens of B. martensi were found in subtropical forests in Eastern Nepal. Baeocera schawalleri sp. n. Holotype, male: Nepal, Taplejung distr., Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (SMNS). Diagnosis. Medium-sized species with aedeagus having very large bulbous portion of median lobe, distal portion of median lobe asymmetrical, with two unequal, narrow sclerotized dorsal lobes and one wider ventral piece; parameres symmetrical, slender und simple; internal sac complex, without flagellum. Basal stria of elytron interrupted. Thoracal and abdominal punctation very fine or absent, punctation on elytra and metasternum coarse. Base of 1st ventrite with row of longitudinal furrows. Description. Length 1.5 mm, width 1.0 mm. Body moderately convex dorsally, very dark reddish-brown to blackish, apex of abdomen and legs paler, antennae ochreous. Eyes large. Antennae with elongate segments III to VI; relative length of antennal segments as follows: III 11, IV 12, V 16, VI 15, VII 16, VIII 13, X 15, XI 25. Segment V 4x as long as wide, as wide as III or VI, somewhat wider than IV; VII about 2.5x as long as wide, distinctly wider than VI, barely wider than VIII; VIII about 2x as long as wide; IX to XI notably wider than VII, XI 2.5x as long as wide. Pronotum with regularly rounded lateral margins: lateral keels not visible in dorsal view; punctation very fine and shallow, visible at 50x magnification. Distal portion of scutellum exposed. Elytra narrowed apically, lateral keels visible near base in dorsal view; sutural area flat, with row of distinct punctures; sutural stria deep, curved near base and extended along basal margin to humeral area, separated from lateral stria by narrow space; lateral stria very densely punctate; discal punctation coarse and dense; in basal half most punctures about as large as intervales, in apical half punctation less dense, most punctures smaller than intervales. Punctation of pygidium very fine, but well visible compared to that of pronotum. Hypomeron and mesepisternum impunctate. Mesepimeral ridge almost 3x as long as interval between its end and mesocoxa. Metasternum with smooth, rather flat centre, elsewhere coarsely punctate; punctures on lateral portion of metasternum not or weakly elongate, smaller and denser near metacoxa than those on anterior portion. Mesocoxal area very narrow, with coarse, not elongate marginal punctures extended laterally along anterior metasternal margin about to level of centre of mesepimeral ridge. Metepisternum 522 IVAN LÖBL somewhat vaulted, at widest point 0.09 mm, narrowed anteriad, its suture deep, punctate, “ somewhat rounded near anterior and posterior angles. Ventrites very finely punctate, without microsculpture. First ventrite with basal row of deep, up to 0.05 mm long furrows. Tibia straight. Segments 1 to 3 of male protarsus enlarged, mesotarsus with segment 1 enlarged. Aedeagus (Figs 112, 113) 0.58 mm long. Median lobe apically asymmetrical, at tip truncate, with two slender dorso-apical lobes. Internal sac bearing numerous spines and denticles, and several teeth-like sclerites. Parameres becoming slender apically and curved near tip. Remarks. Baeocera schawalleri shares essential diagnosic characters with B. insolita Löbl. It may be recognised in having asymmetrical distal portion of the median lobe and coarsely punctate metasternum. The new species was found in a forest remnant with bushes in Eastern Nepal. The wings of the unique specimen were not examined. They are likely fully developed, as in the other dark-coloured species. Baeocera mustangensis sp. n. Holotype male: Nepal, Mustang distr., Lete, 2550m, 2.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 5: as holotype (MHNG) 1 male and Lete, 1971 (Franz) (Coll. H. Franz) 1 male; Mustang distr., 2 km N Kalopani, 2550 m, 1.X.1983 (Löbl & Smetana) (MHNG) 1 male, 1 female; S Lete, 2450-2600 m, 30.IV.- 1.V.1980 (Martens & Ausobsky) (SMNS) 1 male. Diagnosis. Large species of the curtula group with distal portion of median lobe notched ventrally and bearing a bidentate lamina; parameres sinuate in lateral view, narrowed subapically; distal sclerotized portion of ejaculatory duct slender, basal sclerite of internal sac tooth-like, almost symmetrical. Description. Length 2.0 - 2.2 mm, width 1.25 - 1.50 mm. Body as in B. hamifera, rather strongly convex, very dark reddish-brown to black. Apex of abdomen, femora and tibiae more of less dark reddish, tarsi and antennae paler. Pronotum, elytra and pygidium not microsculptured. Eyes rather large. Antennae as in related species. Pronotal centre rather coarsely and very densely punctate, with punctures not well delimited and about as large as intervales, distinct at 12x magnification. Lateral and apical portions of pronotum decidedly finer punctate than centre. Distal portion of scutellum exposed. Elytron with deep, uninterrupted basal stria joined with lateral stria; elytral punctation coarse and dense, much coarser than that on pronotal centre, with many punctures about as large as intervales; punctation on apical portion less coarse than that on middle. Wings fully developed. Pygidium very finely punctate. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum with two rows of coarse setiferous punctures joined by medio-basal transverse row of coarse punctures. Mesocoxal area 0.05-0.07 mm long, margined by coarse punctures extended laterally to mesepimeral ridge. Metepisternum flat, 0.11 - 0.13 mm wide, with straight, punctate suture. Lateral portion of metasternum and Ist ventrite almost impunctate. Basal punctures of 1st ventrite coarse. Pro- and mesotibiae straight, metatibiae somewhat curved. Male protarsus with Ist segment very large, wider than apex of tibia, 2nd segment about as large, 3rd narrower than apex of tibia. Male mesotarsus with Ist segment almost as wide as apex of mesotibia, 2nd segment notably narrower. Aedeagus (Figs 114,115) 0.88 - 1.0 mm long. Median lobe gradually tapering in dorsal view, with asymmetricaly shaped apical portion in lateral view. Extruded portion of flagellum very long, parts of armature of internal sac strongly sclerotized. Remarks. This species has been collected in a Taxus and broad-leaved forest in the Kali Gandaki Valley in Western Nepal. Wider distribution might be expected as it has SCAPHIDIIDAE OF NEPAL 523 fully developed wings. It differs from B. hamifera significantly in the shape of the aedeagus. It may be distinguished from other Old World species of the group also in the pronotal and elytral punctation, in combination with the size of the body. Baeocera thoracica sp. n. Holotype, male: Nepal, Mustang distr., Lete, 2550 m, 2.X.1983 (Smetana & Löbl) (MHNG). Paratypes, 7: as holotype, 1 male; Parbat distr., Ghoropani Pass, 2700 m, 6.X.1983 (Löbl & Smetana) 1 male; Pun Hill at Ghoropani Pass, 3050-3100 m, 8.X.1983 (Löbl & Smetana) 1 male; Sindhupalcok distr., Malemchi, 2800 m, 14. and 17.1V.1981 (Löbl & Smetana) 1 male, 2 females (all MHNG); India, Himachal Pradesh, Simla, Kufri, 16.VII.1989 (Riedel) (SMNS) 1 male. Diagnosis. Member of the satana group. Distal portion of median lobe of aedeagus sinuate, with dorsally notched, not widened apex. Parameres sinuate, without abruptly notched or enlarged areas. Male metasternum simple, metatibia in male with sub- basal tooth. Elytra dark reddish brown, in male darker than pronotum, in female paler than pronotum. Description. Very similar to B. monstrosetibialis and B. dentipes. Length 1.65 - 1.85 mm, width 1.16 - 1.27 mm. Elytra more or less dark reddish-brown in both sexes, thorax and abdomen ochreous in male, blackish in female. Tarsi and antennae usually somewhat paler than tibiae. Relative lenght of antennal segments as follows: III 14, IV 18, V 20, VI 20, VII 24, VII 21, IX 25, X 24, XI 32 (holotype); VII and VIII equally wide, VII about 3.5x as long as wide, VIII 3x as long as wide, XI almost 3x as long as wide. Sutural striae almost parallel, sutural area flat. Humeral hump small. Punctures on elytral disc usually smaller than or as large as intervales. Mesepimeral ridge 2 - 2.5x longer than interval between its end and mesocoxa. Middle portion of metasternum convex, densely and rather coarsely punctate laterally of smooth centre and with several additional coarse punctures near medio-apical process. Lateral portion of metasternum very finely punctate. Mesocoxal area 0.03 - 0.04 mm long, marginal punctures not elongate, extended laterally to level of mesepimeral ridge. Metepisternum vaulted, at widest point almost 0.10 mm, anteriad narrowed, its impunctate suture sinuate, very deep. Basal punctures of Ist ventrite elongate. Male with segments 1 and 2 of protarsus strongly enlarged, 1 almost as wide as apex of protibia, 2 distinctly narrower; segment 3 of protarsus moderately enlarged; male mesotarsus with Ist segment about as large as that of protarsus, 2nd much narrower, 3rd not enlarged. Male metatibia (Fig. 116) with pointed tooth behind basal fourth, beyond inner side flattened, and bearing fine longitudinal keel. Metasternum without sexual characters. Aedeagus (Figs 117 to 119) 0.92 - 0.96 mm long. Apical portion of median lobe curved to left, hardly widened near tip. Remarks. This species may be readily recognised by its sexual characters. It may be distinguished from B. monstrosetibialis also by the colour pattern and by the finer elytral punctation. It occurs in mixed broad-leaved forests in Western and Central Nepal, and in Himachal Pradesh. Baeocera tuberculosa sp. n. Holotype, male: Nepal, Parbat distr., Ghoropani Pass, 2850 m, 5.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 3: as holotype, 1 male, 1 female; Parbat distr., Ghoropani Pass, 2700 m, 6.X.1983 and 10 ridge E Ghoropani Pass, 3000 m, 7.X.1983 (Löbl & Smetana) 1 male (all MHNG). 524 IVAN LÖBL Diagnosis. Member of the satana group with black body except for ochreous prothorax and mesepisternum. Elytron with sutural area narrowed anteriorly and apically. Apex of median lobe of aedeagus strongly enlarged; apical portion of left paramere abruptly narrowed, right paramere sinuate. Description. Similar to B. thoracica from which it differs by the colour pattern, the elytra and the sexual characters. Length 1.65-1.85 mm, width 1.2-1.3 mm. Male with head, apical and basal margins of pronotum, elytra, and most of ventral side of body black. Prothorax ochreous, apical and basal pronotal margins excepted. Mese- pisternum ochreous as prothorax, or pale reddish. Apical abdominal segments very dark brown. Legs dark brown to black, trochanters reddish. Antennae yellowish or pale brown. In female, body entirely black. Elytron with sutural area vaulted and widest in middle, conspicuously narrowed anteriad and apicad; sutural striae very deep, extended along base to humeral hump, separated from lateral stria by wide space. Elytral disc uneven, with very shallow lateral and apical impressions. Elytral punctation fine near sutural stria and on apical portion, with punctures smaller than intervales, that on discal centre coarser, with many punctures about as large as intervales. In male, middle smooth metasternal area impressed, with coarse and dense punctation on each side of impunctate centre, and with conspicuous narrow elongate protuberance. Segments 1 and 2 of pro- and mesotarsus still wider than in B. thoracica, 1st segment of protarsus and mesotarsus as wide as apex of tibia. Metatibia with straight basal fourth, then moderately incurved, in apical two thirds equally wide. Aedeagus (Figs 120 to 122) 1.0 - 1.05 mm long. Median lobe asymmetrical, with apical portion strongly inflexed, and in dorsal view widened. Left paramere abruptly narrowed beyond middle. Remarks. This species exhibits unusual sexual characters. The male may be readily distinguished by its colour pattern from all Baeocera, B. errabunda excepted. Most specimens were found in debris of mixed Rhododendron-oak forest in Western Nepal. Baeocera errabunda sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Goru Dzure Dara, eastern slope, 3350 m, 9.1V.1984 (Löbl & Smetana) (MHNG). Diagnosis. Member of the satana group with black body, ochreous prothorax and mesepisternum excepted. Sutural area of elytron widened in middle. Median lobe of aedeagus with right distal piece not enlarged at apex and moderately longer than left piece, apex of latter pointed. Parameres incurved, distal half of left paramere abruptly narrowed, right paramere dentate. Description. In external characters very similar to B. tuberculosa. Length. 1.65 mm, width 1.15 mm. Whole pronotum ochreous, basal margin only somewhat darkened; trochanters and femora black. Elytral disc almost evenly convex, flattened on small subapical area. Elytral punctuation rather fine, with almost all punctures much smaller than intervales. Median metasternal impression shallow, impunctate, delimited in middle by two keels. Middle portion of metasternum with few moderately coarse punctures. Metatibia straight and gradually stouter from base to mid-length, in apical half somewhat incurved, slender, and flattened on inner side. Aedeagus (Figs 123 to 125) 1.0 mm long. Apical portion of median lobe strongly asymmetrical, near tip strongly inflexed, with weakly sclerotized dorso-apical valve. Remarks. The pale prothorax is possibly a male secondary sexual character, as in B. tuberculosa. The species was found by sifting grass-tufts, fern and moss in Eastern Nepal. SCAPHIDIIDAE OF NEPAL 525 Scaphisoma Leach Scaphisoma Leach, 1817, type species: Silpha agaricina Linnaeus, 1758, by monotypy. Scaphosoma Agazzi, 1846; injustified emendation. Scaphiomicrus Casey, 1900; type species: Scaphisoma pusilla LeConte, by original designation. Pseudoscaphosoma Pic, 1915; type species: Pseudoscaphosoma testaceomaculatum Pic 1915, by monotypy. Scutoscaphosoma Pic, 1916; type species: Scutoscaphosoma rouyeri Pic, 1916, by monotypy. Scaphella Achard, 1924; type species: Scaphosoma antennatum Achard, 1919, by original designation. Scaphomicrus; Achard, 1924 (misspelling). Macrobaeocera Pic 1924; type species: Scaphosoma phungi Pic, 1922, by monotypy. Mimoscaphosoma Pic 1928 (sg); type species: Scaphosoma (Mimoscaphosoma) bruchi Pic, 1928, by monotypy. Scaphisoma is the largest genus of the family in terms of species diversity. It includes about 500 species and is distributed in all major biogeographical regions, Antarctis excepted. Its occurrence on many oceanic islands (i.e., Hawaii, Micronesia, Mascarene Archipelago) suggests high dispersal ability. However, the genus seems to be absent from areas with cool (or cooler) temperate climate such as south of South America or New Zealand where other scaphidiids are encountered. The distribution of the Himalayan species does also suggest their greater thermophily than that of Baeocera: Scaphisoma is notably more speciose but rarely found above the tree-line. Most of the 93 species encountered in the Himalayan region and in other North Indian and Pakistani mountain ranges occur in subtropical and mixed broad-leaved forests and seem to have a wide range of distribution. The Nepalese collections that included 53 species (57% of the Himalayan species of Scaphisoma, of which 21 are new and described below) are likely to be more representative for that area than collections made in other parts of the Himalaya. Additional 12 Himalayan species represented in the MHNG by females, or by males in poor condition, remain to be identified. They are not included in the present study. The genus may be defined by several presumably derived characters states: anten- nomere IV short, basal angles of pronotum extended apically toward metepisternum, metacoxal lines usually distinct, pubescence reduced. Members of this genus exhibit great diversity in aedeagal characters. Several species groups have been distinguished based on this character (i.e., LOBL 1970; 1971; 1979; 1981). The phylogenetic significance of these characters will be investigated in an ulterior study, when also information on Afrotropical and Neotropical species will be available. Only the presence, not the absence, of the hypomeral microsculpture and of the transverse row of punctures on the lateral portions of the metasternum are noted in the descriptions and redescriptions given below. For practical reasons, the species known only from Meghalaya are also included in the key. KEY TO THE HIMALAYAN SPECIES OF Scaphisoma MERE ron with basalistila fe A taser sg EI ta 2 ERE ron wathoutrbasalistHi:=- Cn 27 EL HO withsuninterrupfed!basalsttlaen nee 3 E salpa 9 3 Body uniformly blackish or black, elytron sometimes somewhat paler than pronotum 4 211 22 IVAN LÔBL Pronotum and/or elytra with distinct colour pattern ...................................... 5 Metacoxal area narrow, its margin parallel to metacoxa .............. kaszabianum Löbl Metacoxal area wide, its margin arcuate .............................. kashmirense Achard Elytron dark reddish-brown with wide pale apical portion and with obsolete discal PURELAHEONN etes een sense nenne UC SU DEEE baloo sp. n. Elytron with different colour pattern and distinct discal punctation ..................... 6 Abdominal microsculpture consisting of distinct transverse striae ... clavigerum sp.n. Abdominal microsculpture obsolete or consisting of punctures ......................... 7) Elytron yellowish or ochreous, with narrowly darkened base and one or two small dark spots on central portion of disc (Fig.27) ................................ aurorae Sp. n. Elytron ochreous to reddish-brown, with large dark discal spot extended to sutural stna'or:sutural:margin'(Fig.38) OI I 8 Internal sac of aedeagus without robust sclerite ............................. notatum Löbl Aedeagus with internal sac armed with robust sclerite ......................... jado sp.n. Abdominal microsculpture consisting of transverse striae .............................. 10 Abdominal microsculpture obsolete or consisting of punctures ........................ 13 Antennomere V about 3x as long as very short IV .................... pseudorufum Löbl Antennomere V as long as or moderately longer than IV ............................... 11 Elytron with large sub-basal reddish spot and da i ae SEE IZ) EAN , ... pulchellum Löbl Elytron without particular colour pattern .. SCOLO CR PAS TOI SS EI eee 12 Length below 2 mm. First ventrite with punctation on lateral portion much finer than thakon.centten. LTT RE a assimile curvistria Reitter Length 2 - 2.2 mm. First ventrite evenly, very finely punctate ........... tonkineum Pic Elytron dark reddish-brown, with well delimited yellowish subapical fascia ............ SE no IONI ENTI ROUE NS ae DURS LE CAES TOI flavofasciatum Löbl Colour pattern of.elytron.different:............0. een ner 14 Antennomere IV short, somewhat longer than III, both combined as long as or Shorter: than V. ur... RI LR IRE 15 Antennomere IV long, combined with III longer than V ................................ 20 Head and pronotum black, elytra reddish-brown. Length 2.25- 2.40 mm . SRI I I IRAN 05050468000 ones Tabl Colour;pattern different. Usually smaller species... 16 Antennomere VI as long as or longer than segments III to V combined .................. RIS AA A A antennatum Achard Antennomere VI shorter than segments III to V combined ............................. 17 Pronotum reddish-brown, elytra yellowish, with black basal spot ........... nima sp. n. Pronotum and elytra more or less uniformly dark reddish- brown ..................... 18 Antennomere VI notably longerthan Vi... inquietum sp. n. Antennomere VI as long as or distinctly shorter than V ................................. 19 Antennomere V-about 2x as lon as IV TE adjacens sp. n. Antennomere V about 4x'as long ‘as IV e praesigne sp. n. Length 2.4 - 2.5 mm. Body uniformly black. Antennomere XI more than 4x as long SWIC eee secret ili I ee ee TEN fatuum sp. n. Length not exceeding 2.2 mm.. Body usually not uniformly black. Antennomere XI much less than 4x as long as Wide 5:00... nee ee erie ee DAI Punctation on lateral portion of metasternum much coarser than that on lateral Portion Of Ist;ventrife..........-..-- oO viti Löbl Punctation on lateral portions of both metasternum and of Ist ventrite very fine .... 22 Metacoxalllıneparallelito'metacoxan 2 eee uniforme Löbl 37 38 SCAPHIDIIDAE OF NEPAL 527 IMetacoxalllinerarcualernen. RE ern sea loos ee une 23 Mesocoxal area small, shorter than metacoxal area ...................................... 24 Mesocoxal area fairly large, about as long as metacoxal area .......................... 25 Elytral disc finely punctate, diameters of punctures on centre smaller than those of IM LERVA! SEO nn stash ae ee Russ Dre NA NE AS ese eam DEN aes à AUS rufum Achard Elytral disc coarsely punctate, diameters of punctures on centre larger than those of TECA LS PE PET DL RENO RER US toa acme A BA TINO en RES interjectum sp. n. Median portion of metasternum with wide U-shaped row of coarse punctures .......... TE A RO IO att Men ele SN IRON minutum Achard Netastermnal\Punetatlo NAifere nen ER ee 26 Ventral process of aedeagus robust, oblique. Punctation on medio-basal portion of metasteinum'and'onilist'ventrite very dense ttt. neon puthzi Löbl Ventral process of aedeagus slender and curved. Punctation of median portion on metasternum and on Ist ventrite fairly dense .............................. besucheti Löbl Very small species, length less than 1 mm ..................... minutissimum Champion [Parccispeciesttati castel mm long Rene en 28 Antennomere IV conspicuously short, not or somewhat longer than III, both combinedishonenr than or aboutas lOme; aSiV. 2.2. er 29 Antennomere IV elongate, notably longer than III, both combined usually longer than N 008 RI ATTI EN RER ieee 36 Abdominal microsculpture consisting of punctures...................................... 30 Abdominal microsculpture consisting of transverse striae .............................. 32 Mesepimeral ridge distinct. Elytron with equally large discal punctures ................. TI caen ARA a stele sits Sa icaro falciferum Lobl Mesepimeral ridge obsolete. Elytron with conspicuously irregular punctation ....... 31 Elytral disc flattened, most of dense and coarse punctures situated on latero-central BOTH Se TR o ere US imitator Lobl Elytral disc not flattened, most of dense and coarse punctures situated on central PORHONO TAGES Corre cys eet II ira Note nice mimicum Löbl Elytron with two well delimited reddish spots ..................... quadrimaculatum Pic Elytronimmaculateas sont ne e o nice ace seamen 33 Antennomere VI much longer than V, usually about as long as segments III to V COMITE DES eee to a N o e e I o o A 34 Antennomere VI about as long as V, much shorter than II to V combined .............. LOOSE MARI TA LIA RIA ORIO enden Tara E I SIOE PR I RO peraffine Löbl Length 1.2 - 1.3 mm. Elytron uniformly blackish-brown or black ...... discretum Löbl Length 1.55 - 1.75 mm. Elytron black with pale apical margin ........................ 35 Parameres each equally wide in apical half ............................. fraterculum Löbl Parameres each gradually narrowed sub-apically, with widened apical portion ......... EEC ER NN fratellum sp. n. Pronotum and elytra dark brown to black, each elytron with reddish sub-basal spot or fascia, and with pale apical or subapical area. Metasternum with row of punctures Barallelitormetacoxag HH Weed. II rasen 37 Colourpatterniditferenti =: ur: ana ae Laden ne et ee 39 Elytron with subapical fasciae not extended to touch apical margin (Fig.33) ............ ee De ON OE VIII PIER OO RON IO EEC quadrifasciatum Löbl Entire apical 1/4 to 1/3 of elytron ochreous or yellowish (Fig.34) ..................... 38 Pronotum and elytra densely and coarsely punctate, punctures visible at magni- HC AMON RE 043) UO RR LIE tetrastictum Champion 528 33 SI IVAN LÖBL Pronotum and basal 1/3 of each elytron very finely punctate, punctures not or hardly VISIDICIALIM A PIIBICALIONISOXETA: I RIONI cederholmi Löbl Hypomeron with microsculpture consisting of striae ................................... 40 EHypomeronswithoutimicrosculpture.............22002. 202 2 TE eee 43 Metasternum without row of punctures parallel to metacoxa ......... invalidum sp. n. Metasternum with row of coarse punctures parallel to metacoxa ...................... 41 Punctation near lateral margins of pronotum conspicuously denser and coarser than thagonscontere ee SERI pseudodelictum Löbl Punctation near lateral margins of pronotum not or moderately coarser and denser than onicentre nes ee endeten ee ue DA ONE RIRES 42 Punctation on pronotal base and centre equally or almost equally fine .. argutum Löbl Punctation on pronotal base distinctly coarser than that on pronotal centre .............. RER TOTER ARE EN EHE OR OS Li 0000 malignum Löbl Elytron with more or less dark or black basal spot extended apically along suture and forming triangular or subtriangular pattern... 2... 2.2.2.2. ee 44 Elytralicoloration different \.............2: 22. ee eee EE EERE ERE CRs 48 Metasternum with row of relatively coarse punctures parallel to metacoxa .......... 45 Row of punctures parallel to metacoxa usually absent or formed by very fine PUNC MULES 2 FARBE Neon Heise egw eicie eue ei 47 Elytron without apical or subapical darkened patch. Middle portion of pronotum Garkeme dante en ee RE Re ee cruciatum Champion Elytron with distinct dark subapicalipatch i... 2 eRe 46 Median lobe of aedeagus apically symmetrical, flat, tapering ............ bhareko sp. n. Median lobe of aedeagus apically stick-like, asymmetrical .............. absurdum Löbl Middle portion of pronotum very finely punctate. Most of lateral portion of 1st ventnte notmicrosculptured sa... ee I sikkimense Sp. n. Pronotum coarsely punctate. Most of lateral portion of Ist ventrite microsculptured ... RR caress adie uals areas nine cece caus oubrew e ocio varium Löbl Lateral portion of metasternum without row of punctures parallel to metacoxa ..... 49 Lateral portion of metasternum with distinct row of punctures parallel to metacoxa LE esac AA A ER ABER BERNER ner nan ero aa EL.) 0030000000000 63 Abdominal microsculpture obsolete, or consisting of punctures or of extremely short SRO I EN ec Stee 0000000000050 50 Abdominal microsculpture consisting of distinct transverse striae ..................... SY Mesocoxal area subtriangular. 2.2... eee tee eee ee Reeeers Sil Mesocoxal'area'oval'or’arcuate;........-..-- essen nennen en nee 52 Apical portion of median lobe wide, strongly flattened .............. maindroni Achard Apical portion of median lobe narrow, weakly flattened ..................- spurium Löbl Pronotum and elytron very dark, blackish-brown or black, except for pale elytral APICES RM N ARE TEA EN Rennes bedeli Achard Pronotum and elytra uniformy more or less dark reddish-brown or elytra paler than pronotumi RE LR RSS ARR LILLO RE EEE 33 Median lobe of aedeagus conspicuously thick, with strongly inclined apical portion ... OS OA RE 0 o DIL OL LOT diabolum Löbl Median lobe of aedeagus narrow . 22.222220 TT 55 Aedeagus weakly sclerotized, with median lobe narrowed apically, parameres thin .... D So As eso RER RI eRe eee TI 56 Aedeagus strongly sclerotized, in dorsal view apically widening or subparallel, Parameres1tobust EM RSS ee corbetti Löbl SCAPHIDIIDAE OF NEPAL 529 Parameres straight, symmetrical, widened at middle ....................... suknense Löbl Parameres arcuate, asymmetrical, extremely thin, not widened at middle ................ RS TR ee nee unicolor Achard Lateral portions of metasternum and Ist ventrite coarsely punctate ... fulcratum sp. n. Lateral portions of metasternum and Ist ventrite very finely punctate ................ 58 Pronotum and elytra very dark reddish-brown to black, elytra with narrow apical portionldistinetiy; Palen: Aiea. Meer ee ee RE een en engen eee tes 59 Pronotum and elytra uniformly ochreous or more or less dark reddish-brown, apical elytral margin not or somewhat paler than elytral disc .................................. 60 Medio-basal area of 1st ventrite coarsely punctate beyond basal apophyse .............. Reese melon en mate né Wid bie ony ado eh nie seen EA innotatum Pic Entire first abdominal ventrite very finely punctate, marginal pits of metacoxal area EXCE DLC MMs rence RE PE forcipatum Champion Aedeagus with distal portion of median lobe very short, with distinct sclerotized dorsalivalv.en Body pale u... nennen i garomontium Löbl Aedeagus with distal portion of median lobe long, lacking sclerotized dorsal valve. Bodyadark ne REISE 61 BarametesinattoW.. einen Tn simplicipenis sp. n. Barameresiwiden.... 0.0. MN ER sein eaters luctans Löbl Abdominal microsculpture consisting of punctures ..................................... 64 Abdominal microsculpture consisting of transverse striae .............................. 68 Pronotum and elytra uniformly pale reddish-brown or ochreous ............ aurun Lobl Pronotum and elytra dark reddish-brown or blackish, elytra usually paler than PROMOLITA ae Le I A 65 Mesocoxal area small, 0.03 - 0.04 mm long, with margin parallel to coxal cavity ...... DICCI TTT INTANTO ARIA IE OI SISSI RA ona ates nebulosum Löbl Mesocoxal area larger, with margin arcuate or sub-triangular ......................... 66 Biytmronidarkened!subapically ERP RE cates tees echinatum Löbl Elytron uniformy dark reddish-brown or with darkened base .......................... 67 Entretirstäventnite,very.finelyipunetater nennen nern nefastum Löbl First ventrite with more or less coarsely punctate medio-basal area .... negligens Löbl Sutural striae of elytra strongly converging apically .................................... Sutural striae of elytra parallel, or weakly converging apically ......................... 72 Pronotum and most of elytra uniformy colored, very dark, usually black, elytron with large, sharply delimited pale apical area (Fig.35)................... leucopyga Champion Colounpatterni different. pest IRR ee 70 Sutural stria of elytron curved near base (Fig. 36) ........................ binhanum (Pic) Suturalstmiaroßelytrontangulateinearbasen nr ee: val Pronotum dark reddish-brown, with coarsely punctate base ............ impolitum Löbl Entire pronotum pale, ochreous or yellowish, very finely punctate ......... dohertyi Pic Pronotum and hypomera much paler than ventral meso- and metathoral sclerites, sometimes bicolored. Elytron more or less distinctly maculate ........................ 78 Pronotum, hypomera and entire ventral surface of thorax concolorous ............... TA) Male pronotum ochreous, with black basal pattern (Fig.37), female pronotum dark ICOCISISDIOWOMLO: LIL RE maculigerum Löbl Pronotum in both sexes uniformly pale, or with basal margin very narrowly darkened LECCO RO LEO SORESINA IRR ARA I A SRE on 74 Aedeagus with parameres fairly wide, not lobed ...................... nigrofasciatum Pic Parameres of aedeagus each extended ventrally by large lobe with inner margin SEFONOIVISCIETOLZEGÌ.- 5 me einen ae en TT AR 75 530 89 90 IVAN LÖBL Basal bulb of aedeagus extended dorso-apically, partly overlapping slender apical portion okmediantlobe 2.1... 02.2.0002: sean ces ee Pete atronotatum Pic Basallbulbiof aedeagus not extended apieally..................2.2...... ne 76 Length 1.85 - 2.05 mm. Internal sac of aedeagus with two pairs of large teeth ........... RER ET A RE khasianum Löbl Length 1.60 - 1.85 mm. Internal sac of aedeagus with single pair of large teeth ........ OR LI ii EEE ERR immo cooocoococ cs SINE LOI Antennomere V very long, about 3x as long as relatively short segment IV and 2x as long as segments III and IV combined ................................... onychiorum Löbl Antennomere V not particularly long, about as long as, or moderately longer than IV a PN PP En oc O cd EEE EEE EEE EERE 78 Aedeagus with apical portion of median lobe strongly asymmetrical, overlapped by DATAMOTES RAR M nenne due nen luctuosum Löbl Aedeagus symmetrical, apical portion of median lobe not overlapped by parameres ... EEE IE nie Meine gia o/h. dle Sd 132 0006.05600000066000050000 79 Bulbous portion of median lobe extended dorso-apically, overlapping partly narrow apically portionotpmediamlobey i 6.5 2.2 cosas ee immodicum Löbl Bulbous portion of median lobe not extended ........................................... 80 Median lobe of aedeagus with single undivided dorsal valve .......... flexuosum Löbl Median lobe with two dorsal valves extended by arms or with single valve divided by deep:sutüter....nen.ca ee TEE 81 Each paramere of aedeagus with denticulate expansion ................................ 82 Parameres ofaedeagus not.denticulate 52.2.8. tn oe eee eee eee 83 Basal and central portions of internal sac of aedeagus bearing strongly sclerotized ALMMALUTE I nn le eds ea el armatum Löbl Apical portion of internal sac of aedeagus bearing sclerotized armature ... minax Löbl Paramere of aedeagus not lobed or with minute lobe .................................... 84 Paramere of aedeagus with large membranous lobe ................................... 93 Apical portion of paramere of aedeagus delimited by minute tooth, very narrow ....... BADOO ED SCRE OC EET ERM OR D OM TO Ut db Oo bec b0 oo coalitum sp. n. Apical portion of paramere not delimited by tooth, as wide as, or wider than Subapical portion: 555: ae 85 Apical portion of paramere sharply delimited ................................. alacre sp. n. Apical portion of paramere not sharply delimited ....................................... 86 Inner margin of widened apical portion of paramere sinuate............ scabiosum Löbl Inner margin of apical portion of paramere not sinuate .................................. 87 Internal sac of aedeagus with symmetrical, flat or arcuate central lamina ............ 88 Internal sac of aedeagus without central lamina .......................................... 89 Lamina of internal sac of aedeagus angulate or sinuate anteriorly, strongly sclerotized laterally; apical sclerotized denticles of internal sac arranged in two rows ............... Be tn Shed Sn monde a nics tr RM Seine Si. RO SONORO ee ae nepalense sp. n. Lamina of internal sac of aedeagus tapering apically and evenly sclerotized; apical sclerotized! denticles arranged in single town... ne championi Löbl Basal portion of internal sac of aedeagus with row of strongly sclerotized denticles ... RN NR RER SER 20073500000 prehensor Champion Basal portion of internal sac of aedeagus covered by minute, weakly sclerotized Skale-like or spine-like structures... 90 Internal sac of aedeagus with central sclerotized denticles arranged in two groups pointed toward median line, and with dense apical group of sclerotized denticles ....... SSL A NN sie sic semis mt eee Oe eee eee eee geminatum Löbl SCAPHIDIIDAE OF NEPAL 53] eg Intemalisaciofiacdeapgusidifferente er. ee ene nee 91 91 Paramere of aedeagus with widest point (base excepted) situated far before middle ... ORE DR RER TER ETC GEBR kanchi sp. n. — Paramere of aedeagus parallel-sided up to middle or to apical third ................... 92 92 Arms of dorsal valves of aedeagus very short; proximal end of internal sac strongly inflecetedidorsallysan. aan PR nen ee cs De ten ne see varians sp. n. — Arms of dorsal valves of aedeagus elongate; proximal end of internal sac not or weakiyanflected aan en erden een necopinum Sp. n. 93 Internal sac of aedeagus with long inflected basal tube and with two conspicuous, tooth-like,seleritestinicentren m ee nee pinnigerum Sp. n. — Internal sac of aedeagus lacking basal tube and pair of central tooth-like sclerites ... 94 94 Parameres of aedeagus each widened apically, with large overlapping lobes E RIOT I NONNI netten ne CON LODI — Parameres of aedeagus each evenly wide, apically with small, not overlapping lobes 600 008085 SET RA TARN MECS III EDI ISO RA Re A EBEN indra Löbl NEW RECORDS Scaphisoma rufum Achard Material examined, 6 specimens (males only): Nepal, Kathmandu distr., Nagarjung forest, 1650 m, 2.IV.1981 (Lôbl & Smetana) (MHNG) 1; Sindhupalcok distr., Pokhare NE Barahbise, 2700 m, 2.V.1981 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 4. Distribution. North India, Nepal, Singapore, Japan. Scaphisoma pseudorufum Löbl Material examined, 3 specimens: Nepal, Manang distr., forest W Bagarchhap, 2200 m, 21.1X.1983 (Löbl & Smetana) (MHNG). Distribution. Nepal, India: Darjeeling distr., Thailand. Scaphisoma corbetti Löbl Material examined, 1 male: Nepal, Terai, Bara distr., Amlekhganj, 7-10.X.1970 (Franz) (Coll. H. Franz). Distribution. India: Garhwal and Kumaon, Nepal. Scaphisoma viti Löbl Material examined, 4 specimens: Nepal, Gorkha distr., Arughat-Suteo, 600-700 m, 27.VIL.1983 (Martens & Schawaller) (SMNS, MHNG). Distribution. North Pakistan, Nepal. 532 IVAN LÖBL Scaphisoma quadrifasciatum Löbl Material examined, 11 specimens: India, Uttar Pradesh, Chakrata div., 7000 ft., 1.V11.1932 (Champion) (BMNH) 4; Nepal, Kathmandu distr., Gokarna forest, 1400 m, 31.11.1981 (Löbl & Smetana) (MHNG) 1; Patan distr., Phulcoki (Franz) (Coll. Franz) 1; Taplejung distr., Yamputhin, 1650-1800 m, 26.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 5. Distribution. North Pakistan, North India, Nepal. Scaphisoma fraterculum Löbl Material examined, 28 specimens: Nepal, Patan distr., Phulcoki, 2500-2700 m, 10.V.1981, 13-16.X.1983, 28-30.IV.1984 (Löbl & Smetana) (MHNG) 16; Phulcoki, near Godawari, 1700 m, 10.V.1981 (Löbl & Smetana) (MHNG) 1; Kathmandu distr., Siwapuri Dara, 2500 m, 1.V.1985 (Smetana) (MHNG) 1; Sindhupalcok distr., Gul Bhanjyang, 2600 m, 6.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi, 2800 m, 18.IV.1981 (Löbl & Smetana) (MHNG) 1; Taplejung distr., above Yamputhin, left back of Kabeli Khola, 1800-2000 m, 27-29.IV.1988 (Martens & Schawaller) (SMNS) 4; SE Yamputhin to Yamputhin, 2000-1650 m, 26. and 30.1V.1988 (Martens & Schawaller) (MHNG) 1; Panchthar distr., Dhorpar Kharka, 2700 m, 16.1V.1988 (Martens & Schawaller) (MHNG) 1; Sankhua Sabha distr., Pahakhola, 2600-2800 m, 31.V.-3.V1.1988 (Materns & Schawaller) (SMNS) 1. Distribution. North India, Nepal. Scaphisoma notatum Löbl Material examined, 76 specimens: Nepal, Jumla distr., Dzunda Khola Valley near Talphi, W Jumla, 3000-3500 m, 19.1X.1972 (Franz) (Coll. H. Franz) 1; Parbat distr., Ghoropani Pass, 2850 m, 5.X.1983 (Löbl & Smetana) (MHNG) 1; Ridge east Ghoropani Pass, 3100 m, 7.X.1983 (Löbl & Smetana) (MHNG) 1; Manang distr., Latha Manang W Bagarchhap, 2350 m, 22.IX.1983 (Löbl & Smetana) (MHNG) 1; Gorkha distr., Chuling Khola, 3000-3400 m, 3.VIIL.1983 (Martens & Schawaller) (MHNG) 1; Kathmandu distr., Siwapuri Dara, 2400 m, 30.V.1985 (Smetana) (MHNG) 1; Siwapuri Dara, 2700 m, 24.111.1982 (Rougemont) (MHNG) 5; Siwapuri Dara, 2540 m, 7.X.1981 (Sakai) (Coll. M. Sakai) 2; Patan distr., Phulcoki, 2200-2700 m, VIII.1967, IV-V. 1981, IV.1982, X.1983 (Canadian Nepal Exp., Lôbl & Smetana, A. & Z. Smetana) (CNC, MHNG) 38; Godawari, 13.11.1981 (Rougemont) (MHNG) 1; Sindhupalcok distr., Chaubas, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 1; Gul Bhanjyang, 2600 m, 6.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi, 2800-2900 m, 14. and 17.IV.1981 (Löbl & Smetana) 11; Pokhare NE Barahbise, 2700, 2800, and 3000 m, 2. and 7.V.1981 (Löbl & Smetana) (MHNG) 6; Sankhua Sabha distr., forest above Ahale, 2300 m, 26.11.1982 (A. & Z. Smetana) (MHNG) 3; Taplejung distr., Gunsa Khola betw. Kibla and Amjilera, 2600-2400 m, 12. IX.1983 (Martens & Dams) (SMNS) 1. Distribution. North Pakistan, North India, Nepal. Scaphisoma uniforme Löbl Material examined, 52 specimens: Nepal, Manang distr. forest W Bagarchhap, 2200 m, 21.IX.1983 (Löbl & Smetana) (MHNG) 3; Parbat distr., Ghoropani Pass, N slope, 2700 m, 6.X.1983 (Löbl & Smetana) (MHNG) 5; Kathmandu distr., Siwapuri Dara, 2400 m, 29.IV.1985 (Smetana) (MHNG) 1; Siwapuri, 2700 m, 24.11.1982 (Rougemont) (MHNG) 1; Patan distr., Phulcoki, 2500-2600 m, 13-14.X.1983 and 28-30.IV.1984 (Löbl & Smetana) (MHNG) 32; Phulcoki, 2600 m, 20.1V.1982 (A. & Z.Smetana) 1; Sindhupalcok distr., Pokhare NE Barahbise, 2800 m, 3.V.1981 (Löbl & Smetana) (MHNG)1; Sankhua Sabha distr., Arun Valley, Chichila, 1900-2000 m, SCAPHIDIIDAE OF NEPAL 533 18-20.VI.1981 (Martens & Schawaller) (SMNS) 1; forest NE Kuwapani, 2500 m, 28.11.1982 (A. & Z. Smetana) (MHNG) 2; above Ahale, 2300 m, 26.11.1982 (A. & Z. Smetana) (MHNG) 1; Induwa Khola Valley, 2400 m, 15.1V.1984 (Löbl & Smetana) (MHNG) 1; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 3. Distribution. North India, Nepal. Scaphisoma kaszabianum Löbl Material examined, 106 specimens: Nepal, Manang distr., Bagarchhap, 2200 m, 21.1X.1983 (Löbl & Smetana) (MHNG) 2; Parbat distr., betw. Chitre and Ghandrung, 6.V.1980 (Martens & Ausobsky) (SMNS) 1; Ghoropani, N. slope, 2700 m, 6.X. and Ghoropani Pass, 2850 m, 9.X.1983 (Löbl & Smetana) (MHNG) 6; Patan distr., Godawari, 6000 ft, 7-13.VIII.1967 (Canad. Nepal Exp.) 4; 2 km S Godawari, 1700 m, 19.X.1983 (Löbl & Smetana) (MHNG) 4; Phulcoki, 6600 ft, 13-17.VIII.1967 (Canad. Nepal Exp.) 1; Phulcoki, 2300-2600 m, X.1983 and V.1984 (Löbl & Smetana) (MHNG) 20; Kathmandu distr., Siwapuri, 2100-2300 m, 25.V1.1988 (Martens & Schawaller) (SMNS) 1; Siwapuri Dara, 2400 and 2450 m, 29. and 30.1V.1985 (Smetana) (MHNG) 2; Sindhupalcok distr., Gul Bhanjyang, 2600 m, 6.1V.1981 (Löbl & Smetana) (MHNG) 4; Malemchi, 2800 and 2900 m, 14-18.IV.1981 (Löbl & Smetana) (MHNG) 3; Pokhare NE Bharabise, 2700 m, 2. and 7.V.1981 (Löbl & Smetana) (MHNG) 13; Sankhua Sabha distr., forest NE Kuwapani, 2500 m, 11-15.1V.1982 (A. & Z.Smetana) and 2250 m, 24.1V.1984 (MHNG, CNC) 14; Arun Valley, betw. Mure and Hurure, 2050-2150 m, 17.VI.1988 (Martens & Schawaller) (SMNS) 2; forest S Mangsignma, 2250 m, 12.1V.1984 (Löbl & Smetana) (MHNG) 5; Induwa Khola Valley, 2000 m, 18.1V.1984 (Löbl & Smetana) (MHNG) 12; Panchthar distr., Paniporua, 2300 m, 16-20.IV.1988 (Martens & Schawaller) (SMNS) 10, Ilam distr., Mai Pokhari, 2100-2200 m, 25-27.III.1980 (Martens & Ausobsky) 1; Taplejung distr., above Yamputhin, left bank of Kabeli Khola, 1800-2000 m, 27- 29.1V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. North India (West Bengal: Darjeeling district), Nepal. Scaphisoma unicolor Achard Material examined, 6 specimens (males only): Nepal, Patan distr., Godawari, Phulcoki, 1700 m, 10.V.1981 (Löbl) (MHNG) 1; Sankhua Sabha distr., Induwa Khola Valley, 1750 m, 14.1V.1984 (Löbl & Smetana) (MHNG) 1; Taplejung distr., SE Yamputhin to Yamputhin, 2000- 1650 m, 30.IV.1988 (Martens & Schawaller) (SMNS) 1; above Yamputhin, left bank of Kabeli Khola, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS) 1; betw. Hellok and lower Gunza Khola, 2000-1650 m, 18.III.1988 (Marterns & Schawaller) (SMNS, MHNG) 2. Distribution. Japan, Taiwan, Thailand, Nepal. Scaphisoma besucheti Löbl Material examined, 3 specimens: Nepal, Terai, Hitauri, 9.X.1972 (Franz) (Coll. H. Franz) 2; Parsa distr., Birganj Lothar 450 ft., 5-12.IX.1967 (Canad. Nepal Exp.) (CNC) 1. Distribution. Sri Lanka, India, Nepal. Scaphisoma maindroni Champion Material examined, 17 specimens: Nepal, Terai, Hitauri, 9.X.1972 (Franz) (Coll. H. Franz) 2; Lamjung distr., Bahun danda, 1300 m, 18.IX.1983 (Löbl & Smetana) (MHNG) 4; 534 IVAN LÖBL Marsyandi, 1200 m, Jagat, 11.1V.1980, (Martens & Ausobsky) (SMNS) 4; Gorkha distr., Daronti Khola betw. Doreni and Motar, 900-750 m, 13.VIII.1983 (Martens & Schawaller) (SMNS) 1; Sankhua “ Sabha distr., Dunge Dara N Tumlingtar, 1100 m, 23.11.1985 (A. & Z. Smetana) (MHNG) 3; Lamobagar Gao, 1400 m, 28-31.V.1980 (Wittmer) (NMB) 1; Ilam distr., betw. Ilam and Mai Pokhari, 1400-1600 m, 8.IV.1988 (Martens & Schawaller) (SMNS) 1; Sanishara, 5 km N, 270-300 m, 3- 5.1V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. Pakistan, India, Nepal, Burma, Thailand, Vietnam, China. Remarks. Scaphisoma mutatum Champion was placed in synonymy of S. maindroni (LOBL 1986b). A male syntype of S. mutatum labelled “W.Almora Kumaon U.P. India H.G.C.” is here desingated as lectotype (BMNH); 32 syntypes, sex not examined, from India, Kumaon, Haldwani distr. (BMNH) are designated as paralecto- types. Scaphisoma spurium Löbl Material examined, 26 specimens: India, Uttar Pradesh, Kumaon, Haldwani div. (Champion) (BMNH). Distribution. Sri Lanka, North India: Kumaon and Garhwal. Scaphisoma falciferum Löbl Material examined, 2 specimens: Nepal, Manang distr., forest W Bagarchhap, 2200 m, 21.1X.1983 (Löbl & Smetana) (MHNG). Distribution. Pakistan, North India: Kumaon and Himachal Pradesh, Nepal. Scaphisoma diabolum Löbl Material examined, 1 specimen: Nepal, Sankhua Sabha distr., Arun Valley below Num, 1050 m, 22.IV.1984 (Löbl & Smetana) (MHNG). Distribution. India: Darjeeling district, Nepal. Scaphisoma leucopyga Champion Material examined, 2 specimens: Nepal, Chitawan distr., Chitawan Nat. Park, X.1980 (Rougemont) 1; Kaski distr., betw. Viletadi and Uleri, trek from Pokhara to Ghoropani Pass, 18.IX.1971 (Franz) (Coll. H . Franz) 1. Distribution. Afghanistan, Pakistan, North India, Nepal. Scaphisoma nigrofasciatum Pic Material examined, 3 specimens: Nepal, Gorka distr., Arighat - Suteo, 600-700 m, 27.VII.1983 (Martens & Schawaller) (SMNS, MHNG) 3. Distribution. India, Nepal, Sri Lanka, Mascarene island, Seychelles. SCAPHIDIIDAE OF NEPAL 535 Scaphisoma binhanum (Pic) Material examined, 2 specimens: Nepal, Sankhua Sabha distr., Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 1; Arun Valley, Lamobagar Gola, 1000-1400 m, 27.V.-3.V1.1980 (Holzschuh) (NMB) 1. Distribution. North India, Nepal, Burma, Thailand, Vietnam. Scaphisoma atronotatum Pic Material examined, 67 specimens: Nepal, Kathmandu distr., Gokarna forest, 1400 m, 31.11. - 1.IV.1981, 20.X.1983 (Löbl & Smetana) (MHNG) 62; Patan distr., Phulcoki, Godawari, 1700 m, 10.V.1981 (Löbl & Smetana) (MHNG) 1; Lamjung distr., Marsyandi, 1200 m, Jagat, 11.1V.1980 (Martens & Ausobsky) (SMNS) 1; Sankhua Sabha distr., Arun Valley below Num, 1050 m, 21.1V.1984 (Löbl & Smetana) (MHNG) 1; Taplejung distr., SE Yamputhin to Yamputhin, 2000-1650 m, 26. and 30. IV.1988 (Martens & Schawaller) (SMNS) 1; Ilam distr., Citang Khola Valley, 1750 m, 11-13.1V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. Nepal, Burma, Thailand. Scaphisoma malignum Löbl Material examined, 28 specimens: India, Uttar Pradesh, Kumaon, Haldwani distr. (Champion) (BMNH) 1; Nepal, Parsa distr., Lothar near Birjang, 450 ft., 5-12.IX.1967 (Canad. Nepal Exp.) (CNC) 1; Kathmandu distr., Gokarna forest, 1400 m, 31.II. - 1.IV.1981 (Löbl & Smetana) (MHNG) 16; Sankhua Sabha distr., Arun Valley below Num, 1050 m, 21.1V.1984 (Löbl & Smetana) (MHNG) 2; Arun Valley, Lamobagar Gola, 1000-1400 m, 27.V.-3.VI.1980 (Holzschuh) (NMB, MHNG) 6; Arun Valley, Dunge Dara N Tumlingtar, 1100 m, 23.11.1982 (A. & Z.Smetana) (MHNG) 2. Distribution. North India, Nepal. Scaphisoma aurun Löbl Material examined, 1 specimen: Nepal, Ilam distr., 5 km N Panishare, feet of Siwalik Mts, 270-300 m, 3-5.1V.1988 (Martens & Schawaller) (SMNS). Distribution. Pakistan, South and North India, Nepal. Scaphisoma indra Lob! Material examined, 11 specimens: Nepal, Kathmandu, Nagarjung, Jamacok, 1900- 2100 m, 18.VIII.1983 (Martens & Schawaller) (SMNS, MHNG) 6; Patan distr., Godawari, 13.11.1981 (Rougemont) and 1600 m, 31.11.1984 (Löbl & Smetana) (MHNG) 5. Distribution. North India: Darjeeling district, Nepal. Scaphisoma tetrastictum Champion Material examined, 19 specimens: Nepal, Kathmandu distr., Gokarna forest, 1400 m, 1.VI.1981 (Lôbl & Smetana) (MHNG) 3; Ilam distr., SW Ilam below Parbate, 1250-1450 m, 536 IVAN LÖBL 23.VIIL.1983 (Martens & Dams) (SMNS) 1; India, Uttar Pradesh, Kumaon, Tanakpur (Champion) (BMNH) 14, Kumaon, Ranikhet (BMNH) 1. Distribution. India, Nepal, Burma, Thailand, Vietnam, Taiwan. Scaphisoma varium Löbl Material examined, 3 specimens: Nepal, Sankhua Sabha distr., forestr NE Kuwapani, 2250 m, 24.IV.1984 (Löbl & Smetana) (MHNG) 1; Panchthar distr., Paniporua, 2300 m, 16-20.1V.1988 (Martens & Schawaller) (MHNG) 1; ridge betw. Sheldoti and Paniporua, 2450-2200 m, 29.VIIL.83 (Martens & Dams) (SMNS) 1. Distribution. North India: Darjeeling district, Nepal, Bhutan. Scaphisoma prehensor Champion Material examined, 4 (males only): Nepal, Hitaura, Terai, 9.X.1972 (Franz) (Coll. H. Franz) 1; Ilam distr., 5 km N Sanishare, Siwalik Mts, 270-300 m, 3-5.IV.1988 (Martens & Schawaller) (SMNS, MHNG) 3. Distribution. North India, Nepal. Scaphisoma nefastum Löbl Material examined, 38 specimens: Nepal, Mustang distr., Lete, 2550 m, 2.X.1983 (Löbl & Smetana) (MHNG) 1; Manang distr., forest W Bagarchhap, 2250 m, 22.IX.1983 (Löbl & Smetana) (MHNG) 10; Parbat distr., Chitre, 2400 m, 4.V.1980 (Martens & Ausobsky) (SMNS) 1; Nuwakot distr., Trisuli Khola, Dunche, 21-26.IX.1981 (Pawlowsky & Kuska) (IZK) 1; Kathmandu distr., Siwapuri, 2100-2300 m, 25.VI.1988 (Martens & Schawaller) 3; Siwapuri, 2450 m, 29.IV.1985 (Smetana) (MHNG) 1; Patan distr., 2 km S Godawari, 1700 m, 19.X.1983 (Löbl & Smetana) (MHNG) 1; Phulcoki, 2600 m, 13.X.1983 (Löbl & Smetana) (MHNG) 1; Phulcoki, 6600 ft, 4- 7.VIIL.1967 (Canadian Nepal Exp.) (CNC) 1; Sindhupalcok distr., Pokhare NE Barabhise, 2700 m, 2.V.1981 (Löbl & Smetana) (MHNG) 2; Sankhua Sabha distr., forest NE Kuwapani, 2500 m, 11-15. IV.1982 (A. & Z.Smetana) (MHNG) 3, and 2250 m, 24.IV.1981 (Löbl & Smetana) (MHNG) 2; Induwa Khola Valley, 2000 m, 16.IV.1984 (Löbl & Smetana) (MHNG) 2; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 3; Panchtar distr., Paniporua, 2300 m, 16- 20.IV.1988 (Martens & Schawaller) (SMNS) 1; Taplejung distr., Omje Kharka NW Yamputhin, 2300-2500 m, 1-6.V.1988 (Martens & Schawaller) (SMNS,MHNG) 5. Distribution. North India: Kumaon, Darjeeling district, Nepal. Scaphisoma necopinum Löbl Material examined, 133 specimens: Nepal, Patan distr., Godawari, 7-13.VIII.1967 (Canadian Nepal Exp.) (CNC) 1; 2 km S Godawari, 1700 m, 19-20.X.1983 (Löbl & Smetana) (MHNG) 4; Phulcoki, Godawari, 1700m, 10.V.1981 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., below Sheduwa, 2550 m, 30.11.1982 and 2100-2550 m, 9.IV.1982 (A.& Z. Smetana) (MHNG) 13; Induwa Khola Valley, 2000 m, 16.1V.1984 (Löbl & Smetana) 15; Taplejung distr., Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (SMNS) 3; above Yamputhin, felt bank of Kabeli Khola, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS) 4; Yamputhin, open forest, 1650-1800 m, 26.IV. - 1.V.1988 (Martens & Schawaller) (SMNS, MHNG) SCAPHIDIIDAE OF NEPAL 537 91; Panchtar distr., betw. Paniporua and Hinwa Khola Valley, 3200-1850 m, 27.1V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. North India: Darjeeling district and Sikkim, Nepal. Scaphisoma fortipatum Champion Material examined, 72 specimens: Nepal, Myagdi distr., Myangdi Khola, Muri, S Dhaulagiri, 2100-2300 m, III.1970 (Martens) (SMNS) 1; Kaski distr., Tandarakot, trek Pokhara - Ghoropani, 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz) 3; Kathmandu distr., Gokarna, 1400 m, 31.III. and 1.IV. 1981, 10.IX. and 20.X.1983 (Löbl & Smetana) (MHNG) 18; Gokarna, 1400, 3.VIII.1970 and 3.X.1971 (Franz) (Coll. H. Franz) 3; Nagarjung forest, 1650 m, 2.IV.1981 (Löbl & Smetana) (MHNG) 2; Nagarjung, Jamacok, 1900-2000 m, 18.III. 1983 (Martens & Schawaller) (SMNS) 6; Siwapuri, 2100-2300 m, 25.V1.1988 (Martens & Schawaller) (SMNS) 1; Siwapuri Dara, 2300 m, 3.V.1985 (Smetana) (MHNG) 1; Patan distr., 2 km S Godawari, 1700 m, 19-20.X.1983 and 1600 m, 31.111:1984 (Löbl & Smetana) (MHNG) 8; Godawaki, 1770 m, 19.11.1980 (Martens & Ausobsky) (SMNS) 1; Phulcoki SE Godawari, cca 2000 m, 21.IV.1988 (Brachat) (MHNG) 4; Sindhupalcok distr., Chaubas, 2500 m, 4.1V.1981 (Löbl & Smetana) (MHNG) 3; Burlang Bhanjyang, 2600 m, 5.IV.1981 (Löbl & Smetana) (MHNG) 1; Pokhare NE Barahbise, 2800 m, 2.V.1981 (Löbl & Smetana) (MHNG) 1; Durumtali near Barahbise, 2200-2300 m, 5.VIII.1970 (Franz) (Coll. H. Franz) 1; Sankhua Sabha distr., below Sheduwa, 2550 m, 30.11.1982 (A. & Z.Smetana) (MHNG) 1; bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 3; bottom Arun Valley, betw. Hedangna and Num, 950-1000 m, 6-8.V1.1988 (Martens & Schawaller) (SMNS) 8; Taplejung distr., Tada Khola, Khebang, 1600-1800 m, 2.IX.1983 (Martens & Daams) (SMNS) 1; confluence Kabeli Khola and Tada Khola, 1000-1050 m, 23-25.1V.1988 (Martens & Schawaller) (SMNS) 2; ascent to Khebang from Tada Khola, 1500 m, 25.1V.1988 (Martens & Schawaller) (SMNS) 2; Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (SMNS) 2; Ilam distr., betw. Ilam and Mai Pokhari, 1600-2000 m, 9.1V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. Pakistan, North India, Nepal. Scaphisoma innotatum Pic Material examined, 34 specimens: Nepal, Sankhua Sabha distr., Arun Valley at Num, 1500-1600 m, 10.1V.1982 (A. & Z.Smetana) (MHNG) 1; bottom Arun Valley below Num, 1050 m, 22.1V.1984 (Löbl & Smetana) (MHNG) 27; bottom Arun Valley, betw. Hedangna and Num, 950-1000 m, 6-8.V1.1988 (Martens & Schawaller) (SMNS) 6. Distribution. North India, Nepal, Thailand, Vietnam. Scaphisoma armatum Löbl Material examined, 33 specimens: Nepal, Kathmandu distr., Gokarna, 1400 m, 31.10 and 1.1V.1981, 10.X.1983 (Löbl & Smetana) (MHNG) 10; Patan distr., Godawari, 1600 m, 31.11.84 (Löbl & Smetana) (MHNG) 1; Sindhupalcok distr., Chaubas, 2600 m, 12.1V.1981 (Löbl & Smetana) (MHNG) 1; Rasuwa distr., 1.5 km NE. Bhargu, 2000 m, 12.1V.1985 (Smetana) 2; Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 21.IV.1984 (Löbl & Smetana) (MHNG) 1: below Sheduwa, 2100-2550 m, 9.IV. and 2550 m, 30.11.1982 (A. & Z. Smetana) (MHNG) 5; Induwa Khola Valley, 2000 m, 6.1V.1984 (Löbl & Smetana) (MHNG) 5; Ilam distr., Mai Pokhari, 2100-2200 m, 25-27.111.1980 (Martens & Ausobsky) (SMNS) 3; Taplejung distr., Yamputhin cultural land, 1650-1800 m, 26.1V.1988 (Martens & Schawaller) (SMNS) 5. Distribution. North India, Nepal, Thailand. 538 IVAN LÖBL Scaphisoma minax Löbl Material examined, 18 specimens: Patan distr., 2 km S Godawari, 1700 m, 20.X.1983 (Löbl & Smetana) (MHNG) 2; Godawari, 1770 m, 17.11.1980 (Martens & Ausobsky) (SMNS) 2; Phulcoki SE Godawari, cca 1800 m, 22.1V.1988 (Brachat) (MHNG) 1; Phulcoki, 2700 m, 16.X.1983 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., forest above Ahale, 2300 m, 26.11.1982 (A.& Z.Smetana) (MHNG) 1; Arun Valley, Num-Chichila, 1500-1900 m, 17.VI.1980 (Holzschuh) (MHNG) 1; Arun Valley, betw. Mure and Hurure, 2050-2150 m, 17.V1.1988 (Martens & Schawaller) (SMNS) 4; Taplejung distr., SE Yamputhin to Yamputhin, 2000-1650 m, 26. and 30.1V.1988 (Martens & Schawaller) (SMNS) 2; Yamputhin, 1650-1800 m, 26.IV.-1.V.1988 (Martens & Schawaller) (SMNS) 3; Worebung Pass, 2000 m, 21.1V.1988 (Martens & Schawaller) (MHNG) 1. Distribution. North India: Darjeeling district, Meghalaya; Nepal. Scaphisoma immodicum Löbl Material examined, 3 specimens: India, Himachal Pradesh, Kulu Valley, Naggar, 1850 m, 15.X.1988 (Vit) (MHNG). Distribution. India: Himachal Pradesh. Scaphisoma absurdum Löbl Material examined, 2 specimens: Nepal, Gorkha/Dhading distr., Gorlabesi-Dobhan, 1000-1100 m, 30.VIL.1983 (Martens & Schawaller) (SMNS) 1; Lamjung distr., Marsyandi, 1100- 1250 m, Senghe-Jagat, 11.1V.1980 (Martens & Ausobsky) (MHNG) 1. Distribution. North India: Darjeeling district, Nepal. REDESCRIPTIONS Scaphisoma minutissimum Champion Scaphosoma minutissimum CHAMPION, 1927: 278. Diagnostic characters. Length 0.90 mm, width 0.59 mm. Body ochreous, apices of elytra and abdomen paler. Legs and antennae yellowish. Relative length of antennomeres as follows: III 3, IV 5, V 10, VI 10, VII 13, VIII 9, IX 14, X 14, XI 20 (lectotype); segment V and XI about 3x, VI and VII about 2.5x, VIII 2x as long as wide. Pronotum with weakly arcuate lateral margins, lateral keels barely visible in dorsal view, discal punctation obsolete. Tip of scutellum exposed. Elytron with lateral margin oblique except near base; lateral keel visible in dorsal view; apical margin arcuate; inner apical angle at same level as outer angle; sutural margin not raised; sutural area flat; sutural stria parallel to suture, not curved near base and not extended laterally; discal punctation obsolete. Mesepimeral ridge longer than interval between its end and mesocoxa. Metastenum without any impression, not microsculptured, punctation obsolete (100x magnification). Mesocoxal area 0.04 mm long, with arcuate, very finely punctate margin. Metepisternum strongly narrowed anteriad, at same level as metasternum, with straight suture. First ventrite lacking visible microsculpture, punctation extremely fine (180x magnification). Metacoxal area 0.05 mm long, with margin arcuate, very finely SCAPHIDIIDAE OF NEPAL 539 punctate. Tibia straight. Aedeagus (Fig.126) 0.23 mm long. Median lobe tapering, pointed. Parameres weakly sclerotized apically, curved. Internal sac with very slender flagellum. Type material. Lectotype male, labelled “R.Sarda Gorge Kumaon, U.P. Dec. 1918 H.G.C.” (BMNH), by present designation. Distribution. India: Uttar Pradesh, Kumaon. Remarks. The specimens of Scaphisoma cf. minutissimum recorded from Thailand (L6BL 1990b) belong to another, undescribed species. They may be distinguished from S. minutissimum by the darker body, the coarser elytral punctation and by the different shape of the aedeagus. Scaphisoma tonkineum Pic Scaphosoma tonkineum Pic, 1922: 1; LOBL 1976b: 224. Diagnostic characters. Length 2.0 - 2.1 mm, width 1.43 - 1.48 mm. Body very dark reddish-brown to blackish, with apex of elytra and apical abdominal segments paler. Antennae long, relative length of segments as follows: III 5, IV 13, V 20, VI 28, VII 31, VII 23, IX 28, X 29, XI 35 (lectotype). Pronotum densely and rather coarsely punctate, punctures barely visible at 12x magnification. Tip of scutellum distinct. Elytron with evenly arcuate lateral margin; inner apical angle exceeding level of outer angle; sutural margin not raised; sutural area flat; sutural stria parallel to sutural margin, curved along base and extended beyond middle of basal margin; discal punctation dense, coarser than that of pronotum and distinct at 12x magnification. Mesepimeral ridge somewhat shorter than interval between its end and mesocoxa. Metasternum without microsculpture, punctation dense and coarse on central portion, gradually sparser and much finer laterally. Mesocoxal area 0.08 - 0.10 mm long, with arcuate, coarsely punctate margin. Metepisternum narrowed anteriad. Abdomen with distinct microsculpture consisting of transverse striae. First ventrite very finely punctate. Tibia straight. Male protarsus with enlarged segments 1 to 3. Aedeagus 0.50 - 0.56 mm long (see LOBL 1976b), with median lobe gradually tapering and strongly curved at tip. Parameres slender. Type material. Lectotype, male, labelled “Hoa Binh Tonkin” (MNHN), by present designation. Material examined, 2 specimens. India, Uttar Pradesh, Kumaon, Haldwani distr. (Champion) (BMNH) 1; Nepal, Pokhara - Ghoropani, Sept.-Oct.1977 (Franz) (Coll. H .Franz) 1. Distribution. Vietnam, North India, Nepal. Scaphisoma kashmirense Achard Scaphosoma kashmirense ACHARD, 1920d: 240. Scaphosoma ebeninum CHAMPION, 1927: 278 - syn. n. Scaphisoma kashmirense; LOBL 1970: 764, 1986a: 156, 1986b: 345. Diagnostic characters. Length 2.0 - 2.2 mm, width 1.36 - 1.47 mm. Body blackish, elytra paler than pronotum, apex of abdomen, femora and tibiae reddish- brown, antennae and tarsi ochreous. Relative lengths of antennomeres as follows: III 8, IV 14, V 18, VI 20, VII 27, VIII 19, IX 25, X 24, XI 29. Pronotal punctation sparse and very fine, barely visible at 12x magnification, punctures well delimited, much smaller than intervals. Tip of scutellum exposed. Elytron with lateral keel not visible in dorsal view; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin 540 IVAN LÖBL not raised; sutural area flat anteriorly, weakly vaulted in apical half; sutural stria from apex toward middle of sutural length somewhat diverging with sutural margin, then parallel to it, extended along base to forme uninterrupted basal stria joined with lateral stria; discal punctation fine and sparse, near base similar to that on pronotum, on centre and on apical area less fine, distinct at 12x magnification. Pygidium finely punctate, with microsculpture consisting of punctures. Mesepimeral ridge about as long as interval between its end and mesocoxa. Metasternum not microsculptured, punctation dense and rather coarse on medio-apical portion, very fine and sparse laterally. Mesocoxal area 0.05-0.06 mm long, with arcuate, coarsely punctate margin. Metepisternum at same level as metasternum, strongly narrowed anteriad, suture barely sinuate. Microsculpture of 1st ventrite consisting of distinct punctures on median portion, obsolete on lateral portion. Latter very finely and sparsely punctate. Metacoxal area 0.08 - 0.10 mm long, with arcuate, coarsely punctate margin. Tibia I and II somewhat curved, III straight. Aedeagus as figured in LOBL (1986b). Type material. Lectotype of S. kashmirense, male, labelled “Kashmir” (MNHN), designated in LOBL (1970), examined. Holotype of S. ebeninum, female, labelled “Pindar V. Almora U.P., 8-11.000 ft. July 1920, H.G.C.” (BMNH), examined. Material examined, 10 specimens: India, Kashmir, forest above Pahalgam, 15.X.1977 (Franz) (Coll. H. Franz, MHNG) 2; Himachal Pradesh, Simla distr., Kufri, 24.V1.1975 (Sengupta & party) (ZSI) 1 and Kufri, 16.VIL.1989 (Riedel) (SMNS, MHNG) 6; Nepal, Patan distr., Phulcoki, 2550 m, 29.1V.1984 (Löbl & Smetana) 1. Distribution. North Pakistan, North India, Nepal. Scaphisoma cruciatum Champion Scaphosoma cruciatum CHAMPION, 1927: 275. Diagnostic characters. Length 2.0 mm, width 1.35 mm. Body ochreous, pronotal base and median portion darkened, elytral base strongly darkened, dark pattern extended apically to form triangular spot exceeding mid-length of elytron (Fig.26). Apical portion of elytron, apical abdominal segments, legs and antennae pale ochreous. Relative length of antennomeres as follows: III 8, IV 20, V 30, VI 24, VII 27, VIII 21, IX 28, X 27, XI 33. Pronotal punctation very dense and coarse, visible at 12x magnification. Tip of scutellum exposed. Elytron with lateral keel visible from base to apex in dorsal view; apical margin weakly rounded; inner apical angle at same level as outer angle; sutural margin somewhat raised apically, flat basally; sutural area finely punctate; sutural stria weakly diverging from apex toward anterior 1/5 of sutural length, then parallel to sutural margin, somewhat curved at base, not extended along basal margin; discal punctation very dense, most punctures larger than intervals, near base finer than on centre. Pygidium extremely finely punctate, with distinct microsculpture consisting of punctures. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum microsculptured along posterior margin only, densely and rather coarsely punctate on medio-apical area, very finely and sparsely punctate between mesocoxae and on lateral portion; with 2 shallow medio-apical impressions. Mesocoxal area 0.09 mm long, with inner margin arcuate, outer margin almost oblique, marginal punctures distinct. Metepisternum strongly narrowed anteriad, impressed along suture, latter straight except at angles. Abdomen with distinct microsculpture consisting of transverse striae, entire lst ventrite very finely punctate. Metacoxal area 0.10 mm long, with margin arcuate, coarsely punctate. Type material. Holotype, female, labelled “India: Kumaon Haldwani Distr. H.G.Champion B.M.1930-59” (BMNH), examined. SCAPHIDIIDAE OF NEPAL 541 Remarks. None of the male specimens of different Himalayan species could be in satisfactory way associated with the female holotype of S. cruciatum. Other species with similar colour pattern (S. bhareko, S. sikkimense) differ by their punctation. Scaphisoma championi Löbl Scaphosoma cribripenne CHAMPION, 1927: 277; nec S. cribripenne (Pıc, 1923). Scaphisoma championi LOBL, 1981: 158. Diagnostic characters. Length 1.35 - 1.55 mm, width 0.95 - 1.10 mm. Most of body and femora ochreous, pronotum usually somewhat paler than elytral disc, apical portion of elytra, hypomera, apical abdominal segments, tibiae, tarsi and antennae yellowish or pale ochreous. Antennae as in other species of the group. Pronotum with lateral margins evenly arcuate; lateral keels usually not visible in dorsal view; discal punctation fine nad rather dense, visible at 24x magnification. Tip of scutellum exposed. Elytron with lateral margin arcuate; lateral keel usually visible in dorsal view from base to apex; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin raised; sutural area flat, with a row of rather large punctures; sutural stria gradually, moderately diverging from apex to base, curved near base, not extended laterad of pronotal lobe; punctation coarse and dense on most of elytral surface, finely punctate humeral area excepted, with punctures not well delimited, usually larger than intervals. Pygidium without visible punctation. Mesepimeral ridge about 2x as long as interval between its end and mesocoxa. Median portion of metasternum flattened, without any impression, with microsculpture consisting of transverse striae, distinctly punctate except on portions between mesocoxae and between metacoxae. Lateral portion of metasternum lacking microsculpture, with very dense row of rather coarse punctures parallel to metacoxa, and moderately coarse punctures beyond mesocoxal area; most of metasternal surface extremely finely punctate. Mesocoxal area 0.06-0.07 mm long, with margin parabolic, rather finely punctate. Metepisternum large, flat, narrowed anteriad, at same level as metasternum, its suture more or less rounded. Abdomen with microsculpture consisting of transverse striae. First ventrite extremely finely and very sparsely punctate. Metacoxal area 0.07 - 0.08 mm long, with arcuate, coarsely punctate margin. Tibia straight. Segments 1 to 3 of male protarsus distinctly widened. Aedeagus (Fig. 150) 0.68 - 0.70 mm long. Median lobe with strongly inclined, slender apical portion, and short, narrow valves. Internal sac with rows of medio-apical fairly large. and strongly sclerotized teeth accompanied laterally by membranous scales. Central portion of internal sac with oval disc overlapping fine scales and with two laterally denticulate, strongly sclerotized areas. Base of internal sac armed by two slender, proximally curved sclerites, membranes near basal margin scale-like or papillar. Parameres beyond strongly widened base moderately convergent, then narrowed and strongly curved. Type material. Lectotype of S. cribripenne Champion, male, labelled “India: Kumaon, Haldwani Distr. H.G. Champion B.M.1930-59” (BMNH), by present designation. Paralectotypes: 46 specimens; additional 16 specimens, all with same locatity data as lectotype (BMNG, MHNG). Distribution. India, Uttar Pradesh: Kumaon. Scaphisoma bedeli Achard Scaphosoma bedeli ACHARD, 1920d: 240. Diagnostic characters. Length 1.8 mm, width 1.10 mm. Pronotum very dark, almost black. Elytra blackish-brown, with paler, dark reddish-brown apical 2/7. 542 IVAN LÖBL Apex of abdomen, legs and antennae ochreous. Relative length of antennomeres as follows: III 6, IV 10, V 16, VI 23, VII 28, VII 20, IX 26, X 23, XI 29; segment IV about 2x as long as wide, V almost 4x as long as wide, VI more than 4x as long as wide; VII 3.5x as long as wide; VIII about 3x as long as wide. Pronotum with lateral margins evenly rounded, lateral keels not visible in dorsal view; punctation sparse and very fine, visible at 24x magnification. Tip of scutellum exposed. Elytron with almost evenly arcuate lateral margin; lateral keel not visible in dorsal view; apical margin truncate; inner apical angle about at same level as outer angle; sutural area flat, with a row of very fine punctures; sutural stria parallel to suture, curved near base, not extended laterally of pronotal lobe; punctation fine to very fine, punctures shallow, not well delimited, smaller than intervals. Pygidium with microsculpture consisting of punctures, and with very fine but well delimited punctures. Mesepimeral ridge about as long as interval between its end and mesocoxa. Metasternum with two shallow medio-apical impressions, entire surface with very fine to obsolete punctation. Mesocoxal area 0.07 mm long, with margin arcuate, finely punctate. Metepisternum flat, narrowed anteriad, impressed below level of metasternum, suture almost straight, anterior angle rounded. Entire first ventrite very finely punctate, not microsculptured. Metacoxal area 0.05 mm long, with arcuate, coarsely punctate margin. Following ventrites with microsculpture consisting of points. Type material. Lectotype, female, labelled “Sikkim Kurseong” (NMP), by present designation. Distribution. India: Darjeeling district. Remarks. The specimens recorded as Scaphisoma cf. bedeli in LOBL (1986a) belong to S. varians described below. Non of the recently collected specimens have been found conspecific with the lectotype of S. bedeli. NEW SPECIES Scaphisoma praesigne sp. n. Holotype, male: Nepal, Kathmandu distr., Gokarna forest, 1400 m, 1.IV.1984 (Löbl & Smetana) (MHNG). Diagnosis. Body moderately large, without particular colour pattern. Antennomere IV very short. Basal stria of elytron short. Abdominal microsculpture consisting of punctures. Median lobe of aedeagus asymmetrical, internal sac simple, with extruded flagellum, parameres symmetrical, strongly expanded basally. Description. Length 1.65 mm, width 1.10 mm. Body uniformly reddish- brown, tarsi and antennomeres I to VI paler, antennal club brownish. Relative length of antennomeres as follows: III 4.5, IV 5, V 20, VI 18, VII 22, VII 15, IX 22, X 20, XI 23; IV very small, V about 3.5x as long as wide; VI and VII each about 2.5x as long as wide; VII 2x as long as wide; XI somewhat more than 2x as long as wide. Pronotum with arcuate lateral margins; lateral keels not visible in dorsal view; discal punctation very fine, distinct at 24x magnification. Tip of scutellum exposed. Elytron with arcuate lateral margin; lateral keel not visible in dorsal view; apical margin somewhat rounded; inner apical angle lying at same level as outer angle; sutural margin not raised; sutural stria parallel to suture, curved at base and extended laterally to middle of basal margin; sutural area flat, with row of very fine punctures; discal puctation very fine, expecially on | humeral area. Pygidium extremely finely punctate, apparently without microsculpture (100x magnification). Mesepimeral ridge about as long as intervals between its end and SCAPHIDIIDAE OF-NEPAL 543 mesocoxa. Entire metasternum very finely and sparsely punctate, not microsculptured. Mesocoxal area 0.05 mm long, with arcuate, very finely punctate margin. Metepisternum flat, narrowed anteriad, suture straight except at anterior angle. Entire first ventrite without microsculpture, very finely and sparsely punctate. Metacoxal area 0.07 mm long, with arcuate, finely punctate margin. Following ventrites with microsculpture consisting of punctures. Pro- and metatibia straight, mesotibia weakly curved. Segments 1 to 3 of protarsus weakly widened. Aedeagus (Figs 127, 128) 0.54 mm long. Median lobe and parameres asymmetrical. Apical portion of median lobe gradually narrowed and with curved tip and membranous lateral margin; ventral process protruding from expanded ventral wall. Internal sac simple, with long flagellum. Parameres moderately arcuate, with widened base, and very weakly sclerotized central portion of inner margin. Remarks. This species shares an asymmetrical aedeagus with the members of the unicolor group. It may be readily distinguished from all of them by the very short antennomere IV. Scaphisoma fulcratum sp. n. Holotype, male: Nepal, Patan distr., 2 km S Godawari, 1700 m, 19.X.1983 (Löbl & Smetana) (MHNG). Paratype, male: India, Himachal Pradesh, 10 km NW Sarahan, 1700 m, 7.X.1988 (Vit) (MHNG). Diagnosis. Body moderately large, without distinctive colour pattern. Antennomere IV elongate. Elytron without basal stria. Abdominal microsculpture consisting of striae. Aedeagus symmetrical, median lobe with unpaired dorsal valve and small ventral process, internal sac bearing single slender median sclerite. Description. Length 1.40 - 1.65 mm, width 1.08 - 1.13 mm. Body very dark reddish-brown, apices of elytra, apical abdominal segments, legs and antennae ochreous or yellowish. Relative length of antennomeres as follows: III 6, IV 12, V 16, VI 17, VII 22, VII 15, IX 22, X 22, XI 35; segments HI to VII and XI each about 3x as long as wide; VIII less than 3x as long as wide. Pronotum with evenly arcuate lateral margins; lateral keels not visible in dorsal view; punctation conspicuously dense, coarser near base than that on centre or near apex; punctures near base larger than intervals, those on centre smaller than intervals, visible at 24x magnification. Tip of scutellum exposed. Elytron with arcuate lateral and apical margins; lateral keel not visible in dorsal view; inner apical angle at same level as outer angle; sutural margin not raised; sutural area flat, with row of fine punctures; sutural stria parallel to suture, not curved near base; punctation dense and, except that on humeral area, somewhat more coarse than that on pronotal base, punctures well delimited, mostly smaller than intervals. Mesepimeral ridge somewhat shorter than interval between its end and mesocoxa. Metasternum without microsculpture, lacking depressions; metasternal punctation very dense, relatively coarse, punctures on apical portion larger than intervals. Mesocoxal area 0.05 mm long, margin arcuate, rather finely punctate. Metepisternum weakly convex, narrowed anteriad, impressed along weakly sinuate suture, latter accompanied by coarse punctures. Abdomen with microsculpture consisting of transverse striae. First ventrite coarsely punctate near base, finely punctate near apical margin. Metacoxal area 0.06 mm long, with arcuate, rather finely punctate margin. Tibia straight. Segments | to 3 of male protarsus moderately widened. Aedeagus (Fig.129) 0.38 - 0.55 mm long. Median lobe tapering apically, with single dorsal valve and ostium situated near apex. Internal sac with slender central stick-like sclerite; membranes 544 IVAN LÖBL covered by fine scales. Parameres straight, narrow, moderately narrowed from base toward _ middle third, then evenly wide, not exceeding tip of median lobe. Remarks. Species of uncertain relationship. It may be recognised by the combination of both, external and aedeagal characters. Scaphisoma interjectum sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley, Chichile, forest above Ahale, 2300 m, 26.III.1982 (A. & Z. Smetana) (MHNG). Paratypes, 6: Sankhua Sabha distr., Arun Valley, betw. Mure and Hurure, 2050-2150 m, 17.V1.1988 (Martens & Schawaller) (SMNS, MHNG) 1 male, 1 female; Patan distr., Phulcoki, 2550 m, 15.X.1983 (Löbl & Smetana) (MHNG) 1 male; Phulcoki, 6600 ft., 13.VIII.1967 (Canadian Nepal Exp.) (CNC) 1 female; Manang distr., forest W Bagarchhap, 2200 m, 24.1X.1983 (Löbl & Smetana) (MHNG) 1 male, 1 female. Diagnosis. Body rather small, uniformly brown. Antennomere IV elongate. Elytron with interrupted basal stria. Abdominal microsculpture consisting of punctures. Aedeagus symmetrical, median lobe with short, split dorsal valves, internal sac without flagellum or sclerotized pieces, parameres lobed. Description. Length 1.45 mm, width 1.03-1.08 mm. Body and femora uniformly brown. Apex of abdomen, tibiae, tarsi and antennae yellowish or ochreous. Relative length of antennomeres as follows: III 6, IV 12, V 16, VI 18, VII 24, VIII 18, IX 20, X 19, XI 26 (holotype); IV and VIII 3x as long as wide, VI, VII, and XI each somewhat more than 3x as long as wide. Pronotum with lateral keels not visible in dorsal view; discal punctation very fine, visible at 24x magnification. Tip of scutellum exposed. Elytron with lateral keel visible near base in dorsal view; apical margin weakly rounded; inner apical angle at same level as outer angle; sutural margin raised; sutural area flat or weakly oblique, with row of very fine punctures; sutural stria parallel to sutural margin, curved at base and extended along base to outer 1/3 of basal margin; discal punctation coarse and dense except near base, with many punctures larger than intervals; punctures near base fine and scattered, much smaller than intervals, coarser than those on pronotum. Pygidium extremely finely punctate, with barely visible microsculpture. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum without micro- sculpture, apical depressions obsolete, punctation fine on medio-apical portion, very fine and sparse on lateral portion. Mesocoxal area 0.04 mm long, with arcuate, rather finely and sparsely punctate margin. Metepisternum flat, moderately narrowed anteriad, at same level as metasternum, suture straight, except for rounded anterior angle. First ventrite without microsculture, its punctation very fine laterally, much coarser on median portion. Metacoxal area 0.06 mm long, subtriangular, margin coarsely punctate. Following ventrites with microsculpture consisting of distinct punctures. Tibia slender, straight. Male protarsus with segments 1 to 3 enlarged. Aedeagus (Fig. 130) 0.62 - 0.67 mm long. Median lobe relatively large, with very short apical portion and weakly sclerotized lateral margins, dorsal valves symmetrical, distinctly split medially. Internal sac mostly membranous, bearing very small rounded or pointed scales; in addition with several slender, more or less well delimited, stronger sclerotized pieces. Parameres widened in basal half, each expanded and forming a fairly large inner lobe before middle, slender and very weakly curved in apical half. Remarks. This species shares most diagnostic characters with S. rufum. It may be distinguished by the shorter and wider apical portion of the median lobe, by the parameral lobes situated closer to the base, and by the elytral punctation. SCAPHIDIIDAE OF NEPAL 545 Scaphisoma simplicipenis sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, 2550 m, 29.1V.1984 (Löbl & Smetana) (MHNG). Paratypes, 114: as holotype, 16 males, 28 females; Phulcoki, 1700 m, 10.V.1981 (Löbl) (MHNG) 3 males; Phulcoki, 2500 m, 30.1V.1984 (Löbl & Smetana) (MHNG) 1 male, 6 females; Phulcoki, 2600 and 2700 m, 14. and 15.X.1983 (Löbl & Smetana) (MHNG) 1 male, 1 female; Kathmandu distr., Siwapuri Dara, 2500 m, 1.V.1985 (Smetana) (MHNG) 1 male, 2 females; Sindhupalcok distr., Pokhare NE Barahbise, 2800 m, 2. and 3.V.1981 (Löbl & Smetana) (MHNG) 1 male, 2 females; Parbat distr., Ghoropani Pass, 2850 m, 5.X.1983 and N slope of Ghoropani Pass, 2700 m, 6.X.1985 (Löbl & Smetana) (MHNG) 2 males; Manang distr., forest W Bagarchhap, 2200 m, 21.IX.1983 (Löbl & Smetana) (MHNG) 11 males, 4 females; Latha Manang W Bagarchhap, 2450 m, 23.IX. 1983 (Löbl & Smetana) (MHNG) 1 male, 1 female; Sankhua Sabha distr., forest NE Kuwapani, 2250 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 1 male; Chichila, 2200 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 2 females; Pahakhola, 2550 m, 30-31.V.1988 (Martens & Schawaller) (SMNS, MHNG) 11 males, 18 females; Taplejung distr., above Yamputhin, left bank of Kabeli Khola, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS) 1 male. Diagnosis. Body moderately large, without distinctive colour pattern. Anten- nomere IV elongate. Elytron without basal stria. Abdominal microsculpture consisting of striae. Aedeagus symmetrical, with small basal bulb, robust ventral process, simple internal sac bearing long flagellum, and bases of parameres strongly widened. Description. Length 1.55 - 1.70 mm, width 1.03 - 1.16 mm. Body very dark brown to blackich, elytra sometimes paler than pronotum. Apical abdominal segments reddish-brown or brown, legs and antennae ochreous. Relative length of antennomeres as follows: III 6, IV 10, V 16, VI 23, VII 28, VIII 20, IX 26, X 25, XI 28 (holotype). Segment IV about 2.5x as long as wide, V, VIII and XI each 3x as long as wide, VI 4x as long as wide, VII about 3.5x as long as wide. Pronotum with evenly arcuate lateral margins, lateral keels : not visible in dorsal view; discal punctation sparse and very fine, visible at 24x magnifi- cation. Tip of scutellum exposed. Elytron with central portion of lateral margin oblique; lateral keel not visible, or visible near base in dorsal view; apical margin weakly convex; inner apical angle at same level as outer angle; sutural margin not raised; sutural stria parallel to suture, curved near base but not extended laterally; discal punctation sparse and very fine, similar to that of pronotum or somewhat more distinct. Pygidium with microsculpture consisting of transverse striae. Mesepimeral ridge shorter than interval between its end and mesocoxa. Metasternum with shallow medio-apical impression, entirely very finely punctate, without microsculpture. Mesocoxal area 0.04 mm long, with margin arcuate, finely punctate. Metepisternum narrowed anteriad. First ventrite without microsculpture, very finely punctate. Metacoxal area 0.05 mm long, with rounded, finely punctate margin. Following ventrites with microsculpture consisting of short transverse striae. Tibia straight. Segments 1 to 3 of male protarsus rather strongly enlarged. Aedeagus (Figs 131,132) 0.56 - 0.57 mm long. Median lobe with small basal bulb extended ventro-apically, to forme strong ventral process, and with very long, slender apical portion, with margins parallel from base toward middle, than tapering. Internal sac simple, with long flagellum accompanied laterally by extremely fine, pointed scales. Parameres strongly sclerotized, reaching level of median lobe, at base strongly widened, in lateral view arcuate, in dorsal view irregularly curved. Remarks. This species shares with the species of the unicolor group the median lobe of the aedeagus with small basal bulb with very robust ventral process, and simple internal sac with long flagellum. It differs from members of that group by the symmetrical and strongly sclerotized aedeagus. Scaphisoma fatuum sp. n. Holotype, male: Nepal, Myandi distr., Kali Gandaki Khola, Kopchepani, 1500-1700 m, 15.V.1984 (Holzschuh) (MHNG). 546 IVAN LÖBL Paratypes, 3: as holotype, 2 males, 1 female (MHNG). ; Diagnosis. Large-sized member of the subalpinum group. Antennomere IV fairly long, XI strongly elongate. Basal stria of elytron interrupted. Aedeagus with simple flagellum conspicuously narrowed and incurved apically. Description. Length 2.4 - 2.5 mm, width 1.71 - 1.75 mm. Body black, apex of abdomen, legs and antennae ochreous, except brown basal antennomeres. Antennae long, relative length of segments as follows: III 10, IV 15, V 19, VI 29, VII 34, VII 25, IX 32, X 33, XI 50 (holotype); segments IV, V, VII, VII, and IX each about 3x as long as wide, VI about 4x as long as wide, XI nearly 5x as long as wide. Pronotum with lateral keel not visible in dorsal view, punctation very fine, visible at 24x magnification. Tip of scutellum exposed. Elytron with arcuate lateral margin; lateral keel visible in dorsal view only near base; apical margin rounded; inner apical angle at same level as outer angle; sutural margin not raised, sutural area flat, with a row of fine punctures; sutural stria parallel to sutural margin, curved at base and extended along basal margin toward outer third of basal width; punctation dense and rather coarse, punctures rather well delimited, definitively smaller than intervals. Pygidium very finely punctate, with microsculpture consisting of punctures. Mesepimeral ridge somewhat longer than interval between its end and mesocoxa. Metasternum without microsculpture and without depressions; middle portion flat, rather coarsely and densely punctate apically, elsewhere very finely and sparsely punctate. Mesocoxal area 0.06 mm long, its margin arcuate, rather coarsely punctate. Metepisternum somewhat vaulted, strongly narrowed anteriad, its margin straight or weakly sinuate, impressed below level of metasternum. Abdomen with distinct microsculpture formed by punctures. First ventrite very finely punctate laterally, some- what less finely punctate on middle portion. Metacoxal area 0.10 mm long, with arcuate, coarsely punctate margin. Tibia straight. Male with segments 1 to 3 of protarsus strongly enlarged, narrower than apex of protibia. Segment 1 of male mesotarsus weakly enlarged. Aedeagus (Fig. 133) 1.06 - 1.10 mm. Median lobe with apical portion fairly long, gradually tapering, very slender tip. Internal sac with very long, slender flagellum incurved and narrowed apically. Parameres narrowed from base toward middle, then evenly slender and weakly curved. Remarks. This species is a member of the subalpinum group. It resembles S. antennatum by the conspicuously long antennomere XI. However, it differs in having much longer antennomere IV. Scaphisoma fatuum may be readily distinguished from other members of the group by the large size of the body. Scaphisoma adjacens sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley, Chichila, 2200 m, 24.IV.1984 (Löbl & Smetana) (MHNG). Paratype, male: Nepal, Taplejung distr., above Yamputhin, left bank of Kabeli Khola, open forest, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS). Diagnosis. Member of the subalpinum group. Body fairly large. Antennomere IV very short, XI moderately long. Elytron with basal stria interrupted. Aedeagus with basal two thirds of flagellum extended laterally, and ending as very thin tube; parameres almost evenly narrow. Description. Length 1.85 - 1.90 mm, width 1.23 - 1.27 mm. Body very dark reddish-brown, apex of abdomen, femora and tibiae paler. Tarsi ochreous. Antennomeres I | to IV yellowish, following pale brown. Relative length of antennomeres as follows: III 6, IV 7, V 16, VI 17, VII 25, VIII 18, IX 23, X 22, XI 26 (holotype); segment IV only | | SCAPHIDIIDAE OF NEPAL 547 somewhat longer than wide, V, VI and VII each about 2.5x as long as wide, VIII somewhat more than 2x as long as wide, XI 2x as long as wide.Pronotum with lateral keels not visible in dorsal view; discal punctation rather sparse and fine, visible at 24x magnification. Tip of scutellum exposed. Elytron with evenly arcuate lateral margin; lateral keel not visible in dorsal view; apical margin truncate; inner apical angle exceeding level of outer angle, sutural margin not raised; sutural area flat, with a row of very fine punctures; sutural stria parallel to suture, curved at base and extended laterally, forming shallow basal stria interrupted in humeral area; discal punctation very fine near elytral base, elsewhere about as fine as or coarser than that of pronotum, with punctures well delimited, much smaller than intervals. Propygidium relatively coarsely punctate, pygidium much finer punctate than latter, both without distinct microsculpture (100x). Mesepimeral ridge shorter than interval between its end and mesocoxa. Metasternum without impressions, very finely and sparsely punctate, coarsely punctate on medio-apical portion. Mesocoxal area 0.06 mm long, margin arcuate, densely punctate. Metepisternum strongly narrowed anteriad, with straight margin. First ventrite without microsculpture, very finely punctate. Metacoxal area 0.09 mm long, with arcuate, rather coarsely punctate margin. Tibia moderately curved. Segments 1 to 3 of male protarsus weakly enlarged. Aedeagus (Fig. 134) 0.59 - 0.65 mm long. Median lobe with apical portion fairly wide, moderately long, at tip rounded; ventral process relatively large. Internal sac armed with robust flagellum; flagellum rather slender in apical third and with abruptly narrowed, very thin, transversaly bent end, in basal two thirds extended laterally by asymmetrical, more or less strongly sclerotized lamina. Parameres almost evenly slender from apex to moderately widened base, weakly curved apically. Remarks. This species may be readily distinguished from other members of the subalpinum group by the relative length of the antennomeres, especially by the very short segment IV in combination with weakly elongate segment XI. Scaphisoma inquietum sp. n. Holotype, male: Nepal, Parbat distr., Ghoropani Pass, N slope, 2700 m, 6.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 5: as holotype, 3 females; Mustang distr., 2 km N Kalopani, 2550 m, 1.X.1983 (Löbl & Smetana) (MHNG) 1 female; Jumla distr., Dzuga Khola Valley near Talphi, W Jumla, 3000-3500 m, 19.IX.1972 (Franz) (Coll. H. Franz) 1 male. Diagnosis. Member of the subalpinum group similar to S. adjacens. Median lobe of aedeagus gradually narrowed, flagellum widened basally and forming hook-shaped apophysis, parameres slender beyond middle. Description. Length 1.95 - 2.05 mm, width 1.32 - 1.37 mm. Very similar to S. adjacens and S. uniforme, differs from both in finer elytral and pronotal punctation. Latter almost indistinct at 24x magnification. Relative length of antennomeres as follows: II 6, IV 7, V 15, VI 21, VII 30, VII 19, IX 26, X 25, XI 36 (holotype); IV conspicuously short, less than 2x as long as wide; V about 2.5x as long as wide; VI, VII, and XI each about 3x as long as wide; VIII as wide as VI, less than 3x as long as wide. Apical angle of elytron at same level as outer angle. Pygidium very finely punctate, propygidium with moderately fine punctures. Mesepimeral ridge somewhat shorter than interval between its end and mesocoxa. Metasternum with shallow medio-apical impressions, very finely and sparsely punctate, except for more coarsely punctate medio-apical area. Mesocoxal area 0.10 mm long, margin arcuate, finely punctate. Abdomen with microsculpture consisting of punctures. First ventrite very finely punctate. Metacoxal area 0.05 - 0.06 mm long, 548 IVAN LÖBL inner margin arcuate, outer margin oblique. Tibia weakly curved. Segments 1 to 3 of male _ protarsus weakly widened. Aedeagus (Fig. 135) 0.57 - 0.62 mm long. Median lobe with basal bulb gradually narrowed; apical portion of median lobe almost indistinct in dorsal view, with very short, rather wide tip. Flagellum with wide base asymmetrically extended by a hook-like process, from base to about middle third gradually narrowed, in apical third very slender. Membranes of internal sac covered by distinct scales forming characteristic pattern. Parameres weakly sinuate, wide in basal third, then narrowed, in apical third notably narrower than near base. Remarks. This species may be readily distinguished from other members of the subalpinum group by its aedeagal characters. It also differs from most species by the conspicuously short antennomere IV. Scaphisoma fratellum sp. n. Holotype, male: Nepal, Taplejung distr., above Yamputhin, left bank of Kabeli Khola, bushes, open forest, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS). Paratypes, 3: as holotype, 1 male, 2 females (SMNS, MHNG). Diagnosis. Body moderately large, without particular colour pattern. Antennomere IV very small. Elytron without basal stria. Abdominal microsculpture consisting of striae. Aedeagus symmetrical, with dorsal valve split, abruptly inclined apically and extended by short lamina. Internal sac complex, with robust medio-apical sclerite. Description. Length 1.55 - 1.75 mm, width 1.16 - 1.23 mm. Body black, apex of elytra yellowish or ochreous, apical abdominal segments and legs ochreous, antennae yellowish to pale brown. Relative length of antennommeres as follows: II 5, IV 6, V 11, VI 24, VII 28, VII 15, IX 22, X 22, XI 42 (holotype); segment IV somewhat longer than wide; V about 2x as long as wide, VI and VII each about 4x as long as wide, VIII about 2.5x as long as wide, XI well 4x as long as wide. Pronotum with evenly arcuate lateral margins, lateral keels not visible in dorsal view; punctuation fine and very dense, punctures well delimited, near base somewhat larger than those on centre and near apex, barely visible at 12x magnification. Tip of scutellum exposed. Elytron with lateral margin oblique in middle; lateral keel visible near base only in dorsal view; apical margin arcuate; inner apical angle beyond level of outer angle; sutural angle not raised; sutural area flat, with a row of very fine punctures; sutural stria parallel to suture, not or barely curved near base; discal punctation very dense and rather coarse, most punctures well delimited, about as large as intervals; humeral area finely punctate. Pygidium with obsolete punctation. Mesepimeral ridge as long as or longer than interval between with end and mesocoxa. Metasternum without microsculpture, its median portion flattened apically, densely and finely punctate, punctation becoming gradually finer and sparser laterad. Mesocoxal area 0.05-0.06 mm long, margin arcuate and finely punctate. Metepisternum narrowed anteriad, at same level as metasternum, its suture rounded or oblique in middle. Abdominal microsculpture consisting of distinct transverse striae. First ventrite very finely, sparsely punctate. Metacoxal area 0.10 - 0.11 mm long, margin strongly arcuate, finely punctate. Tibia straight. Segments 1 to 3 of male protarsus strongly enlarged, 1 somewhat narrower than apex of protibia. Aedeagus (Fig. 136) 0.73 mm long. Median lobe with split valves, latter strongly inclined and sclerotized apically, and extended by horizontal, short, notched lobe covering ostium. Ventral process fairly large. Internal sac with robust, long medio- apical sclerite; small central membranous portion bearing short scales; proximal portion | SCAPHIDIIDAE OF NEPAL 549 long, bent basally, with more or less sclerotized long scales or denticle-like structures. Parameres straight from base to widened, incurved apical portion. Remarks. Scaphisoma fratellum is very similar to S. fraterculum Löbl in external characters but may be readily distinguished by the much longer antennae. Both species have very distinct aedeagi, that of S. fratellum has the ostium situated apically and covered by a dorsal lamina, the parameres simple, and the internal sac complex, in a similar way as in S. aurorae. Scaphisoma aurorae sp. n. Holotype, male: Nepal, Parbat distr., Pun Hill at Ghoropani Pass, 3050-3100 m, 8.X.83 (Löbl & Smetana) (MHNG). Paratypes, 31: Parbat distr., as holotype, 1 female; Ridge E Ghoropani Pass, 3100 m, 7.X.1983 (Löbl & Smetana) (MHNG) 2 males, 10 females; Ghoropani Pass, N slope, 2750 m, 5.X.1983 (Löbl & Smetana) (MHNG) 3 males; Ghoropani Pass, 2850 m, 5.X.1983 (Löbl & Smetana) (MHNG) 1 male; Manang distr., Pisang, 3000 m, 3.X.1977 (Deharveng) (MHNG) 1 male; Ghorka distr., Chuling Khola, Djinski Kharka, 3400 m, 4.-5.VIII.1983 (Martens & Schawaller) (SMNS) 2 females; Myandi distr., S Dhaulagiri, Dhorpatan, 3000-3200 m, 16-24.V.1973 (Martens) (SMNS) 1 female; Sindhupalcok distr., between Ghopte and Thare Pati, 3250 m, 23.IV.1985 (Smetana) (MHNG) 1 male; above Shermathang, 2900 m, 26.1V.1981 (Löbl & Smetana) (MHNG) 1 female; Malemchi, 2900 m, 14.IV.1981 (Löbl & Smetana) (MHNG) 1 female; Patan distr., Phulcoki, 2600 m, 13.X.1983 (Löbl & Smetana) (MHNG) 1 female; Sankhua Sabha distr., between Romri La and Pahakhola, 3600-3450 m, 30.V.1988 (Martens & Schawaller) (MHNG) 1 male; Taplejung distr., Simbua Khola Valley, 3100-2900 m, 15.V.1988 (Martens & Schawaller) (SMNS) 1 female; same but near Tseram, 3250-3350 m, 10-15.V.1988 (SMNS, MHNG) 3 males, 1 female. Diagnosis. Body moderately large, with distinctive colour pattern. Antennomere IV slender. Basal stria of elytron complete. Abdominal microsculpture consisting of punctures and of very short striae. Median lobe of aedeagus abruptly inclined apically, with sclerotized valve. Internal sac bearing median sclerites. Parameres slender. Description. Length 1.65 - 1.85 mm, width 1.10 - 1.30 mm. Head and pronotum more or less dark reddish-brown or ochreous. Elytron (Fig. 27) yellowish, with blackish-brown or black base, and in most specimens with two small elongate dark central spots or with larger single dark central spot. Pygidium and propygidium with dark base and pale apices. Hypomeron and legs dark reddish-brown. Thorax and abdomen dark brown to black ventrally, yellowish apex of abdomen excepted. Antennae and tarsi yellowish. Relative length of antennomeres as follows: III 9, IV 15, V 20, VI 21, VII 26, VII 23, IX 25, X 26, XI 33 (holotype); segments IV, V and VI each about 4x as long as wide; VII and XI each 3x as long as wide, VIII about 3.5x as long as wide. Pronotum with lateral margins arcuate, lateral keel not visible in dorsal view; punctation dense, fine and shallow, barely visible at 24x magnification. Tip of scutellum exposed. Elytron with oblique central portion of lateral margin, lateral keel in dorsal view visible only near base; apical margin truncate; inner apical angle at same level as outer angle; sutural margin not raised; sutural area raised, except anteriorly, with row of very fine punctures; sutural stria deep, moderately converging beyond middle, parallel to suture before middle, curved at base and extended laterally to forme uninterrupted basal stria joined with lateral stria; punctation almost evanescent near base, beyond basal third very dense, formed by rather large but very shallow punctures, usually larger than intervals. Pygidium very finely punctate, with microsculpture consisting of punctures and of very short striae. Mesepimeral ridge as long or somewhat longer than interval between its end and mesocoxa. Metasternum not microsculptured, with two median impressions, very finely 550 IVAN LÖBL and sparsely punctate except for more distinctly punctate medio-apical portion. Mesocoxal area 0.05 mm long, its margin subtriangular or arcuate, rather coarsely punctate. Metepisternum strongly narrowed anteriad, with deep straight suture. First ventrite without microsculpture, with punctation sparse, very fine laterally, rather distinct on medio-basal portion. Metacoxal area 0.06-0.07 mm long, margin arcuate, rather coarsely punctate. Following ventrites with microsculpture consisting of punctures. Tibia straight. Segments 1 to 3 of male protarsus strongly enlarged, almost as wide as protibia. Segments 1 to 3 of male mesotarsus moderately enlarged. Aedeagus (Figs 137, 138) 0.78 mm. Median lobe with short, strongly inclined and tapering apical portion, tip visible only in lateral view, apex in dorsal view seemingly truncate. Internal sac long, complex, with two parallel median sclerites. Parameres arcuate, narrowed beyond basal third in dorsal view, evenly slender and almost straight in lateral view. Remarks. This species may be readily recognized by its colour pattern in combination with the complete basal stria of elytron. It is widely distributed for a species occuring only at high altitude. Scaphisoma nima sp. n. Holotype, male: Nepal, Mustang distr., Kalopani, 2550 m, 1.X.1983 (Löbl & Smetana) (MHNG). Paratype, male: as holotype (MHNG). Diagnosis. Body moderately large, with distinct colour pattern. Antennomere IV short. Basal stria of elytron interrupted. Abdominal microsculpture consisting of striae. Aedeagus symmetrical, with small ventral process; internal sac lacking flagellum or robust sclerites; parameres slender, almost straight. Description. Length 1.65 - 1.75 mm, width 1.16 - 1.18 mm. Head, prothorax, femora and tibiae reddish-brown. Elytra ochreous with black basal spot extended on humeral area, and darkened along apical margins. Tarsi, antennae and apical abdominal segments ochreous. Venter of thorax and basal abdominal ventrites dark brown or blackish. Relative length of antennomeres as follows: III 5, IV 7, V 19, VI 24, VII 31, VII 26, IX 28, X 27, XI 28 (holotype); segment IV small, about 2x as long as wide; V to VII each about 3.5x as long as wide; VIII almost 4x as long as wide. Pronotum with arcuate lateral margins; lateral keels not visible in dorsal view; punctation dense and very fine, distinct at 24x magnification. Tip of scutellum exposed. Elytron with very weakly arcuate lateral margin; lateral keel not visible in dorsal view; apical margin somewhat rounded; inner apical angle at same level as outer angle; sutural margin not raised; sutural area raised, very finely punctate; sutural stria weakly convergent from middle or from anterior third of sutural length toward apex, curved at base and extended toward humeral area, forming incomplete basal stria; discal punctation almost obsolete on basal fourth, from there gradually becoming denser and coarser, punctures in apical half large but shallow, larger than intervals. Pygidium extremely finely punctate, with microsculpture formed by transverse striae. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum with two shallow medio-apical impressions, on median area finely and more or less densely punctate, laterally extremely finely and sparsely punctate; without microsculpture. Mesocoxal area 0.04 mm long, with arcuate, rather finely punctate margin. Metepisternum flat, barelly narrowed anteriad, its suture straight except at angles. Abdominal microsculpture consisting of distinct transverse striae absent from latero-basal portion of Ist ventrite. Punctation of 1st ventrite fine or very fine and sparse. Metacoxall | Ä / f SCAPHIDIIDAE OF NEPAL 551 area 0.05 mm long, subparallel, marginal punctures rather coarse. Tibia straight. Segments 1 to 3 of pro- and mesotarsus moderately widened. Aedeagus (Fig. 139) 0.46 - 0.48 mm long. Apical portion of median lobe long, evenly wide, with rounded tip; dorsal valve situated near tip, weakly sclerotized. Basal half of internal sac covered by membranous scales, apical half with long spiny structures; sclerites absent except for thin straight median piece. Parameres almost straight, somewhat convergening, exceeding weakly tip of median lobe. Remarks. This species has the aedeagus similar to that of S. notatum, except for the basal portion of the internal sac which differs significantly. Both species differ drastically by their colour pattern. Scaphisoma jado sp. n. Holotype, male: Nepal, Sindhupalcok distr., Malemchi, 2800 m, 18.1V.1981 (Löbl & Smetana) (MHNG). Paratypes, 15: Sindhulapcok distr., as holotype, 3 females; above Shermathang, 2900 m, 26.1V.1981 (Löbl & Smetana) (MHNG) 1 male, 3 females; below Thare Pati, 3300 m, 10.1V.1981 (Löbl & Smetana) (MHNG) 1 male, 2 females; Nuwakot distr., Trisuli Valley, Gosaikund, 3200 m, 23-26.1V.1973 (Martens) (SMNS, MHNG) 3 males, 2 females. Diagnosis. Body rather large, with conspicuous colour pattern. Antennomere IV slender. Elytron with complete basal stria. Abdominal microsculpture consisting of punctures. Aedeagus symmetrical, dorsal valve of median lobe deeply notched. Internal sac bearing robust sclerite expanded laterally by an apophysis and extended proximally by two slender arms. Parameres with strongly expanded base. Description. Length 1.80 - 2.10 mm, width 1.21 - 1.38 mm. Most of dorsal surface ochreous. Pronotum with large dark discal spot divided in two spots in some specimens. Elytron with large dark brown or black discal spot extended to sutural margin, and darkened along basal margin. Hypomeron ochreous. Thorax ventrally and abdominal ventrites 1 to 3 or 4 dark brown. Legs reddish-brown or ochreous, antennae ochreous to yellowish. Relative length of antennomeres as follows: IH 10, IV 16, V 20, VI 20, VII 27, VIH 20, IX 25, X 25, XI 30 (holotype); segments IV to VI each about 4x as long as wide, VI and VII each about 3x as long as wide, XI not quite 3x as long as wide. Pronotum with evenly acruate lateral margins, lateral keels not visible in dorsal view; punctation fine, dense, more or less distinct at 24x magnification, punctures not well delimited. Tip of scutellum exposed. Elytron relatively strongly narrowed apically; lateral margin almost evenly arcuate; lateral keel not visible in dorsal view; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin not raised; sutural area raised beyond middle or beyond anterior third, very finely punctate; sutural stria deep, converging apically from mid-length of suture, parallel to suture in anterior half, curved at base and extended along base to forme complete basal stria joined with lateral stria; discal punctation dense and fine, with punctures decidedly larger than those of pronotum, many about as large as intervals but very shallow and indistinct on ochreous area. Pygidium very finely punctate, without visible (100x) microsculpture. Mesepimeral ridge shorter than interval between its end and mesocoxa. Metasternum without microsculpture, punctation very fine and sparse laterally, more dense and notably less fine on medio-apical portion. Mesocoxal area 0.04-0.05 mm long, margin arcuate and finely punctate. Metepisternum not or weakly narrowed anteriad, with deep straight suture. First ventrite without microsculpture, punctuation very fine and sparse, except on medio-basal area. Metacoxal area extremely narrow, parallel to coxa, its margin rather coarsely punctate. Apical 552 IVAN LÖBL abdominal segments with microsculpture consisting of punctures. Tibia straight. Segments 1 to 3 of male pro- and mesotarsus moderately enlarged. Aedeagus (Figs 140, 141) 0.63 - 0.68 mm long. Median lobe with long apical portion; dorsal valve strongly sclerotized, deeply notched medially, almost as long as ventral piece; ventral piece strongly inclined, with curved, pointed tip; ventral process well developed. Internal sac armed with robust sclerite extended basally by two narrow arms and with a hook-shaped lateral process. Parameres with conspicuously widened base, gradually narrowed toward tip in lateral view. Remarks. This species can be readily distinguished from S. notatum by the aedeagal characters. Both species live at high altitudes; S. jado is possibly restricted to Central Nepal. Scaphisoma invalidum sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley, Chichila, 2200 m, 24.IV.1984 (Löbl & Smetana) (MHNG). Diagnosis. Body moderately large, with distinct colour pattern. Antennomere IV elongate. Elytron without basal stria. Hypomeron, metasternum and abdomen with microsculpture consisting of striae. Aedeagus symmetrical, with single dorsal valve. Ventral process well developed. Internal sac bearing median sclerite and apical rows of denticles. Description. Length 1.70 mm, width 1.05 mm. Pronotum and hypomera ochreous, former with somewhat darkened centre on each side of pale median line. Elytron yellowish, with black basal spot extended apically to forme a triangular spot, and with blackish apical transverse fascia. Venter of thorax and basal abdominal ventrites dark brown. Apical abdominal segments, legs and antennomeres IV to XI yellowish. Relative length of antennomeres as follows: II 6, IV 15, V 23, VI 23, VII 26, VIII 21, IX 24, X 22, XI 32; segments IV and VI about 4x as long as wide; V almost 5x as long as wide; VII 3x as long as wide; VIII and XI each somewhat more than 3x as long as wide. Pronotum with evenly arcuate lateral margins, lateral keels not visible in dorsal view; punctation dense and very fine, visible at 12x magnification. Tip of scutellum exposed. Elytron with lateral keel visible in dorsal view only near base; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin not raised, sutural area flat, with dense row of rather coarse punctures; sutural stria parallel to suture, curved near base, not extended laterad of pronotal lobe, punctation fine and rather sparse near base, dense and rather coarse elsewhere, most punctures larger than intervals. Pygidium extremely finely punctate, with distinct microsculpture consisting of transverse striae. Hypomeron, mesepisternum, metasternum, metepisternum, and ventrites with microsculpture consisting of striae. Metasternum, except median portion, sparsely and extremely finely punctate; punctures of median portion becoming gradually larger apically. Mesocoxal area 0.04 mm long, with margin arcuate, rather finely punctate. Metepisternum vaulted, suture weakly sinuate. First ventrite sparsely and very finely punctate. Metacoxal area 0.09 mm long, large, its margin strongly arcuate, finely punctate. Protibia straight, mesotibia weakly curved, metatibia weakly sinuate. Aedeagus (Figs 142, 143) 0.57 mm long. Median lobe with slender, arcuate, apical portion pointed in lateral view; ventral process distinctly protruding apically. Internal sac slender, with simple, sinuate median sclerite, two apical groups of small sclerotized denticles, and most of membranes covered by scales. Remarks. This species may be readily distinguished by its colour pattern in combination with the microsculptured hypomeron. The aedeagus resembles that of S. SCAPHIDIIDAE OF NEPAL 553 aurorae by the median lobe having a single dorsal valve, the internal sac armed with a median sclerite and by the narrow parameres. The colour pattern is very similar to that of S. bhareko and S. sikkimense. Scaphisoma baloo sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, 2650 m, 15.X.1983 (Löbl & Smetana) (MHNG). Paratypes, 109: Nepal, as holotype, 5; same but 2700 m, 17; same but 2600 m, 14-16.X.1983, 23; same but 2550 m, 29.X.1984, 14; same but 2500 m, 30.IV.1984, 15; same but 2300 m, 10.IV.1981 (Löbl) 3; Phulcoki, 2600-2650 m, 14.V.1980 (Martens & Ausobsky) (SMNS) 1; Phulcoki, 2600 m, 20.IV.1982 (A. & Z. Smetana) (MHNG) 14; Phulcoki, 21.1.1970 (Franz) (Coll. H. Franz, MHNG) 14; Phulcoki, summit, 4.X.1971 (Franz) (Coll. H. Franz, MHNG) 3. Diagnosis. Body rather large, with distinct colour pattern. Antennomere IV long. Elytra conspicuously narrowed apically, with entire basal striae. Metacoxal area very narrow, largest near metepimeron. Aedeagus symmetrical, with single valve and large protruding ventral process. Internal sac with robust median sclerite and basal setose structures. Parameres with wide base, narrowed toward apical third. Description. Length 2.0 - 2.2 mm, width 1.38 - 1.60 mm. Body dark reddish, elytron darkened subapically and with wide yellowish apical portion (Fig. 28). Apex of abdomen yellowish, legs and antennae yellowish or ochreous. Relative legths of antennomeres as follows: II 12, IV 18, V 23, VI 24, VII 30, VII 25, IX 28, X 28, XI 30; segment III conspicuously elongate, IV, V and VI each about 5x as long as wide; VII 3x as long as wide; VIII about 3.5x as long as wide; XI less than 3x as long as wide. Pronotum with strongly arcuate lateral margins; lateral keels not visible in dorsal view, basal lobe reduced; discal punctation very fine, barely visible at 100x magnification. Scutelum completely or almost completely covered by pronotal lobe. Elytron strongly narrowed apically; lateral margin arcuate in anterior half, oblique beyond middle; apical margin truncate; inner apical angle disinctly exceeding level of outer angle; sutural margin not raised; sutural area flat, conspicuously narrow; sutural stria parallel to suture, curved at base and extended laterally, forming uninterrupted, very fine (almost obsolete) basal stria; discal punctation very fine, punctures extremely shallow, much larger than those on pronotum but usually barely visible at 50x magnification. Mesepimeral ridge indistinct. Metasternum without any impression, not microsculptured; punctation of lateral portion very fine and sparse, dense and distinct at each side of median line. Mesocoxal area 0.05 mm long, with arcuate, finely punctate margin. Metepisternum moderately narrowed anteriad, with suture rounded only at anterior angle, notably deep, widened apically. Abdomen with microsculpture consisting of punctures. First ventrite with middle portion rather densely and fairly finely punctate, lateral portion very finely and sparsely punctate. Metacoxal area very narrow, widening toward metepimeron, with weakly oblique margin. Tibiae straight. Segments 1 to 3 of male pro- and mesotarsi distinctly enlarged. Aedeagus (Figs 144, 145) 0.69 - 0.74 mm. Distal portion of median lobe long, tapering, weakly inclined, with ventral margin sinuate in lateral view; single dorsal valve relatively strongly sclerotized; ostium situated near tip. Ventral process strongly expanded apically. Internal sac with robust, strongly sclerotized piece; membranes in basal half with very fine, dense setose structures. Parameres with wide base, in dorsal view gradually narrowed toward weakly curved apical portion, in lateral view almost straight and evenly wide from tip to level of ventral process. Remarks. This species may be easily distinguished by its colour pattern in combination with the apically strongly narrowed elytra, and with the very narrow 554 IVAN LÖBL metacoxal area. The aedeagal characters suggest possible close relationship with S. ‘ aurorae and S. invalidum. Scaphisoma baloo has reduced wings and appears to be endemic to Phulcoki. Most of the specimens were found in moist plant debris along fallen oak logs. Scaphisoma clavigerum sp. n. Holotype, male: Nepal, Panchthar distr., Dhorpar Kharka, 2700 m, Rhododendron-Lithocarpus forest, 13-16.1V.1988 (Martens & Schawaller) (SMNS). Paratypes, 4: Nepal, as holotype (MHNG) 1 male; India, West Sikkim, Choka, 3050 m, nr. Yuksam, 24.IX.1983 (Sakai) (Coll. M. Sakai, MHNG) 3 females. Diagnosis. Rather large species, with distinct colour pattern. Antennomere IV slender. Basal stria of elytron uninterrupted. Abdominal microsculpture consisting of striae. Aedeagus with strongly sclerotized dorsal valve exceeding level of ventral piece of median lobe. Internal sac bearing robust median sclerite. Parameres with subapical apophysis. Description. Length 1.9 - 2.1 mm, width 1.30 - 1.45 mm. Head, most of pronotal and elytral surface, apical abdominal segments, antennae, tibiae, tarsi and most of femora ochreous or yellowish. Pronotum with two dark brown or blackish discal spots(Fig. 29). Elytral base and venter of thorax dark brown or black, pale hypomeron and metepimeron excepted. Base of femora darkened. Relative length of antennomeres as follows: III 10, IV 17, V 23, VI 30, VII 37, VIII 31, IX 38, X 35, XI 37 (holotype); segments IV, V and VII each almost 4x as long as wide, VI and VIII somewhat more than 4x as long as wide, XI almost 3.5x as long as wide. Pronotal margins arcuate, lateral keels not or barely visible in dorsal view; discal punctation dense and very fine, visible at 50x magnification. Tip of scutellum exposed. Elytron with middle portion of lateral margin oblique; lateral keel in dorsal view visible near base only; apical margin trunctate; inner apical angle exceeding level of outer angle; sutural margin raised; sutural area vaulted, with row of very fine punctures; sutural stria parallel to suture, curved near base and extended laterad to forme uninterrupted, relatively deep basal stria joined with lateral stria; punctation dense, punctures shallow, rather large, diameters mostly as large as or larger than diameters of intervals, punctures near base much smaller than intervals and extremely shallow. Pygidium with sparse and extremely fine punctation. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum not microsculptured; finely, very densely punctate on deeply impressed medio-apical portion, elsewhere sparsely and very finely punctate. Mesocoxal area very narrow, about 0.02 mm long, parallel, finely punctate margin. Metepisternum flat, at same level as metasternum, narrowed anteriad, with straight suture. Abdomen with microsculpture consisting of transverse striae. First ventrite with very fine and sparse punctation, except less finely and more densely punctate medio-basal area. Metacoxal area 0.05 - 0.06 mm long, margin finely punctate, arcuate medially, oblique laterally. Tibia straight, pro- and mesotibia stouter apically. In male, segments | to 3 of protarsus conspicuously widened, | wider, 2 as wide as, 3 somewhat narrower than apex of tibia; segment 1 and 2 of mesotarsus strongly widened, 1 as wide as, 2 narrowed than, apex of tibia, 3 weakly widened. Aedeagus (Figs 146-148) 1.08 mm long. Median lobe gradually narrowed apically, with single, relatively strongly sclerotized dorsal valve, extended to level of tip of parameres; ventral piece short, strongly inclined, with narrow apex. Internal sac armed by robust median sclerite with basal portion extended laterally; membranes bearing small scales. Parameres robust, each somewhat SCAPHIDIIDAE OF NEPAL 555 sinuate in dorsal view, arcuate in lateral view, with wide base and distinct sub-basal apophyse in ventral view. Remarks. This species may be readily distinguished by the conspicuous colour pattern in combination with the uninterrupted basal stria of the elytron and abdominal microsculpture. It is well characterized by the shape of the parameres and by the unusually long dorsal valve of the median lobe. Scaphisoma pinnigerum sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, near Godawari, 1700 m, 10.V.1981 (Löbl) (MHNG). Paratype, male: Patan distr., Phulcoki, 2350 m, X.1977 (Deharveng) (MHNG). Diagnosis. Member of the haemorrhoidale group. Body reddish-brown, elytron paler only at apical margin. Metasternum with distinct row of dense punctures parallel to metaxoca. Abdominal microsculpture consisting of striae. Internal sac of aedeagus bearing two central teeth, medio-apical tube covered by denticles and wide basal duct. Parameres with wide inner lobe. Description. Length 1.85 mm, width 1.20 mm. Body reddish-brown, elytra with very narrow pale apical area; apical abdominal segments, legs and antennae ochreous or yellowish. Antennae similar to those of other species of the group. Pronotum with lateral margins weakly arcuate; lateral keels visible in dorsal view; punctation very fine, barely visible at 12x magnification. Tip of scutellum exposed. Elytron with lateral margin almost evenly arcuate; lateral keel distinct in dorsal view; apical margin arcuate, inner apical angle exceeding level of outer angle; sutural margin barely raised; sutural area flat near base, somewhat vaulted beyond basal third, very finely punctate; sutural stria gradually, weakly diverging from sutural margin anteriad; entire discal punctation dense and coarse, most punctures larger than or as large as intervals; surface along lateral stria impunctate. Pygidium extremely finely punctate. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum without microsculpture; medio-apical portion flattened, very densely, rather coarsely punctate, punctation on centre gradually finer and sparser anteriad. Most of lateral portion of metasternum very finely and sparsely punctate; row of rather coarse punctures parallel to metacoxa very dense, accompanied by additional punctures. Mesocoxal area 0.06 mm long, margin arcuate, rather finely punctate. Metepisternum narrowed anteriad, impressed below level of metasternum, suture straight except on rounded angles. Abdomen with microsculpture consisting of distinct transverse striae. First ventrite sparsely and very finely punctate, several punctures on medio-basal portion somewhat larger. Tibia straight. Segments 1 to 3 of protarsus and segment 1 of mesotarsus rather strongly widened in male. Aedeagus (Fig. 152) 0.44 mm long. Dorsal valves of median lobe rather long and wide, rounded at tip. Internal sac armed with two conspicuous, strongly sclerotized teeth converging anteriad, their base widened and raised; membranes forming long median tube with denticles becoming larger apicad; membranous portion proximal of central teeth covered by small scale-like structures, followed by striate scructure overlapping anterior end of relatively wide basal duct. Parameres curved apically, with wide inner lobe. Remarks. This species is very similar to S. indra from which is difficult to be distinguish by its external characters. However, it may be readily recognised by the structures of the internal sac, especially by the presence of a pair of central sclerotized teeth, in combination with the wide basal duct. 556 IVAN LÖBL Scaphisoma varians sp. n. Scaphisoma cf. bedeli, LOBL 1986a: 205. Holotype, male, Nepal, Sankhua Sabha distr., forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG). Paratypes, 135: as holotype, 14 males, 12 females; Induwa Khola Valley, 2000-2100 m, 16- 17.1V.1984 (Löbl & Smetana) (MHNG) 12 males, 10 females; below Sheduwa, 2550 m, 30.11.1982 (A. & Z. Smetana) (MHNG) 1 male, 1 female, and same but 2100-2550 m, 9.IV.1982, 1 female; forest NE Kuwapani, 2600 m, 15.1V.1982 (A.& Z. Smetana) (MHNG) 5 males, 8 females; Chichila, 2200 m, 4.1V.1984 (Löbl & Smetana) (MHNG) 1 male; Chichila, 2300 m, 26.111.1982 (A. & Z. Smetana) (MHNG) 2 males; Ilam distr., Mai Pokhari, 2100-2200 m, 9-10.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 3 males, 1 female; India, Sikkim, “Rümtek”, ca 2100 m, XI.1981 (Cassagnau) (MHNG) 1 male, 1 female; West Bengal, Darjeeling distr., Algarah, 1800 m, 9.X.1978 (Besuchet & Löbl) (MHNG) 1 male; 3 km N Teesta, 250 m, 10.X.1978 (Besuchet & Löbl) (MHNG) 1 male; betw. Algarah and Labha, 7 km from Algarah, 1900 m, 11.X.1978 (Besuchet & Löbl) (MHNG) 33 males, 32 females. For additional material from the Darjeeling district see LOBL (1986a). Diagnosis. Member of the haemorrhoidale group. Body more or less dark reddish-brown, wide apical portion of elytra and apical abdominal segments paler. Metasternum with sparse row of relatively fine punctures parallel to metacoxa. Abdominal microsculpture consisting of striae. Aedeagus with short pointed dorsal valves of median lobe; Internal sac with two rows of strongly sclerotized, long denticles and with long membranous portion bearing scale-like and denticulate structures. Remarks. This species was characterized and figured in LOBL (1986a). It was hold for possibly conspecific with S. bedeli of which type material has been subsequently examined . Scaphisoma alacre sp. n. Holotype, male: Nepal, Sindhupalcok distr., Malemchi, 2800 m, 14.1V.1981 (Löbl & Smetana) (MHNG). Paratypes, 114: Mustang distr., Lete, 2550 m, 2.X.1983 (Löbl & Smetana) (MHNG) 18 males, 12 females; Kali Gandaki Khola, betw. Mishi and Gomone, 21.IX.1971 (Franz) (Coll. H. Franz, MHNG) 2 females; forest N Lete, 24.IX.1971 (Franz) ((Coll. H. Franz, MHNG) 3 males, 3 females; Lete, 2450-2600 m, 30.1V.1980 (Martens & Ausobsky) (SMNS) 2 males, 1 female; Lete, 24.1X.1971 (Franz) (Coll. H. Franz) 1 female; betw. Lete and Gasa, 25.IX.1971 (Franz) (Coll. H. Franz) 1 male, 4 females: above Shika, 26.[X.1971 (Franz) (MHNG) 2 males; Parbat distr., Chitre, 2400 m, 4.V.1980 (Martens & Ausobsky) (SMNS) 1 male, 1 female; N Ghoropani Pass, 2750 m, 5.X.1983 (Löbl & Smetana) (MHNG) 2 males; Ghoropani Pass, N slope, 2700 m, 6.X.1983 (Löbl & Smetana) (MHNG) 4 males, 2 females; Ghoropani Pass, 2850 m, 5.X.1983 (Löbl & Smetana) (MHNG) 1 female; Rasuwa distr., Dunche, 21-26.IX.1981 (Pawlowski & Kuska) (IZK) 1 male, 2 females; Langtang Khola Valley, 1950 m, 13.1V.1985 (Smetana) (MHNG) 3 males; 1 km NE Bhargu, 2000 m, 12.1V.1985 (Smetana) (MHNG) 1 female; above Bokhajhundo, 1950 m, 11.1V.1985 (Smetana) (MHNG) 1 male; Sindhupalcok distr., Malemchi, as holotype, 16 males, 26 females; Chaubas, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 2 males, 1 female; Patan distr., Phulcoki, 2600 m, 13.X.1983 (Löbl & Smetana) (MHNG) 1 male. Diagnosis. Rather large-sized member of the haemorrhoidale group, with abdominal microsculpture consisting of striae, metasternum with row of punctures parallel to metasternum, discal punctures of elytra well delimited, coarse. Median lobe of aedeagus with slender apical arms of dorsal valves; internal sac with two rows of sclerotized teeth; parameres abruptly widened and moderately curved apically. SCAPHIDIIDAE OF NEPAL 557 Description. Length 1.7 - 1.9 mm, width 1.10 - 1.30 mm. Colour variable, head, pronotum, ventral surface of thorax and basal abdominal ventrites usually dark brown to black, pronotum uniformly more or less dark, or reddish-brown with darkened centre. Elytra in basal 2/3 reddish, usually darkened near sutural striae, sometimes darkened along basal margin and beyond middle; apical third yellowish or ochreous, usually well delimited. Apical abdominal segments, legs and antennae yellowish or ochreous. Antennae with segments IV to XI slender, similar to those of other species of the group. Pronotum with evenly, weakly arcuate lateral margins, lateral keels not visible in dorsal view; punctation dense and fine, well delimited, near base visible at 12x magnification, on centre visible at 20x magnification. Tip of scutellum exposed. Elytron with basal margin weakly arcuate in basal third, then oblique; lateral margin usually not visible in dorsal view, sometimes distinct in apical third; apical margin truncate; inner apical margin exceeding level of outer angle; sutural margin not raised; sutural area flat, with row of fine punctures; sutural stria very weakly converging apically, somewhat curved at base, not extended laterad; discal punctation dense and coarse, punctures well delimited, most distinctly larger than intervals. Pygidium extremely finely punctate. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum not microsculptured. Median portion of metasternum flattened and impressed apically, bearing very dense, rather coarse punctation becoming gradually sparser and finer anteriad. Lateral portion of metasternum with dense row of moderately coarse punctures parallel to metacoxa, with several additional distinct punctures, and with very fine, sparse punctation on most of surface; transverse row not impressed. Mesocoxal area 0.06-0.09 mm long, its margin arcuate, rather finely punctate. Metepisternum moderately narrowed anteriad, near suture impressed below level of metasternum, suture straight except at rounded angles. Abdominal segments with microsculpture consisting of transverse striae. Entire first ventrite sparsely and very finely punctate. Metacoxal area 0.09-0.10 mm long, with arcuate, rather coarsely punctate margin. Tibia straight. Segments 1 and 2 of male pro- and mesotarsus strongly widened, segment 3 moderately widened. Aedeagus (Fig. 153) 0.82 - 0.92 mm long. Dorsal valves with short slender apical arms, ventral piece of median lobe denticulate apically. Internal sac with two apical rows of sclerotized teeth, its central membranous portion very finely denticulate, followed proximally by portion containing long, weakly sclerotized denticles; membranes of basal portion covered by scale-like structures. Parameres straight from base to abruptly widened and moderately curved apical portion, with minute subapical lobe. Remarks. This species may be readily distinguished by the shape of the parameres of the aedeagus. Most specimens may be identified by their colour pattern, in combination with the well delimited discal punctures of the elytra and the relatively fine punctures forming the transverse metasternal row. Scaphisoma nepalense sp. n. Holotype, male: Nepal, Sindhupalcok distr., Pokhare NE Barahbise, 2700 m, 7.V.1981 (Löbl & Smetana) (MHNG). Paratypes, 181: as holotype, and from 2800 m, 2 -3.V.1981, 98 males, 79 females; Durumtali near Barahbise, via Ting Sang La, 2200-2300 m, 5.VIII.1970 (Franz) (Coll. H. Franz) 3 males, 1 female. Diagnosis. Medium-sized member of the haemorrhoidale group, with pale apical portion of elytra. Lateral portion of metasternum with transverse row of punctures. Abdominal microsculpture consisting of striae. Aedeagus with slender arms of dorsal 558 IVAN LÖBL valves. Internal sac with central, weakly sclerotized arcuate plate followed by two rows of teeth. | Description. Length 1.55 - 1.85 mm, width 1.0 - 1.20 mm. Body more or less dark reddish-brown, rarely blackish. Apical fourth to third of elytron yellowish, elytral disc darkened at anterior margin of pale apical area in some specimens. In external characters very similar to S. varians, but elytral punctation less dense and somewhat finer, with most discal punctures about as large as or notably smaller than intervals. Lateral portion of metasternum with distinct transverse row of fine punctures. Aedeagus (Figs 154, 155) 0.71 - 0.74 mm long. Median lobe with slender, apically narrowed arms of dorsal valves; ventral piece long. Internal sac with basal membranes covered by scale-like structures, followed by longer denticles; central portion with elongate, arcuate plate with sinuate apical margin, more strongly sclerotized at lateral margins. Apical portion of internal sac bearing two rows of teeth. Parameres weakly widened at middle, each incurved apically. Remarks. This species may be readily recognised by the shape of the parameres in combination with the characters of the internal sac of the aedeagus. Scaphisoma kanchi sp. n. Holotype, male: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 20.IV.1984 (Löbl & Smetana) (MHNG). Paratypes, 17: as holotype, and from 21.IV.1984, 6 males, 3 females; Arun river at Num, 1500 m, 29.111.1982 and 1500-1600 m, 10.1V.1982 (A. & Z. Smetana) (MHNG) 6 males, 2 females. Diagnosis. Medium-sized species of the haemorrhoidale group with pale apical portion of elytron. Metasternum with row of punctures parallel to metacoxa. Abdominal microsculpture consisting of transverse striae. Aedeagus with distal arms of dorsal valves rather wide and short. Internal sac denticulate in central portion and with spine-like structures in apical portion. Description. Length 1.55 - 1.85 mm, width 1.0 - 1.20 mm. Body dark reddish brown to black, apical fourth of elytron and apical abdominal segments ochreous. In external characters very similar to previous species. Elytron with sutural stria somewhat more converging apically and with discal punctures less delimited. Punctures forming transverse row parallel to metacoxa rather coarse, situated in impressed stria. Aedeagus (Figs 156, 157) 0.79 - 0.81 mm long. Apical portion of median lobe strongly tapering; arms of dorsal valves almost evenly wide. Internal sac with long basal portion divided into two membranous parts covered by fine denticulate structures; centre with a wide tooth accompanied by strongly sclerotized lateral teeth; apical portion of internal sac bearing strong spine-like structures. Parameres strongly narrowed beyond basal third, arcuate, with weakly sclerotized, uneven inner margin. Remarks. This species may be readily distinguished by the aedeagal characters. It differs from S. nepalense, S. prehensor, S. alacre, and S. varians by the transverse row of metasternal punctures situated in an impressed stria. Scaphisoma coalitum sp. n. Holotype, male: Nepal, Ilam distr., betw. Hunse and Mai Pokhari, 1600-2000 m, open land, 9.1V.1988 (Martens & Schawaller) (MHNG). SCAPHIDIIDAE OF NEPAL 559 Diagnosis. Member of the haemorrhoidale group. Body black, large apical portion of elytron pale. Elytral punctation coarse. Metasternum with distinct transverse row of punctures along metacoxa. Abdominal microsculpture consisting of striae. Aedeagus with slender sinuate parameres bearing minute lobe, internal sac armed by apical teeth. Description. Length 1.7 mm, width 1.18 mm. Body black, apical fourth of elytra, apical abdominal segments, antennae and legs ochreous to yellowish. Antennae similar to those of other species of the group. Pronotum with lateral margins weakly arcuate; lateral keels not visible in dorsal view; punctation very fine, distinct at 24x magnification. Tip of scutellum exposed. Elytron with lateral margin arcuate in basal third, almost straight beyond basal third; lateral keel visible in dorsal view from base to apex; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin not raised, sutural area flat, finely punctate; sutural stria weakly diverging from base to anterior third, then parallel to suture, curved along pronotal lobe; discal punctation coarse and dense, most punctures deep, well delimited, larger than or as large as intervals; punctures near sutural stria and apical margin finer. Pygidium extremely finely punctate. Mesocoxal ridge longer than interval between its end and mesocoxa. Metasternum without microsculpture; lateral portion with dense row of rather coarse punctures parallel to metacoxa and with several additional, rather coarse punctures situated close to metacoxa, elsewhere very finely punctate. Medio-apical portion of metasternum weakly impressed, rather densely and distinctly punctate. Mesocoxal area 0.07 mm long, its margin arcuate, finely punctate. Metepisternum narrowed anteriad, impressed along suture, latter rounded only at angles. Abdomen with microsculpture consisting of transverse striae, absent from latero-basal portion of 1st ventrite. First ventrite very finely punctate laterally, with distinct punctures on median portion. Metacoxal area 0.10 mm long, with strongly arcuate, rather coarsely punctate margin. Tibia straight. Segments 1 to 3 of male pro- and mesotarsus widened. Aedeagus (Figs 158-160) 0.86 mm long. Median lobe with short, apically rounded dorsal valves; tip of ventral piece of median lobe truncate in dorsal view. Internal sac bearing several sclerotized apical teeth, subapical sclerotized laminae, scale- like and setose structures covering long membranous portion. Parameres conspicuously slender and sinuate. Remarks. This species is possibly closely related with S. incurvum Löbl from Thailand with which it shares sinuate parameres. It may be distinguished from this by the characters of the internal sac and by the coarser elytral punctation. Scaphisoma coalitum resembles particularly S. minax, but both species exhibit very distinct aedeagi. Scaphisoma sikkimense sp.n. Holotype, male: India, Sikkim (West), Choka, 3050 m, nr. Yuksam, 25.IX.1983 (Sakai) (NSMT). Paratypes, 2: Sikkim (West), Chaozing - Bakkim nr. Yuksam, 2200 - 2670 m, 12.IX.1983 (Sakai) (MHNG) 1 female; Sikkim (West), Bakkim - Choka nr. Yuksam, 2670 - 3050 m, 13.IX.1983 (Sakai) (MHNG) 1 female. Diagnosis. Member of the haemorrhoidale group, with dark triangular elytral spot and metasternum lacking transverse row of punctures parallel to metacoxa. Aedeagus with internal sac bearing two converging subapical teeth, parameres with tuberculate outer margin. Description. Length 1.80 - 1.95 mm, width 1.20 - 1.25 mm. Head and pronotum, including hypomeron, more or less dark reddish-brown. Elytron ochreous or 560 IVAN LÖBL yellowish, with dark brown to blackish triangular basal spot extended at suture to mid- length of elytron. Ventral surface of thorax and Ist abdominal ventrite about as dark as elytral base, following ventrites yellowish or ochreous. Legs and antennae pale reddish brown or yellowish. Antennae similar to those of other species of the haemorrhoidale group. Pronotum with lateral margins arcuate; lateral keel not visible in dorsal view; punctation moderately dense and very fine, barely visible at 24x magnification on most of discal surface, decidedly coarser and more dense near base, many punctures there larger than intervals. Apical portion of scutellum exposed. Elytron with central portion of lateral margin oblique; lateral keels visible in dorsal view only near base; apical margin weakly rounded; inner apical angle exceeding level of outer angle; sutural margin not raised; sutural area flat, with row of very fine punctures; sutural stria weakly converging apically, somewhat curved near base, not extended laterally; discal punctation very dense, rather fine, very shallow, most punctures larger or about as large as intervals, near lateral margin much finer than on centre. Pygidium with microsculpture consisting of transverse striae, extremely finely punctate. Mesepimeral ridge longer than interval between its end and mesocoxa. Metasternum not microsculptured, without medio-apical impression, punctation sparse and very fine on lateral portion, more dense and less fine on middle portion. Metacoxal area 0.05-0.06 mm long, with arcuate, finely punctate margin. Metepisternum flat, weakly narrowed anteriad, somewhat impressed below level of metasternum, suture weakly sinuate to straight with rounded anterior angle. First ventrite very finely punctate, without microsculpture on latero-basal portion, elsewhere with microsculpture consisting of transverse striae. Metacoxal area strongly arcuate, 0.09 mm long, with rather fine marginal punctures. Tibia slender, straight. Segments 1 to 3 of male protarsus notably widened, narrower than apex of protibia. Aedeagus (Figs 161, 162) 0.80 mm long. Median lobe with slender apical portion, dorsal valves short, rather wide. Internal sac complex, with finely denticulate membranes, bearing a pair of subapical, converging teeth and a pair of sub-basal lamellar structures. Parameres weakly curved apically, with tuberculate outer margin, and elongate, narrow inner lobe. Remarks. This species may be distinguished from other members of the group by its colour pattern in combination with the tuberculate outer margin of the parameres and the shape of the sclerotized teeth of the internal sac of the aedeagus. Scaphisoma bhareko sp. n. Holotype, male: Nepal, Patan distr., Phulcoki, 2500 m, 10.V.1981 (Löbl) (MHNG). Paratypes, 307: Nepal, Jumla distr., Dzunda Khola Valley, near Talphi, 3000-3500 m, 19.IX.1972 (Franz) (Coll. H. Franz, MHNG) 4; Mustang distr., 2 km N Kalopani, 2550 m, 1.X.1983 (Löbl & Smetana) (MHNG) 3; Manang distr., Latha Manang W Bagarchhap, 2350-2450 m, 22- 23.1X.1983 (Löbl & Smetana) (MHNG) 58; forest W Bagarchhap, 2200 m, 21.IX.1983 (Löbl & Smetana) (MHNG) 41; Parbat distr., Ghoropani Pass, 2700 and 2850 m, 5. and 6.X.1983 (Löbl & Smetana) (MHNG) 16; Ghoropani Pass, N slope, 2700-2750 m, 5. and 6.X.1983 (Löbl & Smetana) (MHNG) 43; ridge E Ghoropani Pass, 3100 m, 7.X.1983 (Löbl & Smetana) (MHNG) 2; Kaski distr., Dhumpus - Kare near Pokhara (Franz) (Coll. H. Franz) 1; Kathmandu distr., Nagarjung, Jamacok, 1900-2000 m, 18.VIII.1983 (Martens & Schawaller) (SMNS) 1; Siwapuri, 2540 m, 7.X.1981 (Sakai) (Coll. M. Sakai) 3; Siwapuri, 2700 m, 24.11.1982 (Rougemont) (MHNG) 2; Siwapuri, 2500 m, 1.V.1985 (Smetana) (MHNG) 1; Patan distr., Godawari, 6000 ft., 7-13.VIII.1967 (Canad. Nepal Exp.) (CNC) 1; Godawari, Phulcoki, 1770 m, 19.11.1980 (Martens & Ausobsky) and 1700 m, 10.V.1981 (Löbl) (MHNG) 4; Godawari, 13.III.1981 (Rougemont) 1; SE Godawari, cca 1800 m, 22.1V.1988 (Brachat) (MHNG) 1; Phulcoki (Franz) (MHNG) 1; Phulcoki, 2500-2700 m, 10.V.1981, 14-15.X.1983, 28-29.IV.1984 (Lôbl & Smetana) (MHNG) 46; Sindhupalcok distr., near SCAPHIDIIDAE OF NEPAL 561 Shermathang, 1980 (Franz) (Coll. H. Franz) 2; Chaubas, 2500 m, 4.IV.1981 (Löbl & Smetana) (MHNG) 1; Gul Bhanjyang, 2600 m, 6.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi, 2800 m, 14.IV.1981 (Löbl & Smetana) (MHNG) 6; Pokhare NE Barahbise, 2600 and 2700 m, 2., 3. and 7.V.1981 (Löbl & Smetana) (MHNG) 16; Sankhua Sabha distr., forest NE Kuwapani, 2500-2550 m, 11-13. and 15.IV.1982 (A. & Z. Smetana) (MHNG) 4; same but 2350 m, 5.1V.1984 (Löbl & Smetana) (MHNG) 8; Chichila, 2300 m, 26.11.1982 (A. & Z. Smetana) (MHNG) 2; Chichila-Mure, 1900 m, 24.V.1980 (Wittmer) (NMB) 3; Chichila, 2200 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 10; betw. Mure and Hurure, 2050-2150 m, 9-17.V1.1988 (Martens & Schawaller) (SMNS) 4; above Pahakhola, 2600-2800 m, 31.V.1988 (Martens & Schawaller) (SMNS) 1; Induwa Khola Valley, 2100 m, 17.1V.1984 (Löbl & Smetana) (MHNG) 2; Ilam distr., Gitang Khola Valley, 1750 m, 13.1V.1988 m, (Martens & Schawaller) (SMNS) 1; Taplejung distr., SE Yamputhin to Yamputhin, 2000-1650 m, 26. and 30.1V.1988 (Martens & Schawaller) (SMNS) 2; Omje Kharka NW Yamputhin, 2300-2500 m, 1-6.V.1988 (Martens & Schawaller) (SMNS) 4. Diagnosis. Body moderately large, with characteristic colour pattern. Basal elytral stria absent. Metasternum with row of punctures parallel to metacoxa. Abdominal microsculpture consisting of striae. Basal bulb of aedeagus extended dorso-apically, overlapping apical portion of median lobe. Internal sac complex, with paired apical lobes and teeth. Description. Length 1.70 - 2.05 mm, width 1.10 - 1.40 mm. Head and pronotum reddish-brown, medio-basal portion of pronotum more or less darkened, dark area often extended laterally and apically, rarely entire pronotum black. Elytra ochreous with black or blackish brown base, dark basal area extended apically along suture toward apical third, becomming gradually narrower and less dark (Fig. 30). Hypomeron reddish- brown or ochreous. Thorax ventrally and Ist to 3rd abdominal ventrites dark, often blackish-brown or black, following ventrites pale, ochreous or yellowish. Legs and antennae yellowish. Relative length of antennomeres as follows: III 7, IV 18, V 29, VI 23, VII 28, VIII 22, IX 26, X 25, X 30 (holotype); segments IV to VI eachelongate,IV and VI each about 5x as long as wide, V more than 6x as long as wide; VII and XI each about 3x as long as wide, VIII about 4x as long as wide. Pronotum with evenly arcuate lateral margins; lateral keels not visible in dorsal view or distinct only near base; punctation conspicuously coarse and dense near base, visible at 12x magnification, with punctures larger than intervals, those on centre and on apical portion distinctly finer and somewhat less dense. Tip of scutellum exposed. Elytron with lateral margin rounded basally, almost oblique between basal third and apex; entire lateral keel usually visible in dorsal view; apical margin truncate; inner apical angle exceeding level of outer angle; sutural margin not raised; sutural area flat or somewhat vaulted in apical half, with a row of rather coarse punctures; sutural stria gradually, weakly converging apically, hardly curved near base, not extended laterad of pronotal lobe; punctation dense and coarse; punctures near base not or only somewhat larger than those on pronotal base, elsewhere notably larger but not well delimited, distinctly larger than intervals. Pygidium extremely finely punctate. Mesepimeral ridge about 2x as long as interval between its end and mesocoxa. Metasternum not microsculptured, median portion vaulted, with two shallow apical impressions, rather finely and densely punctate. Lateral portion of metasternum very finely and sparsely punctate except for dense row of fairly coarse punctures parallel to metacoxa and for several punctures situated close to it. Mesocoxal area 0.04 - 0.06 mm long, its margin arcuate, coarsely punctate. Metepisternum moderately narrowed anteriad, at same level as metasternum or somewhat impressed, suture straight except for rounded angles. Abdomen with microsculpture of transverse striae. First ventrite fairly coarsely punctate on median portion, very finely punctate laterally. Metacoxal area 0.10 - 0.11 mm long, with arcuate, coarsely punctate margin. Tibia straight. Segment 1 of male protarsus strongly widened but narrower than tibia, following 2 segments moderately widened; 562 IVAN LÖBL segments 1 and 2 of male mesotarsus moderately widened. Lobe of 6st ventrite | conspicuously wide and short, truncate. Aedeagus (Figs 163 to 165) 0.96 - 1.12 mm long.Dorsal membranous wall of basal bulb notably extended apically, and overlapping most of ventro-apical portion of median lobe. Median lobe symmetrical, long, tapering in dorsal view, obliquely inclined, with irregular dorsal margin in lateral view. Internal sac bearing paired apical lobes with strongly sclerotized apices, two slender apical teeth, finely denticulate membranes forming complex structures, and more or less strongly sclerotized central structures. Parameres rather narrow, relatively weakly sclerotized, folded. Remarks. This species is obviously closely related with S. immodicum with which it shares a similar aedeagus. It may be readily distinguished from the latter by the much narrower apical portion of the median lobe. Both species differ drastically in their colour pattern. Caryoscapha Ganglbauer Caryoscapha Ganglbauer, 1899 (sg), type species: Scaphisoma limbatum Erichson, 1845, by original designation. Members of this genus resemble Scaphisoma but exhibit modified apical segment of the maxillary palpus (LOBL 1987a). Four species are assigned to Caryoscapha: C. limbatum (Erichson) disjunctly distributed in Europe and in temperate East Asia, C. seorsum Löbl occuring in Japan, the Nearctic C. americanum Löbl, and the Himalayan C. monticola Löbl. Caryoscapha monticola Löbl Material examined, 1 specimen: Nepal, Manang distr., forest W Bagarchhap, 2200 m, 21.IX.1983 (Löbl & Smetana) (MHNG). Distribution. India: Darjeeling district, Western Nepal. Baeotoxidium Löbl Baeotoxidium Löbl 1971; type species: Baeotoxidium lanka Löbl 1971, by original designation. Baeotoxidium resembles Scaphobaeocera in most characters, including the narrow maxillary galea (Fig. 182) but has elytra without parasutural stria and lacks microsculpture on pronotum, elytra and metasternum. It has a conspicuously thin apical segment of labial palpus (Fig.191). The genus seems to be restricted to the Oriental realm. It includes 7 species, 2 of which occur in the Nepal Himalaya. KEY TO THE SPECIES OF Baeotoxidium 1 Elytron with basal stria uninterrupted, joined to lateral stria ............. siamense Löbl = /Basal-stria of'elytron'absent 1... I I 2, 2 Hypomeron angulate, with longitudinal stria. Antennomere VIII short .................. sui bengalense Löbl SCAPHIDIIDAE OF NEPAL 563 — Hypomeral stria absent. Antennomere VIII elongate ..................................... 3 3 Mesoxocal area margined by row of elongate pits extended laterad of mesocoxa 80 0000 0S Saeed ER AS APS LETTRE CONT RAIN > PRI eres Fe indicum Löbl — Pits margining mesocoxal area not elongate and not extended laterad .................. 4 uelytraldiserdistinetlysbicolored............... green een elegans Löbl SME rali SEUI COLO TE RIETI ER sales 9 5 Mesepimeral ridge indistinct, indicated by extremely fine stria ............... yeti sp. n. = VIESepiIneral Tid oe CaS HMC RI een 6 6 Body blackish. Elytral punctation even, very fine ......................... gagatum Löbl — Body more or less dark reddish-brown. Elytral punctatation uneven, lateral portions moretcoarselyupunetate a. RIT IO AR TR een lanka Löbl NEW RECORDS Baeotoxidium bengalense Löbl Material examined, 74 specimens. Nepal, Parbat distr., Ghoropani Pass, N slope, 2700-2750 m, 5-6.X.1983 (Löbl & Smetana) (MHNG) 5; Kathmandu distr., Shewapuri, 2700 m, 24.III.1983 (Rougemont) (MHNG) 1; Gokarna forest, 1400 m, 10.IX.1983 (Löbl & Smetana) (MHNG) 1; Patan distr., Phulcoki near summit, 2900 m, 4.X.1971 (Franz) (Coll. H. Franz) 3; Phulcoki, 2500-2700 m, IV, V and X. 1981, 1983 and 1984 (Löbl & Smetana) (MHNG) 48; Sindhupalcok distr., Pokhare NE Barahbise, 2800 m, 2-5.V.1981 (Löbl & Smetana) (MHNG) 2; Sankhua Sabha distr., above Sheduwa, 3000 m, 2.IV.1982 (A. & Z. Smetana) (MHNG) 1; Bakan W Tashigaon, 3250 m, 4.1V.1982 (A. & Z. Smetana) (MHNG) 1; Arun Valley, Chichila, 2200 m, 24.1V.1982 (Löbl & Smetana) (MHNG) 1; forest NE Kuwapani, 2350 m, 5.1V.1984 (Löbl & Smetana) (MHNG) 1; bottom Arun Valley below Num, 1050 m, 20-22.1V.1984 (Löbl & Smetana) (MHNG) 4; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 1; Induwa Khola Valley, 2000 m, 16.IV.1984 (Löbl & Smetana) 1; Taplejung distr., Khabeli Khola, Yamputhin, 1650-1800 m, 3- 4.VTIL.1983 (Martens & Daams) (SMNS) 1; left bank of Khabeli Khola, 1800-2000 m, 27- 29.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 2; Hellok in Tamur Valley, 2000 m, 17.V.1988 (Martens & Schawaller) (SMNS) 1; upper Tamur Valley, 1800-2150 m, 19.V.1988 (Martens & Schawaller) (SMNS) 1; descent from Pass Deorali to Hellok, 2600-2000 m, 17.V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. India: Darjeeling distrinct, Nepal. NEW SPECIES Baeotoxidium yeti sp. n. Holotype, male: Nepal, Sindhupallcok distr., Malemchi 2800 m, 14.IV.1981 (Löbl & Smetana) (MHNG). Paratypes, 2: Nepal, Manang distr., Latha Manang W Bagarchap, 2350 m, 22.IX.1983 (Löbl & Smetana) (MHNG) 1 female; Sindhupalcok distr., Gul Bhanjyang, 2600 m, 6.IV.1981 (Lôbl & Smetana) (MHNG) 1 female. Diagnosis. Body ochreous, very finely punctate. Hypomeron not striate. Elytron without basal stria. Flagellum of internal sac with two basal hooks. Description. Length 1.15 - 1.30 mm; width 0.69 - 0.78 mm. Body uniformly ochreous, or apical portion of elytra darkened. Femora, tibiae and antennae hardly paler 564 IVAN LÖBL than body, tarsi and apical abdominal segments notably paler. Relative length of antennal _ segments as follows: III 9, IV 9, V 12, VI 10, VII 17, VIII 9, IX 15, X 16, XI 28; segments II to VI of almost same width; VI and VII about 2.5x as long as wide, VIII less than 2x as long as wide; XI 3x as long as wide. Pronotal punctation extremely fine, hardly visible at 100x magnification. Point of scutellum exposed. Hypomeron concave, lacking longitudinal stria. Elytron weakly narrowed apically; sutural stria deep, curved at base, not extended laterally; sutural margin not raised; discal punctation similar to that on pronotum. Mesepimeral ridge indistinct. Metasternum and Ist ventrite without microsculpture, both very finely punctate laterally. Mesocoxal area 0.04 mm long, with marginal pits not elongate and not extended laterally. Metepisternum flat, large, 0.08 - 0.11 mm wide, apically somewhat narrowed, with straight suture. Basal pits of Ist ventrite weakly elongate. Tibiae straight. Male protarsus with segments 1 to 3 moderately widened. Aedeagus (Fig. 166) 0.26 mm. Median lobe weakly curved. Parameres almost straight, with weakly sclerotized inner apical portion. Internal sac with long flagellum thickened basally and bearing two basal hooks, forming single complete circle when extruded. Remarks. This species may be readily recognised by the combination of the external characters (see the key). Scaphobaeocera Csiki Scaphobaeocera Csiki, 1909; type species: Scaphobaeocera papuana Csiki, 1909, by monotypy. Nesotoxidium Scott, 1922; type species: Nesotoxidium typicum Scott, 1922, by monotypy. Most Scaphobaeocera species may be distinguished from other scaphidiids in having the pronotum and elytra microsculptured and iridescent. However, the genus is defined by the presence of a parasutural stria on each elytron. So far 59 species have been recognised as valid, ranging in Asia from Pakistan and Sri Lanka to Japan, and southeastward to Australia and Micronesia. A few species are known to occur in the Mascarene and Seychelle Islands and in tropical Africa. Twelve species have been found in the Nepalese collections, one of which is described as new below. KEY TO HIMALAYAN SPECIES OF Scaphobaeocera (including species from Meghalaya) [Ely tronochreous Withitwo dark fasciae ee species indet. c Colour. pattern of elytron different ................... Tee TT 2 2 Antennomeres VII and IX conspicuously large, each 4x as long as segments..... VI or AVAL Era lana TER IT ERI zdenae sp.n. tu Antennomeres VIT and IX shorter:.:..""""" "0 TE 3 3, -Antennomere XI as long. as, or shorter than X or IX 380 ee spinigera Löbl = Antennomere XI longer than segments X or EX." here ren + 4 Antennomere XI about as long as segments X and IX combined......................... 5 — Antennomere XI shorter than X and [IX combined "ee" N ee 6 5 Length 1.3 - 1.5 mm. Aedeagus with flagellum widening basally ....stephensoni Löbl — Length 1.2 - 1.2 mm. Aedeagus with flagellum narrowed basally......... dorsalis Lobl 6 Upper portion of hypomeron delimited by longitudinal stria from lower portion...... 7 Ur Hypomeron without longitudinal stria... ur en. ee PA eee eee 9 1 Meso=and metatibiae incurved ee 0 OOO tibialis Löbl SCAPHIDIIDAE OF NEPAL 565 Iibiaegstraisht ES Be DA ARR RE Mauch PA Res 8 Aedeagus with parameres widening apically (lateral view)................ aberrans Löbl Aedeagus with parameres narrowed apically (lateral view) ................ querceti Löbl Punctation on lateral portion of metasternum fairly coarse, notably coarser than that ONE lO Mere LTT cis rae ee eek species indet. a Lateral portion of metasternum about as finely punctate as elytron..................... 10 Antennal segment VIII very small, no more than half as long as segment VII........ 11 Antennal segment VIII longer than half of segment VIL.................................. 12 Pits margining mesocoxal area large, elongate, extended laterad ......... discreta Löbl Pits margining mesocoxal area small, not elongate, not extended laterad ................- ree RO I SO A SARA AR IAA NA RNA NI species indet. b Metasternum and Ist ventrite lacking microsculpture .................................... 13 MetasternumtandAlstiventritesmieroseulptured an seen. 14 Length 0.95 - 1.20 mm. Body pale, ochreous or reddish ...................... nuda Löbl Length 1.45 - 1.55 mm. Body dark reddish brown ....................... fratercula Löbl INICtASteHUMEWI MENSE ORO 15 INEtASternUmMEWINOUHMEdi amnistia, RI 16 Aedeagus with flagellum forming 3 or 4 complete circles ................. difficilis Löbl Aedeagus with flagellum forming 6 complete circles .......................... spira Löbl Basal bulb of aedeagus extended apically beyond level of parameral base, with oblique, abruptly delimited apical wall (lateral view) ........................ nobilis Löbl INCACAUSICIFECIENAR TN ee 17 Median lobe of aedeagus with ventral plate protruding apically above base of PARAMETRI BE N I lamellifera Löbl IMedianlloberofaedeasusilackinsyentrakplater = ea ene eee 18 Ventral processi of aedeagus large, strongly protruding apically ........... tenella Löbl Ventral processi of median lobe small not protruding apically ......................... O Internal sac of aedeagus with flagellum joined to additional basal sclerite ............... 2606 60 26 0 E N I E CINTO I I TA LAER eek a timida Löbl Internal’sac’of’aedeagus’lacking additionalisclerite?-.. ne 20 Hlagellumiotraedeaguswithibasallhook er an nen cognata Löbl Elagellumtof’aedeaguslackingbasallhook.. nn. een. DAI Rlagellumiotacdeagus! spirale tcc E ORE CEE RT mussardi Löbl Flagellum of aedeagus sinuate, not forming complete circles .......... minuta (Achard) NEW RECORDS Scaphobaeocera nuda Löbl Material examined,S specimens: Nepal, Kaski distr., near Pokhara lake, 20.IX.1979 (Franz) (Coll. H. Franz) 1; Patan distr., Godawari, 1600 m, 31.111.1984 (Löbl) (MHNG) 1; Dhanding distr., Thorpu to Kordung, 1300-1400 m, 24.VI.1983 (Martens & Schawaller) (SMNS, MHNG) 3. Distribution. India, Nepal, Thailand. Scaphobaeocera minuta (Achard) Material examined, 4 specimens: Nepal, Sankhua Sabha, Chichila, 2200 m, 4.1V.1984 (Löbl & Smetana) (MHNG) 2; forest S Mangsingma, 2200 m, 11-13.1V.1984 (Löbl & Smetana) (MHNG) 2. 566 IVAN LÖBL Distribution. North India, Nepal, Thailand. Scaphobaeocera dorsalis Löbl Material examined, 1 specimen. Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 21.1V.1984 (Löbl & Smetana) (MHNG). Distribution. North India, Nepal, Thailand, Thaiwan. Scaphobaeocera stephensoni Löbl Material examined, 13 specimens: Nepal, Kathmandu distr., Gokarna forest, 1400 m, 1.IV.1981 (Löbl & Smetana) (MHNG) 2; Sankhua Sabha distr., forest NE Kuwapani, 2500 m, 28.11.1982 (A. & Z. Smetana) (MHNG) 2 and 2250 m, 24.IV.1984 (Löbl & Smetana) (MHNG) 1; Chichila, 2200 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 1; bottom Arun Valley below Num, 1050 m, 21.1V.1984 (Löbl & Smetana) (MHNG) 2; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) 2; Taplejung distr., above Yamputhin, left bank of Kabeli Khola, 1800-2000 m, 27- 29.1V.1988 (Martens & Schawaller) (SMNS) 1; Omje Kharka NW Yamputhin, 2300-2500 m, 1- 6.V.1988 (Martens & Schawaller) (SMNS, MHNG) 2. Distribution. North India (Himachal Pradesh), Nepal. Scaphobaeocera spinigera Löbl Material examined, 27 specimens: Nepal, Sindhupalcok distr., Malemchi, 2800 m, 14. and 17.IV.1981 (Löbl & Smetana) (MHNG) 9; Kathmandu distr., Gokarna forest, 1400 m, 1.1V.1981 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., Arun River at Num, 1500-1600 m, 10.1V.1982 (A. & Z. Smetana) (MHNG) 1; ridge S Mangsingma, 2800 m, 8.1V.1984 (Löbl & Smetana) (MHNG) 1; above Pahakhola, 2600-2800 m, 31.V.-3.VI.1988 (Martens & Schawaller) (SMNS) 1; Ilam distr., Mai Pokhari, 2100-2200 m, 9-10.1V.1988 (Martens & Schawaller) (SMNS) 1; Taplejung distr., SE Yamputhin to Yamputhin, 26. and 30.1V.1988 (Martens & Schawaller) (SMNS) 1; India, Himachal Pradesh, 4 km SW Solan, 1500 m, 8.X.1988 (Vit) (MHNG) 12. Distribution. Pakistan, India, Nepal, Thailand. Scaphobaeocera tenella Löbl Material examined, 3 specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 21-22.1V.1984 (Löbl & Smetana) (MHNG). Distribution. India (Meghalaya), Nepal, Thailand. Scaphobaeocera cognata Löbl Material examined, 8 specimens: Nepal, Kathmandu distr., Nagarjung, Jamacok, 1900-2100 m, 18.VIII.1983 (Martens & Schawaller) (MHNG) 1; Patan distr., Phulcoki, Godawari, 1700 m, 10.V.1981 (Löbl) 1; Sindhupalcok distr., Pokhare NE Barahbise, 2700 m, 7.V.1981 (Löbl & Smetana) 1; Taplejung distr., Yamputhin, 1800 m, 26.IV.-1.V.1988 (Martens & Schawaller) (SMNS) 2; above Yamputhin, left bank of Kabeli Khola, 1800-2000 m, 27-29.1V.1988 (Martens & Schawaller) (SMNS, MHNG) 3. SCAPHIDIIDAE OF NEPAL 567 Distribution. North India (Garhwal and Meghalaya), Nepal. Scaphobaeocera spira Löbl Material examined, 1 specimen: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.1V.1988 (Löbl & Smetana) (MHNG). Distribution. Nepal, Thailand. Scaphobaeocera mussardi Löbl Material examined, 4 specimens: Nepal, Sankhua Sabha distr., forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 1; Chichila, 2200 m, 24.IV.1984 (Löbl & Smetana) (MHNG) 1; forest NE Kuwapani, 2250 m, 24.1V.1984 (Löbl & Smetana) 1; Induwa Khola Valley, 2100 m, 17.1V.1984 (Löbl & Smetana) (MHNG) 1. Distribution. Sri Lanka, India, Nepal, Bhutan. Scaphobaeocera difficilis Löbl Material examined, 99 specimens: India, Himachal Pradesh, 10 km NW Solam, 1700 m, 7.X.1088 (Vit) (MHNG) 35; Barog forest 4 km SW Solam, 1500 m, 8.X.1988 (Vit) (MHNG) 13; Kulu Valley, Naggar 1850 m, 16.X.1988 (Vit) (MHNG) 2; Dalhousie, Shubhash Baoli, 2080 m, 20.X.1988 (Vit) (MHNG) 1; Khajiar E Dalhousie, 1950 m, 21.X.1988 (Vit) (MHNG) 1; Mashabra forest NE Simla, 2100 m, 30.X.1988 (Vit)(MHNG) 1; Nepal, Parbat distr., Ghoropani Pass, 2850 m, 5.X.1983 (Löbl & Smetana) (MHNG) 1; Rasuwa distr., below Gosaikund, 2680 m, 21.X.78 (Cassagnau) (MHNG) 1; Kathmandu distr., Siwapuri Dara, 2400 m, 30.1V.1985 (Smetana) (MHNG) 1; Gokarna forest, 1400 m, 31.11.1981 (Löbl & Smetana) (MHNG) 1; Patan distr., Godawari, Phulcoki, 2000 m, 21.1V.1988 (Brachat) (MHNG) 2; Phulcoki, 2500-2700 m, IV. and V.1981 and 1984, X.1983 (Löbl & Smetana) (MHNG) 13; Sindhupalcok distr., Chaubas, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 1; Gul Bhanjyang, 2600 m, 6.IV.1981 (Löbl & Smetana) (MHNG) 1; Tarke Ghyang, 2650 m, 19.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi, 2800 m, 14. and 17. IV. 1981 (Löbl & Smetana) (MHNG) 4; Pokhare NE Barahbise, 2700 m, 2.V.1981 (Löbl & Smetana) (MHNG) 1; Sankhua Sabha distr., Arun Valley, betw. Mure and Hurure, 2550-2150 m, 3-17.V1.1988 (Martens & Schawaller) (SHNS) 1; Chichila, 2200 m, 4.1V.1988 (Löbl & Smetana) (MHNG) 2; Chitre, 2200-2400 m, 28-29.V.1985 (Holzschuh) (NMB) 1; forest NE Kuwapani, 2450 m, 13.1V.1982 (A. & Z. Smetana) (MHNG) 4; forest S Mangsingma, 2250 m, 12.1V.1984 (Löbl & Smetana) (MHNG) 1; Induwa Khola Valley, 2000 m, 16-17.1V.1984 (Löbl & Smetana) (MHNG) 5; Panchtar distr., Paniporua, 2300 m, 18-20.1V.1988 (Martens & Schawaller) (SMNS) 1; Taplejung distr., Lassetham NW Yamputhin, 3300-3500 m, 6-9.V.1988 (Martens & Schawaller) (SMNS) 1; Ilam distr., Cilang Khola Valley, 11-13.1V.1988 (Martens & Schawaller) (SMNS) 1; Mai Pokhari, 2100-2200 m, 9-10.1V.1988 (Martens & Schawaller) (SMNS) 3. Distribution. Pakistan, India, Nepal, Thailand. Scaphobaeocera timida Löbl Material examined, 115 specimens: Nepal, Jumla distr., Dzunda Khola Valley near Talphi, 3000-3500 m, (Franz) (Coll. Franz) 1; Manang distr., forest W Bagarchap, 2200-2500 m, 22. and 24.1X.1983 (Löbl & Smetana) (MHNG) 22; Latha Manang W Bagarchap, 2350 m, 22.IX.1983 (Löbl & Smetana) (MHNG) 7; Mustang distr., 2 km N Kalopani, 2550 m, 1.X.1983 (Löbl & 568 IVAN LÖBL Smetana) (MHNG) 1; Parbat distr., Ghoropani Pass, 2700-2850 m, 5-6.X.1983 (Löbl & Smetana) (MHNG) 35; Ridge E Ghoropani Pass, 3100 m, 7.X.1983 (Löbl & Smetana) (MHNG) 1; Ghoropani ~ Pass, 3000 m, 19.IX.1971 (Franz) (Coll. H. Franz) 1; Mustang distr., Tukuche, valley to Taksang, ca 3000 m, 23.1X.1971 (Franz) (Coll. Franz) 1; above Shika, Khali Gantaki Valley, 26.IX.1971 (Franz) (Coll. H. Franz) 1; Kaski distr., above Dhumpus, 2100 m, 8-10.V.1980 (Martens & Ausonsky) (SMNS) 1; Kathmandu distr., Gokarna forest, 1400 m, 3.X.1971 (Franz) (Coll. H. Franz) 1; Patan distr., godawari, Phulcoki, 1700 m, 10.V.1981 (Löbl) (MHNG) 1; Phulcoki, 2300-2700 m, V.1981, X.1983, IV.1984 (Löbl & Smetana) (MHNG) 7; Phulcoki, 2600 m, 9.VIII.1970 (Franz) (Coll. H. Franz) 2; Sindhupalcok distr., Chaubas, 2600 m, 5.1V.1981 (Löbl & Smetana) (MHNG) 2; Gul Bhanjyang, 2600 m, 6.1V.1981 (Löbl & Smetana) (MHNG) 2; Malemchi, 2800 m, 14.1V.1981 (Löbl & Smetana) (MHNG) 1; Shermathang, X.1980 (Franz) (Coll. H. Franz, MHNG) 6; Durum Valley near Barahbise, 2200-2300 m, 5.VIII.1970 (Franz) (Coll. H. Franz, MHNG) 2; above Barahbise, 2200-2300 m, 5.VIII.1970 (Franz) (Coll. H. Franz) 1; Sankhua Sabha distr., Chichila, 2200 m, 24.1V.1984 (Löbl & Smetana) (MHNG) 2; forest S Mangsingma, 2200 m, 11.1V.1984 (Löbl & Smetana) (MHNG) 1; above Pahakhola, 2600-2800 m, 31.V.-3.VI.1988 (Martens & Schawaller) (SMNS) 1; Arun Valley, betw. Mure and Hurure, 2050-2150 m, 17.VI.1988 (Martens & Schawaller) (SMNS) 1. Distribution. North India (Himachal Pradesh and Kumaon), Nepal, Bhutan. Scaphobaeocera species indet. c Material examined, 4 females: Nepal, Sankhua Sabha distr., Jaljale Himal, 2530 m, 21.X1.1978 (Casagnau) (MHNG) 1; Therhathum distr., Tinjura Dara, 2450-2850 m, 17.IX.1983 (Martens & Daams) (SMNS) 1; Dankuta distr., Dankuta, 18.IX.1978 (Bhakta) (NMB) 1; Cheavri Bas, 18.IX.1978 (Bhakta) (MHNG) I. Remarks. This is possibly a new species. It resembles much S. alticola, especially in the colour pattern and in having elytron with somewhat shortened sutural striae. It may be distinguished by the relatively much shorter metasternum (with smallest interval between meso- and metacoxa not exceeding 0.10 mm). However, I am reluctant to name it in the absence of males. NEW SPECIES Scaphobaeocera zdenae sp.n. Holotype, female: Nepal, Sankhua Sabha distr., forest NE Kuwapani, 2500 m, 28.11.1982 (A. & Z. Smetana) (MHNG). Diagnosis. Medium-sized species. Antennomeres VII and IX very large, each 4x as long as segment VIII and 6x as long as wide. Hypomeron lacking longitudinal stria. Metasternum without median impression or stria. Description. Length 1.65 mm, width 1.07 mm, dorso-ventral diameter 1.09 mm. Body, antennae and legs almost uniformly dark brown. Pronotum, elytra, and abdominal ventrites microsculptured and iridescent. Punctation extremely fine on both, dorsal and ventral sides of body, setiferous punctures around smooth metasternal centre excepted. Relative length of antennomeres as follows: III 8, IV 14, V 16, VI 10, VII 40, VII 10, IX 40 (segments X and XI missing in both antennae). Segments III to VI each gradually widening apically; III about 1.6x as long as wide; IV and V each 3x as long as | wide; VI about 1.8x as long as wide; VII and IX each 6x as long as wide, parallel-sided; VII about 2x as long as wide. Hypomeron with larger lower portion almost vertical, at an SCAPHIDIIDAE OF NEPAL 569 angle with upper portion; longitudinal stria absent. Metasternum without median impression or stria, distinctly microsculptured laterally. Mesocoxal area 0.03 mm long, with fine marginal pits. Metepisternum flat, parallel-sided, 0.10 mm wide, with straight suture. Basal punctures of 1st ventrite small, not elongate. Tibiae I and II straight, III somewhat curved. Remarks. This species may be easily distinguished from all other members of the genus by the conspicuously large antennal segments VII and IX, each being 4x as long as segments VI or VIII. Scaphoxium Löbl Scaphoxium Löbl, 1979; type species: Toxidium madurense Pic, 1920, by original designation. Scaphoxium may be readily separated from other scaphidiids possessing strongly approximate meso- and metacoxae by the peculiar shape of the basally lobed or extended hypomeron. It may also be distinguished by the curved elongate antennomere III. All Scaphoxium have bodies strongly convex dorsally and ventrally, elytra finely punctate with shortened sutural striae and parameres of aedeagus abruptly narrowed subapically. The genus includes 25 species ranging from India and Sri Lanka to Japan, Australia and Melanesia. Several additional Asian and Afrotropical unidentified species are represented in the collection of the MHNG. KEY TO THE ASIAN SPECIES OF Scaphoxium eee lytronidark,-with two pale’ transverse ASCIAE ee ce ee 2 RE hy ATOMMUMULOTIM VACOlOUTEH' nn. ee 3 2 Length 1.6 - 1.9 mm. Metasternum not impressed at middle ............... zebra (Löbl) — Length 1.25 - 1.40 mm. Metasternum impressed at middle .............. oblitum (Löbl) Ba lrarseispecies2 DS mmlloner. een grande Löbl = SmallenspeeiesJlenethinotiexceeding 1. immo 4 4 Lateral portion of metasternum with relatively coarse punctures .......................... eee E elo DOS UN (Champion), Pateral portion of metasternum very finely punctate nn. senden eee eee cece ene 3) Dinternalsacioffaedeagusimembranouser ren. ee singlanum Löbl Sn intermnalisac/offaedeagus bearine SClerites I een: 6 Be Metasternum flat onconvexcatimiddle: PRES ER Eee 7 Ze Metasterinumeimpressedkatmiddle rn gen ee TTT 18 7) Each paramere of aedeagus with large subapical lobe." "0500 i 8 se arameresjoftaedeagusidiiterent m ee 10 8 Antennomere III as long as IV, segment V as long as VI. Internal sac of aedeagus beaning3Ssshortiseleritese na N REN avidum Löbl — Antennomere III longer than IV, segment V longer than VI. Internal sac of aedeagus WA PAMOLIONOASCIERMES ne ER EN PR NE 9 9 Parameres of aedeagus gradually tapering beyond subapical lobe. Sclerites of internal SAGUNEU ALPEN. ER se ae ee ee Nr i Vaste taiwanum Löbl — Parameres of aedeagus abruptly narrowed beyong subapical lobe. Sclerites of internal SAC MOM UL ATs RE N Eee Bese es topali Löbl 10 Internal sac with apical portion of sclerites uncular ...................... madurense (Pic) 570 IVAN LÖBL MN Scerites/of intemallsacnot uncular 2.2 O TTT P[ TRE 11 . 11 Parameres of aedeagus each bearing minute subapical denticle-like apophysis .......... EN ee Dont nee asien DUT ad dre ROEM RI ARAM AOI A CARCI ac Japonicum Löbl = wukarameres of acdeasus different... TT [[. ES 12 DZ Eensthil>S7mm. Antennomere V longer than Il... en en taylori Löbl Men gthil2immVAntennomereiVias ons as TP eee assamense Löbl 13 Mesosternum with median keel partly dividing median impression ..... sparsum Löbl =)» Mesosterum lacking medianikeel".............=.... 22.002. TETTE 14 14 Parameres of aedeagus with wide, rounded subapical lobe .............. keralense Löbl — Parameres of aedeagus with narrow, pointed subapical lobe ........................... 15 15 Parameres of aedeagus widened or subparallel beyong level of subapical apophysis, MALO WCC MEAT APEX nennen coe cs secs scene esse ee ne eximium Löbl — Parameres of aedeagus tapering from level of subapical apophysis ................... 16 lop Subapicalllobe of parameres minuten... An. ne reductum Löbl — Subapical lobe of parameres long, longer than width of paramere at same level .... 17 17 Antennomere III as long as IV. Internal sac of aedeagus without a pair of elongate basallseleritessnr ee N PME TUE RETTE simulans (Löbl) — Antennomere III longer than IV. Internal sac of aedeagus with a pair of long basal SCIETILCS PE EER NEN ER NET Mais PIO DDA, intermedium Löbl NEW RECORDS Scaphoxium eximium Löbl Material examined, 4 specimens: India, Himachal Pradesh, 10 km NW Sarahan, 1700 m, 7.X.1988 (Vit) (MHNG). Distribution. North India: Himachal Pradesh, Kumaon. Remarks. The armature of the not extruded internal sac of the aedeagus is as in Fig. 167. Scaphoxium taiwanum Löbl Material examined, 1 specimen: Nepal, Sankhua Sabha distr., Induwa Khola Valley, 2000 m, 18.IV.1984 (Löbl & Smetana) (MHNG). Distribution. India, Nepal, Thailand, Taiwan. Scaphoxium sparsum Löbl Material examined, 25 specimens: Kaski distr., Tandarakot, trail Pokhara to Ghoropani, ca 1000 m, 18.IX.1971 (Franz) (Coll. H. Franz, MHNG) 6; hill near Pinta, Kaste Lake, 20.IX.1978 (Franz) (Coll. H. Franz) 2; Kathmandu distr., Rani Ban SE Sanogau, 1500-1600 m, 25.1V.1988 (Brachat) (MHNG) 1; Patan distr., Godawari, 1600 m, 31.11.1984 (Löbl) (MHNG) 1; Phulcoki near Dalikhel, ca 1900 m, 21.IX.1977 (Franz) (Coll. H. Franz, MHNG) 2; Sankhua Sabha distr., bottom Arun Valley below Num, 1050-1100 m, 20-21.1V.1984 (Löbl & Smetana) (MHNG) 9; Arun River at Num, 1500-1600 m, 10.1V.1982 (A. & Z. Smetana) (MHNG) 2; Khandbari, 1700 m, 23.11.1982 (A. & Z. Smetana) (MHNG) 1; Induwa Khola Valley, 2000 m, 18.1V.1984 (Löbl & Smetana) (MHNG) 1. SCAPHIDIIDAE OF NEPAL 571 Distribution. India, Nepal, Thailand. Scaphicoma Motschulsky Scaphicoma Motschulsky, 1863; type species: Scaphicoma flavovittata Motschulsky, 1863, by monotypy. Lepteroscapha Achard, 1921; type species not designated. Members of this genus differ by conspicuously long tarsi and antennae in comparision with all other scaphidiids with similarly strongly approximate meso- and metacoxae. The mouthparts in Scaphicoma (Figs 177, 187, 188) are very similar to those in Toxidium. The genus occurs in tropical Africa and in Asia. Twelwe species habe been recognised, one of which is encountered in the Himalayan region. NEW RECORD Scaphicoma arcuatum (Champion) Material examined, 4 specimens: Nepal, Sankhua Sabha distr., bottom Arun Valley below Num, 1050 m, 22.IV.1984 (Löbl & Smetana) (MHNG) 3; Taplejung distr., Yamputhin, 1800 m, 26.IV. - 1.V.1988 (Martens & Schawaller) (SMNS) 1. Distribution. India: Kumaon, Darjeeling district and Assam, Nepal, Burma, Thailand. Remarks. Itis difficult to distinguish the members of this genus by their aedeagal characters. However, minor distinguishing characters may be found in the shape of the parameres and in the structure of the internal sac. The aedeagus of S. arcuatum is as in Fig. 173. Toxidium LeConte Toxidium LeConte, 1860; type species: Toxidium gammaroides LeConte, 1860, by monotypy. Most of the strongly ventrally vaulted scaphidiids with approximate meso- and metacoxae and with large lateral portions of metasternum were previously assigned to Toxidium. When revising collections, I have transferred many of these species to Scaphobaeocera, Baeotoxidium, Scaphoxium and Scaphicoma. In the Asian and Australian faunas, only two species groups remain to date in Toxidium: the aberrans, and the montanum groups. The latter, for which a new genus is erected below, consists of three described and one new species. The species of the aberrans group share most of the diagnostic characters with T. gammaroides and its New World allies: labrum with marginal row of short thick setae; mandible bidentate apically, with comb-hairs, very fine prosthecal hairs and bunch of premolar hairs (Fig.178); maxillary lacinia slender, setose apically; penultimate segment of maxillary palpus subcylindrical, apical segment tapering, at base almost as thick as preceding segment at apex (Fig. 185); labial palpus (Fig. 190) with segment II about as long as wide, and as wide as I, segment III curved, gradually narrowed apically; eye notched at or above level of its mid-length, antennal insertion fairly distant from clypeal 572 IVAN LÖBL suture; antennomere III slender, straight, similar to the following segments; pronotum with _ basal angles obtuse and not extended, not reaching level of anapleural suture; pair of internal pronotal cavities (or pockets) absent; hypomeron weakly inflexed, not explanate apically; elytral parasutural stria absent, sutural stria usually shortened; prosternal keel sharply raised; mesosternal median keel obsolete or very low, pleural ridge of first ventrite distinct. All species except T. vagans differ from the New World species of Toxidium by the relatively less approximate meso- and metacoxae. All Asian species, including several undescribed species represented in the collection of MHNG, may be also distinguished from the New World members of Toxidium by the ventrally less vaulted body. In addition, these species, T. incompletum and an unidentified species from Thailand excepted, share the unusual character combination of elytron, i. e. shortened sutural and distinct basal striae. In Toxidium, the mesepimeral ridge is usually absent, but it is present in T. indicum. The rare monotypic Scaphischema from the western Mediterranean area shares almost all the diagnostic characters of Toxidium. It is particularly similar to the species of T. aberrans group. KEY TO THE ASIAN SPECIES OF Toxidium (only named species) 1 Most of elytron bright reddish, surface along sutural stria and apical fifth of elytron ES LA ee OE RE spectabile sp. n. — Elytron and pronotum unicolorous, or elytron with small pale transverse fascia ...... 2 2 Metasternum short, mesocoxal area about as long as shortest interval between its marsinländmetacoxa 30. TTT. vagans Löbl — Metasternum long, mesocoxal area much shorter than shortest interval between its marginand metaxoca ........sussenese nenne acted ope nenne TEZTT 3 3m Elytron'withibasalistnia..........n Mena tone a Paso Re eee 4 mew asalistiavohelytrom absentee. PI O incompletum Lobl 4 Lateral portion of metasternum with fairly coarse punctures. Mesepimeral ridge distinch RAS ah ee ee indicum Achard — Entire metasternum very finely punctate. Mesepimeral ridge absent ................... 5 5 Pronotal punctation coarse, distinct at magnification 12x ............. aberrans Achard — Pronotal punctation very fine, hardly visible at magnification 25x ...................... 6 6 Elytron with fairly long sutural stria starting beyond basal 1/8 of sutural length ...... 7 — Elytron with very short sutural stria, visible only near elytral apex ..................... 8 TRE lytralupunctationientirely coarse. rer ee curtilineatum Champion “= WE lyfron\finely;punctate;apically SES ee eee eee styligerum Löbl 8 Elytron with basal stria obsolete laterally; elytral punctation almost obsolete basally ee O INCENERITORI LL daly Se ARS ERS PIEDS NE DE CEE eee pubistylis Löbl — Elytron with basal stria joined to lateral stria; elytral punctation distinct at base ...... 9 9 Mesocoxal area about as long as half of interval between its margin and margin of MÉTACOXA essen TE diffidens Löbl — Mesocoxal area distinctly shorter than half of interval between its margin and METACOXA EEE ER EERENERREN SEHE TRO LTT robustum Pic NEW RECORDS Toxidium curtilineatum Champion Material examined, 7 specimens: Nepal, Lamjung distr., Marsyandi, 1100-1250 m, Senghe-Jagat, 11.1V.1980 (Martens & Schawaller) (SMNS) 1; Kathmandu distr., Gokarna forest, SCAPHIDIIDAE OF NEPAL 578 31.11 - 1.IV. 1981 and 20.X.1983 (Löbl & Smetana) (MHNG) 4; Sankhua Sabha distr., Mure-Num, 1900-1500 m, 25.V.1980 (Wittmer) (NMB) 1; Chichila-Mure, 1900 m, 24.V.1980 (Wittmer) (MHNG) 1. Distribution. North India (Garhwal, Kumaon and Meghalaya), Nepal. Remarks. Someof the Nepalese specimens have a narrow pale transverse fascia on apical half of elytron. The aedeagi of these specimens (Figs 168, 189) are not different from those of the uniformly coloured specimens. No additional distinguishing feature has been found to separate both forms. NEW SPECIES Toxidium spectabile sp. n. Holotype, female: Nepal, Taplejung distr., above Yamputhin, left bank of Kabeli Khola, open forest, 1800-2000 m, 27-29.IV.1988 (Martens & Schawaller) (MHNG). Diagnosis. Elytron with distinct colour pattern, basal stria present in centre of basal margin only, sutural stria obsolete except in apical tenth of sutural length. Metasternum long. Description. Length 2.7 mm, width 1.6 mm. Head black. Pronotum reddish- brown, distinctly darkened at apical margin, somewhat darkened in middle. Elytron bright reddish,reddish-brown near base, blackish along sutural stria and on entire apical fifth. Mesepisternum, metepisternum and lateral portion of metasternum blackish, prosternum, mesosternum and centre of metasternum, ventrites I to IV and legs reddish-brown, apical abdominal segments and antennae ochreous or yellowish. Relative length of antennomeres as follows: III 25, IV 32, V 38, VI 33, VII 34, VII 25, IX 33, X 33, XI 38; segment VII almost 3.5x as long as wide; VII and XI each about 3x as long as wide. Pronotal punctation sparse and very fine, visible at magnification 25x. Exposed portion of scutellum fairly large. Elytron with basal stria visible along middle portion of basal margin, obsolete near pronotal lobe and near lateral margin; sutural stria strongly reduced, distinct in apical tenth of sutural length; discal punctation irregular, much coarser than that of pronotum, most punctures smaller than intervals, punctures near sutural stria arranged in 1 or 2 dense rows. Pygidium extremely finely punctate. Entire ventral side of thorax very finely and sparsely punctate. Metasternal centre flat, lacking impression. Mesocoxal area 0.12 mm long, about as long as half of shortest interval between its margin and metacoxa; marginal pits rather coarse. Metepisternum flat, with straight, deep, punctate suture. Abdominal microsculpture consisting of punctures. Tibia I and III straight, II somewhat curved. Remarks. This species may be readily recognised by its colour pattern. The only other Asian Toxidium which has a well delimited reddish pattern on elytron known to me is an undescribed species from Malaysia. However, it differs conspicuously by much smaller size of the body and by the colour pattern forming a semicircular band. Xotidium gen. n. Type species: Xotidium uniforme sp.n. Gender: neuter. Etymology: anagram of Toxidium. Diagnosis. In general shape similar to Toxidium. Antennal insertion distant from clypeal suture. Mandible multidentate apically. Basal pronotal angles not extended: Elytron with entire basal stria. Metacoxae approximate. Median lobe of aedeagus subcylindrical. 574 IVAN LÖBL Description. Body narrow, similar to that of Toxidium. Pronotum and elytron . not iridescent. Head hypognathous. Eye weakly notched. Antennal insertion about as distant from clypeal suture as from posterior margin of eye. Antenna long, segment III slender, straight, similar to following segments. Mandible multidentate apically, with two comb-hairs, simple prostheca, robust premolar hairs (Fig. 179). Maxillary lacinia very slender, with few lateral hairs, apical hairs absent; galea with marginal rows of robust spine-like hairs (Fig. 186). Segment III of maxillary palpus not thickened, segment IV at base almost as thick as III. Labial palpus 2-segmented, with basal segment subcylindrical, apical segment very slender (Fig. 189). Prothorax with internal setose cavities. Basal angles of pronotum obtuse, not prolongated, not reaching level of anapleural suture. Hypomeron inflexed, not explanate basally. Elytron with entire basal stria joined with lateral and sutural striae; parasutural stria absent. Prosternal keel raised, triangular in lateral view. Mesosternal intercoxal process covering anterior margin of metasternal process. Median mesosternal keel absent. Mesepimeral ridge obsolete. Meso- and metacoxae strongly approximate, median portion of metasternum narrow. First ventrite without pleural stria or ridge. Legs long, meso- and metatibiae with single long apical spur, meso- and metatarsi almost as long as tibiae. Aedeagus with subcylindrical median lobe lacking well delimited apical portion, widened at tip. Parameres with slender basal and wider apical portion. Remarks. The morphological features of this group of species puzzled me for many years. The discovery of an additional Himalayan species made me to reexamine their diagnostic characters. As a result, a new genus is erected. In addition to the type species, the new genus includes Toxidium montanum Löbl and T. pygmaeum Löbl from Sri Lanka, and T. notatum Löbl from Australia (new combinations). To date, I have revised only some of the Afrotropical taxa assigned originaliy to Toxidium: all belong to Scaphobaeocera, Scaphoxium or to an undescribed genus. Hence, the occurrence of Toxidium in Africa has yet to be confirmed. Xotidium may be readily separated from Toxidium and from most species assigned to genera with both, approximate coxae and reduced pubescent of body, by the entire elytral basal stria. Besides Baeotoxidium siamense, only Scaphischema and most of the members of the Toxidium aberrans group exhibit basal stria of elytron, which is usually not joined to the shortened sutural stria (except for that in one undescribed species housed in MHNG). It is probable that this condition is likely apomorphic, while the obtuse, unextended basal angles of the pronotum (shared with Scaphicoma, Toxidium and Scaphoxium) is obviously a plesiomorphic character state. However, both provide useful dichotomous characters. A synapomorphy of Xotidium, Toxidium and Scaphicoma is the position of the antennal insertion; the long tarsi are shared with Scaphicoma, but the shallow eye notch and the presence of the internal prothoracal cavities distinguish the new genus from the others. The mouthparts of Xotidium are unique: the mandible has several apical teeth and bears a premolar group of robust hairs, in combination with the presence of comb-hairs, the maxillary lacinia lacks apical hairs, and the labial palpus is two-segmented, with subcylindrical basal segment and conspicuously thin apical segment. Scaphoxium may be easily separated from Xotidium by the curved 3rd antennal segment, the basally explanate hypomeron, and by the thickened penultimate segment of the maxillary palpus which is much wider at apex than the base of the apical segment. The latter state is found also in Scaphobaeocera and Baeotoxidium (Figs 182, 183). Scaphobaeocera has the elytron almost always microsculptured and iridescent, and exhibits a parasutural stria. Like Baeotoxidium, it also differs from Xotidium by the long, SCAPHIDIIDAE OF NEPAL 575 distinct mesepimeral ridge and by the sutural stria of the elytron not extended along the elytral base. An autapomorphy of Baeotoxidium is the very slender maxillary galea (Fig. 182). KEY TO THE SPECIES OF Xotidium iver body unitormly, brown-or reddish-brown... A eri e 2 SES CAVI STINCHCOLOUT: PALIN re a ATO Ten 3 2 Length 1.10 - 1.25mm. Aedeagus with sinuate, evenly wide flagellum ................... .. Pygmaeum Bi - Length il bi - ne 65mm. ‘Aedeagus with we -shaped, basally thickened flagellum... MOA ee TRE Meret AE EE DARA E II IAS uniforme sp. n. 3 Elytron dark with pale transverse fascia in basal half and with pale apical portion. BroOnotumuMitonmly danke ves. e ae ee sense montanum (Löbl) — Elytron pale, darkened along basal and apical margins, and usually also along sutural margin. Pronotum pale, usually with dark transverse fascia ............ notatum (Löbl) Xotidium uniforme sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Induwa Khola Valley, 2100 m, 17.1V.1984 (Löbl & Smetana) (MHNG). Paratypes, 16: Nepal, Sankhua Sabha distr., Arun Valley, Chichila, 1900-2000 m, bushes near village, 18-20.VI.1988 (Martens & Schawaller) (SMNS, MHNG) 4 males, 5 females; Taplejung distr., SE Yamputhin to Yamputhin, 2000-1650 m, forest mainly Alnus, 26. and 30.1V.1988 (Martens & Schawaller) (SMNS) 1 female; ?Dolakha distr., Jiri - Thorung, 28.V.1976 (Wittmer & Baroni Urbani) (NMB) 1 male; Patan distr., Phulcoki (Franz) (Coll. H. Franz) 1 female; Godawari, 6000 ft. 7-13.VIIL.1967 (Canadian Nepal Exp.) (CNC) 1 male; India, Uttar Pradesh, Kumaon, Rangarh, ca 2000 m, 9.X.1979 (Löbl) (MHNG) 1 female; Himachal Pradesh, 10 km NW Sarahan, 1700 m, 7.X.1988 (Vit) (MHNG) 2 males. Diagnosis. Body relatively large, uniformy brown or dark reddish-brown. Abdomen with microsculpture consisting of punctures. Flagellum of aedeagus U-shaped, thickened at base. Description. Length 1.55 - 1.65 mm, width 0.92 - 1.07 mm. Body uniformly brown or dark reddish-brown. Apical abdominal segments, legs and antennae paler. Relative length of antennomeres as follows: II 23, III 16, IV 19, V 21, VI 21, VII 25, VII 22, IX 27, X 25, XI 28 (holotype). Punctation very fine, that of pronotum, elytron and ventral side of thorax hardly visible at 50x magnification. Sutural area flat. Metasternum lacking median impression. Mesocoxal area 0.05-0.07 mm long, with distinct marginal pits. Metepisternum evenly wide, with straight or somewhat sinuate suture. Abdominal microsculpture consisting of extremely fine punctures. Male protarsus hardly widened. Aedeagus (Figs 170 to 172) 0.67 - 0.80 mm long. Widened apical portion of parameres long. Flagellum of internal sac U-shaped, very thin, thickened basally. Remarks. This species may be readily distinguished by the characters used in the key. 576 IVAN LÖBL Bironium Csiki Bironium Csiki 1909; type species: Bironium longipes Csiki, 1909, by monotypy (longipes Csiki is secondary junior homonym of longipes Reitter, 1880; it is replaced by the juniow subjective synonym Heteroscapha basicolle Pic, 1956). Heteroscapha Achard, 1914; type species: Heteroscapha feai Achard, 1914, by monotypy. Scutotoxidium Pic, 1915; type species: Scutotoxidium nigrolineatum Pic, 1915. Arachnoscaphula Heller, 1917; type species: Arachnoscaphula trisulcata Heller, 1917. This genus is well characterized by the combination of the following characters: antennae, femora and tibiae conspicuously long; antennal insertion situated near upper eye margin; hypomeron vertical; basal pronotal angle rounded, at level of centre of mesepisternum; meso- and metacoxae distant. Most of the 25 recognised species exhibit conspicuous elytral and metasternal punctation, and all of them have elytron with entire basal stria. The range of the genus covers south east Asia and New Guinea. The sole record for the Himalayan region is that of a female specimen of Heteroscapha distinctum Achard (CHAMPION 1927: 272) from “Gopalhara, Darjeeling”. Bironium distinctum is known from Burma and Thailand, and may occur in India. It belongs to a group of which members may be reliably identified only by the male sexual characters. Thus, Champion’s record has to be confirmed. An additional, undescribed species has been found in Eastern Nepal. Bironium nepalense sp. n. Holotype, male: Nepal, Sankhua Sabha distr., Arun Valley, Chichila, 2200 m, 24.IV.1984 (Löbl & Smetana) (MNHG). Paratypes, 2 females: as holotype (MHNG) 1; Chichila, 1900-2000 m, bushes near village (Martens & Schawaller) (SMNS) 1. Diagnosis. Medium-sized species, elytron maculate, vaulted between two discal rows of coarse punctures. Lateral portion of metasternum coarsely punctate. First ventrite with medio-basal protuberance; tibiae striate. Tip of aedeagus inflexed, parameres slender. Description. Length 2.6 - 2.7 mm, width 1.7 - 1.3 mm. Head and pronotum dark reddish-brown, most of pronotal disc darker than head. Elytron blackish-brown or black near base, along sutural stria, at middle, and near apical margin; with oblique reddish fascia on anterior half and with large ochreous spot covering most of apical half. Hypomeron, prosternum and mesosternum reddish, remaining ventral surface of thorax blackish. Basal abdominal ventrites very dark redish brown to blackish, apical segments paler. Antennae and legs reddish or ochreous. Relative length of antennomeres as follows: II 26, III 44, IV 58, V 50, VI 62, VII 64, VII 68, IX 58, X 50, XI 52 (holotype). Pronotum uniformly, very finely punctate, not microsculptured. Exposed portion of scutellum fairly large. Elytron with deep, coarsely punctate sutural stria; sutural area raised; humeral protuberance absent, most of discal surface impunctate or extremely finely punctate; irregular coarse punctation situated near base and on anterior half of lateral portion; two regular long discal rows of coarse punctures lying in depressions separated by vaulted interval, additional short row of coarse punctures situated between sutural stria and inner long row. Propygidium and pygidium with microsculpture consisting of punctures. Hypomeron glabrous, without microculpture. Mesepisternum extremely finely punctate. Mesosternum convex, lacking median ridge, with a pair of large apical punctures, elsewhere extremely finely punctate, not striate. Center of metasternum impressed. SCAPHIDIIDAE OF NEPAL 577 Median metasternal portion with dense transverse row of fairly coarse punctures and two irregular longitudinal rows of similarly coarse punctures. Lateral portion of metasternum very coarsely punctate. Mesepisternal suture deep, coarsely punctate. Ventrites with microsculpture consisting of punctures. Abdominal punctation extremely fine, first ventrite with conspicuous medio-basal protuberance; basal pits coarse, inner larger than outer ones. Tibiae striate, almost straight. In male ventrites without appreciable male sexual characters. Segments 1 to 3 of protarsus distinctly widened, segment 1 somewhat narrower than apex of tibia, following two gradually narrower. Aedeagus (Figs 174 to 176) 0.81 mm long. Median lobe with long, gradually narrowed apical portion; tip inflexed. Parameres almost straight, narrowed subapically, then weakly widened. This species may be readily distinguished from other members of the genus by its colour pattern in combination with the elytral punctation. It resembles B. distinctum and its allies in many characters but differs drastically in having striate tibiae, a large medio- basal protuberance on Ist ventrite, long and inflexed apical portion of median lobe and slender parameres. Mystrix Champion Mystrix Champion, 1927; type species Mystrix termitophilum Champion, 1927, by monotypy. Mystrix includes two species, M. termitophilum known only from the “Haldwani Division of Kumaon” and M. kistneri Löbl from Sumatra. M. termitophilum is the sole member of the termitophilous “Baeoceritae” (ACHARD 1924a). in the Indian sub-continent. The genus may be easily distinguished from other Himalayan scaphidiid genera by the conspicuously pubescent body and flattened femora, and by the combination of the entire eye, the extremely small antennomeres III and IV, the ventrally strongly vaulted body, and the distinct metacoxal lines. A new record of M. termitophilum is a specimen from Nepal, Parsa distr., Birganj Lothar, 450 ft., 13-19.1X.1967, malaise trap (Canadian Nepal Exp.) (CNC). BIOGEOGRAPHY The very diverse Himalayan flora has been characterized in SCHWEINFURTH (1957), and analysed in STAINTON (1972). It belongs, according to MEUSEL & al. (1965) and MEUSEL & SCHUBERT (1971), to the Sino-Japanese phytogeographic realm. DOBREMETZ (1972) found the Himalayan flora distinct from that of the latter, as well as from that of the Indian and Central Asian realms. TROLL (1967) distinguishes five main divisions, viz. the Assam-, Sikkim-, Garhwal-, Punjab-, and Indus-Himalaya. DOBREMETZ (1972) recognised three phytogeographic subregions, the East Himalayan (“Assam Himalaya” i.e., actual Arunachal Pradesh, and most of Bhutan), the West Himalaya including Kashmir and a part of Hindu Kush, and the Central Himalaya which is a transitional area including Garhwal, Kumaon, Nepal, Sikkim, and parts of Bhutan. Four areas are distinguished within Nepal (DOBREMETZ 1972, MARTENS 1979): the relatively narrow Eastern Nepal (between Sikkim and Arun Valley, with moist monsoon climate and absence of xerophilic vegetation), Central Nepal (between Arun Valley and Daulagiri, with less moist monsoon climate in which hygrophilic, mesophilic and xerophilic vegetation co-exist, possibly best characterized by the oak forests (Quercus seme- carpifolia, Q. lanuginosa)), Western Nepal (with dryer seasonal climate and mesophilic and xerophilic vegetation (with Quercus incana, Cedrus deodara, Abies pindrow, Olea 578 IVAN LÖBL cuspidata)), and North-Central Nepal (in rain shadow, north of Daulagiri, Annapurna, Manaslu, etc., with highland steppe vegetation). The high biodiversity of the Himalaya may be explained by both historical and ecological factors. It is a transitional zone inhabited by taxa of southern (Indian), eastern (Chinese, Burmese) and western (Mediterranean, Central Asian) origin (MAnı 1974, MARTENS 1979). An other conspicuous pattern of the Himalaya is the altitudinal range of the vegetation reaching over 6000 m, forming 10 (DOBREMETZ 1972) or 11 (MARTENS 1979) zones. The taxa of presumably southern and eastern origin become gradually less frequent westward and are confined to lower altitudes. The available data on Agathidiini (Leiodidae) and on Scaphidiidae are possibly significant for Himalayan fungivorous beetles. The Agathidiini are a predominantly Holarctic group with more than 400 recognised species. Only a smaller portion of them extends to the tropics or subtropics of south and southeastern Asia (Angelini, pers. commun.). The myxomycetophagous Agathidium species are extremely diverse in the Himalaya where about 110 species have been encountered (LAWRENCE 1989). Most of them appear to be endemic in some parts of the Himalaya, including its western portion and the Hindu Kush - Kohistan ranges. The Scaphidiidae, with roughly 1300 known species, feed on Myxomycetes and on a variety of other fungi. They are most speciose in areas with moist, warm climate, and poorly represented in areas with temperate or dry climate. Nevertheless, like the Agathidiini they are remarkably diverse in the Himalaya. The examined collections include 206 identified scaphidiid species in 18 genera (including those from Meghalaya), and several additional species which have not been identified. However, their distributional pattern is different from that of the Agathidiini. No endemic species occurs in the Hindu Kush - Kohistan ranges, nor in Karakorum and in Kashmir, although 16 species were collected there. Two of them, Scaphium quadraticolle Solsky and Scaphisoma assimile curvistria Reitter, also occur north and northwest of Hindu Kush and Karakorum. All others species were found further eastward, most of them east of the Sutlej river, and their origin is presumably southern or eastern. The number of the species (only the identified ones taken in account), and of the apparently endemic species per area increases from west to east: 35 species with 2 (5.7%) endemics in Himachal Pradesh, 58 species with 6 (10.3%) endemics in Garhwal and Kumaon, 129 species with 38 (29.5%) endemics in Nepal. In the Darjeeling district in West Bengal, including the unrepresentative data from Sikkim and Bhutan, 80 species with 18 (22.5%) endemics have been found. It is hazardous to compare these numbers with those obtained from collections made in 4 days of field work in the Manas National Park in Assam (25 species, with 6, e.g., 24% endemics) or in 11 days of field work in the Khasi and Garo Hills in Meghalaya (where 61 species, with 19, e.g., 31.1% endemic were found, see LOBL 1984; 1986a). The data from Meghalaya and from the Manas National Park are almost exclusively based on material collected by C.Besuchet and the present author on a single trip (see Löbl 1986a) while those from Nepal, Uttar Pradesh and Himachal Pradesh are from the field work of several entomologists in different seasons and during longer periods. Thus, the distributional ranges of the taxa and their endemism as quoted in this paper do rather reflect the effort in field work invested for a particular area than a biological reality. Nevertherless, the data suggest a surprisingly high diversity of the Meghalayan fauna and indicate the following distributional pattern. As in many other groups (i.e., Collembola: Neanuridae, CASSAGNAU 1981; Staphy- linidae: Quediini, SMETANA 1988), the diversity of scaphidiids in general decreases from the east to the west. SCAPHIDIIDAE OF NEPAL 579 According to MANI (1968), nearly 60% of the high altitude beetle species are endemic to the Himalaya and 96% of them are Palaearctic forms. This is certainly not true for the scaphidiids: none of the Nepalese species may be derived from the West Palaearctic stock, and only few (e.g., Scaphidium biundulatum, S. nepalense, S. gurung, Scaphisoma species assigned to the subalpinum group) are probable derivates of the temperate Sino-Japanese fauna. Scaphidiids exhibit high dispersal ability. From the 47 species recorded in South India, 20 (42.6%) occur in the Himalaya, and from the 61 species found to date in Meghalaya 41 (67.2%) occur also in the Himalaya. Actually, 206 species have been identified from the Himalaya and Meghalaya, 55 of these species (26.7%) extend their distributional range eastward beyond political border of India and/or to south India, and 16 species (7.8%) west- and/or southwestward. Collections made in Phulcoki indicate a higher species diversity in the Mahabharat range than in the Inner Himalaya, at approximately the same meridian, and at altitudes ranging from 1600 to 2600 m. Most of the 38 species of scaphidiids inhabiting Phulcoki were found in oak forest, at 2000 to 2600 m altitude. The rate of the endemic scaphidiid species of Phulcoki (Scaphisoma baloo only) is very low in comparision with collections from the Inner Himalayan ranges. It is difficult to speculate on biogeography when phylogeny of the group has not been analysed and the distribution of the taxa is known only partly. However, allopatric speciation may be supposed in groups of closely related vicarious species, such as Scaphidium gurung and S. nepalense, Baeocera microptera and B. puncticollis, or Scaphisoma immodicum and S. bhareko. The number of apparently endemic species in some areas, as in the upper Kali Gantaki Valley or at Ghoropani Pass is relatively high, but little or no information is available from nearby Daulagiri or from the southern slopes of Annapurna. ACKNOWLEDGEMENTS Nicolette Lavoyer made most of the line drawings during her spare time. The author is greatly-indebted to her, and to Ales Smetana for reading and commenting an earlier draft of the manuscript, and to Neil D. Springate for assistance with the English text. The type material was made available through the kindness of Miss N. Berti (Paris), Miss E. De Boise (London), J. Jelinek (Prague), and R. Poggi (Genova). Additional material was obtained from V. Brachat (Miinchen), M. Brancucci (Basel), P. Cassagnau (Toulouse), L. Deharveng (Toulouse), H. Franz (Mödling), S. Hisamatsu (Matsuyama), K. Holzschuh (Wien), J. Jelinek (Prague), J. Martens (Mainz), J. Pawlowski (Krakow), G.M. de Rougemont (London), M. Sakai (Shigenobu), W. Schawaller (Stuttgart), A. Smetana (Ottawa), S. Vit (Geneva), and W. Wittmer (Basel). Their assistance is greatly appre- ciated. Several of these colleagues generously donated specimens from their own collections. I thank all of them most cordially. 4 (a ‘© pas) Z < Ri 580 Sicariciieiatia ihr Fic. 9. Ascaphium ochripes sp. n. SCAPHIDIIDAE OF NEPAL 581 Fics 10 To 13. Colour pattern of elytra in Scaphidium; 10. S. biundulatum Champion; 11. S. fryi Achard; 12. S. in- crassatum Achard; 13. S. nepalense sp. n. 582 IVAN LÖBL Fics 14 To 16. Colour pattern of elytra in Scaphidium; 14. S. gurung sp.n.; 15. S. sylhetense Achard; 16. S. harmandi Achard. SCAPHIDIIDAE OF NEPAL 583 ï } ¢ 2". à r i li i U N Fics 17 TO 20. Colour pattern in Scaphidium; 17. S. sinense Pic; 18. S. semilimbatum Pic; 19. S. rubritarse Pic; 20. S. coomani (Pic). IVAN LÖBL Fics 21 TO 24. Colour pattern in Scaphidium und Hemiscaphium; 21. S. septemnotatum Champion, 22. Scaphidium sp.; 23. S. baconi Pic; 24. H. brunneopictum Achard. 585 SCAPHIDIIDAE OF NEPAL Fics 25 To 28. iatum Champion: FUC ISIC 26 3 25. P. bicolor sp.n. S. aurorae sp.n.; 28. S. baloo sp. n. onium and Scaphisoma Colour pattern in Pseudobir 27 IVAN LÖBL Fics 29 To 32 32% ium Löbl]; _S. var eko Sp. n.: 31 caphisoma; 29. S. clavigerum sp. n.; 30. S. bhar Colour pattern in S S. pulchellum Löbl. SCAPHIDIIDAE OF NEPAL Fics 33 TO 36. Colour pattern in Scaphisoma; 33. S. quadrifasciatum Löbl; 34. S. tetrastictum Champion; 35. S. leu- copyga Champion; 36. $. binhanum (Pic). 588 IVAN LÖBL Fics 37 AND 38. Colour pattern in Scaphisoma, 37. S. maculiger Löbl; 38. S. notatum Löbl. 589 SCAPHIDIIDAE OF NEPAL Fics 39 To 47. Achard; 41. S. 43. S. coomani (Pic); 44. S. septemnotatum Champion; 45. Aedeagi in Scaphidium and Hemiscaphium; 39. S. grande Gestro; 40. S. harmandi rubritarse Pic; 42. S. cyanellum Oberthur; S. cinnamomeum Champion; 46. H. brunneopictum Achard; 47. S. baconi Pic. IVAN LÖBL 590 Fics 48 TO 54. epalense sp. n.; ylhetense Achard; 54. S. holzschuhi sp. n. 3 0) 85 1% ndulatum Champion; 49. S. gurung Sp. n. . S. thakali sp. n.; 53. S.s 2 riatum Champion; 52 = Aedeagi in Scaphidium; 48. S. biu 51. S. biser SCAPHIDIIDAE OF NEPAL 591 56 59 60 57 63 66 67 64 65 62 Fics 55 TO 67. Parameres in Scaphidium and Hemiscaphium; 55. S. grande Gestro; 56. S. gurung sp. n.; 57. S. nepa- lense sp. n.; 58. S. biseriatum champion, 59. S. sylhetense Achard; 60. S. thakali sp. n.; 61. S. harmandi Achard; 62. S. rubritarse Pic; 63. S. cinnamomeum Achard; 64. S. coomani (Pic); 65. S. septemnotatum Champion; 66. S. melanogaster sp. n.; 67. H. brunneopictum Achard. Scale bar = 0.1 mm. IVAN LÖBL 592 Fics 68 TO 71. Internal sac of aedeagi in Scaphidium; 68. S. biundulatum Champion; 69. S. nepalense sp. n.; 70. S. gu- rung sp. n.; 71. S. grande Gestro. Scale bar = 0.1 mm. SCAPHIDIIDAE OF NEPAL 593 Fics 72 TO 75. Internal sac of aedeagi in Scaphidium; 72. S. harmandi Achard; 73. S. rubritarse Pic; 74. S. cya- nellum Oberthur; 75. S. coomani (Pic). Scale bar = 0.1 mm. 594 IVAN LÖBL Fics 76 TO 79. Internal sac of aedeagi in Scaphidium; 76. S. biseriatum Champion; 77. S. thakali sp. n.; 78. S. holz- schuhi sp. n.; 79. S. sylhetense Achard. Scale bar = 0.1 mm. SCAPHIDIIDAE OF NEPAL 595 Fics 80 To 84. Internal sac of aedeagi in Scaphidium and Hemiscaphium; 80. S. septemnotatum Champion; 81. S. ba- coni Pic; 82. S. melanogaster sp. n.; 83. S. cinnamomeum Champion; 84. H. brunneopictum Achard. Scale bar = 0.1 mm. 596 IVAN LÖBL Fics 85 TO 87. Aedeagi in Cyparium, 85. C. montanum Achard: 86. C. bowringi Achard; 87. C. plagipenne Achard. Scale bar = 0.2 mm. 597 SCAPHIDIIDAE OF NEPAL Fics 88 To 91. Aedeagi in Pseudobironium; 88. P. bicolor sp. n., 89. P. almoranum Achard; 90, 91. P. ineptum sp. n. Scale bar = 0.2 mm. 598 IVAN LÖBL Fics 92 To 96. Aedeagi in Pseudobironium; 92 to 94. P. rufitarse sp. n., apical portion of internal sac (93) in detail; 95, 96. P. castaneum Pic, tuberculate margin of internal sac (96) in detail. Scale bar = 0.2 mm (92, 95), and = 0.1 mm (96). 599 SCAPHIDIIDAE OF NEPAL pp Oe Fics 97 To 102. Aedeagi in Baeocera. 97, 98. B. sordidoides sp. n.; 99, 100. B. laminula sp. n., internal sac (100); 101, 2). 0.1 mm (100, 10 102. B. reducta sp. n., internal sac (102). Scale bar = 0.2 mm (98, 99, 101), and 600 IVAN LÖBL 105 108 Fics 103 To 108. Aedeagi in Baeocera; 103 to 105. B. crinita sp. n., internal sac (104), paramere (105); 106 to 108. B. cri- brata sp. n., internal sac (107), paramere (108). Scale bar = 0.1 mm (104, 105, 107, 108), and = 0.2 mm (103, 105). SCAPHIDIIDAE OF NEPAL 601 SS KE 22 N NN ITS SI (E | Fics 109 To 113. Aedeagi in Baeocera; 109 to 111. B. martensi sp. n., internal sac (110), paramere (111); 112 and 113. B. schawalleri sp. n., internal sac with apical portion of median lobe and parameres (113) at higher magnification. Scale bar = 0.1 mm (110, 111), = 0.2 mm (109, 113), and 0.3 mm (112). 602 IVAN LÖBL 118 Fics 114 To 119. Baeocera; 114 and 115. B. mustangensis sp. n., aedeagus; 116 to 119. B. thoracica sp. n., basal portion of male metatibia (116), aedeagus (117), apical half of median lobe and parameres (118, 119). Scale bar = 0.2 mm (114 to 116, 118, 119), and = 0.3 mm (117). SCAPHIDIIDAE OF NEPAL 603 120 7 Bo Ga 7 7 VÀ > Ae A - 7 A = Can 47 27 e —- a Pi LIT n= CRE a -_— TL 4-7 -) VL a SS — 1.7 ESS N N VAS \ SES (y SSA J / \ IL NY POS = Ns _- SS I a (7 Fics 120 To 122. Aedeagus in Baeocera tuberculata sp. n., apical portion of median lobe and parameres (121, 122) at higher magnification. Scale bar = 0.2 mm. 604 IVAN LÖBL Fics 123 To 125. Aedeagus in Baeocera errabunda sp. n., apical portion of median lobe and parameres (124, 125) at higher magnification. Scale bar = 0.2 mm (124), and = 0.3 mm (123). SCAPHIDIIDAE OF NEPAL 605 Fics 126 TO 128. Aedeagi in Scaphisoma; 126. S. minutissimum Champion; 127 and 128. S. praesigne sp. n. Scale bar = 0.1 mm (126), and 0.2 mm (127). 606 IVAN LÖBL RS 1 \ Sy 131 Fics 129 To 131. Aedeagi in Scaphisoma; 129. S. fulcratum sp. n.; 130. S. interjectum sp. n.; 131. S. simplicipenis sp. n. Scale bar = 0.1 mm. SCAPHIDIIDAE OF NEPAL 607 N Ss SSIIIS> = =7 SSS Fics 132 To 134. Aedeagi in Scaphisoma; 132. S. simpicipenis sp. n.; 133. S. fatuum sp. n.; 134. S. adjacens sp.n. Scale bar = 0.1 mm. 608 IVAN LÖBL - Fics 135 To 137. Aedeagi in Scaphisoma; 135. S. inquietum sp. n.; 136. S. fratellum sp. n.; 137. S. aurorae sp. n. Scale bar = 0.1 mm. 609 CA Wh, NA DS : Hig ING Wir = (A (1 i TI - rd < ji La SCAPHIDIIDAE OF NEPAL Fics 138 To 140. Aedeagi in Scaphisoma; 138. S. aurorae sp. n.; 139. S. nima sp. n.; 140. S. jado sp. n. Scale bar 0.1 mm. IVAN LÖBL 610 TR EEE | “a ay ERS Fics 141 TO 143. o sp. n.; 142 and 143. S. invalidum sp. n. Scale bar = Aedeagi in Scaphisoma; 141. Scaphisoma jad 0.1 mm. SCAPHIDIIDAE OF NEPAL 611 Fics 144 To 146. Aedeagi in Scaphisoma; 144 and 145. S. baloo sp. n.; 146. S. clavigerum sp. n. Scale bar = 0.2 mm. IVAN LÖBL 612 149 Fics 147 TO 149. Aedeagi in Scaphisoma; 147 and 148. S. clavigerum sp. n., paramere with apical portion of median lobe (147); 149. S. assimile curvistria Reitter. Scale bar = 0.2 mm. SCAPHIDIIDAE OF NEPAL 613 Fics 150 To 152. Aedeagi in Scaphisoma; 150 and 151. S. championi Löbl; 152. S. pinnigerum sp. n. Scale bar = 0.2 mm. IVAN LÖBL 614 155 56 1 Fics 153 To 156. 0.2 mm. Aedeagi in Scaphisoma; 153. S. alacre sp. n.; 154 and 155. S. nepalense sp. n., paramere (155) in ventral view; 156. S. kanchi sp. n., paramere in ventral view. Scale bar SCAPHIDIIDAE OF NEPAL 615 Za Ging GF A tota . 32° IS SON S > son, 5 > è: se Fics 157 To 159. Aedeagi in Scaphisoma; 157. S. kanchi sp. n.; 158 and 159. S. coalitum sp. n., internal sac (159) in detail. Scale bar = 0.2 mm. 616 IVAN LÖBL 160 161 162 Fics 160 To 162. Aedeagi in Scaphisoma; 160. S. coalitum sp. n., parameres, 161 and 162. S. sikkimense Sp. n., paramere (162) in ventral view. Scale bar 0.2 mm. U —————1 SCAPHIDIIDAE OF NEPAL 617 N A ahh 17. Fics 163 To 165. Scaphisoma bhareko sp. n., aedeagus; internal sac (164) in detail. Scale bar 0.1 mm (164) and 0.2 mm (163). 618 IVAN LÖBL Br = In, 25 169 Fics 166 To 169. Aedeagi; 166. Baeotoxidium yeti sp. n., aedeagus with extruded internal sac; 167. Scaphoxium eximium Löbl, internal sac; 168, 169. Toxidium curtilineatum Champion, detail of internal sac (169). Scale bar = 0.1 mm (166, 167, 169) and 0.2 mm (168). SCAPHIDIIDAE OF NEPAL 619 172 Fics 170 To 172. Xotidium uniforme sp. n., aedeagus; apical portion with extruded internal sac (171). Scale bar = 0.1 mm. IVAN LÖBL 620 175 Fics 173 TO 175. Aedeagi; 173. Scaphicoma arcuatum (Champion); 174-176. Bironium nepalense sp. n., apical portion 0.2 mm (173-175) and 0.1 mm (176). of median lobe (176). Scale bar 621 SCAPHIDIIDAE OF NEPAL Wie 77/25, a EG 4 LEE - \\ Z My \ LA È N GF bruni > UO 77 r = “ry 2, i] / y} oe % 2s GONE lan 1 ° tj = e DE e N ara ANTONINI Lernen L N RN Fics 177 To 181. uniforme sp. n.; 180. Baeotoxidium lanka Löbl; 181. Scaphobaeocera japonica (Reitter). Scale bar 0.1 mm. Mandibles; 177. Scaphicoma arcuatum (Champion); 178. Toxidium vagans Löbl; 179. Xotidium 622 IVAN LÖBL Fics 182 To 186. Maxillae: 182. Baeotoxidium lanka Löbl; 183. Scaphobaeocera japonica (Reiter), 184. Toxidium gammaroides LeConte; 185. Toxidium vagans Löbl; 186. Xotidium uniforme sp. n. Scale bar = 0.1 mm. SCAPHIDIIDAE OF NEPAL 623 \ Fics 187 To 191. | Mouthparts. 187, 188. Scaphicoma arcuatum (Champion), maxilla and labium with palpi; 189. Xotidium uniforme sp. n., labial palpi with hypopharynx; 190. Toxidium gammaroides LeConte, i labium with palpi and hypopharynx; 191. Baeotoxidium lanka Löbl, labium with palpi and hypopharynx. 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A l l | Revue suisse Zool. | Tome 99 | Fasc. 3 | | | p. 629-636 | Genève, septembre 1992 | Morphometric data on the scapula and limb long bones of Arvicola terrestris (Linnaeus, 1758) (Rodentia, Arvicolidae) by Jacint VENTURA* ABSTRACT Data on the morphometry of the scapula and limb long bones of the northern water vole, Arvicola terrestris, according to sex and relative age are given. The sample was formed by 209 specimens (104 males and 105 females) captured in the Spanish Pyrenees. The whole sample was divided into six classes of relative age. The width and functional length of the scapula and limb long bones were measured in each specimen. The results indicate a notable biometric uniformity in the scapula and limb long bones between males and females. The variables with the highest growth rate are the width and length of the scapula. The scapula and limb long bones show their maximum length increase before fourteen weeks of age. The width of the scapula shows a positive allometry against its length up to approximately the first year of age. Thereafter, this relation is isometric. The widths of the humerus and femur show a positive allometric growth against their respective lengths during adulthood. The growth rates of the length of the hind limb bones are higher than those of the forelimb. INTRODUCTION In the large Palaearctic distribution area of Arvicola terrestris, two eco-ethological types of populations are known: the digging populations from central and southwestern Europe, and the aquatic populations found in the remainder of the distribution area. These latter populations show similar habitat requirements and behaviour to the southwestern water vole, Arvicola sapidus, distributed in France and the Iberian Peninsula. In eastern Europe other populations with double habitat preferences (underground or aquatic according to the season) are present (KRATOCHVIL 1983). In the Iberian Peninsula there are only strictly digging populations. Although the information published on A. terrestris is abundant (REICHSTEIN 1982), the bibliographic data on the morphology and morphometry of the girdles and limb skeleton are relatively scarce (BROWN & TwiGG 1969; ScHicH 1971; Bou & CASINOS 1985; Bou et al. 1987; LAVILLE et al. 1989; VENTURA et al. 1991). Likewise, the | information on relative growth of postcranial bones is reduced to the results reported by * Departamento de Biologia Animal (Vertebrados), Facultad de Biologia, Universidad de Barcelona, Avda. Diagonal, 645, 08028-Barcelona, Spain. 630 JACINT VENTURA BROWN & Twicc (1969) and VENTURA ef al. (1991) on the os coxae. The aim of this study is to give biometrical data, according to sex and relative age, on the scapula and limb long _bones of an Iberian population of A. terrestris. MATERIAL AND METHODS The sample analysed was formed by 209 specimens (104 males and 105 females) captured between 1983 and 1984 in the Spanish Pyrenees (Aran Valley, Lérida). The specimens were distributed into six classes of relative age (O-V) according to the type of coat and stage of moult, morphological characteristics of the skull and sexual stage (VENTURA & GOSALBEZ 1990). The intervals of age corresponding each category are as follows (VENTURA 1988): class O: 0-3 weeks; class I: 3-6 weeks; class II: 6-10 weeks; class III: 10-14 weeks; class IV: specimens older than 14 weeks, but before the end of their first winter; class V: specimens that have wintered at least once. The following measurements were taken (VENTURA 1990): SW: width of the scapula; SL: functional length of the scapula; HW: width of the humerus; HL: functional length of the humerus; UL: functional length of the ulna; RL: functional length of the radius; FW: width of the femur; FL: functional length of the femur; TL: functional length of the tibia. All measurements were taken with vernier calipers. The following indices were considered (VENTURA 1990): SI = (SW/SL)%; HI = (HW/HL)%; FI = (FW/FL)%; LI = [(HL+RL) / (FL+TL)] %. The variation percentages between the means of consecutive age classes were calculated by the formula (VENTURA 1990) VP =[(x;,1 -X;) + 100] /x;, where x; and X;,j are the averages corresponding to the age classes considered. The significance of the differences between two sample means was calculated by the Student's t-test (SOKAL & ROHLF 1981). With the logarithmic values of the data, regressions were calculated by the least squares method (SOKAL & ROHLF 1981). The significance of the differences between two regression coefficients was determined by the comparison of their confidence intervals at 95%. RESULTS The intersexual comparisons of the averages of each parameter in all age classes reveal the existence of a high biometric uniformity between males and females. Only significant differences are present in UL in classes IV (males: x=19.03, sd=0.55, n=19; females: x=18.65, sd=0.52, n=22; t=2.2719, p<0.05) and V (males: x=19.61, sd=0.51, n=17; females: x=19.22, sd=0.55, n=16; t=2.1136, p<0.05) and in RL in class I (males: x=15.17, sd=0.26, n=6; females: x=14.61, sd=0.35, n=9; t=3.3369, p<0.01). In the indices considered, only SI in class 0 (males: x=68.17, sd=1.16, n=4; females: x=61.96, sd=4.66, n=4; t=2.5863, p<0.05) and LI in classes I (males: x=83.91, sd=1.84, n=3; females: x=86.38, sd=1.49, n=8; t=2.3171, p<0.05) and IV (males: x=82.83, sd=1.7, n=20; females: x=81.21, sd=1.43, n=22; t=2.1108, p<0.05) show significant differences. From these results, the data obtained in males and females were treated together (table 1). With the aim of determining the growth rate of all the variables considered, each one was correlated against the body weight, which was used as an approximate indicator of age (PELIKAN 1972; VENTURA & GOSALBEZ 1990). The confidence intervals of the regression coefficients (b) reveal that, except between HW and FL, and UL and RL, the MORPHOMETRIC DATA OF ARVICOLA TERRESTRIS 631 TABLE 1. Values of the measurements and indices taken in the scapula and limb long bones in Arvicola terrestris, according to the relative age. Variable Age n x sd min. max. or index class SW 0 8 6.52 0.53 5.6 7.0 I Bi 7.85 0.58 6.8 9.0 II 42 9.30 0.52 7.9 10.7 II 42 10.64 0.69 9.4 12.0 IV 43 12.23 0.78 10.4 13.8 V 41 13.13 0.68 11.6 14.7 SL 0 8 10.04 0.57 8.7 10.6 I 31 11.58 0.73 10.2 12.9 II 42 13.64 0.63 12.3 14.8 II 42 15:55 0.74 14.0 15.9 IV 43 1721 0.79 152 18.8 V 41 18.46 0.64 17.0 19.8 HW 0 9 1.43 0.11 12 1.6 I 31 1.52 0.08 1.4 1.7 II 42 1.68 0.09 15 1.9 III 42 1.85 0.11 1.6 2.1 IV 43 2.05 0.13 1.8 23 V 41 2.25 0.15 2.0 2.6 HL 0 7 13.54 0.88 11.7 14.8 I 31 15.19 0.73 13.5 16.7 II 42 17.01 0.63 15.5 18.7 III 42 18.51 0.67 16.8 20.3 IV 43 19.73 0.66 18.2 21.3 V 41 20.63 0.70 18.6 22.0 UL 0 6 14.38 0.70 13.1 15.2 I 14 15.66 0.44 15.0 16.4 II 26 16.80 0.69 15.7 19.3 II 36 17.80 0.59 16.7 19.0 IV 41 18.83 0.56 Well 20.0 V 33 19.42 0.56 18.3 20.4 RL 0 6 13.62 0.76 12.3 14.7 I 15 14.81 0.40 14.0 15.4 II 37 16.03 0.48 15.0 16.7 II 38 17.12 0.50 16.0 18.4 IV 42 18.00 0.54 16.8 19.0 V 40 18.63 0.51 17.6 19.8 FW 0 9 1.81 0.18 15 2.0 I 31 2.07 0.15 1.8 2.4 II 42 2.35 0.13 Dall 2.6 II 42 2.58 0.76 2.0 3.0 IV 43 2.89 0.20 25 3.4 V 41 3.07 0.21 2:7 3.4 FL 0 U 14.07 0.81 12.3 15.0 I 29 15.74 0.82 14.3 172 632 JACINT VENTURA TABLE 1. (continuation) Variable Age n x sd min. max. “or index class III 42 20.03 0.76 18.0 21.8 IV 43 21.69 0.75 20.0 23.0 V 41 23.04 0.75 ALT 24.9 TL 0 6 17.70 0.32 17.4 18.3 I 23 19.17 0.94 Ion 21.2 II 34 21.62 0.78 20.2 23.4 III 37 23.12 0.87 20.1 24.4 IV 43 24.49 0.78 21.9 25.7 V 40 25.83 0.83 24.4 Dal! SI 0 8 65.07 4.61 53.37 70.10 I 31 67.82 2.68 61.95 73.73 II 42 68.17 1.79 58.09 72.66 III 41 68.40 2.39 63.51 123 IV 42 70.94 2.67 64.07 76.40 V 41 12 2.98 64.40 79.00 HI 0 7 10.65 0.66 9.70 11.85 I 31 10.03 0.51 9.26 11.27 II 42 9.89 0.55 8.52 11.61 IH 42 9.98 0.48 9.04 11.11 IV 43 10.41 0.62 9.04 11.70 V 41 10.92 0.68 9.52 12.63 FI 0 6 13.17 0.88 12.19 14.28 I 29 13.16 0.64 11.73 14.46 II 42 13.08 0.72 11.50 14.77 III 42 12.89 0.79 9.52 14.35 IV 43 13.31 0.89 11.36 15.53 V 41 13.32 0.85 11.54 14.80 LI 0 4 86.97 1.31 85.26 88.59 I 11 85.70 1.93 81.32 87.88 II 30 83.56 123 80.67 85.71 III 33 82.37 1.38 79.38 86.32 IV 42 81.69 1.65 77.20 88.10 V 39 80.33 1.26 77.40 82.63 slopes of the remaining regression are significantly different (table 2). SW is the variable that shows the highest slope against body weight. The length of the proximal bones of each limb shows higher regression coefficients than that of the distal ones. Moreover, the femur shows a higher growth rate than the humerus, and the tibia grows faster than the ulna and radius (table 2). The lengths of the scapula and limb long bones generally show a progressive diminution of the variation percentages of the means between consecutive age classes (table 3), higher values appearing in 0-I (HL, UL and RL) or in I-II (SL, FL and TL) intervals. From this last interval, the variation rates diminish until class V. The variation percentages of the widths of the scapula, humerus and femur decrease from the lowers (O- | | MORPHOMETRIC DATA OF ARVICOLA TERRESTRIS 633 TABLE 2. Values of the regression equations (log y = b log x + log a) for the correlations between body weight (x) and all the measurements considered (y). Confidence intervals of b (95%) y loga b r n minor major SW 0.0513 0.5388 0.5153 0.5623 0.9528 207 SE 0.2233 0.4810 0.4646 0.4974 0.9702 207 HW —0.4952 0.3845 0.3622 0.4068 0.9207 208 BE 0.6138 0.3241 0.3119 0.3363 0.9643 206 UL 0.7915 0.2290 0.2146 0.2434 0.9288 156 RL 0.7544 0.2378 0.2248 0.2508 0.9375 178 FW —0.4188 0.4168 0.3949 0.4387 0.9337 208 EE 0.5176 0.3894 0.3757 0.4031 0.9689 204 THE 0.7700 0.2956 0.2816 0.3096 0.9509 183 TABLE 3. Variation percentages between the means of consecutive age classes in the measurements and indices considered. Variable or index 0-I I-II II-II III-IV IV-V SW 20.40 18.47 14.41 14.94 7.36 SE 15.34 17.79 12.28 10.67 7.26 HW 6.30 10.53 10.11 10.81 9.76 HL 12.19 11.98 8.82 6.59 4.56 UL 8.90 7.28 5.95 5.79 3.13 RL 8.74 8.23 6.80 5.14 113.50 FW 14.36 13.53 9.79 12.01 6.23 FL 11.87 14.48 11.15 8.29 6.22 TL 8.30 12.78 6.94 5.93 5.47 SI 4.22 0.52 0.34 3.71 0.25 HI —5.82 —1.40 0.91 4.31 4.90 FI —0.08 —0.61 —1.45 3.26 0.07 LI —1.46 —2.50 —1.42 —0.82 —1.66 I in SW and FW, and I-II in HW) to the II-III intervals, relatively high values appearing in | the next (table 3). The indices considered show particular variation patterns with relative age. SI progressively increases its averages from classes O to V (table 1), specially in the III-IV interval (table 3). The differences between the mean values obtained in the extreme categories (O, V) as well as in classes III and V are significant (O-V: t=4.7801, p<0.001; | III-V: t=4.5689, p<0.001). 634 JACINT VENTURA The HI averages diminish significantly between classes 0 and II (t=3.2935, p<0.01). In the interval between classes II and V the mean of this index increases significantly -(t=7.5767, p<0.001) (table 1). This increase can be verified when the relatively high percentages of variation (4-5%) in the last two age intervals are taken into account (table 3). The FI index follows a similar variation pattern to that of HI, although FI shows a smaller range (table 1). Thus, the average values diminish, albeit not significantly, between classes 0 and III, and from this latter one the means of FI increase significantly until class V (t=2.3859, p<0.05). Nevertheless, this increase is relatively less marked than that observed in HI in the II-V interval. The LI averages decrease from classes 0 to V with relatively constant rate (tables 1 and 3). The differences between the means of these age classes are clearly significant (t=10.0079, p<0.001). DISCUSSION Although significant differences between males and females are observed in some parameters, due to the higher biometrical similarity in most of the intersexual comparisons performed, it can be stated that there is no sexual dimorphometry in the scapula and limb long bones in the population analysed. Taking into account the values of the regression coefficients and their confidence intervals, the width of the scapula is the parameter that shows highest growth rate. The long bones of the hind limb grow faster than those of the forelimb. Likewise, in each limb, the proximal bones show the highest growth rate. The variation percentages between the averages of consecutive age classes reveal that the scapula and the limb long bones show their maximum length increase before fourteen weeks of age, approximately. The widths of the scapula, humerus and femur show a relatively high increase starting from fourteen weeks of age. In the correlation against the body weight, these parameters show significantly higher growth rate than the corres- ponding lengths. The means of SI in adult specimens (classes IV and V) are about 71%. This value is lower than that obtained in A. sapidus (75.1-76.1%; VENTURA 1990). These differences can be attributed to functional factors determined by the different eco-ethological characteristics of the two species. Thus, the relatively longer scapula shown by underground rodent species (SCHICH 1971; LAVILLE et al. 1989) may be the factor that determines the interspecific differences detected. The variation pattern of SI with relative age in A. terrestris is, in general terms, similar to that obtained in A. sapidus (VENTURA 1990). In both species, the width of the scapula shows a positive allometry against length up to, approximately, the first year of age. Thereafter, this relation is isometric. The averages of HI in adult specimens vary between 10.4-11%. These values are notably higher than those detected in A. sapidus (8.3-8.6%; VENTURA 1990).This inter- specific relation coincides with the results obtained by CABRERA-MILLET (1980). Taking into account the data reported by Bou er al. (1987)., the differences between these two species may be due to the relatively higher width that show the proximal bones of the fore limb in underground rodent species. Likewise, digging species tend to have shorter limb | bones (BOU et al. 1987). | During the first ten weeks of life, approximately, the length of the humerus shows a positive allometry against width. Thereafter, the allometric growth is favourable to the MORPHOMETRIC DATA OF ARVICOLA TERRESTRIS 635 width. This variation pattern is similar to that reported in A. sapidus (VENTURA 1990), but the increase that occurs during adulthood is higher in A. terrestris. The average of FI in adult specimens is about 13.3%. This percentage is clearly higher than that observed in A. sapidus (11.3-11.7%; VENTURA 1990). These results coincide with the data of Bou et al. (1987), which indicate that underground rodent species tend to have relatively shorter limb bones. During the fourteen first weeks of life, the length and width of the femur vary isometrically, existing a relative light increase of the length. From this age on, the averages of FI increase due to the positive allometry favourable to the width during adulthood. The variation pattern of FI with relative age is similar to that reported in A. sapidus, but the range of variation is higher in this species (VENTURA 1990). The mean values of LI in adult specimens vary between 80.3-81.7%. These percentages are notably higher than those observed in A. sapidus (74.1-74.9%; VENTURA 1990). This interspecific relation coincides with the results reported by CABRERA-MILLET (1980). According to SCHICH (1971) and CABRERA-MILLET (1980), the aquatic rodents show relative longer hind limbs. This fact determines the lower averages of LI observed in A. sapidus in relation to A. terrestris. The regression coefficients obtained in the correlations of each limb long bone length against the body weigth indicate that the hind limb shows a siginificant higher growth rate than the forelimb. The variation pattern of LI until fourteen weeks of age, approximately, is similar to that observed in A. sapidus (VENTURA 1990). During adulthood LI averages diminish lighter in both species. The results obtained indicate some differences in the growth patterns of the scapula and limb long bones between the two Palaearctic species of Arvicola. The precise determination of these patterns and their adaptative and phylogenetic significance will be stated when studies on this matter on the aquatic populations of A. terrestris are undertaken. ACKNOWLEDGEMENTS The author wishes to express his gratitude to J. Gosälbez and M. J. Löpez-Fuster (Barcelona) for their technical assistance. LITERATURE Bou, J. & A. Casinos 1985. Scaling of bone mass to body mass in insectivores and rodents. Pp. 61- 64. In: Functional Morphology in Vertebrates (R. DUNCKER & G. FLEISCHER eds.). Gustav Fisher Verlag, Stuttgart. Bou, J., A. Casinos & J. OCANA 1987. Allometry of the Limb Long Bones of Insectivores and Rodents. J. Morphol. 192: 113-124. Brown, J.C. & G.I. TWIGG 1969. Studies on the pelvis in British Muridae and Cricetidae (Rodentia). J. Zool., Lond. 158: 81-132. CABRERA-MILLET, M. 1980. Estudio morfolögico del esqueleto locomotor de los roedores ibéricos. Tesis de Licenciatura. Universidad Autönoma de Madrid. KRATOCHVIL, J. 1983. Variability of some criteria in Arvicola terrestris (Arvicolidae, Rodentia). Acta Sc. Nat. Brno 17: 1-40. 636 JACINT VENTURA LAVILLE, E., A. Casinos, J.P. Gasc, S. RENous & J. Bou 1989. Les mécanismes du fouissage chez Arvicola terrestris et Spalax ehrenbergi: étude fonctionnelle et évolutive. Anat. Anz. 169: 131-144. PELIKAN, J. 1972. Arvicola terrestris (L.) indexes of reproduction in Czechoslovakia. Acta Sc. Nat. Brno 11: 1-50. REICHSTEIN, H. 1982. Arvicola terrestris (Linnaeus, 1758)-Schermaus. Pp. 209-252. In: Handbuch der Säugetiere Europas, 2/1 (J. NIETHAMMER & F. KRAPP eds.). Akademische Verlagsgesellschaft, Wiesbaden. SCHICH, J. 1971. Funktionelle Deutung anatomischer Baumerkmale am Achsen-und Gliedmas- senskelett der Schermaus Arvicola terrestris scherman (Shaw, 1801). Säugetierkdl. Mitt. 19: 305-338. SOKAL, R.R. & F.J. ROHLF 1981. Biometry: the principles and practice of statistics in biological research. Second ed. Freeman and Company, San Francisco, 859 pp. VENTURA, J. 1988. Contribuciôn al conocimiento del género Arvicola Lacépéde, 1799, en el nordeste de la Peninsula Ibérica. Tesis Doctoral. Universidad de Barcelona. VENTURA, J. 1990. Datos biométricos sobre los huesos largos y la escäpula de Arvicola sapidus Miller, 1908 (Rodentia, Arvicolidae). Bol. R. Soc. Esp. Hist. Nat. (Sec. Biol.) 86: 55- 64. VENTURA, J. & J. GOSALBEZ 1990. Reproductive cycle of Arvicola terrestris (Rodentia, Arvicolidae) in the northeast of the Iberian Peninsula. Z. Sdugetierkunde 55: 383-391. VENTURA, J., J. GOSALBEZ & V. GOTZENS 1991. The os coxae of a digging form of the northern water vole Arvicola terrestris (Rodentia, Arvicolidae). Anat. Histol. Embryol. 20: 225-236. Revue suisse Zool. | Tome 99 | Fasc.3 | p. 637-643 Genève, septembre 1992 | Eine neue Bolitobius-Art aus Pakistan (Coleoptera, Staphylinidae) 9. Beitrag zur Kenntnis der Tachyporinen von Michael SCHÜLKE* Mit 17 Textfiguren ABSTRACT A new Bolitobius-species from Pakistan (Coleoptera, Staphylinidae). — The description of Bolitobius besucheti spec. nov. from Pakistan is given with illustrations of distinctive characters. A Lectotype of Bolitobius bicolor (Cameron) is designated, and the distribution of both species is shown. Anläßlich der Beschäftigung mit der Bolitobius setiger — Gruppe, die im Himalayagebiet und in Ostasien verbreitet ist, wurden auch alle anderen beschriebenen Taxa der Gattung aus dem betreffenden Faunengebiet sowie eine größere Menge unbearbeiteten Materials untersucht. So fand sich unter einer größeren Anzahl von Bolitobius aus dem Himalaya, die mir dankenswerter Weise von den Kollegen des Genfer Museums (Cl. Besuchet und I. Löbl; (MHNG)) zur Bearbeitung zur Verfügung gestellt wurden, eine neue Art aus der Verwandschaft von Bolitobius bicolor (Cameron), die im nachfolgenden beschrieben werden soll. Neben den genannten Genfer Kollegen gilt mein besonderer Dank Miss E. DeBoise und Mr. R. Aldridge (British Museum, Natural History — BMNH) für die Ausleihe von Typenmaterial Camerons. Bolitobius besucheti spec. nov. Material: Hototypus-d: Pakistan: Swat, s/Utrot; 14.V.1983, 2500-2600 m, Besuchet - Löbl; Holotypus-d, Bolitobius besucheti spec. nov., M. Schülke det. 1990 (rot) (MHNG); Paratypen- d 2:19, gleiche Daten, (MHNG); 14, 19, gleicher Fundort, 13.V.1983 (MHNG, cSCHU); 14, Pakistan: Swat, s/Miandam, 2300 m, 10.V.1983, Besuchet - Löbl (MHNG); 26, Pakistan: Chitral, s/Madaglasht, 27.V.1983, 2900-3050 m, Besuchet - Löbl (MHNG, cSCHÜ); alle mit Etikett: Paratypus-d 9, Bolitobius besucheti spec. nov., M. Schülke det. 1990/91 (rot). * Schulzestr. 26, D - (0) 1100 Berlin, Bundesrep. Deutschland. 638 MICHAEL SCHÜLKE Kopf und Halsschild dunkelbraun bis schwarz, die Hinterecken des Halsschildes ‘etwas heller durchscheinend. Vorderrand des Clypeus und Mitte der Stirn etwas aufgehellt. Flügeldecken einschließlich der Epipleuren rotbraun, Hinterleib dunkelbraun bis schwarz, die Hinterränder der Tergite III-VI aufgehellt, das Tergit VII mit heller Hinterhälfte, die Hinterleibsspitze ganz hell. Fühlerglieder 1 und 2 gelb, die folgenden Glieder dunkler werdend, bräunlich, das Endglied heller. Beine einfarbig gelbbraun, die Mundwerkzeuge von gleicher Farbe. Größe: 8 - 8,5 mm, Vorderkörperlänge 3,25 - 3,5 mm. Im Habitus B. bicolor sehr ähnlich, weshalb sich die folgende Beschreibung im wesentlichen an für die Unter- scheidung beider Arten wesentlichen Merkmalen orientiert. Kopf wie bei allen anderen Arten der Gattung gebaut, Augen mäßig hervortretend. Labrum (Abb. 6) quer, wie bei anderen Arten der Gattung querreihig beborstet (Anzahl größerer Tastborsten 19). Mentum (Abb. 7) stark quer, deutlich mehr als doppelt so breit wie lang, mit zwei Paaren langer Tastborsten an den Seiten und einem Paar etwas kürzerer Borsten in der Mitte. Fühler schlank, Glied 1 dreimal so lang wie breit, Glied 3 länger und etwas breiter als Glied 2, alle folgenden Fühlerglieder länger als breit, Glied 10 so lang wie breit, das Endglied mit ausgeprägtem Sexualdimorphismus, beim Männchen gestreckt, 1,75 mal so lang wie breit, beim Weibchen kürzer, nur 1,35 mal so lang wie breit. Halsschild quer, etwa um ein Viertel breiter als lang, wie bei allen anderen Arten der Gattung mit vier langen Tastborsten an Vorder-, Hinter- und Seitenrändern. Flügeldecken etwas länger als breit (Flügeldeckenseitenrandlänge zu -breite 1,09 : 1), trotz deutlicher Schultern nach hinten etwas erweitert, die größte Breite kurz vor dem Hinterrand erreichend. Auf den Flügeldecken befinden sich zahlreiche Borstenpunkte in denen lange Tastborsten inserieren (Abb. 8). Diese Borstenpunkte sind mehr oder weniger reihig angeordnet. Neben der Flügeldeckennaht befindet sich eine Suturalreihe, die aus 14-18 Borsten- punkten besteht. Am Außenrand der Flügeldecken befindet sich eine aus etwa 12 Borstenpunkten bestehende Lateralreihe und am Hinterrand eine aus 6 - 8 Borstenpunkten bestehende Apikalreihe. Auf der Scheibe der Flügeldecken sind weitere etwa 30 Borsten- punkte in vier undeutlichen Reihen angeordnet. Eine weitere Borstenreihe befindet sich dicht unterhalb des Flügeldeckenaußenrandes auf den Epipleuren. Dort sind jederseits 20- 25 kleinere Borsten angeordnet. Hinterleib gestreckt, wie bei anderen Arten der Gattung gebaut. Das II. Tergit mit einer breiten unpunktierten Mittelzone, sonst kräftig und weitläufig beborstet. Auf den folgenden Tergiten ist die Beborstung gleichmäßiger, nur vor dem Tergithinterrand bleibt in der Mitte jeweils eine punktfreie Zone erhalten. Hinter- rand von Tergit VII mit deutlich entwickeltem Hautsaum. Beine schlank und gestreckt. Mikroskulptur! Kopf nur andeutungsweise quermaschig mit etwa 2-3 Maschen/ 10 um chaginiert, Halsschild mit sehr feinem quergerieftem Chagrin von etwa 5 Maschen/ 10 um. Flügeldecken noch enger als der Halsschild skulpturiert (etwa 7-8 Maschen/ 10um), Hinterleibstergite etwa so dicht wie der Halsschild, quermaschig chagriniert. Männchen: Beim Männchen sind die Vordertarsen erweitert, Sternit VII (Abb. 3) mit einzelnen Borsten in der Mitte des Hinterrandes, Sternit VIII ähnlich wie bei Bolitobius castanueus (Steph.) gebildet (Abb. 4), der Hinterrand in der Mitte mit einem abgerundet dreieckigen Vorsprung. In der Mittellinie in der hinteren Hälfte mit einem langgestreckten Feld wenig dicht stehender kurzer, kräftiger Borsten, Sternit X schlank (Abb. 5). Aedoeagus (Abb. 1) ähnlich dem von B. cingulatus Mannh. mit mäßig gestrecktem, apikal zugespitztem Medianlobus (Innenstruktur des Medianlobus, Abb. 2). BOLITOBIUS-ART AUS PAKISTAN 639 SSS — 5 nn ra: = DA ) LE > » » Sy SA ,— N > >> LZ — In È Ex ? ABB. 1-5. Bolitobius besucheti spec. nov., Pakistan, Swat 1) Aedoeagus, PT (ZNr. 85), 2) Aedoeagus, Medianlobus-Innenstruktur PT (ZNr. 85), 3) d-Sternit VII, HT (ZNr. 91), 4) d-Stemit VIII, PT (ZNr. 87), 5) 8 -Sternit X, PT (ZNr. 86) - Maßstab 0,5 mm. 640 MICHAEL SCHÜLKE ABB. 6-11. Bolitobius besucheti spec. nov., Pakistan, Swat 6) Labrum, PT (ZNr. 119), 7) Mentum, PT (ZNr. | 120), 8) Flügeldecke, PT (ZNr. 121), 9) 9 -Tergit VIII, PT (ZNr. 122), 10) 9 -Sternit VIII, PT (ZNr. 123), 11) 2-Tergit X, PT (ZNr. 116) - Maßstab 0,25 mm (6,7), 0,5 mm (8-11). BOLITOBIUS-ART AUS PAKISTAN 641 Weibchen: beim Weibchen ist das Sternit VIII am Hinterrand einfach breit abgerundet (Abb. 10), das Tergit VIII ebenfalls ohne besondere Auszeichnungen (Abb. 9). Tergit X relativ schlank (Abb. 11). Verbreitung: Bisher nur aus den pakistanischen Provinzen Swat und Chitral aus Höhenlagen zwischen 2300 und 3050 m bekannt. Wahrscheinlich ist die Art in ähnlichen Höhen im Westhimalaya weiter verbreitet (Abb. 17). Derivatio nominis: Herrn Dr. CI. Besuchet (Museum d'Histoire naturelle Genève) als Entdecker der Art gewidmet. Differentialdiagnose: Leider sind mir bisher von Bolitobius bicolor (Cameron) nur weibliche Tiere bekannt, so daß ein Vergleich der männlichen Sexualcharaktere vorerst ausbleiben muß. Beide Arten lassen sich aber auch im weiblichen Geschlecht deutlich unterscheiden: B. besucheti spec. nov. Flügeldecken (Abb. 8) schlanker, etwa so breit wie der Halsschild, auf der Scheibe mit etwa 30 in vier undeutlichen Reihen angeordneten Borstenpunkten, Apikalreihe aus 7- 8 Borstenpunkten bestehend. Flügeldeckenepipleuren wie die Flügeldecken gefärbt. Fühler gestreckt, die vorletzten Glieder länger als breit oder so lang wie breit (Glied 10), Fühlerglieder 3-10 bräunlich. Sternit und Tergit VIII des Weibchens am Hinterrand +/- einfach abgerundet (Abb. 10, 9), Tergit X schlanker (Abb. 11). Mentum mit schärferen Hinterecken und einem Paar kleinerer Tastborsten in der Mitte (Abb. 7). Mikroskulptur auf dem Kopf erloschen und wie auf dem Halsschild etwas weitläufiger als bei B. bicolor (Cameron). Bisher in Höhenlagen zwischen 2300 und 3050 m gefunden. B. bicolor (Cameron) Flügeldecken (Abb. 13) etwas breiter als der Halsschild, auf der Scheibe mit etwa 25 in drei Reihen angeordneten Borstenpunkten. Auf der Innenseite der Flügeldecken befinden sich neben der Suturalreihe nur wenige Einzelborsten. Die Apikalreihe besteht nur aus 5 Borsten. Die Flügeldeckenepipleuren schwarz. Fühler kurz, die vorletzten Glieder deutlich quer, Fühlerglieder 3-10 schwarz. Sternit VIII des Weibchens am Hinterrand etwas zugespitzt (Abb. 15), Tergit VII (Abb. 14) am Hinterrand in der Mitte mit einem abgerundeten Mittelvorsprung. Tergit und Sternit VIII breiter als bei 2. besucheti spec. nov., Tergit X ebenfalls wesentlich breiter (Abb. 16). Mentum mit breit abgerundeten Hinterecken (Abb. 12) und ohne zusätzliches drittes Borstenpaar in der Mitte. Mikroskulptur auf dem Kopf deutlicher und dichter (5 Maschen / 10 um), auf dem Halsschild ebenfalls dichter (7-8 Maschen/ 10 um), sonst wie bei B. besucheti. Die bisherigen Funde stammen soweit bekannt aus Höhenlagen unter 2000 m. Bolitobious bicolor (CAMERON, 1926) Typen: Bolitobius bicolor (Cameron) wurde von Dehra Dun und Sijla Gad (Chakrata District) beschrieben. Mir lag aus dem British Museum (Natural History) ein Syntypus (2) mit folgender Etikettierung vor: Syntype (rund mit blauem Rand); Dehra Dun, Dr. 642 MICHAEL SCHÜLKE ABB. 12-16. Bolitobius bicolor (Cameron), Indien, Chaubattia 12) Mentum, (ZNr. 118), 13) Flügeldecke (ZNr. | 124), 14) 9 -Tergit VIII (ZNr. 125), 15) 2 -Sternit VIII (ZNr. 126), 16) 2 -Tergit X (ZNr. 117) - | Maßstab 0,25 mm (12), 0,5 mm (13-16). | BOLITOBIUS-ART AUS PAKISTAN 643 Cameron, 11-7-1921; M. Cameron, Bequest., B.M. 1955-147.; Wood (rotten); Bryocharis bicolor (Cam., P.M. Hammond det. 1989, Syntype. Dieser Syntypus wird hiermit als Lectotypus designiert, er wurde von mir mit einer Etikette: Lectotypus-9, Bryocharis bicolor Cameron, 1926, des. M. Schülke 1991, versehen. Weiteres Material lag mir von folgenden Fundorten vor: Indien, Kumaon (UP), Chaubattia Pres. Rassikhet. env. 1800 m, 14-13-X-79, I. Löbl (19, cSCHU); Pakistan: Hazara, Malkandi, 1500 m, 3.V1.1983, Besuchet - Löbl (19, MHNG). Damit scheint B. bicolor über den Westhimalaya weit verbreitet zu sein und hier vor allem in den Vorgebirgen der Siwalik Range vorzukommen (Abb. 17). BI Bolitobius bicolor (CAMERON), überprüftes Material O Bolitobius bicolor (CAMERON), nicht überprüft ® Bolitobius besucheti spec. nov. Pakistan ABB. 17. Verbreitung von Bolitobius besucheti spec. nov. und B. bicolor (Cameron). Bolitobius bicolor (Cameron) und B. besucheti spec. nov. sind wahrscheinlich Schwesterarten. Auf Grund des Fehlens von männlichen Exemplaren von B. bicolor Cameron kann ich z.Z. aber nicht ausschließen, daß bicolor doch zur B. setiger (Sharp) - Gruppe gehört. Beide Artengruppen sind nach ektoskelettalen Merkmalen im weiblichen Geschlecht nicht zu unterscheiden. Ein Indiz für die Zugehörigkeit zur B. setiger-Gruppe könnte eventuell das Fehlen eines dritten Borstenpaares auf dem Mentum sein (bei den anderen Arten der B. castaneus (Stephens) - Gruppe sind drei Borstenpaare vorhanden, die allerdings anders als bei B. besucheti angeordnet sind). Wer in seiner Sammlung eventuell männliche Exemplare von Bolitobius bicolor (Cameron) besitzt, sei hiermit gebeten diese Tiere zur Klärung zur Verfügung zu stellen. LITERATUR CAMERON, M. 1926. New species of Staphylinidae from India. — Trans. Ent. Soc. London, 74: 171-191. i x i | > # FRITTA fl CN } oe i “ir > di ‘ u By Maral cé u © ANR | | | | Revue suisse Zool. | Tome 99 | Fasc. 3 p. 645-653 | Geneve, septembre 1992 | Two new species of Laureola Barnard, 1960 from India and Vietnam (Crustacea, Oniscidea, Armadillidae) par D.H. KWON*, F. FERRARA** and S. TAITI** With 5 figures ABSTRACT Two species of Laureola Barnard, 1960 (Armadillidae), L. indica from India, and L. vietnamensis from Vietnam, are described as new. These records fill the distributional gap of the genus, previously known from Africa and Australia. The genus Laureola was established by BARNARD (1960: 53) to accommodate three species previously ascribed to the genus Akermania Collinge, 1919 and three new species, all from southern Africa. The definition of the genus is very short and incomplete, and also contains a mistake ("No flange or tooth ventrally on epimera of 15* and 24 peraeon segments", actually present). Some years later VANDEL (1973a), discussing species from Australia, gave a new definition of Laureola and instituted the new genus Praelaureola. Curiously all Barnard's species fit Vandel's definition of Praelaureola while none can be placed in Laureola sensu Vandel, 1973 so that SCHMALFUSS & FERRARA (1983) synonymized Praelaureola Vandel, 1973 with Laureola Barnard, 1960 and instituted the new genus Pseudolaureola for Laureola sensu Vandel. Pseudolaureola includes three species: P. wilsmorei (Nicholls and Barnes, 1926) from western Australia, P. atlantica (Vandel, 1977) from St. Helena, and P. hystrix (Barnard, 1958) from Madagascar. Since no type species was selected for Pseudolaureola, Laureola atlantica Vandel is here designated as such. SCHMALFUSS & FERRARA (1983) also suggested that Laureola might correspond to Echinodillo Jackson, 1933 which at present includes two species: E. montanus Jackson, 1933 from Marquesas Islands, and E. cavaticus Green, 1963 from Tasmania. The problem of this synonymy can only be clarified with a general comparative study, which has not yet been possible to undertake. * Department of Biology, Inje University, Kimhae 621-749, Republic of Korea. ** Centro di Studio per la Faunistica ed Ecologia Tropicali del Consiglio Nazionale delle Ricerche, Via Romana 17, 50125 Firenze, Italy. 646 D.H. KWON, F. FERRARA & S. TAITI Nine species are currently ascribed to the genus Laureola: L. paucispinosa (Barnard, - 1949), L. longispina (Barnard, 1956). L. miacantha (Barnard, 1960), L. bivomer Barnard, 1960, L. hiatus Barnard, 1960, L. rubicunda Barnard, 1960, L. silvatica (Vandel, 1973). L. canberrensis (Vandel, 1973), and L. dubia Schmalfuss and Ferrara, 1983. The first six species populate southern Africa, L. dubia Principe and Sao Tomé islands in the Gulf of Guinea, and L. silvatica and L. canberrensis New South Wales, Australia. The discovery of two new species of Laureola from southern India and Vietnam is of particular interest since these two areas fill a good part of the distributional gaps between the species of Laureola. The material is desposited in the collections of the Hungarian Natural History Museum, Budapest (HNHM), the Muséum d'Histoire Naturelle, Genève (MHNG), and the Museo Zoologico "La Specola" (Sezione del Museo di Storia Naturale) dell'Università, Firenze (MZUF). Genus Laureola BARNARD, 1960 Synonyms: Praelaureola VANDEL, 1973a: 150. Paralaureola VANDEL, 1973b: 142 (nomen nudum). Type-species: Akermania paucispinosa Barnard 1949, by original designation. Laureola indica sp. nov. (Figs 1-3) Specimens examined. - India, Travancore: 1 2 holotype, Palnis supérieurs, petite Shola au-dessus de Pumbarai, 2000 m, sous bois, 29.11.1927, leg. J. Carl (MHNG); 2 2 2 paratypes, Palnis supérieurs, Kukkal-Shola, ca. 2000 m, sous bois pourri, 1.1V.1927, leg. J. Carl (MHNG); 1 2 paratype, same data (MZUF); 5 2 2 paratypes, Vallée de Vattavadai (entre Palnis et Anaimalais), grande forêt primaire dans la partie supérieure de la vallée, env. 1800-1850 m, 10.1V.1927, leg. J. Carl (MHNG); 1 4,1 9, paratypes, same data (MZUF). Description. — Maximum size, 6.5 x 4.5 mm. Colour yellow, brown or reddish brown. Dorsum with short spiniform tubercles arranged as follows: 5 on cephalon; 1+5 on pereonite 1; 5 on each of pereonites 2-7; 1 in the middle of each of pleonites 3-5. One line of noduli laterales per side, arranged on lateral tubercles of each pereonite. Eyes with 16- 17 ommatidia. Cephalon with frontal lamina distinctly protruding above vertex, with upper margin raised in the middle, forming a triangular process frontally excavated. Pereonite 1 with posterior margin concave at the sides, postero-lateral corner rounded; a | small rounded transversal tooth on ventral surface of epimera. Epimera of pereonites 2-4 triangular; 5-7 and pleonites 3-5 with short points at posterior corners. Pereonite 2 | ventrally with a large quadrangular lobe more or less parallel to the anterior margin, | clearly separated in the distal part from outward directed epimeron. Telson with triangular | distal part, sinuous sides. Antennule with many superimposed aesthetascs near apex. | Antenna with second article of flagellum four times as long as first. Mandible with molar | penicil composed of several plumose setae of increasing length, each arising from a | common stem. Outer branch of maxillule with 10 teeth apically simple; inner branch with | two unequal penicils. Maxilliped with no particular modifications. Uropods with protopod | triangular, exopod inserted near medial margin, not surpassing tip of protopod. Male. All pereopods without apparent sexual specializations. Pleopod 1 exopod | small, much wider than long, with largely rounded distal part; endopod with apical part | bent outwards, without particular modifications. Pleopod 2 exopod slightly shorter than | endopod; endopod with styliform distal part. ; LAUREOLA FROM INDIA AND VIETNAM 647 FIG. 1. Laureola indica sp. nov., 9 paratype: A) lateral view; B) cephalon and pereonite 1 in dorsal view; C) the same in frontal view; D) left epimera of pereonites 1-3 in ventral view; E) pereonite 7, pleon, telson and uropods; F) antennule. 648 D.H. KWON, F. FERRARA & S. TAITI FIG. 2. Laureola indica sp. nov., 9 paratype: A) antenna; B) mandible; C) maxillule; D) maxilliped; E) | ? uropod. LAUREOLA FROM INDIA AND VIETNAM 649 Fic. 3. Laureola indica sp. nov., d paratype: A) pereopod 1; B) pereopod 7; C) pleopod 1 endopod; D) pleo- pod 1 exopod; E) pleopod 2. 650 D.H. KWON, F. FERRARA & S. TAITI Etymology. - The name of the species refers to India, where the specimens were collected. Remarks. — The easiest characters to distinguish species of Laureola are number, shape and disposition of dorsal spiniform tubercles. L. indica is readily distinguished from southern African species by the fewer tubercles (total 44 vs 81 or more), from L. dubia and L. canberrensis by the different number of tubercles and also by the shape of the telson with triangular instead of rectangular distal part (from L. dubia also by the shape of uropodal protopod, distally acute vs rounded); from L. silvatica by the number (46 vs 73) and disposition (presence of median spine) of tubercles. Laureola vietnamensis sp. nov. (Figs 4, 5) Specimens examined. - Vietnam: 1 2 holotype, Cuc Phuong, Prov. Ninh binh, 14.V.1966, beaten from bushes in forest, leg. Topal (HNHM). Description. — Dimensions: 8 x 5 mm. Colour yellowish (faded by conservation?). Dorsum with long spiniform tubercles arranged as follows: 2+5 on cephalon; 7 on pereonite 1; 9 on pereonites 2-6; 7 on pereonite 7; a short median one on pleonites 3 and 4; a long one on basal part of telson. Pereonites 2-7 with a pair of sublateral triangular projections, directed forwards, which overlap the preceding pereonite. One nodulus lateralis per side on each lateral tubercle of pereonites, in subapical position. Globose eyes with 16 ommatidia. Cephalon with frontal lamina distinctly protruding above vertex and with a long triangular process in the middle. Pereonite 1 with posterior margin straight and largely rounded posterolateral corner; a small rounded tooth on ventral surface of epimera near posterior margin; epimera directed outwards. Pereonites 2-7 and pleonites 3-5 with lateral margins of epimera elongated, tapering, directed horizontally outwards. Epimera of pereonite 2 ventrally with a large quadrangular lamellar lobe. Telson with narrow triangular distal part, apex subacute. Antenna with long fifth article of peduncle, slightly exceeding middle of pereonite 1 when pushed back; flagellum with second article over three times as long as first. Buccal pieces as in L. indica. Uropods with protopod triangular, long exopod inserted on medial margin, distinctly surpassing tip of protopod but not reaching tip of telson. Etymology. — The species is named after Vietnam where the specimen was | collected. | Remarks. - L. vietnamensis differs from all the other species by the number and | disposition of tergal tubercles. It is similar to L. longispina, L. miacantha, L. bivomer, and | L. hiatus, all with a triangular telson and long dorsal spiniform tubercles. Among other | characters, L. vietnamensis is readily distinguished from these and L. indica by the lack of | tubercles on pleonite 5 and the long triangular process in the middle of the frontal lamina. | It differs from L. silvatica, L. dubia and L. canberrensis by having a median tubercle on | the pereonites, pleonites 3-4 and telson, by the frontal process and the length of tubercles. | From L. dubia and L. canberrensis, it also differs in the shape of telson (triangular vs | rectangular) and uropods. ACKNOWLEDGEMENTS We wish to thank Dr B. Hauser (MHNG) and Dr L. Forrö (HNHM) for the loan of | material. LAUREOLA FROM INDIA AND VIETNAM 651 Fic. 4. Laureola vietnamensis sp. nov., 9 holotype: A) lateral view; B) cephalon and pereonites 1-4 in dorsal view. 652 D.H. KWON, F. FERRARA & S. TAITI FIG. 5. Laureola vietnamensis sp. nov., 2 holotype: A) cephalon and pereonite 1 in dorsal view; B) cephalon in frontal view; C) left epimera of pereonites 1-3 in ventral view; D) pereonite 7, pleon, telson and uropods (position of spiniform tubercles on pereonite 7 is represented by circles); E) pleonite 5, telson and uropods in dorsal view; F) telson and uropods in ventral view. LAUREOLA FROM INDIA AND VIETNAM 653 RESUME Deux nouvelles espèces de Laureola Barnard, 1960 (Armadillidae), L. indica d'Inde et L. vietnamensis du Viét-Nam sont décrites. Ces données permettent d'étendre la distribution du genre, connu jusqu'à présent seulement d'Afrique et d'Australie. REFERENCES BARNARD, K.H. 1960. Terrestrial Isopoda from the Transvaal. Ann. Natal Mus. 15: 45-55. SCHMALFUSS, H. & F. FERRARA. 1983. Terrestrial isopods from West Africa. Part 3: Family Armadillidae Verhoeff, 1917. Monitore zool. ital., (N.S.) Suppl. 18: 111-157. VANDEL, A. 1973a. Les Isopodes terrestres de l'Australie. Etude systématique et biogéographique. Mém. Mus. natn. Hist. nat., Paris, (N.S.) (Ser. A, Zool.) 82: 1-171. — 1973b. Les Isopodes terrestres (Oniscoidea) de la Mélanésie. Zool. Verh. 125: 1-160. Revue suisse Zool. | Tome 99 | Fasc. 3 p. 655-672 Genève, septembre 1992 Palpigrades cavernicoles et endogés de Thaïlande et des Célèbes (1ère note) par Bruno CONDE* Avec 11 figures ABSTRACT Endogean and cave dwelling Palpigrades from Thailand and Sulawesi (First contribution). — The first Palpigrades known from Thailand were described by H. J. HANSEN (1901) as Koenenia angusta and K. siamensis. Two other species, the endogean Koeneniodes spiniger and the troglomorphic Eukoenenia thais, were recently established (CONDÉ 1984, 19885). The present paper deals with six species, four of them being previously undescribed. Palpigrades are recorded for the first time from Sulawesi. Les premiers Palpigrades connus de Thaïlande ont été récoltés par Th. Mortensen sur l'île de Koh Chang, dans le Golfe de Siam, sous des pierres, les 7 et 17 janvier 1900. Les deux espèces reconnues par H.J. Hansen et attribuées au genre Koenenia étaient inédites: Eukoenenia angusta (Hansen, 1901) et E. siamensis (Hansen, 1901). Eukoenenia angusta a été retrouvée par P. Leclerc en 1976, sur la plage de Wanakhon, au sud de Prauchuap Khiri Khan, dans le milieu interstitiel (CONDÉ 1988a). Cette espèce a été rencontrée à Pondichéry (ssp. tamula Remy) et sur Sri Lanka par P. REMY (1960, 1961) et je l'ai fait connaître de Varanasi (Bénarès) (ssp. hindua Condé, 1989). Dans l'humus forestier de la région de Chiang Dao, entre 700 et 1000 m d'altitude, L. Deharveng a récolté, en décembre 1980, les premiers spécimens de Koeneniodes spiniger Condé, 1984. Au cours d'une visite du réseau supérieur de la grotte dite Tham Chiang Dao, le 25 décembre 1980, L. Deharveng et A. Gouze ont capturé une femelle adulte de la première espèce troglomorphe connue du Sud-Est asiatique, Eukoenenia thais Condé, 1988b, dont les relations avec l'espèce endogée E. siamensis, non perçues lors de sa description, sont établies dans le présent travail. Nous poursuivons ici l'étude des très nombreux matériaux rassemblés depuis 1985 par L. Deharveng et P. Leclerc, en Thaïlande et à Sulawesi, les Palpigrades étant signalés pour la première fois de cette grande île. * Musée de Zoologie de l'Université et de la Ville de Nancy, 34, rue Sainte-Catherine, F-54000 Nancy. 656 BRUNO CONDE Six espèces des genres Eukoenenia (4) et Koeneniodes (2) ont été reconnues, parmi lesquelles deux seulement (E. thais, K. spiniger) étaient déjà décrites. Tous les spécimens sont déposés au Muséum d'Histoire naturelle de Genève, Dépar- tement des Arthropodes et d'Entomologie II. Je rends hommage à la mémoire du regretté M. J. Chevelu pour le talent et la patience qu'il a déployés dans la mise au net de l'abondante et difficile 1llustration de cette note. 1°. Eukoenenia deleta n. sp. THAILANDE. Province de Phangnga, chef-lieu de Tha Put, village de Ban Tham Thong Lang. Grotte de Tham Nam, rivière souterraine d'environ 600 m de long, milieu très humide 20.VII.87, P. Leclerc leg: 1 femelle adulte, perdue au cours des manipulations de montage et d'orientation. — Grotte de Tham Phet, réseau très voisin du précédent, 18.VII.87, P. Leclerc leg: 1 immature A. Longueurs. — Corps: 0,92 mm (extension); bouclier prosomien: 0,27 mm; basitarse IV: 100 um; patte IV, à partir du tibia: 0,34 mm; B/bta: 2,78; bta/ti: 0,88. Prosoma. — Organe frontal médian presque 1 fois 3/4 (1,70) aussi long que large, à branches latérales ovalaires, terminées par une pointe très nette. Organes latéraux comprenant 4 éléments fusiformes acuminés à gauche et 3 seulement à droite. Bouclier avec au moins 8 + 8 soies assez développées. Segment libre avec 3 + 3 phanères, les intermédiaires les plus longs. Une seule soie deuto-tritosternale. Cheliceres avec 7 dents à chaque mors et 2 phanères rigides sur l'article basal. 50 pm (A,B) 100 pm (C) FIG. 1. Eukoenenia deleta n. sp. immature A holotype de Tham Phet: A. Organe frontal médian. B. Organe latéral gauche. C. Basitarse de la patte locomotrice IV. Explication des lettres dans le texte. 1 Province et chef-lieu traduisent respectivement changwat et amphoe; ban signifie village et tham grotte. PALPIGRADES CAVERNICOLES 657 Pedipalpes et pattes locomotrices I et IV. Longueurs relatives des articles: pédi- palpes: ti (non mesurable), bta 1= 40, bta 2= 57, ta 1= 33, ta 2= 47, ta 3= 61; pattes I: ti= 110, bta 1 + 2= 88, bta 3= 53, bta 4= 46, ta 1= 42, ta 2= 43, ta 3= 153; pattes IV: ti= 131, bta= 116, ta 1= 63, ta 2= 91. Aux pattes I, la soie raide du basitarse 3, environ 1 fois 1/4 aussi longue que le bord tergal de l'article, est insérée un peu au-delà du milieu du bord sternal. Aux pattes IV, le basitarse est plus court que le tibia (116/131, bta/ti= 0,83) et, mesuré au niveau de r, environ 4 fois 1/2 (4,67) aussi long que large; la soie raide (r) est épaisse, exactement 2 fois plus courte que le bord tergal de l'article (58/116), et insérée aux 4/6 distaux de ce bord (65/116, t/er= 1,78), son apex dépassant le bord distal de l'article. Les 2 autres phanères sont les soies sternales distales (esd), insérées presque au même niveau, et de longueurs subégales (72, 75). Opisthosoma. — Au sternite III, st; est deux fois plus court que sf, et stz. De IV à VI, les a, et les a; (53-55) sont longs et relativement grêles, égaux à un peu plus des 2/3 de l'écartement des aj. Une seule paire de phanères au sternite VII. Segments VIII à XI avec chacun 8 soies (4 + 4). Le XIe, un peu moins long que large (75/80), est sensiblement égal à la somme de IX et X (77). Seul le premier article du flagelle est présent; en y incluant l'anneau intermédiaire il est environ 2 fois 1/3 aussi long que XI (179/80), avec un verticille apical de 8 longues soies, suivi d'une couronne de 16 phanères épineux. Discussion. Nous admettons que l'immature de Tham Phet est cospécifique de l'adulte, malencontreusement perdu, de Tham Nam, les deux cavités appartenant au même réseau. Ses caractères, parmi lesquels la forme de l'organe frontal médian, la présence de 3 et 4 éléments aux organes latéraux et la position distale de la soie raide du basitarse IV permettront d'identifier l'adulte correspondant et justifient de nommer ce stade juvénile. L'existence de 3 ou 4 éléments aux organes latéraux dès le stade À, n'est connue que dans les populations occidentales (Basses-Alpes, Savoie) de E. spelaea (Peyerimhoff) (CONDÉ 1977: 666); l'augmentation du nombre de ces récepteurs sensoriels étant une caracté- ristique des lignées troglobies, nous pouvons attribuer E. deleta à l'une d'elles. On notera aussi la taille relativement grande de cet immature qui égale déjà celle de la femelle adulte de E. lyrifer, par exemple. Le nom spécifique (du latin deletus, a, um, détruit) rappelle la perte, particulièrement malheureuse, du seul spécimen adulte. 2°. Eukoenenia thais Condé, 1988b THAILANDE. Province de Chiang Mai, chef-lieu de Chiang Dao, village de Ban Tham Chiang Dao. Grotte de Tham Chiang Dao, réseau supérieur, à 150 m du point de capture de l'holotype, 10.VII.85, L. Deharveng leg.: 1 mâle immature (C). Longueurs. — Les nombres entre parenthèses se rapportent à la femelle holotype. Corps 0,80 mm (1,07 en extension médiocre); bouclier prosomien 0,27 mm (0,36); basitarse IV: 158 um (223); patte IV, à partir du tibia 0,47 mm (0,62); B/bta: 1,69 (1,61); bta/ti: 0,95 (1). Prosoma. Le bord antérieur du bouclier est endommagé, ne permettant pas de lire l'organe frontal médian. L'organe latéral gauche comprend 9 éléments (13 chez l'holotype). Bouclier dorsal avec 10 paires de soies courtes. Cinq soies deuto-tritosternales (2 + 1 + 2) en W. Chélicères avec 8 dents à chaque mors. 658 BRUNO CONDE Pedipalpes et pattes locomotrices I et IV. Longueurs relatives des articles (holotype entre parenthèses): pédipalpes: ti= 72 (95), bta 1= 31,5 (40), bta 2= 34 (46), ta 1= 24 (26), ta 2= 23 (29), ta 3= 36 (42); pattes I: ti= 83 (114), bta 1 + 2= 64 (82), bta 3= 35 (45), bta 4= 28 (36), ta 1= 21 (24), ta 2= 23 (28), ta 3= 74 (95); pattes IV: i= 86 (116), bta= 82 (117), ta l= 35 (43), ta 2= 41 (49). A la patte IV gauche, la seule bien orientée, la soie raide du basitarse est brisée dès la base. Son insertion est un peu plus distale que chez l'holotype, t/er= 4,37 (5,58). Opisthosoma. — Papille génitale correspondant à la variante 3 (CONDÉ 1984: 387). Remarque. — Ce spécimen est le deuxième représentant connu de l'espèce. 3°. Eukoenenia lyrifer n. sp. THAILANDE. Province de Chiang Rai, chef-lieu de Mae Sai, village de Ban Tham. Grotte de Tham Kukan, 24.VI.86, P. Leclerc leg.: 1 femelle adulte. Longueurs. — Corps: 1,07 mm (extension); bouclier prosomien: 0,27 mm; basitarse IV: 100 um; patte IV, à partir du tibia: 0,32 mm; B/bta: 2,70; bta/ti: 0,88. Prosoma. — Organe frontal médian environ 2 fois aussi long que large; ses branches, à bord externe convexe et à bord interne subrectiligne ou légèrement concave, arrondies à l'apex. Organes latéraux non dénombrables à gauche, mais formés à droite de 8 éléments fusiformes acuminés!. esd esp 50 um (A,B) 100 pm ‘(C,D) FIG. 2. Eukoenenia lyrifer n. sp. femelle adulte holotype de Tham Kukan: A. Organe frontal médian. B. Organe lateral droit. C. Basitarses 3 et 4 de la patte locomotrice I. D. Basitarse de la patte locomotrice IV. Explication des lettres dans le texte. 1 Le comptage du groupe de droite n'a été possible qu'à l'aide du dispositif de contraste interférentiel (Nomarsky-Nachet). PALPIGRADES CAVERNICOLES 659 Bouclier avec 10 + 10 soies relativement courtes. Segment libre avec 3 + 3 phanères, les intermédiaires les plus longs. 7 soies deuto-tritosternales (3 + 1 + 3), insérées suivant un W dont les branches médiales sont très courtes. Chélicères avec 9 dents à chaque mors. Deux longs phanères rigides, terminés en une petite palette denticulée, sur l'article basal. Pédipalpes et pattes locomotrices I et IV. Longueurs relatives des articles: pédipalpes: ti= 119, bta 1= 44, bta 2= 56, ta 1= 35, ta 2= 44, ta 3= 56; pattes I: ti = 116, bta 1 + 2= 95, bta 3= 56, bta 4= 47, ta 1= 28, ta 2= 38, ta 3= 121; pattes IV: tibia= 133, bta= 117, ta 1= 48, ta 2= 66. Aux pattes I, la soie raide du basitarse 3, un peu plus longue que le bord tergal de l'article (61/56), est insérée à peine en deçà du milieu du bord sternal. Aux pattes IV, le basitarse est plus court que le tibia (117/133, bta/ti= 0,88) et, mesuré au niveau de r, environ 5 fois aussi long que large; la soie raide r est environ 1 fois 3/5 plus courte que le bord tergal de l'article (75/117, t/r= 1,56) et est insérée un peu au- delà du 1/4 proximal de ce bord (32/117, t/er= 3,65). Les 6 autres phanères sont la soie grêle tergale (grt, 61) qui s'insère très près de r, un peu plus distalement à droite et un peu plus proximalement à gauche; la soie grêle latérale (gla, 65); les 2 paires de soies sternales, les éléments de chaque paire légèrement inégaux (48, 52, 56), étant presque superposés en vue latérale. Opisthosoma. — Tergites II à VI sans fy; les t] (19,5) et les #3 (21) plus courts que l'écartement des 7] (28) et que la distance f; — fz (23,5). Tergite VII sans s. Segments VIII avec 9 (4 + s + 4), IX à XI avec 8 (4 + 4) phanères. Premier volet génital avec 11 + 11 soies (a, présents); à la rangée distale, les phanères ne peuvent être mesurés avec précision, en raison de leurs orientations; a, et a semblent un peu plus courts que a3 et a4, ces derniers étant du reste mieux orientés. Au deuxième volet, les x sont plus grêles que y et z, mais sensiblement de même longueur. Au bord interne de chaque moitié du volet, une sclérification sinueuse dessine, avec sa symétrique, un motif rappelant une lyre. Examiné en contraste interférentiel, l'ensemble prend un aspect assez différent dans lequel on reconnaît néanmoins les branches de la lyre. Le réceptacle séminal n'a pu être observé. Sternites IV à VI avec 2 + 2 phanères épais (a), a,) compris entre une seule paire de soies beaucoup plus courtes et un peu plus grêles (s). Les a; sont plus courts que leur écartement (environ 1 fois 1/3 en V). Une paire de plages réfringentes en IV et V. Sternite VII avec une seule paire de phanéres a, | fois 2/5 plus courts que leur écartement, et une paire de s. Affinités. E. lyrifer est, par les dimensions et par un certain nombre d'autres caractères, intermédiaire entre l'espèce endogée E. siamensis (Hansen) et l'espèce troglobie évoluée E. thais Condé (tableau I). Elle permet ainsi de préciser la position systématique de cette dernière que nous n'avions pu établir à l'époque de sa description (CONDÉ 1988b: 741). TABLEAU I B PIV. BtaIV B/bta t/r t/er bta/ti L/lbta o. lat. (mm) (mm) (um} E. siamensis — 0,21 SIT - 1,07 — 0,69 3,75 3 E. lyrifer 0272082100 eth Wake) Som) 20887755 8 E. cf. lyrifer 027220352121 2235,1:90975522 70.9857 26:90 6 E. thais 0,36 0,62 223 PCM 27E es Sel 9 13 660 BRUNO CONDE FIG. 3. Eukoenenia lyrifer n. sp., femelle adulte holotype de Tham Kukan: A. Premier et deuxième volets génitaux. B. Sclérification du deuxième volet observée en contraste interférentiel. Explication des lettres dans le texte. Parmi les ressemblances avec E. thais, on peut retenir la forme de l'organe frontal médian, le nombre et la disposition des phanères du deuto-tritosternum; l'insertion de la soie raide (r) du basitarse IV qui à une exception près est un peu plus proximale que celle de la soie grêle tergale (grt); la chétotaxie des sternites opisthosomiens IV a VI, comprenant une seule soie grêle latérale (s) et celle des quatre derniers segments. Les différences concernent essentiellement le nombre d'éléments aux organes latéraux (8 au lieu de 13), le moindre allongement des appendices (L/l=5 au lieu 9 pour bta IV), la sclérification (lyriforme) du 2e volet génital et la taille beaucoup plus faible (bta IV 100 um au lieu de 223). PALPIGRADES CAVERNICOLES 661 Le nom spécifique (du grec latinisé /yra, ae, lyre et du latin fero, porter) rappelle la sclérification du deuxième volet génital. 4°. Eukoenenia cf. lyrifer THAILANDE. Province de Chiang Mai, chef-lieu de Chiang Dao, village de Ban Tham Chiang Dao. Grotte de Tham Chiang Dao, réseau guano, sur la paroi, 16.VII.85, P. Leclerc leg: 1 femelle adulte. Longueurs. — Les nombres entre parenthèses se rapportent à la femelle holotype de E. lyrifer. Corps: 0,84 mm, opisthosome contracté (1,07 en extension); bouclier prosomien: 0,27 mm (0,27); basitarse IV: 121 um (100); patte IV, à partir du tibia: 0,35 mm (0,32); B/bta: 2,23 (2,70); bta/ti= 0,98 (0,.88). Prosoma. — Bouclier comme l'holotype, sauf 6 éléments au lieu de 8 à chaque organe latéral et la forme un peu différente de l'organe médian. Pédipalpes et pattes locomotrices I et IV. Longueurs relatives des articles (holotype de E. lyrifer entre parenthèses). pédipalpes: ti= 130 (119), bta 1= 52 (44), bta 2= 63 (56), ta 1= 38 (35), ta 2= 42 (41), ta 3= 60 (56); pattes I: ti= 127 (116), bta 1 + 2= 102 (95), bta 3= 55 (56), bta 4= 48 (47), ta 1= 25 (28), ta 2= 33 (38), ta 3= 119 (121); pattes IV: ti= 141 (133), bta= 139 (117), ta 1= 57 (48), ta 2= 70 (66). Longueurs comparées des trois appendices mesurés à partir du tibia: 385 (351), 509 (501), 407 (364). 5Oym(A,B) 100 um (C,D) FIG. 4. Eukoenenia cf. lyrifer n. sp., femelle adulte de Tham Chiang Dao. A. Organe frontal médian. B. Un élément de l'organe latéral droit. C. Basitarse de la patte locomotrice IV. D. Région marginale du premier volet génital et portion postérieure du deuxième volet. Explication des lettres dans le texte. 662 BRUNO CONDE Le basitarse IV est à peine plus court que le tibia (bta/ti= 0,98) et sa soie raide est . relativement plus courte et insérée plus proximalement: t/r 1,90 (1,56), t/er= 5,52 (3,65), ces valeurs rapprochant ce spécimen de la femelle holotype de E. thais (2,27; 5,58). Opisthosoma. — La marge postérieure du premier volet génital présente une profonde échancrure subtriangulaire médiane, séparant les paires de phanères aj et ay, comme chez E. thais (CONDÉ, loc. cit., fig. 12, B). Le deuxième volet, observé de trois quarts, présente une sclérification lyriforme. Remarque. En dépit de certaines ressemblances, et de la proximité du lieu de récolte, il n'est pas possible de rapporter ce spécimen à E. thais, d'autant que l'on connaît à présent l'immature C de cette espèce qui le dépasse déjà en taille (bta IV: 158 contre 121 um) et en différenciation (9 éléments contre 6 aux organes latéraux). Comme E. lyrifer, cette forme paraît intermédiaire entre l'endogée E. siamensis et l'espèce troglomorphe E. thais. Une comparaison entre les représentants des autres groupes zoologiques récoltés respectivement dans le réseau supérieur et dans le réseau inférieur (guano) de cette vaste cavité pourrait éclairer les relations entre les peuplements de ces deux biotopes. 5°. Eukoenenia maros n. sp. SULAWESI. Province de Sulawesi-sud, chef-lieu de Maros, village de Kappang. Grotte de Gua Tanette (réseau Kiri), à environ 800 m de l'entrée, 6. VII.88, P. Leclerc leg.: 1 femelle adulte. Longueurs. Corps: 0,98 mm (extension moyenne); bouclier prosomien: 0,25 mm; basitarse IV: 133 um; patte IV à partir du tibia: 0,40 mm; B/bta: 1,87; bta/ti: 0,88. Prosoma. Organe frontal médian environ 2 fois aussi long que large; ses branches, à bords subrectilignes, terminées par une très courte pointe mousse. Organes latéraux comprenant 5 éléments dont un plus petit à gauche, 4 à droite; tous sont foliacés, larges et terminés par une très courte pointe mousse. Bouclier portant 10 + 9 soies minuscules, difficiles à dénombrer; le phanère latéral droit de la rangée postérieure n'a pas été vu. Segment libre avec 2 + 2 phanères; le médial (t;) manque, l'intermédiaire (?,) étant presque 2 fois plus long que le latéral (#3) (65/34). 10 soies deuto-tritosternales en 2 rangées, une antérieure de 2 phanères, et une postérieure sinueuse de 3 + 1 + 4. Chélicères avec 9 dents à chaque mors. Trois longs phanères rigides, terminés en une petite palette denticulée, sur l'article basal. Pédipalpes et pattes locomotrices I et IV. Longueurs relatives des articles: pédipalpes: ti= 112, bta l= 43, bta 2= 55, tal= 33, ta 2= 38, ta 3= 55; pattes I: ti= 130, btal + 2= 84, bta 3= 59, bta 4= 42, tal= 43, ta 2 = 40, ta 3= 128; patte IV: tibia= 158, bta= 130, tal= 49, ta 2= 53. Aux pattes I, la soie raide du basitarse 3, un peu plus longue que le bord tergal de — l'article (125/118), est insérée un peu au-delà du 1/3 distal du bord sternal (60/102, s/er= . 1,70). | Aux pattes IV, le basitarse est plus court que le tibia (130/158, bta/ti= 0,88) et, | mesuré au niveau de r, presque 9 fois aussi long que large (L/l= 8,8); la soie raide r est | presque 2 fois 1/2 plus courte que le bord tergal de l'article (106/255, t/r= 2,40) et est | insérée un peu en deçà du milieu de ce bord (117/255, t/er= 2,18). Les 6 autres phanéres | sont la soie grêle tergale (grt, 98), la soie grêle latérale (gla, 103) insérée pres de la | PALPIGRADES CAVERNICOLES 663 100pm (Lo) | FIG. 5. |. Eukoenenia maros n. sp., femelle adulte holotype de Gua Tanette. Bouclier prosomien et tergite du segment libre. o.m = organe frontal médian, o./ = organe latéral. précédente et, un peu plus distalement, les 2 paires de soies sternales (esp, esd) de longueurs comparables entre elles (100 - 110). Opisthosoma. Tergites III à VII avec une rangée réduite à une paire de phanères latéraux (tz) particulièrement longs (55 um et égaux aux 3/4 environ de leur écartement en IV) et 664 BRUNO CONDE épais, mais progressivement atténués vers l'apex, comprise entre une paire de phanères . courts et grâles (s). Segment VIII avec 7 (3 + s + 3), IX et X avec 6 (3 + 3) et XI avec 8 (4 + 4) phanères. Premier volet génital avec 10 + 10 soies (les a3 manquent); à la rangée distale, les a, sont un peu plus courts que les a, (28 — 35), leurs embases sensiblement équidistantes; les ory wate Fic. 6. Eukoenenia maros n. sp., femelle adulte holotype de Gua Tanette: A. Organes frontal médian et latéraux droit (à gauche) et gauche, vus par la face ventrale. — B. Soies du deuto-tritosternum. — C. Basitarse 3 de la patte locomotrice I. — D. Basitarse de la patte locomotrice IV. Explication des lettres dans le texte. PALPIGRADES CAVERNICOLES 665 ay sont plus longs (ca 45) et relativement plus grêles, la distance a, — a, étant au moins double de la distance a; — a. En arrière de a; et a), deux lobes subtriangulaires prolongent la portion centrale de la marge; ils sont hyalins, avec des épines cuticulaires éparses. Deux orifices parasagittaux (g,) s'ouvrent sur la face interne. Le deuxième volet A,B 50 pm C,D 25 um FIG. 7. Eukoenenia maros n. sp., femelle adulte de Gua Tanette: A. Premier et deuxième volets génitaux. — B. Deuxième volet génital, les lobes apicaux non représentés. — C. Lobes apicaux et processus sétiforme du deuxième volet génital. — D. Coupe optique au niveau des sclérifications et du réceptacle du deuxième volet. Explication des lettres dans le texte. 666 BRUNO CONDE porte sur chaque moitié les 3 soies habituelles (x, y, z) qui sont de longueur et de calibre . comparables. Un lobe subtriangulaire hyalin, à pubescence éparse et à bord latéral externe épineux, prolonge chaque moitié du volet. De la face dorsale du lobe gauche se détache un diverticule à région basale subcylindrique, se terminant par un court segment sétiforme. Cette formation a été observée debout sur le lobe droit, avant la mise à plat de la préparation. Le bord interne de chaque moitié du volet est un peu épaissi et rejoint une sclérification qui, avec sa symétrique, évoque une sorte de coupe au voisinage de l'orifice génital. En profondeur, on distingue un réceptacle circulaire, à paroi épaissie, de 5 um de diamètre seulement. Deux groupes latéraux antérieurs de 3 gros orifices glandulaires (g;). Les sternites IV à VI sont en majeure partie épilés; les phanères qui subsistent sur la moitié droite sont longs et grêles, une discrimination entre ceux des séries a et s étant impossible à présent; on ne discerne aucun massif glandulaire sous-jacent et aucune plage réfringente n'a été vue au voisinage du plan sagittal. J'ai compté 4 + 4 poils en IV, V et VI, ceux de la paire médiane étant plus espacés l'un de l'autre que les suivants. Sternite VII avec 2 + 2 poils, les latéraux plus courts que les autres. Affinités. Eukoenenia maros est une espèce troglomorphe, comparable, de ce point de vue, à E. thais Condé, 1988b, du réseau supérieur de la grotte dite Tham Chiang Dao, en Thaïlande septentrionale. L'allongement des appendices est cependant un peu moins accentué, le basitarse IV étant encore plus court que le tibia correspondant (0,82 au lieu de 1), quoique ses proportions soient approchantes chez les deux espèces (environ 9 fois aussi long que large); en revanche, sa longueur, en valeur absolue, est discriminante (133 et 223 um), d'autant que le basitarse d'un mâle juvénile de E. thais atteint déjà 158 um. Les organes latéraux (5 + 4 éléments) sont modestes, face aux groupes de 8 à 13 éléments observés chez quelques espèces. E. thais notamment, mais on connaît aussi des formes troglomorphes, telle E. gasparoi Condé, ne possédant que 3 éléments de chaque côté. La présence, sur les tergites de l'opisthosome, d'une seule paire de phanères f, considérés comme 1; d'après leur position latérale, est partagée par un très petit nombre d'espèces de Eukoenenia, toutes troglomorphes (draco zariquieyi Condé, patrizii Condé, gasparoi Condé, naxos Condé). On peut encore mentionner les soies minuscules du bouclier prosomien qui sont communes à gasparoi, thais, naxos et semblent accompagner souvent les modifications précédentes. Parmi les caractères séparant E. maros de E. thais, on peut encore retenir la forme de l'organe frontal médian, l'absence de t; au segment libre du prosoma, la présence de 3 phanères rigides sur l'article basal des chélicères, la position de r et gla au basitarse IV, l'absence de a3 au premier volet génital, l'aspect des sclérifications du deuxième volet et la chétotaxie des sternites IV à VI. 6°. Koeneniodes leclerci n. sp. THAILANDE. Province de Hua Hin, chef-lieu de Pran Buri, village de Ban Khung Tanot (massif de Kao Sam Roi Yot). Grotte de Tham Sai, réseau fossile de 250 m environ, avec avens d'effondrement, milieu humide, 26.VII.87, P. Leclerc leg.: 1 femelle adulte. Longueurs. — Corps: 1,07 mm (en extension); bouclier prosomien: 0,25 mm; basitarse IV: 88,6 um; patte IV, à partir du tibia: 0,28 mm; B/bta: 2,82; bta/ti: 0,89. Prosoma. — Organe frontal médian à branches assez trapues, terminées par une courte pointe. Organes latéraux formés chacun de 4 éléments fusiformes acuminés, formant une rangée subrectiligne. Bouclier dont les soies minuscules ne sont pas dénombrables par PALPIGRADES CAVERNICOLES 667 transparence. Segment libre avec 3 + 3 phanères, les intermédiaires (#9) 1 fois 3/4 (1,74) aussi longs que les latéraux (tz). 5 soies deuto-tritosternales sur un V. Chélicères avec 9 dents à chaque mors. Deux phanères rigides, terminés en une petite palette denticulée, sur l'article basal. esp 50 pm (A,B) 100 um (C, D) Fic. 8. Koeneniodes leclerci n. sp., femelle adulte holotype de Tham Sai: A. Organe frontal médian. B. Organes latéraux, 2 éléments du gauche et les 4 éléments du droit, ces derniers à région basilaire contractée. C. Basitarses 3 et 4 de la patte locomotrice I. D. Basitarse de la patte locomotrice IV. Explication des lettres dans le texte. Pédipalpes et pattes locomotrices I et IV. Longueurs relatives des articles: pédipalpes: ti= 95 (+); bta 1= 40; bta 2= 55; ta 1= 29; ta 2= 42; ta 3= 56; pattes I: ti= 113; bta 1 + 2= 92; bta 3= 56; bta 4= 41; ta 1= 31; ta 2= 37, ta 3= 114; pattes IV: ti= 114; bta= 102; ta 1= 41; ta 2= 66. Aux pattes I, la soie raide du basitarse 3, de méme calibre que la soie gréle tergale, est sensiblement égale à la longueur du bord tergal de l'article (58/56) et est insérée vers le quart distal du bord sternal (39/53, s/er= 1,35), son apex atteignant la région basilaire du tarse 1. Aux pattes IV, le basitarse est plus court que le tibia (102/114, bta/ti= 0,89) et, mesuré au niveau de r, environ 5 fois aussi long que large; la soie raide r est robuste, environ 1 fois 3/4 plus courte que le bord tergal de l'article (58/102, t/r= 1,75) et est insérée un peu au-dela des 3/5 proximaux de ce bord (44/102, t/er= 2,31). Les 6 autres phanères sont la soie grêle tergale (grt, 35), insérée un peu plus distalement que r; la soie gréle latérale (gla, 48), insérée entre r et grt; les deux paires de soies sternales (51,46/48, 43), les phanéres de chaque paire étant insérés presque au méme niveau. Opisthosoma. — Tergites II a VI sans fp (ty, tz, 5); les t, un peu plus courts que les #3 et que la distance ty — tz (43/51/50 en II). 668 BRUNO CONDE 50pm Fic. 9. Koeneniodes leclerci n. sp., femelle adulte holotype de Tham Sai: Volets génitaux de profil. g,= orifices glandulaires du deuxième volet; r= réceptacle séminal. Explication des autres lettres dans le texte. Tergite VII avec 4 + 4 phanères, la paire médiane la plus courte. Segments VIII avec 11 (5 +1s + 5), IX avec 9 (? 1 sternal), X et XI avec 8 (4 + 4) phanères. Premier volet génital avec 7 + 7 soies, 4 + 4 ventrales, disposées en paires et 3 +3 constituant la rangée distale. A cette rangée, les phanères a], a, et a3 sont de longueurs croissantes (15, 25, 33), les a3 étant comparativement plus grêles. Entre les 3 paires les plus antérieures, on distingue un groupe de 7 (3 + 1 + 3) minuscules phanères à base renflée (pbr) et tige égale au plus à la base. Le deuxième volet porte sur chaque moitié les 3 soies habituelles les x plus longues que y ou z (40/25). Au sternite III, les st, et sf; sont semblables entre eux. Au bord antérieur du sternite IV, un groupe médian de 6 gros phanères coniques (gm) disposés sur 2 rangs superposés (4 + 2) et, en arrière, une rangée de 4 courts phanères bacilliformes (a), a), comprise entre une paire de soies plus longues et plus gréles (s1, 52); sternite V identique, moins le groupe médian; sternite VI avec une rangée antérieure de 9 phanères bacilliformes et une rangée postérieure de 4 phanères semblables, cette dernière homologue de la rangée unique des sternites IV et V. 669 PALPIGRADES CAVERNICOLES \ 100 pm Fic. 10. Koeneniodes leclerci n. sp., femelle adulte holotype de Tham Sai: Sternites IV à VI de l'opisthosome et détail du groupe médian de phanères du sternite IV. Explication des lettres dans le texte. Affinités. L'espèce la plus voisine est K. deharvengi Condé, 1981, de Mindoro (Philippines), connue par la seule femelle holotype. Chez cette dernière, le groupe médian du sternite IV est beaucoup plus important (19 phanères), mais on retrouve les 4 phanères bacilliformes de la rangée postérieure; ces derniers sont également plus nombreux en VI (8), sans atteindre la densité constatée chez leclerci (13). Il existe aussi des ressemblances 670 BRUNO CONDE 100 yum(A), 5Oum (B,C) Fic. 11. Koeneniodes spiniger Condé, 1984, femelle adulte de Tham Soi Yok Noi: A. Volets génitaux et sternites IV à VI. B. Lobes apicaux du premier volet. C. Apex des phanères y et z du deuxième volet. 81» 82, 83= massifs glandulaires; sj, s,= soies gréles; sf, st3= phanères latéraux du sternite III. Explication des autres lettres dans le texte. PALPIGRADES CAVERNICOLES 671 au niveau des organes latéraux du prosome (4 éléments); des cheliceres (9 dents); du basitarse IV, sur lequel la soie grêle tergale (grt) est distale par rapport à la soie raide (r); du premier volet génital avec 7 + 7 phanères principaux. Aucune confusion n'est cependant possible entre les deux espèces, l'absence des groupes latéraux de phanères plumeux aux sternites V et VI de leclerci étant le critère saillant; s'y ajoutent la brievete des soies du bouclier prosomien, la position et la longueur de la soie grêle tergale du basitarse IV, la rudimentation des phanères médiaux du premier volet génital et le détail de la chétotaxie des sternites IV à VI. 7°. Koeneniodes spiniger Condé, 1984 THAILANDE. Province de Kanchanaburi, chef-lieu de Nam Tok. Près de l'entrée de la grotte de Tham Soi Yok Noi, sous une pierre enfoncée dans une terre plus ou moins noire, sous une litière assez humide, en forêt primaire, 17.VI.86, P. Leclerc leg.: 1 femelle adulte. Longueurs. — Les nombres entre parenthèses se rapportent à la femelle holotype. Corps: 0,82 mm (0,96), l'une et l'autre en extension médiocre; bouclier prosomien: 0,27 mm (0,23); basitarse IV: 82,5 um (80,2); B/bta: 3,27 (2,85); bta/ti= 0,83 (0,85). Prosoma. — Au basitarse IV: t/r= 1,35 (1,23); t/er= 2,57 (2,48). La soie grêle antérieure gla (55) est insérée à égale distance de r (70) et de la soie grêle tergale grt (64), tandis qu'elle est plus près de r que de grt chez l'holotype (1984, fig. 4B). Opisthosoma. — Au premier volet génital, le phanère grêle nommé a, dans la diagnose originale (1984, fig. 6A) ressemble en fait aux autres soies ventrales qui seraient ainsi au nombre de 6 paires (au lieu de 5); le très long phanère marginal, inséré à proximité de az, est considéré ici comme un ay. Deux lobes subtriangulaires, aigus et hyalins, non vus chez le type, prolongent le volet au-delà des phanères a, — a4 (fig. 11 B). Les gros phanères y et z du deuxième volet sont tous nettement bifides à l'apex; les deux pointes sont glabres, l'une, plus courte et plus grêle que l'autre, n'ayant pas été observée chez le type (fig. 11 C). Les sternites IV, V et VI portent chacun une rangée de 4 gros phanères surmontant de volumineux massifs glandulaires. Les limites des sternites sont effacées, comme on l'a déjà noté pour l'holotype, et les rangs de phanères sont proches les uns des autres, l'ensemble formant une protubérance médio-ventrale caractéristique. Il ne s'agit donc pas d'un artefact, résultant d'une déformation de l'opisthosome, comme on pouvait le craindre lors de la description du type, mais bien d'un complexe glandulaire associant trois segments, comme chez d'autres espèces de Koeneniodes. BIBLIOGRAPHIE CONDÉ, B. 1977. Nouveaux Palpigrades du Muséum de Genève. Rev. suisse Zool. 84 (3): 665-674. — 1981. Palpigrades des Canaries, de Papouasie et des Philippines. Revue suisse Zool. 88 (4): 941-951. — 1984. Palpigrades d'Europe, des Antilles, du Paraguay et de Thaïlande. Revue suisse Zool. 91 (2): 369-391. 672 BRUNO CONDE — 1988a. Palpigradida, in R.P. Higgins and H. Thiel ed. Introduction to the Study of Meiofauna, Smithsonian Inst. Press, Washington D.C., London: 425-427. — 1988b. Nouveaux Palpigrades de Trieste, de Slovénie, de Malte, du Paraguay, de Thaïlande et de Bornéo. Revue suisse Zool. 95 (3): 723-750. — 1989. Palpigrades (Arachnida) endogés de l'Inde et de Sumatra. Revue suisse Zool. 96 (2): 411-424. HANSEN, H.J. 1901. On six species of Koenenia, with remarks on the order Palpigradi. Ent. Tidskr. 22: 193-240. REMY, P. 1960. Palpigrades de la Région de Pondichéry (Inde). Bull. Mus. natn. Hist. nat. Paris, 2e serie, 32 (3): 230-234. — 1961. Les Palpigrades de Ceylan et leur écologie. Revue fr. Entomol. 28 (2): 112-119. | | | Revue suisse Zool. Tome 99 Fasc. 3 | p. 673-712 | Genève, septembre 1992 | New and interesting mites from the Geneva Museum LXIII. A survey of the Oribatid fauna of Senegal (Acari: Oribatida) by S. MAHUNKA* With 102 figures ABSTRACT Thirty Oribatid species are enumerated from Senegal, fourteen of them are new to science, two also representing new genera: Senilochthonius gen. n. (Haplochthoniidae) and Chaunoproctellus gen. n. (Chaunoproctidae). Taxonomic investigations are presented concerning ther family Haplochthoniidae (partial redescription of Haplochthonius simplex Willmann, 1930 and H. sanctaeluciae Bernini, 1973, and description of a new species from Greece) and the "areolata"-group in the genus Galumnella (with the redescription of G. subareolata Mahunka, 1969), furthermore, the description of a new genus and species: Trichogalumnella hauseri gen. et sp. n. (Galumnidae) from Rhodesia is presented. INTRODUCTION My research aiming at the exploration of the Oribatid fauna of the Ethiopian Region were mostly concentrated to East, Central and South Africa, since no adequate material originating from West Africa was available. This is why I am especially grateful to Dr. B. Hauser, curator of the Arthropoda Collection, Muséum d'Histoire naturelle, Geneva, for his kindness in allowing me to study a large material from Senegal, during my stay in Geneva in 1985. This material, collected by several collectors (R. Mussard, S. and P. Hainard, P. Strinati) at various times and collecting sites in Senegal, proved to be highly interesting and very rich, well beyond expectation. I have identified a total of 30 species, 14 of which are new to science, two also representing two new genera (Senilochthonius: Haplochthoniidae) * Zoological Department, Hungarian Natural History Museum, Baross utca 13, H-1088 Budapest, Hungary. 674 S. MAHUNKA and Chaunoproctellus: Chaunoproctidae). While studying this material several taxonomic problems have been encountered. Thus it was inevitable to revise all the taxa of Haplochthoniidae. The comparative studies revealed a new species, also representing a new genus, while two species are related to species known mostly from the Palaearctic Region. The species that I mentioned from Greece under the name Haplochthonius simplex, is in fact an independent, new species, whose description follows hereunder. The description of a new species in the genus Galumnella also focused my attention to some problems; consequently, I had to revise the whole "areolata"-group. I give a redescription of G. subareolata Mahunka, 1969. The species mentioned by me as G. areolata Balogh, 1961 from Rhodesia, is in fact a new taxon, for which the establishment of a new genus is necessary. Though the material is large, it is still insufficient for zoogeographical conclusions. Nevertheless, it is interesting to note that besides the Central African elements (Allonothrus monodactylus, Africacarus calcaratus) there ae also some species showing links with Palaearctic elements (Haplochthonius sanctaeluciae, Medioppia subpectinata, Xylobates lophotrichus). In the descriptions I generally apply the terminology used in NORTON & BEHAN- PELLETIER (1986) based on Grandjean's work. Measurements given correspond to extremes observed in the present material; length is measured from the rostral apex to the furthermost opposite point of the body. The pilosity of the parts of the body and of the legs are expressed in formulae. The sequence of the anogenital formula is: number of genital, aggenital, anal and adanal setae. LIST OF LOCALITIES Se-72/1 = Senegal, Rufisque (port 30 km de Dakar), au pied d'un Baobab (Adansonia digitata, Bombacacées), 21.11.1972. leg. R. Mussard. Se-72/2 = Senegal, Ziguinchor (Casamance), 12°30N. — 16°15W au Sud de la Gambie, 20.11.1972. leg. R. Mussard. Se-72/3 = Senegal, Rufisque (port 30 km de Dakar), vieux baobab pourri, tamisage et appareil Winkler, VII. 1972. leg. R. Mussard. Sen-73/1 = Sénégal, Rufisque, 2.VII.1973. leg. P. Strinati. Sen-76/1 = Sénégal, Casamance, Ziguinchor, sol sableux ferrugineux, forêt secondaire, env. 20 m, prélèvement de terre, 9.XI.1976. leg. S. et P. Hainard. Sen-76/2 = Sénégal, Casamance, Ziguinchor, sol sableux ferrugineux, forêt à Cola cordifolia, env. 20 m, prélèvement de terre, 9.XI.1976, leg. S. et P. Hainard. Sen-76/3 = Sénégal, Casamance, Ziguinchor, sol sableux-vaseux, salé, mangrove à Avicennia nitida, 9.X1.1976. leg. S. et P. Hainard. Sen-77/1 = Sénégal, Nianing, baobab pourri, 1.V.1977. leg. R. Mussard. Sen-77/2 = Sénégal, Rufisque, prélèvement de bois décomposé de baobab, 7.IX.1977. leg. R. Mussard. Sen-77/3 = Sénégal, Nianing, bois pourri de baobab, 7.IX.1977. leg. R. Mussard. Sen-77/4 = Sénégal, Nianing, (baobab pourri), 28.IX.1977. leg. R. Mussard. Hel-75/1 = Péloponnèse: au bord de la route de Krestena à Andritsena, 230 m, prélèvement de terre sous Acer monspessulanum, 19.1V.1975. leg. B. Hauser. The-76/25 = Grèce (Acarnanie): prélèvement de terre au pied de Quercus sp., près Astakos, 120 m, 15.V.1976, leg. B. Hauser. Rho-69/1 = Rhodesia: Inyanga, Umtali, 27.11.1969, leg. R. Mussard. ORBATID FAUNA OF SENEGAL 675 LIST OF IDENTIFIED SPECIES Haplochthoniidae van der Hammen, 1959 Senilochthonius baobab gen. n., sp. n. Locality: Sen-77/3. Haplochthonius graecus sp. n. Localities: The-76/25, Hel-75/1. Haplochthonius sanctaeluciae Bernini, 1973 Localities: Sen-76/2: 1 specimen, Sen-77/2: 2 specimens, Sen-77/3: 2 specimens, Sen-77/4: 2 specimens. Haplochthonius simplex (Willmann, 1930) Locality: Sen-77/3: 3 specimens. Mesoplophoridae Ewing, 1917 Mesoplophora africana Balogh, 1958 Locality: Sen-76/1: 2 specimens. Lohmanniidae Berlese, 1916 Torpacarus omittens Grandjean, 1950 Locality: Sen-76/1: 2 specimens. Epilohmanniidae Oudemans, 1923 Epilohmannia pallida Wallwork, 1962 Locality: Sen-76/1: 2 specimens. Nothridae Berlese, 1885 Nothrus senegalensis sp. n. Locality: Sen-76/2. Trhypochthoniidae Willmann, 1931 Allonothrus monodactylus Wallwork, 1960 Locality: Sen-76/2: 8 specimens. Malaconothridae Berlese, 1916 Malaconothrus heterotrichus sp. n. Locality: Sen-76/1. Damaeolidae Grandjean, 1965 Fosseremus quadripertitus Grandjean, 1965 Locality: Sen-76/1: 1 specimen. Oppiidae Grandjean, 1954 Graptoppia mussardi sp. n. Localities: Sen-77/1, Sen-77/4. Insculptoppia crenata sp. n. Locality: Sen-76/2. Karenella foveolata sp. n. Locality: Sen-76/2. Multioppia calcarata sp. n. Locality: Sen-77/2. Paroppia senegalensis (Mahunka, 1975) Localities: Se-72/1: 15 specimens, Se-72/2: 1 specimen, Se-72/3: 8 specimens, Sen-77/1: 2 specimens, Sen-77/2: 1 specimen, Sen-77/3: 5 specimens. 676 S. MAHUNKA Medioppia subpectinata (Oudemans, 1901) 5 Localities: Sen-77/3: 1 specimen, Sen-77/4: 2 specimens. Oppiella nova (Oudemans, 1902) Localities: Sen-73/1: 1 specimen, Sen-77/3: 3 specimens. Uroppia hainardorum sp.n. Locality: Sen-76/2. Chaunoproctidae Balogh, 1961 Chaunoproctellus rugosus gen. n., sp. n. Localities: Sen-77/1, Sen-77/2. Oribatulidae Thor, 1929 Baobabula mussardi Mahunka, 1975 Localities: Se-72/1: 4 specimens, Se-72/3: 11 specimens. Perscheloribates minimus sp. n. Locality: Se-72/2. Scheloribates exiguus sp. n. Locality: Sen-76/3. Scheloribates fimbriatus Thor, 1930 Locality: Sen-77/2: 10 specimens. Scheloribates laevigatus (C.L. Koch, 1836) Locality: Sen-77/3: 9 specimens. Haplozetidae Grandjean, 1936 Xylobates lophotrichus (Berlese, 1904) Locality: Sen-76/1:2 specimens. Ceratozetidae Jacot, 1925 Africacarus calcaratus Wallwork, 1965 Locality: Sen-77/3: 2 specimens. Oribatellidae Jacot, 1925 Oribatella ceylanica (Oudemans, 1915) Locality: Se-72/2: 1 specimen. Galumnidae Jacot, 1925 Allogalumna sinornata sp. n. Locality: Se-72/2. Galumna coronata sp. n. Locality: Se-72/2. Galumnella apiculata sp. n. Locality: Sen-76/1. Trichogalumnella hauseri gen. n., sp. n. Locality: Rho-69/1. DESCRIPTIONS The genus Haplochthonius Willmann, 1930 GRANDIEAN (1947) published a very good redescription of the genus in connection with his research on the system of the group Enarthronota. He examined two species which belong to this genus (H. simplex Willmann, 1930 and H. sanctaeluciae Bernini, ORBATID FAUNA OF SENEGAL 677 1973*) but he did not see the type specimens. BERNINI (1973), who described the second species, did not give any new data regarding the knowledge of the genus and he did not discuss the epimeral chaetotaxy. Since then some authors mentioned one of the two species, but no relevant data were disclosed. Both species belong to a uncommon group, therefore, it was very unexpected, to find three Haplochthonius species in these small samples, and even, in one sample, all three together. Because of the significant geographical distribution and the very special biotopes I compared these specimens with both species collected in the Mediterranean Region. I have found, that one species is unambiguously identical with H. sanctaeluciae and one species with H. simplex (sensu GRANDJEAN). The third specie stands very far from both preceding ones and the establishment of a new genus is inevitable for it. I re-examined also a series of specimens which was published by me (MAHUNKA 1977, 1982b) under the name of H. simplex from Greece, however, they did not prove to be conspecific with GRANDJEAN's species, therefore, I describe it as a new species. I examined in every case some of the important characters: the position of notogastral cupules, the epimeral and the anogenital chaetotaxy. I found, that among these species great differences exist: simplex graecus gen. n. sanctaeluciae (sensu Grandj.) form of not. setae simple simple simple widened position of cupule near near partly far near epimeral setal formula 3-2-3-4 3-2-3-3 3-2-3-3 3-2-2-3 number of genital setae U 7 9 i The variability of the epimeral setal formula is most remarkable and it querries the value of this character in other relatively primitive groups, as the family Brachych- thoniidae. The most important character is the number of the genital setae. On this ground it would be inconsequent to order the new species in the same genus, thus I establish a new genus. Senilochthonius gen. n. Diagnosis: Family Haplochthoniidae. Habitus and notogastral chaetotaxy similar to that of the genus Haplochthonius. Notogastral cupules originating in some cases far from the median setae (in the case of setae d, and h,). Epimeral setal formula: 3-2-3-3. Nine pairs of genital setae present. All legs with one claw. Type species: Senilochthonius baobab sp. n. * At that time as "espèce de Sainte-Lucie": GRANDJEAN 1947. 678 S. MAHUNKA Fics 1-5. Senilochthonius baobab gen. n., sp. n. — 1: dorsal side; 2: anogenital region; 3: notogaster from lateral view; 4: trichobothrium; 5: coxisternal region. Senilochthonius baobab sp. n. Measurements: — Length: 316 um, width: 119 um. Prodorsum: Weakly chitinized, no transversal ridge between the rostral and in front of the lamellar setae. All prodorsal setae thin, but ciliate. Sensillus (Fig. 4) ORBATID FAUNA OF SENEGAL 679 comparatively short, flabellate, its laminate head wide, the surface spiculate. Both pairs of exoborthridial setae long, they are not shorter than the interlamellar ones, lamellar setae three times longer than the length of the exobothridial setae. Notogaster: Cerotegument ornamented by very fine reticulation. All setae thin, comparatively short, their surface finely roughened; setae dj and e) not longer than half diameter of segments NM, or NM». The cupules originating far from setae c,, dj and hj, only at setae e) and fj they stand close to them (Fig. 1). Surface of tergites with some fine rugae, being stronger on the pygidium. Lateral margin of pygidium also waved. Pleural region (Fig. 3): Apophysis Te well separated, but not high. Lyrifissures im an ip originating near to notogastral tergites. Coxisternal region (Fig. 5): Epimeral setal formula: 3-2-3-3 (!). All setae long. Anogenital region (Fig. 2): Anogenital setal formula: 9-0-4-4. Material examined: Holotype: Sen-77/3. Holotype. MHNG!., Remarks: Asl discussed previously this new species stands very far from any known Haplochthonius Willmann, 1930 species. I give hereunder the description of the other examined species, being also new for science. Haplochthonius graecus sp. n. Measurements. — Length: 311-326 um, width: 157-165 um. Prodorsum: Weakly chitinized, but a fine transversal line observable in the lamellar relgion. All setae simple, nearly equal in length. Sensillus flabellate, compara- tively wide. Notogaster (Fig. 6): Notogastral setae very short (dj, e 39-45 um) but stick- shaped, their surface finely roughened. Notogastral cupule originating — with the exception of one pair on Na-tergite — very near to the median setae. Coxisternal region: Epimeral setal formula: 3-2-3-3. All setae simple (Fig. 9). Anogenital region: As shown in Fig. 7. Material examined: Holotype: The-76/25; 26 paratypes: from the same sample; 1 paratype: Hel-75/1. Holotype and 17 paratypes: MHNG, 9 paratypes (1156-PO-85): HNHM£. Remarks: The heretofore known Haplochthonius Willmann, 19303 species may be distinguished by the following key: 1 (2) Notogastral setae wide, setae an, also wider than other anal setae (Figs 12-14). sanctaeluciae Bernini, 1973 2 (1) Notogastral setae thin, filiform or stick-shaped. All anal setae similar to each other. 3 (4) Epimeral setal formula: 3-2-3-4, all setae long (Figs 15-20) simplex (Willmann, 1930) sensu GRANDJEAN, 1946 4 (3) Epimeral setal formula: 3-2-3-3, all setae short graecus sp. n. 1 MHNG = deposited in the Muséum d'Histoire naturelle, Genève. 2 HNHM = deposited in the Hungarian Natural History Museum, Budapest with identification number of the specimens in the Collection of Arachnida. 3 I have not investigated the Haplochthonius clavatus (Hammer, 1958) from South America. 680 S. MAHUNKA Fics 6-11. Haplochthonius graecus sp. n. — 6: dorsal side; 7: anogenital region; 8: tarsus of leg I; 9: coxisternal . region; 10: notogaster from lateral view; 11: pygidium. | | | | | ORBATID FAUNA OF SENEGAL 681 TA 14 UA (a ut: DTA) Fics 12-14. Haplochthonius sanctaeluciae Bernini, 1973 — 12: dorsal side; 13: coxisternal region; 14: tricho- bothrium. Torpacarus omittens Grandjean, 1950 Measurements. — Length: 631-648 um, width: 271-285 um. The specimens from Senegal are completely identical with the description and figures given by GRANDIEAN (1950), except some round areae porosael arranged in transversal bands on the notogaster, which are absent on the South American specimens. . But they are highly variable (Figs 21-22) and do not justify the separation of a new taxon. | The sutures mt, nt or pt also vary. Examined material: Sen-76/1: 2 specimens. Nothrus senegalensis sp. n. Measurements. — Length: 688-720 um, width: 322-348 um. Prodorsum: Rostral and lamellar setae arising on tubercles connected by transversal laths. Both pairs of setae and the interlamellar ones also ciliate, spathulate. 1 WALLWORK (1962) also observed them on his specimen collected in Ghana. 682 S. MAHUNKA Fics 15-20. mann, 1930 — 15: dorsal side; 16: coxisternal region; 17: tarsus of leg I; Haplochthonius simplex Will 9: notogaster from lateral view; 20: trichobothrium. 18: pygidium; 1 ORBATID FAUNA OF SENEGAL meer, = 7 nee | C £ i ° a 6 R S|, | nn Fics 21-22. Torpacarus omittens Grandjean, 1950 — 21-22: dorsal views of different specimens. 684 S. MAHUNKA 25 Fics 23-26. Nothrus senegalensis sp. n. — 23: dorsal side; 24: ventral side; 25: distal end of sensillus; 26: noto- gastral seta. Prodorsal surface areolate. Sensillus (Fig. 25) very long (240 um), much longer than distance between bothridia. Notogaster: Surface also areolate, but laterally pustulate. Notogastral setae — with the exception of k, — also dilated (Fig. 26). Significant differences in their lengths (Fig. 23) exist: c; more than twice as long as ©», pı (91 Him) nearly two and a half times longer than h,. Setae hy (275 um) thin, gradually narrowing distally, with a slightly flagellate end. Coxisternal region: Epimeral setal formula: 5-4-4-6 (it was observable only in one specimen!). All setae more or less dilated. ORBATID FAUNA OF SENEGAL 685 Anogenital region: Inner margin of genital plates rugose, some larger longitudinal rugae also observable (Fig. 24). Genital setae also dilated. Two pairs of anal, three pairs of adanal setae present, all dilated, however, an, much larger than the others. Legs: Alllegs monodactylous. Material examined: Holotype: Sen-76/2; 1 paratype: from the same sample. Holotype: MHNG, paratype (1157-PO-85): HNHM. Remarks: The new species belongs to the "palustris"-group, however, on the ground of the number of claws it stands nearest to Nothrus mystax Mahunka, 1985, but, the ratio of the notogastral setae in the latter one is different. Malaconothrus heterotrichus sp. n. Measurements. — Length: 414-429 um, width: 182-200 um. Prodorsum: Rostral and lamellar setae thick, but rostral ones smooth and thinner than the latter. Interlamellar setae thinner but longer than sensillus, both pairs ciliate basally. Two pairs of ridges observable, one stronger around the bothridium, bending inwards, the other weaker, laterally, directed to lamellar setae. Some large foveolae visible basally and medially (Fig. 29). Pori m well visible, large, insertion of exobothridial setae scarcely observable. Notogaster: All notogastral setae ciliate basally; with the exception of setae e,, hy and ps, all short, dilated on their basal part (Fig. 27). Setae eo, hy and ps, much longer than the others (Fig. 32). Lyrifissure ip opening always transversally. Coxisternal region: Setae A slightly dilated, blunt. Epimeral setae different in lengths: setae /c minute, /b much longer than Ja. Setae 3b and 3c and 4c slightly dilated, well ciliate. Cerotegument between the anterior and posterior sternal plates well granulate. Anogenital region: Five (sometimes six) pairs of dilated and slightly pilose genital setae present. One or two minute anal setae hardly recognizable. All three pairs of adanal setae strong, ad, much longer but thinner than ad, and ad3. Legs: Solenidium vj comparatively short, simply bent anteriorly. Setae d and 1’ of tibia I and the setae of genu and femur well dilated and ciliate basally (Fig. 33). These setae also dilated on the other legs. Material examined: Holotype: Sen-76/1; 8 paratypes: from the same sample. Holotype and 5 paratypes: MHNG, 3 paratypes (1158-PO-85): MNHM. Remarks: The new species belongs to a species group ("plumosus"-group) that can be characterized by the dialted and ciliate prodorsal or notogastral setae. These species are: plumosus Willmann, 1929 — Java robustus Hammer, 1958 — South America keriensis Hammer, 1966 — New Zealand neoplumosus Balogh & Mahunka, 1969 — South America variosetosus Hammer, 1971 — Fiji pachypilus Hammer, 1972 — Tahiti ensifer Mahunka, 1982 — Ethiopia 686 S. MAHUNKA Fics 27-33. Malaconothrus heterotrichus sp. n. — 27: dorsal side; 28: ventral side; 29: lateral part of prodorsum from dorsal view; 30: lateral part of prodorsum from lateral view; 31: seta dj; 32: seta eg; 33: leg I ORBATID FAUNA OF SENEGAL 687 On the ground of the notogastral heterotrichy the new species stands nearest to M. variosetosus, however, it is distinguished from the latter by the very thick lamellar and interlamellar setae, by the sensillus, and by the shape of adanal and genital setae. Graptoppia mussardi sp. n. Measurements. — Length: 196-200 um, width: 90-96 um. Prodorsum: Rostrum widely rounded, rostral setae arising on the dorsal surface, thicker and longer than the lamellar on interlamellar ones. Well developed costula present, narrowing basally. A convex transcostula present, but becoming thin medially. Three pairs of light spots present in the interlamellar region. Exobothridial setae thinner, but not shorter than interlamellar ones. Sensillus short, its head strongly, but asymmetrically clavate, with 10-11 lateral branches. Notogaster: Elongate. Ten pairs of notogastral setae present. Setae ta minute, all others short, nearly equal in length, stick-shaped, some of them, in posteromarginal position, arising from small tubercles (Fig. 34). Lateral part of podosoma: Pedotecta I small, II-III absent, discidium small, but sharply pointed and steeply projecting from the surface. Exobothridial region (Fig. 38) granulate. Coxisternal region: Borders between the 1. and 2. epimeres hardly observable, sejugal borders wide (Figs 36-37). Epimeral surface with some polygonal fields. Epimeral setae short, setae /c originating far from pedotecta 1, setae 3c and 4c (!) arising from tubercles. Between epimeres 3 and also 4 a wide median field present. Anogenital region: Anogenital setal formula 5-1-2-3, aggenital, adanal and anal setae nearly equal in length. Setae ad, in postanal, setae ad; in preanal position, the latter nearly in a transversal line along with the aggenital setae. Lyriffisure iad in adanal position. Legs: All solenidia of tarsus I (Fig. 35) short, w | arising on a tubercle. Solenidium w | of tibia II also short and blunt, directed laterally (Fig. 39). Material examined: Holotype: Sen-77/4; 1 paratype: Sen-77/1. Holotype: MHNG, paratype (1159-PO-85): MNHM. Remarks: The new species stands very near to G. foveolata (Paoli, 1908) and G. africana Mahunka, 1987. It is distinguished from both species by the presence of setae ta on the notogaster and by the number of lateral branches of the sensillus. Insculptoppia crenata sp. n. Measurements. — Length: 295-312 um, width: 164-171 um. Prodorsum: Ratio of prodorsal setae: ro>in>le>ex (Fig. 43).All setae ciliate, but setae ro also slightly thicker than the ohters. Dorsal surface with a pair of sharp lines running from bothridium towards lamellar setae. Between interlamellar setae 3 (4) pairs of irregular spots and among them a well-visible, short, longitudinal lath (Fig. 40) present. Sensillus fusiform, unilaterally ciliate. Ciliae (or branches) different in length. Notogaster: Nine pairs of short notogastral setae present, setae ta represented only by their alveoli. 688 S. MAHUNKA Fics 34-39. n. — 34: dorsal side; 35: leg I; 36: ventral side; 37: coxisternal region; 38: Graptoppia mussardi sp. prodorsum from lateral view; 39: leg II. ORBATID FAUNA OF SENEGAL 689 Fics 40-43. Insculptoppia crenata sp. n. — 40 dorsal side; 41: ventral side; 42: leg I; 43: prodorsum from lateral view. 690 S. MAHUNKA Lateral part of prodorsum: Surface well granulate. Pedoctecta 1 small, pedotecta 2 absent, discidium without sharp spur. Setae /c originating far from pedotecta 1. Coxisternal region: Epimeral borders well observable. Sejugal borders with a pair of characteristic tubercles, directed backwards. Epimeral surface ornamented by polygonal network. Epimeral setae short and simple (Fig. 41). Anogenital region: Anogenital setal formula: 5-1-2-3. All setae simple and short. Setae ad; originating in postanal position and directed slightly outwards. Legs: Tibia of leg I (Fig. 42) without spur. Solenidium w | long, w > blunt, vj also blunt but v7, filiform, much longer than v ;. Material examined: Holotype: Sen-76/2; 2 paratypes: from the same sample. Holotype and 1 paratype: MHNG, 1 paratype (1160-PO-85): MNHM. Remarks: The new species stands nearest to /. fusiformis (Wallwork, 1961) from Ghana, however, the latter has no median laths between the light spots of the interbothridial region and the head of sensillus is narrower than in the new species. Karenella foveolata sp. n. Measurements. — Length: 271-300 um, width: 157-174 um. Prodorsum: Rostrum widely rounded, rostral setae arising laterally far from each other. All prodorsal setae simple, setiform, setae in minute, setae ex represented only by their alveoli. Between the interlamellar setae a characteristic formation present, which consists of one pair of short, longitudinal laths and between them two pairs of round spots. Sensillus long, its head asymmetrically clavate, barbed distally. Notogaster: Its surface ornamented by large but shallow foveolae, their margin hardly observable, indistinct. Ten pairs of setae present, nine pairs of them characteristically widened basally (Fig. 44), and barbed distally, one pair (ta) minute, originating very near to lyrifissura ia. Lateral part of podosoma: Exobothridial region granulate, some stronger rugae also observable (Fig. 47). Pedotecta 1 normal, discidium weakly developed, without sharp spur. Setae 4c originating very far from the acetabulum of leg IV. Setae Jc arising also on the epimeral surface (Fig. 45). Coxisternal region: A strong sejugal band observable, other epimeral borders — with the exception of bo, — not or only partly observable. Epimeral surface ornamented by a few polygonal fields or spots. Epimeral setae different in length, but all thin and simple. Anogenital region: Anogenital setal formula 5-1-2-3. Seate ad, like notogastral ones, dilated basally, all others thin and simple. Legs: Tibia of leg I with long and strong spur dorsally, both solenidia arising on it (Fig. 46). All solenidia of tibia and tarsus of leg II blunt, directed forwards. Material examined: Holotype: Sen-76/2; 1 paratype: from the same sample. Holotype: MHNG, 1 paratype (1161-PO-85): HNHM. Remarks: Thenew species is well characterized by its notogastral sculpture and by the shape of its notogastral setae. On this around it may be well distinguishable from all related taxa. In my opinion Karenella lanceosetoides (Balogh, 1960) and the new species ORBATID FAUNA OF SENEGAL 691 Fics 44-47. Karenella foveolata sp. n. — 44: dorsal side; 45: ventral side; 46: leg I; 47: prodorsum from lateral view. 692 S. MAHUNKA present transitional forms from Karenella Hammer, 1962 to Corynoppia Balogh, 1983, therefore, the latter probably will have to be synonymized with Karenella, and Karenella should be placed close to the Stachyoppia — Striatoppia group. Multioppia calcarata sp. n. Measurements. — Length: 271-302 um, width: 147-158 um. Prodorsum: Rostrum widely rounded, rostral setae geniculate, originating near to each other on the drosal surface. In front of them a transversal lath present. Lamellar setae slightly shorter and thinner than interlamellar ones. Sensillus asymmetrically clavate, with 9-10 long branches and its peduncle with 7-8 short cilia on each side. Exobothridial region (Fig. 53) well granulate. Notogaster: Twelve pairs of characteristic notogastral setae present (Fig. 48), setae ta reduced, their insertion also invisible. Lateral part of podosoma: Pedotecta with a very long and strong spur anteriorly (Fig. 51). Pedotecta II small, discidium sharply pointed, slightly curved backwards. Coxisternal region: All epimeral setae comparatively short, some of them ciliate. Setae /c originating far from pedotecta I. Epimeral surface ornamented by polygonal reticulation, epimeral borders in parts hardly observable. On the sejugal borders one pair of round tubercles (Fig. 49) present, directed posteriorly. Anogenital region: Allsetae short and simple. Anogenital setal formula: 5-1-2-3. Lyrifissure iad long. Legs: Tibia of leg I (Fig. 52) without tubercles. Solenidia w ; and w 5 standing far from each other. Solenidium w | of leg II (Fig. 50) long, filiform. Material examined: Holotype: Sen-77/2; 22 paratypes: from the same sample. Holotype and 14 paratypes: MHNG, 8 paratypes (1162-PO-85): MNHM. Remarks: The new species is well characterized by the anterior spur of its pedotecta I and the completely reduced setae ta. On this ground it may be well distin- guished from all heretofore known Multioppia Hammer, 1961 species. Paroppia senegalensis (Mahunka, 1975) comb. nov. This species belongs without doubt to the genus Paroppia Hammer, 1968. On the ground of the newly examined specimens I give some complementary figures (Fig. 54-57). The original description is acceptable, but the prodorsal and notogastral setae are slightly more rigid than they have been figured (MAHUNKA, 1975: 289, fig.: 1-2). Examined material: Se-72/1: 15 specimens, Se-72/2: 1 specimen, Se-72/3: 8 specimens, Sen-77/1: 2 specimens, Sen-77/2: 1 specimen, Sen-77/3: 5 specimens. Uroppia hainardorum sp. n. Measurements. — Length: 369-395 um, width: 209-225 um Prodorsum: Rostrum conical, rostral setae arising near to the rostral apex, close to each other, curved inwards. Lamellar and interlamellar setae similar, nearly equal ORBATID FAUNA OF SENEGAL 693 Fics 48-53. Multioppia calcarata sp. n. — 48: dorsal side ; 49: ventral side; 50: leg II; 51: pedotecta 1; 52: leg I; 53: prodorsum from lateral view. S. MAHUNKA 694 Fics 54-57. Paroppia senegalensis (Mahunka, 1975) — 54: ventral side; 55: leg I; 56: leg II; 57: prodorsum from lateral view. ORBATID FAUNA OF SENEGAL 695 Fics 58-63. Uroppia hainardorum sp. n. — 58: dorsal side; 59: epimeral borders and genital plate; 60: ventral side; 61: leg I; 62: trichobothrium; 63: prodorsum from lateral view. 696 S. MAHUNKA in length (Fig. 63). A strong transversal costula in front of lamellar setae and a sharp longitudinal line present, the latter running anteriorly from bothridium, slightly conver- gent. Between interlamellar setae some short laths present. Senillus (Fig. 62) gradually thickened, with long branches of various lengths. Exobothridial region strongly granulate. Notogaster: Ten pairs of notogastral setae present, setae fa originating far from dorsosejugal suture, setae p standing near to each other. Setae te — r3 well ciliate (Fig. 58). Lateral part of prodorsum: Lateral margin of prodorsum bent characteristically posteriorly. Setae /c originating far from pedotecta 1 and much shorter than 3c and 4c. Discidium small. Coxisternal region (Fig. 60): Apodemes and epimeral borders well observable, composing a dense network. Setae — with the exception of 3c and 4c — short and simple, the latter two pairs with long cilia. Epimeres ornamented by irregular spots. Anogenital region: Genital plates (Fig. 59) hollowed out at their anterior median margin. Anogenital setal formula: 5-1-2-3. Genital and aggenital setae simple, thin, all others thicker and with strong cilia. Setae ad; originating far from ad), anteriorly, only slightly behind setae ag. Lyrifissure iad in adanal position, but near to the posterior corner of anal plates. Legs: All tarsi (Fig. 61) gradually narrowed anteriorly, without a bulbiform basal part. Claws short, comparatively small. Tibia of leg I without process. Material examined: Holotype; Sen-76/2; 1 paratype: from the same sample. Holotype: MHNG, paratype (1163-PO-85): HNHM. Remarks: The new species stands very near to the type species of the genus Uroppia Balogh, 1983 [U. acusiensis (Wallwork, 1961)] described from Ghana. They differ from each other by the following characters: U. acusiensis! U. hainardorum sp. n. 1. Smaller species: measurements: 1. Bigger species: measurements: 312-341 x 175-206 um 369-395 x 209-225 um 2. Setae 4b ciliate 2. Setae 4b smooth. 3. Setae ad; arising nearer to 3. Setae ad; arising farther from apodemes 4 than setae ag. apodemes 4 than setae ag. 4. Lyrifissure iad in apoanal 4. Lyrifissure iad in adanal position. position. Chaunoproctellus gen. n. Diagnosis: Family Chaunoproctidae. Similar to Chaunoproctus. Lamellae with very large, wide cuspis, translamella narrow. Dorsosejugal suture interrupted medially. Ten pairs of notogastral setae and five pairs of pori present. Epimeral setal formula: 3-1-3-3. Anal and genital apertures originating far from each other, the distance being greater than the length of anal plates. Anogenital setal formula: 6-1-2-2. Lyriffisure iad in adanal position. All legs tridactylous. Type species: Chaunoproctellus rugosus sp. n. 1 Based on WALLWoRK's description only. ORBATID FAUNA OF SENEGAL 697 FIGs 64-67. Chaunoproctellus rugosus gen. n., sp. n. — 64: dorsal side; 65: lamellar region; 66: notogastral seta; 67: humeral part of notogaster. Remarks: The new taxon stands near to the genus Chaunoproctus Pearce, 1906 btu the latter has three pairs of adanal setae and its dorsosejugal suture is not interrupted medially. Chaunoproctellus rugosus sp. n. Measurements: — Length: 397-494 um, width: 276-350 um. Prodorsum: Rostrum conical. Rostral setae long, thin, arising on the cuspis of tutorium. Lamellae thick, of a complicate structure and sculpture (Fig. 65). Lamellar setae 698 S. MAHUNKA arising on their cup-shaped cuspis, bacilliform, spiculate. Rostral region, before trans- lamellae, ornamented by polygonal sculpture, interlamellar region smooth. Interlamellar setae also spiculate, slightly thicker than lamellar ones (Fig. 70). Bothridium protruding laterally, sensillus short, its head clavate and spiculate. Notogaster: A small humeral projection present, its surface and a longer posteriorly directed band (Fig. 67) rugose. Whole surface irregularly foveolate. Ten pairs of dilated notogastral setae (Fig. 64) present, setae ps, and ps; much shorter than the others. Five pairs of pori present, anterior one (Pa) longer than the others. Lateral part of podosoma: Pedotecta 1 an 2 well developed, its surface also polygonate. Among the lateral porose areae only the humeral one (Ah) visible, instead of the sublamellar one (A/) only a light spot visible. The whole surface of this region rugose. Coxisternal region (Fig. 68): Only a short part of apodeme 1 and the sejugal one is visible. Epimeral borders also absent. All setae thin, comparatively long, surface ornamented by irregular spots. Anogenital region: Surface foveolate, but the anterior part, near to the genital aperture, with some irregular spots, similar to the epimeral surface. Anogenital setal formula: 6-1-2-2, all setae short and thin, setae gg characteristically bent inwards. Legs: All legs tridactylous. All trochanters and femora finely rugose or striolate, the other segments smooth. The third and fourth tibia very long, therefore the posterior two pairs of legs much longer than the anterior pairs. Porose areae visible. Tibia of leg I with long process (Fig. 69), bearing solenidium |, @ originating also on a small tubercle. Material examined: Holotype: Sen-77/2: 1 paratype: from the same sample, 1 paratype: Se-72/2. Holotype and 1 paratype: MHNG, 1 paratype (1164-PO-85): HNHM. Remarks: In addition to the generic characters, the new species may also be well separated from the other species belonging to the family Chaunoproctidae by the rugose humeral projection and the shape of lamellae. Baobabula mussardi Mahunka, 1975 In the original description of the genus I erroneously referred to the areae porosae as respiratory organs. I must correct this: It has four pairs of large sacculi. I remarked that the "sacculi" are very hardly observable, because the "sacks" are mostly round and have a struture like pori (connections of tracheae?) well visible in them (Figs 71-72). In spite of this change the validity of the genus is unambiguous, but it might belong to the alliance of Constrictobates Balogh et Mahunka, 1966. Perscheloribates minimus sp. n. Measurements. — Length: 239-281 um, width: 135-177 um. Prodorsum: Rostrum obtuse, rostral setae arising far from each other, in marginal position. Lamellae well developed, a pair of short, bent interlamellar lines present, prelamellae short, not reaching to the insertion of rostral setae (Fig. 75). A transversal band observable behind the rostral setae. Sensillus large, directed outwards, clavate, its head rarely spinose. ORBATID FAUNA OF SENEGAL 699 Fics 68-70. Chaunoproctellus rugosus gen. g. n., sp. n. — 68: ventral side; 69: leg I; 70: prodorsum from lateral view. Notogaster: Dorsosejugal suture arched. Notogastral setae reduced, only setae Dj visible, all others represented by their alveoli. Four pairs of minute sacculi present. Lateral part of podosoma: As shown in (Fig. 77). Coxisternal region: Some spots on the epimeral surface visible. Epimeral setae simple, setae /c arising on the outer border of epimer 1. 700 S. MAHUNKA Fics 71-74. Baobabula mussardi Mahunka, 1975 — 71: notogaster; 72: sacculi Sh and seta te Africacarus calcaratus Wallwork, 1965 — 73: posterior part of notogaster; 74: coxisternal region. Anogenital region: Allsetae short and thin, no essential difference among their lengths (Fig. 76). Material examined: Holotype: Se-72/2; 6 paratypes: from the same sample. Holotype and 4 paratypes: MHNG, 2 paratypes (1165-PO-85): HNHM. Remarks: The difference among the genera Perscheloribates Hammer, 1973, Ischeloribates Corpus-Raros, 1980 and some other related genera is rather uncertain, however, the new species strongly resembles Perscheloribates clavatus Hammer, 1973, therefore I placed it in this genus. It differs from the type species by the presence of setae Pp}, the short prelamellae and the straight transversal line behind the rostral setae (the latter is well arched in clavatus). ORBATID FAUNA OF SENEGAL 701 Fics 75-77. Perscheloribates minimus sp. n. — 75: dorsal side; 76: ventral side; 77: prodorsum from lateral view. Scheloribates exiguus sp. n. Measurements. — Length: 394-420 um, width: 204-227 um. Prodorsum: Rostrum apex truncate. Ratio of prodorsal setae ro, portent à considérer les eukoenéniens comme des formes ayant subi une contraction du développement postembryonnaire, attestée par la perte d'un stade et l'absence définitive de vésicules. Dans la lignée prokoenénienne, l'acquisition d'une troisième paire de fusules sur le premier volet génital du mâle et l'absence d'organe frontal, propres à Triadokoenenia, seraient des caractères dérivés uniques dans le groupe (autapomorphies). Des six genres actuels de Palpigrades, A/lokoenenia est, de loin, le plus mal connu, car il ne repose que sur deux femelles et un juvénile de Guinée, et n'a jamais été retrouvé. C'est pourquoi, à l'occasion de cette très brève revue des principaux types de Palpigrades, il nous a semblé utile de faire connaître un spécimen juvénile encore inclassable avec certitude, mais plus proche d’Allokoenenia que de tout autre genre, découvert dans une grotte de Thaïlande. Ce spécimen et celui faisant l'objet de l'addendum sont déposés au Muséum d'Histoire naturelle de Genève, Département des Arthropodes et d'Entomologie II. La mise au propre de l'illustration est due au talent du regretté M.J. Chevelu pour la figure 1 et à celui de Mme E. Seraoui pour la figure 2 et des compléments à la figure 1; nous lui adressons nos vifs remerciements. ? Allokoenenia sp. Thaïlande. Province de Hua Hin, chef-lieu de Pran Buri, village de Ban Khung Tanot (massif de Kao Sam Roi Yot). Grotte de Tham Sai, réseau fossile de 250 m environ, avec avens d'effondrement, milieu humide, 26.VII.87, P. Leclerc leg.: 1 immature A. Longueurs. — Corps: 0,64 mm (extension); flagelle 0,20 mm; bouclier prosomien: 0,17 mm; basitarse IV: 62,5 um; patte IV, à partir du tibia: 0,20 mm; B/bta = 2,77; bta/ti = 0,91. Prosoma. — Organe frontal médian environ 1 fois 1/2 aussi long que large (1,57), ses branches écartées l'une de l'autre, élargies en palette et terminées en une pointe mousse. Un seul élément fusiforme, acuminé, à chaque organe latéral. PALPIGRADE DE THAILANDE 743 Vil D 50 um (A-D) 100 pm (E) FIG. 1. ? Allokoenenia, immature A de la grotte de Tham Sai, Thaïlande: A. Organe frontal médian. B. Phanère de l'organe latéral. C. Basitarse de la patte locomotrice IV gauche. D. Basitarse de la patte locomotrice IV droite. E. Opisthosome, face sternale, à partir du VIe segment, et trois articles du flagelle. Explication des lettres dans le texte. 744 BRUNO CONDE Bouclier portant des soies minuscules, non dénombrables avec certitude en vue ventrale. Segment libre avec 3+3 phanères, l'intermédiaire (t,) beaucoup plus long que le latéral (tz) (92/63). Deux courtes soies deuto-tritosternales dans le plan sagittal. Chélicères avec 7 dents aux mors. Patte locomotrice IV. Longueurs relatives des articles: ti = 79; bta = 72; ta, = 39; ta, = 48. Le basitarse est un peu plus court que le tibia (72/79, bta/ti = 0,91) et, mesuré au niveau de r, un peu plus de 3 fois (3,13) aussi long que large; la soie raide (r) est épaisse, égale aux 4/5 environ du bord tergal de l'article (94/121,5; t/r = 1,29), et est insérée un peu au-delà des 2/5 proximaux de ce bord (45/121,5; t/er = 2,70); les soies sternales distales (esd) sont de longueurs comparables (62,67). A droite, il existe un quatrième phanère, inséré plus distalement que r, qui peut représenter soit la soie grêle tergale (grt), soit la soie grêle latérale (g/a), l'une comme l'autre normalement absentes chez l'immature A. Opisthosoma. — Tergites III à VI avec une paire de phanères latéraux (f;), égaux au 1/4 environ de leur écartement (24/95) en III et au 1/8 seulement (13/106) de IV a VI. Une paire de soies s, plus longues que les #; (40-44), de III a V. Sternite II avec une rangée de 2+2 soies longues et gréles, sternite III avec 2+2 soies, les latérales beaucoup plus longues que les sublatérales, Sternites IV à VI avec chacun deux paires de poils (a], a) un peu plus épais que les autres phanères sternaux; les a, sont légèrement plus courts que les a, (38/42) et en moyenne une fois et demie plus courts que leur écartement (1,42-1,60). Les segments IX à XI sont de largeur subégale, beaucoup plus étroits que le VIII. Les IX et X portent chacun un verticille de 4+4 longues soies; le XIe n'en possède que 3+3, celles de la paire tergale plus de 5 fois plus courtes (5,2) que les autres. Le XIe segment est extraordinairement allongé, un peu plus de 2 fois 1/2 aussi long que large (2,66) et 2 fois plus long que les IX et le Xe ensemble. Face sternale, vers le tiers antérieur, on voit une dépression subcirculaire à parois plissées vers le centre de laquelle s'ouvre un orifice légèrement ovalaire. Une telle formation, de nature glandulaire sans doute, n'a jamais été signalée. L'anneau basal supportant le flagelle est dépourvu de phanères. Le flagelle compte 7 articles dont les longueurs relatives sont 15, 14, 17, 16, 16, 19, 6. Les deux premiers sont moniliformes, les suivants plus allongés, le 6e surtout, le dernier étant minuscule. Aucun ne possède de verticille subapical d'expansions spiniformes, mais seulement 6 ou 7 soies, grêles et courtes sur les 5 premiers, et 6 très longues sur les 2 derniers. DISCUSSION ET CONCLUSION La série typique d’Allokoenenia, sans désignation d'holotype, se compose de deux femelles adultes et d'un individu juvénile au dernier stade (B) récoltés dans l'humus à Mamou, Guinée française. Faute d'élément de comparaison, le stade A de l'unique espèce, Allokoenenia afra, étant inconnu, l'attribution générique du spécimen thaïlandais ne peut être rigoureusement établie et il serait également imprudent de lui attribuer un nom spécifique. On retiendra cependant que la forme générale des segments IX a XI est conforme à la définition du genre: «Abdominis segmentum nonum undecimo latitudine subaequale». Toutefois, l'allongement du XIe est unique chez les Palpigrades, de même que la formation, présumée glandulaire, de la face sternale du même segment; on objectera qu'il pourrait s'agir de caractères transitoires propres à la larve À, décrite pour la première fois. L'aspect du flagelle supporterait aussi l'attribution suggérée, par comparaison avec la figure VI (5) de SILVESTRI (1913). PALPIGRADE DE THAILANDE 745 FIG. 2. ? Allokoenenia, immature À de la grotte de Tham Sai, Thaïlande. Articles 4 à 7 du flagelle. Aucun argument biogéographique ne peut infirmer la détermination proposée ici sous réserve, bien au contraire, car beaucoup de genres de Palpigrades ont une vaste répartition dans la zone intertropicale. Koeneniodes, par exemple, décrit comme Allokoenenia de Guinée (Kakoulima, non loin de Mamou) est bien représenté dans la région madécasse et dans la région orientale, et le spécimen étudié ici a été récolté avec un Koeneniodes inédit dont la description est en cours de publication! . L'examen, presque 80 ans après la description d'Allokoenenia, d'un spécimen qui pourrait lui appartenir et serait alors son quatrième représentant connu, est révélateur de l'état de nos connaissances sur le groupe, d'autant que le cas n'est pas isolé, beaucoup d'espèces n'ayant jamais été revues depuis leur découverte. La faible densité des populations de Palpigrades, en général, et plus encore leur extrême fragilité lors de la capture et du transport, explique qu'aucun élevage n'ait été mené à bien et que leur tenue en captivité n'ait pas excédé quelques semaines (P. Weygoldt comm. pers.). ADDENDUM Cette note était déjà rédigée lorsque P. Leclerc m'a communiqué un second représentant de l'espèce étudiée ici. J'en propose donc une brève description comparative. ! Koeneniodes leclerci Condé 1992, Revue suisse Zool. 99(3): 666. 746 BRUNO CONDE Thaïlande. Province et District de Yala, près du village de Ban Na Tham, Grotte de Tham Sam Pao To, 14.7.91, P. Leclerc leg.: 1 mâle juvénile (C). Longueurs. — Corps: 0,95 mm; flagelle manquant; bouclier prosomien: 0,26 mm; basitarse IV: 102 um; patte IV, à partir du tibia: 0,32 mm; B/bta = 2,55; bta/ti = 0,93. Prosoma. — 2 éléments à chaque organe latéral. Phanères f, à #3 du segment libre: 62, 57, 48. Deuto-tritosternum: 2+2 soies. Chélicères: 8 dents à chaque mors. Basitarse IV avec 6 phanères (une seule esp): t/r = 1,48; t/er = 2,43; gla/grt (insérées presque au même niveau) = 0,79. Opisthosoma. — Aire génitale correspondant à la variante 3 (CONDE 1984: 387). Une paire de soies s de IV a VI, un peu plus courtes que les a, et a, subégaux (27/34,5). Le segment XI presque 2 fois 1/2 plus long que large (2,4) et une fois 1/2 plus long que les IX et X ensemble (1,52). La dépression à parois plissées, décrite chez l'immature A, est clairement visible. On notera que bien que les deux spécimens proviennent de grottes relativement profondes, humides et abritant des troglobies, ils ne présentent aucun des caractères liés à ce milieu. BIBLIOGRAPHIE BÔRNER, C. 1901. Zur äusseren Morphologie von Koenenia mirabilis Grassi. Zool. Anz. 24: 537-556. CONDE, B. 1965. Présence de Palpigrades dans le Milieu Interstitiel Littoral. C.R. Acad. Sci. Paris, 261: 1898-1900. ConDe, B. 1984. Palpigrades (Arachnida) d'Europe, des Antilles, du Paraguay et de Thaïlande. Revue suisse Zool. 91 (2): 369-391. CONDÉ, B.V.N. 1988. Palpigradida in: Introduction to the Study of Meiofauna, R.P. Higgins and H. Thiel eds. Smithsonian Institution Press, Washington D.C., London: 425-427. CONDE, B. 1991. Prokoenenia millotorum Remy, type du nouveau genre Triadokoenenia (Arachnida, Palpigradida). Bull. Mus. natl. Hist. nat., Paris, 4e ser. 13, sec. A (3-4): 351-360. GRASSI, B. 1885. Intorno ad un nuovo Aracnide Artrogastro (Koenenia mirabilis) che crediamo rappresentante d'un nuovo ordine (Microteliphonida). Natural. Siciliano, 4: 127-133, 162-168. HANSEN, H.J. 1901. On six species of Koenenia, with remarks on the order Palpigradi. Ent. Tidskr. 22: 192-240. MILLOT, J. 1942. Sur l'Anatomie et l'Histophysiologie de Koenenia mirabilis Grassi (Arachnida Palpigradi). Revue fr. Ent. 9 (2): 33-51. MILLOT, J. 1943. Notes complémentaires sur l'Anatomie, l'Histologie et la Répartition géographique en France de Koenenia mirabilis Grassi (Arachnida Palpigradi). Revue fr. Ent. 9 (3- 4): 127-135. MonNIOT, F. 1966. Un Palpigrade Interstitiel. Leptokoenenia scurra n. sp. Revue Ecol. Biol. Sol. 3 (1): 41-64. PEYERIMHOFF, P. de. 1902. Découverte en France du genre Kaenenia (Arachn. Palpigradi). Bull. Soc. ent. Fr. 1902: 280-283. ROWLAND, J.M. and W.D. Sıssom, 1980. Report on a fossil Palpigrade from the tertiary of Arizona, and a review of the morphology and systematics of the order (Arachnida: Palpigradida). J. Arachnol. 8: 69-86. RUCKER, A. 1901. The Texan Koenenia. Amer. Nat. 35 (416): 615-630. SILVESTRI, F. 1913. Nuovi generi e specie di Koeneniidae. Boll. Lab. Zool. gen. e agr., Portici, 7: 211-217. WHEELER, W. 1901. A singular arachnid (Koenenia mirabilis Grassi) occuring in Texas. Amer. Nat. 34: 837-850. Revue suisse Zool. Tome 99 Genève, décembre 1992 Fasc. 4 p. 747-754 | Les dents des sous-familles Hypostominae et Ancistrinae (Pisces, Siluriformes, Loricariidae) et leur valeur taxonomique ” par Sonia MULLER*** et Claude WEBER*** ABSTRACT The teeth of the subfamilies Hypostominae and Ancistrinae (Pisces, Siluri- formes, Loricariidae) and their taxonomical value. — Tooth number and shape vary greatly among hypostomines and ancistrines. Some extreme forms characterize taxa. Morphologic and meristic studies of teeth of Hypostomus Lacépède and Ancistrus Kner indicate that differences at species or species group level may be obtained among the numerous standard toothed species. INTRODUCTION Selon la classification d'ISBRÜCKER (1980 modifiée), plus de 600 espèces réparties en 90 genres forment la famille néotropicale très diversifiée des Loricariidae. Deux sous- familles intéressent cette étude, les Hypostominae et les Ancistrinae, qui regroupent chacune près du tiers des espèces. La plupart d'entre elles sont connues de descriptions anciennes à caractère général. L'étude morphologique des espèces de Hypostomus et d'Ancistrus nous a amené à observer, notamment sur le matériel type, les caractères dentaires qui sont souvent négligés dans ces descriptions. Seules quelques formes remarquables de dents caractérisent depuis longtemps certains taxa, comme Cochliodon Heckel, 1854. Plus récemment, GOSLINE (1947: 85) souligne l'intérêt de la denture à un niveau plus élevé et rapproche par une forme "ster&otypee" de dent les Plecostominae (Hypostominae et Ancistrinae), les Hypopto- pomatinae et les Loricariinae. Actuellement, les auteurs omettent rarement la description de la denture qui, comme le rappelle SCHAEFER (1986: 9, 30), montre une grande variation de nombre et de forme entre les espèces. * Travail présenté à Zoologia 92. ** Recherche bénéficiant du soutien du Fonds national suisse de la recherche scientifique, requête n° 31-9443.88. *** Muséum d'Histoire naturelle, case postale 434, CH-1211 Genève 6. 748 SONIA MULLER ET CLAUDE WEBER Selon la classification suivie, Cochliodon est le seul genre qui se distingue des autres Hypostominae par une grosse dent en forme de cuillere. Chez les Ancistrinae, qui comptent quatre tribus, les Pseudacanthicini regroupent des espèces à prémaxillaires solidaires et à denture réduite. Parmi les Ancistrini, plusieurs genres sont aussi caractérisés par leur denture particulière: Panaque Eigenmann & Eigenmann, 1889 (comparable à celle de Cochliodon) et, récemment décrits, Scobinancistrus Isbrücker & Nijssen, 1989, Oligancistrus Rapp Py-Daniel, 1989 ainsi que Hypancistrus Isbrücker & Nijssen, 1991. Toutes ces formes se dégagent nettement de la denture courante, présente chez la majorité des espèces de Hypostominae et d' Ancistrinae, qui n'a fait jusqu'à présent l'objet d'aucune étude morphologique comparative. La dent "standard" est généralement donnée filiforme, en forme de Z et bicuspidée. En dépit de son uniformité apparente, il nous a semblé utile de l'étudier chez Hypostomus Lacépède, 1803 et chez Ancistrus Kner, 1854, genres riches (respectivement 140 et 54 espèces décrites) et largement distribués. MATÉRIEL ET MÉTHODES Nous avons examiné les dents de plus d'un millier de spécimens dont les types de la grande majorité des espèces nominales des genres Hypostomus et de toutes celles d'Ancistrus sauf quatre, qui sont introuvables ou disparus. De nombreuses investigations ont été faites dans les genres voisins. Les dents sont dénombrées sur les deux mâchoires. Celles érigées étant généralement usées ou brisées, ce sont des dents de remplacement matures relevées ou prélevées de la pre- mière rangée au centre de la mandibule gauche qui sont choisies pour l'étude morphologique. Beaucoup ont été figurées et mesurées, au moyen d'un projecteur de profil NIKON V12 ou d'un microscope WILD M20, et photographiées à l'aide d'une loupe binoculaire NIKON SMZ10 équipée d'un boitier FX 35A et d'un appareil de mesure de pose UFX 12. La figure 1 présente les termes utilisés et les mesures prises sur une dent standard. VARIATION ET ONTOGÉNIE Nombre: Chez les Hypostomus examinés, le nombre de dents d'une mâchoire varie de 6 (chez H. roseopunctatus Reis, Weber & Malabarba) à 125 (Hypostomus aff. regani (Ihering), LS: 187 mm) et chez Ancistrus de 18 (Ancistrus groupe bufonius, LS: 16 mm) à près de 130 (A. megalostomus Pearson, LS: 81 mm). Pour la population de Hypostomus auroguttatus Natterer & Heckel étudiée et pour toutes celles d'Ancistrus, le nombre de dents augmente en fonction de la taille des spécimens comme il l'avait été montré chez deux espèces (MULLER 1990). Cette augmentation, qui diminue ou cesse pour une taille propre à chacune, est plus importante chez les espèces à dents nombreuses. Pour la majorité des populations examinées, le nombre moyen de dents comptées sur le dentaire est semblable à celui du prémaxillaire chez les individus de même taille. Ce n'est toutefois pas le cas pour quelques populations d'Ancistrus à dents en nombre élevé. Forme: Pour les différentes espèces des deux genres, les dents ne subissent pas de changement de forme majeur durant l'ontogénie. A l'exception de quelques cas (voir ci- après), la dent possède toujours une couronne comportant une cuspide majeure et une cuspide mineure du côté externe de la mâchoire. La couronne a une taille très variable à l'intérieur de chacun des genres étudiés. DENTS DE HYPOSTOMUS ET D'ANCISTRUS 749 TABLEAU |. Constantes d'allométries de la hampe (b1) et de la couronne (b2) en fonction de la LS, de la couronne en fonction de la hampe (b3) (régressions en coordonnées logarithmiques); rapports de la hampe dans la LS (rl) et de la couronne dans la hampe (r2) (grossissement 56x) pour 4 populations d'espèces Hypostomus auroguttatus Natterer & Heckel (n=15; LS=78-135 mm), H. aff. derbyi (Haseman) (n=13; LS=75-151 mm), Ancistrus cf. bufonius (Valenciennes) (n=6; LS=43-81 mm) et A. hoplogenys (Günther) (n=10: LS=30-109 mm). espece constantes d'allométries rapports morphométriques | b3 | r2 H. auroguttatus 2.11-2.67 (2.38; 0.18) | 1.00-1.41 (1.22: H. aff. derbyi 2.27-3.26 (2.69; 0.32) 1.39-1.88 (1.65; A. cf. bufonius 2.07-2.59 (2.29; 0.20) 1.23-1.42 (1.28; A. hoplogenys 1.76-2.75 (2.16; 0.32) 2.41-3.62 (2.84; La hauteur de la hampe et la longueur de la couronne, caractères morphometriques utilisés pour la comparaison des espèces, ont une croissance allométrique différente selon les populations étudiées (voir tableau 1). Dans chaque genre, les espèces choisies sont éloignées et présentent des caractères dentaires distincts. Hypostomus auroguttatus se caractérise par une allométrie négative particulièrement forte pour la longueur de la couronne dentaire par rapport à la LS et à la hampe, dont l'allométrie est positive. Pour les couronne ane cuspide principale < cuspide laterale Fic. 1. Schéma des mensurations d'une dent standard. 750 SONIA MULLER ET CLAUDE WEBER deux populations d'Ancistrus, ces deux caractères ont une allométrie négative par rapport à _la LS. La couronne croît toutefois proportionnellement plus que la hampe pour la population à longue couronne dentaire (Ancistrus cf. bufonius (Valenciennes)), carac- téristique aussi observée chez d'autres espèces d'Ancistrus à longue couronne. RÉSULTATS TAXONOMIQUES Chez Hypostomus, les dents présentent d'importantes variations en nombre, des formes diverses ainsi qu'une relation entre le nombre et la forme. Ces éléments permettent la caractérisation d'espèces et, en corrélation avec d'autres caractères, la formation de groupes d'espèces: La forme particulière de la couronne dentaire (cuspide principale plus longue et cuspide latérale réduite et parfois absente; fig. 2d) est le principal caractère de la diagnose différentielle de A. piratatu Weber avec H. boulengeri (Eigenmann & Kennedy). Un groupe d'espèces, que nous nommons regani, se distingue par un nombre élevé de dents (de 37 à 125) à couronne longue, associé à une mandibule large. Il se répartit sur une zone s'étendant du Rio Parnaiba au bassin du Parana, incluant des affluents du Rio Sao Francisco, ainsi que le Rio Pilcomayo et le Rio Bermejo supérieur. Entre autres espèces, on y trouve H. auroguttatus (fig. 2e). Un autre groupe que nous appelons plecostomus est caractérisé par une mandibule moyenne, garnie de dents à couronne courte (fig. 2f: Hypostomus aff. derbyi (Haseman)). Il rassemble un grand nombre d'espèces largement distribuées dans l'aire de répartition du genre, parmi lesquelles se trouvent H. boulengeri, H. commersonii Valenciennes et H. plecostomus (Linné). H. microstomus Weber et H. roseopunctatus possèdent des dents massives en très faible nombre (de 6 à 16) associées à une petite mâchoire (pl. 1a; WEBER 1987: 282, fig. 7). Ces deux espèces, qui montrent beaucoup d'affinités, occupent les bassins voisins du Parana et de l'Uruguay. Le même type de denture se retrouve chez deux autres espèces: H. margaritifer (Regan) occupant le bassin du Parana, mais avec des dents en plus grand nombre (de 16 à 27), et H. carinatus (Steindachner) provenant de l'Ama- zone, avec seulement 12 à 16 dents comptés sur l'holotype qui diffère toutefois complète- ment des trois autres par la livrée. Chez Ancistrus, l'examen des dents donne également des résultats intéressant espèces et groupes d'espèces. Par leur nombre très faible (de 23 à 48 pour des individus de 17 à 130 mm), elles sont caractéristiques d'une espèce non-décrite (A) du bassin amazonien . Par leurs formes uniques dans le genre, elles distinguent deux autres nouvelles espèces également amazoniennes. La première (B) possède des dents à cuspides homo- logues (fig. 2g) comme chez certains Peckoltia (fig. 2a) et Delturus. La seconde (C) se caractérise par une dent à cuspide latérale absente ou minuscule (fig. 2h), comme c'est parfois le cas chez H. piratatu. La longueur de la couronne est particulièrement variable dans ce genre; toute espèce confondue, son rapport dans la longueur de la hampe va de 0, 8 à 5, 9. Ces extrêmes sont Justement observés chez les nouvelles espèces B et C. Certaines espèces se rapprochent par les caractères dentaires. Les couronnes les plus longues (moins de 1,4 dans la hauteur de la hampe) et un nombre très élevé de dents se rencontrent chez des espèces qui présentent aussi d'autres caractères en commun; nous les regroupons sous le nom groupe bufonius. La dent à très longue couronne possède une hampe de taille généralisée (2,1 à 3,2, moy. 2,5 dans la LSx1/56) et s'inscrit grossièrement dans un carré (fig. 2i: A. bufonius; pl. 1c: A. megalostomus). Ce type de denture se DENTS DE HYPOSTOMUS ET D'ANCISTRUS 751 ay È, f e de % Ir FIG. 2. 8) j Vue laterale de la dent et vue dorsale de la couronne dentaire. a: Peckoltia oligospila Günther (LS: 88; 20x); b: Chaetostoma lineopunctatum Eigenmann & Allen (LS: 64; 20x); c: Cochliodon cochliodon (Kner) (LS: 224; 10x); d: H. piratatu Weber (LS: 244; 10x); e: Hypostomus auroguttatus Natterer & Heckel (LS: 120; 10x); f: H. aff. derbyi (Haseman)(LS: 145; 10x); g: Ancistrus sp. B (LS: 110; 20x); h: Ancistrus sp. C (LS: 110; 20x); i: A. bufonius (Valenciennes) (LS: 103; 20x); j: A. hoplogenys (Günther)(LS: 100; 20x); k: A. lithurgicus Eigenmann (LS: 64; 20x). LS: longueur standard en mm; x: grossissement de la vue latérale; couronne agrandie 2 fois par rapport à la dent sauf pour C. cochliodon. 752 SONIA MULLER ET CLAUDE WEBER retrouve dans d'autres genres, notamment Chaetostoma Tschudi, 1845 (fig. 2b). La distribution de ce groupe semble limitée à l'Est des Andes péruviennes et boliviennes, dans les affluents de l'Ucayali et du Madre de Dios. Le type d'Ancistrus caucanus Fowler, seul exemplaire que nous ayons pu examiner pour cette espèce, possède également des dents à très longues couronnes (1,1 dans la hampe) mais en nombre moins élevé (environ 50 pour une LS de 52 mm), comparable à celui des nombreuses espèces à denture moyenne que l'on trouve largement distribuées dans l'aire de répartition du genre. A l'inverse du groupe bufonius, on observe des dents peu nombreuses à couronne courte (plus de 2 dans la hauteur de la hampe) et hampe habituelle (1,8 à 3,2, moy. 2,3 dans la LS x 1/56) chez les espèces du groupe hoplogenys, distribuées dans le bassin de l'Amazone ainsi que dans celui de l'Orénoque et le plateau guyanais (fig. 2j et pl. 1b: A. hoplogenys (Günther)). À. lithurgicus Eigenmann (fig. 2k) de l'Essequibo et À. macro- phthalmus (Pellegrin) de l'Orénoque possèdent aussi une couronne courte, mais s'en distinguent par leurs dents fines et longues (LS/hampe x 1/56: 1,2 à 2, moy. 1, 5). DISCUSSION La denture des populations à dent standard étudiées montre une variabilité ontogénique non négligeable du nombre de dents, qui s'ajoute à la variabilité individuelle souvent importante. La comparaison du nombre moyen de dents par classes révèle chez Hypostomus et Ancistrus une grande variation selon les espèces. Certaines populations (chez Ancistrus groupe bufonius) ont un nombre de dents significativement différent sur le dentaire et le prémaxillaire. Il pourrait s'agir de carac- téristiques spécifiques, certaines ayant plus de dents sur le prémaxillaire, d'autres sur le dentaire. Nous attendons l'examen d'un plus grand nombre de populations, comprenant des individus de toute taille, pour le préciser. La variabilité morphologique est faible durant l'ontogénie. Nous notons toutefois des allométries caractéristiques de la longueur de la hampe de la dent et de sa couronne. En comparaison, on observe parmi d'autres genres des modifications importantes. Chez Cochliodon, les dents des très jeunes individus sont comparables par la forme avec celles habituellement rencontrées chez les autres Hypostominae. Elles ne sont remplacées par les dents à grosse couronne sans cuspide latérale (fig. 2c) qu'au delà de 60 mm. de LS (LILYESTROM 1984: 44). Dans plusieurs populations de Liposarcus pardalis (Castelnau), nous avons observé que les dents standard sont remplacées par des dents unicuspidées à couronne fortement allongée dès 245 mm de LS. Chez Hypostomus et Ancistrus, la longueur de la couronne est très variable à l'intérieur des genres, ce qui n'est pas le cas pour d'autres tels Pterygoplichthys Gill, 1858, Liposarcus Günther, 1864 et Glyptoperichthys Weber, 1991 récemment révisés. Les mesures prises sur les dents permettent de caractériser des groupes d'espèces et, souvent associées à d'autres éléments morphologiques ou morphométriques, précisent certaines affinités. Plusieurs caractères sont généralement corrélés: la taille des mâchoires, le nombre de dents et la longueur de la couronne. Dans une grande mâchoire, on trouve généralement un nombre élevé de dents à longue couronne et inversément. Nous observons aussi des exceptions à l'image de deux espèces colombiennes, H. varimaculatus (Fowler) du bassin du Caqueta et A. caucanus du Cauca, qui présentent une couronne longue avec un nombre de dents et une mâchoire tout à fait ordinaires. DENTS DE HYPOSTOMUS ET D'ANCISTRUS 753 PLANCHE | Bouche et couronne dentaire. a: Hypostomus roseopunctatus Reis, Weber & Malabarba (paratype, MAPA 23125; LS: 190); b: Ancistrus hoplogenys (Günther) (syntype, BMNH 1849.11.8:89; LS: 86,5) (photo bouche: G. Dajoz); c: Ancistrus megalostomus Pearson (syntype, CAS 64614; LS: 82,9 mm) (photo bouche: C. Ratton). 754 SONIA MULLER ET CLAUDE WEBER Enfin, l'examen systématique des dentures permet la discrimination de plusieurs espèces de Hypostomus et d'Ancistrus par leurs caractères uniques au sein des genres. Dans le cas de H. piratatu, les caractères dentaires sont les seuls discriminants et une étude électrophorétique comparative de certaines populations sympatriques devrait, le cas échéant, confirmer leur validité. REMERCIEMENTS De nombreux instituts ont mis à notre disposition aide et matériel; qu'ils soient tous remerciés ici. Nous sommes reconnaissants à Volker Mahnert de ses judicieux conseils et encouragements. Nous remercions également Béatrice Naef de sa précieuse aide technique lors de la préparation de cette étude, ainsi que Florence Marteau et Gilles Roth, dessinateurs au Muséum. BIBLIOGRAPHIE GOSLINE, W. A. 1947. Contribution to the classification of the loricariid catfishes. Arg. Mus.nac. Rio de Janeiro, 33: 79-138. i ISBRÜCKER, I.J.H. 1980. Classification and catalogue of the mailed Loricariidae (Pisces, Siluri- formes). Versl. Techn. Geg., Inst. Taxon. Zool. (Zool..Mus.), Univ. Amsterdam, 22: 1-181. LILYESTROM, C. G. 1984. Consideraciones sobre la taxonomia de las especies del genero Cochliodon Heckel en Venezuela (Pisces, Loricariidae). Revta UNELLEZ de Ciencias y Technotosta ser. Prod. Agricola, Guanare, Venezuela. 2 (2): 41-53. MULLER, S. 1990. Etude méristique et morphométrique d’Ancistrus piriformis Muller et Ancistrus pirareta Muller (Pisces, Siluriformes, Loricariidae). Revue suisse Zool. 97: 153-168. SCHAEFER, S. A. 1986. Historical biology of the loricariid catfishes: phylogenetic and fonctional mor- phology. Thèse, Université de Chicago: i-x, 1-198, 46 fig. WEBER, C. 1987. Hypostomus microstomus sp. nov. et autres poissons-chats cuirassés du Rio Parana (Pisces, Siluriformes, Loricariidae). Arch. Sc. Genève 40: 273-284. | Revue suisse Zool. | Tome 99 Fasc. 4 | p. 755-770 Geneve, décembre 1992 | | Evolutionary aspects of development, life style, and reproductive mode in incirrate octopods (Mollusca, Cephalopoda)* by Sigurd v. BOLETZKY ** With 7 figures ABSTRACT The incirrate octopods are defined as a monophyletic group by a number of characters unknown in the cirrate octopods or in other cephalopods. The biologically most significant incirrate feature is the incubation of eggs by the female. The morphological and behavioural characters underlying this special mode are analysed with regard to developmental processes and their modification related to life style evolution. 1. INTRODUCTION Common inshore species of the genus Octopus Lamarck, 1798 are popular models in comparative invertebrate biology (YOUNG, 1971; WELLS, 1978); in such a context they can be viewed as ‘typical cephalopods’. When attention is focussed on diversity within the class Cephalopoda, however, the common octopuses turn out to be rather ‘special’ cephalopods. This observation raises the question of respective systematic positions: what is special about which group of octopods ? 1.1. SYSTEMATICS Within the cephalopod subclass Coleoidea Bather, 1888 (=Endocochleata, Dibran- chiata), the order Octopoda Leach, 1818 is a well defined taxon. It contains two suborders, the Cirrata Grimpe, 1916 (better known as ‘finned octopods') and the Incirrata * Travail présenté a Zoologia 92. ** C.N.R.S., URA 117, Laboratoire Arago, F-66650 Banyuls-sur-Mer (France) 756 SIGURD V. BOLETZKY Grimpe, 1916, to which the common octopuses belong. YounG (1989) proposed a new -classification, with an infraclass Octobrachia containing two orders, the Cirroctopoda (=Cirrata) and the Octopoda (=Incirrata). For practical reasons, the old classification is used here, especially to avoid confusion between Octopoda Leach, 1818 and Octopoda Young, 1989 (cf. quotation from YOUNG, loc. cit. in Discussion). 1.2. PHYLOGENETIC BACKGROUND To appreciate the evolutionary significance of incirrate characters with regard to functional adaptation and phyletic conservation of patterns, the incirrates must of course be viewed in comparison to their supposedly closest relatives, the deap-sea cirrate octopods. The respective positions of the fossil Proteroctopus of the Middle Jurassic and Palaeoctopus of the Late Cretaceous are not discussed here, because the available morphological data are insufficient (see ENGESER, 1988 for a review). The characters of living octopods can nevertheless be scrutinized in the greater framework of coleoid phylogeny, starting out from the Vampyromorpha (cf. YOUNG, 1989). This taxon was formerly included in the Cirrata because of the great similarity in arm morphology. When the so-called retractile filaments of Vampyroteuthis were recognized as an additional pair of rudimentary arms, the Vampyromorpha were made an order of its own (PICKFORD, 1939, 1949). The evolutionary history of endocochleate cephalopods can be traced back to the lower Devonian (BANDEL et al. , 1983; BANDEL & BOLETZKY, 1988). For the present purpose it is sufficient to consider the extant coleoids in relation to those fossil coleoids that had ten arms of similar length and structure, a crucial feature that NAEF (1923) used for the definition of a hypothetical 'Protodibranchus'. This arm pattern disappeared with the extinction of the belemnites at the end of the Cretaceous. The 'belemnoid' arm crown morphology is important for our understanding of coleoid phylogeny because it is the only one from which one can derive the respective patterns of 1) the decapodan cuttlefishes and squids, which have the fourth arm pair modified as tentacles, and 2) a group provisionally named 'Vampyropoda' (=Octopodi- formes Berthold & Engeser, 1987; name preoccupied) to include the Vampyromorpha Pickford, 1939 with their modified second arm pair and the Octopoda Leach, 1818 (lack of probably the second arm pair). The important point here is that these two modifications must have occurred independently, 1. e. at two different speciation events, because a simultaneous occurrence of mutually exclusive modifications is inconceivable (Fig. 1a). Prerequisite to this phylogenetic deduction is the existence of unambiguous identities and positional relationships of brachial appendages allowing one to recognize modifications for a given pair of arms (see BOLETZKY, 1992 for a review). These criteria are concerned with the ‘integration level’ of distinct appendages; they do not involve the specialisations at the next lower level which comprises the armature of the arms and tentacles (suckers, hooks, cirri). Given the situation described above, any discussion of phylogenetic systematics of the Coleoidea has to cope with (only) two possibilities: either the decapods and the belemnoids are sister groups for which the 'Vampyropoda' are the outgroup, or the 'Vampyropoda' and the belemnoids are sister groups, the decapods being the outgroup (Fig. 1b). The question of which one of these arrangements is true lies outside the scope of this paper. EVOLUTIONARY ASPECTS OF OCTOPODS 757 2. INCIRRATE CHARACTERS In addition to the absence of brachial cirri warranting the name of the taxon, and hectocotylization of one of the ventro-lateral arms, the Incirrata show a series of characters unknown in the Cirrata or in any other coleoid cephalopod (cf. NAEF, 1923, 1928); the most conspicuous of these are: 1. absence of muscular fins ('finlessness') 2. presence of Kölliker's organs in hatchling skin 3. partial modification of egg case (chorion stalk) 4. reduced encapsulation of egg case (chorion stalk only) 5. egg-care behaviour (or ovovivipary) in the female. Although none of these characters is known in any other cephalopod, the question remains whether they are uniquely derived (apomorphic) characters of the Incirrata, or whether they (or some of them) could be autapomorphic characters of the octopodan ancestor that were subsequently eliminated in the Cirrata (so they would be plesiomorphic at the level of the Incirrata). So far nothing seems to indicate that the absence of the above characters in the Cirrata could be the result of such an elimination. An equally important question is how closely related these characters are to one another. Here one has to consider several variables; a behavioural one, namely the post- hatching life style as compared to the adult mode of life, and two morphometrical variables, egg size and body proportions. The behavioural features necessarily lead to the question of how to define the ancestral life style from which the modes of living incirrates must be derived. Before this question can be addressed, the incirrate characters listed above have to be scrutinized in some detail. Ad I. The absence of muscular fins in the Incirrata is not total if embryonic development is taken into account. Fin rudiments do appear during organogenesis, in close positional relation to the shell sac as is typical for the coleoids (Fig. 2). The incirrate shell sac is very small from the beginning; it becomes drawn out laterally during early development (APPELOF, 1898). The resulting transverse tube finally splits into two independant tubes which become embedded in the muscular tissue of the mantle (and can finally disappear, as in Argonauta). During this shell sac differentiation the fin rudiments gradually smooth out (NAEF, 1928). What might be taken as rudimentary fins in preserved hatchlings viewed in the scanning electron microscope are fixation artefacts due to shrinkage (Fig. 3); although the position of these ostensible 'buds' corresponds to the location of the shell stylets, no trace of fin tissue is histologically detectable. The character ‘absence of muscular fins' could be rephrased to emphasize the rudimentation of the whole fin-shell complex, assuming homology of fin rudiments in Cirrata and Incirrata (BOLETZKY, 1982 a). When viewed against the pattern of an unpaired transverse shell sac and associated fins (as present in cirrates), the bipartite shell sac of incirrates suggests a correlation between two apomorphic characters, namely subdivision of the shell sac and truncation of fin differentiation, i. e. 'finlessness'. Ad 2. The majority of incirrate hatchlings have special tegumentary organs; they were observed in Argonauta embryos by KOLLIKER (1844) and fully described in other incirrates by QUERNER (1927) and more recent authors (e. g. FIORONI, 1962) (Fig. 3). The absence of Kölliker's organs in Octopus briareus (Fig. 7) and O. maya (BOLETZKY, 1973) 758 SIGURD V. BOLETZKY can be viewed as the result of total suppression in specifically modified integument ‘morphogeneses. So far no trace of these organs has been found in embryos of cirrate octopods (BOLETZKY, 1982 b). Ad 3. During vitellogenesis, cephalopod oocytes may take on a markedly elongate form, but it is only in the Incirrata that this elongation leads to the differentiation of a distinct chorion stalk (Figs 4, 5). The elongate form of the incirrate chorion is also meaningful with regard to embryonic movements that occur in all incirrates so far studied, with the exception of Argonauta ; this feature could be a side effect of the primary modification of late oogenetic processes (BOLETZKY & FIORONI, 1990) (Fig. 4). Ad 4. In the incirrates the 'cement' secreted by the oviducal glands (FROESCH & MARTHY, 1975) ‘encapsulates’ only part of the chorion stalk (Fig. 5). This ‘partial egg encapsulation’ contrasts with the complete encapsulation of the cirrate eggs; it thus appears as an apomorphic character linked with the formation of a chorion stalk (character 3). Ad 5. Although visual stimulation by other egg masses can induce spawning in many coleoid cephalopods, post-spawning egg care exists only in the incirrate octopods (BOLETZKY, 1986). This unique protective behaviour must be related to the absence of protective encapsulation (character 4). In other words, characters 3, 4 and 5 are clearly connected, forming an apomorphic complex of features named the "incubating mode" of reproduction. In the pelagic genus Ocythoe, incubation exists in the 'pure' form of ovovivipary (NAEF, 1923). For two families, the Alloposidae and the Amphitretidae, egg- care is not yet documented (Hochberg, pers. comm.). Before approaching the question of possible relations between the above ‘incubation complex’ and the characters 1 and 2 (see C.), it is necessary A. to review life styles and morphometrics at different stages of the incirrate life cycle, and B. to see whether characters | and 2 are correlated. A. LIFE STYLES AND MORPHOMETRICS A.1. Life styles a) ADULT LIFE STYLES IN THE INCIRRATA Eight incirrate families are recognized if the Idioctopodidae Taki, 1962 are included in the Amphitretidae Hoyle, 1886 (cf. HOCHBERG ef al., 1992; disregard the erroneous statement in BOLETZKY, 1978-79, p. 107). Only the Octopodidae Orbigny, 1840 are clearly benthic at the adult stage. R. E. YOUNG (pers. comm. to HOCHBERG et al., 1992) suggests that adult Alloposus mollis Verrill, 1880 (of the monotypic family Alloposidae Verrill, 1882) may also be benthic. The remaining six families are pelagic; these are the Argonautidae Tryon, 1879, the Tremoctopodidae Tryon, 1879, the Ocythoidae Gray, 1849 (these three families were grouped with the Alloposidae in a tribe called Argonautida by Rosson, 1932), the Vitreledonellidae Robson, 1930, and the two "ctenoglossan" families Bolitaenidae Chun, 1911 and Amphitretidae Hoyle, 1886. EVOLUTIONARY ASPECTS OF OCTOPODS 759 In all these families, reproduction takes place in midwater, and the eggs apparently remain with the female until the young hatch out. The most elaborate mode of egg care is achieved by the female Argonauta which produces a calcified "brood shell". However, along with housing the egg mass, this pseudoconch serves as a floater; the animal keeps an air bubble in the apex and thus obtains neutral buoyancy (BOLETZKY, 1983). Moreover the brood shell supports the brachial membrane in a food detective function (YOUNG, 1960). A much simpler form of egg carrier is produced by Tremoctopus ; as in Argonauta, the calcified structures are secreted by the dorsal arms (NAEF, 1923). In Eledonella pygmaea (family Bolitaenidae), the whole arm crown of the female forms a brood chamber (YOUNG, 1972). Male sexual behaviour in pelagic incirrates can be only partly inferred from the structure of the copulatory arm (hectocotylus). In the "Argonautida" sensu ROBSON (1932), the morphologically and morphometrically extreme differentiation of the hectocotylus seems correlated with the capacity to autotomize. In the benthic Octopodidae, females always spawn on the bottom. Generally single eggs or egg strings are cemented to the wall or ceiling of the den occupied by the female. In a few octopodid species, the females carry egg masses loose and thus can move about while brooding the eggs (see HOCHBERG et al., 1992). b) POST-HATCHING LIFE STYLES IN THE INCIRRATA As far as is known (cf. HOCHBERG et al., 1992), the juveniles of pelagic families live in midwater (including the Alloposidae; see above). In the benthic Octopodidae, the representatives of the subfamily Bathypolypodinae Robson, 1931 probably stay on the bottom throughout their life (cf. Bathypolypus arcticus, as observed by O'Dor & MACALASTER, 1983). If the new arrangement proposed by Voss (1988) is accepted, the new subfamilies Graneledoninae and Pareledoninae are entirely holobenthic (Hochberg., pers. comm.). The subfamilies Octopodinae Grimpe, 1921 and Eledoninae Gray, 1849 include numerous species characterized by the same 'holobenthic' mode as Bathypolypus, while others have a planktonic post-hatching phase; the mode of life of the latter species can be named 'merobenthic'. Their young animals are actively foraging carnivores that feed on both living planktonic prey and drifting food items (facultative scavenging). They generally remain in midwater until they have grown larger. In some species, newly- hatched animals show temporary settling between phases of active swimming (BOLETZKY, 1977). A.2. Morphometrics a) BODY PROPORTIONS OF INCIRRATE HATCHLINGS The hatchlings of pelagic incirrates are characterized by short arms (generally less than 1/3 of total length) with few suckers. This feature again appears in the newly hatched animals of merobenthic octopodids (Fig. 6), although in the larger hatchlings each of these relatively short arms may carry up to 15 suckers. In contrast, the hatchlings of holobenthic octopodids have arms at least as long as the rest of the body, with more than 20 suckers per arm (Fig. 7). 760 SIGURD V. BOLETZKY The body proportions of planktonic hatchlings gradually change due to the positive . allometric growth of the arms. In the young merobenthic octopodids, body proportions thus become similar to those of the 'crawl-away' hatchlings of holobenthic species. In Octopus vulgaris (and probably in the majority of merobenthic octopodids) the young animals, having reached these body proportions, gradually change from continuous swimming to the adult-type bottom life, which includes only occasional excursions into the water column (ITAMI et al., 1963). This drastic change contrasts with the condition of pelagic incirrates, in which juvenile arm growth is not accompanied by a thorough modification of life style. b) INCIRRATE EGG SIZES AND HATCHLING FEATURES Within the Incirrata, the size of a single ovum varies from 0.8 mm in Argonauta spp. to 35 mm in Graneledone sp. (HOCHBERG et al., 1992). Among the pelagic incirrates, the variation spans only from 0.8 to about 4 mm, however. In contrast, egg sizes vary from about 1.5 mm to 35 mm in the Octopodidae. RoBSON (1932, p. 25) once expressed egg lengths as percentages of adult mantle- lengths, but his erroneous egg index for Eledone cirrosa prevented him from realizing the great difference between Eledone moschata (egg length ca 15% of adult mantle length) and E. cirrhosa (ca 5%; cf. Fig. 5); adults of the two species are similar in size. This index becomes particularly interesting when absolute egg sizes are similar among species with very different adult sizes. There are several octopodid species that produce eggs measuring about 5 to 8 mm; in the larger species the embryos become planktonic hatchlings with short arms and less than 15 suckers per arm, whereas in the smaller species embryos of the same size end up as benthic hatchlings with long arms and more than 20 suckers per arm (cf. A.2.a). In fact, the ostensibly ‘intermediate’ egg sizes fall under the same categories as the ‘very large’ and the ‘Very small’ eggs and are distinguishable by the relative egg size, or egg index (BOLETZKY, 1974, 1977). An index smaller than 10% is indicative of the merobenthic mode, whereas an index greater than 10% reflects holobenthic conditions (occasional behavioural peculiarities in newly hatched animals notwithstanding). An exception is Octopus fitchi, a very small species in which the eggs (ca 5 mm) are large relative to the adult mantle-length (ca 30 mm: egg index ca 16); the arms are stout and almost as long as the rest of the body like in hatchlings of holobenthic species, but each arm carries less than 17 suckers, and the post-hatching life style is clearly planktonic (HOCHBERG et al., 1992). B. CORRELATION OF CHARACTERS 1 AND 2 a) FINLESSNESS AND THE BENTHIC LIFE STYLE That the incirrates lack fins was long interpreted as a result of adaptation to benthic life (NAEF, 1923). The absence of fins in the pelagic incirrates was then naturally viewed as a condition conserved from a finless benthic ancestor, assuming that fins lost in that ancestor were not "reinvented" in its pelagic descendants. However, a causal relationship between the benthic life style of an octopus and the absence of muscular fins has never been shown to exist. In fact, cuttlefish and sepiolid squids demonstrate that fins may be indispensable even on the bottom as can be seen when these animals bury in soft EVOLUTIONARY ASPECTS OF OCTOPODS 761 substrates: at the outset of burying they can remain on the spot only because the fin movements counteract the propulsive effect of the funnel jet by which substrate particles are blown up. Fins can be expected to have disappeared from the morphogenetic program only if they were incompatible with the functional morphology corresponding to a given life style. There is no indication of such incompatibility in relation to the benthic life style as it appears in the Octopodidae. b) FINLESSNESS AND THE PELAGIC LIFE STYLE Among the pelagic incirrates, one condition may appear incompatible with the presence of fins; this is the presence of a brood shell in female Argonauta. However, one cannot reasonably assume that this highly elaborate female structure represents an ancestral incirrate condition. A feature really incompatible with the presence of fins could be character 2 of our list, i. e. presence of Kölliker's organs in the juvenile skin. The way these organs function, especially when they evaginate and spread the setal tufts, suggests that they would interfere with fin activity if fins still existed along with them. Provided that tuft spreading occurs under higher nervous control and has a parachute effect in midwater when the animal remains motionless (BOLETZKY, 1978-79), the establishment of Kölliker's organs can be considered in relation to the pelagic life style. At the level of the incirrate ancestor, this of course holds only for small body sizes at which the tufts can generate enough drag to slow sinking. In other words, the formation of Kölliker's organs is likely to reflect an originally juvenile adaptation to pelagic life. Finlessness thus appears as the obligatory counterpart of the juvenile 'setaceousness', in other words characters 1 and 2 of our list are probably correlated. C. CORRELATION OF CHARACTER COMPLEXES Nothing so far mentioned provides an indication of any relationship between the complex of characters 1 and 2 and the complex of characters 3, 4 and 5 of our list. Such a link appears only when hatching is considered. In the merobenthic and pelagic incirrates, Kölliker's organs play an essential, though passive, role during hatching (BOLETZKY, 1978- 79). The setal cores of these organs provide a "shingle" structure to the hatchling skin and thus prevent its slipping back into the chorion when the animal makes the stretching movements necessary to work itself through the hatch opening (which is produced by enzymes released from the hatching gland). Notwithstanding exceptions like Scaeurgus unicirrhus where short arms are used during hatching (BOLETZKY, 1984), the role of the setal cores seems essential in the small young having very short arms that remain passive during hatching (in contrast to the holobenthic octopodids where the crawl-away hatchlings always use their long arms to work themselves out of the chorion). However, the shingle structure of the skin is effective only if the hatch opening has a solid edge. This condition is fulfilled by the relatively thick, stiff chorion of incirrate eggs. Another question is whether this particular function during hatching is the primitive function of Kölliker's organs. The complex structure of these organs, especially the elaborate musculature that permits repeated spreading and retraction of the tufts during post-hatching life, and the fact that the organs cover also the arms where they are not needed for hatching, suggest that their function during hatching is a secondary adaptation 762 SIGURD V. BOLETZKY superimposed on a primary function related to the post-hatching mode of life. A ‘ prerequisite of this secondary adaptation must have been the modification of the encapsulation process, which changed from the complete encapsulation seen in the cirrates to partial encapsulation of the egg. This evolutionary transformation is conceivable only in combination with a special timing of egg release allowing the follicular chorion attachment to be drawn out into a distinct chorion stalk. Internal fertilization, a likely prerequisite, was already achieved in the octopodan ancestor, as demonstrated by the Cirrata (cf. VILLANUEVA, 1992); Vampyroteuthis appears to have external fertilization similar to the decapods (PICKFORD, 1949). 3. DISCUSSION Starting from the feature 'finlessness' through the related 'setaceousness' and its implications in both the post-hatching life style and the hatching mechanism, our survey arrives at the question of the evolutionary origin of the incirrate mode of reproduction. Considering the constraints placed on egg shaping and timing of egg release, attention is naturally drawn to ovovivipary, the special incubation mode of Ocythoe. Could this mode represent the primitive condition from which the post-spawning egg care was derived? Is the inverse process more likely? Or is the 'intermediary' condition of Argonauta, where eggs are released only after the first cleavage stages (NAEF, 1928), closer to the primitive condition from which the other two were derived? These questions inevitably raise the problem of the ancestral life style under which incubation became established. Given that seven of the eight living incirrate families are pelagic, it appears likely that the 'most generalized' life style represents the ancestral condition. But this hypothesis remains very vulnerable as long as it is only based on the respective numbers of extant families representing the pelagic or the benthic life style. An indication supporting the above hypothesis could be the existence of a pelagic juvenile phase in many octopodids (merobenthic species). This juvenile phase is likely to be a conserved feature that stems from a pelagic ancestor. Advantages of this conservation could have been greater availability of small prey animals in midwater (BOLETZKY, 1977, 1981) and low selective pressure in a relatively 'simple' open water environment where the limited behavioural repertoirs of very small juveniles suffice (BOLETZKY, 1987). Thus the planktonic juvenile phase would have been eliminated in the holobenthic species. It is indeed easier to imagine an evolutionary parallelism resulting from convergent suppressions of the pelagic phase than the inverse, namely independently emerging pelagic juvenile phases. With the latter hypothesis, it would be particularly difficult to explain why the planktonic hatchlings tend to be so similar, and why they resemble so closely those of the pelagic incirrates. One may of course argue that perhaps the pre-octopodid ancestor was already characterized by juvenile life style switching; this would have allowed the holopelagic life cycle of the majority of incirrates to emerge through a paedomorphic 'abbreviation' by suppression of the ancestral adult mode. This could be the hypothesis underlying the comment of YOUNG (1989, p. 235-236) on a cryptic incirrate character related to the receptor system of the statocyst: "The division of the crista into nine sections is a unique apomorphic feature of the order Octopoda; it is not present in Cirroctopoda, which presumably never possessed it. The feature was possibly developed to provide for the wide EVOLUTIONARY ASPECTS OF OCTOPODS 763 range of frequency of turning during walking and swimming. It is surprising to find that the crista is still so divided in all the pelagic octopods examined”. The question can of course be reversed: is it surprising to find that the crista is so divided in all the pelagic incirrates ? Not if one assumes that the benthic octopodids are derived from a pelagic ancestor. Efforts should now be concentrated on the identification of sister group relationships within the incirrates. Do the octopodids have an immediate common ancestor with one of the other incirrate subgroups, or is the octopodid lineage derived from a basic dichotomy so that the Octopodidae were the sister group of all other incirrates ? Two variants of a peculiar behavioural feature in some pelagic incirrates deserve special attention. One is the use of the own brood shell as a buoyant device by female Argonauta (cf. 2.A.1.a), the other is the use that male Ocythoe make of empty tests of doliolids and salps as drifting ‘homes’ (NAEF, 1923). The great similarity of these behavioural patterns suggests that the typical arm posture of a benthic octopodid sitting in its den is homologous to the respective attitudes of female Argonauta and male Ocythoe in their pelagic 'homes'. It is conceivable that the behaviour pattern corresponding to such a ‘rafting’ mode of life provided the initial condition for the establishment of an adult benthic mode. The inverse process seems conceivable only if the supposed benthic ancestor already had a planktonic juvenile phase. In conclusion, the most generalized life style in incirrates is characterized by active swimming and drifting; ontogenetically this is an elaboration of a pelagic juvenile phase. This phase has probably been eliminated in many species of the benthic family Octopodidae. To derive the wide variety of incirrate modes from a holobenthic ancestor, through repeated ‘invention of the pelagic juvenile phase, seems rather problematic. Egg incubation in incirrates may thus be surmised to have emerged in the adaptive context of a bentho-pelagic or pelagic life style. SUMMARY This paper reviews the common features of the octopodan subgroup Incirrata from an evolutionary point of view, raising questions of functional adaptation and co-adaptation of morphological and behavioural characters. The most conspicuous difference between incirrate octopods and other cephalopods is the ‘incubating mode’ of reproduction (post- spawning egg care or ovovivipary). As the cirrate octopods, which are the likely sister group of the incirrates, show no signs of incubation, the evolutionary origin of this novel mode of incirrate reproduction can only be ‘reconstructed' through careful weighing of the relative importance of characters that are more or less closely related to reproduction. Developmental features provide particularly interesting cues (e. g. truncation of fin development, formation of special tegumentary organs) allowing one to approach the question of the ancestral life style from which the pelagic and benthic modes of extant incirrates must be derived. ACKNOWLEDGMENTS I sincerely thank Dr. F. G. Hochberg (Santa Barbara Museum) for his critical reading of the manuscript and for his stimulating suggestions. 764 SIGURD V. BOLETZKY Decapoda Vampyropoda (Vampyromorpha + Octopoda) Belemnoidea ext. Decap. Belemn. Vampyrop. Decap. Belemn. Vampyrop. Fic. 1 a. Phylogenetic relationships between the living coleoid groups Decapoda (five arm pairs, fourth pair modified) and Vampyropoda (five arm pairs, second pair modified) via the extinct Belemnoidea (five arm pairs without distinct modifications). The trichotomy is shown unresolved in terms of sister group relationships. b. The two conceivable sister group relationships (the theoretical third one, supposing Decapoda + Vampyropoda with Belemnoidea as the outgroup, is inconceivable for morphological reasons, as explained in the text). FIG. 2 Lateral view of a live embryo of Octopus vulgaris in its chorion, the large outer yolk sac (at right) and the chorion stalk (at left) are not shown. At this advanced organogenetic stage (stage XII of Naef, 1928), one can recognize the organ complexes surrounding the dark yolk mass: a voluminous buccal mass (b), the stubby arms (a), the funnel tube (f), and the cap-like mantle with the fin rudiments (arrow) overlying the rudimentary shell sac (arrow head). Fic. 3 Preserved hatchling of Octopus vulgaris in caudo-dorsal view (SEM), with the posterior mantle apex at the lower left. The elevations marked by arrowheads correspond roughly to the position of fin rudiments in decapod embryos, but they are fixation artefacts due to shrinkage (see text). The small arrows point at some of the small elevations producing the "shingle" structure of the hatchling skin; the tips of the setal cores of Kölliker's organs have broken through the skin surface only in the nuchal region (upper right). EVOLUTIONARY ASPECTS OF OCTOPODS 765 766 SIGURD V. BOLETZKY FIG. 4 Live embryo of Argonauta argo, With pigmented eyes (e), tightly enclosed in its chorion, with the chorion stalk (arrow) at the upper right. The outer yolk sac lies at the stalk side, the mantle of the embryo at the side of the micropyle (arrow head). At organogenetic stages, incirrate embryos normally show inverse orientation inside the chorion (see text). E EVOLUTIONARY ASPECTS OF OCTOPODS AGH Fic. 5 Egg strings from the octopodids Eledone cirrhosa (left and middle) and Octopus vulgaris (right). The arrow points at the embedding site of a long chorion stalk in the 'cement' secreted by the oviducal gland; the egg string in the middle shows a 'tidier' arrangement with deep chorion stalk embedding (arrow head) and formation of a central cement axis, similar to that of the Octopus vulgaris egg string (central axis not visible). Despite the great difference in egg size, hatchlings of both species have short arms (cf. Fig. 6) and live for some time in the plankton (see text). 768 SIGURD V. BOLETZKY EVOLUTIONARY ASPECTS OF OCTOPODS 769 REFERENCES APPELLÖF, A. 1898. Über das Vorkommen innerer Schalen bei den achtarmigen Cephalopoden (Octopoden). Bergens Mus. Aarb. 12: 1-15. BANDEL, K. & S. v. BOLETZKY. 1988. Features of Development and Functional Morphology required in the Reconstruction of Early Coleoid Cephalopods. Zn: Cephalopods - Present and Past (Wiedmann, J. & J. Kullmann eds.). Schweizerbart'sche Verlagsbuchhandlung, Stuttgart. pp. 229-246. BANDEL, K., J. REITNER & W. STÜRMER. 1983. Coleoids from the Lower Devonian Black Slate ("Hunsriick-Schiefer") of the Hunsrück (West Germany). N. Jb. Geol. Paläont. Abh. 165: 97-417. BERTHOLD, T. & T. ENGESER. 1987. Phylogenetic analysis and systematization of the Cephalopoda (Mollusca). Verh. naturw. Verein Hamburg , n. s. 29: 187-220. BOLETZKY, S. v. 1973. 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The embryonic development of the octopus Scaeurgus unicirrhus (Mollusca, Cephalopoda). Vie Milieu 34(2/3): 87-93. BOLETZKY, S. v. 1986. Encapsulation of cephalopod embryos: a search for functional correlations. Amer. Malac. Bull. 4(2): 217-227. BOLETZKY, S. v. 1987. Juvenile behaviour. /n : Cephalopod Life Cycles, vol. II (P. R. Boyle ed.). Academic Press London. pp. 45-60. BOLETZKY, S. v. 1992. The arm crown in cephalopod development and evolution: a discussion of morphological and behavioral homologies. Amer. Malac. Bull. 10(1): ...-... BOLETZKY, S. v. & P. FIORONI. 1991. Embryo inversions in incirrate octopods: the state of an enigma. J. Ceph. Biol. 1(2): 37-57. FIG. 6 Scanning electronmicrograph of an Octopus vulgaris hatchling in dorsal view. Note the short arms with a few relatively large suckers (upper right). FIG. 7 A live Octopus briareus hatchling with its long arms and numerous suckers adhering to the substrate (glass dish) during crawling. This is a bottom living 'miniature octopus' hatched from a very large egg, showing no signs of pelagic life style. 770 SIGURD V. BOLETZKY ENGESER, T. 1988. Fossil "Octopods" - A Critical review. Jn: The Mollusca, vol. 12: Paleontology and Neontology of Cephalopods (Clarke, M. R. & E. R. Trueman, eds.). Academic Press, San Diego. pp. 81-87. FIORONI, P. 1962. Die embryonale Entwicklung der Kölliker'schen Organe von Octopus vulgaris Lam. Rev. suisse Zool. 69: 497-511. FROESCH, D. & H.-J. MARTHY. 1975. The structure and function of the oviducal gland in octopods (Cephalopoda). Proc. R. Soc. Lond. B. 188: 95-101. HOCHBERG, F. G., M. Nixon & R. B. TOLL. 1992. Order Octopoda Leach, 1818. In: "Larval" and Juvenile Cephalopods: A Manual for Their Identification (Sweeney, M. J., Roper, C. F. E., Mangold, K. M., Clarke, M. R. & Boletzky, S. v., eds.). Smithson. Contr. Zool. 513: 213-279. ITAMI, K., Y. IZAWA, S. MAEDA & K. NAKAI. 1963. Notes on the Laboratory Culture of the Octopus Larvae. Bull. Jap. Soc. Scientif. Fish. 29: 514-520. KOLLIKER, A. 1844. Entwickelungsgeschichte der Cephalopoden. Meyer und Zeller, Ziirich. 180 pp. NAEF, A. 1923. Die Cephalopoden. Fauna Flora Golf. Neapel, 35 (1-1): XIV + 863 pp. NAEF, A. 1928. Die Cephalopoden. Fauna Flora Golf. Neapel, 35 (1-2): IX + 357. O'Dor, R. K. & E. G. MACALASTER. 1983. Bathypolypus arcticus. In : Cephalopod Life Cycles, vol. 1 (P. R. Boyle ed.). Academic Press, London. pp. 401-410. PICKFORD, G. E. 1939. The Vampyromorpha. A new Order of Dibranchiate Cephalopoda. Vestnik Cs. Zool. Spol. Praze 6-7: 346-358. PICKFORD, G. E. 1949. Vampyroteuthis infernalis Chun - An archaic dibranchiate cephalopod. II. External anatomy. Dana-Rep. 32: 133 pp. QUERNER, F. R. v. 1927. Die Köllikerschen Büschel jugendlicher Octopoden, nebst einigen Bemerkungen zur Histologie der Haut dieser Formen. Z. Zellforsch. 4: 237-265. Rosson, G. C. 1932. A Monograph of the Recent Cephalopoda, Part II. British Museum (Natural History), London. 359 pp. VILLANUEVA, R. 1992. Continuous spawning in the cirrate octopods Opisthoteuthis agassizii and O. vossi: features of sexual maturation defining a reproductive strategy in Cephalopods. Mar. Biol. (in press). Voss, G. L. 1988. Evolution and Phylogenetic Relationships of Deep-Sea Octopods (Cirrata and Incirrata). /n: The Mollusca, vol. 12: Paleontology and Neontology of Cephalopods (M.R. Clarke and E. R. Trueman eds.). Academic Press, San Diego. pp. 253-276. WELLS, M. J. 1978. Octopus. Physiology and behaviour of an advanced invertebrate. Chapman & Hall, London. 417 pp. Young, J. Z. 1960. Observations on Argonauta and especially its method of feeding. Proc. zool. Soc. Lond. 113: 471-479. Young, J. Z. 1971. The anatomy of the nervous system of Octopus vulgaris. Clarendon Press, Oxford. 690 pp. Young, J. Z. 1989. The angular acceleration receptor system of diverse cephalopods. Phil. Trans. R. Soc. Lond. B 325: 189-237. YOUNG, R. E. 1972. Brooding in a Bathypelagic Octopus. Pacif. Sci. 26(4): 400-404. Revue suisse Zool. Tome 99 Fasc. 4 | p. 771-791 | Genève, décembre 1992 | | l Abundance, species richness, host utilization and host specificity of insect folivores from a woodland site, with particular reference to host architecture by Yves BASSET* and Daniel BURCKHARDT** With 3 figures ABSTRACT We studied the local abundance, species richness, host utilization and host specificity of insect folivores associated with 10 deciduous woody and 10 perennial herbaceous plant species growing in a woodland site in the Swiss Jura. Regional species richness of insect folivores on their hosts, inferred from compilation of insect faunas for Central Europe, was highly correlated with local species richness, as estimated by a 7-month field survey. Woody hosts sustained more insect species and a higher proportion of chewers, than herbaceous plants which, in turn, sustained a higher proportion of leaf miners. Overall insect abundance was not affected by plant architecture. On average, 56 % of the species feeding upon a particular host were specialists, with no apparent effect of plant architecture, but proportions for herbaceous plants fluctuated from 0 to 100 %. At the regional scale, 85 % of the variance in herbivore species richness was explained by the height of the host, its taxonomic relatedness and its leaf water content. At the local scale, 88 % of the variance could be explained, with a significant contribution of host phenology. The variances explained by models describing other local variables - such as herbivore abundance, number of specialist species, proportions of ectophagous/endophagous species and leaf palatability - were generally lower, with, sometimes, a significant contribution of sampling effort. Although confirmation will be needed from more extensive studies, particularly encompassing plants growing in other habitats, we suggest that the influence of variables such as leaf water and host phenology may have been underestimated as predictors of herbivore species richness. “ 3 Ch. Chesnaie, 1219 Chatelaine, Switzerland; present address: Bishop Museum, 1525 Bernice Street, PO Box 19000A, Honolulu, Hawaii 96817 0916, USA. te Département d'Entomologie, Muséum d'Histoire Naturelle, 1211 Genéve 6, Switzerland. 772 YVES BASSET AND DANIEL BURCKHARDT INTRODUCTION Since the seminal paper of SOUTHWOOD (1961) about the number of insect species associated with various British trees, there has been growing interest in the host-related determinants of insect species richness on particular plant species. These determinants include the abundance of the host and its distribution; its age of establishment; the number of habitats within which it grows; its taxonomic relatedness and isolation; its architecture and biomass; its leaf size and shape; and whether it is deciduous or evergreen (SOUTHWOOD, 1961; LAWTON and SCHRÔDER, 1977; LAWTON and PRICE, 1979; STRONG and LEVIN, 1979; BIRKS, 1980; CLARIDGE and WILSON, 1981, 1982; NEUVONEN and NIEMELÀ, 1981; RiGBY and LAWTON, 1981; FOWLER and LAWTON, 1982; KARBAN and RICKLEFS, 1984; KENNEDY and SOUTHWOOD, 1984; CORNELL, 1985; LEATHER, 1986; CORNELL and KAHN, 1989; JONES and LAWTON, 1991). Most authors examined the whole herbivore community associated with particular plants, but some studies concerned restricted herbivore groups and/or restricted host taxa (above references). However, few studies attempted to predict the guild structure of herbivore communities (as represented by the number of species of chewers, sap-suckers, leaf miners and gall formers) from host-related variables, with the notable exception of CORNELL and KAHN (1989) analyzing data for British trees (KENNEDY and SOUTHWOOD, 1984). Among other things, these authors concluded that models centered on the specialist/generalist dichotomy may prove to be more satisfactory in explaining the organization of herbivore communities than models derived from the relative importance of guild categories. CORNELL (1989) also advised paying attention to the ratio between ectophages (i.e., chewers) and endophages (i.e., other guilds) on plant species. To date, there is no published attempt to examine the number of specialist species, or their relative proportion, within a particular plant community. Usually, information about herbivore loads on particular host plants is compiled from published insect faunas. This practice provides reliable material for insect species richness analyses (e.g., NIEMELA and NEUVONEN, 1983; LEATHER, 1990) but presents several impediments to estimating the proportion of specialist and generalist species feeding on a particular plant using the entomocentric information provided by such literature. First, hosts for polyphagous species may not be extensively listed, hosts may be identified by their generic names only or by general quotations (such as "on herbaceous plants" or "on various trees and shrubs", etc.), and the quality of information is generally poorer for herbaceous plants than for trees (LEATHER, 1986). Second, in these faunas the apparent sampling effort (which is confounded with plant citation) is not identical among hosts: often sampling/mention of hosts of economic importance and/or of extended distribution is prevalent (see NIEMELA and NEUVONEN, 1983). Some correction for the sampling intensity is needed (e.g., KARBAN and RICKLEFS, 1983) but this is less important if species- area effects are to be demonstrated (REY ef al., 1981). However, sampling effort may be crucial when investigating the relative proportion of specialist species, because increasing sampling effort is likely to append a high number of generalist species to a particular host (i.e., if sampling is unlimited, all highly-generalist species present locally may eventually be collected on a certain plant). Most studies of host-related deteminants in herbivore species richness used multiple regression for statistical analysis. Three points are worth noting regarding the models generated by such analyses. First, certain attributes of foliage, which may influence herbivores feeding on leaves, may not explain much of the variance of insects associated with the wood, reproductive organs and roots of the host. Care must be taken to exclude those from the analyses if particular attention is given to the predictive value of SPECIFICITY OF WOODLAND INSECTS 773 independent variables related to host-foliage characteristics. Second, it is well documented that both species richness (e.g., CORNELL, 1985) and insect host-range (e.g., FOX and Morrow, 1981) differ when measured at local and regional scales. Therefore, as far as possible, it may be important to consider models predicting these two variables at both scales. Thirdly, woody and herbaceous plants are thought to display different sets of chemical defences and/or different apparency to herbivores (e.g., FEENY, 1976). Thus, host-related determinants of herbivore species richness may also differ between these two categories of hosts (see LEATHER, 1986). Our study aimed at documenting both the regional and local species richness of insect folivores associated with several host plants, with particular reference to host architecture. We sought to identify the major host-related attributes which appear to contribute significantly to herbivore species richness on these hosts. As far as possible, we tried to control and to reduce differences in sampling effort among host plants. In particular, we asked the following questions: (a) do models explaining insect species richness differ when considered at the regional and local scale?; (b) do patterns of abundance, seasonal distribution, species richness, composition of guilds and host specificity of folivore communities differ according to plant architecture?; (c) what are the best models for the prediction of the above patterns in the light of host-related attributes? We use the term "specialist" in the usual meaning of herbivores feeding upon a single species, genus or family of plants, while "generalist" refer to insects feeding on several plant families (e.g., NEUVONEN and NIEMELÄ, 1981). MATERIAL AND METHODS STUDY SITE AND HOST PLANTS The study site encompassed the upward slopes of the Swiss Jura situated in the Vendöme - Bonmont area: Grande Cöte and Petite Cöte de Bonmont, Combe de la Mey, Les Deplumeaux (Vaud, Switzerland, approximately 46° 25' N, 6° 09' E). Sampling was restricted to an altitude of 600 - 750 m. The woodlands in this area represent a mosaic of associations including principally Lathyro-Quercetum, Coronillo-Quercetum, Cardamino- Fagetum and Luzulo-Fagetum (HAINARD and TCHEREMISSINOFF, 1973). The host plants investigated included 10 deciduous woody plants (trees and shrubs) and 10 perennial herbaceous plants (Table 1). Since it was not always possible to identify hosts with certitude in the field, some of them were combined into "aggregates" (Table 1). Hereafter, the host plants and aggregates are designed by their generic names. All hosts were relatively common and widespread within the study area. As far as possible, they were chosen as representatives of different plant families growing in similar ecological situations, inside woods and/or on their margins. INSECT SAMPLING Sampling and other field analyses were performed by the senior author from April to October 1990 (11 sampling occasions). Hosts, and, for trees, only accessible branches (< 2 m high), were visually searched at random and during day-time for insect herbivores. Foliage insects, galls and mines were counted and collected: insects associated with wood, flowers, seeds and roots were not considered. As far as possible, insects were reared (when collected as juvenile instars) and identified. Due to difficulties in their identification, thrips 774 YVES BASSET AND DANIEL BURCKHARDT TABLE 1. Host plants investigated and the number of associated species of insect folivores recorded from the literature and collected during the survey. For aggregates, the first named species indicates the most common host. Hosts Plant family Nospecto Nospecie a) Woody hosts Salix caprea L. and hybrids Salicaceae 446 46 Corylus avellana L. Betulaceae 186 31 Fagus silvatica L. Fagaceae 122 38 Quercus petraea Lieblein - Q. robur L. - Q. pubescens Willd. aggr. Fagaceae 445 48 Crataegus oxyacantha L. - C. monogyna Jacq. aggr. Rosaceae 180 35 Sorbus aria Crantz and hybrids Rosaceae 100 31 Acer opalus Miller and hybrids Aceraceae 83 31 Fraxinus excelsior L. Oleaceae 80 25 Lonicera xylosteum L. Caprifoliaceae 88 20 Viburnum lantana L. Caprifoliaceae 35 18 b) Herbaceous hosts Arum maculatum L. Araceae 4 3 Paris quadrifolia L. Liliaceae 5 3 Dentaria heptaphylla Villars Cruciferae 27 13 Lathyrus vernus Bernh. Leguminosae 53 8 Euphorbia amygdaloides L. Euphorbiaceae 41 5 Mercurialis perennis L. Euphorbiaceae 7 6 Orthilia secunda L. Pyrolaceae 5 3 Melittis melisophyllum L. Labiatae 8 8 Galium odoratum (L.) Scop. Rubiaceae 17 5 Solidago virgaurea L. Compositae 45 7 and imagines of aleyrodids were counted only. Sample time and number of sampling occasions were identical for each host. However, the foliar area sampled varied among hosts: one sampling occasion consisted of > 20 samples (one different plant each) for herbaceous hosts and 10 samples (one different branch or branchlet of 30 - 50 leaves) for woody hosts. In each sample, the number of young and mature leaves/leaflets were counted. For each host and sampling occasion the mean leaf area (measurements of five leaves, totals of lower and upper sides) was determined with a transparent grid. This enabled an estimation of the leaf area sampled in each sample. Herbivore abundances were corrected by total leaf area sampled and expressed as number of individuals per 500 em? of leaf area (i.e., the sample size closest to the average sample size for all hosts). This allowed us to compare herbivore abundance between host plants and to compute an index of herbivore abundance averaged from all sampling occasions (variable Index, see below). EVALUATION OF INSECT HOST-SPECIFICITY Insect host-specificity was derived from the relevant literature (see next section) and, in cases of difficult identification, deduced from field observations. In addition, chewers SPECIFICITY OF WOODLAND INSECTS 775 were tested for feeding on the 20 hosts. Two or more individuals, kept within glass vials with 100 % R.H., were presented with fresh leaves (young or mature) of study hosts for 24 h. Feeding was scored as follows: 0 (no feeding), 1 (attempting to feed), 10 (small consumption of foliage) and 100 (extensive consumption of foliage). A logarithmic scale was chosen to emphasize regular and extensive feeding. Prior to testing, insects were fed with leaves from their presumed host and allowed to reach late instars in order to avoid inclusion in the analyses of "incidentals" (insects which rest on the foliage and do not feed; variable Incid, see below), and poor correspondence between oviposition and performance which may occur in some instances (e.g., THOMPSON, 1988). Because host- specificity indicated by these feeding tests refers to laboratory conditions, it was preferred, when available, to use the information found in the literature, which is usually relevant to field conditions. Because of problems with identification of aphids, the species of this group could not be assigned to either specialists or generalists according to the literature. Since most aphids are specialists (BORNER, 1952), all species collected were assigned to this category. VARIABLES RETAINED FOR STATISTICAL ANALYSES We inferred the regional species richness of herbivores from the literature and the local species richness from our field sampling. The data were analyzed with forward step- wise multiple regression analyses. The following dependent variables were considered. First, an estimate of the number of insect folivores associated with the 20 hosts at the regional scale was obtained by scanning the following insect faunas from Central Europe (in a few cases Northern Europe) and their host plant records (variable Nospecto): Heteroptera (STICHEL, 1955-1962); Auchenorryhncha (OSSIANNILSSON, 1978-1983); Sternorrhyncha (BORNER, 1952; VONDRACEK, 1957; ZAHRADNIK, 1963; KOSZTARAB and KOZAR, 1988); Coleoptera (FREUDE er al., 1966-1983); Symphyta (Lorenz and Kraus, 1957); "Microlepidoptera" (SWATSCHEK, 1958; HANNEMANN, 1977; PALM, 1989; HERING, 1932); "Macrolepidoptera" (BECK, 1960; FoRSTER and WOHLFAHRT, 1955, 1960, 1981); leaf miners (HERING, 1957); and gall formers (BUHR, 1964-1965). This literature also provided estimates of the regional richness in chewers, sap-suckers, leaf miners and gall formers for each plant species (variables Chwto, Sapto, Mineto, Gallto). Since we did not identify thrips, they were disregarded to ensure valid comparisons between regional and local scales. When generic host-records were available only, they were assumed to apply to the plant species considered. An estimate of the local species richness of all insect folivores (and of particular insect guilds) within the study site was provided by the number of species recorded on each host during all sampling occasions (variables Nospecie (Tablel), Chw, Sap, Mines and Galls). Other dependent variables included Index and Incid, already defined, the local number of specialist species (Speciali), the local percentage-ratio of specialist species (Ratiospe, %), of ectophagous and endophagous species (Ratioect and Ratioend, %), and the mean scores for each host in feeding experiments for young leaves, mature leaves and all leaves combined (Scoyoung, Scomatu and Scotot, units = relative scores). Since feeding tests involving insects and plants from which they were collected were excluded from the analysis, these scores represented a measure of plant palatability relatively independent of the number of chewers collected from the host. The independent variables used in the analyses are defined in Table 2, and some only, due to space limitations, are detailed further in Appendix 1. Water content and specific weight of young and mature leaves were determined by oven-drying at 100 °C [20 776 YVES BASSET AND DANIEL BURCKHARDT discs punched from each host, collected on the same day, at noon, in late May (young -leaves) and early August (mature leaves)]. Despite the difficulty of characterising host biochemistry, we attempted to use two different variables related to host chemical defenses. First, an index of the diversification of plant chemical defences was provided by the variable Chemic, which represented the number of broad categories of chemical compounds present in the host and known to be active against herbivores. The choice of the chemical categories followed ROSENTHAL and JANZEN (1979) and included: toxic non- protein amino-acids, cyanogenic compounds, alkaloids, glucosinolates, terpenoids, saponins, flavonoid pigments, tannins, other phenols, coumarins, cardenolides, phyto- ecdysones and accumulation of silica. This information was abstracted from HEGNAUER (1962-1989), GIBBS (1974) and RAFFAUF (1970). Second, an estimation of the isolation of host chemical defences was provided by the variable Disschem, which was the similarity measure (euclidean distance) with an average and hypothetical host possessing the most common categories of chemical defences (i.e., the categories found in more than 10 of the 20 study plants). In July 1990, the abundance of the 20 host-species was recorded in 50 plots of 10 x 10 m within the study site. These plots were representative of locations sampled and provided estimations of the local abundance of hosts (Locabund), their local distribution (Locadist) and aggregation (Aggrega). The variable Compac represented a measure of the compactness of host foliage. For herbaceous hosts, this was defined as the leaf area of the whole plant and for woody hosts as the total leaf area supported by 50 cm of branch. Both dependent and independent variables were log (x+1) transformed (natural base) in order to satisfy the assumption of normality. In most cases, transformation of data improved the fit to the models. The step-wise multiple regressions were computed with «- to-enter/remove = 0.150 (BENDEL and ALFIFI, 1977) and are presented with their adjusted coefficient of determination. Variables with a tolerance value < 0.2 were deleted from the model, to avoid multicollinearity problems. More conservative models were also computed, with the regression being halted after the last variable with a significant (p < 0.05) parameter entered the model. RESULTS “COMPOSITION OF HERBIVORE FAUNAS The estimation of regional species richness of insect folivores as recorded in the literature from Central Europe was highly correlated with the estimation of local species richness provided by the field survey (Table 1; r = 0.85, p < 0.001). Regional and local species richness also correlated with the total number of arthropod herbivores (insects and mites) recorded from corresponding British tree genera studied by KENNEDY and SOUTHWOOD (1984) (r = 0.98, p < 0.001 and r = 0.91, p < 0.01, respectively). The species richness of chewers, sap-suckers, leaf miners and gall formers was similarly well correlated between local and regional scales (r = 0.75, 0.88, 0.78 and 0.76, respectively, p < 0.001 in all cases). Although these general correlations appear to be good, Table 1 suggests that assessments of local herbivore richness may have been underestimated by the field survey on the following plants: Salix, Quercus, Lathyrus, Euphorbia and Solidago. In particular, few Chrysomelidae and Symphyta were collected from Quercus and Salix; no leaf miners from Lathyrus; few leaf miners and Chrysomelidae from Euphorbia; and no Lepidoptera from Solidago. It may also be of interest to note that, with SPECIFICITY OF WOODLAND INSECTS Sele) TABLE 2. Independent variables determined for each host and used in multiple regression analyses. Variables indicated with '*' were determined empirically. Coding variable Description Lwe *, Lwcy * leaf water content of young and mature leaves [% DW] Slw *, Slwy * specific leaf weight of young and mature leaves leaves [g-4 x cm-2] Chemic number of broad categories of chemical defences (see text) Disschem dissimilarity of chemical defences with an average and hypothetical host (see text) Congensp taxonomic relatedness: no. of congeneric species in AESCHIMANN and BURDET (1989) Confamsp taxonomic isolation: no. of confamilial species in AESCHIMANN and BURDET (1989) Locasp * no. of congeneric species present within the study site (1) Abund abundance of woody hosts in Switzerland (Anon., 1988) [1000 m3] Locabund * local abundance (total no. individual censued, see text) Aggrega * local aggregation (coefficient of variation for local abundance, see text) Distger distribution in West Germany (no. of 10 x 10 km squares in HAEUPLER et al., 1988) Distsw distribution in Switzerland (no. sampling sectors in WELTEN and RUBEN SUTTER, 1982) Distsg distribution in West Germany and Switzerland (Distger + Distsw) (2) Locadist * local distribution (no. plots in which the host was present, see text) Height maximum height in AESCHIMANN and BURDET (1989) [cm] Leafar * leaf area of a single leaf/leaflet [cm2] Shape leaf shape: 1 = entire; 2 = dented; 3 = lobed; 4 = dented-lobed Width * ratio max. width to max. length of leaf/leaflet Compac * foliage compactness (leaf area within 50 cm of branch, see text) [cm2] Pheno * no. of days during which young leaves (leaves pale green and of tender texture) were observed within the study site in 1990 Hairs * no. of hairs per cm2 of area (leaf underside) Habitats no. of habitats occupied by the host in AESCHIMANN and BURDET (1989) Areatot * total sum of leaf area sampled [m2] (1) Determined on the basis of field notes and on the distribution atlas of WELTEN and RUBEN SUTTER (1982) (2) Swiss sampling sectors represented in average 100 km?. the exception of Solidago, these plants are represented by many congeneric species which are often mentioned by their generic names in published insect faunas. At both the regional and local scales, woody plants sustained significantly more herbivore species than herbaceous plants (t = 5.61, p < 0.001 and t = 7.96, p < 0.001, respectively). The proportion of incidental insects collected was lowest in Arum, Crataegus, Euphorbia and Solidago (between 0 and 0.7 % of the total number of individuals censused, Table 3) and highest in Paris, Orthilia, Mercurialis and Melittis (between 8.7 and 48.0 %). In total, 2.2 % of insects collected from study hosts were considered to be incidentals. The distribution of species within the different guilds, as recorded during the field survey, is indicated for each host in Fig. 1. At the regional scale, corresponding data were tested by contigency table analysis for non-uniformity. This showed that the distribution of species was significantly non-uniform among woody hosts (G-test, G = 121.0, p < 0.001), as already noted by CORNELL and KAHN (1989) for British trees, and also among 778 YVES BASSET AND DANIEL BURCKHARDT TABLE 3. Local dependent variables recorded from host plants: index of herbivore abundance, number and percentage of specialist species, number of incidental insects collected, total number of individuals censused and mean scores obtained in feeding experiments (young and mature leaves, all leaves combined and number of insect species tested). Host Index (s.e.) Speciali (Ratiospe) Incid Censused Scoyoung Scomatu Scotot n Salix 1.94 (0.56) 33 (71.7) 14 405 35.3 36.1 35.6 49 Corylus 0.99 (0.17) 14 (45.2) 19 343 19.6 25.6 22.8 47 Fagus 2.26 (0.42) 18 (47.4) 19 793 45.2 36.8 42.1 68 Quercus 3.44 (0.41) 30 (62.5) 24 1046 36.8 22.1 29.3 57 Crataegus 3.88 (0.66) 24 (68.6) 1 343 31.0 18.5 26.1 51 Sorbus 1.87 (0.25) 15 (48.4) 9 643 24.3 20.6 22.6 47 Acer 2.68 (0.42) 17 (54.8) 15 1445 37.1 28.0 33.3 70 Fraxinus 0.95 (0.14) 12 (48.0) 6 342 30.4 8.6 20.5 62 Lonicera 1.54 (0.35) 17 (85.0) 11 183 12.9 14.1 13.5 53 Viburnum 1.72 (0.45) 12 (66.7) 18 687 10.5 15.7 135 52 Arum 0.59 (0.17) 0 (0) 0 17 0.2 0 0.1 35 Paris 0.44 (0.10) 3 (100.0) 2 23 0.2 8.0 3.2 36 Dentaria 1.33 (0.32) 9 (69.2) 15 174 0.6 10.1 4.1 41 Lathyrus 0.91 (0.22) 2, (25.0) 2 30 23.9 12.2 18.3 40 Euphorbia 13.89 (2.81) 4 (75.0) 2 469 5.9 13.5 8.9 43 Mercurialis 0.29 (0.07) 4 (66.7) 4 23 15.1 6.2 11.7 47 Orthilia 0.47 (0.16) 1 (33.3) 1 9 1.5 6.9 4.3 31 Melittis 0.34 (0.07) 2 (25.0) 12 25 0.7 1.8 1.0 33 Galium 1.17 (0.39) 2 (40.0) 1 27 0.5 0.1 0.4 37 Solidago 7.34 (0.87) 6 (85.7) 6 855 0.1 15.7 6.9 32 herbaceous hosts (G = 49.2, p < 0.01). The same type of distribution was also tested for non-uniformity between an average woody host and an average herbaceous host (sum of all species per guild in each host category). This revealed a significant influence of host architecture on the distribution of species within guilds (G = 34.4, p < 0.001): the proportion of leaf mining species was high in herbaceous plants, while that of chewers was high on woody hosts. Similar analyses were difficult to perform with local data because the frequencies were too low in many cells. However, comparison of average woody and herbaceous hosts showed no significant effect of host architecture at the local scale (G = 3.820, 01122); At the regional scale, the distribution of species within ectophagous and endophagous types was non-uniform among woody and herbaceous hosts (G = 46.0, p < 0.001 and G = 19.8, p < 0.05, respectively), and non-uniform between the average woody and herbaceous hosts (G = 4.5, p < 0.05). The latter sustained a high proportion of endophages. At the local scale the trends were different: the distribution was uniform among woody plants (G = 8.2, p = 0.516), non-uniform among herbaceous plants (G = 33.3, p < 0.001), and uniform between host categories (G = 0.1, p = 0.722). HERBIVORE ABUNDANCE, HOST UTILIZATION AND INSECT HOST-SPECIFICITY Mean abundances of insect folivores, as determined during the whole interval of the field survey, are presented in Table 3. They did not differ significantly between woody and herbaceous hosts (t = 0.38, p = 0.708), but the most extreme values were recorded on SPECIFICITY OF WOODLAND INSECTS 779 Salix Corylus Fagus Quercus Crataegus 257- 118- 60-11 84- 69- 29- 4 61- 45- 12-4 213- 125- 78- 29 93- 47- 36- 1 21 1 1 1 Sorbus Acer Fraxinus Lonicera Viburnum 39- 29- 32-0 30- 39- 9-5 27- 45- 5-3 36- 26- 19-7 8- 21- 5-1 Arum un Dentaria Lathyrus Euphorbia 18- 2- 7-0 17- 10- 24- 2 17- 12- 10- 2 Mercurialis Orthilia Melittis Galium li C]CHEWERS [}SAP-SUCKERS E LEAF-MINERS [ll GALL-FORMERS Ectophages Endophages Fic. 1. Distribution of the number of species per guild, as recorded for each host during the field survey. Regional data are indicated below each host. the latter (highest values: Euphorbia and Solidago; lowest values: Mercurialis and Melittis). Seasonal abundance of herbivores was significantly non-uniform among woody plants and among herbaceous plants (unpubl. data; G = 1659.7 and G = 3090.2, respectively, p < 0.001). Seasonal distribution was also non-uniform between average woody and herbaceous hosts (G = 1172.7, p < 0.001): herbivore abundance was high in late April - early May on the former, decreased in mid-July and rose again by mid- October, whereas on the latter, it increased until the end of June, fell sharply by early July and increased again later, but not as markedly as previously. The number of species collected on young foliage, mature foliage and on both foliage types is indicated in Fig. 2. This provided only a rough idea of host utilization (1.e., the use 780 YVES BASSET AND DANIEL BURCKHARDT of young and mature foliage), because sampling effort differed between foliage types (on ‘average 4 out of 11 sampling occasions were concerned with young foliage), and collection of an herbivore on a particular type of foliage does not imply that it can use it successfully. Contingency tests showed that "host utilization" was uniform among woody and herbaceous hosts (G = 9.8, p = 0.365 and G = 5.8, p = 0.759, respectively), and also between average woody and herbaceous hosts (G = 0.03, p = 0.865). A different assessment of host utilization by insect chewers was given by the scores obtained during feeding experiments (Table 3). In some instances, young foliage was apparently more palatable than mature foliage (Fagus, Quercus, Crataegus, Sorbus, Acer, Fraxinus, Lathyrus and Mercurialis), in other cases the opposite applied (Corylus, Lonicera, Viburnum, Paris, Dentaria, Euphorbia, Orthilia and Solidago), and there was no clear trend in others (Salix, Arum, Galium and Melittis). Paired t-tests indicated that palatability was not significantly different between young and mature foliage within woody (t = 1.97, p = 0.08) and herbaceous hosts (t = 0.97, p = 0.356). However, the palatability of both foliage types was much higher in woody than in herbaceous hosts (t = 5.79, p < 0.001). Salix Corylus Fagus Quercus Crataegus Sorbus Acer Fraxinus Lonicera Viburnum Arum CE Paris (TA Dentaria Lathyrus SS Euphorbia IE Mercurialis EEE Orthilia CEE Melittis Galium Solidago No. sp. collected 0 10 20 30 40 50 60 O on young foliage El on mature foliage M on both foliage types FIG. 2. Number of species collected on different types of foliage during the field survey. The number of specialist species (and their percentage of the total number of species) collected on each plant is indicated in Table 3. The highest specialist ratios were found on Paris, Solidago, Lonicera, Euphorbia, while the lowest occurred on Arum, Lathyrus and Melittis. Overall, these ratios were not significantly different between woody and herbaceous hosts (t = 0.72, p = 0.486), and, on average, amounted to 55.9 + 5.38 % (s.e.) when all hosts were considered. Similarily, contigency table analysis did not reveal any influence of plant architecture on the distribution of species within specialist and generalist types (G = 0.6, p = 0.439). The scores for chewer species tested in feeding experiments are summarized in Fig. 3. Few species accepted more than 5 hosts as suitable for feeding. Both this data set and the former indicate that most of the insects collected were specialists. SPECIFICITY OF WOODLAND INSECTS 781 60 56 di 40 O 38 LU (= (dp) LL = 22 © 20 14 10 7 4 0 100* 100 200 300 400 500 >500 CLASS OF SCORES Fic. 3. Scores of feeding experiments for chewing species. The first class of score (100*) refers to the number of species which could only be tested once, on the host from which they were collected. MULTIPLE REGRESSION ANALYSES A correlation matrix for selected dependent variables is presented in Table 4. A weak positive correlation existed between the overall abundance of herbivores and the palatability of mature leaves. This suggested that this last factor was important for large build-ups of herbivore populations during the growing season. The proportion of specialists was not correlated significantly with herbivore abundance. The species richness and the proportions of specialists and ectophages were positively correlated with the palatability of leaves, particularly that of young leaves. The results of multiple regressions analyses are detailed in Table 5, with particular reference to the composition of herbivore communities at the regional and local scales. For the former, 85 % of the variance was explained by the height of the host, its taxonomic relatedness, water content of young leaves, and, to a lesser extent, by its taxonomic isolation. At the local scale, 88 % of the variance could be explained, with height and host phenology entering first and second in the regression. In general, the predictors of models for regional and local data were different. The local species richness of chewers could only be significantly accounted for by the total leaf area sampled, a measure of the sampling effort, and water content of young leaves. Table 6 summarizes the multiple regressions with particular reference to plant architecture. The predictors were also quite different between host categories. For woody hosts, 58 % of the variance could be accounted for by the number of habitats in which the host grows and its height (regional data). In contrast, 69 % of the variance was explained in herbaceous plants by leaf width. The number of insect specialists collected on these hosts depended also upon leaf width. 782 YVES BASSET AND DANIEL BURCKHARDT TABLE 4. Lower correlation matrix (Pearson coefficient) for selected dependent variables. Index Ratiospe Scoyoung Scomatu Ratioect Ratiospe 0.35 — — — — Scoyoung 0.18 0.29 = — — Scomatu 0.45 * 0/69 ONDES ~ — Ratioect 0.06 0.29 0.602 0.54 * - Ratioend 0.26 0.15 0.16 0.25 0.78 *** Nospecie 0.25 0.35 0:82,27 0722, OST Nospecto 0.49 * 0.40 OT OTIS) to 0.42 pO Osa p< OO asp < 0.001 TABLE 5. Summary statistics of stepwise multiple regression analyses performed on the 20 hosts, with particular reference to the composition of herbivore communities at the regional and local scales. The order of the variables in the final equation follows the step in which they entered the regression. Data in brackets indicates the R of the conservative model (see text), and the corresponding number of variables which entered the regression (same order, starting from the left). Dep. var. Regression equation R2 F-test (R2, n) (a) Regional data Nospecto 11.79 + 0.60 Height*** + 0.49 Congensp** - 2.95 Lwcy* + 0.20 Confamsp 0.85 28. O*** (0.83, 3) Chwto 6.60 + 0.48 Height*** + 1.96 Pheno*** - 3.78 Width** - 3.22 Lwc 0.83 DAS (0.80, 3) Sapto 14.09** + 0.53 Height*** - 3.40 Lwcy** + 0.25 Congensp* 0.90 5082 = Mineto 13.05 - 3.56 Lwc* + 0.92 Habitats + 0.37 Congensp* + 0.34 Confamsp* + 0.29 Height* 0.76 eps (0.60, 2) Gallto -0.10 + 0.28 Height** + 0.37 Congensp* -3.23 Disschem 0.60 10.4*** (0.53, 2) (b) Local data Nospecie -2.98** + 0.40 Height*** + 0.78 Pheno** 0.88 VL u Chw 11.06 + 0.50 Areatot** - 3.53 Lwcy** 0.72 DATE ive Sap -5.05*** + 0.31 Height*** + 0.88 Pheno*** + 0.72 Chemic** 0.93 SOS — Mines 13.94*** - 3.34 Lwc*** + 1.11 Habitats** 0.69 DIETE ~ Galls 0.03 + 0.18 Height** - 2.04 Disschem* 0.64 17:05 Significance of parameter estimates: *p < 0.05, ** p < 0.01, *** p < 0.001 Table 7 details the models relevant to local independent variables. Herbivore abundance was related principally to the number of habitats in which the host grows, its distribution in Switzerland and its local aggregation. The number of specialist species was dependent upon the height and the phenology of the host. The proportions of specialists, ectophages and endophages could not be adequately explained by our variables. The palatability of the different types of foliage depended on several predictors, among which were more particularly sampling effort and leaf water. Not surprisingly, sampling effort was the most significant predictor explaining the number of incidental insects collected, but there were also weak contributions from host aggregation, chemical defences and pubescence. SPECIFICITY OF WOODLAND INSECTS 783 TABLE 6. Summary statistics of stepwise multiple regression analyses performed with particular reference to host architecture. Presentation follows Table 5. Dep. var. Regression equation R? F-test 5 (R2, n) (a) Woody plants Nospecto -0.84 + 2.22 Habitats* + 0.50 Height* 0.58 USE - Nospecie 15.21** - 2.87 Lwc** 0.56 12.4** - Speciali Not significant — - - (b) Herbaceous plants Nospecto 4.74*** - 5.76 Width** 0.69 20.8** - Nospecie 0.23 + 0.20 Confamsp* -0.22 Distsg* + 0.54 Pheno 0.75 RÉF (06722) Speciali 2.54*** - 3.57 Width* 0.53 11.0* - TABLE 7. Summary statistics of stepwise multiple regression analyses performed for local dependent variables. Presentation follows Table 5. Dep. var. Regression equation R? F-test (R2,n) Index 5.57** + 1.68 Habitats*** - 0.62 Distsw** - 0.42 Aggrega* - 0.18 Leafar 0.56 TAO (0.51, 3) Speciali -5.43** + 0.38 Height*** + 1.22 Pheno** 0.79 SE = Ratiospe Not significant — - — Ratioect 34.17** - 7.24 Lwcy* 0.25 TSE — Ratioend Not significant = — = Scoyoung 13.46 + 0.50 Height*** - 3.98 Lwcy* + 1.73 Chemic 0.70 STE (0/662) Scomatu 17.50* + 0.35 Areatot* -3.73 Lwcy* - 1.62 Chemic* 0.66 Bee Scotot 9.35 - 3.46 Lwc** + 0.38 Areatot* + 0.74 Pheno 0.74 ISOTTA (0571562) Incid 1.40 + 0.48 Areatot*** - 0.44 Aggrega - 1.49 Chemic* + 0.11 Hairs 0.76 SES (DCE AL) DISCUSSION SAMPLING PROCEDURE The limitations of our study are straightforward. Local estimates of species richness rely on data obtained during only a 7-month field survey. The number of sibling species may have been underestimated in the field, but there is no reason to believe that this factor was biased towards particular host plants. Sampling bias appeared identical among hosts, but insects associated with trees were collected from low branches only (sampling effort as measured by the total leaf area sampled is a different problem which is addressed below). The insect data are valid for common species, easy to discover on the foliage during day- time. Some herbivores may spend most of their time hidden in the leaf litter, and may feed on their hosts during short periods only. Those species, along with nocturnal and highly active ones, are likely to have been underestimated by the field survey. Increasing the 784 YVES BASSET AND DANIEL BURCKHARDT number of sampling occasions might also have increased the contrast in species richness . between particular hosts. Furthermore, our data set is relevant only to part of a particular plant community, namely plants growing in woodlands of the collinean level. Inclusion of other plant species, particularly those growing in different habitats, might have altered the patterns described. REGIONAL AND LOCAL PATTERNS OF SPECIES RICHNESS The general correlation between estimates of species richness at the regional and local scales was good. In particular, estimates of local species richness were close to those of regional richness for plants which, presumably, are infrequently surveyed by entomologists and/or cited in insect faunas (e.g., Melittis, Mercurialis, Arum). However, the predictive models for herbivore species richness at the regional and local scales differed greatly in terms of the relative contribution of the significant variables. Some ecological factors - such as local aggregation and phenology of the host - may act as significant selective agents, influencing locally the commonness and rarity of certain species. However, their influence may be cancelled out at the regional scale, 1.e., when host attributes are examined over a larger number of ecological situations and habitats. CORNELL (1985) pointed out that if local interactions are strong, then correlation between species richness at the regional and local scales should be weak. However, the regional data of our study do not formally represent regional data sensu CORNELL (1985), since both distributional and insect data did not relate to the entire geographical range of host plants, because of the lack of suitable data (see below). INFLUENCE OF PLANT ARCHITECTURE ON OBSERVED PATTERNS The slight increase in herbivore density observed on most woody hosts in mid- October corresponded probably to searching for hibernation sites and, thus, appeared to be directly related to plant architecture. Herbaceous plants sustained a higher proportion of leaf mining species, and, more generally, of endophages, than did woody hosts. However, these trends were more marked for regional data than for local data. Concealment of species in herbaceous plants is likely to be related to several factors. Amongst others, avoidance of predators and parasitoids and/or weather effects may be crucial when the architecture of the host is relatively simple. At the local scale, herbivore abundance and proportion of specialists were not significantly different between the two host categories. Although the last observation was unexpected, the variance in the proportion of specialists was much higher in the case of herbaceous than woody hosts. This suggests that biochemical conditions for herbivores are more varied in the former than in the latter. Indeed, out of the 14 species which could be considered as highly generalist (feeding score > 500), only one was collected from herbaceous plants. Different authors have emphasized that non-apparent plants, such as herbs, may have evolved towards chemical diversification (e.g., FUTUYMA, 1976; SCRIBER and FEENY, 1979). At both regional and local scales, herbivore species richness was differently explained on woody and herbaceous hosts but there was no obvious contribution of plant biochemistry (hovewer, see discussion of this variable below). HOST-RELATED VARIABLES AND THEIR PREDICTIVE VALUES The effects of certain variables, which were determined locally, such as water content of young and mature leaves and host phenology, were significant at the regional scale. SPECIFICITY OF WOODLAND INSECTS 785 However, it remains to be seen if their predictive value is really significant when intraspecific differences are accounted for, when average values are determined over most of the area of distribution of hosts, and when host plants growing in different habitats are included in the analyses. Sampling effort, as represented by the total leaf area sampled per host, had a significant effect in the models describing the local species richness of chewers, the number of incidental insects collected and the palatability of mature leaves. That effect was primarily due to sampling of much smaller foliage areas for herbaceous hosts than for woody hosts (Appendix 1), a deliberate procedure to maximise time investment during sampling. After removing the effect of sampling, in testing residuals as dependent variable, leaf water and leaf area only had a significant effect on the local species richness of chewers, but the variance explained was low (R? = 0.30). These conclusions differ slightly from those of KARBAN and RICKLEFS (1984) who found that, after removing the effect of sampling effort, Lepidoptera species richness could not be explained significantly by chemical traits (including leaf water) of the foliage of 33 deciduous tree species in Ontario. The height and the leaf water of the host were important predictors in the various models tested. They may reflect differences in plant architecture, but also seem to possess intrinsic predictive power. This was demonstrated by introducing a dummy variable accounting for plant architecture: this variable did not enter in the regressions describing species richness at the regional scale, and when it entered in other models, it did not totally cancel the effect of height and leaf water. No satisfactory estimation of total leaf area for woody hosts, which would probably have been highly correlated with height, could be introduced in the models. Total leaf area did not significantly affect herbivore species richness on herbaceous plants. Since leaf water is usually correlated with leaf nitrogen and limits its assimilation, this factor is important in the nutrition of insect herbivores (MATTSON and SCRIBER, 1987). However, leaf water was negatively correlated with species richness. This may be related to the observation that leaf water content often reflects different strategies of plant chemical defences, such as the distribution of carbon- and nitrogen-based secondary compounds (MATTSON and SCRIBER, 1987). Alternatively, plants with low water content may grow in sunny or xeric conditions, which, in turn, may be more suitable for sylvestral herbivores exploiting cool and shady habitats, such as reported here. Since leaf water is likely to vary through seasonal and daily cycles (SCRIBER, 1977), the analyses presented here, which are based on discrete leaf water measurements, should be confirmed by more extensive studies. NIEMELÄ and HAUKIOJA (1982) presented evidence that the extent of the shoot-growth period in deciduous trees affects the seasonal distribution of Macrolepidoptera species richness. Furthermore, RAUPP et al. (1988) showed that the distribution of some generalist Lepidoptera is strongly influenced by phenological differences in host suitability. In our study, an estimation of the number of days during which young leaves were available at the study site proved useful for predicting the regional species richness of chewers, the local species richness of folivores in general, of sap-suckers and of specialists. Young leaves were still available on certain tree species after most of the foliage had matured (particularly on Salix), or secondary shoot-growth occurred during summer. These "islands" of young foliage often sustained high grazing damage, as reported in NIEMELÄ and HAUKIOJA (1982), and might have increased herbivore species richness locally. These observations remain to be tested at the regional scale. They are unlikely to be general and will probably depend on the particular host plants included in the analyses, since the palatability of young leaves was not always higher than that of mature leaves. 786 YVES BASSET AND DANIEL BURCKHARDT Taxonomic relatedness and isolation, as represented by the variables Congensp and _Confamsp, contributed significantly to several models at the regional scale. These observations concur with other studies (see introduction). Since Congensp and Congenfam accounted for additional and different proportions of the variance in the models, with the former variable being often more significant, this suggests that their effects may be more important respectively for monophagous and oligophagous insects. The "local" taxonomic isolation of the host appeared less important and, in contrast with other studies (e.g., KENNEDY and SOUTHWOOD, 1984), the number of plant species recorded within the plant order of the host explained less variance than the previous variables. Many authors studying models of herbivore species richness on their hosts considered that at least some of the variance left unexplained could be attributed to the chemical defences of the host (e.g., CLARIDGE and WILSON, 1982; JONES and LAWTON, 1991). In our study, this factor influenced marginally some of the models tested, but this need not imply that it has little influence on herbivore species richness. Other variables, such as taxonomic relatedness and isolation (CLARIDGE and WILSON, 1982) may better describe the influence of host biochemistry. The number of broad chemical categories accounted for a small proportion of the variance in local species richness of sap-suckers and in the palatability of the foliage. However, it was a very crude index of the diversity of chemical defences, since it did not contain information about the concentration levels and the chemical diversification of secondary compounds within the categories of defences defined. The chemical dissimilarity with an hypothetical host accounted for some variance in the regional species richness and the local number of gall forming species. This variable is as good as the choice of the average host. In this simple analysis, the average host turned to be equivalent to Quercus (Appendix 1), and, therefore, it was not surprising that the variable Disschem influenced the model describing the number of gall forming species, which are well-diversified on Quercus (e.g., BUHR, 1964-1965). The regional distribution of host did not contribute to the variance in herbivore species richness at the regional scale. Although species-area relationships account for a significant proportion of the variance (20-90 %) in species richness (references cited in the introduction), recent analyses by CLARIDGE and EVANS (1990) showed that using more accurate distributional data may result in a sharp decrease of the variance explained. Currently, distribution maps of all study hosts are not available for Europe, or for Central Europe. We chose the Swiss and German data as a substitute for regional distribution because they were accurate and concerned adjacent areas. Data covering more extended and adjacent regions of Europe would have probably increased the importance of host regional distribution for herbivore species richness. Another related variable, local host aggregation, was important in some models, whereas local host distribution and abundance were insignificant. Although most of the hosts analysed included plants growing in woodlands only, the number of habitats occupied by the host was also important in several of our models, as reported in other studies. Inclusion in the analysis of other plants growing in various habitats would have probably increased further the significance of this factor. Lastly, the age of establishment of the host could not be considered in our analyses, since similar data as are available for trees (BIRKS, 1980) were not available for shrubs and herbaceous plants. PREDICTIVE MODELS FOR INSECT TROPHIC CATEGORIES AND HOST-SPECIFICITY To date, the information compiled for British trees by KENNEDY and SOUTHWOOD (1984) represents the best regional-scale data of this kind. CORNELL and KAHN (1989) used SPECIFICITY OF WOODLAND INSECTS 787 this data set to predict insect species richness by trophic categories. They found that between 54 % and 83 % of the variance, depending on insect guild, could be explained by host abundance, taxonomic isolation, age of establishment and "evergreenness". The present study suggests that an additional part of the variance could be accounted for by leaf water-content and host phenology. Inserting leaf water in the model describing species richness of British data (7 tree genera were common to both studies) resulted in an improvement from 38 % of variance explained (only tree abundance significant) to 62 % (tree abundance first, leaf water second). ZWÖLFER and BRANDL (1989) suggested that the organization of endophytic herbivore communities is more structured than that of ectophytic herbivore communities. Assuming that a highly structured community should also be highly predictable, our data neither support nor refute their hypothesis, as proportions of endophagous and ectophagous species were both difficult to predict. Most information available regarding the proportion of specialist species feeding on particular host plants has been obtained from compilation of insect faunas. For instance values reported by NEUVONEN and NIEMELA (1981), ROWELL-RAHIER (1984) and LEATHER (1985) range from 10 % to 66 %. The high proportions of specialists reported in the present study (average 56 %) result probably from the bias of the sampling procedure towards abundant species, which are more likely to be specialists (see GASTON and LAWTON, 1988). Although the effect of sampling effort was controlled in our study, no model could be satisfactorily fitted to account for the proportion of specialists. This contrasts with the variance explained in herbivore and specialist species richness (88 % and 79 %, respectively) and emphasizes how much is still to be learnt in the field of insect-plant interactions. Despite the significant effects of sampling effort, the number of incidental insects collected appeared to be related to host aggregation and chemical defences. These factors may be important in processes of host colonization by introduced insect species. In conclusion, this study revealed that some host-related attributes, rarely considered in other studies of a similar kind, may have some predicting power in models describing the organisation of herbivore communities on their host plants. Other host-related attributes which were not considered here, such as the history of interactions between insects and their hosts, and the genetic variability of the host population (MADDOX and RooT, 1990), may also have significant effects on herbivore communities. Furthermore, this is likely to be true of other variables not directly related to the host, such as the local abundance and foraging efficiency of predators/parasitoids, and the history of speciation within insect taxa. Integration of all of these factors, if possible, should greatly improve our understanding of insect-plant interactions. ACKNOWLEDGMENTS We are indebted to M. Krähenbuhl for botanical advice and access to botanical literature, and to E. Danzig, M. Döberl, A. F. Emelyanov, P. Scherler and S. E. White- bread for identification of the material. N. D. Springate checked the manuscript, which benefited from comments by R. Brandl, R. H. Cowie, S. R. Leather and M. Rowell- Rahier. The senior author acknowledges gratefully the Muséum d'Histoire Naturelle de Genève for financial support towards travel expenses. YVES BASSET AND DANIEL BURCKHARDT 788 O'S 90 6€ 60 8r TC GE 16 eG cal L'ST CE OLI 007 8 LT 97 vol 861 L'8I Lvl ya TAINAN A mm Nm mm T8 c6 67 09 66 96 v8 68 c9 IL Tel CCI cs O£T vol 8cl sc vol COT C91 0c 0 vo0 vv 0 69°0 1240) 670 seo LT'O 09°0 080 OL'0 19'0 070 tcl L9'0 590 09°0 69°0 9L'0 0S'0 OI TE J0J291V SJeJIQUH ououd © 1 ie © © no nn mn CO = G 67 CI € 08€ C8C LIT 1270 CLT dal ITT 101 [4073 OIC €8C ECE coE 8 Tee 1270 SCC 687 OTE Ive 9YLI LSST vST Lvv VISI 1440 CCL OI 8871 SAIT trl ETET 6€07 I 009 5881 IrsI €107 LIOT 9107 6 891 6 8€I 6 tcl S'S8c 6°0€I € 6SI V VIC S'OST 8 8c€ 0’E69 Crs E Cel Cc 981 S'OSI 9971 6 6€C L'ITC t'S9 8 ECC erch 877 VOL LIT GS Ich val SLI erc Lvy'0 85T°0 Lyv0 8Sc 0 Lyv'0 Lyv'0 Lrr'O OTSO cE9°0 Lyÿ'0 8570 S9€°0 Lrr'O 8570 S9€°0 Lrr'O 0 8570 8570 S9£'0 06 Of OS OC SE 09 SE 09 Of OS OST 007 OOSE 0071 00SI 007 0007 OOSE 006 0001 yryrnwonw rounwowonnwowoeworwrt © Ot WIpiM dswejuoD dsusZuoy Isıpe9o] MSISIQ MÖISIQ = e80188y punqeoo] woayossiq IUT) JUSIOH "| XIGNaddV S'E8 C'E8 9'8L (475 L'6L 99 LOL 6 6L 618 8 98 €09 EOE 6SL L9 S'L9 Les cs9 SOL rL9 9°99 KOM] ‘Z OJge,L Ul se SJIUn pue soqeitea BUIPOD ‘SosAeue uOIss21391 ajdrpnu ur posn sofgeLieA Juspusdapur paJ9919S OTL L'69 0'OL 6 19 9'YL S'IL 8 89 0'08 USL 6 88 €79 T's) ÿ 59 £'8s 975 1229 SIG Sys S'I9 LA MT 03Dpuos LIDO) SION ALU EZIO) SIIDLAINIAO A D1g10ydn7 SNAKYIDT DIADJUIG SDA UnA WNUANGIA D1991U0T SNUIXDA ] 109 SNQUOS SNTIDIDLI SNIAENG SNSD + snjd10) MIIDS) SPECIFICITY OF WOODLAND INSECTS 789 REFERENCES AESCHIMANN, D. and BURDET, H. 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Fiora isa siro si CANNES #3 ng N be uit by sisi i Lar | | | | | | | Revue suisse Zool. | Tome 99 | Fasc. 4 | p.793-819 | Genève, décembre 1992 | | | | Notes sur quelques Brycon des bassins de l'Amazone, du Parana-Paraguay et du Sud-Est brésilien (Pisces, Characiformes, Characidae) par J. GERY* & V. MAHNERT** Avec 15 figures ABSTRACT Notes on some Brycon (Pisces, Characiformes, Characidae) from the Amazon, Parana-Paraguay and South-Eastern Brazil systems. — Most of the types of the Brycon spp. from the river systems enumerated in the title have been studied, together with additional material from the Amazon and La Plata system. From the Amazon basin, four species are recognized and redefined in the group with the well-known colour-pattern of B. melanopterus, viz. B. cephalus, B. melanopterus, B. bicolor and an undescribed species from the Rio Madeira basin. In Paraguay, two species have been recently collected and are redescribed, viz. Brycon orbignyanus and B. microlepis, often called B. hilarii by authors. It is shown that the true B. hilarii, together with B. opalinus and B. orthotaenia, are restricted to the South-East of Brazil. INTRODUCTION Jusqu'au catalogue de HOWES (1982), seules des études sporadiques, la plupart partielles et surtout consacrées aux espèces de l'Amérique centrale et du Nord-ouest de l'Amérique du Sud, avaient été publiées sur le genre de Characidae Brycon Müller & Troschel, 1844 (espèce-type par désignation de Jordan & Evermann: Brycon falcatus): STEINDACHNER (1876, discussion de Megalobrycon); MEEK & HILDEBRAND (1916, Panama); EIGENMANN (1922, Colombie et Equateur, avec la collaboration de Hildebrand); HILDEBRAND (1938, Panama); DAHL (1943, 1971, Colombie); CAMPOS (1950, collection du Departement de Zoologie de Sao Paulo); BÖHLKE (1958, Equateur); MYERS & WEITZMAN * F-24200 Sarlat, France. ** Muséum d'Histoire naturelle, CP 434, CH-1211 Genève 6, Suisse. 794 J. GERY & V. MAHNERT (1960, liste des espèces décrites depuis 1910); et GÉRY (1964, clé des espèces du «groupe “falcatus» et 1978, clé compilée des espèces). La systématique des quelque 40 ou 45 espèces valables (sur environ 60 décrites), et particulièrement des espèces guyano-amazoniennes et du système de la Plata (Paraguay- Parana), présentait donc encore beaucoup d'incertitudes. HowEs (op. cit.) a eu le mérite, non seulement de citer in extenso les références des noms publiés dans le genre Brycon au sens large, de revoir et de donner une description complémentaire de onze des types du British Museum et du Muséum national de Paris, mais aussi de grouper les espèces de manière assez satisfaisante, la biogéographie recoupant d'assez près l'anatomie. Grâce à ce travail, les characologistes disposent d'une base leur permettant d'entreprendre la révision de tel ou tel groupe intéressant ou mal connu. Dans ce travail, nous discuterons certains Brycon amazoniens et paraguayens, en terminant par des remarques sur Brycon opalinus et les espèces de l'est brésilien. Nous employons les méthodes de comptage et de mesures de Howes (op. cit.), avec deux exceptions: les écailles longitudinales sont comptées jusqu'à la fin de la ligne latérale (y compris celles qui s'étendent sur la caudale, généralement au nombre de 5 ou 6), et les écailles transversales jusqu'à la base de la ventrale et non jusqu'à la ligne ventrale médiane. (1) NOTE SUR CERTAINS Brycon DE L'AMAZONIE CENTRALE Le groupe dont l'approche est la plus délicate paraît être celui des trois ou quatre Brycon habitant la cuvette centrale amazonienne. Il est important de pouvoir mettre un nom scientifique sur ces espèces, en raison de l'intérêt économique potentiel de l'une d'entre elles, le «Matrinchao». Cette forme est récoltée près de Manaus, où elle n'est pas rare; elle vit en général avec une espèce jumelle (ou considérée comme telle) appelée «Jatuarana», souvent confondue, ces deux espèces étant le plus souvent appelées Brycon melanopterus (cf GÉRY 1978, WERDER 1983, WERDER & SOARES 1984). Le Matrinchao, étudié depuis plus de dix ans à la Station de Pisciculture de l'Institut de Recherches . amazoniennes (INPA), s'est révélé être d'une croissance très rapide (taille «portion» atteinte en quelques mois, pour une taille maximale d'environ 40 cm). On peut donc espérer un excellent rendement commercial avec des techniques voisines de celles de la trutticulture, d'où l'intérêt de fixer son statut taxonomique et de permettre le choix de la forme à élever en toute connaissance de cause. Les spécimens de Brycon ayant servi à la présente étude, en dehors des types, sont déposés au Département d'Ichtyologie du Muséum d'Histoire naturelle de Genève. Nous remercions M. Ulrich Werder qui a attiré notre attention sur le sujet traité dans cette note, et qui a mis à notre disposition le matériel biologique nécessaire. (a) Caractères des Brycon récoltés dans l'Amazonie centrale: les deux principales espèces ont le patron de coloration caractéristique de B. melanopterus auct., à savoir une large bande noire oblique partant de la base de la nageoire anale pour se terminer sur le lobe supérieur de la caudale, une tache humérale et une série de lignes longitudinales le long des rangées d'écailles. Cette coloration commune, ainsi qu'un nombre très voisin d'écailles et de rayons aux nageoires, expliquent qu'elles aient été souvent confondues par les zoologistes, tandis que les pêcheurs locaux les distinguent parfaitement. Un examen attentif révèle en effet quelques différences: NOTES SUR BRYCON 795 Fic. 1. Patron de coloration (schématique) de Brycon spp. de l'Amazonie centrale: en haut le Jatuarana, en bas le Matrinchao La bande noire du Jatuarana commence bien en avant des ventrales, passe sous le pédicule caudal et est pratiquement continue avec la tache caudale; la base de l'anale est droite, le diamètre de la tache humérale correspond à celui de la pupille, les nageoires ne sont pas marquées (sauf l'anale et la caudale), et les lignes longitudinales sont formées par des points foncés situés au centre des écailles. /n vivo, il existe une petite tache rouge au sommet de l'opercule et les ventrales sont jaunes. Chez le Matrinchao, la bande noire commence au-dessus des premiers rayons de l'anale, et s'interrompt généralement au niveau du pédicule caudal, surtout chez les spécimens adultes; la base de l'anale est légèrement convexe, le diamètre de la tache humérale correspond à celui de l'œil, le menton, la base des nageoires dorsale et de 796 J. GERY & V. MAHNERT l'adipeuse, les pectorales et les ventrales sont marquées de noir (de même que l'anale et la _caudale), et les lignes longitudinales, en zigzag, sont formées par le bord foncé des écailles. /n vivo, la joue et l'opercule sont rouges, ainsi que la base des ventrales. Les différences anatomiques sont résumées dans le tableau suivant: Jatuarana Matrinchao L.S. maxi. ca 250 mm ca 400 mm Plus grande hauteur au niveau des ventrales entre P. et V. Squamae 13-14/66-69/6-7 13-14/65-67/8-9 Branchiospines 10-13/14-16 14-16/15-21 Pectorales 1, 11-13, courtes 1, 13-14, longues Canaux de la L.L. courbés vers le bas généralement (a age Egal) ou bifurqués trifurqués ler postorbitaire égal ou à peine plus égal ou à peine moins haut (de infraorbitaire) haut que la pupille que le diamètre oculaire Vertèbres précaudales 22-23 25 Vertèbres caudales 22-23 21 Supraneuralia 8-9 10 Le nombre des écailles au-dessous de la ligne latérale (jusqu'aux ventrales), le nombre des branchiospines, le nombre des vertèbres et des supraneuralia (sur radio- graphies), et la forme des canaux de la ligne latérale, semblent discriminants (à condition, pour les canaux, de comparer des individus de même taille, car le nombre de canaux semble augmenter avec l'âge chez les Brycon; dans le cas du Jatuarana, plus petit, la structure des canaux, plus simple, pourrait être qualifiée de néoténique). On note aussi une tête un peu plus longue chez le Matrinchao, ainsi qu'un espace interorbitaire moins convexe et croissant relativement plus vite que la longueur de la tête (allométrie majorante, fig. 2). En revanche, aucune différence significative n'a été notée pour les caractères souvent utilisés chez les Brycon, tels que le nombre des rayons de l'anale ou la forme et le nombre des dents (d'après la comparaison point par point de 12 ex., 6 de chaque espèce, sympatriques du Lac Janauca près de Manaus, aimablement confiés pour examen par M. Ulrich Werder, et de la mesure, pour étude des allométries, de 11 autres Jatuarana et de 21 autres Matrinchao). (b) Identification des espèces Par ordre de priorité historique!, les espèces suivantes ont été décrites du bassin amazonien, ainsi que des Guyanes et de l'Orénoque: Chalceus amazonicus Spix in Spix & Agassiz, 1829 («Amazone») Brycon falcatus Müller & Troschel, 1844 (Guyana; type du genre Brycon) Brycon pesu Müller & Troschel 1845 (Guyana; type du genre Holobrycon, rejeté par Howes, loc. cit.) Brycon carpophagus Valenciennes in Cuv. & Val., 1849 (Essequibo, Amazone) Brycon brevicauda Günther, 1864 (Rios Tocantins et Capim) Py ! L'espèce la plus ancienne, Chalceus opalinus Cuvier, traitée plus loin, n'appartient pas à la faune amazonienne. J. GERY & V. MAHNERT Matrinchäo et Jatuarana y= -3.246 + 0.533x R = 1.00 Interorbitaire y= - 1.165 + 0.460x R = 0.99 20 40 60 80 L. tête Fic. 2. 797 (ol Matrinchäo ® Jatuarana 100 Espace interorbitaire avec allométrie majorante comparée à la longueur de la tête chez le Matrinchao. Megalobrycon cephalus Günther, 1869 (Haute Amazonie; type du genre Megalobrycon, rejeté par Howes, loc. cit.) Brycon capito Cope, 1872 (Rio Ambyiacu, Haute Amazonie) Megalobrycon melanopterus Cope, 1872 (id.) Megalobrycon erythropterus Cope, 1872 (id.) Brycon longiceps Steindachner, 1879 (Venezuela) Brycon stolzmanni Steindachner, 1879 (Rio Maranon) Brycon stuebelii Steindachner, 1882 (Iquitos, Haute Amazonie) Brycon bicolor Pellegrin, 1909 (Orénoque) Brycon siebenthalae Eigenmann, 1912 (Aruka River, Guyana) Brycon coquenani Steindachner, 1917 (Rio Coquenan, Venezuela) Brycon pellegrini Holly, 1829 (Manaus) Brycon matrinchao Fowler, 1941 (R. Parnaiba ou Amazone) Brycon coxeyi Fowler, 1943 (Rio Pastazza, Haute Amazonie) D'après Howes (op. cit.), B. amazonicus serait un nomen dubium, B. capito un juvénile non identifiable, B. stuebelii un synonyme de B. falcatus, B. matrinchao un synonyme de B. brevicauda et B. pellegrini un synonyme de B. cephalus. En vue d'attribuer au Jatuarana et au Matrinchao un nom adéquat, il faut éliminer les espèces dont le patron de coloration est différent: c'est le cas de: 798 J. GERY & V. MAHNERT N NS NOTES SUR BRYCON 799 — B. falcatus et de B. brevicauda (voir plus loin), qui ont une marque caudale en forme de croissant couvrant de façon symétrique les deux lobes, et non seulement le lobe supérieur et la partie moyenne de la nageoire — B. pesu, très différent — B. carpophagus (Val.) (non CASTELNAU, 1855 pl. 34 fig. 3, qui est une autre espèce comme l'a montré HOWES (op. cit.), peut-être ce que les auteurs appellent B. orthotaenia, voir plus loin) et B. siebenthalae, tous deux de Guyana et apparemment synonymes: 1ls n'ont aucune marque caudale — et de B. erythropterus, stolzmanni, longiceps, coquenani et coxeyi, de Haute Amazonie et du Vénézuéla, qui tous ont une tache au milieu du pédicule caudal, s'étendant sur la nageoire chez B. erythropterus et coxeyi (B. erythropterus fait l'objet d'un commentaire dans ce travail — voir plus loin). Deux des trois espèces qui restent à la disposition du réviseur, en raison de leur patron de coloration, B. cephalus et B. bicolor, ont été examinées par HOWES (op. cit.), qui en a donné une bonne description complémentaire. B. bicolor, dont nous avons nous- méme revu les types, peut, au moins provisoirement, étre exclu de la discussion en raison de son patron de coloration (pas de tache humérale et pas de bande au-dessus de l'anale), et de la localité de capture (Orénoque). La troisième espèce, B. melanopterus (Cope, 1872), a été figurée et caractérisée par son descripteur, qui en a souligné deux traits typiques, a savoir la bande oblique continue sur le bord inférieur du pédicule et la tache rouge du sommet de l'opercule, ainsi que par le premier réviseur FOWLER (1907), qui donne 13 comme nombre des branchiospines inférieures: ces caractères sont ceux qui viennent d'être fournis pour le Jatuarana. Avant examen des types, on peut admettre que le nom scientifique le plus ancien de cette forme amazonienne est B. melanopterus (Cope, 1872). Comme il a été établi que les deux espèces du bassin central étaient différentes, il s'ensuit que le Matrinchao doit s'appeler Brycon cephalus (Günther, 1869). Ceci est corroboré par le patron de coloration décrit par HOWES (op. cit.), à savoir les lignes en zigzag entre les écailles et non a travers leur centre, la grande tache humérale, la bande noire anale ne s'étendant pas en avant de la nageoire, la coloration des nageoires, ainsi que par quelques caractéres méristiques (en dépit de la mauvaise condition des 2 syntypes). Compte tenu du fait que HOWES (op. cit.) a dénombré les écailles transversales jusqu'à la ligne médiane ventrale, et les écailles de la ligne latérale sans les pores situés sur la base de la caudale, on peut vérifier que les comptes d'un des types de B. cephalus (?13/ 58 /?12, ce qui correspond à 13/ 58+6 ou 8 /9) ne s'écartent pas sensiblement de ceux du Matrinchao. Quant à la différence dans le compte des rayons pectoraux (i, 12 au lieu de i, 13 ou 14), elle peut aussi s'expliquer par une différence d'appréciation dans le dénombrement des très petits rayons, souvent rudimentaires, du côté ventral de la nageoire. Ces quelques incertitudes ont pu être levées par G. Howes (in litt., 30.9.1982) qui a bien voulu nous confirmer l'hypothèse de l'attribution du Matrinchao à l'espèce B. cephalus: «It is my opinion, from a re-examination of the two syntypes and from your key, that all specimens listed on p. 18 of my catalogue are conspecific with those types. Fics 3-5. Brycon cephalus, Lago do Castanho (3); Brycon erythropterus, Iquitos (4); Brycon melanopterus, Alto Guapore a Vila Bela (5). 800 J. GERY & V. MAHNERT Furthermore, in their gill-rakers counts, colouration and other characters you list they .conform to what you term Matrinchao.» (c) Le raisonnement ci-dessus procède par Elimination et son caractère circulaire est patent, mais il semble inévitable. En désignant B. melanopterus et B. cephalus comme noms respectifs des deux Brycon communs de la cuvette amazonienne, il aboutit à la réfutation de la synonymie suggérée par HOWES (op. cit.): erythropterus = melanopterus = cephalus, hypothèse que l'un de nous avait lui-même émise (Géry, 1978) a propos de erythropterus seul. En fait, il existe bien 3 espèces. L'examen de spécimens provenant de Haute Amazonie et de Bolivie montre qu'une forme très proche de B. cephalus possède plus d'écailles en ligne latérale: cette forme doit être appelée par le nom le plus ancien, B. erythropterus (Cope, 1872) (à nouveau par élimination). Comme pour les espèces précédentes, on peut relever des incertitudes concernant quelques caractères décrits par Cope (1872) (l'espèce n'a jamais été révisée, ni même vue par FOWLER 1907 à l'occasion de la révision des espèces de Cope): les individus du Maranon et de l'Ucayali, observés par nous, sont plus hauts (hauteur 2,75-3,45 dans la L.S. au lieu de 4), avec la tête plus longue (3,10-3,70 dans la L.S., au lieu de 4). La formule des écailles transversales (13-15 / 8-10) ne diffère guère de celle de B. cephalus, mais les écailles sont plus nombreuses en ligne latérale: 69-79 au total, alors que B. cephalus ne dépasse pas 68 ou 69 écailles; les pores de la ligne latérale ne sont pas bifurqués chez de petits spécimens de la localité typique, mais ils le sont chez de grands individus du bassin du Rio Madeira en Bolivie; le nombre des vertèbres précaudales (25-26) correspond à celui de B. cephalus, tandis que le nombre des supraneuralia (8) correspond à celui de B. melanopterus (tableau II); la coloration rappelle tout à fait celle d'un Salminus (une tache humérale ovale, des lignes en zigzag le long du corps, entre les rangées d'écailles, dont la netteté dépend probablement de la préservation et de l'àge de l'individu, et une tache caudale allongée horizontalement, rigoureusement symétrique et se continuant sur les rayons médians); les grands individus de Bolivie mentionnés plus haut ont les nageoires paires, ainsi que les derniers rayons de l'anale, noirs ou noirätres. Les caractères différentiels des trois espèces apparaissent dans la clé qui termine cette note. D'après les récoltes à notre disposition, il semblerait que B. cephalus ne soit pas présent à Iquitos, mais dans l'Ucayali, et que B. erythropterus n'existàt pas à Manaus, les deux espèces pouvant étre vicariantes. Dans cette hypothèse (fort fragile), la localité typique de 8. cephalus («Upper Amazon») devrait être restreinte a l'Ucayali. (d) Une quatrième espèce encore non nommée, dont le patron de coloration s'approche singulièrement de celui de B. cephalus et B. melanopterus, a été récoltée dans le Rio Aripuana, près de l'Ilha do Castanhal, par l'équipe ichtyologique de l'INPA, il y a une douzaine d'années. Bien que, par plusieurs caractères méristiques et la présence de points au centre des écailles, elle puisse être attribuée à B. brevicauda, il pourrait s'agir d'une forme nouvelle en raison de la tache caudale ne prenant que le lobe supérieur (la tache est en croissant chez B. brevicauda). Les 5 petits (101-126 mm LS) exemplaires conservés à l'INPA sont décrits ci-après, en vue d'une éventuelle et future description officielle de cette intéressante forme (un sixième individu provenant du Lago Amana — latéral du Rio Japura —, récolté en 1980, pourrait aussi appartenir à cette espèce). Nous laissons le soin de nommer cette espèce, s'ils le jugent opportun, à nos collègues de Manaus, qui ont le matériel à leur disposition. Brycon sp., 5 ex., 101-126 mm LS (Collections de l'INPA, Manaus): Hauteur (au niveau de la dorsale) 2,60-2,75 et tête (sans membrane) 3,50-3,75 dans la LS; dorsale NOTES SUR BRYCON 801 sensiblement au milieu du corps, un peu en avant chez les plus grands, très légèrement en arrière chez les plus petits; œil 3,0-3,45 (allométrie minorante), espace interorbitaire 2,40- 2,75 (allométrie majorante), maxillaire 2,85-3,05, museau, plutôt pointu, 3,50-3,65 et partie post-oculaire (horizontalement jusqu'à l'opercule) 2,45-2,65, dans la longueur de la tête; pédicule caudal 1,1 à 1,4 fois plus long que haut. Squamae 51-57 (en comptant tous les pores, bi- ou trifurqués, de la ligne latérale, laquelle est basse sur le pédicule), 9 ou 10/5 1/2 ou 6 en une rangée transversale entre dorsale et ventrale, 17-18 autour du pédicule et 18-20 en série prédorsale; 11,9 rayons à la dorsale, i11,22(1) à 24(1) rayons à l'anale, dont la base est droite, et 1,12-13 rayons à la pectorale, qui est relativement longue, mais n'atteint pas tout à fait l'origine de la ventrale. Dents au nombre de 10 ou 11 à la rangée externe du prémaxillaire, 24-26 au maxillaire, 11 à la rangée mandibulaire externe (4 fortes dents pentacuspidées et 7 plus petites) et environ 12 à la rangée postérieure et interne, plus une dent conique en arrière de la symphyse, bien développée (de chaque côté des mächoires); premier postorbitaire (infraorbitaire 4) de hauteur égale ou supérieure à 1/2 diamètre oculaire; branchiospines au nombre de 14 / 15-17. Patron de coloration voisin de celui de B. melanopterus: une tache humérale ronde ou ovale d'environ 1/2 à 2/3 du diamètre oculaire; une large tache pédiculaire symétrique se continuant sur le lobe supérieur de la caudale, le lobe inférieur seulement grisâtre; une bande noire sur la partie inférieure de l'abdomen et le long de la base de l'anale, à partir de la racine de la ventrale, ne s'étendant pas sur la partie inférieure du pédicule caudal; une série de points au centre des écailles formant des lignes longitudinales; base des rayons anaux, ainsi que l'extrémité du lobe anal, noires, dorsale et adipeuse incolores (la dorsale avec un fin liseré distal), pectorales incolores, pointe de la ventrale noirâtre. TABLEAU I. Principaux caractères de deux espèces, Brycon sp. (Rio Aripuana) et B. bicolor (2 des types) Brycon sp. B. bicolor (Aripuana) (Orénoque) Nb. d'ex. 5 ex. 2 ex. L.S. (mm) 101-126 112-122 Hauteur 2,60-2,75 2.60 Téte 3.50-3,75 3,45-3,55 Dorsale env. milieu env. milieu Oeil 3,0-3,45 3,45-3,60 Espace interorb. 2,40-2,75 2,30-2,60 Maxillaire 2,85-3,05 2,85-2,90 Museau 3,50-3,65 3,45-3,70 Pédicule caudal 1,1-1,4 1,0-1,3 Squamae long. 51-57 62-66 Sq. tr. 9 ou 10/5 1/2 ou 6 14/6-7 Sq. pédic. 17-18 18-20 Sq. prédors. 18-20 - Anale 111,22 - 24 111,24 - 25 Pectorale 1,12-13 1,13-14 Dents pmx. ext. 10 ou 11 7-8 maxillaires 24-26 26-29 mdb. int. 12? 22-23 Branchiospines 14/15-17 13-15 / 15-16 Tache humérale 1/2 à 2/3 de l'œil = pupille 802 J. GERY & V. MAHNERT Par rapport à B. brevicauda Günther, 1864, dont l'un de nous (J.G.) avait donné en - 1964 une description complémentaire, et dont les types ont été révisés par HOWES (op. cit.), cette espèce diffère non seulement par la coloration mentionnée plus haut, mais aussi par certains caractères: hauteur plus grande, maxillaire plus court mais avec plus de dents, écailles et branchiospines un peu moins nombreuses. Par rapport à B. bicolor (également révisé par HOWES (op. cit.), et revu par nous), qui a la même coloration caudale que 2. melanopterus et B. cephalus, mais qui, selon Howes, en est distinct, les différences semblent se résumer à la dimension de la tache humérale et la présence d'une bande au- dessus de l'anale, et surtout à des écailles nettement moins nombreuses (9-10/51-57/5 1/2-6 au lieu de 14/62-66/6-7) et des dents mandibulaires internes moins nombreuses (environ 12 au lieu de 22-23). Il s'agit néanmoins, apparemment, d'une espèce assez proche de B. bicolor Pellegrin. Le tableau I donne les caractères des deux espèces (mesures prises par le même opérateur (J.G.). L'un des trois exemplaires types de B. bicolor, N° 87.746, est en mauvais état: 1l n'en a pas été tenu compte dans le tableau et dans la clé ci-après: (e) CLÉ DE DÉTERMINATION DES ESPECES CITÉES la. Une bande noire oblique allant de la base de la nageoire anale au lobe supérieur de la caudale, interrompue ou non sur le pédicule caudal; généralement pas plus de70lecailles perforees (total) en ligne latérale "Re EEE 2a. Sq. 9-10/51-57/51/2-6, 17-18 circumpédiculaires, 18-20 en série prédorsale; hauteur 2,60-2,75 dans la L.S.; 24-26 dents maxillaires (lignes longitudinales formées de points au centre des écailles; base de l'anale droite; pectorales 1,12-13, n'atteignant pas tout à fait les ventrales; branchiospines 14/15-17)................... EEE A Brycon sp. (aff. bicolor) 2b. Sq. 12-14/62-70/6-9 1/2, 18-23 circumpédiculaires et 23-26 en série prédorsale ......... 3a. Hauteur 2,60; 26-29 dents maxillaires (sq. 14/62-66/6-7; pectorales 1,13-14; tacherhumeraleregalesalapupille) nenn. ee Brycon bicolor (Pellegrin) 3b. Hauteur 2,95-3,30 en général; pas plus de 20-22 dents maxillaires ........................ 4a. Sq. tr. 6 ou 7 de la LL aux ventrales; pectorales i,11-13, courtes, n'atteignant pas les ventrales; canaux de la ligne latérale bifurqués; premier postorbitaire étroit, pas plus d'!/2 ceil; plus grande hauteur au niveau des ventrales; base de l'anale droite; lignes longitudinales formées de points au centre des écailles; tache humérale égale à la pupille, nageoires paires généralement incolores, bande anale commençant avant les ventrales et se continuant sous le pedicule; in vivo, une tache rouge à la en supérieure de l'opereuler.. ee ER .. Brycon melanopterus (Cope) 4b. Sq. tr & 91/2 de la übe aux - ventrales; ‘pectorales le) 14, longues, atteignant généralement les ventrales; canaux de la LL trifurqués; premier postorbitaire développé, sa hauteur presque aussi grande que le diamètre oculaire; plus grande hauteur entre pectorales et ventrales; base de l'anale convexe; lignes longitudinales formées de dessins en zigzag entre les écailles; tache humérale presque aussi grande que l'œil; nageoires paires généralement noiratres; bande anale commençant un peu avant la nageoire et ne s'étendant pas sous le pédicule; in vivo, ri postérieure de la tête rouge (surtout la joue) .............. ee .. Brycon cephalus (Günther) lb. Une bande noire au- -dessus de la base de l'anale et une tache pédiculaire médiane se continuant sur les rayons caudaux médians; 69-79 écailles perforées en ligne latérale (sq. tr. 13-15/8-10, 22-24 circumpédiculaires, 24-26 prédor- NOTES SUR BRYCON 803 sales; hauteur 2,75-3,45 — parfois moins ? — dans la L.S.; 15-18 dents maxillaires; pectorales i,13-14, n'atteignant pas les ventrales; canaux de la ligne latérale bi- ou trifurques; branchiospines 13-15/16-18; lignes longitudinales formées de dessins en zigzag entre les écailles; tache humérale presque aussi grande que Iceilsnageoiresspaires ioncees nn EE ck ER CEE Brycon erythropterus (Cope) (2) NOTE SUR LES Brycon DU PARAGUAY. Les différentes expéditions du Muséum de Genève au Paraguay (voir GERY et al., 1987, pour la liste des stations), ont permis de récolter 2 espèces de Brycon. L'une, rare au Paraguay mais, semble-t-il, commune dans le Rio de la Plata, est conforme à Brycon orbignyanus (Valenciennes, 1849) dont nous donnons plus loin la synonymie et la description complémentaire (surtout sous forme de tableau). La seconde espèce récoltée, à écailles petites et nombreuses, a été citée de façon confuse, soit comme Brycon microlepis Perugia, soit comme Brycon hilarii (Valen- ciennes), parfois comme deux espèces, ainsi que l'indiquent les listes bibliographiques suivantes (qui ne comprennent la plupart du temps que des citations sans examen des spécimens). Il s'agit, d'après l'examen des types respectifs, d'une seule espèce, et incontestablement de B. microlepis. B. hilarii semble ne pas exister au Paraguay (une description complémentaire commentée de cette espèce figure dans la note (3). a) Brycon microlepis Perugia Brycon microlepis Perugia, Annali Mus. civ. Stor. nat. Genova, 2(18): 149, 1897 (Alto Paraguay); BOULENGER, id., 2(19): 127, 1898 (id.); EIGENMANN, MCATEE & WARD, Ann. Carnegie Mus., 4: 153, 1907 (cit.); EIGENMANN, Rep. Princeton Univ. Exp. Patagonia, 3: 431, 1910 (cit.); BERTONI, Cat. sist. Vert. Paraguay: 11, 1914 (cit.); PEARSON, Proc. Calif. Acad. Sci., ser. 4, 23: 109, 1937 (cit. Paraguay); BERTONI, Revta Soc. cient. Paraguay 4 (4)): 55, 1939 (cit.); FOWLER, Arg. Zool. S. Paulo, 6 (2): 338, 1950 (cit.); GERY, Characoids of the World: 338, 1978 (clé); Howes, Bull. Br. Mus., 43 (1): 34 et 36, 1982 (cit.). Chalceus hilarii, non Valenciennes, KNER, Denkschr. k. Akad. Wiss. Wien, 18: 10-11, 1860 (Rio Cujaba). Brycon hilarii, non Valenciennes, BOULENGER, Boll. Mus. Zool. Anat. comp. Torino, 15 (370): 3, 1900 (Corumba); EIGENMANN & OGLE, Proc. U.S. natl. Mus., 33: 30, 1907 (Paraguay); EIGENMANN, MACATEE & WARD, Ann. Carnegie Mus., 4 (7): 153 (cit.); EIGENMANN, Rep. Princeton Univ. Exp. Patagonia, 3(4): 55, 1911 (cit.); BERTONI, Cat. sist. Vert. Paraguay: 11, 1914 (cit.); FOWLER, Proc. Acad. nat. Sci. Philad. 84: 357, fig. p. 346, 1933 (Descalvados, Mato Grosso); PEARSON, Proc. Calif. Acad. Sci., (4) 23: 109, 1937 (cit. Paraguay); BERTONI, Revta Soc. cient. Paraguay, 4 (4): 55, 1939 (cit.); CAMPOS, Pap. avuls. Dept. Zool..S. Paulo, 9 (10): 140, 1950 (Mato Grosso p.p.); FOWLER, Arg. Zool. S. Paulo, 6 (2): 336-337, 1950 (cit.). Brycon hilarlii (sic), non Valenciennes, EIGENMANN & KENNEDY, Proc. Acad. nat. Sci. Philad., 55: 523-524, 1903 (Arroyo Trementina). 15 exemplaires ont été récoltés: Bassin du Rio Paraguay: — 1 ex., MHNG 2053.97, 250 mm L.S., Arroyo Tagatya-mi, est de Puerto Max, coll. 20- 22/10/1979 — 3 ex., MHNG 2156.45 et 46, 216-270 mm L.S., Concepcion, gué de l'Arroyo Tagatya-mi, coll. C. Weber et C. Dlouhy, 10.10.1983 — 2ex., MHNG, 2239.26, 255-280 mm L.S., Gué de l'Arroyo Tagatya-guazu, coll. 21-22.10.85 804 J. GERY & V. MAHNERT — 1 ex., MHNG 2239.27, 177 mm L.S., Concepcion: Laguna Negra, 15 km E. de Paso Bareto, - coll. 18.10.1985 - 3 ex., MHNG 2396.26, 155-195 mm L.S., Concepcion, Estancia Primavera, coll. 31.10.1987 = 1 ex., MHNG 2053.96, 210 mm L.S., Concepcion: Estancia Estrella, Rio Apa, coll. 16.10.79 = 2 juv. (1 coloré a l'alizarine), MHNG 2358.58, L.S. max. 97 mm, coll. H. Bleher, 3.1987 Bassin du Rio Parana: 2 ex., 310-343 mm L.S., en face de Puerto Iguazu, coll. C. Dlouhy 1.2.1987 (# 100188 et 100304) Et 3 ex. extra-territoriaux (bassin du haut Paraguay): 2 ex., 146-163 mm L.S., Lago Sinha Mariana, 30 km en aval de Barao de Melgaco, Rio Cuiaba, coll. Géry & al., 30/11/1979 1 ex., env. 150 mm L.S., Barao de Malgaco, Rio Cuiaba, coll. K. Silimon, Mus. Cuiaba, 1978. 6 FIGs 6-7. Brycon microlepis Perugia, holotype (MSNG 36916), Bahia Negra. NOTES SUR BRYCON 805 Description des exemplaires paraguayens: L.S. maximale 343 mm. Hauteur du corps 2,8-3,3, longueur de la tete 3,7-4,5, distance prédorsale 1,9-2,0 et distance préventrale 2,0-2,2 dans la L.S. Diamètre oculaire 3,6-4,3, espace interorbitaire 2,1-2,5 et longueur du maxillaire 3,0-3,3 dans la longueur de la tête. D.i1,9, A.iv,23-26, les rayons garnis de crochets chez les mâles adultes, P.1,13-14 et V.i,7; 74-82 écailles perforées (total) en ligne latérale, laquelle se continue sur le rayon caudal médian, 15-17 / 8-9 (jusqu'aux ventrales) transversales, 26-28 autour du pédicule caudal. Canaux de la ligne latérale courbés vers le bas ou bifurqués chez les spécimens de 95-150 mm L.S., à 3 ou 4 branches (190 mm L.S.), 5 ou 6 branches (210-265 mm L.S.) et jusqu'à 10 branches chez le spécimen de 343 mm L.S., la complexité des canaux étant en rapport avec l'âge. Dents prémaxillaires 7-10 / 2-3 / 7-9, maxillaires 14-15, mandibulaires environ 17, dents symphysaires internes présentes. Hauteur du 4e interorbitaire plus faible que le diamètre oculaire, plus petit (95 mm L.S.) à plus grand (250 mm L.S.) que le diamètre pupillaire. Branchiospines au nombre de 14-16 en bas du premier arc branchial. Vertèbres au nombre de 45-46, dont 24-25 précaudales et 21-22 caudales. Une tache ronde humérale, plus grande que la pupille; une bande longitudinale variable, commençant au-dessus du tiers postérieur de l'anale, ou au début du pédicule, et se continuant sur la nageoire caudale jusqu'à son extrémité ou presque. Le lobe caudal médian est prolongé en pointe chez les individus de taille inférieure à 200 mm L.S., les lobes caudaux sont rouges (au moins chez l'individu du Rio Apa). Comparaison du type de B. microlepis avec les récoltes récentes: Type de Spécimens B. microlepis du Paraguay RS: 116 mm jusqu'à 340 mm Haut. 3,4 2,8-3,2 Tete 39 3,7-4,5 Prédors. 1,8 1,9-2,0 Préventr. 2,0 2,0-2,2 Oeil 3,4 3,6-4,3 Esp. I.-O. 2,4 DAS Maxill. 2,85 3,0-3,3 A. 111,25 iv,23-26 Sq. long 80 74-82 Sq. transv. 16/9 15-17/8-9 Sq. pédic. 28 26-28 Pmx. 10/4/7 7-10/2-3/7-9 Mx. 19 14-15 Mdb. 9+? 17 Brsp. 7 14-16 Les caractères méristiques relevés sur ces spécimens récents correspondent à ceux de Brycon microlepis Perugia, 1897, de Bahia Negra, Chaco Boreale, Alto Paraguay, comme le montrent la description complémentaire suivante du holotype et la comparaison des deux échantillons, tandis que les discordances dans certaines proportions peuvent s'expliquer par les allométries, l'holotype de B. microlepis étant un individu juvénile: 806 J. GERY & V. MAHNERT Holotype de Brycon microlepis Perugia, N° MSNG 36916, Bahia Negra, L.S. 116 - mm; hauteur 3,4, tête 3,3, distance prédorsale 1,8, et distance préventrale 2,0 dans la L.S. Diamètre oculaire 3,4, espace interorbitaire 2,4, et maxillaire 2,85 dans la longueur de la tête. D.11,9; A.111,25; P.1,12, longues, atteignant les ventrales. Squamae 16 / 80 (75 + 5) /9, 28 circumpédiculaires; Canaux de la ligne latérale courbés vers le bas ou bifurqués. Dents prémaxillaires 10 / 3 / 7, maxillaires 19, mandibulaires 4 (6-7 cuspides) + 5 triscuspidées à coniques; présence d'une paire de dents internes symphysaires. Branchiospines au nombre de 17 en bas du premier arc branchial. Vertèbres au nombre de 46, dont 25 précaudales et 21 caudales. Supraneuralia au nombre de 10. En conclusion, le nom correct des individus du Paraguay est Brycon microlepis Perugia, 1897, seule espèce (en dehors du Nord-Ouest de l'Amérique du Sud) à posséder jusqu'à 82 écailles (au total) en ligne latérale. En effet, l'examen des types de Brycon hilarii (Valenciennes) montre que B. microlepis a été mal identifié comme B. hilarii par les rares auteurs qui ont pu étudier des spécimens paraguayens (BOULENGER 1900, EIGENMANN & KENNEDY 1903 et FOWLER 1932). Ces auteurs ont probablement été trompés par le chiffre très exagéré («quatre- vingt») du nombre des écailles longitudinales donné par Valenciennes, ainsi qu'il ressort de la description du type de l'espèce et de sa description complémentaire. Nous donnons plus loin, à propos des Brycon du Sud-Est brésilien, les éléments qui nous permettent d'avancer cette hypothèse. (b) Brycon orbignyanus (Valenciennes, in CUVIER & VALENCIENNES, 1849) Chalceus orbignyanus Valenciennes, in CUVIER & VALENCIENNES, Hist. nat. Poiss., 22: 249, 1849 (La Plata); ? KNER, Denks. Akad. Wiss. Wien, 18: 11, 1860 (Rio Guaporé); GÜNTHER, Cat. Fishes B.M., 5: 333, 1864 (cit) Brycon orbignyanus, EIGENMANN & EIGENMANN, Proc. U.S. natl Mus., 14: 55, 1891 (cit.); BERG, Anales Mus. nac. Buenos Aires, 4 (2a)1:123-125 (Rio Parana, Rio de la Plata); BOULENGER, Boll. Mus. Zool. Anat. comp. Univ. Torino, 12 (279):4, 1897 (Bolivia); EIGENMANN, Cat. Fishes S. Am.: 431, 1910 (cit.); BERTONI, Fauna Paraguaya, Peces: 11, 1914 (cit. Paraguay); DEVINCENZI, An. Mus. nac. Montev. (2), 1(5):174, 1924 (Rio Uruguay); MARELLI, Mem. Mrio O. Publ. 1922-1923: 559, 1924 (Prov. Buenos Aires, Rio Parana, Rio de la Plata); DEVINCENZI & BARATTINI, Album ictiologico del Uruguay, 2a Ser., Pl.13 fig. 2, 1928 (Uruguay); BERTONI, Revta Soc. cient. Paraguay, 4 (4): 55, 1939 (cit. Paraguay); RINGUELET, Not. Mus. La Plata, 5, Zool. (34): 105, 1940 (Rosario); DEVINCENZI, in DEVINCENZI & TEAGUE, An. Mus. Hist. nat. Montev. (2), 5(4):73, 1942 (Uruguay, fig.); Pozzi, GAEA, 7(2): 254, 1945 (Rio de la Plata, Rio Parana, Rio Paraguay); THORMALEN DE Gil, Revta Mus. La Plata (N.S.), 5 (Zool.): 351-440, 1949 (monographie); DE BUEN, Publ. Client. S.O.Y.P. (2):83, 1950 (cit. ?); FOWLER, Arg. Zool. S. Paulo 6 (2): 337, 1950 (cit.); CORDINI, Mrio. Agric. y Ganad., Publ. misc., 410: 32, 37, 1955 (Rio Parana); RINGUELET & ARAMBURU, Mrio Asuntos Agrarios Bs. As. N° 119: 13, 1957 (Parana-Plata); MARTINEZ ACHENBACH & BONETTO, An. Mus. Prov. C. Nat. Fno. Ameghini, 1(2): 10, 1957 (Rio Parana medio); GNERI & NANI, Suma geogr., Peuser, 5: 248, 1960 (fig.); RINGUELET & ARAMBURU, Agro, Ano 3 (7): 31, 1962 (cit.); MARINI & Lopez, Eval. Rec. nat., 7: 81, 1963 (cit.); RINGUELET, ARAMBURU & ARAMBURU, Los Peces argentinos de agua dulce: 135-137, 1967 (Rio de la Plata, Rio Parana); GERY, Characoids of the World: 339, 1978 (clé); PANATTIERI & BARCO, CYTA, Mrio. Agric. Ganad. Santa Fe, 16: 20-22, 1980 (Sta Fe); BONETTO, CANON VERON & ROLDAN, Ecosur 8 (6): 29-40, 1981 (Rio Parana); HOWES, Bull. Br. Mus., 43(1): 38, 1982 (cit.); PANATTIERI & DEL BARCO, CYTA, Mrio. Agric. Ganad. Santa Fe, 29: 32-33, 1982 (Sta. Fe); OLDANO & TABLADO, Stud. neotrop. Fauna Environm. 20 (1): 49-58, 1985 (Rio Parana medio). Chalceus rodopterus Valenciennes, in CUVIER & VALENCIENNES, Hist. nat. Poiss., 22: 249, 1849 (Buenos Aires); LAHILLE, Revta Mus. La Plata, 6: 269, 1895 (Rio Santiago). Brycon rodopterus, EIGENMANN & EIGENMANN, Proc. U.S. natl Mus., 14: 55, 1891 (cit.). NOTES SUR BRYCON 807 Brycon lineatus Steindachner, Sber. Akad. Wiss. Wien, 53: 211, pl. 2, 1866 (La Plata); EIGENMANN & EIGENMANN, Proc. U.S. natl. Mus., 14: 55, 1891 (cit.); EIGENMANN, Cat. Fishes S. Am.: 431, 1910 (cit.); POZZI, Gaea, 7(2): 254, 1945 (Rio de la Plata, Rio Parana, Rio Paraguay); FOWLER, Arg. Zool. S. Paulo 6 (2): 337, 1950 (cit.); RINGUELET & ARAMBURU, Agro, Ano 3 (7): 31, 1962 (cit.); MARINI & Lopez, Eval. Rec. nat., 7: 81, 1963 (cit.). Brycon orthotaenia (non Giinther, 1864), GUNTHER, Ann. Mag. nat. Hist., London, (5) 6: 13, 1880 (Rio de la Plata). A propos de cette espece, HOWES (op. cit.: 39) ne donne pas de liste synonymique parce que: «... until the types of all these species (orthotaenia, rodopterus, lineatus) have been compared any such compilation will be virtually useless». Ayant pu étudier les types en question, nous avons proposé cette synonymie (sans tenir compte de certaines orthographes simplificatrices), et confirmé que B. rodopterus (fondé sur des individus juvéniles) et B. lineatus (fondé sur un exemplaire tératologique ?) sont bien des synonymes de B. orbignyanus. D'après la liste des publications et nos récoltes, cette espèce ne remonterait pas dans le Rio Paraguay et s'arrêterait au Parana moyen. Les citations suivantes de Brycon orbignyanus concerneraient d'autres espèces: CAMPOS, Pap. avuls. Depto Zool., S.P., 9 (10): 139, 1950 (rio Aguapei, rio Piracicaba, rio Mogi-Guagu); Howes, Bull. Brit. Mus. (N.H.), zool. Ser., 43 (1): 38, 1982 (rio das Velhas). 4 exemplaires ont été récoltés (Lac Itaipu, bassin du Rio Parana): — 2 ex., MHNG 2476.96 et 97, 105-152 mm L.S., Canendiyu, lac Itaipu à la hauteur du Salto Guiara, coll. C. Dlouhy, 20.6 et 18.7.1989 — 1 ex., MHNG 2152.56, 198 mm L.S., Canendiyu, lac Itaipu à la hauteur de l'arroyo Pyra-Pita, coll. C. Dlouhy — lex. en face de Puerto Iguazu, coll. 1.2.1987. B. orbignyanus, espece bien connue surtout depuis la monographie de THORMALEN de GIL (1949), est caractérisé de la façon suivante: hauteur 3,0-3,5 et tête 3,1 (juvéniles) à plus de 4 fois dans la longueur standard (tête particulièrement courte pour un Brycon); dorsale insérée un peu en arrière du milieu du corps; ceil 3,5 (juv.)-4,4, espace interorbitaire 2,3 (adulte)-2,9, et maxillaire 2,9-3,2 dans la longueur de la tête; squamae 56-63 en ligne latérale (totale), 11-12/6-7 jusqu'aux ventrales en série transversale, et 20-22 autour du pédicule caudal; 111,25-26 rayons à la nageoire anale, 1,13-14 aux nageoires pectorales; une vingtaine de dents maxillaires; 17-18 branchiospines sur l'arc branchial inférieur; 28 vertèbres précaudales et 20-21 vertèbres caudales, 11-12 supraneuralia. Le nombre élevé des vertèbres précaudales et des supraneuralia nous paraît être caractéristique de l'espèce (voir aussi plus loin à propos de B. lineatus et tableau M!. La coloration après fixation ressemble à celle de B. erythropterus, d'Amazonie supérieure, qui a plus d'écailles et moins de vertèbres et de supraneuralia (tableau II). Nous donnons sous forme de tableau hors-texte (tableau III) les caractères relevés sur les types et sur deux exemplaires du Rio Parana récoltés dans le lac Itaipu et en face de Puerto Iguazu. (Le type de B. lineatus (NMW 62943, rio de la Plata), examiné à notre demande par H. Ahnelt et C. Weber, semble ne différer en rien de B. orbignyanus, sauf par le moindre nombre des vertèbres précaudales (21) et des supraneuralia (8) (tableau Il): il s'agit apparemment d'un spécimen tératologique). Le lectotype de Brycon orbignyanus ! D'une manière générale, le nombre des vertèbres et des supraneuralia nous paraît être un élément supplémentaire en vue d'une meilleure compréhension des affinités des Brycon spp. Le recours systématique à la radiographie, quand elle est réalisable et sur des individus préservés dans leur intégralité, semble particulièrement prometteur dans ce groupe, d'autant plus que la dissection est impossible sur les spécimens-types. 808 J. GÉRY & V. MAHNERT TABLEAU II. “ Comptes des vertèbres et des supraneuralia (SUPNEUR) de certains Brycon (PRECAUD= vertèbres précaudales; CAUD=vertebres caudales; TOT=total des vertebres). ESPECES PRECAUD CAUD TOT SUPNEUR B. cephalus Cano Yarina (1 ex.) 25 21 46 Lago Janauaca (2 ex.) 25 21-22 46-47 10 Lago Castanha (1 ex.) 25 21 46 B. melanopterus Iquitos (3 ex.) 22-23 22-23 44-45 8-9 Manacapuru (2 ex.) 22-23 22-23 45 Guapore (1 ex.) 22 22 44 B. erythropterus Iquitos (3 ex.) 25-26 21-22 46-48 8 B. orbignyanus Syntypes (2) 28 20-21 48-49 Paraguay (1 ex.) 28 20 48 11-12 B. rodopterus Syntypes (2) 28-29 20-21 49 B. lineatus Holotype 21 20 41 8 B. travassosi Holotype 28 21 49 12 B. microlepis Holotype 25 21 46 Paraguay (6 ex.) 24-25 21-22 45-46 10 TABLEAU III. Principaux caractères des types de B. orbignyanus, B. rodopterus, B. travassosi, et d'exemplaires paraguayens de B. orbignyanus. Caractères B. orbignyanus B. rodopterus B. travassosi Paraguay 2 syntypes 2 syntypes holotype 2 specimens (La Plata) (B. Aires) (R. Bodoquena) (Parana) L.S. 144-161 mm 78-81 mm 212 mm 190-355 mm L.S./Haut. 3,45-3,50 ca. 3,65 2,8 3,0-3,1 L.S./Tête 3,9-4,0 3,0-3,1 4,2 3,8-4,1 L.S./Pred. 9 1,8 1,8 1,8-2,0 Tête/Oeil 3,85 3,50-3,70 3,8 4,2-4,4 Tête/IntOr. 2,55-2,60 2,80-2,90 2,45 2,30 Tête/Max. 39, 29 3,18 39. Anale 111,25-26 111,25-26 1v,22 111,24-26 Pectorales 1112) 1,14 1,13 1,13-14 Squamae 12/62 12/56 12/59 11-12/61-63 Sq. pedic. 2 2 20 DI Dents max. 20 14-16 17 16-17 Brsp. inf. 18 117 19 19 Vertèbres 28+20+21 28-29+20-21 28+21 28+20 Supneuralia — - 12 11-12 NOTES SUR BRYCON 809 Fics 8-10. Brycon orbignyanus (Val.), sp&cimens types (MNHN A 9835), La Plata. 810 J. GERY & V. MAHNERT Fics 11-13. Brycon rodopterus (Val.), syntypes (MNHN A 9834), Buenos Aires. NOTES SUR BRYCON 811 (Valenciennes) est ici désigné: il s'agit du plus grand exemplaire, portant le N° MNHN 9835, d'environ 161 mm L.S. Le second exemplaire (même N°, environ 144 mm L.S.) est le paralectotype. Les deux Brycon du Paraguay se distinguent aisément par le nombre des écailles: la. Sq. 15-17/74-82 (tot.) / 8-9 (jusqu'aux ventrales), 26-28 autour du pédicule caudal Re IN EP AR feines Sa eas B. microlepis lb. Sq. 11-12/56-63 (tot.) / 7 (jusqu'aux ventrales), 22 autour du pédicule caudal ...... re IT re ER N RE muets ence B. orbignyanus (c) A propos de Brycon travassosi Campos, 1950 Brycon travassosi Campos, Pap. avuls. Zool. Sao Paulo, 9 (10): 141-142, 1950 (Rio Bodoquena, Pantanal, Mato Grosso do Sul); MYERS & WEITZMAN, Stanford Ichthyol. Bull., 7 (4):103 (cit.); BRITSKI, Pap. avuls. Zool. Sao Paulo, 22 (19): 202, 1969 (rev. type); GERY, Characoids of the World: 342, 1978 (clé); Howes, Bull. Br. Mus., 43(1): 45, 1982 («Possibly a synonym of B. orbignyanus»). Il semble pertinent de discuter le statut taxonomique de Brycon travassosi Campos, 1950, du Pantanal du Mato Grosso (bassin du Rio Paraguay), espèce dont nous avons pu examiner l'holotype et unique spécimen connu, grace à H.A. Britski: Holotype en alcool, 212 mm de longueur standard, Rio Bodoquena, Estado do Mato Grosso, Brasil; Lauro Travassos Filho col. 1941, N° MZUSP 3811 (exemplaire en assez mauvais état, cavité branchiale ouverte, abdomen éviscéré). Le nombre des vertèbres, des supraneutralia et les structures dentaires de cette espèce (voir tableaux II et III) sont en faveur de son appartenance au «type B. orbignyanus» («groupe B. orbignyanus» partim, de HOWES op. cit.:46), qui semble bien caractérisé par les vertèbres précaudales au nombre de 28-29, et les dents larges, jusqu'à 5 cuspides). L'habitus, le nombre des écailles, et la plupart des proportions, ne sont pas très différents de ceux de B. orbignyanus. Toutefois le moindre nombre de rayons anaux (iv, 22 au lieu de iii ou iv, 25-26), ainsi que la hauteur du corps, paraissent séparer les deux espèces, à un niveau taxonomique que le manque de matériel récent de B. travassosi ne nous permet pas d'apprécier. (3) NOTE ET REMARQUES SUR LES SPECIMENS-TYPES DES Brycon DU SUD-EST BRÉSILIEN Le souci d'identifier correctement nos spécimens paraguayens nous a amenés à examiner les types des régions voisines, et particulièrement les types du Muséum national d'Histoire naturelle (Paris): outre ceux déjà cités, Chalceus hilarii, C. orbignyanus et C. rodopterus, nous avons revu Chalceus opalinus Cuvier, 1819, Chalceus carpophaga non Valenciennes, CASTELNAU, 1855 (ainsi que Chalceus carpophaga Valenciennes, 1849), Chalceus devillei, et Brycon reinhardti (paratype); nous avons pu connaître les principaux caractères de B. lineatus (déjà cité) et du spécimen déterminé comme Brycon opalinus par KNER, 1860 et nommé Brycon nattereri par GÜNTHER, 1864, grâce à C. Weber et H. Ahnelt; Brycon bahiensis et Brycon orthotaenia ont été révisés par HOWES (op. cit.): leurs caractères sont bien connus; enfin BRITSKI (1969) a donné une description complémentaire de Brycon travassosi Campos, 1950. Seul Brycon lundii ne nous est pas familier, mais nous admettons volontiers avec HOWES (op. cit.) qu'il s'agit d'un synonyme de B. orthotaenia. Le résultat de notre examen suit, fondé uniquement sur l'étude de spécimens anciens: il ne s'agit donc pas d'une révision, mais de remarques taxonomiques et zoogéographiques, faute de matériel récent approprié. 812 J. GÉRY & V. MAHNERT Fics 14-15. Brycon travassosi Campos, holotype (MZUSP 3111), Rio Bodoquena, Estado do Mato Grosso. (a) Chalceus opalinus CUVIER, Mem. Mus. Hist. nat., 5, 1819, texte p. 351, fig. 1 pl. 26. C'est le premier Brycon (au sens moderne) décrit. Sa localité typique est difficile à préciser; en effet, le seul exemplaire cité par Cuvier aurait été envoyé par A. de St-Hilaire: NOTES SUR BRYCON 813 «Ce poisson est originaire des rivières du Brésil, d'où il a été envoyé... par M. Auguste de Saint-Hilaire.» Ce holotype incontestable est assez bien figuré, mais sa description est succincte (sq. 45; A.20; P.14; tr. 12 ou 13 au-dessus de la ligne latérale d'après la figure) et sans mention de la taille, indiquant seulement qu'elle est à peu près celle du Chalcée à grandes écailles (Chalceus macrolepidotus Cuvier, 1818). L'holotype de cette dernière espèce, donné comme étant long de 7 pouces (env. 180 mm) sur 21/4 de haut (env. 57 mm), mesure en réalité env. 250 mm de longueur standard (un peu moins de 10 pouces) sur env. 60 mm de haut (moins de 2 pouces 1/2). Dans l'Histoire naturelle des Poissons, VALENCIENNES (1849 pp. 244-246) écrit: «... j'ai sous les yeux les exemplaires qui ont servi à l'établissement (de Chalceus opalinus)...» et il donne dans sa description «A.28» alors que Cuvier n'avait compté que 20 rayons anaux. L'exemplaire cité en premier est manifestement celui qui a servi à Valenciennes pour sa description complémentaire: «Nous en possédons un exemplaire long de dix-sept pouces (env. 435 mm) originaire du Rio Tiquilenhonha (= Jequitinhonha); il a été donné par M. A. de Saint-Hilaire, en août 1822. Un autre exemplaire a été rapporté de Rio Janeiro par M. de Lalande». Mais c'est ce dernier examplaire qui est mentionné comme holotype par BERTIN (op. cit.): «1 ex., N° A.8613, Holotype — Rio de Janeiro, Delalande. 320 mm, sec sur verre, bon état». Bertin se fonde sur l'opinion de Eigenmann, cité par lui p. 18, qui pense que les localités des types respectifs de Chalceus fasciatus («Rio de Janeiro, Brésil — P.A. Delalande») et de Chalceus opalinus («Rio San Francisco, Brésil — A. Saint-Hilaire») ont été inversées. Le Poisson étudié par Cuvier, et par conséquent l'unique type, est bien l'exemplaire désigné par Bertin, et ceci pour plusieurs raisons (nous remercions vivement M. Laurent Lauzanne de l'aide qu'il a apportée à la solution de ce problème): L'exemplaire du Jequitinhonha est trop grand (17 pouces) pour être le type de Cuvier; de plus, il semble bien, comme Valenciennes le mentionne expressément, avoir été donné au Muséum après le mémoire de CUVIER (1819) (où ne figurent que deux espèces récoltées par le botaniste A. de Saint-Hilaire dans le Rio S. Francisco, soit Chalceus fasciatus (?) et le «Serrasalme» piraya, un nom latinisé ultérieurement). De plus, l'exemplaire N° A.8613 a bien l'anale «un peu altérée», il est à peu près de la même longueur que le type de Chalceus macrolepidotus et il correspond bien à la figure de CUVIER (1819, PI. 26 fig. 1). De toutes façons, la discussion est assez académique car les deux spécimens du Muséum, holotype N° A.8613, L.S. 263 mm, «Rio de Janeiro» et N° A.8612, L.S. 355 mm, «Rio Jequitinhonha» paraissent bien appartenir à la même espèce. Description complémentaire de Chalceus opalinus Cuvier, 1819 (les valeurs du N° A.8612, indiquées entre parenthèses): L'holotype de B. opalinus (A.8613) est un spécimen sec, empaillé et verni, monté sur plaque de verre, d'environ 310 mm de longueur totale et 263 mm de L.S. (355 mm de L.S., spécimen également sec). Le pédicule caudal est assez abîmé, ainsi que la nageoire anale, où l'on compte toutefois nettement plus de rayons que Cuvier, probablement 111,23 (intacte, probablement 111,24); squamae 47 (47 ?) a droite, 49 (45) à gauche, 7/4 (7/4-5) transversales jusqu'aux ventrales, les canaux de la ligne latérale seulement bifurqués, 14? (16?) écailles prédorsales et 14? (15?) écailles circumpédiculaires (ces deux comptes trés incertains); pectorales i,14 (i,13), la longueur de la nageoire 65% (?) de la distance 814 J. GERY & V. MAHNERT pectorale-ventrale; la hauteur du corps (proportion peu fiable chez les spécimens secs) est ‘faible, 3,75 (3,80) dans la L.S.; les distances prédorsale et préventrale font 54% (52% et 51%), la base de l'anale 20,5% (22,5%) (1,4 fois (1,25) dans la longueur de la tête), et la hauteur du pédicule 9% (?), de la L.S.; la tête est plutôt courte, 3,5 (3,6) fois dans la L.S.; espace interorbitaire 3,33 (2,8), orbite 5,45 (5,40), maxillaire 3,0 (3,1), mandibule 2,1 (2,1) et museau 4,33/3,65 (4,3/3,5) (projection/oblique) dans la longueur de la tête. Le grand sousorbitaire fait 78% (84%) de la joue. Les dents sont petites, tricuspidées pour la plupart; au prémaxillaire, 10-11 (10-11) externes, 2 internes (2), serrées contre les externes et disposées entre les 1-2 et 2-3 internes; 9 (9-8) internes, la 3e légèrement en avant; le maxillaire, relativement long, porte 25 (25) dents à droite et 28 (26) dents à gauche, en comptant les racines des dents brisées; mandibule difficile à étudier sur le type (mieux visible sur le N° A.8612); probablement 10 ou 11 (16) dents dans la rangée externe, les médianes usées, et une dizaine (17?) de dents postérieures en une rangée interne; une dent conique symphysaire du côté gauche seulement (id., en crochet). Aucun patron de coloration n'est actuellement visible sur le type, mais on croit distinguer une tache à la fin du pédicule sur l'autre spécimen. Outre la localité typique, cette espèce pose d'autres questions, notamment celle de savoir pourquoi elle a été si rarement mentionnée après CUVIER & VALENCIENNES (op. cit.), et surtout presque jamais récoltée. En dehors des catalogues de EIGENMANN & EIGENMANN (1891), EIGENMANN (1910) et HOWES (op. cit.), B. opalinus n'a été cité que par MUELLER & TROSCHEL (1844), qui ont assigné Chalceus opalinus à leur nouveau genre Brycon, KNER (1860) et GUENTHER (1864). Seul KNER a pu en étudier des spécimens (2 ?) ex. secs récoltés par Natterer à Irisanga, Rio Mogi-Guaçu), que GUENTHER (1864), sans avoir vu le type de B. opalinus, a pensé être différents et nommé B. nattereri. KNER (1860) avait souligné que ses exemplaires correspondaient parfaitement à la figure et au compte des écailles tels que donnés par CUVIER (sq. 8/50/4), mais que l'anale avait 23 rayons et que la base de la caudale était marquée d'une grande tache noire. Ce sont ces deux caractères différentiels qui ont motivé la création de Brycon nattereri fondé sur les exemplaires de Kner. Rappelons que les 2 ex. du Muséum de B. opalinus peuvent être définis par sq. 7/45- 49/4-5, A.iii (ou iv), 23-24 et peut-être par la présence d'une tache à la fin du pédicule caudal. La description complémentaire de Brycon bahiensis Günther, 1864, faite par HOWES (op. cit.), fait penser que les trois espèces (B. opalinus, nattereri et bahiensis) pourraient n'être qu'une seule et unique forme. (b) Brycon hilarii (Valenciennes in Cuvier & Valenciennes, 1949). Résumé de la description originale et historique: Chalceus hilarii Valenciennes, in CUVIER & VALENCIENNES, 1849 (Histoire naturelle des Poissons, tome 22, p. 246). «... M. de Saint-Hilaire avait rapporté avec l'espèce précédente.» (c'est-à-dire le grand individu de Chalceus opalinus, voir plus haut) «un autre Chalcee... ce poisson a la tête plus courte et plus grosse. L'intervalle entre les yeux plus large et plus convexe. L'ceil plus grand, beaucoup plus rapproché du bout du museau. La bouche a des intermaxillaires plus longs, et les maxillaires plus courts; … grosseur considérable des dents, etc.» «... nous aurions pu tirer la dénomination spécifique de ce poisson de la petitesse relative de ses écailles, qui sont beaucoup plus petites que dans les espèces précédemment décrites» (Chalceus macrolepidotus, C. ararapeera et C. opalinus). «... Une large tache noire couvre la base de la queue, les lobes de la caudale et l'anale. Nous avons un exemplaire rapporté du Rio San Francisco, long de vingt pouces». NOTES SUR BRYCON 815 «M. de Castenau nous en a procuré plusieurs autres pris dans l'Amazone et dans d'autres rivières du Brésil...» (2 exemplaires conservés jusqu'à nos jours). BERTIN (1948) cite les spécimens décrits par Valenciennes de la façon suivante: «Syntype A.8616, 450 mm, sec, bon état; Rio S. Francisco, A. St. Hilaire (1822)» Syntypes A. 9893-9894, 2 ex. secs, en bocal, Amazone, Castelnau». Ces exemplaires, en mauvais état, proviennent en réalité de Salinas, bassin du Rio Jequitinhonha (CASTELNAU, 1855: 68), et non de l'Amazone. Description complémentaire de Brycon hilarii (Valenciennes): revu en 1980, l'exemplaire sec de L.S. 415 mm, N° A.8616-81-25-71, Rio San Francisco, St. Hilaire 1922, est désigné ici comme lectotype. Il a les caractères principaux suivants: hauteur environ 4, tête environ 3,85 et distance prédorsale environ 2,15 dans la L.S.; le museau est court, ainsi que le maxillaire, environ 3,8 dans la tête, avec environ 20 dents; D.11,9(1); A. ?? iii, 20 avec une erreur de 2 rayons en plus ou en moins; P ?, courtes, n'atteignant pas les ventrales. Squamae certainement pas moins de 60 et pas plus de 65, 13 ou 14 /environ 10 en transversale; une paire de dents coniques mandibulaires. Patron de coloration indistinct. Les 2 exemplaires de Castelnau, N° A. 9893 et A.9894, conservés à sec en flacon, sont en très mauvais état. Le moins mal préservé, N° A.9894, environ 170 mm L.S., a peut-être 11/62/5 écailles (sur la figure de CASTELNAU, 1855, pl. 36 fig. 1, on ne compte que 46 écailles). Le patron de coloration figuré par Castelnau ne semble pas comporter de tache caudale. Il est impossible de dire s'il s'agit de la même espèce, qui doit être caractérisée par le seul lectotype (voir plus loin les remarques sur la distribution de l'espèce). Nous avons du mal à expliquer l'énorme différence entre le chiffre de VALENCIENNES (op. cit.) pour les écailles longitudinales («quatre-vingt») et celui relevé sur l'holotype (60 à 65, compte tenu des erreurs de comptage). Il ne peut s'agir d'une substitution de spécimen, le N° A.8616 représentant incontestablement le type de l'espèce, étant donné sa taille et les données d'époque qui l'accompagnent. Un lapsus calami semble pouvoir être écarté, Valenciennes ayant insisté sur la petitesse relative des écailles (les espèces décrites en même temps que Chalceus hilarii, c'est-à-dire Chalceus orbignyanus. C. rodopterus et C. carpophaga (ex. de Castelnau, non type) ont respectivement 62, environ 56 et environ 50 écailles en ligne latérale, ce qui n'est guère moins que le compte réel de B. hilarii). Enfin la confusion avec Salminus hilarii (faite par BERTIN, 1948) en raison de l'homonymie, est impossible à envisager de la part de Valenciennes. Le type de cette dernière espèce est en effet relativement petit (165 mm L.S.) et les paralectotypes les plus grands (maximum 345 mm L.S.), d'ailleurs conservés en alcool, sont encore loin d'atteindre la taille du type de Chalceus hilarii (415 mm L.S.). La marge du nombre d'écailles en ligne latérale est de 60-69 chez ces Salminus (cf. GÉRY & LAUZANNE 1990), très différente du chiffre de 80. L'erreur ne peut être due, selon nous, qu'à une mauvaise transcription en toutes lettres, sur le manuscrit définitif, du chiffre (60 ??) mentionné dans les notes originales. (c) Brycon carpophagus (non Valenciennes), CASTELNAU 1855 (pp. 68-69: «Chalceus carpophagus Cuv. Val.», Pl. 34 (fig. 3) HOWES (op. cit.: 17), d'après l'examen de K.A. Banister, a émis l'hypothèse que le spécimen décrit par CASTELNAU, 1855, sous le nom de «Chalceus carpophagus Cuv. Val.», n'était pas conspécifique avec le type de l'espèce, et que ses caractères seraient plus 816 J. GERY & V. MAHNERT voisins de deux de B. orthotaenia. Notre étude des spécimens du Muséum nous permet de - corroborer cette hypothese. Le spécimen récolté par Schomburgk en Guyane anglaise, cité en premier par VALENCIENNES in CUVIER & VALENCIENNES (1949: 252-253), est ici désigné comme lectotype de Chalceus carpophagus (bien qu'il ait été cité en second comme syntype par BERTIN, loc. cit., après l'exemplaire de Castelnau, cité en premier). Ce type (déjà revu par Howes, loc. cit., chiffres entre parenthèses), est un spécimen en alcool de 285 (280) mm de L.S., N° A.9832, caractérisé de la façon suivante: hauteur 2,8 (34%) et tête (assez courte) 4,2 (24%) dans la longueur standard; dorsale en avant du milieu du corps, distance prédorsale à 51% (53%) de la L.S.; A.111,23 (iv,23), P.1,14 (1,15), n'atteignant pas les ventrales; squamae en ligne latérale 60, à 2 près (58 +écailles caudales); 11/7 (12/9 jusqu'à la ligne médiane) transversales, environ 22 (?) prédorsales et 18 (9) circumpédiculaires; museau de la longueur moyenne; maxillaire avec une vingtaine de dents (22), mandibule avec une paire de dents postérieures symphysaires très petites; grand sousorbitaire couvrant presque toute la joue, sauf une zone nue de 2-3 mm; aucune coloration visible. Le spécimen désigné ici comme paralectotype est en mauvais état, mou et sans écailles. Il porte le N° 98 (81.25.3.1), de L.S. de 254 mm environ et a été récolté par Montravel dans «le fleuve Amazone». Les quelques caractères observables (anale, poches écailleuses etc.) plaident pour la conspécificité. C'est ici l'occasion de signaler que le type de Brycon siebenthalae Eigenmann, 1912, 170,5 mm de L.S., N° 53353, provenant de l'Aruka River en Guyana, et revu au Field Museum de Chicago il y a 25 ans par l'un de nous (J.G.), présente tous les caractères de B. carpophagus (contrairement à la mention de la clé, le spécimen n'a pas les rayons caudaux médians prolongés). Comme il provient du même territoire, il s'agit très probablement d'un synonyme. Sa denture a été figurée in GÉRY (1978), mais l'espèce n'est pas citée dans la clé et la synonymie probable n'est pas mentionnée. Le patron de coloration est encore visible: une grande tache ovale horizontale dans la région humérale, une série de lignes entre les écailles et pas de marque caudale. Les pectorales et les ventrales sont noirâtres. Le troisième exemplaire, indiqué en premier par BERTIN (1948) comme syntype (probablement parce qu'il avait été figuré par CASTELNAU (1855), mais postérieurement à la première description, ce qui supprime un argument pour le choix du lectotype), appartient à une espèce différente: Chalceus carpophagus non Val., CASTELNAU, 1855, N° A.8615, coll. Castelnau, «Rio de Sabara», Minas Gerais (= Ril das Velhas à 50 km au sud de Lagoa Santa). L'exemplaire sec d'environ 310 mm de L.S. est caractérisé de la fagon suivante: hauteur 3,4 et téte (courte) 4,5 dans la longueur standard; dorsale légèrement en avant du milieu du corps, distance prédorsale à 50,5% de la L.S.; A. 111,28 ou 29, P. 1,12, n'atteignant pas les ventrales; squamae en ligne laterale 50; 11/4 1/2 transversales, 20 ou 22 prédorsales et 18 (?) circumpédiculaires; canaux de la ligne latérale ramifiés; museau court; maxillaire avec environ 24 dents, mandibule avec une paire de dents postérieures symphysaires non réduites; grand sous-orbitaire couvrant presque toute la joue (sauf une zone nue de 1-2 mm); aucune coloration visible. Méme en tenant compte du fait que la comparaison entre un spécimen sec et un spécimen conservé en alcool est hasardeuse, on doit conclure que, par rapport a B. carpophagus, l'exemplaire de Castelnau est beaucoup plus allongé, avec beaucoup moins d'écailles longitudinales et au-dessous de la ligne latérale, et une anale nettement plus longue. Il pourrait représenter la même population que celle qui a été appelée Brycon lundii par LUETKEN (1874), car les caractères et les localités typiques sont identiques, à NOTES SUR BRYCON 817 condition d'admettre que Lütken ait compté toutes les écailles avec canaux, y compris celles qui bordent le posttemporal et le supracleithrum (la seule spèce du Rio Sao Francisco avec 60 ou plus écailles de la ligne latérale est B. hilarii, très différent, qui figure pour mémoire dans le tableau suivant). Nous admettons aussi provisoirement la synonymie avec B. orthotaenia (bassin du Rio Itapicuru), en attendant de pouvoir étudier un matériel abondant du Rio Sao Francisco. Le tableau suivant montre les similitudes entre ces espèces: type de type de B. carpophagus B. lundii B. orthotaenia B. hilarii sensu Castelnau d'après Lütken d'après Howes LS 310 ca 380 330 415 Haut. 30% ca 35% 33% 25% PD 50,5% ca 51% 53% 47% Tête 22% ca 23% 21% 26% Sq. 11/50/4 1/2 11/59-61/8 10/48/6 1/2 13-14/60-65/10 (fig. 10/55/5) A. 111,28 111,27-29 111,26 111,20? PS 1,12 1,13-14 il ? Mx. 24 x 22 ca 20 En résumé, il pourrait n'exister dans le Sud-Est et l'Est du Brésil (Haut Parana, Rio Sao Francisco, fleuves cötiers etc.) que trois Brycon: Une forme à anale longue (A. iii, 26-29 env.), B. orthotaenia Günther, 1864 (synonyme probable B. lundii), qui correspond également au B. carpophagus (non Valenciennes) de Castelnau, localisée dans le Rio Itapicuru et le Rio Sao Francisco (y compris le Rio das Velhas). Deux formes à anale plus courte (A. iii ou iv, 20-24 env.), B. opalinus (Cuvier, 1819) (synonymes probables B. nattereri Günther, 1864 — pour opalinus sensu Kner — et B. bahiensis Günther, 1864), qui habite du Rio Mogi-guaçu à «Bahia», et B. hilarii Valenciennes, 1849, du Rio Sao Francisco. Toutes deux semblent rares, peut-être en danger dans certains rios, et sont (ou ont été) peut-être sympatriques dans le Rio Itapicuru. Tous deux se distinguent aisément par le nombre des écailles: 7/45-49/4-5 pour B. opalinus contre 13-14/60-65/10 pour B. hilarii. On notera que deux de ces formes font partie du groupe d'espèces amazoniennes («Brycon falcatus-group») de Howes (op. cit.) dont il faut étendre la distribution. Quant à B. opalinus, il semble bien faire partie du même groupe, mais l'absence de matériel non historique ne nous permet pas de préciser ses caractères anatomiques. REMERCIEMENTS Nos remerciements vont à tous ceux qui nous ont aidés, particulièrement à Madame G. Arbocco (Museo civico di Storia naturale «Giacomo Doria», Genova), M.-L. Bauchot (Muséum national d'Histoire naturelle, Paris) et MM. R. Ahnelt (Naturhistorisches Museum, Wien), H.A. Britski (Museu de Zoologia da Universidade, Sao Paulo), L. Lauzanne (MNHN), C. Weber (Muséum d'Histoire naturelle, Genève, lequel a bien voulu examiner pour nous des exemplaires du Muséum de Vienne), et U. Werder (à l'époque INPA, Manaus). 818 J. GERY & V. MAHNERT REFEFENCES CITEES “ BERTIN, L., 1948. Catalogue des Types de Poissons du Muséum national d'Histoire naturelle. 3e Partie. Ostariophysaires (Characiniformes, Gymnotiformes). /mp. Colas, Bayeux, 42 pp. BÔHLKE, J.E., 1958. Studies on fishes of the family Characidae-N° 14. A report on several extensive recent collections from Ecuador. Proc. Acad. nat. Sci. Philad. 110: 1-121. BOULENGER, G.A., 1900. Viaggio del Dr. A. Borelli nel Matto Grosso e nel Paraguay. III. Boll. Mus. Zool. Anat. comp. Torino 15 (370): 1-4. BRITSKI, H.A., 1989. Lista dos tipos de Peixes das Coleçoes do Departamento de Zoologia da Secrataria da Agricultura de Sao Paulo. Pap. avuls. Depto. Zool. S. Paulo, 22 (19): 197-215. Campos, A. Amaral, 1950. Sobre a subfamilia Bryconinae. Especies existentes nao colecao de peixes do Departamento de Zoologia de Sao Paulo. Pap. avuls. Depto. Zool. S. Paulo 9(10): 137-143, fig. 1. CASTELNAU, F. de, 1855. Animaux nouveaux ou rares recueillis pendant l'expédition dans les parties centrales de l'Amérique du Sud etc. II. Poissons. Paris: i-xii, 1-112, pls. 1-50. Cope, E.D., 1872. On the fishes of the Ambyiacu River. Proc. Acad. nat. Sci. Philad. (1971): 250- 294, pls. ili-xvii. DAHL, G., 1943. New or rare Fishes of the Family Characinidae from the Magdelena System. K. Pfysiogr. Sällsk. Lund Förh. 12(18) (1942): 215-220. DAHL, G., 1971. Los Peces del norte de Colombia. /nDeReNa, Bogota (Characoidei: 100-156). EIGENMANN, C.H., 1910. Catalogue of the fresh-water fishes of tropical and south-temperate America. Repts Princeton Univ. Exp. Patagonia, 3 (1) (Zool.): 375-511. EIGENMANN, C.H., 1922. The fishes of Western South America, part. I. Mem. Carnegie Mus. 1): 1- 246, 21 figs., 35 pls. EIGENMANN, C.H. & R.S. EIGENMANN, 1891. A catalogue of the fresh-water fishes of South America. Proc. U.S. natl. Mus. for 1891, 14: 1-81. EIGENMANN, C.H. & C.H. KENNEDY, 1903. On a collection of fishes from Paraguay, with a synopsis of the American genera of Cichlids. Proc. Acad. nat. Sci. Philad. 55(2): 497-537. FOWLER, H.W., 1907. Further knowledge of some heterognathous fishes, Part I. Proc. Acad. nat. Sci. Philad. 58 (3): 431-483, figs 34-60. FOWLER, H.W., 1932. Zoological results of the Matto Grosso Expedition to Brazil in 1931. — Fresh Water Fishes. Proc. Acad. nat. Sci. Philad. 84: 343-377. GERY, J., 1964. Poissons characoides nouveaux ou non signalés de l'Ilha do Bananal, Brésil. Vie et Milieu, Supp. 17: 447-471. GERY, J., 1978. Characoids of the World. T.F.H. Publ. Inc., Neptune N.J. (1977): 672 pp. GERY, J. et L. LAUZANNE, 1990. Les types des espèces du genre Salminus Agassiz, 1829 (Ostariophysi, Characidae) du Muséum d'Histoire naturelle de Paris. Cybium 14(2): 113-124. GERY, J., MAHNERT, V. & DLOUHY, C., 1987. Poissons Characoïdes non Characidae du Paraguay (Pisces, Ostariophysi). Revue suisse Zool. 94 (2): 357-464. GUENTHER, A., 1864. Catalogue of the Fishes in the British Museum. Vol. 5 (fam. 2, Characinidae: 278-380). HILDEBRAND, S.E., 1938. A new catalogue of the fresh-water fishes of Panama. Field Mus. nat. Hist., Zool. Ser. 22(4): 215-359, figs 2-13. Howes, G., 1982. Review of the genus Brycon (Teleostei: Characoidei). Bull. Br. Mus. nat. Hist., Zool. Ser. 43(1): 1-47. KNER, R., 1860. Zur Familie der Characinen, III. Folge der ichthyologischen Beiträge (2. Abt.). Denkschr. k. Akad. Wiss. Wien,18: 9-62, pls. 1-8. LUETKEN, C.R., 1874. Characinae novae Brasiliae centralis. Overs. K. danske Widensk. Selsk. Forho. 1873 (3): 127-143. NOTES SUR BRYCON 819 MEEK, S.E. & S.F. HILDEBRAND, 1916. The fishes of the fresh waters of Panama. Publ. Field Museum (Zool. Ser.) 10: 217-374 pls. 6-32. MUELLER, J. & F.H. TROSCHEL, 1844. Synopsis generum at specierum familiae Characinorum (Pro- dromus descriptionis novorum generum et specierum). Archiv Naturgesch. 10 (1): 81-99. MYERS, G.S. & S.H. WEITZMANN, 1960. Two new fishes collected by General Thomas D. White in eastern Colombia. Stanford Ichthyol. Bull. 7 (4): 98-103. STEINDACHNER, F., 1876. Die Süsswasserfische des südöstlichen Brasilien II. Sitzber. k. Akad. Wiss. Wien 74: 559-694, pl. 1-13. THORMAHLEN DE GIL, A.L., 1949. Una Contribucion al Estudio del Pirapita (Brycon orbignyanus). Revta. Mus. La Plata (N.S.), 5; Zool.: 351-440. WERDER, U., 1983. Age determination by scale analysis in juvenile Matrincha (Brycon cf. melanopterus Müller & Troschel, Teleostei: Characoidei), a tropical characin from Central Amazon. Amazoniana 7 (4): 445-464. WERDER, U. & G.M. SOARES, 1984. Age determination by sclerite numbers, and scale variation in six fish species from the Central Amazon (Osteichthyes, Characoidei). Amazoniana 8 (3): 395-420. = wire, à Revue suisse Zool. Tome 99 Fasc. 4 p. 821-834 Genève, décembre 1992 Some new data on the Nicoletiidae (Insecta: Zygentoma) from Europe and Asıa Minor by ** kK Luis F. MENDES : With 29 figures and 1 map ABSTRACT In this paper, several samples of the Nicoletiidae genera Coletinia and Lepidospora are studied. The material has been collected in Sardinia, peninsular Italy, Yugoslavia, Greece, Austria and Turkey. Lepidospora (L.) escherichi is redescribed upon Ipiro and Levke island specimens and Lepidospora escherichi sensu Wygodzinsky (1980), from Kos and Rhodes islands, is considered as a new taxon, Lepidospora (L.) wygodzinskyi n. sp. INTRODUCTION The Nicoletiidae (Zygentoma) of Europe and mediterranean basin are represented by elements of two subfamilies. The plesiomorphic Protrinemurinae (MENDES, 1988) are known to occur only in aegean Greece — the troglobiont Protrinemura mediterranea has been described from Iraklia (MENDES, op. cit.) — and in Turkey — Trinemophora bitschiana (WYGODZINSKY, 1959), probably an edaphic species, is known from the vicinity of Izmir. The Coletiniinae, with the exception of some rare edaphic Lepidospora s.s. noticed from Sicily, peninsular Italy (Tuscany), Greece, Turkey and Israel, belong to the genus Coletinia, restricted to the northern and eastern mediterranean basin, which species are edaphic or cavernicolous. Nowadays, the known distribution of the Nicoletiidae Coletiniinae in this area, if we consider the reliable determinations only, is as follows: PORTUGAL — C. mendesi Wygodzinsky, edaphic (WYGODZINSKy, 1980). SPAIN — C. capolongoi Wygodzinsky, troglobic (WYGODZINSKY, 1980); C. asymetrica Bach et al. edaphic (BACH, MENDES & GaAJU, 1985); C. mendesi Wygodzinsky, edaphic (BACH, MENDES & GAJU, 1985). FRANCE — C. jeanneli (Silvestri), troglobic (SILVESTRI, 1938). * Centro de Zoologia do Instituto de Investigaçao Cientifica Tropical — R. da Junqueira 14, P- 1300 Lisboa Portugal. ** Fellow of the INIC — PL 2. 822 LUIS F. MENDES CORSICA — C. corsica (Chopard), edaphic (CHOPARD, 1924). SARDINIA — C. subterranea | (Silvestri), edaphic (WYGODZINSKY, 1980). PENINSULAR ITALY — C. subterranea (Silvestri), edaphic (SILVESTRI, 1902) (WyGODZINSKy, 1980); C. maggii (Grassi), edaphic (PARONA, 1888) (GRASSI & ROVELLI, 1889) (WYGODZINSKY, 1980) (MENDES, 1981). SICILY — C. maggii (Grassi), edapahic (GRASSI, 1887) (GRASSI & ROVELLI, 1889); C. subterranea (Silvestri), edaphic (WYGODZINSKY, 1980); C. setosula Wygodzinsky, (?) edaphic (WYGODZINSKY, 1980); L. (L.) grassii (Escherich), edaphic (ESCHERICH, 1905) (WYGODZINSKY, 1980). MALTA — C. maggii (Grassi), edaphic (MENDES, 1981). YUGOSLAVIA — C. maggii (Grassi), edaphic (WYGODZINSKY, 1980). BULGARIA — C. bulgarica (Kosaroff), (?) edaphic (KOSAROFF, 1939). GREECE (Corfu island) — C. subterranea (Silvestri), edaphic (SILVESTRI, 1908); — L. (L.) escherichi Silvestri, edaphic (SILVESTRI, 1908). TURKEY — C. longissima Mendes, troglobic (MENDES, 1988); L. (L.) aquilonaris (Wygodzinsky), edaphic (Wygodzinsky, 1959) ? (PACLT, 1974) ? (WyGop- ZINSKY, 1980). ISRAEL — L. (L.) silvestrii Wygodzinsky, edaphic (WYGODZINSKY, 1942). Further samples composed by female or juvenile specimens only, have been pointed out (WYGODZINSKY, 1980) to SPAIN (Coletinia sp I and sp II, both cavernicolous), to PENINSULAR ITALY (Coletinia sp HI and Lepidospora sp) and to YUGOSLAVIA (Coletinia sp IV), edaphic. The sample of C. subterranea refered to the Eastern Pyrenean, France, by DENIS (1923), corresponds undoubtedly to any other species in the genus; it has been pointed, indeed, «... ne montre pas d’appendice en languette a l’Ant. II gauche...», a quite typical feature of the SILVESTRI's 1902 species. In the present paper, we deal with several samples of Coletinia and with a few specimens of Lepidospora s.s.; unfortunately, many of the former are composed by females only, which unables an accurate determination. The specimens have been collected in the Sardinia island, peninsular Italy, Yugoslavia (Dalmatia), Greece (ionian area) and Turkey. Lepidospora escherichi Silvestri “redescribed” by WYGODZINSKY (1980) upon Kos and Rhodes material, is considered as a new taxon, L. (L.) wygodzinskyi n. sp.; almost topotypical material from the Silvestri's greek species (collected in Levkäs island and in the western Ipiros, geographically quite close to Corfu), allow the redescription of Lepidospora escherichi, which description was based upon immature specimens only. The material is deposited in the following entomological collections: Centro de Zoologia do Instituto de Investigagao Cientifica Tropical (CZ), E. Christian Collection (ECC), Gruppo Speleologico Sassarese (GSS), Museum d'Histoire naturelle de Genéve (MG), Museo civico di Storia Naturale “Giacomo Doria” (MGD), Museo regionale di Scienze Naturali di Torino (MT), Forschungsinstitut und Museum Senckenberg (MS) and Universita di Genova (UG). We are deeply greatfull to the colleagues who so kindly lent or offered the specimens which are the aim of this paper: Dr. E. Christian from Vienna, Austria; Dr. G. Gardini from Genova, Italy (UG); Dr. P.M. Giachino from Torino, Italy (MI); Dr. G. Grafitti from Sassari, Sardinia, Italy (GSS), Dr. B. Hauser from Genever, Switzerland (MG); Dr. R. Poggi from Genova, Italy (MGD) and Dr. H. Schröder from Frankfurt-Main, Germany (MS). Coletinia longissima Mendes, 1988 Examined material: TURKEY — Silifke, Astim Dilek Magara, 11/VII/1988, col. F. Gardini, R. Rizzario & S. Zoia, 36 4 29 2 6 immature specimens (UG) 14 19 (CZ). This troglobic species was quite recently described (MENDES, 1988) upon material from the same area; the types were collected some 20 Km NE of Silifke though in a NICOLETIIDAE FROM EUROPE AND ASIA MINOR 823 distinct cave (the Asthma Cave), quite probably part of the same cave-system. The specimens under study fully agree with the original description though some of the adults (of both sexes) are clearly longer than the firstly known ones; the longer d and @ attain a body length of 18.9 mm (5 mm more than in the bigger type-specimen) and one of the studied 2 9, which terminal filament is not damaged, shows a total length of 39.0 mm; the cerci of this same specimen, quite probably undamaged, are as long as 19.2 mm. Coletinia maggii (Grassi, 1887) Nicoletia phytophila PACLT, 1961 nec Gervais, 1844 n. syn. Examined material: AUSTRIA — Vienna, Stadtpark (=city park) under stones near composts, 15/V/1990, col. H. Gross, 25 3 (CZ); ibid. 28/V/1990, 14 (ECC). ITALY — Lippiano (near Arezzo), ?/VIII/1909, col. Dr. Andreini, 18 (MGD); Varazze (Liguria occ.), 28/11/1959, n° 188, 16 (MS). The sample from Varazze (MS) was determined as Nicoletia phytophila by PACLT (1961); during a short visit to the Senckenberg Museum and thanks to the kindness of Dr. Schröder, we had the chance to study this only slide which includes one typical male of Coletinia maggii. Nicoletia phytophila enters, so, as a new synonymy of the Grassi's species. The geographical distribution of this taxon, known till now to extend from Malta and Sicily through continental Italy to Dalmatia, is clearly widened northwards and the species reported for the first time in Central Europe; the presence of typical adult males of Coletinia maggii in northeastern Austria, collected in a garden (outside greenhouses) may correspond to a well established introduced population or may represent no more than the lack of carefully carried Zygentoma prospections in Slovenia, Karintia and Styria. The actual finding of Coletinia maggii in Vienna problemizes again the taxonomic position of the “Nicoletia” registered in Central Europe and particularly of those noticed from Hungary (PACLT, 1959 and 1974), which status has never been clarified: Nicoletia phytophila Gervais, 1844 or Coletinia maggii (Grassi, 1887) ? Coletinia subterranea (Silvestri, 1902) Examined material: ITALY (SARDINIA) — Campeda, 28/1V/1908, col. A. Dodero, 18 (MGD); Daieri, 2/V/1908, col. A. Dodero, 23 d (MGD). The species was known from Sardinia exclusively by the specimens collected in Bosa and reported by WYGODZINSKY (1980). The males now pointed to Campeda and to Daieri agree closely to the redescription of the species (WYGODZINSKY, op. cit.). Coletinia cf. subterranea (Silvestri, 1902) Examined material: ITALY (SARDINIA) — Voragine “Sa Nurra ’e Leone”, Monte Coazza, Dorgali, Nuoro, 8/XII/1985, col. R. Loru, n° 810 Dorgali, 12 (GSS): Grotta Pisanu (o di Gurennoro), Monte Coazza, Dorgali, Nuoro, 3/1/1987, col. G. Grafitti, n° 872 Dorgali, 19 (GSS); no precise location, no date, col. A. Dodero, 19 (MGD). Italy (Sicily) — Ficuzza, 25/V/1906, col. A. Dodero, 19 (MGD). The registered samples are considered under C. cf. subterranea (Silvestri), on account of some minor differences and due to the absence of collected males, though the main morphological characteristics agree fairly with those presented in the redescription of 824 LUIS F. MENDES the species (WYGODZINSKY, 1980). The only 2 from Ficuzza is uniformly yellowish brown and no differences on integument colour have been observed; the faint pigmentation must correspond, as WYGODZINSKY (op. cit.) refered, to a not fully developed specimen — the body length attained 7.4 mm. Both © © from the Dorgali district and the only specimen from the MGD collection, with the general characteristics of the species (the ovipositor is thin and long, extending behind the IX stylets by 2.75-3.6 times these stylets length), present, however, a few setae in the hind border of the Xth urotergite (Fig. 1); furthermore, the 22 from Monte Coazza caves, collected both in dark area, in stalagmitic soil or in stalagmitic concretions and 200-300 m far off the entrance, though not very well preserved, seem to be more thin and delicate than the typical specimens of Coletinia subterranea. Coletinia sp. A Examined material: YUGOSLAVIA — Krk island, near the chapel of Sveti Fusca, NW of the village Vrh, under a big stone in Quercus ilex bush, 20 m inland of the shore, 5 m asl, 28/VII/1987, col. E. Christian, 19 19 immature (CZ); Ibid., Sveti Fusca Bay, west of Vrh, Quercus pubescens wood, under stones, 3 m asl, 26/VII/1988, col. E. Christian, 19 immature (CZ). Both samples from the Krk island include females and one only is an adult specimen; its ovipositor is short and robust, 20-23 articulated and extending behind the IX stylets by about twice their own length; the subgenital plate is parabolic, rounded in the distal mar- gin and setose, and the cephalic capsule shows a “capolongoi-type” chaetotaxy (Fig. 2); the bigger © Xth urotergite (Fig. 3) remains that of Coletinia subterranea (WYGODZINSKY, 1980) though its posterior depression seems not so clear; it is completely different from the Xth tergite of Coletinia maggii — noticed from the Ombla valley, in Dalmatia (WYGODZINSKy, 1980) — as it is from the same tergite of Coletinia sp. III — collected in the neighbouring Sistiana (WYGODZINSKY, op. cit.) — the only Nicoletiidae in the area. Only the eventual capture of adult males in Dalmatia and/or Istria, will allow to know the exact taxonomic status of these samples. Coletinia sp. B Examine material: ITALY — Voltaggio, 27/VI/1908, col. A. Dodero, 19 (MGD). This only ? with 8.2 mm of body length, collected a few miles north of Genova, belongs quite probably to Coletinia maggii; the ovipositor is 23-25 articulated and extends beyond the IXth stylets by about 2.5 times their own length; presents a subgenital plate very alike those I observed in other © © of this species, shows a frontal chaetotaxy typical of this species, with short setae and a few macrochaetae; the Xth urotergite (Fig. 4) presents, however, a greater number of lateral marginal setae and much stronger apical setae in the under surface. Coletinia spp. Examined material: FRANCE — Galerie du Pré a Vincent, St. Didier au Mont d'Or, Rhone, 11/VI1/1986, col. M. Meyssonnier, 1 young & (MG). ITALY — Voltaggio, 20/IX/1909, col. F. Sotari, 1 juv (MGD). ITALY (SARDINIA) — Doliali, Dorgali, Gr. Pisanu, 15/X1/1985, n° 215 SINU, col. P.M. Giachino, 1 young d (MT). The registered specimens are too young to allow a more detailed determination; the young d from Sardinia was accompanied by the following label: “Nicoletia jeanneli NICOLETIIDAE FROM EUROPE AND ASIA MINOR 825 Silvestri det. P.M. Giachino 1986”; though devoided of sexual secondary characteristics, which will allow a specific determination, it is much more probable that this specimen will correspond to the species wich adults have been collected in this same cave, Coletinia cf. subterranea as stated before. Lepidospora (L.) escherichi Silvestri, 1908 Examined material: GREECE — Leucade (= Leukäs or Levkäs), au-dessus de Phryni, oliviers et macchia, 27/111/1971, col. V. Mahnert, n° GR-71/36, 18 19 (MG) 13 19 (CZ); Epire (= Ipiros), au nord de Kestrion, 30 m, sous arbustes, 5/V/1973, col. V. Mahnert, n° EP-73/96, 16 (MG). Fics 1-4. Fig. 1. Coletinia cf. subterranea 9, specimen from the MGD Xth urotergite; Fig. 2. Coletinia sp. A 9, frontal chaetotaxy of the adult specimen; Fig. 3. Ibid., Xth urotergite; Fig. 4. Coletinia sp. B ©, Xth urotergite. Scales: 0.1 mm. Lepidospora escherichi was described from Corfu (=Kerkira) (SILVESTRI, 1908) and, as latter reported (WYGODZINSKY, 1980), all the type material was immature; in this same paper, WYGODZINSKY “redescribes” the species upon adult males and females collected in the aegean island of Kös and points its presence (based on further young specimens) in the not very far Rhodes island. On account of the several similarities presented by the Corfu and the aegean specimens it was stated (WYGODZINSKY, 1980: 23): “... In order not to continue burdening the literature with names of unrecognizable species, I have redescribed Lepidospora escherichi from adults... collected on Kös, which, although not topotypical is LUIS F. MENDES 826 NICOLETIIDAE FROM EUROPE AND ASIA MINOR 827 reasonably close... to make specific identity a distinct possibility...”. The study of the adult specimens collected in the Levkäs island and in western littoral Ipiros, both geographically much closer to the type locality of Corfu, and the conspicuous dissimilarities presented by the ionian and the aegean specimens, led us to consider the material obtained in Phryni and in Kestrion as conspecific with L. escherichi, and the specimens from K6s and Rhodes (the aegean islands) as part of a not yet named taxon (while described in detail under the name of L. escherichi) we will analyse afterwards. Based on the ionian insular and western mainland specimens considered above, and having in mind that the only real description of this species is, indeed, the original one, based in non-adult material, we will proceed, right away, to its redescription. Body length: 5.1-5.2 mm (4) 5.0-5.4 mm (2); thorax length (1.9-2.0 mm (dé) 1.8- 1.9 mm (2); thorax width: 1.5-1.6 mm (6) 1.6-1.7 mm (9). Body limuloid, elongated. Hypodermal pigment absent, the general colour yellowish white or ivory. Macrochaetae yellowish brown, the stronger ones apically biphid. Scales typical, whitish, with abundant rays, little while clearly visible and very abundant. Head without special features, the frons with a few strong and long macrochaetae and numerous shorter and thiner setae, as in Fig. 5. Antennae of 2 without special characteristics. Pedicellus of d (Figs 6 and 7) with a strong distal inner apophysis, longer than wide at base, conical, provided with a very little anteapical conule and with an inner subovoid fovea; distal area of the apophysis extending beyond the level of the first flagellar article. Mandibles typical, with numerous acute and sclerotized distal teeth, as in Fig. 8. Maxillae as in the remaining species of the genus; galea with two strong apical conules, the lacinia equally long and with an inner pectinated area which extends close to the distal outer tooth, the bigger one of the pair. Maxillary palp as in Fig. 9, with a few strong macrochaetae in the 2nd and in the distal dorsal area of the 3rd article; distal article about 5 times longer than wide and 1/5 longer than the penultimate (n/n-1 = 1.26-1.33). Labium typical, the labial palp as in Fig. 10. Nota scaly, with thin and short setae and isolated macrochaetae along the lateral borders (Fig. 11) and with 2+2 short setae in the posterior margin; among the scales, in the disc, abundant minute isolated thin cilia. Legs robust, the tibia of P III (Fig. 12) about 4 times longer than wide, with one only dorsal (with the exception of the distal dorsal short ones) and 4 ventral thin macrochaetae, clearly shorter (ca 3/4) than the tibial diameter; femur with a pair of ventral distal and an isolated dorsal distal macrochata. Praetarsus without special features, the lateral claws longer than the empodium, this one clearly setulated. Urotergites I-VIII with 4+4 posterolateral (or 2+2 posterolateral and 2+2 close infralateral) macrochaetae, the external ones accompanied in the outer and ventral area by 5-6 thin setae, as in Figs 13 and 14; in the Ist urotergite, 1+1 or 2+2 short and delicate sublateral macrochaetae, in the remaining II-VIII tergites 2+2 stronger and longer macrochaetae wide spaced as in Fig. 15. IXth urotergite without strongly produced posterolateral lobes, with 1+1 sublateral macrochaetae only (Fig. 16), its lateroventral chaetotaxy as in Fig. 17, the inner macrochaeta of each pair distinctly shorter than the Fics 5-10. Lepidospora (L.) escherichi Silvestri, 1908. Fig. 5. Chaetotaxy of the frons, clypeus and labrum; Fig. 6. d pedicellus of antenna; Fig. 7. Ibid., from one other specimen; Fig. 8. Mandibles; Fig. 9. Maxillary palp; Fig. 10. Labial palp. Scales: 0.1 mm. 828 LUIS F. MENDES NICOLETIIDAE FROM EUROPE AND ASIA MINOR 829 outer one mainly in the d. Urotergite X of the 9 as in fig. 18, clearly emarginated, the angle formed by the inner margins of the apical notch of 90-100°; lateral margins with a row of 5-7 setae; macrochaetae of the posterolateral angles long and stout, longer than the marginal setated area. Xth d urotergite (Figs 19 and 20) deeply emarginated, the sides of the emargination forming an angle of ca. 50°, the posterolateral lobes triangular and wide; dorsal surface with a lateral row of strong setae and 1+1 similar antedistal setae, the terminal macrochaeta absent; ventral surface provided with 8-10 sclerotized brownish conules, the most anterior longer and clearly thiner, the remaining ovoid; apical peg the largest. Urosternite I with a triangular sternite and 1+1 wide lateral coxites, typical. Urosternites II-VII with a pair of stylets (Figs 21 and 22), entire, with 1+1 short and stout submedian macrochaetae close each other besides several thiner setae; those from the vesicles (in sternites II-VI) are longer, those from the pseudovesicles (in the VIIth) as long as the remaining; disc of the urosternites scaly, with isolated tiny cilia. Coxite VII of & (Fig. 23) with the submedian area protruding between the stylets and with about 8-10 stout and long macrochaetae, almost all clearly longer than the submedian ones of the preceeding sternites (twice longer and acute pointed — those of the II-VII are apically biphid); longer macrochaetae as long as 2/3-3/4 of the distance between the most external pair. [Xth stylets stout, much longer and stronger than the preceeding pairs, provided with some very strong setae and with two ventral long macrochaetae (Fig. 24). Paramera as in Figs 24 and 25, cylindrical, 4.5-5 times longer than wide, shorter than the level of the insertion of the stylet hind macrochaeta, attaining the length of 2/3 of the IXth stylets. Subgenital plate of the ? (Fig. 26) widely parabolic, with an irregular row of marginal short setae. Ovipositor strong, fusiform (Fig. 26), extending beyond the apex of the IXth stylets by their own length. Gonapophyses with 11 articles, the VIIIth clearly stronger, their chaetotaxy as in Figs 27 and 28. Cerci of d and all the terminal filaments of £ with the usual chaetotaxy, the terminal median filament with very strong and elongate ventral macrochaetae. Dorsal surface of 4 terminal filament (Fig. 29) with 10-12 sclerotized pegs in the basal area, arranged in 2 (exceptionally 3) longitudinal rows. Discussion: L. escherichi Silvestri, seems to approach, among the described mediterranean species, L. aquilonaris (at least the type material from Alexandretta, in as much as the sample from the european Turkey — PACLT, 1974 — has not been confirmed as belonging to this same species); the shape of the d Xth urotergite and its specialized chaetotaxy are quite alike; however, this so briefly described turkish taxon (WYGODZINSKY, 1959, as Lepidospora silvestrii aquilonaris) shows a Xth d urotergite less deeply emarginated, with a distinct while similar specialized chaetotaxy and the sclerotized pegs are clearly thiner than in the ionian SILVESTRI’s species; furthermore, the number of pegs in the basal dorsal terminal filament is clearly higher in the turkish species. Relatively to the aegean species which has been “redescribed” as L. escherichi (see afterwards), the differences are quite evident; they concern several features as: the shape Fics 11-17. Lepidospora (L.) escherichi Silvestri, 1908. Fig. 11. Hind border and posterolateral area of mesonotum; Fig. 12. P III; Fig. 13. Detail of the chaetotaxy of Ist urotergite; Fig. 14. Infralateral chaetotaxy and external sublateral macrochaeta of the VIIIth urotergite; Fig. 15. VIth urotergite; Fig. 16. IXth urotergite; Fig. 17. Ibid., infralateral chaetotaxy. Scales: 0.1 mm. LUIS F. MENDES NICOLETIIDAE FROM EUROPE AND ASIA MINOR 831 of the d pedicellar apophysis; the shape of the lateral lobes of the d Xth urotergite and the number and shape of the ventral pegs; the distinct elongation of the tibias (mainly Ti III) and the dorsal chaetotaxy of the femur; the number and mainly the length and robustness of the hind border macrochaetae in the d VIIIth urosternite; the number of sclerotized pegs in the ¢ terminal median filament; the more elongated ovipositor; the length of the posterolateral Xth urotergal macrochaetae in the 2. Lepidospora (L.) wygodzinskyi n. sp. Lepidospora escherichi WYGODZINSKY, 1980 nec Silvestri, 1908. As refered before, the samples of Lepidospora collected in the aegean islands of Kös and Rhodes (which correspond to the only known material of this new taxon), has been considered by WYGODZINSKY (1980) as Lepidospora escherichi and the material from Kös — the specimens from Rhodes are all immature — served as the base to his “redescription” of the SILVESTRI's species. The text and figures presented are quite detailed and are sufficient to characterize the new L. wygodzinskyi. The species is named after its describer, who noticed and figured accurately the main morphological characters as pointed, Dr. P.W. Wygodzinsky. It is, furthermore, an hommage to a great entomologist, desceased in the 27th January 1987, whose first publication on thysanurans (in 1939) was the start to an extraordinarily scientific career (see TERAN, 1987 and SCHUH & HERMAN, 1988) with a production of almost 250 papers; 70 among these, concern Microcoryphia and/or Zygentoma and include one of the largest contributions to the knowledge of these two orders: the original description of 11 genera and 146 species of Microcoryphia and of 16 genera and 80 species of Zygentoma in addition to an endless list of redescriptions. As far as we know, the mediterranean Lepidospora are completly allopatric, extending from Sicily and peninsular Italy to Greece, Turkey and Israel (MAP 1). The palestinian taxon (WYGODZINSKY, 1942) seems much more closely allied to the species described from the Kurdistan and southern Iraq as already stated (MENDES, 1985), on account of the much higher number of sensorial pegs in the ventral ¢ Xth urotergite. The status of the sicilian and of the mainland italian Lepidospora populations remains, as discussed by WYGODZINSKY (1980), quite problematic, as neither from Sicily nor from Tuscany adult specimens are actually known — nor even the validity of L. grassi has been definitely established, as no further material has been studied after the description of Nicoletia grassi (ESCHERICH, 1905). As already discussed (WYGODZINSKY, 1980) the species from Kös seems particularly close to the ionian L. escherichi and to the turkish L. aquilonaris by the type of chaetotaxy in the Xth urotergite of d, but it is completely distinct on account of the shape of the d antennal pedicellar apophysis and of the morphology of the lateral lobes of the d Xth urotergite; both these features seem to be species-specific. Fics 18-24. Lepidospora (L.) escherichi Silvestri, 1908. Fig. 18. Xth urotergite of ©, dorsal view; Fig. 19. Xth urotergite of d, dorsal view; Fig. 20. Ibid., detail of the ventral pegs; Fig. 21. Vth urosternite; Fig. 22. VIIth urosternite; Fig. 23. VIIIth urosternite of à ; Fig. 24. Genital area of d. Scales: 0.1 mm. 832 LUIS F. MENDES Fics 25-29. Lepidospora (L.) escherichi Silvestri, 1908. Fig. 25. Paramerum; Fig. 26. Genital area of 9 and subgenital plate; Fig. 27. VIIIth gonapophyses, distal articles; Fig. 28. IXth gonapophyses, distal articles; Fig. 29. Pegs of the 3 dorsal basal terminal filament. Scales: 0.1 mm. NICOLETIIDAE FROM EUROPE AND ASIA MINOR 833 11 MAP 1. Known distribution of the Coletiniinae of Europe and Asia Minor. Genus Coletinia (n. 1-11); genus Lepidospora (n. a-f). 1. C. asymetrica; 2. C. bulgarica; 3. C. capolongoi; 4. C. corsica; 5. C. jeanneli; 6. C. longissima; 7. C. maggii; 8. C. mendesi; 9. C. setosula; 10. C. subterranea; 11. Coletinia spp. a. L. (L.) aquilonaris; b. L. (L.) escherichi; c. L. (L.) grassi; d. L. (L.) silvestrii; e. L. (L.) wygodzinskyi; f. Lepidospora s.s. spp. REFERENCES BACH DE Roca, C., L.F. MENDES & M. GAJU-RICART 1985. 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Biography and bibliography. Petr Wolfgang Wygodzinsky (1916-1987). J. New York entomol. Soc. 96 (2): 227-244. SILVESTRI, F. 1902. Materiali per lo studio dei Tisanuri. III. Nuove specie di Nicoletia. Boll. soc. entomol. ital. 33: 223-227. — 1908. Materiali per lo studio dei Tisanuri. X. Su alcuni Tisanuri di Corfu. Boll. Lab. Zool. gen. agr. Portici 2: 381-393. — 1038. Descrizione di una nuova specie di Nicoletia vivente in una grotta della Francia (Insecta, Thysanura). Rev. franc. Entomol. 5: 188-193. TERAN, A.L. 1987. Dr. Pedro Wygodzinsky. 1916/1987. Nota necrolögica. Acta zool. Lill. 39 (1): 97- 98. WYGODZINSKY, P. 1942. Second contribution towards the knowledge of Diplura and Thysanura from Palestine. Rev. brasil. Biol. 2 (1): 29-46. — 1959. Beitrag zur Kenntnis der Machilida und Thysanura der Türkey. Opusc. entomol. 24: 36-54. — 1980. A survey of the Nicoletiinae of Europe (Nicoletiidae, Thysanura, Insecta). Amer. Mus. Novit. 2695: 1-24. | Revue suisse Zool. Tome 99 | Fasc. 4 | p. 835-858 | Genève, décembre 1992 | Mayflies from Israel (Insecta; Ephemeroptera) I.- Heptageniidae, Ephemerellidae, Leptophlebiidae & Palingeniidae by Michel SARTORI! With 45 figures ABSTRACT This paper is the first part of a work dealing with the mayfly fauna of Israel. Eleven species are reported here. The most diversified family is the Heptageniidae with six species belonging to six different genera: Rhithrogena znojkoi (Tshernova), Epeorus zaitzevi Tshernova, Ecdyonurus asiaeminoris Demoulin, Electrogena galileae (Demoulin) (comb. nov.), Afronurus kugleri Demoulin and Heptagenia samochai (Demoulin) (comb. nov.). E. zaitzevi is new for the fauna of Israel. The male of H. samochai is described for the first time and the synonymy with H. lutea Kluge (syn. nov.) is proposed. Eggs of the six species are described and illustrated. Keys are provided for nymphs and adults. Ephemerellidae are represented by a single species, Ephemerella mesoleuca (Brauer). Leptophlebiid species are: Paraleptophlebia submarginata (Stephens), Choroterpes (Ch.) picteti Eaton and Choroterpes (Euthraulus) ortali nov. sp. described at all stages. New features to distinguish the nymphs of the Mediterranean Euthraulus species are provided. One species of Palingeniidae has been found in the collections of Bet Gordon Museum in Deganya: Palingenia orientalis Chopra. The female of this species is described for the first time. P. orientalis disappeared from the investigated area in the early fifties. Some geographical data are given on the distribution of the species inside and outside the investigated area, as well as some ecological observations. For instance, underwater emergence is reported for the first time in the genus Afronurus. INTRODUCTION The mayfly fauna of the Near East or Levant is still not well known. In a work devoted to the northern part of this area (mainly Turkey, Syria, Lebanon), KocH (1988) lists the main important literature references available until now for this region. Concerning Israel, the study and collecting of Ephemeroptera can be divided into three main periods. “ This work was supported by a grant of the Swiss Academy of Sciences. | Museum of Zoology, Palais de Rumine, P.O. Box 448, CH-1000 Lausanne 17, Switzerland. 836 MICHEL SARTORI The first one (1930-1960) is related to the work of Y. Palmoni from Bet Gordon ~ Museum, who collected some mayflies, mainly in Lake Kinneret and Lower Jordan River area. The second one (1967-1971) is the important contribution by SAMOCHA (1972), who dealt with a general survey of the Ephemeroptera of this country. With the help of this material, DEMOULIN (1973) published the first taxonomical work and described four new species of the family Heptageniidae. The third one (1977-1990) is related to the work of the Inland-waters Ecological Service laboratory (IES) with the help of the River Surveillance System of the Nature Reserves Authority, Israel. During that time, more than 5'500 samplings have been made in more than 2'000 stations. Recently, I had the opportunity to study the important collection of Dr. M. Samocha, deposited at Tel-Aviv University, as well as the huge collection of the Institute of Life Sciences in Jerusalem. The whole material was available through the courtesy of Dr. R. Ortal (Department of Zoology, The Hebrew University of Jerusalem and Nature Reserves Authority, Israel) who Isincerely thank for his cooperation. Moreover, I had the opportunity to travel twice to Israel in 1990 and 1991 and to collect material in the most important localities. Almost twenty years after DEMOULIN (1973), I have to repeat the following foreword. Some localities where the material comes from, as well as some places where I had the opportunity to travel and collect material (mainly in the Golan Heights) are actually within the boundaries of Israel (cease-fire line of June 1967), but the property of such territories is always controversial. This first contribution deals with the families Heptageniidae, Ephemerellidae, Leptophlebiidae and Palingeniidae. The remaining families will be published later by other specialists. INVESTIGATED AREA Nine among the eleven species belonging to the above mentioned families are only found in the North Eastern part of the country, i.e. mainly Hula Valley and Golan (map 1). This area is the only one to provide suitable habitats for the Heptageniidae, Ephemerellidae and most of the Leptophlebiidae. The fact that no representatives have been found in the streams and rivers of Lower and Upper Galilee (except two tributaries of Lake Kineeret, N. Amud and N. Zalmon), which offers also a good diversity of habitats, is probably due to the generally higher water temperatures of these streams (BROMLEY, 1988), as well as to human impacts on the environment (R. Ortal, comm. pers.). Other areas, such as Judea, Samaria, Negev and the coastal plain, do not possess cold permanent running Waters, and so are only colonized by Caenidae and Baetidae species. Concerning the two other taxa, the palingeniid species has been found in the Lower Jordan River (between Lake Kinneret and the Dead Sea), whereas one leptophlebiid species, belonging to the genus Choroterpes (subgenus Euthraulus) has been found only in some restricted places near the Dead Sea (N. Arugot, En Gedi). HEPTAGENIIDAE As already mentioned, this family has been worked by DEMOULIN (1973). Some complements are brought to the fore in this study. The Israeli fauna is peculiar in that it is composed of six genera, each comprising only a single species. MAYFLIES FROM ISRAEL 837 Map I. Investigated area and detail of the North Eastern part with the main important watercourses. Dotted lines: temporary waters. 1: Nahal Senir (Hasbani River); 2: Nahal Dan, 3: Nahal Hermon (Banias); 4: Nahal Iyon (Ayoun River); 5: Upper Jordan River; 6: Nahal Meshushim; 7: Nahal Yadudiyya; 8: Nahal Daliyyot; 9: Nahal Amud. KEY FOR THE IDENTIFICATION OF THE NYMPHS This key is based on the one proposed by HEFTI & TOMKA (1989) for the European genera of Heptageniidae. 1 2 Nymph with two cerci, but no terminal filament ......................... Epeorus zaitzevi Nymphswithitwoicereiland oneiterminalfllament nn 2 ne 2. External borders of pronotum elongated caudally (fig. 5) ... Ecdyonurus asiaeminoris External broders of pronotum not elongated caudally .................................... 3 First gill lamellae expanded ventrally and bigger than the other ones ............... FO OSIO COLOUR POON OCR CHEE DIR D SS OE CI LIRA COCR RCE MELEE TI AAT CRE ars Rhithrogena znojkoi First gill lamellae not expanded ventrally and shorter than the other ones ............. 4 Galea and lacinia with a row of setae on the ventral side (fig. 4); head trapezoid; VIIth pair of gills with a tuft of filaments (fig. 6) ................. Heptagenia samochai Galea and lacinia with scattered hairs on the ventral side (fig. 3); head rounded; VIIth pair of gills without a tuft of filaments (figs 8-9) ................................... 5) Anterior margin of labrum with a median notch (fig. 1); inner face of the femora with a distinct dark spot; VIIth gill broad and rounded (fig. 8) ....... Afronurus kugleri 838 MICHEL SARTORI — Anterior margin of labrum concave, without a median notch (fig. 2); inner face of femora with a transversal dark band; VIIth gill almost pointed (fig. 9) ................... BRE .….….…. Electrogena galileae KEY FOR THE IDENTIFICATION OF THE IMAGOES i) Upperface offemora with a.distinct red to violet spot St e 2 — Upper face of femora without such spot (Caution! on lower face, a spot may be PLCSEMO VASE eee tel init eme LC 606856055 5206080666660 9002002: 3 2 Tergites uniformly coloured; male and female genitalia as in fig. 11 and fig. 16 TES DECTIVE VE M ee nee sin ROIO Rhithrogena znojkoi — Tergites with posterior margin bordered with a darker narrow band (fig. 38); male gsenitalasasın MSO et EEE Epeorus zaitzevi 3 Transversal veins in the costal and subcostal fields broader than the others, tinted Withivioleti(fig:21)} ann are Dane AMP 4 — Transversal veins in the costal and subcostal fields normally build, brown or yellow, bußnevertinted in Violet 72:41:22 le a A ER 5 4 Transversal veins in the costal field sinuous (fig. 21); male and female genitalia as in nes LE san onl all Sy sees III Heptagenia samochai — Transversal veins in the costal field not sinuous; male and female genitalia as in fig. fanden ern Ecdyonurus asiaeminoris 5 Body colour reddish-brown; male and female genitalia as in fig. 15 and fig. 20 à db bp BORD PAR Rea OS IE HO ANS RE RE CEE N IT Electrogena galileae — Body colour light brown or even yellowish; male and female genitalia as in fig. 14 ATT Bt IBAN nn RE ee elica RON de SO ARE Afronurus kugleri Epeorus zaitzevi Tshernova, 1981 Epeorus sp. SAMOCHA, 1972 Epeorus sp. DEMOULIN, 1973 Epeorus znojkoi BRAASCH, 1978a nec TSHERNOVA, 1938 Epeorus zaitcevi KAZANCI & BRAASCH, 1988 (injust. emend.) Epeorus zaicevi KOCH, 1988 (injust. emend.) Material examined: 19 subimago, 16 N, more than 300 L from 122 samples in 7 localities along the Dan River. For a description of the d, see TSHERNOVA (1981) and KAZANCI & BRAASCH (1988). For a description of the nymphs, see BRAASCH (1978a). Despite the lack of d imagoes, we agree with Braasch's opinion (in KocH, 1988), that specimens illustrated by DEMOULIN (1973) belong to E. zaitzevi. The nymphs are similar to those described by BRAASCH (1978a) (sub. nom. E. znojkoi). Moreover, a peculiar feature of this species is the presence of a narrow dark band on the posterior margin of each abdominal tergite (see TSHERNOVA, 1981 fig. 3 p. 225, and also fig. 38). This character is visible both in nymphal and winged stages. As for all known species of the genus Epeorus (DEGRANGE, 1960), the eggs of E. zaitzevi bear no peculiar exochorionic structures, nor polar cap (fig. 39). 2-3 micropyles visible in the equatorial area. MAYFLIES FROM ISRAEL 839 BE. zaitzevi A Rh. znojkoi X E. mesoleuca (\- MAP 2. Distribution of E. zaitzevi, Rh. znojkoi and E. mesoleuca. For explanations, see map 1. Distribution: E.zaitzevi has been found only in the headwater of the Jordan (map 2) and seems to be exclusively restricted to the torrential part of the Dan River (see also POR et al., 1986). In the visited localities, this species is rather common but is never abundant. So far known also from Caucasus (Armenia) (TSHERNOVA, 1981), Turkey (KAZANCI & BRAASCH, 1988) and from Syria (KOCH, 1988). Rhithrogena znojkoi (Tshernova, 1938) Ecdyonurus znojkoi TSHERNOVA, 1938 Rhithrogena sp. SAMOCHA, 1972 Epeiron znojkoi DEMOULIN, 1973 Rhithrogena sp. DEMOULIN, 1973 Epeorus znojkoi TSHERNOVA, 1981 Rhithrogena znojkoi THOMAS & DIA, 1982 Material examined: 1d, 23 N, 77 L from 43 samples in 9 localities along the Banias, Dan and Hasbani Rivers. The adults of Rh. znojkoi have been redescribed by THOMAS & DIA (1982), and the nymphs and eggs by SARTORI & SowA (1992), where its affinities and proper status have been quoted. In the investigated area, Rh. znojkoi can be easily separated from all other species, both in adult and nymph stages. Differential diagnoses are available in SARTORI & 840 MICHEL SARTORI Oo Fıc. 1-9. Heptageniidae nymphs. A. kugleri (1, 8), E. galileae (2, 3, 9), H. samochai (4, 6), E. asiaeminoris (5, 7). 1, 2: labrum; 3, 4: maxilla; 5: right half of the pronotum; 6-9: 7th gill. MAYFLIES FROM ISRAEL 841 14 15 Fics 10-15. Heptageniidae male genitalia in ventral view. 10: E. zaitzevi; 11: Rh. znojkoi; 12: E. asiaeminoris; 13: H. samochai; 14: A. kugleri; 15: E. galileae. Fig. 10 redrawn after KAZANCI & BRAASCH, 1988. 842 MICHEL SARTORI . SOWA (op. cit.). The eggs of Rh. znojkoi are easily recognizable by the tooth-like shape of the exochorionic structures (fig. 40). Distribution: Rh. znojkoi seems to be restricted to the Hula Valley, and particularly to the Dan and Hasbani Rivers, where this species is never abundant (map 2). Known also from Caucasus (Armenia) (TSHERNOVA, 1938), Lebanon (THOMAS & DIA, 1982; Dra, 1983; MOUBAYED, 1986), Turkey (KAZANCI & BRAASCH, 1988; KocH, 1988) and Syria (KOCH, 1988). Ecdyonurus asiaeminoris Demoulin, 1973 Ecdyonurus galileae SAMOCHA, 1972 nec DEMOULIN, 1973! Material examined: 2d,6%,79N, more than 260 L from 75 samples in 10 locali- ties along the Dan, Hasbani and Banias Rivers. This species belongs to the so-called “venosus group”, and is really easy to recognize, especially in larval stages. The seventh pair of gills bears also a tuft of tracheous filaments, as the six previous ones (fig. 7). Only one species shares the same feature: E. insignis Eaton, an inhabitant of European rivers. The transversal veins tinted with violet in the fields C and Sc of the forewings also allow separation of the winged stages. E. asiaeminoris presents no clear affinities with other members of the genus Ecdyonurus known to occur in Near- and Middle East, although some similar characters with E. ornatipennis Tshernova could be found (TSHERNOVA, 1938; BRAASCH, 1980a). The eggs of E. asiaeminoris are characterized by medium size KCTs, regularly arranged on the chorionic surface, except on one pole where they are bigger and all closer (fig. 41). Small rounded tubercles are also found on the surface. 7-8 micropyles present in the sub- equatorial area. Distribution: this species seems to be restricted to the Hula Valley, and especially Hasbani River after the confluence with the Dan River (map 3). The species occurs really scarcely in Banias and Tel Dan for instance, but is abundant in Dan river from Dan 5 locality (see Por et al., 1986; ALLAN et al., 1988 for localisation of the sites). E. asiaeminoris has never been quoted outside this area, and is unknown from Turkey, Lebanon or Syria for instance. Afronurus kugleri Demoulin, 1973 Afronurus kugleri SAMOCHA, 1972 “ Afronurus kugleri DIA, 1983 Afronurus kugleri MOUBAYED, 1986 Afronurus kugleri KOCH, 1988 Material examined: 278, 222,91 N, more than 450 L from 119 samples in 43 localities along 17 watercourses. A. kugleri is the second species of this genus known from the Mediterranean area. It shares some common features with its closest relative A. zebratus (Hagen) known exclusively from Corsica and Sardinia. The main differences between these two species concern the galea-lacinia of the nymphs. In A. kugleri, the distal part of the galea-lacinia bears 13-15 combs, the median ones with 9-10 teeth (fig. 22), whereas in A. zebratus, 19- 21 combs can be found, the median ones with 15-18 teeth (fig. 23). The peculiarities of egg exochorionic structures of A. zebratus (GAINO & MAZZINI, 1987; GAINO et al., 1987) can also been observed in A. kugleri. The whole surface of the chorion is covered with KCTs (fig. 42), and resembles what is found in A. zebratus. MAYFLIES FROM ISRAEL 843 BE. galileae Ci = X E. asiaeminoris % A H. samochai ( N ré j I LE ) D — & (È N. DA \ \ ) I) \ { ) i 4 \ i / i ; 2 7 EFT a J one 4 U 4 A MENSE, / a Ne a / { Map 3. Distribution of E. galileae, E. asiaeminoris and H. samochai. For explanations, see map 1. Distribution: A. kugleri is the most widespread Heptageniidae in Israel. It occurs in the Hula valley, the Golan Heights as well as in the Upper Jordan River (map 4). Known also from Lebanon (DiA, 1983; MOUBAYED, 1986) and from Turkey and Syria (Koch, 1988). A. kugleri seems to be more tolerant than other Heptageniidae, especially with regards to the temperature, i.e. this species can colonize other streams and rivers where the other ones are missing for (N. Meshushim, N. Samakh, N. Yahudiyya, N. Zavitan for instance). During our researches in the field, as well as during the rearings, we had the opportunity to observe the subimaginal emergence of A. kugleri. It was surprising to find this species has an underwater emergence. The nymphs begin to molt a few centimeters below the water level (under artificial conditions), the subimagoes then “swim” rapidly towards the surface for emergence. Until now, underwater emergence was known to occur only in the genus Electrogena (KIMMINS, 1941 and pers. obs.). The fact that A. kugleri (and probably also A. zebratus) exhibits the same behaviour also confirms the strong morphological and biochemical relationships between these two genera (HEFTI & TOMKA, 1989). Electrogena galileae (Demoulin, 1973) comb. nov. Ecdyonurus golanicus SAMOCHA, 1972 nomen nudum Ecdyonurus galileae DEMOULIN, 1973 Ecdyonurus galileae Dia, 1983 Ecdyonurus galileae MOUBAYED, 1986 Ecdyonurus galileae KOCH, 1988 844 MICHEL SARTORI AA. kugleri Map 4. Distribution of A. kugleri, P. submarginata and Ch. picteti. For explanations, see map 1. Material examined: 56d, 142, 162 N, more than 270 L from 96 samples in 28 localities along 14 watercourses. By its distinctive features both in nymphal and adult stages, E. galileae clearly belongs to the genus Electrogena Zurwerra & Tomka, 1985. But a comparative study of E. galileae with its close relatives is still not possible at the moment. This is mainly due to the great number of species only partly described during the last decade from nearby areas, mainly Caucasus and Transcaucasus (BRAASCH, 1978b, 1980a, 1980b, 1983). Never- theless, the status of E. galileae as species propria is certainly good for it can be distinguished from all other species described before 1973. The eggs of this species are typical for the genus and resemble those of E. grandiae for instance (GAINO et al., 1987). They are characterized by medium size KCTs and rounded tubercles covered by granular ground matrix. 4-5 micropyles are found in the equatorial area (fig. 43). Distribution: In Israel, E. galileae occurs mainly in the Hula valley, where it is particulary abundant in the sources regions, such as Tel Dan springs, Banias (map 3). Also reported from Lebanon (Dia, 1983; MOUBAYED, 1986) and Syria (KocH, 1988). But I am not sure the citations outside Israel are correct. The examination of undetermined material from Lebanon (A. Dia & A.G.B. Thomas leg.) indicates that there are at least two species belonging to the genus Electrogena in this country, neither of them related to E. galileae. . Heptagenia samochai (Demoulin, 1973) comb. nov. Sigmoneuria samochai SAMOCHA, 1972 Sigmoneuria samochai DEMOULIN, 1973 Heptagenia lutea KLUGE, 1987 (syn. nov.) MAYFLIES FROM ISRAEL 845 Fics 16-20. Heptageniidae female genitalia in ventral view. 16: Rh. znoijki; 17. E. asiaeminoris; 18. H. samochai; 19: A. kugleri; 20: E. galileae. Material examined:374,379%,19 N, 94 L from 31 samples in 17 localities along the Dan, Hasbani, Banias and Upper Jordan Rivers. The d imago of this species was not known from DEMOULIN (1973). We give here below its first description. Size: body length: 11.9 - 14.5 mm; forewing: 11.3 - 12.0 mm; cerci: 22.0 - 25.0 mm. Eyes uniformly greyish. Antennae with yellowish-brown pedicel and whitish funicule. Thorax more or less uniformly yellowish-brown. A small, dark elongated spot on the metapleurites. Forelegs light brown, with blackish junction between tarsi and tibiae. Middle and hindlegs yellowish. Wings translucent. Longitudinal veins brown. Transversal veins of the costal and subcostal fields tinted with violet or black. In the proximal part of the forewing, transversal veins of the costal field slightly sinuous, those of the subcostal field tinted in violet and more or less quadratic (fig. 21). 846 MICHEL SARTORI 21 22 23 Fics 21-24. 21. Proximal part of the costal and subcostal fields of the forewing in H. samochai; 22: comb of the galea-lacinia in A. kugleri; 23: comb of the galea-lacinia in A. zebratus; 24: cubito-anal field of the forewing in P. orientalis. Abdomen colourless, except tergites I-II whitish and segments VIII-X whitish- brown. Cerci whitish, every other junction strongly coloured in violet. Genitalia (fig. 13): general colour whitish. Styliger plate with a median smooth concave incision. On each penis lobe, ventral tooth long and regularly curved. Titillators short, pointed and straight. The eggs of H. samochai are similar to those found in other species, such as H. sulphurea (Miiller) or H. coerulans (Rostock) for instance (DEGRANGE, 1960). They are characterized by medium size KCTs, bigger on the polar cap, and a great number of micropyles (8-12) in the equatorial area (fig. 44). As already mentioned by BRODSKY (1980) and by KLUGE (1989b) the genus Sigmoneuria has no phylogenetic existence, and therefore has to be regarded as a junior synonym of Heptagenia. H. samochai is a very distinct species, both in imaginal and larval stages, and presents no clear affinities with species such as H. coerulans or H. longicauda (Stephens). By the shape of the cross veins in the C and Sc fields, H. samochai resembles H. perflava Brodsky, 1930 but can be separated on the shape of the penis lobes and titillators. Another related species is H. lutea Kluge, described from Caucasus, Armenia, Azerbajidzan, Georgia and Iran (KLUGE, 1987). I had the opportunity to compare H. samochai with specimens coming from Elbourz mountains in Iran (near Bujnurd, coll. F. Schmid). These specimens completely fit the description given by KLUGE (op. cit.), and therefore can be regarded as H. lutea. The only differences found between H. samochai and H. lutea are the shape of the cross veins in the subcostal field, triangular in H. lutea, whereas quadratic in H. samochai as well as MAYFLIES FROM ISRAEL 847 the margin of the styliger plate, smoother in H. lutea than in H. samochai. We could compare only the male imagoes but strong affinities are also visible in the nymphal stage, especially in the labrum and gill shape (see figs. 42-45 & 50 in KLUGE, 1987). Moreover, it is also Kluge's opinion (in litt.) that 7. lutea has to be regarded as a junior synonym of H. samochai. Distribution: A. samochai is mainly restricted to the Hula valley and the Upper Jordan River (map 3). Outside Israel, H. samochai is also found in Georgia, Armenia, Kimea and Elbourz montains (KLUGE, 1987, 1989a). EPHEMERELLIDAE This family is represented in the investigated area by a single species. Ephemerella mesoleuca (Brauer, 1857) Ephemerella maculocaudata IKONOMOV, 1961b Ephemerella sp. n. SAMOCHA, 1972 Ephemerella mesoleuca KocH, 1988 Material: 69,9 N, 23 L from 14 samples in 6 localities along the Hasbani, Banias and Upper Jordan Rivers. The specimens from Israel perfectly fit the diagnosis of E. maculocaudata Ikonomov, 1961 given by SOLDAN (1982) on the basis of material collected in Bulgaria. But recently, STUDEMANN & TOMKA (1989) have proposed the synonymy between these two species. Although some variations such as the size and position of the transversal dark band on the cerci can be noticed between populations (see also ALBA-TERCEDOR, 1991) they have to be regarded as belonging to the same species E. mesoleuca (D. Studemann, comm. pers.) Distribution: E. mesoleuca has been found mainly in the Upper Jordan River (map 2). It is also known from Banias and Hasbani Rivers where Samocha collected it, but this species has not been found again in these localities since 1971, where it has probably disappeared from. Found also by KocH (1988) in Syria (Orontes) but evidence of its distribution in other Levantine countries, mainly Lebanon and Turkey, is not yet established. LEPTOPHLEBIIDAE In his work, SAMOCHA (1972) reported three leptophlebiid species from Israel, none of them specifically identified, and belonging to the genera Paraleptophlebia and Choroterpes (subgenera Choroterpes and Euthraulus). In the examined material, only these three taxa have been found, each of them represented by a single species. Paraleptophlebia submarginata (Stephens, 1835) Paraleptophlebia sp. SAMOCHA, 1972 Material examined: 1d,4L from only 4 samples in 4 small watercourses (En Abu Fakusa, En Jalabina, En Qusbyie, Nahal Gamla). The Israeli specimens completely fit the diagnosis for this species, and therefore present no differences with European populations. Both nymphs and imagoes are easy to recognize. 848 MICHEL SARTORI Distribution: probably one of the less abundant species in Israel. P. “ submarginata seems to be restricted to the Golan Heights (map 4), and has not been found in other places, especially in the Hula valley. Outside the country, the species is known from Iran (BRAASCH, 1981) and Turkey (KAZANCI, 1986). This species is widespread in Europe (PUTHZ, 1978). Choroterpes (Choroterpes) picteti Eaton, 1871 Choroterpes (s.s.) SP. n. SAMOCHA, 1972 Material examined: 56,59,20 N, 35 L from 12 samples in 8 localities along 8 watercourses. This species, thought to be new by SAMOCHA (1972), is in fact the same species which occurs in Europe. Its distinctive features from other new species described recently from the Mediterranean area, such as Ch. volubilis Thomas & Vitte, 1988 (Morocco) or Ch. borbonica Belfiore, 1988 (southern Italy) are well marked, especially the shape of the penis lobes and the colouration of the forewings. Distribution: Ch.picteti has been found in the Golan Heights and also in the Hula valley (map 4). Known also from Lebanon (DIA, 1983) and Turkey (KAZANCI, 1984). Choroterpes (Euthraulus) ortali nov. sp. Choroterpes (Euthraulus) sp. SAMOCHA, 1972 Material examined: 14 imago holotype (with its nymphal exuviae); 26 d imagoes, 14 subimago, 3? © imagoes, 29 © subimagoes, 15 nymphs paratypes: Israel, Dead Sea Area, Nahal Arugot (En Gedi), -300 m below sea level, 12.V.1991 (coll. M. Sartori). Other paratypes: 16 imago, 16 subimago, 2 nymphs, same locality, 15.V.1990 (coll. R. Ortal & M. Sartori); 3 nymphs, same locality, 6.XII.1990 (coll. R. Ortal). Other material (not type specimens): more than 500 larvae and nymphs from the same wadi (coll. Nahal Arugot project); 1 larva, Ein Doyuq, 8.1II.1970 (coll. Avrahami) and 1 larva, En Mishmar, 25.11.1970 (coll. Gasith). All specimens preserved in alcohol except one d paratype partly in microscopic preparation (wings, genitalia). Holotype and most of the paratypes housed in the Musée de Zoologie, Lausanne. 1d imago, 1d subimago, 19 imago, 1? subimago and 4 nymphs paratypes are deposited in the Museum of Zoology of the Tel-Aviv University. Description Nymph Sizes (without cerci): d nymphs up to 6.3 mm; 2 nymphs up to 7.5 mm. Head medium brown; thoracic sclerites dark brown. Legs yellowish-brown. Upper face of femora with an elongated dark spot in the distal part. Abdominal tergites greyish- brown, sternites yellowish-brown. Cerci light brown, proximal part darker. Anterior margin of the labrum with a very smooth emargination, and with two rows of fine setae (fig. 25). Antero-lateral projection of the lacinia with a small projection. 2nd/3rd segments of the maxillary and labial palps ratio about 1.7 (figs 27-28). Hypopharynx as in fig. 26. Legs with the same kind of ornamentation as in Ch. (Eu). arabica (SARTORI & GILLIES, 1990). Hind tibiae with a row of 24-27 external bristles (fig. 30). Tarsal claws moderately hooked, with 13-14 teeth. Gills typical of the subgenus Euthraulus, with gill I lanceolate, and gills II-VII with two lamellae plate-like and terminated in three slender, subequal processes. Lateral margin of terga VII-IX as in fig. 29. IXth sternite of male and female nymphs as in fig. 31 and fig. 32 respectively. MAYFLIES FROM ISRAEL 849 28 "rem 27 29 31 32 Fics 25-32. Ch. (Eu.) ortali nov. sp. Nymph. 25: labrum; 26: left half of the hypopharynx; 27: labial palp; 28: maxillary palp; 29: lateral margin of abdominal tergites VII-IX; 30: bristles on the outer margin of hind tibiae; 31: sternite X in male nymph; 32: sternite X in female nymph. 850 MICHEL SARTORI 33 Figs 33-37. Ch. (Eu.) ortali nov. sp. Male imago (33-35), female imago (36), male subimago (37). 33: hind wing; 34: genitalia in ventral view; 35: detail of the penis lobes in ventral view; 36: subanal plate; 37: genitalia in ventral view. MAYFLIES FROM ISRAEL 851 Male imago Size: body length: 7.8 - 8.2 mm; forewing: 6.4 - 6.6 mm; cerci: 8.8 - 9.4 mm. Upper part of the eyes brownish-red. Thoracic sclerites dark brown. Femora of all legs greyish-brown with an elongated spot in the distal part of the upper face, as in the nymphs. Femoro-tibial joints medium brown. Tibia and tarsi yellowish-brown. Abdominal terga greyish-brown, sterna lighter except the IXth medium brown. Forewings translucent, except basis of C, Sc and R veins medium brown, and pterostigmatic area milky. Pterostigma with 9-12 simple transversal veins. Rs vein forked a little bit nearer to base of wing than attachment of vein MP, to MP}. MA forked over half of distance from base to margin. MP vein asymmetrical. 4 longitudinal veins in the cubital area. Hindwings with a rounded and symmetrical process; subcostal vein reaching the costa immediately behind the process (fig. 33). Genitalia Subgenital plate medium brown; forceps yellowish-brown. Hind margin of the subgenital plate regularly convex, without incision. First segment of the forceps very broad in the proximal half, suddently reduced in the apical half, forming a small rounded inner process (fig. 34). Penis lobes well developed, scissor-like, pointed at their apex which bears about 5 small spines (fig. 35). Female imago Size: body length: 5.8 - 6.2 mm; forewing: 6.5 - 6.8 mm; cerci: 6.0 - 7.5 mm. Thorax dark brown, wings entirely translucent. Abdominal tergites medium brown, sternites greyish-brown. Subanal plate elongated with a median incision (fig. 36). Male subimago General colour as for the d imago, but more contrasted. Wings entirely greyish, except the proximal part of the forewings greyish-brown. Genitalia as in fig. 37. Female subimago General colour of the abdomen reddish brown. Wings greyish. Eggs General shape ovoid. Length: 170-180 um, width: 95-105 um. Exochorionic structures with cross and asteroid costae (fig. 45). In the middle of them, a small adhesive element is present which is bigger on one pole. Aiea ni t 16s In nymphal stage, Ch. (Eu.) ortali shares some common features with other representatives of the Mediterranean area. It can be separated from Ch. (Eu.) lindrothi (Peters, 1980), Ch. (Eu.) balcanica (Ikonomov, 1961) and Ch. (Eu.) assimilis Gaino & Sowa, 1985 by the shape of the labrum, especially the less marked median emargination. Other mouth parts rather similar, as well as gills. Legs ornamentation can also provide good features to separate these four species, especially on the hind legs. The shape and number of the bristles on the outer margin of the hind tibiae are rather constant in each species. In Ch. ortali, there are 24-27 long, subparallel bristles, rounded at the apex, whereas they are shorter, less numerous and enlarged at the apex in Ch. assimilis, and much longer, less numerous, and pointed at the apex in Ch. lindrothi. In Ch. balcanica, they seem to be shorter and less numerous (see IKONOMOV, 1961a, fig. 9, p. 6). 852 MICHEL SARTORI In winged stages, Ch. ortali can be compared with certainty only to Ch. lindrothi and Ch. arabica, for male imagoes of Ch. balcanica and Ch. assimilis are still unknown. Genitalia of Ch. ortali are quite different from those of Ch. lindrothi, especially in the shape of the penis. It presents some affinities with Ch. arabica, but can be separated by the shape of the first segment of the forceps, as well as by the outer margin of penis lobes. 3 subimago of Ch. ortali can be distinguished from the one of Ch. assimilis by the shape of the penis lobes. On the basis of wing venation, distinctive features are hard to find between Ch. ortali and Ch. balcanica, although the subcostal vein of the hind wing seems longer in Ch. balcanica than in Ch. ortali. Finally, the eggs of Ch. ortali are easily distinguishable from those of Ch. assimilis (no exochorionic structures), Ch. lindrothi and Ch. arabica (polar cap, size and shape of the costae). Distribution: Ch. ortali has been found to occur widely in one tributary of the Dead Sea, Nahal Arugot, where this species is rather common. Prospections in the nearby wadi Nahal David where surprisingly fruitless. Information on the type locality is available in FURTH (1983). Known so far only from that area. Derivatio nominis: this species is named after Dr. Reuven Ortal (Jeru- salem) for his tremendous help during the whole study. PALINGENIIDAE The occurence of a Palingeniidae species in the Jordan River has been known since the mid-thirties (BODENHEIMER, 1935). But the specific identification of the population living in this watercourse was still enigmatic. BODENHEIMER (1935) reported the mass-flight of the gigantic species Palingenia jordanica, but unfortunately gave no description of this mayfly. In 1937, the same author proposed the synonymy of P. jordanica with P. orientalis Chopra. This synonymy has been done in fact by Y. Palmoni (1897-1971), who was the founder and director of “Bet Gordon”, the A.D. Gordon Agriculture and Nature Study Institute at kibbuz Deganya (LULAV, 1972). In 1939 specimens were sent to Dr. G. Ulmer for examination, and returned identified as P. longicauda (Olivier). Later on, other material was sent to Prof. V. Landa who determined them as P. sublongicauda Tshernova. This specific name has been used by SAMOCHA (1972) and remained until now. Thanks to the courtesy of S. Lulav, the actual director of Bet Gordon, we had the opportunity to study the collections of this institution. Among them, 19 Palingeniidae specimens are still available. They are all 9 9. No nymphs nor d d are preserved. However, these specimens could be related to Chopra's species, especially on the basis of their peculiar forewing venation. Palingenia orientalis Chopra, 1928 Palingenia jordanica BODENHEIMER, 1935 nomen nudum! Palingenia orientalis BODENHEIMER, 1937 Palingenia sublongicauda SAMOCHA, 1972 nec Tshernova, 1949 Material examined: 10 2 2: Israel, Lower Jordan River, Deganya (outlet of Lake Kinneret), -210 m below sea level, 7.1V.1944. Other specimens, all from the same place: 1 9 29.11.1935, 7 2 9 27.11.1944, 1 9 29.11.1947 (coll. Y. Palmoni). Description (dried specimens) Size: body length (without cerci): 22-31 mm; forewing: 24-29 mm; cerci broken. MAYFLIES FROM ISRAEL 853 Fics 38-41. Female subimago (37) and eggs (38-41). 38-39: E. zaitzevi; 40: Rh. znojkoi; 41: E. asiaeminoris. General coloration yellowish brown. Head yellowish; eyes black, as well as intraocellar area. Thorax medium brown, legs yellowish. Wings uniformiy milky, longitudinal veins (especially C, Sc, Rand MA) tinted in yellow. Abdominal tergites I-VI yellowish brown, VII-X light brown, without specific patterns; sternites uniformly light yellow. Cerci light brown. Forelegs short, twisted but not really atrophied. Middle and hindlegs normally built, the hind ones longer. On all tarsi, double claws, one hooked, the other obtuse. Inner border of the forewings regularly curved, without any winding. Vein MA forked over half of the distance from base to margin. 2 ICuA present. Vein CuA clearly bifid, CuA, long and regularly curved whereas CuA, short and reaching the border almost in a straight line (fig. 24). In Palingeniidae, wing venation, especially in the forewing, gives good criteria even to distinguish the taxa at the specific level (DEMOULIN, 1965; Soldan, comm. pers.). The examined specimens present the same peculiar cubito-anal venation as mentioned by GRAVELY (1920) and CHoPRA (1928). Moreover, they belong without any doubt to the genus Palingenia: regularly curved forewing, head without cephalic processes, vein MA 854 MICHEL SARTORI Fics 42-45. Eggs. 42: A. kugleri; 43: E. galileae; 44: H. samochai; 45: Ch. (Eu.) ortali. forked over half the distance from base to margin. Among the species belonging to that genus, P. orientalis can be easily separated from P. longicauda (Olivier), P. fuliginosa (Georgi), P. sublongicauda Tshernova and P. apatris Demoulin, in which vein CuA) is always much longer and not attached to CuA, as in P. orientalis. This peculiar branching of CuA is found for instance in most of Anagenesia species (DEMOULIN, 1965). But in this genus, MA vein is always forked nearer the base than margin. Bionomy.- The following information is based on the unpublished notes of Y. Palmoni. In Israel, P. orientalis was located in the Lower Jordan river, i.e. downstream to Lake Kinneret. One peculiarity of P. orientalis was to emerge and fly in the early morning (between 5 and 8 a.m.). In comparison, SOLDAN (1978) reports for P. sublongicauda early morning or evening emergences, whereas for P. longicauda and P. fuliginosa, only evening emergences have been reported (RUSSEV, 1987; SOLDAN & LANDA, 1986; pers. obs.). Most of the occurences reported here come from the Jordan River near Deganya (outlet of Lake Kinneret). P. orientalis has been observed from 1935 to 1950. Emergence MAYFLIES FROM ISRAEL 855 and flight took place from end of March (earliest date: 27.11.1944) until the beginning of May (latest date: 5.V.1939). Specimens began to fly before 6 a.m. until 7.30 or 8 a.m. On the 27th of March 1944, Palmoni noted that at 9 a.m., about 10% were still alive. Since 1950 (4.V.1950), no more specimens have been collected, nor observed, in spite of a survey in the area for several years. According to ORTAL & Por (1978), as well as information communicated by R. Ortal (Jerusalem) the disappearance of P. orientalis is related to human impact along the Jordan watercourse, where the suitable habitats for the larvae have been destroyed by new buildings for irrigation or electric power stations. P. orientalis was described from populations in Seistan (eastern Iran) and was never quoted since that time. The Jordan River population extends far to the west of the original distribution area of this species. No Palingenia species have been reported inbetween. The only Palingeniidae known from Middle East is Mortogenesia mesopotamica (Morton) from Iraq, but this species has few morphological affinities with representatives of the genus Palingenia (DEMOULIN, 1965). P. orientalis has to be considered as having disappeared from Israel as well as Jordan. Evidence of remaining populations in the area or even elsewhere is not known. BIOGEOGRAPHY OF THE STUDIED ELEMENTS Although Israel belongs to the Palaearctic region, it is situated near the border of the Afrotropical region to which it is connected through the Rift Valley (see Por, 1975; TCHERNOV, 1988). One species probably originated from Afrotropical region, the leptophlebiid Ch. (Eu.) ortali, found in the Dead Sea area. The subgenus Euthraulus is well represented by several species in Africa, as well as in the Oriental region (PETERS & EDMUNDS, 1970). This species is also the only one of the studied families to be present outside the northern part of the country. The genus Palingenia, represented here by the species P. orientalis, is a west Palaearctic element. All the other species are more or less restricted to the headwaters of the Jordan, the Hula Valley, and some streams in the Golan Heights, as for Israeli stoneflies species (BROMLEY, 1988). They are all Palaearctic elements although their origin is probably different. The two leptophlebiid species Ch. (Ch.) picteti, P. submarginata as well as the ephemerellid E. mesoleuca probably have an European origin, and they reach their most south-eastern boundaries in the Levant. In contrast, none of the heptageniid species are known from Europe. They can be divided in two main categories. First, species endemic to the Levant, such as E. galileae, E. asiaeminoris and A. kugleri. If E. asiaeminoris probably has a west Asiatic origin, E. galileae could be a Pontic element, whereas an African origin for A. kugleri is probable. Secondly, Caucaso-Levantine species such as E. zaitzevi, Rh. znojkoi and H. samochai. These species are distributed from the foothills of the Caucasus, through Turkey, Syria, Lebanon and they reach their southern boundary in the headwaters of the Jordan. ACKNOWLEDGMENTS At the end of this study, I am grateful to Dr. Amnon Freidberg (Museum of Zoology, Tel-Aviv) for the loan of Demoulin's type material, as well as to S. Lulav (Bet Gordon) for the loan of precious specimens of P. orientalis and to Dr. J van Tol (National Museum of Natural History, Leiden) for allowing me comparison with P. apatris. My sincere thanks 856 MICHEL SARTORI also to Prof. G. Lampel (Fribourg) for his russian translation of Kluge's description of H. lutea, and to Denise Studemann and Dr. Peter Landolt (Fribourg) for helpful discussion on the Ephemerella species. My gratitude to Dr. Tomas Soldän (Ceske Budejovice) for helpful advices, for allowing me comparison with other Palingenia species, and for his hospitality during my short visit in his laboratory. I am indebted to Dr. Heather J. Bromley (Jerusalem) for improving the English. The SEM photographs have been realized at the Electronic Microscopy Center of the Lausanne University. My stays in Israel in May 1990 and 1991 were made possible through a grant of the Swiss Academy of Sciences (travel funds), as well as financial support by the Museum of Zoology in Lausanne. The stay and travelling in Israel were arranged by the Nature Reserves Authority wardens, especially by Dror Pevzner and Hillel Glassmann during my stay in the kibbouz Ortal (Golan), and by Gershon Goldstein and Avi Reuven during my stay in Tel Dan. Special thanks also to all the staff of the En Gedi Nature Reserve, especially to Ya'al Dolev for accomodation. I am also grateful to Avi Reuven (She'ar Yashuv) and Glassmann's family (Harashim) for their friendly hospitality. Last but not least, nothing could have been realized without help and licence for collecting material in the Reserves provided by Reuven Ortal (Jerusalem), as well as for all accomodation and help during my field trips in Israel. RESUME x Cet article constitute la première partie d'un travail consacré à l'étude des Ephéméroptères d'Israél. Onze espèces sont citées. La famille la plus diversifiée est celle des Heptageniidae avec six genres renfermant chacun une seule espèce: Rhithrogena znojkoi (Tshernova), Epeorus zaitzevi Tshernova, Ecdyonurus asiaeminoris Demoulin, Electrogena galileae (Demoulin) (comb. nov.), Afronurus kugleri Demoulin et Heptagenia samochai (Demoulin) (comb. nov.). E. zaitzevi est nouveau pour la faune d'Israël. Le mâle de H. samochai est décrit pour la première fois, et la synonymie avec H. lutea Kluge (syn. nov.) est proposée. Les œufs des six espèces sont décrits et illustres. Des clés de determination pour les mâles et les larves sont inclues. Les Ephemerellidae sont représentés par une seule espèce, Ephemerella mesoleuca (Brauer). Les espèces de Leptophlebiidae sont: Paraleptophlebia submarginata (Stephens), Choroterpes (Ch.) picteti Eaton et Choroterpes (Euthraulus) ortali nov. sp. qui est décrite à tous les stades. De nouveaux critères pour distinguer les larves du sous-genre Euthraulus sont proposés. Une espèce de Palingeniidae a été découverte dans les collections du Bet Gordon Museum à Deganya: Palingenia orientalis Chopra dont la femelle est décrite pour la première fois. Cette espèce a disparu de la région concernée au début des années "50. Des informations géographiques sont également données sur la distribution de ces espèces à l'intérieur et en dehors de l'aire d'étude, de même que certaines observations écologiques. Par exemple, l'émergence sous l'eau est signalée pour la première fois dans le genre Afronurus. REFERENCES ALBA-TERCEDOR, J. 1991. Sobre el conocimiento de los Ephemerellidae ibericos: primera cita de Ephe- merella maculocaudata Ikonomov, 1961 (Insecta: Ephemeroptera). Eos 66(2): 209-214 (1990). ALLAN, J.D., G.N. HERBST, R. ORTAL & Y. REGEV 1988. Invertebrate drift in the Dan River, Israel. Hydrobiologia 160: 155-163. BODENHEIMER, F.S. 1935. Animal Life in Palestine. L. MAYER ed., Jerusalem, 506 pp. BODENHEIMER, F.S. 1937. Prodomus faunae Palestinae. Mem. Institut d'Egypte, Imp. Institut français d'Archéologie orientale, Le Caire, 286 pp. MAYFLIES FROM ISRAEL 857 BRAASCH, D. 1978a. Epeorus znojkoi Tschern. und /ron magnus n.sp. (Heptageniidae, Epheme- roptera) and dem Kaukasus. Entomol. Nachr. 22(5): 65-70. BRAASCH, D. 1978b. Neue Ecdyonurus-Arten (Ephemeroptera, Heptageniidae) aus dem Kaukasus und aus Transkaukasien (Sowjetunion). 1. Entomol. Nachr. 22(10): 145-159. BRAASCH, D. 1980a. Neue Ecdyonurus-Arten (Ephemeroptera, Heptageniidae) aus dem Kaukasus und aus Transkaukasien (Sowjetunion). 2. Entomol. Nachr. 24(2): 23-28. BRAASCH, D. 1980b. Neue Ecdyonurus-Arten (Ephemeroptera, Heptageniidae) aus dem Kaukasus und aus Transkaukasien (Sowjetunion). 3. Entomol. Nachr. 24(7): 103-110. BRAASCH, D. 1981. Eintagsfliegen aus Anatolien und Iran (Insecta, Ephemeroptera). Faun. Abh. Mus. Tierk. Dresden 8(6): 75-79. BRAASCH, D. 1983. Ecdyonurus armeniacus n. sp. aus der Armenischen SSR (UdSSR) (Ephe- meroptera, Heptageniidae). Reichenbachia 21(13): 87-90. Bropsky, K. 1980. Mountain torrent of the Tien Shan. A faunistic-ecology essay. Monographiae Biologicae 39, W. Junk, The Hague, 311 pp. BROMLEY, H.J. 1988. A note on the Plecoptera of Israel. Israel. J. Entomol. 22: 1-12. CHOPRA, B. 1928. The Indian Ephemeroptera (Mayflies). Part I. The suborder Ephemeroidea: Families Palingeniidae and Polymitarcidae. Rec. Ind. Mus. 29: 91-138 (1927). DEGRANGE, Ch. 1960. 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Recherches sur la faunistique, l'écologie et la zoogeographie de trois réseaux hydrographiques du Liban: l’Assi, le Litani et le Beyrouth. Thèse Doctorat d'Etat, Univ. Toulouse, 496 pp. ORTAL, R. & F.D. Por 1978. Effect of hydrological changes on aquatic communities in the Lower Jordan River. Verh. Internat. Verein. Limnol. 20: 1543-1551. PETERS, W.L. & G.F. EDMUNDS Jr. 1970. Revision of the generic classification of the Eastern Hemisphere Leptophlebiidae (Ephemeroptera). Pacific Insects 12(1): 157-240. POR, F.D. 1975. An outline of the zoogeography of the Levant. Zoologica Scripta 4: 5-20. POR, F.D., H.J. BROMLEY, CH. DIMENTMAN, G.N. HERBST & R. ORTAL 1986. River Dan, headwater of the Jordan, an aquatic oasis of the Middle East. Hydrobiologia 134: 121-140. PUTHZ, V. 1978. Ephemeroptera 256-263 in: J. ILLIES ed. Limnofauna Europaea, G. Fisher Verlag, Stuttgart 532 pp. RUSSEV, B. 1987. Ecology, life history and distribution of Palingenia longicauda (Olivier) (Ephe- meroptera). Tijd. 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TOMKA 1989. Contribution to the study of European Ephemerellidae (Ephemeroptera). III. Synonymy of Ephemerella maculocaudata Ikonomov, 1961 syn. n. with Ephemerella mesoleuca (Brauer, 1857). Mitt. schweiz. ent. Ges. 62(1/2): 129-130. THOMAS, A.G.B. & A. DIA 1982. Ecdyonurus (?) znojkoi Tschernova, 1938: redescription et appartenance générique reelle. (Ephemeroptera, Heptageniidae). Bull. soc. Hist. Nat. Toulouse 118: 297-303. TCHERNOV, E. 1988. The paleobiogeographical history of the southern Levant: 159-250, in : Y. YOM- Tov & E. TCHERNOV eds. The zoogeography of Israel. Monographiae Biologicae 62, W. Junk, The Hague, 600 pp. TSHERNOVA, O. 1938. Zur Kenntnis der Ephemeropteren Ost-Transkaukasiens. Trudy zool. Inst. Baku 8: 55-64. TSHERNOVA, O. 1981. On the systematics of adult mayflies of the genus Epeorus Eaton, 1881 (Ephemeroptera, Heptageniidae). Rev. Entomol. URSS 60(2): 323-336 (in Russian). | | | Revue suisse Zool. Tome 99 Fasc. 4 p. 859-953 Genève, décembre 1992 | | Monographie der Scydmaenidae (Coleoptera) von Sabah (NO-Borneo) von Herbert FRANZ * Mit 108 Abbildungen ABSTRACT Monography of the Scydmaenidae (Coleoptera) of Sabah (NE-Borneo). — Apart from 6 already known species, 109 new species are described and illustrated in the genera Scydmaenus (32 spp.), Loeblites (1 sp.), Syndicus (1 sp.), Horaeomorphus (3 spp.), Euconnus (69 spp.) and Protoscydmus (1 sp.). Three new monotypic genera and each a new subgenus in the genera Scydmaenus and Euconnus respectively are erected. VORWORT In dieser Monographie sind die von Burckhardt, Löbl und Smetana in den Jahren 1987 und 1988 im Kinabalu-Nationalpark in Sabah gesammelten Scydmaeniden be- arbeitet. Neben 6 schon bekannten Arten wurden 109 Arten als für die Wissenschaft neu beschrieben. Davon gehören 38 zur Gattung Scydmaenus, 1 Art zu Loeblites, und je eine zu Syndicus, Syndicomorphus und Borneosabahia, 3 zu Horaeomorphus, 69 Arten zu Euconnus und eine zu Protoscydmus. Die von Reitter aus Borneo beschriebenen Arten aus den Genera Euconnus und Scydmaenus habe ich nicht gesehen, der Verbleib der Typen ist mir unbekannt. Die im Deutschen Entomologischen Institut in Eberswalde verwahrten Syntypen der von Schaufuss beschriebenen beiden Agathelor-Arten wurden mir in freundlicher Weise zugesandt, die eine hat sich zahlreich im Material des Genfer Museums wiedergefunden. * Jakob-Thoma-Str. 3B, A-2340 Mödling, Österreich. 860 HERBERT FRANZ Ich bin dem Muséum d'Histoire Naturelle de Genève und dort vor allem Herrn Dr. I. - Löbl zu großen Dank dafür verpflichtet, daß mir das dort verwahrte Material aus Sabah zur Bearbeitung anvertraut worden ist. Es handelt sich um erheblich über 1000 Exemplare, wobei allerdings von einer Reihe von Arten große Serien, von anderen nur Einzel- exemplare vorliegen. Da von einigen Spezies offenbar nur 2 2 gesammelt wurden, die eine Determination bis zur Art unmöglich machen und anderseits zahlreiche Arten in nur einem Exemplar vorhanden sind, liegt die Vermutung nahe, daß trotz des Umfanges der vorliegenden Ausbeute noch keineswegs der ganze Artenbestand des Untersuchungs- gebietes erfaßt ist. Man muß demnach damit rechnen, daß der tatsächliche Artenbestand an Scydmaeniden des Nationalparks von Sabah die 115 dort derzeit nachgewiesenen Arten noch um einiges überschreitet. Alle in der vorliegenden Arbeit behandelten Taxa sind am Ende der Mongraphie nach Gattungen geordnet aufgezählt. Alle Holotypen werden im Museum in Genf (cMG), eine Anzahl von Paratypen in meiner Sammlung (cF) verwahrt. Meine Sammlung wird nach meinem Ableben dem Naturhistorischen Museum in Wien übergeben. Gattung Scydmaenus Latreille Untergattung Scydmaenus Latr. s. str. Scydmaenus novaehollandiae Lhoste Orig. Diagn.: LHOSTE (1938), Arb. morph. taxon. Ent. 12, p. 113-114, fig. 2,6. Die Art ist nach 2 Ex. aus Niederländisch-Indien, jetzt Indonesien ohne genauere Fundortangabe beschrieben. Mir liegen nun 36 Exemplare vor, die A. Smetana sowie Bruckhardt u. Löbl im Kinabalu-Nat. Park gesammelt haben. Von 2 d d von Poring Hot Springs, 495 m, 21.u.24.8.1988 wurden Penispräparate angefertigt, die mit den von Lhoste angefertigten Abbildungen übereinstimmen. Eines dieser beiden dd und 5 weitere Exemplare befinden sich in meiner Sammlung, alle anderen in der Sammlung des Mus. in Genf. Scydmaenus vestitoides Reitter Orig. Diagnose: REITTER (1913), Ent. Mitt. II (9), p. 268. FRANZ (1985): Mitt. Münch. ent. Ges. 74, p. 110-111. Von dieser Art befinden sich 41 Ex. in dem mir vorliegenden Material aus Sabah. Die Determination ist durch Anfertigung von Penispräparaten gesichert. Scydmaenus minangkabauensis Blattny Orig. Diagnose: BLATTNY (1926) Suppl. Ent. 14, p. 3 - 4, fig. 2. FRANZ, H. (1984), Sber. Ost. Akad. Wiss., Math. nat. Kl. Abt. I, Bd. 193, p. 104. Von dieser Art befinden sich in dem Material aus Sabah 12 Ex., die Determination ist durch Anfertigung von Penispräparaten gesichert. Scydmaenus (s. str.) kinabaluensis nov. spec. MATERIAL: Holotypus, 6, Mount Kinabalu 1750 m (lg. Burckhardt u. Löbl, cMG); ebenda 1 Paratypus (cF); 4 Paratypen, Mt. Kinabalu, 2600 m (1 g. Burckhardt u. Löbl, 2600 m (cMG) und ebenda 1 Ex., (cF), SCYDMAENIDAE VON SABAH 861 DIAGNOSE: Gekennzeichnet durch schlanke Fühler und Beine, gedrungene Gestalt, sowie fein punktierte, nach hinten gerichtet behaarte Flügeldecken. BESCHREIBUNG: Long. 2,00 bis 2,20 mm, lat. 0,80 bis 0,90 mm, Rotbraun, gelblich behaart. Kopf von oben betrachtet um ein Viertel breiter als lang, von den Augen zur Basis geradlinig verschmälert, die Schläfen doppelt so lang wie der Augendurchmesser. Die schlanken Fühler zurückgelegt die Halsschildbasis erreichend, beim © alle Glieder länger als breit, beim d 6 und 7 breiter als lang, das spitz-eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild beim 8 kaum merklich, beim 9 deutlich gestreckt, ohne Basalgrübchen. Flügeldecken zusammengenommen an der Basis nur wenig breiter als die Halsschildbasis, ohne Basalimpression und ohne Schulterbeule, fein und dicht punktiert, schräg nach hinten gerichtet behaart. Flügel verkümmert. Beine schlank und lang, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 1) distal dorsalwärts gekrümmt, Ostium penis lang, dorsal gelegen, halb so lang wie der Peniskörper. Aus ihm ragt der Ductus ejaculatorius dorsalwärts heraus, er ist nach hinten gebogen und überragt den Hinterrand des Apex penis ein wenig. Der Apex ist am Hinterrand dreieckig ausgeschnitten. Scydmaenus (s. str.) pseudovestitoides nov. spec. MATERIAL: Holotypus d (Penispräparat) und 6 Paratypen, Sabah, Poring Hot Springs, 9.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 6 Paratypen, (cF). DIAGNOSE: Robust und stark gewölbt, fein punktiert und dicht behaart. Der Penis in Lateralansicht dem des Sc. vestitoides ähnlich. BESCHREIBUNG: Long. 2,00 mm, lat. 0,80 bis 0,90 mm. Schwarz, die Extremitäten rotbraun gefärbt, gelblichgrau behaart. Kopf von oben betrachtet gerundet-querrechteckig, die Schläfen schwach zur Basis konvergierend, Stirn und Scheitel äußerst fein punktiert und behaart. Fühler dick, zurückgelegt die Halsschildbasis nicht ganz erreichend, ihr Basalglied doppelt so lang wie breit, 3 bis 6 isodiametrisch bis leicht gestreckt, 7 und 8 breiter als lang. 9 doppelt so breit wie 8, wie auch 10 schwach quer, das spitz-eiförmige Endglied nicht ganz so lang wie die 2 vorletzten zusammen. Halsschild kugelig gewölbt, isodiametrisch bis leicht gestreckt, vor der Mitte am breitesten, fein punktiert und dicht, abstehend behaart, mit 2 Basalgrübchen. Flügeldecken zusammen an der Basis etwas breiter als die Halsschildbasis, stark gewölbt, ohne Schulterbeule und Basalimpression, fein und dicht punktiert, lang abstehend behaart, mit 2 Basalgrübchen. Flügel verkümmert. Beine dick, Vorderschenkel dicker als die der Mittel- und Hinterbeine, Schienen gerade, Vordertarsen des d schwach verdickt. Penis (Abb. 2) wie bei Sc. vestitoides Reitt. annähernd in der Längsmitte im stumpfen Winkel nach oben geknickt, Apex penis breit abgestutzt, sein Hinterrand beiderseits mit einem feinen Stachel, die Seiten davor stumpfwinkelig erweitert. In der Mitte des Peniskörpers befindet sich dorsal ein ovales Fenster, in dem 2 durch eine Querverbindung aneinandergefügte kleine Drüsen liegen. Beide sind mit einem Ausführungsgang versehen, die beiden Ausführungsgänge sind distal ebenfalls miteinander verbunden. Scydmaenus (s. str.) trapeziceps nov. spec. MATERIAL: Holotypus d, Sabah, Poring Hot Springs, 500 m, 8.8.1987 (lg. Burckhardt u. Löbl, cMG); Paratypus 1 Ex. Sabah, Mt. Kinabalu, 1580 m, 27.4.1987 (lg. Burckhardt u. Löbl, cF). 862 HERBERT FRANZ u SE res Y5rım 15mm ABB. 1-8. 1: Scydmaenus kinabaluensis nov. spec., Penis in Lateralansicht. 2: Scydmaenus pseudovestitoides nov. spec., Penis in Dorsalansicht. 3: Scydmaenus trapeziceps nov. spec., Penis in Lateralansicht. 4: Scydmaenus bukitulari nov. spec., Penis in Dorsalansicht. 5: Scydmaenus crockerensis nov. spec., Penis in Lateralansicht. 6: Scydmaenus borneoensis nov. spec., Penis in Dorsalansicht. 7: Scyd- maenus borneoi nov. spec. Penis in Lateralansicht. 8: Scydmaenus fraternus nov. spec., Penis in Lateralansicht. SCYDMAENIDAE VON SABAH 863 DIAGNOSE: Sehr ausgezeichnet durch nahezu geradlinig trapezförmig von der Basis zum Vorderrand verschmälerten Kopf, ferner durch beim d nicht verbreiterte Vor- dertarsen und medialwärts gekrümmte Mittel- und Hintertibien. BESCHREIBUNG: Long. 2,30 bis 2,40 mm, lat. 0,90 mm. Dunkel rotbraun, graubraun behaart. Kopf von oben betrachtet trapezförmig, Augen flach, nahe dem Vorderrand der Stirn gelegen, diese in der Mitte mit einem flachen Längskiel. Fühler am Vorderrand des Kopfes nahe beieinander eingefügt, zurückgelegt die Halsschildbasis Knapp erreichend, ihr Basalglied doppelt so lang wie breit, breiter als die folgenden, 3 bis 5 eineinhalbmal so lang wie breit, 6 leicht gestreckt, 7 und 8 breiter als lang, 9 3 mal so breit wie 8, 10 noch etwas breiter als 9, schwach quer, das Endglied etwas asymmetrisch, um ein Viertel länger als breit. Halsschild etwas länger als breit, vor seiner Längsmitte am breitesten und hier etwas breiter als der Kopf, mit 2 Basalgrübchen, abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit Andeutung einer Basalimpression, lang, nach hinten gerichtet behaart. Beine kurz, Vordertarsen des d nicht verbreitert, Mittel- und Hintertibien mediodistal ausgerandet. Penis (Abb. 3) füllhornförmig, distal ventralwärts gebogen, an der Basis ringförmig stärker sklerotisiert, dahinter mit einem pilzförmigen Druckregulierungsorgan. Von diesem zieht ein Muskelstrang distalwärts zum letzten Drittel des Penis. Dieses ist durch eine Membran gegenüber dem vorderen Teil des Peniskörpers abgegrenzt. Distal davon liegt eine Reihe von 3 Kammern, deren letzte mit einem Ausführungsgang in das Ostium penis mündet. Scydmaenus (s. str.) bukitulari nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Mt. Kinabalu Nat. Park, Bukit Ular Trail, 1750 m, 29.4.1987 (lg. Smetana, cMG); Paratypus, Mt. Kinabalu, 2600 m, 2.5.1987 (1g. Burckhardt u. Löbl, cF); 2 Paratypen Mt. Kinabalu Nat. Park. Sumit Trail, 1850 m (lg. Smetana, cMG). DIAGNOSE: Durch verhältnismäßig lange Fühler, 4 Basalgrübchen des Halsschildes und nicht verbreiterte Vordertarsten des d gekennzeichnet. BESCHREIBUNG: Long. 1,80 bis 1,90 mm, lat. 0,80 mm. Schwarzbraun, bräunlich behaart. Kopf von oben betrachtet um ein Viertel länger als breit, mit schwach gerundeten Schläfen, diese doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halschildbasis etwas überragend, ihre Basalglied doppelt, das 2. knapp doppelt so lang wie breit, 3,4 und 5 um die Hälfte länger als breit, 6 isodiametrisch, 7 und 8 breiter als lang, die 3-gliederige Keule schlank, Glied 9 und 10 etwas weniger als die Hälfte breiter als 7, beide um ein Drittel länger als breit, das spitz-eiförmige Endglied etwas kürzer als die beiden vorletzten zusammen. Halsschild kugelig, annähernd so lang wie breit, etwas vor seiner Mitte am breitesten, mit 4 Basalgrübchen. Flügeldecken zusammen schon an der Basis wesentlich breiter als die Halsschildbasis, ohne Basalimpression und ohne Schulterbeule glatt und anliegend behaart. Beine verhältnismäßig schlank, Schenkel schwach verdickt, Schienen gerade, Vorder- tarsen des d nicht verbreitert, Penis (Abb. 4) von oben betrachtet zylindrisch, seine Basalöffnung dorsobasal, das Ostium penis terminal gelegen. Seine Dorsalwand im letzten Fünftel medial ausgeschnitten, die beiden Seiten aber weit distalwärts vorspringend. In das Ostium penis ragt aus dem Penisinneren ein Sklerotinstab vor, an dessen Basis sich eine kleine runde Blase befindet. Neben dem dicken Stab verläuft von oben und hinten 864 HERBERT FRANZ betrachtet rechts ein dünner zweiter Stab, während links ein kurzer nach hinten gebogener ‘Stachel vorragt. Die das Ostium seitlich begrenzenden Seitenteile der Peniswand sind außen an ihrem abgerundeten Ende mit einem kleinen Zahn bewehrt. Das Operculum ist rechteckig mit vorspringenden Hinterecken, es überragt die Seitenteile der Dorsalwand des Penis. Scydmagnus (s. st.) crockerensis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Crocker Range, 1350 m, km 60 Kinabalu. Tambunan, 17.5.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Durch breiten Kopf und annähernd gerundet- trapezförmigen Halsschild an Armatoscydmaenus erinnernd, aber durch bei d ungezähnte Hinterschenkel und anderen Penisbau davon verschieden. BESCHREIBUNG: Long. 1,50 mm, lat. 0,60 mm, Rotbraun, gelblich behaart. Kopf von oben betrachtet quer-fünfeckig mit parallelen Schläfen, diese doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis ein wenig überragend, ihr Basalglied eineinhalbmal so lang wie breit, die folgenden Glieder schmäler, 2 bis 6 deutlich gestreckt, 7 und 8 schwach quer, 9 eineinhalbmal, 10 doppelt so breit wie 8, das eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild so lang wie breit, im vorderen Drittel seiner Länge am breitesten, von da gerade zur Basis verengt, flach gewölbt, mit 2 Basalgrübchen, ziemlich dicht behaart. Flügeldecken oval, schon an der Basis zusammen breiter als die Halsschildbasis, mit seichter Basalimpression, fein punktiert und nach hinten gerichtet behaart. Beine mittellang, Schenkel mäßig verdickt, Hinterschenkel beim d ohne Zahn, Schienen gerade. Penis (Abb. 5) in der Anlage tonnenförmig, der Apex an der Basis verbreitert, dorsal wulstförmig vorgewölbt, seine Dorsalwand weit nach hinten vorgezogen und sichelförmig nach oben gebogen, das Ostium penis umfassend. In dieses ragt von vorne der Ductus ejaculatorius heraus, dessen Ende beinahe den Hinterrand des Penis erreicht. Er entspringt ungefähr in der Längsmitte des Peniskörpers in einer quergestellten, schwachen Erweiterung, eine Samenblase fehlt. Scydmaneus (s. str.) borneoensis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus £ Sabah, Poring Hot Springs, 500 m, 7.5.1987 (lg, Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch stark queren Kopf, isodiametrischen Halsschild und kurze Beine. BESCHREIBUNG: Long. 1,80 bis 2,00 mm, lat. 0,80 mm. Schwarz, Extremitäten rotbraun. Bräunlich behaart. Kopf von oben betrachtet breiter als lang, Schläfen zur Basis leicht konvergierend, spärlich mit abstehenden Haaren bedeckt. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied doppelt so lang wie breit, 2 bis 6 isodiametrisch, 7 und 8 breiter als lang, 9 doppelt so breit»wie 8, 10 noch etwas breiter, beide breiter als lang, das Endglied spitz -kegelförmig, nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild isodiametrisch, vor der Mitte am breitesten, seitlich abstehend, auf der Scheibe schütter und anliegend behaart, mit 2 Basalgrübchen. Flügeldecken kurzoval, schon an der Basis zusammen breiter als die Halsschildbasis, stark gewölbt, fein punktiert und abstehend behaart, Basalgrübchen und Schulterbeule fehlend. SCYDMAENIDAE VON SABAH 865 , 11 I 1 i ' i 1 1 ' i Wa 2 lu La 7 a = ABB. 9-16. 9: Scydmaenus brevitarsis (Schaufuss), Penis in Dorsalansicht. 10: Scydmaenus densepunctatus nov. spec., Penis in Dorsalansicht. 11: Scydmaenus (Mimoscydmaenus) crockeri nov. spec., Penis in Lateralansicht. 12: Scydmaenus (Mimoscydmaenus) complexipenis nov. spec., Penis in Dorso- lateralansicht. 13: Scydmaenus (Mascarensia) dissimilis nov. spec., Penis in Lateralansicht. 14: Scydmaenus (Eustemmoides) punctatus nov. spec., Penis in Dorsalansicht. 15: Scydmaenus (Eustem- moides) alessmetanai nov. spc., Penis in Dorsalansicht. 16: Scydmaenus (Eustemmoides) silvicola nov. spec., Penis in Dorsalansicht. Beine kurz und kräftig, Schenkel mäßig verdickt, Schienen gerade. Penis (Abb. 6) von oben betrachtet zylindrisch, sein Apex spitzwinkelig -dreieckig, die Basalöffnung dorsobasal gelegen, mit stark sklerotisiertem Rahmen, Ostium penis termi- nal gelegen, bis ins distale Drittel der Penislänge nach vorne reichend. Im Ostium penis ist der spiralig aufgerollte Ductus ejaculatorius sichtbar. Er entspringt in einem Trichter im 866 HERBERT FRANZ vorderen Drittel der Penislänge, von wo ein gerades Rohr bis zum Ostium führt, wo dieses ‘ spitzwinkelig in den Ductus ejaculatorius mündet. Von der Basalöffnung des Penis zieht ein weites Rohr distalwärts und mündet hinter dem Trichter in den Ductus. Scydmaenus borneoi nov. spec. MATERIAL: Holotypus & (Penispräparat), Sabah, Poring Hot Springs, 500 m, 8.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 1 Paratypus 9, 8.5.1987 (lg. Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch von oben betrachtet runden Kopf, schlanke Fühlergeißel und länglichen Halsschild. ohne Basalgrübchen. BESCHREIBUNG: Long. 1,80 mm, lat. 0,80 mm. Rotbraun, hellgrau behaart. Kopf von oben betrachtet kreisrund, klein, anliegend behaart. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied eineinhalbmal so lang wie breit, 2 bis 8 isodiametrisch bis leicht gestreckt, 7 und 8 nicht asymmetrisch, 9 um die Hälfte breiter als 8, 10 noch etwas breiter, beide schwach quer, das spitz-eiförmige Endglied so lang wie die beiden vorletzten zusammen. Halsschild um ein Sechstel länger als beit, seitlich gleichmäßig gerundet, dicht, nach hinten gerichtet behaart, ohne Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, seitlich gleichmäßig gerundet, ohne Schulterhöcker und ohne Basalimpression, lang, nach hinten gerichtet behaart. Beine kräftig, ziemlich kurz, Vordertarsen des d nicht verbreitert. Penis (Abb. 7) lang- gestreckt, schwach dorsalwärts gekrümmt, in einer Spitze endend. Seine Basalöffnung dorsobasal gelegen, darunter befindet sich ein scheibenförmiges Druckregulierungsorgan. Dahinter befindet sich in der vorderen Hälfte des Peniskörpers eine Reihe von 3 Kam- mern, an die der bis an das Penisende heranreichende Ductus ejaculatorius anschließt. Sein Ende ragt ein wenig aus dem Ostium penis nach oben heraus. Scydmaenus (s. str.) fraternus nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Sabah, Crocker Range, 1270 m, 60 km von Kota Kinabalu nach Tambunan, 17.5.1987 (lg. Burckhardt u. Löbl cMG); Paratypus 9 Sabah, Poring Hot Springs, 550 bis 600 m, 9.5.1987 (lg. Burckhardt u. Löbl, cF); ebenda, 31.8.1988 (lg. Smetana, cMG). DIAGNOSE: Mit S. tenuicornis Schauf. und allotenuicornis m. sehr nahe verwandt, die Fühler aber kürzer und dicker, ihre Keule scharf abgesetzt. Penis abweichend gebaut, auch dem Sc. paratenuicornis sehr ähnlich, aber bedeutend größer als dieser. Die Stellung der Art innerhalb der Untergattung Sydmaenus s. str. ist als provisorisch anzusehen. BESCHREIBUNG: Long. 1,20 bis 1,30 mm, lat. 0,50 bis 0,60 mm. Rotbraun, fein gelblich behaart. Kopf von oben betrachtet rundlich, etwas breiter als lang, die Schläfen doppelt so lang wie der Augendurchmesser. Füher schlank, zurückgelegt die Halsschildbasis um die beiden letzten Glieder überragend, ihr Basalglied zweieinhalbmal so lang wie breit, 2 und 5 eineinhalbmal so lang wie breit, 3, 4 und 6 leicht gestreckt, 7 und 8 schwach quer, 9 um ein Drittel breiter als 8, 10 noch’ein wenig breiter, beide um ein Drittel länger als breit, das spitz-eiförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild ein wenig länger als breit, seitlich gleichmäßig gerundet, nur so breit wie der Kopf, ziemlich dicht punktiert, ohne Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, stark gewölbt und seitlich stark gerundet, ohne Basalimpression und ohne Schulterbeule, seicht punktiert und nach hinten gerichtet behaart. Flügel verkümmert. SCYDMAENIDAE VON SABAH 867 Beine lang und schlank, Schenkel sehr schwach verdickt, Schienen gerade. Penis (Abb. 8) im Bau dem des Sc. tenuicornis und allotenuicornis sehr ähnlich, sein Apex gerade, kürzer und breiter als bei Sc. allotenuicornis, die aus dem Ostium penis herausragenden sklerotisierten Differenzierungen von der Vergleichsart abweichend gebaut. Subgenus Agathelor Schaufuss SCHAUFUSS (1884): Annali. Mus. civ. Genova (2) 1 XXI, p. 420-422. FRANZ (1984): Sber. Ost. Akad. Wiss., Math. nat. Kl. Abt. I, 193, p. 133-135. Schaufuss hat Agathelor als Gattung auf A. deplanatum Schauf. und brevitarse Schauf- begründet. Von beiden Arten befindet sich ein als Syntpus bezeichnetes Exemplar im Deutsch. Ent. Inst. in Eberwalde. Beide Ex. sind 9 £ und tragen Patriazettel mit dem Text “Borneo Sarawak 1865 — 66 coll G. Doria”, der mit der Fundortangabe in der Original- diagnose übereinstimmt. Ob im Museum von Genua weiteres Typenmaterial verwahrt wird, ist mir unbekannt, Schaufuss gibt in der Beschreibung an “In Mus. Civ. Jan.”. Ich habe (FrAnz 1984) nach den beiden in Eberswalde verwahrten Syntypen Neu- beschreibungen angefertigt und festgestellt, daß beide in das Genus Scydmaenus gehören, konnte aber damals nur vermuten, da mir kein d vorlag, daß beide in das Subgenus Armatoscydmaenus gehören. Nunmehr habe ich in der großen Ausbeute aus Sabah von Agathelor brevitarsis auch 3 4 auffinden können, die tatsächlich gezähnte Hinterschenkel besitzen, womit erwiesen ist, daß A. brevitarsis zu Armatoscydmaenus gehört. Dies gestattet es, für A. brevitarsis unter der Untergattung Armatoscydmaenus Schauf. eine Neubschreibung zu geben. Leider hat sich von Agathelor deplatum Schaufuss in der Ausbeute von Sabah kein weiteres Material gefunden, sodaß von dieser Art nach wie vor nur der im Deutschen Ent. Inst. verwahrte Syntypus (2) vorliegt. Dieses Tier weicht aber von den bekannten Armatoscydmaenus-Arten so weit ab, daß mit zeimlicher Sicherheit festgestellt werden kann, daß Agathelor deplanatum kein Armatoscydmaenus ist. Ich habe daher das Genus Agathelor Schauf für A. deplanatus Schaufuss als Typusart aufrechterhalten und begründe diese Entscheidung nachstehend. Scydmaenus (Agathelor) deplanatus Schaufuss SCHAUFUSS (1884): Annali. Mus. civ. Genova (2) 1 (XXD, p. 421 FRANZ (1984: Sber. Ost. Akad Wiss. Math. nat. Kl. Abt 1193, p. 134. Die zitierte Neubeschreibung ist durch einige ergänzende Daten zu vervollständigen. Halsschild und Flügeldecken sind ziemlich dicht und lang behaart, die Haare sind aber beim Umpräpariren an der Körperberfläche angeklebt und täuschen, da sie vorwiegend nach hinten gerichtet sind, eine längsrissige Skulptur vor. An einer Stelle sind die Haare aber abgeschabt, wodurch eine feine und schüttere Punktierung des Untergrundes sichtbar ist. An einer anderen Stelle sind die Haare quergestellt, was ebenfalls beweist, daß der Eindruck einer längsrissigen Struktur eine Täuschung ist. Agathelor deplanatus ist, wie schon der Autor durch die Namensgebung zum Ausdruck gebracht hat, durch eine außerordentlich flache Körperform ausgezeichnet. Dieses Merkmal unterscheidet ihn von den Arten des Subgenus Armatoscydmaenus. Ein weiteres Unterscheidungsmerkmal ist der vor der Basis ausgeschweifte Halsschild, wodurch die Hinterecken des Halsschildes scharf gewinkelt sind. Weiters sind die Beine außer- ordentlich kurz, wodurch allein schon die Entwicklung eines Zahnes an den Hinter- schenkeln unmöglich ist. Vor der Basis des Halsschildes stehen zwei sehr kleine, schwer 868 HERBERT FRANZ sichtbare Grübchen. Diese stehen nebeneinander vor dem Schildchen, nicht, wie bei den Armatoscydmaenus-Arten zur Seite gerückt. Alle diese Merkmale lassen erkennen, daß A. deplanatus nicht in das Subgenus Armatoscydmaenus gehört, sondern in ein eigenes Subgenus der Gattung Scydmaenus zu stellen ist. Subgenus Armatoscydmaenus Franz Scydmaenus (Armatoscydmaenus) brevitarsis (Schaufuss) SCHAUFUSS (1884): Annali. Mus. civ. Genova (2) 1 (XXI) p. 421-423 (Agathelor) FRANZ (1984): Sher. Ost. Akad. Wiss., Math. nat. KI. Abt. I, 193, p. 134. MATERIAL: Syntype 2 Sarawak (Deutsches Ent. Inst.); Borneo, Sabah, 43 (Penispräparat) 19, Kinabalu Nat. Park, Poring Hot Springs, 485 m, (lg. Smetana, CMG); ebenda 23 19 (Penispräparat, cF); Mount Kinabalu, 1500 m, 1790 m, 21.4.1987 (lg. Burckhardt u. Löbl, cMG), 38 19 (CMG), 26, cr). DIAGNOSE: Sehr ausgezeichnet durch dunkel rotbraune Färbung, beim d gezähnte Hinterschenkel und stark einwärts gebogene, mediodistal abrupt verbreiterte Hinter- schienen. BESCHREIBUNG: Long. 1,60 bis 2,20 mm, lat. 0,70 bis 0,80 mm. Dunkel rotbraun, fein und dicht gelblich behaart. Kopf von oben betrachtet gerundet-fünfeckig, breiter als lang, Augen groß, Schläfen ein wenig länger als der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied zweieinhalbmal so lang wie breit, 3 bis 6 isodiametrisch, 7 und 8 breiter als lang, nicht deutlich asymmetrisch, 9 doppelt so breit wie 8, 10 noch etwas breiter, das eiförmige Endglied kürzer als 9 und 10 zusammen. Halsschild flach gewölbt, so lang wie breit, im vorderen Drittel seiner Länge am breitesten, von da zur Basis geradlinig verengt, sehr fein punktiert und aufgerichtet behaart, mit Andeutung einer Schulterbeule und Basalimpression. Beine kurz, besonders die Tarsen sehr kurz, Hinterschenkel des d scharf gezähnt, Hinter- schienen des d medialwärts gebogen und distal stark verbreitert. Penis (Abb. 9) am apikalen Ende abgestutzt, nicht wie bei den bisher bekannten Arten des Subgenus Armatoscydmaenus zugespitzt und stark S-förmig gekrümmt. Im Inneren des Peniskörpers liegt vor dessen Längsmitte eine kapuzenförmige Blase, an die distal eine nierenförmige Blase anschließt. Diese mündet in den Ductus ejaculatorius, der an der Basis sehr stark erweitert ist. Der Ductus ejaculatorius reicht in der Ruhelage bis an das apikale Ende des Penis heran, ist dickwandig und sein Lumen überragt dünnwandig ein wenig den dickwandigen Teil. Neben dem Ductus liegt von oben und hinten betrachtet links ein großer Stachel, der das Penisende nicht erreicht. Das Ostium penis liegt dorsoapikal. Es reicht an den Seiten des Peniskörpers bis zu dessen distalem Viertel, weiter medial jedoch fast bis zur Längsmitte. Scydmaenus (Armatoscydmaenus) trapezicollis Lhoste LHOSTE (1938): Arb. morph. taxon. Ent. Berlin Dahlem 5, 124-125. fig. 18, 19. FRANZ (1984): Sber. Ost. Akad. Wiss. Math. nat. Kl. Abt I, 193, p. 135. Ich habe Sc. trapezicollis (l.c.) als fragliches Synonym zu Ag. deplanatum gestellt, was nach dem nunmehrigen Stand des Wissens unrichtig ist. Nach Lhoste ist A. trapezicollis mit 1,50 mm Körperlänge größer als Ag. deplanatum mit nur 1,25 mm. Auch sind die Beine bei Ag. deplanatum noch kürzer als nach Lostes Zeichnung bei trapezicollis. Die Halsschildseiten sind bei trapezicollis vor der Basis nicht ausgeschweift. SCYDMAENIDAE VON SABAH 222.07 Pers ooo { CES ABB. 17-24. 17: Scydmaenus (Eustemmoides) cuneipenis nov. spec., Penis in Dorsalansicht. 18: Scydmaenus (Eustemmoides) thermarum nov. spec., Penis in Dorsalansicht. 19: Scydmaenus (Eustemmoides) furcatus nov. spec., Penis in Dorsalansicht. 20: Scydmaenus (Eustemmoides) burckhardtloebli nov. spec., Penis in Dorsalansicht. 21: Scydmaenus (Eustemmoides) sabahi nov. spec., Penis in Dorsalansicht. 22: Scydmaenus (Eustemmoides) allosabahensis nov. spec., Penis in Lateralansicht. 23: Scydmaenus (Eustemmoides) parasabahensis nov. spec., Penis in Dorsalansicht. 24: Scydmaenus (Eustemmoides) sabahensis nov. spec., Penis in Dorsolateralansicht. 869 870 HERBERT FRANZ Der Penis ist bei der Lhoste-schen Art wie bei der armatus-regularis-Gruppe S-förmig _ gekrümmt, nicht wie bei Sc. brevitarsis und latipes nahezu gerade, wodurch trapezicollis der armatus-Gruppe näher steht als den auch noch durch die Verbreiterung der Hintertibien abweichenden Arten brevitarsis und latipes. Nach der von Lhoste gegebenen Abbildung ist zu erwarten, daß Sc. trapezicollis im männlichen Geschlecht gezähnte Hinterschenkel besitzt und daß dies von Lhoste übersehen worden ist. Scydmaenus (? Armatoscydmaenus) laticeps nov. spec. MATERIAL: Nur Holotypus 9, Sabah, Poring Hot Springs, 500 m, 8.5.1987 (lg. Burckhardt u. Löbl cMG). DIAGNOSE: Ausgezeichnet durch den sehr breiten Kopf, sehr flachen Körper und die kurzen Fühler und Beine, die die Art in das Subgenus Armatoscydmaenus verweisen. Da jedoch kein ¢ vorliegt, ist unbekannt, ob dieses gezähnte Hinterschenkel besitzt. BESCHREIBUNG: Long. 1,80 mm, lat. 0,60 mm, Rotbraun, gelblich behaart. Kopf von oben betrachtet gerundet-fünfeckig, um die Hälfte breiter als lang, die Schläfen 3 mal so lang wie der Durchmesser der kleinen Augen, Stirn und Scheitel flach gewölbt, fein aber dicht punktiert und behaart. Fühler zurückgelegt nur die Halsschildbasis erreichend, ihr Basalglied um die Hälfte länger als breit, 2 leicht gestreckt, 3 bis 8 breiter als lang, nur halb so lang wie die gesamten Fühler, 9 und 10 fast so lang wie breit und zusammen fast so lang wie das eiförmige Endglied. Halsschild am Vorderrand am breitesten und hier nur wenig breiter als der Kopf, seitlich gerundet zur Basis verengt, fein, abstehend behaart, sehr fein punktiert, mit 4 Basal- grübchen. Flügeldecken oval, flach gewölbt, deutlich und dicht punktiert, abstehend behaart, ohne Schulterbeule und Basalimpression, Schildchen erkennbar. Beine kurz, Schenkel schwach verdickt, Schienen, besonders die der Vorder- und Hinterbeine distal verbreitert. Die Art unterscheidet sich von den übrigen bekannten Aymatoscydmaenus-Arten durch den ungewöhnlich breiten Kopf, die kurzen Fühler und die von oben betrachtet breiten Schienen. Scydmaenus (Androscydmaenus) nov. subgen. In der Ausbeute von Sabah liegt ein Scydmaenus-3 vor, das durch den Bau des männlichen Kopulationsapparates von allen mir bekannten Scydmaenus-Arten so weit abweicht, daß es mit ihnen nicht in nähere verwandtschaftliche Beziehungen gebracht werden kann. In allen äußeren Merkmalen ist das Tier jedoch dem Genus Scydmaenus zugehörig. Es besitzt vor allem die für die Gattung Scydmaenus charakteristische Fühlerbildung, das distal ausgerandete 1. Fühlerglied, an dessen Basis die Fühler nach oben abgeknickt werden können, und das kleine, asymmetrisch gebaute 7. und 8. Fühlerglied. Der Kopf ist gerundet quer-fünfeckig mit flachen, seitlich nicht vorstehenden Augen. Es besitzt einen seitlich gleichmäßig gerundeten Halsschild und seitlich gerundete Flügeldecken ohne deutliche Basalimpression und Schulterbeule. Der männliche Kopulationsapparat weist jedoch nicht wie normal einen annähernd tonnenförmigen Peniskörper auf, sondern dieser teilt sich in der Mitte in 2 Äste, zwischen denen, weit vor dem Penisende, das Ostium penis liegt. Die beiden Seitenteile können nicht als Parameren gedeutet werden, da sie ihrer Anlage nach nie die Funktion von Tastorganen erfüllt haben können. SCYDMAENIDAE VON SABAH 871 Scydmaenus (Androscydmaenus) densepunctatus nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Sabah, Kinabalu Nat. Park, HQ. Livagu River, 1500 m, 17.5.1987 (lg. Smetana, cMG). DIAGNOSE: Außer durch den Bau des männlichen Kopulationsapparates durch dichte Punktierung des Halsschildes und der Flügeldecken sowie durch schlanke Fühler und Beine gekennzeichnet. BESCHREIBUNG: Long. 1,90 mm, lat. 0,70 mm, Rotbraun, hell bräunlich behaart. Kopf von oben betrachtet in der Anlage querrechteckig, die Stirn aber zwischen den Fühlerwurzeln stumpfwinkelig vorspringend, ihr Mittelteil glatt und glänzend, die Seitenteile punktiert. Augen groß, schwach vorgewölbt, Schläfen nur wenig länger als der Augendurchmesser. Fühler schlank, zurückgelegt die Halsschildbasis um die beiden letzten Glieder überragend, das Basalglied doppelt so lang wie breit, an der Spitze ausgerandet, Glied 3, 4 und 5 schmäler als 2, ebenfalls doppelt so lang wie breit, die Länge des 6. Gliedes seine Breite um die Hälfte übertreffend, 7 und 8 klein, asymmetrisch, eben merklich breiter als lang, 9 um die Hälfte breiter als 8, 10 eineinhalbmal so breit wie 9, das Endglied nicht ganz so lang wie 9 und 10 zusammen. Drittes Glied der Maxil- lartaster ebenso lang wie der Augendurchmesser, das Endglied nicht deutlich erkennbar. Halsschild um ein Fünftel länger als breit, seitlich gleichmäßig gerundet, dicht punktiert und anliegend behaart. Basalgrübchen von der Grundpunktur nicht unterscheidbar. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, ohne Schulterbeule und Basalimpression, seitlich gleichmäßig gerundet, dicht punktiert und anliegend behaart. Beine schlank, Schenkel keulenförmig verdickt, Schienen gerade, wie auch die Tarsen schlank. Penis (Abb. 10) dreiteilig, der mittlere Teil kürzer als die beiden Seitenteile, nur die Spitze des Ductus ejaculatorius weiter nach hinten vorragend. Er setzt sich im Inneren des Penis ein kurzes Stück nach vorne fort und erweitert sich dann zu einer runden Blase, vor der eine breit becherförmige Blase steht. In der basalen Hälfte des Penis sind die beiden Seitenteile mit dem Mittelteil des Penis verschmolzen, die Basalöffnung liegt dorsobasal und ist von einem breiten Sklerotinrahmen umgeben, der die ganze Breite der Penisbasis einnimmt. Auch die Seitenteile des Penis sind von einem Sklerotinrahmen umgeben. Dieser trennt durch eine Querleiste einen vorderen Abschnitt der Seitenteile von einem kleineren distalen, der über das apikale Ende des Mittelteils hinausragt. Die Querleisten tragen medial einen scharfen schräg zur Mitte und nach hinten gerichteten Zahn. Subgenus Mimoscydmaenus Franz FRANZ (1986): Revue suisse Zool. 93, p. 967-970. Das Subgenus Mimoscydmaenus wurde monotypisch auf Scydmaenus (Mimoscyd- maenus) chiangmaii Franz errichtet, der in Thailand und auf der Halbinsel Malakka verbreitet ist. Das Auftreten zweier weiteren Arten in NO-Borneo ist einer von einer Reihe von Beweisen, daß zwischen der Fauna des äußersten Südostens des asiatischen Festlandes und dem Nordosten von Borneo enge faunistische Beziehungen bestehen. Scydmaenus (Mimoscydmaenus) crockeri nov. spec. MATERIAL: Holotypus d ( Penispräparat), Sabah, Crocker Range, 1350 m, 60 km von Kota Kinabalu Tambunan, 13.5.1987 (lg. Burckhardt u. Löbl, cMG). 872 HERBERT FRANZ ns \ ano. 1/5mm 26 ABB. 25-32. 25: Scydmaenus (Eustemmoides) sabahanus nov. spec., Penis in Lateralansicht. 26: Scydmaenus (Eustemmoides) filipenis nov. spec., Penis in Lateralansicht. 27: Scydmaenus (Eustemmoides) parafilipenis nov. spec., Penis in Dorsolateralansicht. 28: Scydmaenus (Eustemmoides) allofilipenis nov. spec., Penis in Dorsalansicht. 29: Scydmaenus (Eustemmoides) bidentipenis nov. spec., Penis in Dorsalansicht. 30: Scydmaenus (Eustemmoides) frater nov. spec., Penis in Dorsalansicht. 31: Scydmaenus (Eustemmoides) poringensis nov. spec., Penis in Dorsalansicht. 32: Scydmaenus (Eustemmoides) livagui nov. spec., Penis in Dorsalansicht. SCYDMAENIDAE VON SABAH 873 DIAGNOSE: Dem Sc. chiangmaii äußerlich sehr ähnlich, im Bau des männlichen Kopulationsapparates aber stark abweichend. BESCHREIBUNG: Long. 1,80 mm, lat. 0,80 mm. Graubraun, die Extremitäten gelb- braun, grau behaart. Kopf von oben betrachtet fast Kreisrund, stark gewölbt, in der Mitte mit einer Längsfurche, mit kleinen, weit nach vorne gerückten Augen, Schläfen 3 mal so lang wie der Augendurchmesser. Fühler körperlang, fadenförmig, alle Glieder sehr langgestreckt. Halsschild eineinhalbmal so lang wie breit, zum Vorderrand viel stärker verengt als zur Basis, kaum breiter als der Kopf, mit 2 sehr kleinen Basalgrübchen, sehr fein staubartig behaart. Flügeldecken zusammengenommen schon an der Basis ein wenig breiter als der Halsschild, seitlich schwach gerundet, ohne Spur einer Basalimpression und ohne Schulterhöcker, äußerst fein punktiert und nach hinten gerichtet behaart. Flügel verkümmert. Beine sehr lang, Schenkel keulenförmig verdickt, Schienen gerade, wie auch die Tarsen sehr dünn. Penis (Abb. 11) gedrungen gebaut. Unter der Basalöffnung befindet sich ein Druck- ausgleichsorgan, das nicht wie meist bei den Vertretern der Gattung Scydmaenus pilzförmig sondern flach kappenförmig ist und schräg am dorsalen Ende eines dorsoventral verlaufenden Scklerotinbandes haftet. Von dem Druckregulierungsorgan zieht ein Bündel von Muskelsträngen, die sich annähernd in der Penismitte befinden, zu einer zweikammerigen Blase. Die basale dieser Blasen ist kleiner und schräggestellt, sie sitzt einer größeren nierenförmigen Blase auf, die ihrerseits in den nicht genau im Präparat erkennbaren Ductus ejaculatorius mündet. Das äußerste Ende des Ductus ejaculatorius ist am apilaken Rand des Penis sichtbar. Dorsal davon steht ein starker Stachel nach oben und hinten ab. Scydmaenus (Mimoscydmaenus) complexipenis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Poring Hot Springs, 500 m, 7.5.1987 (Ig. Burckhardt u. Löbl, cMG); 1 Paratypus ebenda (cF); 1 Paratypus $ ebenda, 480 m, 29.8.1988 (lg. Smetana, cMG). DIAGNOSE: Dem Sc. crockeri sehr ähnlich, aber der Körper etwas breiter, die Flügel- decken mit deutlicher Schulterbeule, die Beine kürzer, der Penis anders geformt. BESCHREIBUNG: Long. 1,60 bis 1,70 mm, rotbraun, schütter gelblich behaart. Kopf von oben betrachtet fast kreisrund, mit einem Längseindruck auf der Stirn, die Schläfen dreimal so lang wie die kleinen, weit vorne stehenden Augen. Fühler körperlang, fadenförmig, alle Glieder sehr lang und dünn. Halsschild um ein Viertel länger als breit, nur wenig breiter als der Kopf, zum Vorderand stärker als zur Basis verengt, ohne Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis mit deutlicher Schulterbeule, deutlich punktiert und abstehend behaart. Flügel verkümmert. Beine langgestreckt und schlank, aber etwas kürzer als bei Sc. crockeri, Schenkel keulenförmig verdickt, Schienen und Tarsen sehr schlank. Penis (Abb. 12) asymmetrisch mit dorsolateral gelegener, von einem stark sklerotisierten Rahmen umfaßter Basalöffnung und dorsoapikalem Ostium penis. Im Penisinneren liegt knapp vor dessen Längsmitte die kugelige Samenblase, die distal einen weiten Ausführungsgang besitzt. Dieser ist durch einen stark sklerotisierten Querbalken fixiert, an dessen distalem Rand 2 dicke Schläuche entspringen. Von diesen ist der von oben und hinten besehen linke dünner und S-förmig gekrümmt. Er reicht bis nahe an das Penisende 874 HERBERT FRANZ heran und trägt terminal eine lange Geißel. Der rechte ist dicker und basalwärts umgebogen. An der Biegungsstelle entspringt ein kurzes Rohr, aus dem ein sichelförmig gekrümmter, blind endender Schlauch nach hinten abzweigt. An seinem Hinterende inseriert ein schwanzförmiger Fortsatz. Subgenus Mascarensia Franz Scydmaenus (Mascarensia) dissimilis nov. spec. MATERIAL: Nur Holotypus d Sabah, Poring Hot Springs, 500 bis 600 m, (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Dem Sc. similis Schaufuss ähnlich, aber etwas kleiner, die Fühler länger und schlanker, der Halsschild mit 2 Basalgrübchen, die Flügeldecken kürzer, der Penis abweichend gebaut. BESCHREIBUNG: Long. 1,10 mm, lat. 0,50 mm, bräunlichgelb, sehr fein behaart. Kopf von oben betrachtet gerundet-fünfeckig, breiter als lang, die Schläfen nahezu parallel, fast doppelt so lang wie der Augendurchmesser. Stirn und Scheitel flach gewölbt, glatt und glänzend, kahl, Fühler zurückgelegt die Halsschildbasis ein wenig überragend, ihre beiden ersten Glieder knapp doppelt so lang wie breit, 3 bis 5 deutlich gestreckt, 6 klein, länger als breit, 7 und 8 asymmetrisch, schwach quer, 9 doppelt so breit wie 8, 10 noch etwas breiter, beide eben merklich gestreckt, das spitz-eiförmge Endglied so lang wie die beiden vorletzten zusammen. Halsschild kaum merklich länger als breit, sehr wenig breiter als der Kopf, etwas vor seiner Längsmitte am breitesten, sehr fein punktiert (80-fache Vergrößerung), mit 2 Basalgrübchen. Flügeldecken zusammen an der Basis nur wenig breiter als die Halsschildbasis, kurzoval, nur so lang wie der Kopf und Halsschild zusammen, fein punktiert, mit schwer erkenn- barer Behaarung, ohne Basalimpression und ohne Schulterbeule. Flügel entwickelt. Beine schlank, Hinterschenkel des d mit einer Grube versehen. Penis (Abb. 13) langgestreckt, in seiner Längsmitte stark eingeschnürt, hinter der Ein- schnürung liegt dorsal das große Ostium penis. In diesem befindet sich eine annähernd isodiametrische, an der Basis in ihrer ganzen Breite konkav ausgeschnittene Blase, an die distal ein breiter Gang anschließt, der nach kurzem Verlauf von oben und hinten betrachtet nach rechts biegt, sich dabei verschmälert und an der Außenseite der Biegung mit Zähnchen besetzt ist. Subgenus Eustemmoides Franz FRANZ (1957); Koleopt. Rasch. 35, p. 3-6. Das Subgenus Eustemmoides habe ich auf E. alluvialis aus dem Tschadgebiet im subtropischen Afrika errichtet, es ist in der äthiopischen Region und darüber hinaus in den warmen Klimagebieten der Erde weiter verbreitet. Die Fauna von Sabah enthält, wie die Bearbeitung des mir von dort vorliegenden Materiales zeigt, eine ganz außerordemtliche Vielfald von Arten dieses Subgenus. Scydmaenus (Eustemmoides) punctatus nov. spec. MATERIAL: Holotypus & (Penispräparat), Sabah, Poring Hot Springs, 600 m, bei Bat Cave, 10.5.1987 (lg. Burckhardt u. Löbl, cGM); Paratypus d (Penispräparat), Sabah, Poring Hot Springs, 500 m, 10.5.1987 (1g-Burckhardt u. Löbl, cF)); ebenda 500 m, 6.5.1987 (1g. Burckhardt u. Löbl, cMG). SCYDMAENIDAE VON SABAH 875 DIAGNOSE: Gekennzeichnet durch die deutliche Punktierung des Halsschildes und der Flügeldecken. BESCHREIBUNG: Long. 1,20 bis 1,30 mm. lat 0,60 mm. Rotbraun, gelblich behaart. Kopf von oben betrachtet gerundet-fünfeckig, breiter als lang, die Schläfen leicht zur Basis konvergierend, doppelt so lang wie der Augendurchmesser, Führer zurückgelegt die Halsschildbasis knapp erreichend, doppelt so lang wie der Augendurchmesser; ihr Basalglied annähernd so lang wie breit, 3 bis 5 leicht gestreckt, 6 bis 8 breiter als lang, 9 und 10 quadratisch, 9 nicht ganz, 10 reichlich doppelt so breit wie 8, das spitz-eiförmige Endglied knapp so lang wie 9 und 10 zusammen. Halsschild eben mehrlich länger als breit, etwas vor seiner Längsmitte am breitesten, an der Basis nicht ganz so breit wie die Flügeldecken zusammengenommen, mit 2 kleinen Basalgrübchen, zwischen der deutlichen Punktierung nur schwer sichtbar behaart. Flügeldecken schon an der Basis zusammen wesentlich breiter als die Hasschildbasis, ohne Schulterhöcker und nur mit Andeutung einer Basalimpression, deutlich und ziemlich dicht punktiert und ziemlich eng anliegend behaart. Flügel entwickelt. Penis (Abb. 14) von oben betrachtet dreimal so lang wie breit, in der Längsmitte etwas schmäler als an den beiden Enden. Seine Basalöffnung dorsobasal gelegen, mit stärker sklerotisierter Umrandung, dahinter mit pilzhutförmigem Druckregulierungsorgan. In der Längsmitte des Penis befindet sich ein von oben besehen kreisförmig abgegrenzter Raum, in dem eine große, distal mit einem Ausführungsgang versehene Blase gelegen ist. Es ist nicht erkennbar, ob der Ausführungsgang distal einen Ausgang besitzt. Der distale Teil des Penis ist zweiteilig. Er besteht aus 2 in der Sagittalebene zusammenstoßenden Lappen, die nur laterobasal mit dem Peniskörper verbunden sind. Jeder dieser Lappen ist durch einen tiefer Einschnitt in 2 Teile gespalten, deren vorderer medialwärts zu einem Sklerotinstab verschmälert ist. Die beiden Stäbe berühren einander in der Sagittalebene, vor der Berührungstelle der distalen Lappen. Sie scheinen bei der Kopula ausein- andergespreizt werden zu können. Scydmaeus (Eustemmoides) alessmetanai nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Kinabalu Nat. Park. Poring Hot Springs, 485 m, 27.8.1988 (lg. Smetana, cMG). DIAGNOSE: Gekennzeichnet durch stark gewölbten Körper, nahezu kahle Körperoberfläche und starken Glanz. BESCHREIBUNG: Long. 1,50 mm, lat. 0,60 mm. Kastanienbraun, oberseits fast kahl. Kopf klein, von oben betrachtet gerundet-rautenförmig, so lang wie breit, schütter abstehend behaart, Augen klein, flach. Fühler zurückgelegt die Halsschildbasis knapp erreichend, ihr Basalglied doppelt so lang wie breit, 3 bis 6 sehr wenig länger als breit, 7 und 8 breiter als lang, 9 um ein Drittel breiter als 8, um ein Fünftel länger als breit, 10 so lang wie 9, zur Spitze verbreitert und da doppelt so breit wie das vorhergehende, das spitz- eiförmige Endglied etwas kürzer als die beiden vorletzten zusammen. Halsschild um die Hälfte länger als breit, etwas breiter als der Kopf mit den Augen, ungleichmäßig gerundet, schütter, anliegend behaart, ohne Basalgrübchen. Flügeldecken an der Basis zusammen nur so breit wie die Halsschildbasis, stark gewölbt, aber seitlich nur schwach gerundet, sehr schütter und eng anliegend behaart, ohne Schulterbeule und ohne Basalimpression, Flügel entwickelt. Beine ziemlich kurz, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 15) von oben betrachtet in der Anlage lang-rechteckig, mit scharf abgesetztem, schmalem Apex. Basalöffnung dorsobasal gelegen, sehr stark quer, mit 876 HERBERT FRANZ breitem, stark sklerotisiertem Rahmen. Ostium penis apikal in breitem Bogen aus der Dorsalwand des Penis ausgeschnitten, von dem langen schmalen Apex weit überragt. Der Hinterrand des Peniskörpers seitlich weit über die Basis des Apex nach hinten ragend. Die Hinterecken des Peniskörpers spitz. Hinter der Basalöffnung des Penis liegen hintereinander 2 Kammern, an die distal ein dickes Rohr anschließt. Dieses gabelt sich beim Austritt aus dem Ostium, der von oben und hinten besehen linke Ast setzt sich im Ostium leicht verdickt mit einem Knick zur Mitte fort und reicht in der Ruhelage bis zur Längsmitte des Apex penis. Der rechte Ast ist kürzer und ebenfalls beim Austritt in das Ostium verdickt. Ganz recht liegt ein langer Stachel, der von der Basis des Ostiums bis zum Ende des Apex reicht. Scydmaenus (Eustemmoides) silvicola nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Poring Hot Springs, nr Bat Cave, 10.5.1987 (lg. Burckhardt u. Löbl, cMG); Kinabalu Nat. Park, Poring Hot Springs, 485 m, 29.8.1988 (lg. Smetana) 1 Paratypus (CMG und 1 Paratypus (cF)). DIAGNOSE: Durch geringe Größe, spärliche Behaarung und durch das im Verhältnis zum 10. bedeutend kleinere 9. Fühlerglied ausgezeichnet. BESCHREIBUNG: Long. 1,10 mm, lat 0,50 mm, kastanienbraun, sehr fein hellbraun behaart. Kopf von oben betrachtet gerundet-fünfeckig, wenig breiter als lang, die Schläfen parallel, eineinhalbmal so lang wie der Durchmesser der flachen Augen. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied knapp doppelt, das 2. eineinhalbmal so lang wie breit, 3 bis 6 annähernd so lang wie breit, 7 und 8 quer, 9 um die Hälfte breiter als 8, 10 doppelt so breit, beide kugelig, nicht ganz so lang wie breit, das gerundet-kegelförmige Endglied nicht ganz so lang wie die beiden vorhergehenden zusammen. Halsschild leicht gestreckt, seitlich gleichmäßig gerundet, stark gewölbt, glatt und glänzend, mit 2 Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit schwach angedeuteter Schulterbeule, sehr fein punktiert und fein anliegend behaart. Beine kurz, Vorderschenkel stärker verdickt als die der beiden anderen Beinpaare. Penis (Abb. 16) langgestreckt, seine basale Hälfte breiter als die distale. Die Basalöffnung ist sehr stark quer, dorsobasal gelegen, Ostium penis dorsoapikal. Hinter der Basalöffnung liegt im Penisinneren ein pilzförmiges Druckregulierungsorgan. Hinter der Längsmitte des Penis liegen im Penisinneren hintereinander 3 Blasen, von denen die vorderste gerundet- dreieckig ist und basalwärts eine langgestreckte “Haube” trägt. Die 2. Blase ist kugelförmig und starkwandig. Sie ist doppelt so breit wie die 3., die ebenfalls kugelig und starkwandig ist. Auf der von oben und hinten besehen linken Seite der 3 Blasen befindet sich ein breiter Stachel. Der Apex penis ist am Hinterende breit abgerundet. Scydmaenus (Eustemmoides) cuneipenis nov. spec. MATERIAL: Holotypus d (Penispräparat) Sabah, Mount Kinabalu HQ at Livagu River, 1500 m, 16.5.1987 (lg. Smetana, cMG). DIAGNOSE: Durch schlanke Gestalt und lange Fühler einem Mimoscydmaenus ähnlich, von diesem Subgenus aber durch den Besitz einer zwar schlanken, aber deutlich abgesetzten Fühlerkeule und durch einfach keilförmig gebauten Penis verschieden. BESCHREIBUNG: Long. 1,65 mm, lat. 0,65 mm. Rotbraun, gelblich behaart. Kopf von oben betrachtet gerundet-fünfeckig, die fast parallelen Schläfen eineinhalbmal so lang wie der Augendurchmesser. Fühler zurückgelegt die Mitte der Flügeldecken 877 SCYDMAENIDAE VON SABAH ABB. 33-37. spec. Penis in Dorsalansicht. 36: 33: Loeblites sabahensis nov. spec., Penis in Dorsalansicht. 34: Horaeomorphus loeblianus nov. Syndicomorphus magnus, nov. spec., Penis in Lateralansicht. spec., Penis in Dorsalansicht. 35: Syndicus kinabalui nov. 878 HERBERT FRANZ erreichend, alle Fühlerglieder mit Ausnahme des 7. und 8. gestreckt, diese so lang wie breit, 9 bis 11 die deutlich abgesetzte Keule bildend. Halsschild um etwa ein Fünftel länger als breit, seitlich gleichmäßig gerundet, fein punktiert und behaart. Flügeldecken zusammen breiter als die Halsschildbasis, ohne deutliche Schulterbeule und Basalimpression, fein punktiert und schräg abstehend behaart. Penis (Abb. 17) lang keilförmig mit dorsolateral gelegener Basalöffnung. Im Penisinneren liegt in der Längsmitte eine Folge von 3 Blasen, an die distal ein Ausführungsgang anschließt, der im distalen Drittel des Penis endet. Der Ausführungsgang ist dreigegliedert: auf ein kurzes basales Stück folgt ein verkehrt L-förmiger Abschnitt, zuletzt wieder ein gerades Stück, das aber schmäler ist als das basale. Der Winkel des L wird durch ein dünnes Rohr überbrückt. Scydmaenus (Eustemmoides) thermarum nov. spec. MATERIAL: Nur Holotypus & (Penispräparat) Sabah, Poring Hot Springs, 500 m, 7.5.1987 (lg Burckhardt u. Löbl, CMG). DIAGNOSE: Gekennzeichnet durch hell rotbraune Färbung, schlanke, aber hoch- gewölbte Gestalt, gestreckten Halsschild mit 2 kleinen Basalgrübchen und dicht punktierte Flügeldecken. BESCHREIBUNG: Long. 1,80 mm, lat. 0,65 mm. Hell rotbraun, gelblich behaart. Kopf von oben betrachtet nur wenig breiter als lang, die parallelen Schläfen nicht ganz doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis knapp erreichend, ihr Basalglied zweieinhalbmal so lang wie breit, 2 leicht gestreckt, 3 bis 6 isodiametrisch, 7 und 8 breiter als lang, 9 um die Hälfte breiter als 8, wie auch 10 so lang wie breit. Das spitz-eiförmige Endglied kürzer als die beiden vorletzten zusammen. Halsschild um knapp ein Fünftel länger als breit, nur wenig breiter als der Kopf, vor der Längsmitte am breitesten, fein, nach hinten gerichtet behaart, mit 2 seichten Basal- grübchen. Flügeldecken länglichoval, seitlich flach gerundet, zusammen nur wenig breiter als die Halsschildbasis, dicht punktiert, schräg nach hinten gerichtet behaart. Flügel verkümmert. Beine mittellang, Schenkel schwach keulenförmig verdickt, Schienen gerade. Penis (Abb. 18) in dem einzigen Präparat immatur, langgestreckt, der Apex spitzwinkelig- dreieckig. Hinter der dorsobasal gelegenen Basalöffnung liegt im Penisinneren eine kleine, stark sklerotisierte Blase in einem viereckigen Rahmen. An sie schließt distal der dicke S- förmig gekrümmte Ductus ejaculatorius an. Er verläuft zunächst sagittal, biegt dann in der Mitte des Penis im rechten Winkel, von hinten und oben betrachtet nach rechts und läuft dann in einem Zweidrittelkreis wieder nach links, wobei sich das Rohr allmählich verengt. In dem scharfen Winkel entspringt ein Sklerotinstachel, der zunächst nach rechts gerichtet ist, um dann im stumpfen Winkel nach hinten zu verlaufen. Scydmaenus (Eustemmoides) furcatus nov. spec. MATERIAL: Es liegen insgesamt 17 Exemplare der Art vor, wovon 11 einschließlich des Holotypus d (Penispräparat) von Sabah, Kinabalu Nat. Park, Poring Hot Springs, 485 m, 28.8.1988 (lg. Smetana) in cMG und 6 Exemplare vom selben Fundort (2 Penispräparate) in cF aufbewahrt werden. DIAGNOSE: Dem an späterer Stelle beschriebenen Sc. sabahensis ähnlich, aber kleiner und schlanker, der Kopf fast so lang wie breit, der Halsschild länger als breit und der Penis ganz anders gebaut. SCYDMAENIDAE VON SABAH 879 BESCHREIBUNG: Long. 1,20 mm, lat. 0,55 m. Dunkel rotbraun, gelblich behaart. Kopf von oben betrachtet nur wenig breiter als lang, die Schläfen parallel, doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit sehr scharf abgesetzter Keule, ihr Basalglied doppelt so lang wie breit, 2 leicht gestreckt, 3 bis 6 annähernd quadratisch, 7 und 8 breiter als lang, 9 bis 11 die scharf abgesetzte Keule bildend, 9 und 10 etwas breiter als lang, das spitz-eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild länger als breit, nicht breiter als der Kopf, an der Basis fast so breit wie die Basis der Flügeldecken gemeinsam, mit 2 kleinen Basalgrübchen, sehr fein punktiert und behaart. Flügel voll entwickelt. Beine schlank, Vordertarsen des d nicht verdickt, Schienen gerade. Penis (Abb. 19) 3 mal so lang wie breit, sein Apex spitzwinkelig - dreieckig, seine Spitze schmal aufgebogen. Im Penisinneren befindet sich knapp vor der Penismitte eine Blase, die einem kurzen Rohrstutzen aufsitzt. Dahinter folgt beiderseits der Sagittalebene je ein Sklerotinstab, beide Stäbe sitzen distal einem sklerotisierten Querbalken auf. Hinter diesem Balken folgt das Ostium penis, in das von links ein Chitinstab hineinragt, der sich an der Spitze in eine zweispitzige Gabel verzweigt. Die beiden Spitzen sind nach vorne umgebogen. Scydmaenus (Eustemmoides) burckhardtloebli nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Poring Hot Springs, 500 m, 7.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 484 m, 2 Paratypen, 28.8.1988 (lg. Smetama, cMG); 2 Paratypen ebenda, 20.8.1988 (lg. Smetama, cF). Dragnose: AuBerlich dem später beschriebenen Sc. allofilipenis ähnlich, aber noch etwas schlanker, der Penis anders geformt. BESCHREIBUNG: Long. 1,00 mm, lat. 0,35 bis 0,40 mm. Dunkel rotbraun, sehr fein gelblich behaart. Kopf von oben betrachtet so lang wie breit, rundlich, Fühler zurückgelegt die Hals- schildbasis knapp erreichend, ihr Basalglied nur um ein Viertel länger als breit, 2 doppelt so lang wie breit, 3 bis 6 isodiametrisch, klein, 7 und 8 stark quer, 9 um die Hälfte breiter als 3, breiter als lang, 10 viel größer, um die Hälfte breiter als 9, quadratisch, das spitz- eiförmige Endglied etwas länger als die beiden vorletzten zusammen. Halsschild länger als breit, ein wenig breiter als der Kopf, schwach gleichmäßig gerundet, anliegend behaart, mit 2 Basalgrübchen. Flügeldecken sehr regelmäßig länglich-oval, an der Basis zusammen nur so breit wie die Halsschildbasis, glatt und glänzend, ohne Basalimpression und ohne Spur eines Humeralhöckers. Flügel verkümmert. Beine zimlich lang, Schenkel mäßig verdickt, Schienen gerade. Penis (Abb. 20) langgestreckt, sein Apex sehr spitzwinkelig. Das Ostium penis bis zum apikalen Ende reichend. Die Basalöffnung dorsobasal gelegen, knapp dahinter liegt ein pilzförmiges Druckregulierungsorgan und dahinter in der Dorsalwand ein durchsichtiges gerundet-viereckiges Fenster. Distal der Penismitte liegen im Penisinneren zwei langgestreckte stark sklerotisierte Körper, deren einer am Hinterrand einen zur Seite gerichteten Zahn trägt. Scydmaenus (Eustemmoides) sabahi nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Mount Kinabalu Nat. Park, HQ at Liwagu- River, 1500 m, 17.5.1987 (lg. Smetana, cMG); 3 Paratypen Mount Kinabalu (lg. Burckhardt u. Löbl, cMG); 2 Paratypen, Crocker Range, 1370-1500 m, (lg. Burckhardt u. Löbl, cMG); Kinabalu, HQ 880 HERBERT FRANZ Bukit Ular Trail, 1700 m, 29.4.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 1 Paratypus (cF); 3 ‘ Paratypen, Kinabalu 1500 bis 1650 m und 2600 m (lg. Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch breiten Kopf, relativ lange Fühler und schlanke Beine sowie deutliche Punktierung von Halsschild und Flügeldecken. BESCHREIBUNG: Long. 1,40 bis 1,80 mm, lat. 0,60 mm. Kastanienbraun, die Extre- mitäten heller, bräunlich behaart. Kopf groß, um ein Viertel breiter als lang, fast so breit wie der Halsschild, mit schwach gerundeten, kurz und fein behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis um das Endglied überragend, ihr Basalglied 3 mal so lang wie breit, 2,3 und 5 doppelt, 4 und 6 eineinhalbmal so lang wie breit, 7 und 8 breiter als lang, 9 und 10 leicht gestreckt, das spitz-eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild seitlich stark gerundet, fast so breit wie lang, fein punktiert und anliegend behaart, ohne Basalgrübchen. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, stark gewölbt und seitlich gerundet, ohne Basalimpression und ohne Schulterbeule, kräftig punktiert, fein und anliegend behaart, ohne Basalgrübchen. Beine lang, Schenkel keulenförmig verdickt, Schienen gerade, Vordertarsen des 4 nicht verbreitert. Penis (Abb. 21) ziemlich langgestreckt, sein Apex am Ende abgerundet-dreieckig, vom Peniskörper nur schwach abgesetzt. Die Basalöffnung dorsobasal gelegen, knapp dahinter liegt ein pilzförmiges Druckregulierungsorgan, von dem ein Bündel von Muskelsträngen zu zwei hintereinander liegenden Blasen zieht. Von diesen ist die basale quer kappen- förmig, die distale lang eiförmig. Die letztere ragt in der Ruhelage ein wenig in das langovale Ostium hinein, das bis zum Penisende reicht. An seinem Vorderrand liegt ein quergestellter Sklerotinkomplex, der nach hinten und links außen einen starken Zahn entsendet. Rechts davon durchsetzt diesen Komplex der mit der langeiförmigen Blase kommunizierende Ductus ejaculatorius. Er verschmälert sich distalwärts zu einer engen Düse und reicht bis zur Penisspitze. Scydmaenus (Eustemmoides) allosabahensis nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Mount Kinabalu Nat. Park, Poring Hot Springs, 485 m, 28.8.1988 (Ig. Smetana, cMG). DIAGNOSE: Wesentlich kleiner und weniger robust als Sc. sabahi. BESCHREIBUNG: Long. 1,20 mm, lat. 0,60 mm. Kastanienbraun, gelblich behaart. Kopf von oben betrachtet etwas breiter als lang mit parallelen Schläfen, so breit wie der Halsschild. Fühler zurückgelegt die Halsschildbasis knapp überragend, ihr Basalglied zweieinhalbmal, das 2. doppelt so lang wie breit, 3, 4 und 6 leicht gestreckt, 5 um ein Viertel länger als breit, 7 und 8 breiter als lang, nicht asymmetrisch, 9 und 10 quer, eng an die vorhergehenden anschließend, das eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild isodiametrisch, seitlich stark gerundet, vor seiner Längsmitte am breitesten, vor der Basis mit 2 großen Grübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit seichter Basalimpression, sehr fein punktiert und fein, abstehend behaart. Flügel entwickelt. Beine ziemlich lang, Schienen gerade, Vordertarsen des d leicht verbreitert. Penis (Abb. 22) langgestreckt, seine Basis bei seitlicher Betrachtung durch eine Ein- schnürung von der Dorsalseite hinter der Basalöffnung deutlich abgesetzt. In der distalen Hälfte des Peniskörpers befindet sich der Genitalapparat, bestehend aus 2 hintereinander liegenden Blasen und aus dem Ductus ejaculatorius. Die basale Blase ist gerundet- SCYDMAENIDAE VON SABAH 881 kegelförmig und besitzt in der distalen Hälfte ein Lumen, das ventral weiter nach vorne reicht als dorsal. Die distale Blase ist so breit wie lang, in seitlicher Ansicht quadratisch und besitzt eine verdickte Wand. Der Ductus ejaculatorius ist von einer dicken Sklerotinhülle ummantelt, er reicht bis nahe an das abgerundete Penisende. Scydmaenus (Eustemmoides) parasabahensis nov. spec. MATERIAL: Holotypus (Penispräparat) und 3 Paratypen, Kinabalu Nat. Park, Poring Hot Springs, 500 bis 600 m, 9.5.1987 (lg. Smetana, c MG); ebenda 3 Paratypen, (4 Penispräparat) (1g. Smetana, cF). DIAGNOSE: Dem Sc. sabahanus nahe stehend, aber robuster, der Halsschild länger als breit, die Flügeldecken länger oval, die Beine kräftiger, der Penis ganz anders gebaut. BESCHREIBUNG: Long. 1,65, lat. 0,65 mm. Dunkel rotbraun, gelblich behaart. Kopf von oben betrachtet wenig breiter als lang mit parallelen Schläfen. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied 3 mal, 3 bis 6 eineinhalbmal so lang wie breit, 7 und 8 etwas breiter als lang, 9 und 10 doppelt so breit wie 8, schwach quer, das eiförmige Endglied wenig kürzer als die beiden vorletzten zusammen. Halsschild um ein Viertel länger als breit, wenig vor seiner Längsmitte am breitesten, zum Vorderrand nur wenig stärker verengt als zur Basis, mit 2 kleinen Basalgrübchen, anliegend behaart, fast glatt. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, seitlich gleichmäßig gerundet, fein punktiert, nur mit Andeutung einer Schulterbeule. Flügel verkümmert. Beine kurz und kräftig, Schenkel mäßig verdickt, Schienen fast gerade. Penis (Abb. 23) in den basalen 4 Fünfteln von oben betrachtet fast parallelseitig, im distalen Fünftel dreieckig zur Spitze verengt. Ostium penis die distale Hälfte der Dorsalseite des Penis einnehmend, ein mächtiger sklerotisierter Zahn entspringt an seinem Basalrand und verbreitert sich hinter diesem auf 2 Drittel der Penisbreite. Er reicht bis über die Basis des Apex penis nach hinten. In seiner Längsmitte tritt auf seiner medialen Seite der dünne Ductus ejaculatorius aus. Er verläuft nach einer S-förmigen Krümmung gerade zur von oben und hinten betrachtet rechten Seite des Apex und überragt dessen Rand um ein bedeutendes Stück. Scydmaenus (Eustemmoides) sabahensis nov. spec. MATERIAL: 25 Ex., Sabah, Kinabalu Nat. Park, Holotypus d (Penispräparat) Poring Hot Springs und Langanan River (lg. Burckhardt u. Löbl, sowie Smetana, CMG); Langanan River, 850 m (lg. Smetana, CMG); 18 Ex., Poring Hot Springs, cF). BESCHREIBUNG: Long. 1,40, lat. 0,65. Dunkel rotbraun, gelblich behaart. Kopf etwas breiter als lang, Schläfen wenig länger als der Augendurchmesser, zur Basis konvergierend, Fühler mit scharf abgesetzter Keule, ihr Basalglied dicker als die folgenden, um ein Drittel länger als breit, 2 bis 6 annähernd isodiametrisch, 7 und 8 breiter als lang, 9 mehr als doppelt so breit wie 8, isodiametrisch, 10 etwas breiter als 9, schwach quer, das eiförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild so lang wie breit, vor der Mitte am breitesten, zum Vorderrand viel stärker verengt als zur Basis, abstehend behaart, ohne Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, fein punktiert und abstehend behaart, ohne Basalimpression und ohne Schulterbeule. Beine kräftig und mäßig lang. 882 HERBERT FRANZ (D Wiis At 2 \ \ i i 1 hi 39 5 ABB. 37-40. 37: Borneosabahia mirifica nov. spec. a bis d Penis in verschiedenen Stellungen (Erläuterungen im Text). 38: Euconnus (s. str.) pseudosukhotanus nov. spec., Penis in Dorsalansicht. 39: Euconnus (s. str.) allosukhotanus nov. spec., Penis a) in Drosal - b) in Lateralansicht. 40: Euconnus (s. str.) kina- baluanus nov. spec., Penis in Dorsalansicht. SCYDMAENIDAE VON SABAH 883 Penis (Abb. 24) zweieinhalbmal so lang wie breit mit dorsoapikal gelegenem Ostium, an dessen Basis eine querovale, vom eigentlichen Ostium durch einen Querbalken getrennte Öffnung liegt. Von dem Querbalken zweigt an seinem von oben und hinten betrachtet rechten Ende ein dicker Zahn im rechten Winkel nach hinten ab. Vom linken Rand des Ostiums ragen 2 spitze Stachel in sein Inneres vor. Unter diesen ist das Operculum sichtbar, das vor der rechten Hinterecke tief kreisförmig ausgeschnitten ist. In diesem Ausschnitt ist ein sehr langer gerade nach hinten gerichteter Stachel sichtbar. Scydmaenus (Eustemmoides) sabahanus nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Sabah, Mount Kinabalu, 1550 bis 1600 m, 24.4.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Äußerlich dem Sc. sabahi sehr ähnlich und von ihm nur durch den Bau des männlichen Genitalapparates sicher unterscheidbar. BESCHREIBUNG: Hinsichtlich der äußeren Merkmale kann auf Sc. sabahi m. verwiesen werden. Penis (Abb. 25) langestreckt, sein Basalteil vom übrigen Peniskörper etwas abgesetzt. Dieser ist bei lateraler Betrachtung im flachen Bogen nach oben gekrümmt. In seinem Inneren liegt eine große, schwach sklerotisierte nierenförmige Blase, an die ein schwach S-förmig gekrümmter Ausführungsgang anschließt. Dieser ragt mit seinem Hinterende ein wenig über den Apex penis hinaus und entsendet kurz davor einen kurzen Schlauch dorsalwärts aus dem Ostium penis heraus. Dahinter stehen zwei kleine Stachel schräg nach vorne und oben. Von der Penisbasis zieht ein mächtiger Sklerotinbalken an der nieren- förmigen Blase vorbei bis ins distale Drittel des Peniskörpers. Scydmaenus (Eustemmoides) filipenis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Crocker Range, 1200 m, km 60 Kota Kinabalu Tambunan, 4.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda 1 Paratypus (cF). DIAGNOSE: Äußerlich dem Sc. sabahensis gleich, von ihm nur durch den Bau des männlichen Kopulationsapparates zu unterscheiden. BESCHREIBUNG: Long. 1,50 mm, lat. 0,70 mm. Rotbraun, gelblich behaart. In den äußeren Merkmalen dem E. sabahensis gleich. Penis (Abb. 26) sehr schlank und langgestreckt, flach dorsalwärts gebogen, in einer scharfen Spitze endend. Der basale Teil des Penis im einzigen Präparat undurchsichtig, distal der Längsmitte befindet sich ein S-förmig gebogener, mit der Spitze distalwärts orientierter Stachel, an dem ventral ein runder Sklerotinkörper hängt. Distal von dem beschriebenen Stachel liegt ein zweiter nur leicht gebogener, schwächer sklerotisierter Stachel. Beide ragen in der Ruhelage nicht aus dem Penisinneren heraus. Scydmaenus (Eustemmoides) parafilipenis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Kinabalu Nat. Park, Poring Hot Springs, area eastern Ridge, 850 m, 28.8.1988 (lg. Smetana, cMG); 3 Paratypen ebenda (CMG); 3 Paratypen (2 Penispräparate) (lg. Smetana, cF). DIAGNOSE: AuBerlich dem Sc. filipenis sehr ähnlich, aber der Kopf größer, der Halsschild etwas länger, mit 2 Basalgrübchen, der Penis anders gebaut. BESCHREIBUNG: Long. 1,40 mm, lat. 0,70 mm. Dunkel rotbraun, gelblich behaart. Penis (Abb. 27) langgestreckt, schwach dorsalwärts gekrümmt, sein Apex abgerundet, an den beiden Seiten der Dorsalwand des Penis befindet sich, über die basalen drei Viertel 884 HERBERT FRANZ der Penislänge reichend, ein Sklerotinstreifen. Die Streifen erinnern an Parameren um so _ mehr, als sie an dem Rahmen der Basalöffnung wurzeln. In der Mitte des Peniskörpers liegt eine kugelige Blase, die distal in einen langen Ausführungsgang mündet, der bis ins Ostium penis reicht und dort mit einer düsenförmigen Verengung endet. Zu den beiden Seiten des Ausführungsganges liegt je ein langestreckter schmaler Körper, der linke von ihnen mündet in ein dickes Rohr, das fast bis an das Hinterende des Ostiums heranreicht. Scydmaenus (Eustemmoides) allofilipenis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Poring Hot Springs, 600 m, nr. Bat Cave 10.5.1987 (Ig. Bruckhardt u. Löbl, CMG); Kinabalu Nat. Park, Poring Hot Springs, 485 m, 25.8.1988, lg. Smetana, 1 Paratypus (CMG und 1 Paratypus (cF)). DIAGNOSE: Im Penisbau dem Sc. parafilipenis ähnlich, aber viel kleiner und schlanker als dieser. BESCHREIBUNG: Long. 1,20 mm, lat. 0,40 mm. Dunkel rotbraun, bräunlich behaart. Kopf wenig breiter als lang, die Schläfen nur so lang wie der Durchmesser der großen Augen. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Basalglied mehr als doppelt so lang wie breit, 2 leicht gestreckt, die folgenden Glieder klein, 9 nicht ganz doppelt so breit wie 8, 10 noch etwas breiter, das spitz-eiförmige Endglied kürzer als die beiden vorletzten zusammen. Halsschild länger als breit, kaum breiter als der Kopf, seitlich gleichmäßig gerundet, glatt, sehr fein behaart, ohne Basalgrübchen. Flügeldecken langoval, zusammen schon an der Basis ein wenig breiter als die Halsschildbasis, mit angedeutetem Schulterwinkel und einer Basalimpression. Flügel entwickelt. Beine ziemlich lang und schlank. Penis (Abb. 28) von oben betrachtet biskottenförmig, seine Basalöffnung dorsobasal gelegen, das Ostium penis apikal. In der Längsmitte des Penis liegt eine runde Blase, an deren distalem Ende ein Ausführunggang entspringt, der zunächst durch eine lange, becherförmige Halterung hindurchtritt und danach bis an das Penisende reicht. Zu den beiden Seiten der Blase liegt ein schmales lappenförmiges Sklerotingebilde, an das distal beiderseits der Sagittalebene ein bogeförmig zur Mitte gekrümmter großer Lappen anschließt. Die beiden großen Lappen umschließen den dünnen distalen Teil des Ductus ejaculatorius. Das Hinterende des Apex penis ist bogenförmig. Scydmaenus (Eustemmoides) bidentipenis nov. spec. MATERIAL: 25 Ex., davon 17 in cMG und 8 in cF, Kinabalu Nat. Park, 500 bis 1750 m. Holotypus d (Penispräparat) 1500 m, 2.4.1987 (Burckhardt u. Löbl (CMG); d u. andere Paratypen, 1750 m, 27.4.1987 (lg. Burckhardt u. Löbl (cF). DIAGNOSE: Gekennzeichnet durch schlanke Fühler und Beine, sowie durch einen gestreckten Halsschild mit sehr kleinen Basalgrübchen. BESCHREIBUNG: Long. 1,40 mm, lat. 0,70 mm, hell rotbraun, gelblich behaart. Kopf von oben betrachtet querrundlich, die Schläfen doppelt so lang wie die flachen Augen. Fühler schlank, zurückgelegt die Halsschildbasis um das lange Endglied überragend. Alle Glieder bis auf das 7. und 8. länger als breit, die beiden ersten zweieinhalbmal so lang wie breit, das 9. und 10. nicht ganz doppelt so breit wie lang, um ein Viertel länger als breit, das spitz-eiförmige Endglied so lang wie die beiden vorletzten zusammen. SCYDMAENIDAE VON SABAH 885 Halsschild von oben betrachtet kurzoval, stark gewölbt, vor der Mitte am breitesten und hier wenig breiter als der Kopf mit den Augen, glatt und glänzend, fein und anliegend behaart, mit 2 kleinen Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, gleichmäßig gerundet und stark gewölbt, fein punktiert und anliegend behaart, ohne Basalimpression und ohne Schulterbeule. Flügel voll entwickelt. Beine schlank, Schenkel schwach keulenförmig verdickt, Schienen gerade, Vordertarsen des 4 nicht verbreitert. Penis (Abb. 29) tonnenförmig, der Apex aber zur Spitze kegelförmig verschmälert. Die Basalöffnung basal, das Ostium penis dorsoapikal gelegen. Hinter der Basalöffnung liegt ein pilzförmiger Druckregulierungsapparat, dahinter in der basalen Hälfte des Penis eine sehr große langovale Blase mit apikalem dünnen Ausführungsgang. Dieser ist nahe seinem Ursprung in einer ringförmigen Halterung gefaßt, die Teil einer großen von oben und hinten betrachtet links liegenden Sklerotinplatte ist. Hinter der Halterung setzt sich der Ductus ejaculatorius als dickes Rohr apikalwärts fort und endet sichelförmig mit einer scharfen Spitze. Parallel dazu entspringt in der großen Sklerotinplatte ein dicker Sklerotinstrang, der ebenfalls vor der Penisspitze sichelförmig endet. Zwischen den beiden sichelförmig endenden Strängen liegt ein dritter kürzerer und schmälerer, der ebenfalls sichelförmig endet. Die Penisspitze ist abgerundet zweispitzig. Scydmaenus (Eustemmoides) frater nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Kinabalu Nat. Park, Poring Hot Springs, 28.8.1988 (1g. Smetana, cMG); Ebends, Paratypus 3 21.6.1988 (lg. Smetana, cF). DIAGNOSE: Gekennzeichnet durch relativ gestreckte Gestalt, vor allem aber durch den Bau des männlichen Kopulationsapparates, der verwandtschaftliche Beziehungen zu Sc. poringensis erkennen läßt, weshalb ich der Art den Namaen frater gegeben habe. BESCHREIBUNG: Kopf relativ schmal, fast so lang wie breit, Schläfen eineinhalbmal so lang wie der Augendurchmesser, parallel. Fühler zurückgelegt die Halsschildbasis knapp überragend, Glied 1 und 2 gestreckt, 3 bis 6 quadratisch, 7 und 8 breiter als lang, 9 und 10 mehr als doppelt so breit wie 8, schwach quer, das spitz eiförmige Endglied fast so lang wie die leiden vorletzten zusammen. Halsschild etwas länger als breit, vor seiner Längs- mitte am breitesten und hier etwas breiter als der Kopf, fein behaart, ohne Basalgrübchen. Flügeldecken an der Basis zusammen nur wenig breiter als die Halsschildbasis, stark gewölbt, seitlich gleichmäßig gerundet, ohne Basalimpression und ohne Schulterbeule, fein punktiert und kurz behaart. Flügel voll entwickelt. Beine kräftig, für dieVerwandtschaftsgruppe relativ lang, Schenkel keulenförmig verdickt, Schienen gerade, Vordertarsen des d verbreitert. Penis (Abb. 30) nach oben gekrümmt, in der distalen Hälfte verbreitert, zum apikalen Ende gerundet zu einer Spitze verengt, die Basalöffnung dorsobasal gelegen, Ostium penis dorsal im Bereich der Penismitte gelegen, sein Rahmen gegen den Apex verflachend. Aus dem Ostium ragt ein starker Stachel von oben und hinten besehen schräg nach links hinten. Der Ductus ejaculatorius entspringt unter dem Stachel und ist distal düsenförmig verengt. Scydmaenus (Eustommoides) poringensis nov. spec. MATERIAL: Holotypus d (Penispräparat), Kinabalu Nat. Park, Poring Hot Springs, 475 m, 21.8.1988 (Lg. Smetana, cMG); Paratypus d (Penispräparat), ebenda, 480 m, 26.8.1988 (lg. Smetana, cF). 886 HERBERT FRANZ DIAGNOSE: Durch kurze Beine mit dicken Schenkeln und durch die Penisform sehr gekennzeichnet. BESCHREIBUNG: Kopf von oben betrachtet etwas breiter als lang, Schläfen leicht zur Basis konvergierend, eineinhalbmal so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis knapp erreichend, ihr Basalglied doppelt so lang wie breit, 2 leicht gestreckt, 3 bis 6 isodiametrisch, 7 und 8 breiter als lang, 9 doppelt so breit wie 8, 10 noch etwas breiter, beide so lang wie breit, das spitz-eiförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild etwas länger als breit, vor seiner Längsmitte am breitesten, glatt und glänzend, mit 2 kleinen Basalgrübchen. Flügeldecken zusammen nur wenig breiter als die Halsschildbasis, stark gewölbt, seitlich gleichmäßig gerundet, ohne Basalimpression und ohne Schulterbeule, sehr fein und seicht punktiert, nach hinten gerichtet behaart.. Beine kurz, Schenkel stark verdickt, Schienen gerade, Vordertarsen des ¢ verbreitert. Penis (Abb. 31) leicht dorsalwärts gebogen, vor dem Apex stumpfwinkelig erweitert, dahinter beiderseite ausgebuchtet. Der Hinterrand des Apex mit einer sehr stumpf- winkeligen Spitze. Die Basalöffnung dorsobasal gelegen, mit einem stark sklerotisierten Rahmen. Hinter der Basalöffnung liegt in der Sagittalebene eine kaputzenförmige Blase und dahinter ein etwa doppelt so breiter sklerotisierter Körper. Dieser liegt in der Ruhelage zum Teil vor, zum Teil hinter dem Basalrand des Ostium penis. Dieses ist langeiförmig und endet im Niveau der stumpfwinkeligen Erweiterung der Penisseiten. Nahe hinter seiner Basis endet der breite sklerotisierte Körper und entsendet links nach hinten einen langen, stumpfen Zahn. Ein zweiter sehr schmaler Zahn liegt vor dem Hinterende des Ostiums, das distal breit umrahmt ist. Scydmaenus (Eustemmoides) livagui nov. spec. MATERIAL: Nur Holotypus d (Penispräparat) Kinabalu Nat. Park, HQ. at Livagu River, 1500 m, 16.5.1987 (Ig. Smetana, CMG). DIAGNOSE: In der Körperform und namentlich im Bau des männlichen Kopulations- apparates mit Sc. poringensis verwandt. BESCHREIBUNG: Long. 1,60 mm, lat. 0,70 mm. Dunkel rotbraun, gelblich behaart. Kopf von oben betrachtet sehr wenig breiter als lang, die zur Basis konvergierenden Schläfen nur wenig länger als der Durchmesser der großen, flach gewölbten Augen. Fühler zurückgelegt die Halsschildbasis erreichend, ıhr Basalglied etwa doppelt so lang wie breit, 2 und 3 leicht gestreckt, 4 bis 6 isodiametrisch, 7 und 8 breiter als lang, 9 und 10 schwach quer, das spitz-eiförmige Endglied so lang wie die beiden vorletzten zusammen. Halsschild etwas breiter als lang, mit 4 Basalgrübchen. Flügeldecken zusammen etwas breiter als die Halsschildbasis, seitlich mäßig gerundet, fein punktiert und behaart. Beine kräftig, Schenkel keulenförmig verdickt, Schienen gerade, Vordertasen des d kaum merklich verdickt. Penis (Abb. 32) langgestreckt, von oben betrachtet bis zur Apikalpartie parallelseitig, zum Apex leicht erweitert, der Spitzenbereich schwach abgesetzt mit seitlich vortretenden Ecken, der Hinterrand breit abgestutzt. Ostium penis auf der Dorsalseite gelegen, zwei Drittel der Dorsalseite des Penis einnehmend. In seinem Inneren ist ein langer S-förmig gekrümmter Sklerotinstab vorhanden. Er wurzelt an der Basis des Ostiums mit einer stark verbreiterten Basis und endet mit dem schmaleren Ductus ejaculatorius düsenförmig verschmälert knapp vor dem Penisende. Vor der Basis des Ostiums befindet sich eine SCYDMAENIDAE VON SABAH 887 n d 2 « IS LTS 1/10mm ABB. 41-44. 41: Euconnus (s. str.) simillimus nov. spec., Penis a) in Dorsal- b) in Dorsolateralansicht. 42: Euconnus (s. str.) latus nov. spec., Penis in Dorsalansicht. 43: Euconnus (s. str.) paenetypicus nov. spec., Penis in Dorsalansicht. 44: Euconnus (s. str.) paeneglaber nov. spec., Penis a) in Dorsal-, b) in Lateralansicht. 888 HERBERT FRANZ - kurzovale Blase, die mit einem kurzen und dicken Ausführungsgang in die basale Erweiterung des dicken Sklerotinstabes mündet. ANMERKUNG: Da von mehreren der vorstehend beschriebenen Eustemmoides-Arten nur 1 6 vorliegt, ist die nachstehende Bestimmungstabelle unvollständig und es empfielt sich zur Sicherung der Determination Penispräparate anzufertigen. BESTIMMUNGSTABELLE DER Eustemmoides-ARTEN VON SABAH 1 Fühler fast halb-körperlang, alle Glieder bis auf das 7. und 8 viel länger als breit ....... LE APE AR A PARIS RPC ge wis hag a Hh Mey EE SRE e PR, cuneipenis n. SP. =) Fublersktirzer undidicker i Ri PRO SR ER 2 2 Elügeldecken.dicht und kräftis;punktiert”. LI Re Res 3 — Flügeldecken höchstens schütter und unauffällig punktiert ............................... 5 Fühler kurz, zurückgelegt die Halsschildbasis nicht erreichend ........................... + Fühler lang, zurückgelegt die Halsschildbasis deutlich überragend ........................ BO IAT Ml ee ee LE A BEE sabahi n. sp. und sabahanus n. sp. 4 Kleiner, long. 1,20 mm, Kopf fast so breit wie der Halsschild ............. punctatus n. sp. — Größer, long. 1,40 mm, Kopf deutlich schmäler als der Halsschild .... thermarum n. sp. 5 9. Fühlerglied nur halb so groß wie das 10 . ee SUVICOANESPN — 9. und 10. Fühlerglied in der Größe nur wenig verschieden ee CT 6 6 Kôrperoberfläche nahezu kahl, Kopf ein wenig länger als breit ...... alessmetanai n. Sp. — Kòrperoberseite mindestens stellenweise deutlich behaart, Kopf nicht länger als breit 7 7 sNorperlange unter 1,20 mme ee ee SEE 8 = JKorperlange 1,20 mimi und darüber... Eten TI PIONEER 10 8 Fühler zurückgelegt die Halsschildbasis deutlich überragend, Körperlänge knapp 1,20 TS en ehe see linie ee en a III furcatus n. sp. — Fühler zurückgelegt die Halsschildbasis knapp erreichend ............................... 9 976. Fühlerglied langer als breit, Körperlänge um 120mm E Re ere — 6. Fühlerglied annähernd isodiametrisch, Körperlänge um 1,00 mm ....................... EIA IRAN) QI el e Ni bue D N St een burckhardtloebli n. sp. 10 Körperlänge 1,50 bis 1,60 mm, Halschildbasis mit Punktgrübchen ..................... 11 — Körperlänge unter 1,30'mm cenere TIT Ce ER 12 MSEaISShId'mIt #'Punktsrübehen TO livagui n. sp. a Halsschildmit 2/Punktsrübehen un. EI poringensis n. Sp. 12##und'8Eühlerglied nicht asymmetrisch.." "CRC TC eee eee EEE EEE 13 — 7. und 8. Fühlerglied asymmetrisch”.......2. cc D TETE TL ES 14 13 2. und 3. Fühlerglied mehr als eineinhalbmal so lang wie breit, Vordertarsen des d niehtsverbreitert®en. ILL bidentipenis n. sp. — 2. und 3. Fühlerglied weniger als eineinhalbmal so lang wie breit, Vordertarsen des d VI AO RES RR eo dae A A a il 0000050000000 frater n. sp. 14 Dem Armatoscydmaenus brevitarsis durch gedrungenen Körperbau, trapezförmigen Halsschild und kurze Beine sehr ähnlich, aber die Hinterschenkel des d ungezähnt undidemPenisiganz anders celonmt RSA E eee NE re filipenis n. sp. — Andere Merkmalskombinationen ............ sabahensis n. sp. und allosabahensis n. sp. SCYDMAENIDAE VON SABAH 889 Gattung Loeblites Franz In Sabah kommt eine zweite Art der bisher monotypischen Gattung Loeblites vor. Loeblites sabahensis nov. spec. MATERIAL: Holotypus d (Penispräparat) und 4 Paratypen d, Poring Hot Springs, 500 m, 7.5.1987 (Burckhardt u. Löbl, CMG); 19 Paratypus, ebenda, 11.5.1987 (Burckhardt u. Löbl, cMG); 20 1? Paratypen, ebenda (lg. Burckhardt u. Löbl) 11.5.1987 (cF); 3d Paratypen, Kibongol V. 7 km N Tambunan, 700 m, 20.5.1987 (Burckhardt u. Löbl, cMG); 14 (Paratypus) Poing Hot Springs, 490 m, area Kipungit, 14.8.1988 (lg. Smetana, cF); 19 Crocker Range, 1270 m, 60 km NE Kota Kinabalu-Tambunan, 17.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Dem L. mastigicornis m. sehr ähnlich, aber rotbraun gefärbt, am Halsschild und namentlich auf den Flügeldecken viel länger behaart, der Halsschild viel kürzer und nur um etwa ein Viertel länger als breit, seine Basalgrübchen viel größer, die Flügeldecken mit deutlicher, lateral von einem Schulterhöcker begrenzter Basalimpression sowie mit einer großen Depression hinter dem Schildchen beiderseits der Naht. BESCHREIBUNG: Long. 2,20 bis 2,40 mm, lat. 0,70 bis 0,90 mm. Dunkel rotbraun, stark glänzend, die Palpen und Tarsen heller gefärbt, braun behaart. Kopf mit den stark vorgewölbten Augen fast so breit wie lang, mit deutlichen Supraantennalhöckern. Fühler beim d länger als beim 9, beim d fast körperlang, beim 9 die Längsmitte der Fliigeldecken erreichend, beim d Glied 1 der Fühler etwas mehr als doppelt so lang wie breit, 3 und 4 mehr als 3 mal, 5 bis 8 fiinfmal so lang wie breit, 9 etwas, 10 und 11 noch kürzer. Halsschild im vorderen Viertel seiner Lange am breitesten, von da zur Basis leicht ausgeschwungen verengt, mit 4 großen Basalgrübchen, oberseits schiitter, an den Seiten dicht und struppig abstehend behaart. Flügeldecken schon an der Basis zusammen wesentlich breiter als die Halsschildbasis, deutlich punktiert und lang abstehend behaart, mit nach hinten verflachter Basal- impression und hinter dem Schildchen beiderseits der Naht mit großer, flacher Impression. Beine lang, Schenkel keulenförmig verdickt, Schienen gerade, Tarsen sehr schlank. Penis (Abb. 33) in der Form dem des L. mastigicornis sehr ähnlich, schwach sklerotisiert. Parameren die Basalöffnung des Penis distal und lateral umfassend, das Penisende überragend, ohne Tastborsten. Im Penisinneren ist hinter der Basalöffnung ein zungen- förmiger horizontal gelegener Sklerotinkörper erkennbar, bei einem Paratypus ist an seiner Stelle ein ungleich stark sklerotisierter, unregelmäßig geformter Körper vorhanden. Gattung Horaeomorphus Schaufuss Horaeomorphus loeblianus nov. spec. MATERIAL: Holotypus 4 und ein Paratypus, Sabah, Mount Kinabalu Nat. Park, Poring Hot Springs, area Eastern Ridge Tr., 790 m, 17.8.1988 (lg. Smetana, cMG); 1 d und 4 Ex. Paratypen, Poring Hot Springs, 550 bis 600 m, 9.5.1987 (lg. Burckhardt u. Löbl, cF); 5 Paratypen, ebenda, 480 bis 530 m, 8.-9.5 1987 (Ig. Smetana, CMG); 25 (Penispräparat) Paratypen 7 km N Tambunan und Langanan Fall, 900-950 m, 13.5.1987 (lg. Burckhardt u. Löbl, cMG); 24 und 4 Ex. (Paratypen) ebenda, (CMG); Poring Hot Springs, Laganan River, 850 bis 950 m, 9.-20.5.1987 (CMG). DIAGNOSE: Gekennzeichnet durch dicke, sehr kurze, zurückgelegt die Halsschildbasis nicht erreichende Fühler, zwischen den hoch aufgewölbten Supraantennalhöckern tief eingesenkte Stirn und mediodistal ausgerandete, in der Ausrandung steif behaarte Mittelschienen. 890 HERBERT FRANZ BESCHREIBUNG: Long. 2,00 bis 2,10 mm, lat. 0,80 bis 1,00 mm. Dunkel rotbraun, rotgelb behaart. Kopf von oben betrachtet in der Anlage lang-dreieckig, im Bereich der knapp vor der Kopfbasis stehenden Augen am breitesten, die Schläfen sehr kurz, abstehend beborstet, die Stirn zwischen den großen Supraantennalhöckern tief eingesenkt. Fühler dick, zurückgelegt die Halsschildbasis nicht erreichend, ihre 5 ersten Glieder annähernd isodiametrisch, 7 bis 10 breiter als lang, gegen das 10. an Breite zunehmend, das gerundet- kegelförmige Endglied so lang wie die beiden vorletzten zusammen, alle Glieder lang abstehend behaart. Halsschild wenig länger als breit, im vorderen Drittel seiner Länge am breitesten und hier breiter als der Kopf mit den Augen, in der vorderen Hälfte kurz und steif aufgerichtet, hinter der Mitte noch dichter und lang, nach hinten gerichtet behaart, mit 4 Basalgrübchen, auf der Scheibe dicht punktiert. Flügeldecken länglichoval, stark gewölbt, an der Basis zusammen nur wenig breiter als die Halsschildbasis, mit runder, lateral von einer Humeralfalte begrenzter Basalimpression, fein punktiert, die Punktierung von der Mitte nach hinten gerichteten Behaarrung weitgehend verdeckt. Flügel voll entwickelt. Beine mäßig lang, Schenkel keulenförmig verdickt, Vorderschienen mediobasal leicht aus- gerandet, die Mittelschienen wesentlich stärker, in der Ausrandung mit steifen Borsten besetzt. Penis (Abb. 34) etwa doppelt so lang wie von oben besehen breit, von der Längsmitte zur Spitze allmählich verengt, die Parameren lang und parallel, die Penispitze ein wenig überragend, mit einigen terminalen Tastborsten. Etwas vor der Längsmitte des Penis liegt sagittal ein stark sklerotisierter, annähernd isodiametrischer Komplex, der aus um eine zentrale Achse angeordneten Körpern besteht. Zu der Achse sind hintereinander 2 Paare von Schalen spiegelbildlich angeordnet. Die Achse ist offenbar der Ductus ejaculatorius, der in der Ruhelage bis knapp vor die Spitze des Penis reicht und dort zu einer Düse verengt ist. Horaeomorphus sabahensis nov. spec. MATERIAL: Holotypus d (Penispräparat). Crocker Range, 1550-1650 m, 16.5.1987 (Ig. Burckhardt u. Löbl, cMG); Paratypus d (Penispräparat, ebenda (cF); 9 Paratypus, Mount Kinabalu Nat. Park, NO Silau-Silau Tr., 1540 m, 14.8 bis 1.9.1988 (lg. Smetana, cMG). DIAGNOSE: Relativ groß und schlank, nur die Fühler kurz und dick. Halsschild und Flügeldecken ziemlich fein punktiert, aber dicht und lang behaart. BESCHREIBUNG: Long. 2,60 bis 2,80 m, lat. 1,00 bis 1,20 mm, schwarz, die Beine dunkel rotbraun, bräunlich behaart. Kopf von oben betrachtet gerundet-dreieckig mit großen Supraantennalhöckern, da- zwischen eingesenkter Stirn und unter den Fühlerwurzeln stehenden Augen, Schläfen sehr kurz. Fühler sehr dick und zurückgelegt die Halsschildbasis wenig überragend, beim d die ersten 6 Glieder annähernd quadratisch, die folgenden viel breiter als lang, und bis zum 10. Glied an Breite zunehmend, das Endglied kegelförmig, so lang wie breit. Beim 2 alle Glieder bis auf das Endglied breiter als lang. Halsschild etwas länger als breit, im vorderen Drittel seiner Länge am breitesten und hier breiter als der Kopf mit den Augen, vor der Basis ausgeschwungen, mit 4 in einer Querfurche stehenden Grübchen, besonders an den Seiten dicht und abstehend behaart. Flügeldecken länglichoval, stark gewölbt und schon an der Basis zusammen viel breiter als die Halsschildbasis, mit strichförmig lateral von einem Schulterhöcker begrenzter Basalimpression, sehr fein punktiert und dicht, schräg abstehend behaart. Flügel entwickelt. SCYDMAENIDAE VON SABAH 891 Beine beim d etwas länger als beim 2, Schenkel mäßig verdickt, Schienen gerade. Penis in dem einzigen vorhandenen Präparat stark geschrumpft, in den Umrissen dem des H. loeblianus ähnlich, etwas länger und schlanker, die Parameren zur Spitze verbreitert, mit zahlreichen terminalen Tastborsten versehen. Horaeomorphus punctatissimus nov. spec. MATERIAL: Holotypus d, Sabah, Kinabalu Nat. Park, HQ at Livagu Rv., 1500 m, 30.4.1987 (lg. Smetana, cMG); Paratypus $, Kinabalu N HQ. 1550 m, 2. bis 4.9.1988 lg. Bright, cMG); Sabah, Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl (CMW); Mount Kinabalu Nat. Park, HQ at Livagu Rv., 1500 m, 30.4.1987 (lg. Smetana, cF). DIAGNOSE: Sehr ausgezeichnet durch flach gewölbte Gestalt, sehr dichte Punktierung der ganzen Oberseite sowie einen langen Sporn an den Trochanteren der Hinterbeine der à. BESCHREIBUNG: Long. 2,40 bis 2,60 mm, lat. 0,80 bis 0,90 mm, rotbraun, bräunlich behaart. Kopf von oben betrachtet annähernd gerundet-dreieckig, länger als breit, im basalen Viertel seiner Länge im Niveau der kleinen Augen am breitesten, die Augen an den Kopfseiten gelegen, wenig vorragend, die Schäfen nur halb so lang wie der Augendurchmesser, hinter den Fühlerwurzeln mit 2 Grübchen, schütter punktiert, nur die Schläfen spärlich behaart. Fühler ziemlich schlank, allmählich zur Spitze verdickt, die ersten 6 Glieder gestreckt, mit gegen das 6. abnehmender Länge, 7 isodiametrisch, 8 und 9 wenig, 10 stark quer, das Endglied kegelförmig, so lang wie breit. Halsschild flach gewölbt, vor seiner Mitte am breitesten und hier wesentlich breiter als der Kopf mit den Augen, vor der Basis mit 2 Grübchen, dicht punktiert und abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, flach gewöbt, dicht punktiert und nach hinten gerichtet behaart, mit breiter Basalimpression und Andeutung einer Schulterbeule. Flügel voll entwickelt. Beine mittellang, Hinterschenkel stark verdickt, Hinterschienen stark medialwärts gekrümmt, die Trochanteren der Hinterbeine beim d mit einem langen, parallel zur Schenkelachse liegenden Dorn. Bei dem einzigen vorliegenden d wurde bei der Prä- paration kein Aedeagus gefunden. BESTIMMUNGSTABELLE DER Horaeomorphus-ARTEN BORNEOS 1 Tibien, besonders die der Hinterbeine stark medialwärts gekrümmt, die Trochanteren der Hinterbeine beim d mit einem langen Sporn, Oberseite sehr dicht und grob punktiert, Körperlänge 2,40 mm und darüber ................... punctatissimus nov. spec. — Tibien höchstens schwach medialwärts gekrümmt, Trochanteren der Hinterbeine beim CONTE: SDOLH PUS LS NINE TEAMS NEE tee Ne SUIS SE RE EN 2 2 Sehr kleine Art (long 1,50 mm), mit dicht und kräftig punktierter Oberseite ........ ee A OBEN I MORTA CI NE SR sno sarawakensis Franz — Größere Arten mit weniger kräftiger oder ohne Punktierung der Oberseite ............. 3) 3 Große Art, Körperlänge 2,60 bis 2,80 mm. Schenkel mäßig verdickt, Schienen gerade u RAA AIR MR cc Re ERROR sabahensis nov. spec. — Kleinere Art, Körperlänge 2,00 bis 2,10 mm, Mittelteil des Kopfes mediodistal ausgerandet, in der Ausrandung grob punktiert (Sabah) ............ loeblianus nov. spec. 892 HERBERT FRANZ Gattung Syndicus Motschulsky Syndicus kinabalui nov. spec. MATERIAL: Holotypus 4 und Paratypus d, Mount Kinabalu Nat. Park, Poring Hot Springs, 485 bis 490 m, 24. und 27.8.1988 (lg. Smetana, CMG); Paratypus d ebenda. 27.8.1988 (lg. Smetana, cF). DIAGNOSE: Durch die sehr stark verdickten Schenkel und sehr feine Punktierung von Kopf, Halsschild und Flügeldecken ausgezeichnet. BESCHREIBUNG: Long. 2,50 bis 2,60 mm, lat. 0,80 bis 1,00 mm. Kopf von oben betrachtet doppelt so breit wie lang, die Stirn gewölbt, ohne Spur von Supraantennalhöckern, Augen flach gewölbt, aus der Kopfwölbung kaum vorstehend, Oberseite des Kopfes sehr fein punktiert und kaum erkennbar behaart (80-fache Vergrößerung). Fühler zurückgelegt die Halsschildbasis nicht ganz erreichend, ihre ersten 4 Glieder leicht gestreckt, 5 isodiametrisch, 6 bis 9 breiter als lang, das 10. und 11. Glied zusammen gerundet-kegelförmig, das 11. dem 10., wie auch bei den anderen Arten der Gattung, wie eine Eichel dem Fruchtbecher aufsitzend. Halsschild um ein Fünftel länger als breit, breiter als der Kopf mit den Augen, von der Mitte zur Basis sehr stark verengt, vor dieser mit 4 seichten Grübchen, die mittleren voneinander durch einen Längskiel getrennt, die Scheibe dicht und fein punktiert, kurz und steif abstehend behaart. Flügeldecken schon an der Basis zusammen wesentlich breiter als die Halsschildbasis, mit einer runden, nach hinten offenen Basalimpression und Andeutung einer Schulterbeule, sehr fein punktiert, schräg nach hinten gerichtet behaart. Flügel entwickelt. Beine kurz, Schenkel sehr stark keulenförmig verdickt, Schienen gerade, Tarsen kurz. Penis (Abb. 35) langgestreckt, von oben besehen lang-eiförmig, das Penisende schmal abgerundet. Parameren das Penisende erreichend, ohne Tastborsten. Im Penisinneren liegt sagittal der Ductus ejaculatorius, der hinter der Basalöffnung in einem Komplex von 3 stark sklerotisierten Drüsen entspringt. Zu den beiden Seiten dieses Komplexes befinden sich sklerotisierte Falten der Präputialsackwand. Gattung Syndicomorphus nov. gen. In meiner Monographie der Gattung Syndicus (FRANZ 1971) habe ich nachgewiesen, daß die Vertreter der Gattung Syndicus eigentlich 11-gliederige Fühler besitzen. Ich führte aus: “Untersucht man die Fühler der verschiedenen Syndicus-Arten genauer, so sieht man, daß diese eigentlich nicht 10- sondern 11-gliederig sind. Das 11. Glied sitzt dem 10. wie eine Eichel ihrem Fruchtbecher (Cupula) auf, wobei es bei den einzelnen Arten zu einer mehr oder weniger weitgehenden Verschmelzung kommt”. Das 10. und 11. Glied sind zusammen gerundet-konisch oder kurz-eiförmig. Nun liegt mir aus Sabah neben dem schon beschriebenen Syndicus kinabalui eine durch bedeutendere Größe und lange und schlanke Fühler ausgezeichnete Art vor, bei der auch das 10. Glied sehr scharf begrenzt ist und keine Spur eines ihm aufsitzenden 11. Gliedes zeigt. Diese Art weicht von den Syndicus-Arten so weit ab, daß es gerechtfertigt ist, für sie ein eigenes Genus zu errichten. Es erinnert im Habitus stark an Mastigus und auch an Loeblites, unterscheidet sich aber von dem letzteren durch robusteren Körperbau und kürzere Fühler, sowie von beiden Genera durch nur 10-gliederige Fühler. Typusart ist der nachfolgend beschriebene Syndicomorphus magnus nov. spec. SCYDMAENIDAE VON SABAH 893 Syndicomorphus magnus nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Sabah Kinabalu Nat. Park, Bukit Utar Trail, 1700 m, 29.4.1987 (1g. Smetana, cMG). DIAGNOSE: Auffällig groß, long. 3,20 mm, lat. 1,20 mm. Schwarz, schwarz-braun behaart. BESCHREIBUNG: Durch die bedeutende Größe, die schwarze Farbe und 10-gliederige Fühler sehr ausgezeichnet. Kopf mit den vorgewölbten Augen um ein Drittel breiter als lang, die Stirn zwischen den Fühlerbasen flach eingedellt, schütter punktiert und behaart. Fühler fadenförmig, zurückgelegt die Längsmitte der Flügeldecken knapp erreichend, ihr Basalglied leicht gestreckt, 2 schmäler und kürzer als 1, 3 einhalbmal, 4 knapp doppelt so lang wie breit, 5 bis ausschließlich 9 zweieinhalbmal so lang wie breit, das Endglied ist spitz-oval und erreicht die Länge der 3-fachen Breite. Halsschild im vorderen Drittel seiner Länge am breitesten und hier deutlich breiter als der Kopf mit den Augen, um ein Drittel breiter als lang, stark gewölbt, auf der Scheibe lang, schütter und anliegend behaart. Flügel verkümmert. Beine lang und ziemlich schlank, Schenkel keulenförmig verdickt, Schienen gerade, Tarsen schlank. Penis (Abb. 36) langgestreckt, von der Seite besehen doppelt so lang wie breit, in einer scharfen Spitze endend. Parameren schlank, die Penisspitze nicht erreichend, ohne Tastborsten. Das Penisinnere ist undurchsichtig. Genus Borneosabahia nov. gen. Mit Euconnus nahe verwandt, die Fühler allmählich zur Spitze verdickt, mit sehr großem keulenförmigen Endglied. Der Kopf gerundet-dreieckig, lang und nach hinten gerichtet behaart, die Augen in den stark nach hinten verlagerten Fühlerfurchen versenkt. Der Halschild halb so breit wie die Kopfbasis, Kiefertaster klein, das 4. Glied pfriemenförmig, schwer sichtbar. Halsschild konisch, kaum breiter als der Kopf, Hinterhüften breit getrennt, Mittelhüften durch einen hoch aufgewölbten Mesosternalkiel voneinander gechieden. Typusart: Borneosabahia mirifica nov. spec. Borneosabahia mirifica nov. spec. (Abb. 37) MATERIAL: Holotypus 5 (Penispräparat, Crocker Range, 1500 m, Mount Kinabalu, 20.4.1987 (1g. Burckhardt u. Löbl, cMG); ebenda, 1000 bis 1500 m, 24 Ex. (Paratypen) lg. Burckhardt u. Löbl (cMG u. cF, mehrere Penispräparate). DIAGNOSE: Sehr ausgezeichnet durch kurze, dicke Fühler mit auffallend großem, keulenförmigen Endglied, bei Betrachtung von oben gerundet-dreieckigen Kopf, mit lang nach hinten gerichtet behaartem Scheitel, durch kleinen gerundet-viereckigen Halsschild mit 4 untereinander durch eine Querfurche verbundene Basalgrübchen, durch ovale ziemlich flach gewölbte Flügeldecken mit nach hinten verflachter Basalimpression und schlanke Beine. BESCHREIBUNG: Long. 1,90 bis 2,20 mm, lat. 0,78 bis 0,90 mm, schwarzbraun bis dunkel rotbraun, braun behaart. Kopf von oben betrachtet gerundet-dreieckig, am Scheitel lang, nach hinten gerichtet behaart, die Stirn wulstförmig emporgewölbt, mit tiefen, nach hinten allmählich verflachten Fühlergruben und in diesen an den Kopfseiten eingesenkten Augen. Fühler 894 HERBERT FRANZ - kurz und dick, allmählich zur Spitze verbreitert, mit großem keulenförmigem Endglied, zurückgelegt die Halsschildmitte wenig überragend, ihr Basalglied um die Hälfte länger als breit, 2 leicht gestreckt, die folgenden bis zum 10. zunehmend breiter als lang und zunehmend breiter. Maxillarpalper schlank, ihr 4. Glied pfriemenförmig, sehr klein. Halsschild konisch, so lang wie breit, an der Basis kaum breiter als der Kopf mit den Augen vor der Basis, mit 4 großen durch eine Querfurche verbundene Grübchen, abstehend, an den Seiten struppig behaart. Flügeldecken schon an der Basis zusammen viel breiter als die Halsschildbasis, flach gewölbt, dicht und schräg nach hinten abstehend behaart. Flügel voll entwickelt. Beine schlank, Schenkel schwach verdickt, Schienen gerade, Hinterhüften breit getrennt, die Hinterbrust glatt und glänzend. Penis (Abb. 37a-d) stark sklerotisiert, fast rechtwinkelig gekrümmt, in einem kleinen dreieckigen Apex endend. Dieser ist in seiner Lage zum Peniskörper verstellbar. In Abb. 37a ist der Penis ausgestülpt in Ventralansicht dargestellt, in Abb. 37b in Dorsalansicht, in Abb. 37c in Lateralansicht in Ruhestellung und in Abb. 37d in Erektion. Die Basalöffnung besitzt keinen sklerotisierten Rahmen, die Parameren sind an der Basis vor der Basalöffnung verbunden. Der Ductus ejaculatorius ist großenteils dünnhäutig und in den Präparaten nicht sichtbar. Nur das verbreiterte Ende, das zum Teil auf der Dorsalseite des Penis aus dem Peniskörper herausragt, ist stärker sklerotisiert. Es deutet die Lage des Ostium penis an, die wie die Basalöffnung keinen sklerotisierten Rahmen besitzt. Dieser Mangel wie die geringe Differenzierung des ganzen Kopulationsapparates bezeugen eine niedere Entwicklungsstufe der Gattung. Die geringe Formbeständigkeit des männlichen Kopulationsapparates verleitet dazu, morphologische Unterschiede zu vermuten. Das Vorhandensein einer sehr großen Zahl von Belegexemplaren hat es ermöglicht, zahlreiche Penispräparate anzufertigen und damit die Einheitlichkeit des Baues des Kopulationsapparates bei allen undersuchten Individuen sicherzustellen. Gattung Euconnus Thomson Untergattung Euconnus Thomson s. str. Econnus (s.str.) pseudosukhotanus nov. spec. MATERIAL: Holotypus d (Penispräparat), Mount Kinabalu Nat. Park, HQ at Livagu, 1500 m, 15.5.1987 (Ig. Smetana, CMG); Paratypus d (Penispräparat), Mount Kinabalu, 1900 m, 28.4.1987 (1g. Burckhardt u. Löbl, CMG); 4 Paratypus (Penispräparat), ebenda, 1500 m, 29.4.1987 (cF); Mount Kinabalu Nat. Park, HQ Silau-Silau, 1550 m, 4.9.1988 (lg. Smetana, cF); ebenda Paratypus d (Penispräparat, CMG). Dazu zahlreiche weitere Ex. wahrscheinlich dieser Art (CMG). DIAGNOSE: Dem E. sukhotanus aus Thailand ähnlich. Gekennzeichnet durch mit Ausnahme des Halsschildes kahle Körperoberseite, großen Kopf, lange Fühler mit lockerer 4 gliederiger Keule und stark keulenförmig verdickte Vorderschenkel. BESCHREIBUNG: Long. 1,80 bis 2,20 mm, lat. 0,80 bis 0,95 mm, Rotbraun, glänzend, Halsschild braun behaart. Kopf von oben betrachtet queroval, mit den großen, flach gewölbten Augen breiter als der Halsschild. Fühler zurückgelegt ungefähr die Halsschildbasis erreichend, mit langer, lockerer, 4 gliederiger Keule, ihre beiden ersten Glieder um ein Drittel länger als breit, 3 bis 7 leicht gestreckt, 8 bis 10 nicht ganz doppelt so breit wie 7, um ein Fünftel länger als breit, das Endglied eiförmig, etwas länger als das vorletzte. Halsschild leicht gestreckt, glatt und glänzend, ohne Basalgrübchen, ziemlich gleichmäßig gerundet und struppig abstehend behaart. Flügeldecken stark gewölbt, schon an der Basis SCYDMAENIDAE VON SABAH 895 zusammen breiter als die Halsschildbasis, glatt und glänzend, ohne Basalimpression und ohne Schulterbeule, die Naht hinter der Basis leicht eingetieft, Schildchen nicht erkennbar. Flügel voll entwickelt. Beine ziemlich lang, Vorderschenkel des 3 stark verdickt, Schienen vor der Längsmitte mediodistal ausgerandet. Penis (Abb. 38) stark sklerotisiert, von oben betrachtet kurzoval, der Apex zweispitzig, die Basalôffnung groß, von einem stark sklerotisierten Rahmen umgeben. Bei seitlicher Betrachtung dem E. paeneglaber nov. spec. (Abb. 44b) ähnlich. Parameren das Penisende nicht erreichend. Im Penisinneren liegt sagittal hinter der Längsmitte eine kleine runde Blase, die distal mit 2 divergierenden Stützen mit einem ambossförmigen Sklerotinkörper verbunden ist. Dieser sitzt unmittelbar vor dem apikalen Penisende einem M-förmigen Sklerotinband auf. Euconnus (s. str.) allosukhotanus nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Crocker Range, 1600 m, km 51 von Kota Kinabalu Tambunan, 18.5.1987 (lg. Burckhardt u. Löbl, cMG); Paratypus d (Penispräparat) vom selben Fundort (lg. Burckhardt u. Löbl. cF); 4 Paratypus (Penispräparat) vom selben Fundort (CMG). DIAGNOSE: Äußerlich dem E. pseudosukhotanus fast vollständig gleich, im Penisbau jedoch von ihm sehr verschieden. BESCHREIBUNG: Long. 2,20 mm, lat 0,80 mm. Dunkel rotbraun, Palpen und Beine heller gefärbt, Halsschildseiten braun behaart. Äußerlich unterscheidet sich die Art von E. pseudosukhotanus durch nur eineinhalbmal so langes wie breites |. Fühlerglied und isodiametrische Glieder 3 bis 6, sowie durch außerordentlich stark keulenförmig verdickte Vörderschenkel des à. Penis (Abb. 39a, b) ganz anders gebaut als bei der Vergleichsart. Peniskörper von oben betrachtet gerundet-rechteckig, um ein Fünftel länger als breit, mit sehr großer, gerundet- querrechteckiger, von einem stark sklerotisierten Rahmen umgebener Basalöffnung, Para- meren gerade, dünn, nicht sklerotisiert, die Basis des Apex penis erreichend, mit je 2 terminalen Tastborsten. Apex aus 2 sehr spitzwinkelig-dreieckigen, an der Basis medial gemeinsam queroval ausgeschnittenen Teilen bestehend. Operculum den Pensikörper ein wenig überragend, sein Hinterrand bogenförmig, zwischen den beiden Teilen des Apex sichtbar. Im Penisinneren sind infolge von Lufteinschlüssen im Präparat keine skleroti- sierten Gebilde erkennbar. Euconnus (s. str.) kinabaluanus nov. spec. MATERIAL: Holotypus d (Penispräparat), Mount Kinabalu, 1500 m, 25.4.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Dem E. pseudoukhotanus äußerlich fast völlig gleich, des Penis aber ganz anders geformt. BESCHREIBUNG: Long 2,00 mm, lat. 0,75 mm. Dunkel rotbraun, Fühlergeißel, Palpen und Beine heller gefärbt, Halsschild sehr kurz, wie geschoren braun behaart. Stimmt in den äußeren Merkmalen so weit mit E. pseudosukhotanus überein, daß es genügt, die geringfügigen Unterschiede anzuführen: Zweites Fühlerglied nur um ein Viertel länger als breit, Halsschild ein wenig länger als breit, Mittelschienen des & medialwärts gekrümmt. Penis (Abb. 40) sehr gedrungen gebaut, von oben betrachtet in der Anlage sehr kurzoval, der Apex vom Peniskörper deutlich abgesetzt, dreieckig, von oben und hinten betrachtet 896 HERBERT FRANZ ABB. 45-51. 45: Euconnus (s. str.) smetanaensis nov. spec., Penis in Dorsalansicht. 46: Euconnus (s. str.) apicefurcatus nov. spec., Penis in Dorsalansicht. 47: Euconnus (Borneoconnus) laticlava nov. spec., Dorsalansicht des Vorderkörpers. 48: Euconnus (Borneoconnus) laticlava nov. spec. Penis in Dorsalansicht. 49: Euconnus (Borneoconnus) sabahanus nov. spec., Vorderkörper in Dorsalansicht. 50: Euconnus (Borneoconnus) sabahanus nov. spec., Penis in Dorsalansicht. 51: Euconnus (Borneo- connus) eremita nov. spec. Habitusbild. SCYDMAENIDAE VON SABAH 897 links von einer sehr spitzwinkelig -dreieckigen, horizontalen Platzte zum Teil überlagert. Basalöffnung groß, mit einem breiten, stark sklerotisierten Rahmen umgeben. Dieser Rahmen springt basal beiderseit mit einer abgerundeten Ecke vor und ist distal breit abgerundet. Die Parameren entspringen hier, sie sind dünnhäutig, breit und erreichen die Basis des Apex nicht. Tastborsten sind an ihnen nicht erkennbar. Vor dem Ostium penis befindet sich im Penisinneren eine querovale, stark sklerotisierte Blase. Aus dem Ostium penis ragt nahe der Sagittalebene ein dicker Stab über den Apex penis vor, ein zweiter tritt von oben und hinten betrachtet nahe dem rechten Seitenrand nach hinten aus dem Peniskörper aus. Euconnus (s. str.) simillimus nov. spec. MATERIAL: Holotypus d (Penispräparat), Mount Kinabalu, 1550 m, 29.4.1987 (Burckhardt u. Löbl, cMG); ebenda, 13 Ex., wahrscheinlich dieser Art (CMG); ebenda 8 Ex., 1150 m 27.4.1987, ohne Penispräparat nicht als Paratypen bezeichnet, 27.4.1987 6 (CMG); ebenso 15 Ex. cMG und 6 Ex (CB): DIAGNOSE: Da sich von E. kinabaluanus keine äußeren Unterschiede feststellen lassen, erübrigt sich eine Wiederholung der für diese Art angeführten Merkmale. BESCHREIBUNG: Die Penisform weicht von der Vergleichsart stark ab (vgl. Abb. 41). Penis von oben betrachtet gerundet-rechteckig, stark sklerotisiert, die Basalöffnung sehr groß, mit sehr breitem sklerotisiertem Rahmen. Die Dorsalwand ist apikal im Bogen über die ganze Penisbreite tief ausgeschnitten, im Ausschnitt ist eine horizontale Sklerotinplatte sichtbar, die medial von hinten tief eingeschnitten ist und beiderseits des Einschnittes breit dreieckig nach hinten vorspringt. In dem Einschnitt liegt eine spitzwinkelig-dreieckige Platte, die der Lage nach dem Operculum entspricht. Die Parameren erreichen das Hinterende des Penis nicht, sie tragen je 2 terminale Tastborsten. Bei dorsolateraler Betrachtung (Abb. 42 b) sieht man, daß die unter dem bogenförmigen Ausschnitt der Dorsalwand sichtbare mediale Platte tiefer liegt als die Dorsalwand und dem Operculum homolog ist. Noch tiefer liegt der sagittal angeordnete Ductus ejaculatorius, der hinter der Penismitte in eine kleine, stark umwandete Blase mündet und seitlich beiderseits von einem schräg nach hinten und zur Mitte gerichteten Sklerotinstab begleitet ist. Euconnus (s. str.) latus nov. spec. MATERIAL: Holotypus d (Penispräparat) Mount Kinabalu, 1550 bis 1750 m, 29.4.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 6 Ex. Determination fraglich, (CMG und cF). DIAGNOSE: Dem E. simillimus n. sp. weitgehend gleich, der Körper etwas breiter, die Fühler etwas länger, der Penis anders gebaut. BESCHREIBUNG. Mit Rücksicht auf die fast völlige Übereinstimmung in den äußeren Merkmalen mit E. simillimus genügt es, eine Penisbeschreibung zu geben. Penis (Abb. 42) gedrungen gebaut, von oben betrachtet in der Anlage kurz rechteckig, mit sehr großer, mit einem sehr breiten Sklerotinrahmen versehener Basalöffnung. Die Parameren an deren distalem Rand inserierend, das Penisende erreichend, mit je 2 terminalen Tastborsten versehen. Apex penis nicht abgesetzt, sein Hinterrand in breitem Bogen ausgeschnitten. In dem Auschnitt ist das Operculum sichtbar, dessen Hinterrand mit dem Hinterende der Penisseitem koinzidiert, zwischen ihnen nahezu gerade verläuft, in der Sagittalebene aber schmal ausgerandet ist. In der Ausrandung sind die Enden von 2 Skerotinstäben sichtbar. Vor der Ausrandung steht im Penisinneren ein becherförmiges Sklerotingebilde, aus dessen basalem Rande eine kugelförmige Blase nach vore ragt. 898 HERBERT FRANZ Euconnus (s. str.) paenetypicus nov. spec. MATERIAL: Holotypus d (Penispräparat) und 5 Paratypen, Poring Hot Springs, 25. bis 29.8.1988 (Ig. Smetana, CMG); ebenda 2 Paratypen (lg. Smetana, cF). DIAGNOSE: Weitgehend in den äußeren Merkmalen mit den aus dem Raum von Sabah beschriebenen Arten aus dem Subgenus Euconnus s. str. übereinstimmend, von diesen jedoch durch behaarte Schläfen und durch mit einer Querfurche verbundene Basalgrübchen des Halsschildes, sowie durch viel geringere Größe verschieden. BESCHREIBUNG: Long. 1,20 bis 1.40 mm. lat. 0,50 bis 0,55 mm. Rotbraun, Schläfen und Hasschildseiten braun behaart. Kopf von oben betrachtet queroval, flach gewölbt, die Schläfen etwas länger als der Augendurchmesser, dicht und steif abstehend behaart. Fühler zurückgelegt die Hals- schildbasis um das Endglied überragend, mit breiter, 4 gliederiger Keule, ihr 2. Glied um die Hälfte länger als breit, 3 bis 7 isodiametrisch, 8 bis 10 nicht ganz doppelt so breit wie 7, leicht gestreckt, das Endglied eiförmig, kürzer als die beiden vorletzten zusammen. Halsschild so lang wie breit, stark gewölbt, zum Vorderrand etwas stärker als zur Basis verengt, auf der Scheibe glatt und glänzend, an den Seiten struppig abstehend behaart, vor der Basis mit 2 großen und tiefen Grübchen, die bisweilen durch eine seichte Querfurche verbunden sind. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, glatt und glänzend, kahl, mit tiefer, lateral durch eine Humeralfalte begrenzter Basalimpression in einer im Halbkreis weit über sie hinaus auf die Flügeldecken reichenden Verflachung gelegen. Beine schlank, Vorderschenkel stärker verdickt als die der beiden anderen Beinpaare. Penis (Abb. 43) in dem einzigen vorliegenden Präparat bei der Entfernung von Lufteinschlüssen stark beschädigt und ein Teil der Parameren und des Penisinneren zerstört. Der Hinterrand des Penis ist gerade, die beiden Seiten springen aber spitzwinkelig vor. Von den beiden Paramen ist nur eine zum Teil erhalten, ihr distales Ende ist fußförmig verbreitert und im Spitzenbereich mit einer Anzahl von Borsten besetzt. Im Penisinneren befindet sich ein großer, von oben betrachtet etwa trapezförmiger Komplex, von dem hinter der Penisbasis ein tropfenförmiges Gebilde stark sklerotisiert erhalten ist. Euconnus (s. str.) paeneglaber nov. spec. MATERIAL: Holotypus 9 (Penispräparat) und 2 Paratypen, Kinabalu Nat. Park, Poring Hot Springs, 486 bis 495 m, 25.8.1988 (lg. Smetana, cMG) ebenda 2 Paratypen (1g. Smetana, cF). DIAGNOSE: Gekennzeichnet durch bis auf die Halsschildseiten kahlen Körper, lange, schlanke Fühler mit langer, 4-gliederiger Keule und schlanke Beine mit schwach ver- dickten Schenkeln. BESCHREIBUNG: Long. 1,50 mm, lat. 0,60 mm. Rotbraun, glatt und glänzend, Hals- schildseiten bräunlich behaart. Kopf von oben betrachtet annähernd kreisrund mit flachen Augen. Fühler zurückgelegt die Körpermitte erreichend, ihre 4-gliederige Keule fast so lang wie die Geißel, das Basalglied dicker als die folgenden, 2 um die Hälfte länger als breit, 3 bis 6 isodiametrisch, 7 breiter als lang, die Keule doppelt breiter als die Geißel, 8 bis 10 annähernd so lang wie breit, das eiförmige Endglied kürzer als die beiden vorletzten zusammen. Halsschild so lang wie breit, ungefähr so breit wie der Kopf, zum Vorderrand stärker als zur Basis verengt, stark gewölbt, vor der Basis ohne Grübchen, die Seiten abstehend behaart. SCYDMAENIDAE VON SABAH 899 Flügeldecken schon and der Basis zusammen beträchtlich breiter als der Halsschild, stark gewölbt, nur mit sehr kleiner Basalimpression, ohne Schulterbeule. Beine schlank, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 45 a, b) stark sklerotisiert, oberseits flach, ventral stark gewölbt, sein Apex in der Längsmitte tief eingeschnitten, das Operculum zungenförmig. Basalöffnung sehr groß, gerundet-querrechteckig, Parameren von oben betrachtet sehr breit, das Penisende nicht erreichend, mit 2 langen terminalen Tastborsten. Im Penisinneren liegt hinter der Basalöffnung eine stark sklerotisierte herzförmige Blase mit einem ins Ostium penis führenden Ausführungsgang. Lateral an diesem befindet sich eine langovale Drüse. Vor dem Ostium liegt quer zur Sagittalebene ein doppelt wellenförmig gebogenes Sklero- tinband. Der Apex penis besteht aus zwei breiten nach außen gebogenen Teilen, die durch einen basal spitzwinkelig zusammenlaufenden Spalt getrennt sind. Euconnus (s. str.) smetanaensis nov. spec. MATERIAL: Hototypus d (Penispräparat), Kinabalu Nat. Park Poring Hot Springs, 490 m, 20.8.1988 (lg. Smetana, cMG); ebenda 3 Ex. 20-21.8.1988 (CMG); ebenda, 1 Ex. 21. u. 25.8.1988 (lg. Smetana, cF). DIAGNOSE: Viel kleiner als die anderen Arten des Subgenus aus Sabah. Penisbau von den anderen Arten stark abweichend. BESCHREIBUNG: Long. 1,30 mm, lat. 0,60 mm. Braunschwarz, Tibien und Tarsen rotbraun, Palpen hellgelb. Stark glänzend, Halsschild braun behaart. Kopf von oben betrachtet mit den großen Augen queroval, so breit wie der Halsschild. Fühler zurückgelegt die Halsschildbasis erreichend, mit lockerer, 4 gliederige Keule, das Basalglied dicker als die folgenden, 2 gestreckt, 3 bis 7 kugelig, 8 knapp doppelt so breit wie 7, wie auch 9 und 10 nicht ganz so lang wie breit, das eiförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild so lang wie breit, zum Vorderrand stark, zur Basis nur wenig verengt, seitlich abstehend behaart, vor der Basis ohne Grübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, glatt und glänzend, mit kleiner, lateral von einer Humeralfalte begrenzter Basalimpression. Beine schlank, Vorderschenkel aber beim d sehr stark keulenförmig verdickt, Schienen gerade. Penis (Abb. 45) von oben betrachtet mehr als doppelt so lang wie breit, mit sehr großer Basalöffnung, sein Apex aus 2 am Ende gegeneinander gekrümmten Spitzen bestehend. Zwischen der Basis derselben ragt der Ductus ejaculatorius ins Freie. Auch das Operculum ist zweiteilig, jeder Teil endet in einer stumpfen Spitze, die lateralwärts gedreht ist. Die Parameren sind gerade und tragen je eine terminale Tastborste. Sie erreichen nur die Längsmitte des Apex penis. Euconnus (s. str.) apicefurcatus nov. spec. MATERIAL: Holotypus d (Penispräparat) und 2 Paratypen (davon 14 Penispräparat), Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Bruckhardt u. Löbl, cMG); ebenda 23 4 (Penispräparate, cF); Kinabalu Nat. Park, area Langanan 485 m, 28.8.1988 (lg. Smetana, CMG); Poring Hot Springs, 480 m, 27.8.1988 (Ig. Smetana, CMG); Mont Kinabalu, 1450 bis 1550 m, 23.5.1987 (Ig. Burckhardt u. Löbl, cF). DIAGNOSE: Kleiner als E. pseudosukhotanus, Fühler viel kürzer, Penis ganz anders gebaut. 900 HERBERT FRANZ ; BESCHREIBUNG: Long. 1,70 bis 1,80 m. Kastanienbraun, die Beine hell rotbraun, Halsschild bräunlich behaart. Kopf von oben betrachtet rund, etwas breiter als lang, flach gewölbt, Augen flach, Fühler zurückgelegt die Halsschildbasis nicht erreichend, mit lockerer viergliederiger Keule, ihr 2. und 7. Glied um etwa ein Drittel länger als breit, 3 bis 6 isodiametrisch, 8 bis 10 doppelt so breit wie 7, so lang wie breit, das eiförmige Endglied wesentlich kürzer als die beiden vorletzten zusammen. Halsschild kaum merklich länger als breit, kugelig, kurz und abstehend behaart, ohne Basalgrübchen, so breit wie der Kopf mit den Augen. Flügeldecken sehr stark gewölbt, schon an der Basis zusammen wesentlich breiter als die Halsschildbasis, sehr fein punktiert, glänzend. Flügel verkümmert. Beine ziemlich kurz, Vorderschenkel des 9 stark verdickt, Schienen dünn, die der Mittelbeine mediodistal leicht ausgeschnitten. Penis (Abb. 46) von oben betrachtet mehr als doppelt so lang wie breit, der Peniskörper nahezu parallelseitig, die Basalöffnung nicht ungewöhnlich groß, die Parameren die Basis des Apex penis erreichend, mit je 2 terminalen Tastborsten. Apex penis aus zwei fußförmig endenden Teilen bestehend, Operculum zungenförmig, nur die Längsmitte des Apex erreichend. Vor dem Ostium penis stehen im Penisinneren 2 kleine in der Ruhelage distalwärts gerichtete Stachel. BESTIMMUNGSTABELLE DER IN SABAH VORKOMMENDEN ARTEN AUS DEM SUBGENUS Eucon- nus S. Str. I Klein!Körperlänse 1320) bis) 1 AQ mim 22.0 nenn. ONE Scene eee eens 2 =. Großer, Körperlänge über ll, SO min = 5 2 Schlafen unbehaart, Halsschild ohne Basalgrübchen .................. smetanaensis n. Sp. — Schlafen behaart, Halsschild mit 4 durch eine Querfurche verbundenen Grübchen...... Sc 108 RT a RI IR ce paenetypicus n. sp. 3) Körperlänge 1,50 bis 1,80 mm, robust gebaut... 2... ace ce Se 4 — » Körperlänge mindestens 1,80.mm (meist über 220 mm). LE 5 4 Körperlänge 1,50 mm, Fühler zurückgelegt die Körpermitte erreichend ................... KERN ea te Se fac tente né AS et een paenaglaber n. sp. — Körperlänge 1,70 bis 1,80 mm, Fühler zurückgelegt die Halsschildbasis nicht Enreichendir. an. A ee RE NE Tan apicefurcatus n. sp. 5 Mittelschienen des d sehr stark medialwärts gekrümmt (hierher 3 nur durch den Penisbau unterscheidbare Arten simillimus n. sp., latus n. sp. und kinabaluanus n. sp.) ... =, Mittelschienen ganz gerade... ea RE eee ER RER Eee 6 6 Vorderschenkel des d sehr stark keulenförmig verdickt, halb so breit wie der Kopf lang, mittlere Fühlergeißelglieder isodiametrisch ................... allosukhotanus n. Sp. — Vorderschenkel des d weniger stark verdickt, mittlere Glieder der Fühlergeißel leicht DÉSERT e N pseudosukhotanus n. Sp. Untergattung Borneoconnus nov. subgen. Die Arten der neuen Untergattung der Gattung Euconnus Thomson sind durch beim dé monströses 9. Fühlerglied der 3 gliederigen Fühlerkeule, durch langgestreckten Kopf mit kleinen Augen und langen, bärtig behaarten Schläfen, durch konischen Halsschild mit großen Basalgrübchen und durch den Bau des männlichen Kopulationsapparates gekenn- zeichnet. Dem Aedeagus fehlt eine deutlich abgegrenzte Apikalpartie. 901 SCYDMAENIDAE VON SABAH ABB. 52-57. 52: Euconnus (Borneoconnus) eremita nov. spec., Penis in Ventralansicht. 53: Euconnus (Napochus) kinabalui nov. spec., Penis in Dorsalansicht. 54: Euconnus (Napochus) mirus nov. spec., Penis in (Napochus) allomirus nov. spec., Penis in Dorsalansicht. 47: Euconnus (Napochus) layangensis nov. spec., Penis in Dorsalansicht Dorsalansicht. 55: Euconnus (Napochus) paramirus nov. spec., Penis in Dorsalansicht.56: Euconnus 902 HERBERT FRANZ - Zur Typusart bestimme ich die nachfolgend an erster Stelle beschriebene Euconnus (Borneoconnus) laticlava. Euconnus (Borneoconnus) laticlava nov. spec. MATERIAL: Holotypus d (Penispräparat) und 2 Paratypen (16 Penispräparat), Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda d (Penispräparat) (cF); Kinabalu Nat. Park, area Langanan, 485 m, 22.8.1988 (1g. Smetana, CMG); Poring Hot Springs, 480 m, 27.8.1988 (lg. Smetana, CMG); Mount Kinabalu, 1450 bis 1550 m, 23.5.1987 (lg. Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch von oben betrachtet rautenförmigen Kopf, der so lang ist wie der Halsschild und durch den Fühlerbau beim d (Abb. 47). BESCHREIBUNG: Long. 1,70 bis 1,80 mm, lat. 0,55 bis 0,60 mm. Hell rotbraun bis .dunkelbraun, gelblich behaart. Kopf um ein Drittel länger als mit den flachen Augen breit, mit diesen so breit wie der Halsschild in seiner Längsmitte, an den Seiten steif abstehend behaart, die Schläfen zweieinhalbmal so lang wie der Augendurchmesser, Fühler zurückgelegt nur die Hals- schildmitte erreichend, das Basalglied doppelt so lang wie breit, das 2. Glied schwach, die folgenden stärker quer, das 8. sehr kurz, das 9. beim d so breit wie das 3. bis 8. Glied lang, mehr als doppelt so breit wie lang, nach außen stärker erweitert als nach innen, distalwärts gebogen und in einer Spitze endend, das 10. mit schrägen Seiten, das Endglied gerundet-kegelförmig, halb so breit wie das 10. Halsschild schmal konisch, so lang wie an der Basis breit, die Seiten steif abstehend behaart, mit 2 großen, in die Quere gezogenen Basalgrübchen. Flügeldecken langoval, schon an der Basis zusammen wesentlich breiter als der Hals- schild, mit tiefer, scharf begrenzter Basalimpression, fein und ziemlich dicht punktiert, anliegend behaart. Flügel verkümmert. Beine kurz, Schenkel etwas verdickt, Schienen fast gerade. Penis (Abb. 48) von oben betrachtet glockenförmig mit dorsobasal gelegener, mit einem breiten Sklerotinrahmen versehener Basalöffnung und dorsoapikal gelegenem Ostium penis. Parameren gerade, den Penisseiten eng anliegend, das Penisende nicht ganz erreichend, im Spitzenbereich mit mehreren langen Tastborsten. Ein Apex penis ist nicht vorhanden, vielmehr schließt die Dorsalwand des Pensis geradlinig ab. Eine ventral nach hinten vorragende Spitze ist wohl als Operculum zu deuten. An Stelle normaler Genitalorgane ist im Inneren des Penis nur ein von der Basis bis ins distale Drittel des Penis reichender dicker Balken zu erkennen. Im Bereich des Ostiums sind übereinander 4 horizontale Ebenen erkennbar: zuunterst das “Operculum”, darüber eine horizontale Platte, die nur wenig über die Dorsalwand des Penis vorragt und von deren von oben und hinten betrachtet rechter Seite ein schmaler Sklerotinstab querüber nach links gerichtet ist. In der 3. Ebene liegt ein Sklerotinrahmen, der rechts schmal und links breit ist und eine große querovale Öffnung umfaßt, die vielleicht das Ostium penis ist. Die oberste Ebene wird durch die Dorsalwand des Penis gebildet. Euconnus (Boreoconnus) sabahanus nov. spec. MATERIAL: Holotypus & und 256 Paratypen (Penispräparat) sowie 142 Paratypen, Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl cMG); ebenda 96 69 Paratypen (cF); ebenda, ca. 1000 m, 28 5? Paratypen, cMG (1g. Smetana, cMG); ebenda, 439 (Paratypen), 27.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, d 49 Paratypen, Livago Ricer, 1490 bis 1500 m (lg. Burckhardt u. Löbl, CMG); Poring Hot Springs, 98 72 Paratypen (lg. Burckhardt u. Löbl (cF); Mount Kinabalu, 2000 m, 29 Paratypen (CMG); Crocker Range, 7 km N Tambunan, 700 m, 20.5.1987 (CMG). SCYDMAENIDAE VON SABAH 903 DIAGNOSE: Fühler mit scharf abgesetzter, 3 gliederiger Keule, das 9. Glied beim dé asymmetrisch, außenseitig erweitert, das 10. groß, regelmäßig rechteckig. Halsschild konisch mit 2 großen Basalgrübchen (Abb. 49). BESCHREIBUNG: Long. 1,15 bis 1,30 mm. Kastanienbraun, die Beine heller, braun behaart. Kopf von oben betrachtet länglich-eirund, im Niveau der kleinen, flachen Augen am breitesten, von da zur Basis in flachem Bogen verengt, die Schläfen dreimal so lang wie der Augen- durchmesser, schräg abstehend steif und kurz, nach hinten gerichtet behaart. Fühler zurück- gelegt die Halsschildbasis nicht erreichend, mit sehr scharf abgesetzter, 3 gliederiger Keule, ihr Basalglied und das 2. etwas länger als breit, 3 bis 8 klein, breiter als lang, das 9. Glied sehr breit, beim d asymmetrisch, rechts viel stärker erweitert als links, aber viel weniger stark als bei E. laticlava, 10 rechteckig, fast so lang wie breit, das eiförmige Endglied ist nur so lang wie das 10. und viel schmäler als dieses. Hals nur ein Drittel so breit wie die Halsschildbasis. Halsschild konisch, länger als breit, mit 2 sehr großen und tiefen Basalgruben, seitlich abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit runder Basalimpression, ohne Schulterbeule, nach hinten gerichtet behaart. Flügel verkümmert. Penis (Abb. 50) im Bau dem des E. laticlava ähnlich, von oben betrachtet 3 übereinander gelegene Ebenen erkennen lassend, zu oberst eine rechteckig begrenzte, am wenigsten weit distalwärts reichende, die von einem sklerotisierten Rahmen umgeben ist. Über ihr liegen die Parameren. Darunter folgt eine zweite distal im flachen Bogen begrenzte mittlere Platte, unter der das Ostium penis liegt und unter der sich der Ductus ejaculatorius befindet. Er entspringt in einer länglichovalen Blase unter der Basalöffnung des Penis und verläuft zunächst S-förmig gekrümmt bis nahe an das Penisende, biegt dann von oben und hinten betrachtet im flachen Bogen nach rechts und weiter in einem weiten Bogen über die rechte Penisseite wie der nach vorne und zurück zur Penismitte. Dabei verschmälert er sich zu einem dünnen Rohr. Unter ihm folgt auf der dritten Ebene das Operculum, dessen Hinterrand gerundet-rechteckig ist und nur wenig über die mittlere Platte nach hinten vorragt. Die Basalöffnung liegt dorsobasal und besitzt einen breiten Sklerotinrahmen, von dessen distaler Seite entspringen die Parameren, die am distalen Ende 3 lange Tastborsten tragen. Je 2 von diesen sind sehr lang und am Ende zur Sagittalebene gebogen. Sie überragen das Penisende. Euconnus (Borneoconnus) eremita nov. spec. MATERIAL: Nur Holotypus 4 (Penispräparat), Mount Kinabalu, 2995 m, Paka Cave, 5.2.1987 (lg. Smetana, cMG). DIAGNOSE: Größer und vor allem breiter als E. laticlava, die Fühler länger, die Geißelglieder von einander wenig verschieden, die Schläfen stark gerundet, auch die Flügeldecken stärker gerundet als bei den Vergleichsarten, nur wenig länger als Kopf und Halsschild zusammen (Abb. 51). BESCHREIBUNG: Long. 1,90 mm, lat. 0,60 mm. Hell rotbraun, gelblich behaart. Kopf gerundet- rautenförmig, nicht ganz so lang wie der Halsschild, der Hals weniger als halb so breit wie der Vorderand des Halsschildes, die Schläfen zu ihm gerundet verengt, seitlich abstehend behaart, die Stirn unter der Fühlern flach, leicht eingetieft. Fühler zurückgelegt beinahe die Halsschildbasis erreichend, ihre Geißel deutlich länger als die Keule, die beiden ersten Fühlerglieder um die Hälfte länger als breit, das 3. bis 6. Glied fast so lang wie breit, 7 und 8 etwas breiter und deutlich quer, die Keule fast 4 mal so breit wie sie. Glied 9 und 10 fast symmetrisch, 9 nach außen distal auf das Dreifache verbreitert, 10 trapezförmig noch etwas breiter als 9 und distal schwach verbreitert, das eiförmige Endglied etwas schmäler als das vorhergehende. 904 HERBERT FRANZ - Halsschild an der Basis nur wenig breiter als am Vorderrand, abstehend, an den Seiten dichter als auf der Scheibe behaart, vor der Basis mit 2 mäßig großen Grübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, seitlich gleichmäßig gerundet, mit kleiner, mäßig tiefer Basalimpression, abstehend behaart. Flügel verkümmert. Beine verhältnismässig länger als bei den beiden anderen Arten der Untergattung, Schenkel schwach verdickt, Mittel- und Hinterschienen schwach medialwärts gebogen. Penis (Abb. 52) von oben betrachtet annähernd zylindrisch mit basal gelegener Basal- öffnung und terminal gelegenem Ostium penis. Parameren das Penisende nicht ganz erreichend, mit je 3 langen terminalen Tastborsten. Im Penisinneren befindet sich vor der Penismitte eine kleine länglichovale Blase mit breitem, nur bis zum distalen Drittel des Penis reichendem Ausführungsgang. An diesen schließt von oben und hinten betrachtet rechts ein breites sklerotisiertes Feld an, an dem keine weiteren Differenzierungen erkennbar sind. Der Hinterand der Dorsalwand des Penis ist bandförmig stärker sklero- tisiert. Unter ihm ragt eine weitere sklerotisierte Fläche vor, die am Hinterrand einen eiförmig gekrümmten Stachel nach rechts entsendet. Darunter findet sich in einer dritten Ebene eine dritte Sklerotinplatte, die vielleicht das Operculum repräsentiert. Ein Apex penis ist auch bei dieser Art nicht entwickelt. Untergattung Napochus Reitter Euconnus (Napochus) kinabalui nov. spec. MATERIAL: Holotypus d (Penispräparat) Poring Hot Springs, 550 bis 600 m, 9.5.1987 (lg. Burckhardt u. Löbl, cMG); 2 Paratypen Mount Kinabalu, 2600 m, 1.5.1987 (lg. Burckhardt u. Löbl (cF); ebenda. 2 Paratypen (CMG); 2 Ex. Paratypen, Mount Kinabalu, 1500 m, 21. u. 15.5.1987 (Ig. Burckhardt u. Löbl, cMG); ebenda 2 Paratypen, 25.5.1987 (cF); 1 Paratypus, ebenda, 1750 m, 24.4.1987 (CMG); Mont Kinabalu, Sumit Tr. Podok, 2300 bis 2499 m, 29.4.1987 (lg. Smetana, cMG); 1 Paratypus, Crocker Range, Kota Kinabalu, Tambunan, 18.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Klein, hell rotbraun, Kopf isodiametrisch, Fühler zurückgelegt die Hals- schildbasis um die letzten 2 Glieder überragend, Halsschild isodiametisch, Flügeldecken mit langer Humeralfalte. BESCHREIBUNG: Long. 1,00 bis 1,20 mm, lat. 0,45 bis 0,50 mm, hell gelbbraun, gelblich behaart. Kopf isodiametrisch-rund mit stark vorgewölbten Augen und steif abstehend behaarten Schläfen. Fühler mit lockerer, 4 gliederiger Keule, zurückgelegt die Halsschildbasis um die beiden letzten Glieder überragend, die beiden ersten Glieder leicht gestreckt, 3 bis 7 klein, schwach quer, 8 bis 10 rundlich, nicht ganz so breit wie lang, das Endglied rundlich, nicht ganz so lang wie breit. Halsschild konisch, isodiametrisch mit 2 Basalgrübchen, ohne basale Querfurche, abstehend, an den Seiten dichter und struppig behaart. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, seitlich stark gerundet, um ein Viertel länger als Kopf und Halsschild zusammen, fein punktiert, schräg abstehend behaart, mit tiefer, von einer schrägen Humeralfalte scharf begrenzter Basalimpression. Flügel ent- wickelt. Beine mit schwach verdickten Schenkeln und geraden Schienen. Penis (Abb. 53) von oben betrachtet etwas mehr als doppelt so lang wie breit, seine Basalöffnung weit auf die Dorsalseite gerückt, die Parameren an ihr mit der ganzen Breite ansetzend, das Penisende nicht annähernd erreichend, im Spitzenbereich mit je 2 Tastborsten. Am Hinterrand der Basalöffnung zieht ein breites Sklerotinband quer über die SCYDMAENIDAE VON SABAH 905 YSmm 1/5mm 110mm Inn u? if 62 1/5mm ABB. 58-63. 58: Euconnus (Napochus) valdeobscurus nov. spec., Penis in Dorsalansicht. 59: Euconnus (Napochus) parakinabalui nov. spec., Penis in Dorsalansicht. 60: Euconnus (Napochus) fuscus nov. spec., Penis in Dorsalansicht. 61: Euconnus (Napochus) funestus nov. spec., Penis in Dorsalansicht. 62: Euconnus ? (Napochus) sabahinanus nov. spec., Penis in Dorsalansicht. 63: Euconnus (Napochus) borneoi nov. spec., Penis in Dorsalansicht. 906 HERBERT FRANZ ‘ganze Dorsalwand des Penis. Hinter diesem Querband ist der Penis größtenteils durchsichtig, so daß der Ductus ejaculatorius von außen zu sehen ist. Er ist lang trichterförmig und am distalen Ende zu einem sehr dünnen Rohr verengt, das zur Seite gebogen ist. Der Apex penis ist vom Peniskörper nur schwach abgesetzt, distalwärts fast nicht verschmälert, sein Hinterrand gerundet, in der Mitte aber in einer kleinen Spitze vorspingend. Euconnus (Napochus) mirus nov. spec. MATERIAL: Holotypus 4 (Penispräparat) Mount Kinabalu Nat. Park, Poring Hot Springs area below Langanan Fall, 800 m, 12.5.1987 (lg. Smetana cMG); 2 Paratypen, Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG und cF); Mont Kinabalu, 2600 m, April 1987 (Ig. Burckhardt u. Löbl, cF); 5 Paratypen, Mount Kinabalu, 1500 bis 1600 m, 25.4.1987 (lg. Burckhardt u. Löbl, CMG); Poring Hot Springs, 550 bis 600 m, 9.6.1987 (1g Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch geringe Größe, tiefe Basalfurche des Halsschildes und eine große, beide Flügeldecken umfassende Verflachung hinter der Basalimpression. BESCHREIBUNG: Long. 0,95 mm bis 1,00 mm, lat. 0,50 mm. Schwarzbraun mit heller brauner Extremitäten, lang schwarzbraun behaart. Kopf von oben betrachtet queroval, mit großen, flach gewölbten Augen und langer abstehender Behaarung der Schläfen. Fühler zurückgelegt die Halsschildbasis erreichend, mit lockerer 4 gliederiger Keule une schlanker Geißel, ihre beiden ersten Glieder gestreckt, 3 bis 7 sehr klein, etwas breiter als lang, 8 bis 10 im distalen Drittel ihrer Länge am breitesten, von da zur Spitze abrupt verengt, das Endglied leicht gestreckt, sein Spitzenteil wie eine Eichel der Cupula aufsitzend. Halsschild konisch, so lang wie breit, an der Basis kaum breiter als der Kopf mit den Augen, struppig abstehend behaart, mit tiefer basaler Querfurche, in dieser mit 4 Grübchen. Flügeldecken kurzoval, an der Basis zusammen so breit wie die Halsschildbasis, distal aber stark verbreitert, mit breiter, lateral von einer langen Humeralfalte begrenzter Basal- impression, bis zur Mitte verflacht, fein punktiert und behaart. Flügel voll entwickelt. Beine kurz und schlank. Penis (Abb. 54) gedrungen gebaut, aus einem von oben besehen gerundet-kurz- rechteckigen Peniskörper und einem dünnhäutigen, zungenförmigen Apex bestehend. Parameren die Basis des Apex penis etwas überragend, mit je einer lateral vor der Spitze stehenden Tastborste. Basalöffnung ohne stark sklerotisierten Rahmen, nur sein Hinterrand durch Sklerotinkörnchen markiert. Im Penisinneren ist ein dessen größten Teil erfüllender Sklerotinkomplex vorhanden, der in halbkreisförmiger Begrenzung vom Hinterrand der Basalöffnung bis in den Bereich des Apex penis reicht. Dieser ist spitz- zungenförmig, das Operculum etwas kürzer und abgerundet-zungenförmig. Innerhalb des Sklerotinkomplexes im Penisinneren fallen zwei annähernd kugelige, etwa in der Längsmitte des Penis, spiegelbildlich zur Sagittalebene liegende Zysten auf. Die von oben und hinten besehen linke entsendet basalwärts ein annähernd dreiblättriges Gebilde, das rechte ein breites Sklerotinband. Lateral von den beiden Zysten steht auf beiden Penisseiten ein langgestreckter Sklerotinkörper. Euconnus (Napochus) paramirus nov. spec. MATERIAL: Holotypus 3 (Penispräparat), Sabah, Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, | Paratypus (cF); 2 Paratypen, Mount Kinabalu, 2600 m, April 1987 (lg. Burckhardt u. Löbl, cF); 1 Paratypus, Mount Kinabalu Nat. Park, HQ at Livago River, 1500 m, 1.9.1988 (lg. Smetana, cMG). SCYDMAENIDAE VON SABAH 907 DIAGNOSE: Dem E. (Napochus) mirus sehr ähnlich, aber die Fühler länger, zurückgelegt die Halsschildbasis überragend, ihre Keule länger als die Geißel, die Flügeldecken länger als Kopf und Halsschild zusammen, der Apex penis (Abb. 55) etwas kürzer, die im Penisinneren liegenden Zysten exakter kugelförmig mit einem opaken Kern, zwischen ihnen steht in der Sagittalebene an Stelle des dreiblättrigen Gebildes ein Sklerotintrichter, dessen Hals den Apex penis nicht erreicht. Euconnus (Napochus) allomirus nov. spec. MATERIAL: Holotypus d (Penispräparat). Sabah, Mount Kinabalu, 1500 m, 25.4.1987 (le. Burckhardt u. Löbl, cMG): Paratypus 2, Kinabalu 1550 m, 28.4.1987 (lg. Bruckhardt u. Löbl, cF). DiAGnosE: Äußerlich dem Euconnus mirus ähnlich, aber wesentlich größer, die Fühlerkeule länger als die Geißel, der Halsschild mit sehr tiefer basaler Querfurche, die Flügeldecken ohne deutliche Punktierung, der Penis ganz anders geformt. BESCHREIBUNG: Long. 1,20 mm, lat. 0,50 mm. Dunkel rotbraun, gelbbraun behaart. Kopf von oben betrachtet gerundet-rautenförmig mit großen, stark vorgewölbten Augen und bärtig behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis erreichend, mit lockerer, 4 gliederiger Keule, diese länger als die Geißel, ihre beiden ersten Glieder so lang wie breit, 3 bis 7 sehr klein, 8 bis 10 reichlich doppelt so breit wie die vor- hergehenden, breiter als lang, mit dem für die Untergattung Napochus typischen Bau, das Endglied länger als die beiden vorletzten, sein distaler Teil schmäler als der basale, dem er eine Eichel ihrer Cupula aufsitzt. Halsschild konisch, etwas breiter als lang, an der Basis breiter als der Kopf mit den Augen, seitlich struppig abstehend behaart, mit auffällig tiefer Basalfurche. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, fein behaart, ohne deutlich erkennbare Punktierung. Flügel voll entwickelt. Beine schlank, Vorderschenkel etwas stärker verdickt als die der beiden anderen Beinpaare. Penis (Abb. 56) sehr eigenartig gebaut. Peniskörper von oben betrachtet oval mit einem langen, schmalen Apex und einem großen halbkreisförmig begrenzten Operculum. Basalöffnung sehr groß mit stark sklerotisiertem Rahmen, der an der Basis zu einem großen dreieckigen Lappen verbeitert ist. Mit der Basalöffnung sind die Parameren fest verwachsen, sie schließen an den dreieckigen Lappen mit einer Öse an, sind an der Basis breit, dann tief ausgebuchtet, zu letzt außer gerade, der Innenrand aber schwach gebogen, so daß die Parameren zur Spitze verschmälert sind. Sie tragen in der postbasalen Ausbuchtung 2 steife, schräg nach innen und hinten gerichtete Borsten. Im Spitzenbeich stehen terminal 2 lange Borsten und davor medial eine sehr kurze. Der Apex wurzelt unmittelbar am Hinterrand der Basalöffnung, die vom Vorderrand des Penis fast bis in dessen Längsmitte verschoben ist. Nach der trichterförmigen Basis ist der Apex sehr stark eingeschnürt und verläuft dann parallelseitig bis zur Spitze, vor der beiderseits eine kleine Ecke vorspingt. Innerhalb des Apex verläuft der Ductus ejaculatorius, an der Ein- schnürungstelle als enges, dahinter als weites Rohr. Euconus (Napochus) layangensis nov. spec. MATERIAL: Holotypus d, Kinabalu Nat. Park, Layang Layang, 2600 m, 2. bis 8.5.1987, in Falle (Ig. Smetana, cMG). DIAGNOSE: im Rahmen des Subgenus Napochus etwas abweichend, weil das Endglied der Fühler nicht deutlich in einen breiteren basalen und einen schmäleren distalen Teil gegliedert ist, sonst aber durchaus den Merkmalen des Subgenus Napochus entsprechend. 908 HERBERT FRANZ BESCHREIBUNG: Long. 1,60 mm, lat. 0,70 mm. Hell rotbraun, gelblich behaart. Kopf von oben betrachtet queroval, mit den großen vorgewölbten Augen so breit wie die Halsschildbasis, flach gewölbt, an den Schläfen und am Hinterkopf fein und abstehend behaart, Fühler mit lockerer, 4 gliederiger Keule, zurückgelegt die Halsschildbasis knapp erreichend, ihr Basalglied und das 2. doppelt so lang wie breit, 3 bis 7 klein, annähernd isodiametrisch, 8 doppelt so breit wie 7, 9 und 10 noch ein wenig breiter, alle 3 schwach quer, das gerundet-kegelförmige Endglied etwas länger als die beiden vorletzten zusammen. Halsschild kurz, etwas breiter als lang, konisch, mit 2 Basalgrübchen, fein punktiert und dicht, steif aufgerichtet behaart. Flügeldecken zusammen schon an der Basis etwas breiter als die Halsschildbasis, dicht und steif abstehend behaart. Flügel entwickelt. Beine schlank, Schenkel schwach verdickt. Penis (Abb. 57) mit von oben betrachtet langovalem Peniskörper, spitzwinkelig- dreieckigem Apex und den Apex überragendem Operculum. Parameren die Mitte des Apex kaum erreichend, schlank, mit je 2 terminalen Tastborsten. Basalöffnung des Penis breit, nur ihr Basalrand stark sklerotisiert. Im Inneren des Peniskörpers befindet sich ein ausgedehnter sklerotisierter Komplex, in dem von einem zentralen kompakten Bereich 4 große sichelförmig gekrümmte Stachel distalwärts ausgehen. Von oben betrachtet links befinden sich neben einander 2 nach links außen gekrümmte Stachel und rechts der Sagittalebene ein nach rechts und daneben außen ein nach links gekrümmter Stachel. Der Komplex wird distalwärts durch eine Querleiste begrenzt. Euconus (Napochus) quinquearticulatus nov. spec. MATERIAL: Holotypus 9, Mount Kinabalu Nat. Park, 1550 bis 1650 m, HQ, 24.4.1987 (Ig. Smetana, cMG); Paratypus 2, Pondok Lowii, 2300 bis 2400 m, 28.8.1988 (lg. Smetana, cF). BESCHREIBUNG: Long. 2,10 mm, lat. 1,10 mm. Schwarz, Palpen, erste Fühlerglieder und Beine dunkel-rotbraun, braun behaart. Kopf von oben betrachtet gerundet-rautenförmig, mit sehr großen, seitlich vorgewölbten Augen, so breit wie lang, am Scheitel schütter, weißlich, nach hinten, an den Schläfen seitlich abstehend behaart. Fühler mit scharf abgesetzter 5 gliederiger Keule, die Geißelglieder leicht gestreckt, das Basalglied und das 2. etwas dicker als die folgenden, 7 doppelt so breit wie 6, 8 bis 10 noch etwas breiter, fast so breit wie lang, das Endglied um ein Drittel länger als das vorletzte, sein distaler Teil eichelartig vom basalen abgeschnürt. Halsschild konisch, an der Basis doppelt so breit wie am Vorderrand und hier breiter als lang, mit basaler Querfurche, in dieser mit 3 seichten Grübchen, auf der Scheibe schütter, an den Seiten dichter, abstehend behaart. Flügeldecken kurzoval, sehr breit, schon am Vorderrand zusammen um die Hälfte breiter als die Halsschildbasis, mit einer lateral von einer Schulterbeule begrenzten Basal- impression, lang und abstehend behaart. Flügel voll entwickelt. Beine schlank, Schenkel sehr schwach verdickt. Euconnus (Napochus) ventriculosus nov. spec. MATERIAL: Nur Holotypus 2, Mount Kinabalu, 1550 bis 1650 mm, 24.4.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Wie die vorhergehende Art durch 5 gliederige Fühlerkeule ausgezeichnet, alle Keulenglieder deutlich gestreckt, das letzte, wie für das Subgenus kennzeichnend, 2- SCYDMAENIDAE VON SABAH 909 ABB. 64-71. 64: Euconnus (Napoconnus) cephalotes nov. spec., Penis in Dorsalansicht. 65: Euconnus (Napo- connus) parallelipenis nov. spec., Penis in Dorsalansicht. 66: Euconnus (Napoconnus) valdepullus nov. spec., Penis in Dorsolateralansicht. 67: Euconnus (Napoconnus) langananensis nov. spec., Penis in Dorsalansicht. 68: Euconnus (Napoconnus) elongatior nov. spec., Penis in Lateralansicht. 69: Euconnus (Napoconnus) tambunanus nov. spec., Penis in Dorsolateralansicht. 70: Euconnus (Napo- connus) proceripenis nov. spec., Penis in Dorsalansicht. 71: Euconnus globicollis nov. spec. Penis in Lateralansicht. 910 HERBERT FRANZ ‚teilig, mit schmälerem distalem Teil, mit schmälerem Kopf, der Hinterkopf beulenförmig über den Hals vorgewölbt, die Flügeldecken sehr stark bauchig verbreitert. BESCHREIBUNG: Long. 1,90 mm, lat. 1,0 mm. Schwarzbraun, Palpen, Fühler und Beine hellbraun (? immatur). Kopf von oben betrachtet gerundet-rautenförmig, mit wenig vorragendem Apex, Scheitel, Hinterkopf und Schläfen dicht und abstehend behaart, der Scheitel als Beule ziemlich spitz über den Hinterkopf basalwärts vorragend. Hallschild breiter als lang, sein Basalrand aber nur um ein Drittel breiter als der Vorderrand, die Seiten steif abstehend behaart, vor der Basis mit 3 großen Grübchen. Flügeldecken sehr stark bauchig erweitert, zusammen schon an der Basis wesentlich breiter als die Halsschildbasis, mit seichter, lateral von einer Humeralfalte begrenzter Basalimpression, sehr fein punktiert und schütter behaart. Flügel voll entwickelt. Beine schlank, Schenkel kaum verdickt. Euconus (Napochus) burckhardtianus nov. spec. MATERIAL: Nur Holotypus & und Paratypus ©, Sabah, Poring Hot Springs, 500 m, 7.5.1987 (lg. Burckhardt u. Löbl, cMG); und 10.5.1987, (cF). DIAGNOSE: Eine durch ihre Größe und die relativ langen Fühler auffallende Art, die im Bau des männlichen Kopulationsapparates den Arten mit nach oben gebogenem Apex penis zuzuordnen ist. Einen ähnlich gebauten Penis besitzt von den Napochus-Arten aus Sabah vor allem N. parakinabalui, der aber wesentlich kleiner ist und kürzere Fühler besitzt, darüber hinaus auch mit E. (Napochus) fricatoris Schauf. aus Singapore und seinen Verwandten, die aber wesentlich größer sind und bei denen das 8. bis 10. Fühlerglied nur wenig breiter als lang ist. BESCHREIBUNG: Long. 1,60 mm, lat. 0,80 mm, braungrau, braun behaart. Kopf von oben betrachtet rundlich, etwas breiter als lang, die großen Augen seitlich stark vorgewölbt, die Oberseite lang und abstehend, die Schläfen sehr dicht und seitlich steif abstehend behaart. Fühler dick, zurückgelegt die Halsschildbasis nicht ganz um das Endglied überragend, ihr 2. Glied quadratisch, 3 bis 7 breiter als lang, 8 und 9 doppelt so breit wie 7, so lang wie breit, 10 sehr schwach quer, das Endglied etwas länger als breit, sein distaler Teil schwach stufenförmig abgesetzt und etwas schmäler als der basale. Halsschild konisch, etwas breiter als lang, an der Basis ein wenig breiter als der Kopf mit den Augen, dicht, an den Seiten sehr dicht und struppig abstehend behaart, vor der Basis mit einer beiderseits von einem Grübchen begrenzten Querfurche. Flügeldecken zusammen schon an der Basis viel breiter als die Halsschildbasis, sehr lang und abstehend behaart, mit tiefer, länglicher Basalimpression. Flügel voll entwickelt. Schenkel schwach verdickt, Schienen schlank. Penis im einzigen vorliegenden Präparat sehr stark geschrumpft und deshalb nicht abbildbar, mit großer, von einem sklerotisierten Rahmen umgebener, dorsal nahe der Penisbasis gelegener Basalöffnung. Parameren am Präparat nicht erhalten, Apex wie bei E. parakinabalui aufgebogen, breit zungenförmig, Operculum flach, annähernd so lang wie der Apex, sein Hinterrand spitzbogenförmig. Euconnus (Napochus) valdeobscurus nov. spec. MATERIAL: Holotypus ¢, Mount Kinabalu, 1550 bis1650 m, 24.4.1957 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus, Mount Kinabalu, Nat. Park, Poring Hot Springs, 450 bis 500 m, 30.5.1988 (lg. Smetana, cF). DIAGNOSE: Ausgezeichnet durch tief schwarze Körperfarbe und dichte, schwarz- braune Behaarung. SCYDMAENIDAE VON SABAH 91] BESCHREIBUNG: Long. 1,50 mm, lat. 0,70 mm. Schwarz glänzend, die Beine dunkel rotbraun. Kopf von oben betrachtet rundlich, mit den flach gewölbten Augen ein wenig breiter als lang, dicht und lang schwarzbraun behaart, Fühler zurückgelegt die Halsschildbasis knapp erreichend, die breite Keule scharf abgesetzt, die beiden ersten Glieder um ein Viertel länger als breit, 3 bis 7 klein, schwach quer, 8 bis 10 dreimal so breit wie 7, beim d stark beim © schwach quer, das Endglied an der Spitze abgerundet, so lang wie breit, in der Längsmitte stufig auf die halbe Breite verschmälert. Halsschild breiter als lang, an seiner Basis deutlich breiter als der Kopf mit den Augen, dicht und abstehend behaart, beim d mit 3 in einer seichten Querfurche stehenden Grübchen, beim 2 ohne solche. Flügeldecken schon an der Basis zusammen wesentlich breiter als die Halsschildbasis, stark gewölbt und seitlich stark gerundet, mit tiefer, lateral von einer langen Humeralfalte scharf begrenzter Basalimpression, fein und seicht, aber dicht punktiert, schräg abstehend behaart. Beine sehr schlank, Schenkel kaum verdickt, Schienen im distalen Drittel leicht ver- breitert, Tarsen sehr zart. Penis (Abb. 58) von oben betrachtet breit, der Peniskörper gerundet-trapezförmig, der Apex spitzwinkelig-dreieckig, an seiner Basis nur ein Drittel so breit wie der Peniskörper, die Basalöffnung dorsobasal gelegen, das Ostium penis offenbar terminal, der Ductus ejaculatorius scheint bis zur Penisspitze zu reichen, er entspringt am distalen Ende des Peniskörpers aus einem stark sklerotisierten Komplex, der von einer durchsichtigen Cuticula umhüllt ist. Parameren sind an dem einzigen vorliegenden Präparat nicht vorhanden, sie sind vermutlich bei der Präparation verloren gegangen. Euconnus (Napochus) parakinabalui nov. spec. MATERIAL: Holotypus d (Penispräparat) und 3 Paratypen, Mount Kinabalu, 2600 m, April 1987 (lg, Burckhardt u. Löbl cMG); ebenda 2 d (Penispräparat und 8 Paratypen (cF); 1 Paratypus Mount Kinabalu 1450 bis 1500 m (cMG); 2 Paratypen Crocker Range, 1559 bis 1650 m, 16.5.1987 (cMG); 2 Paratypen, Mount Kinabalu, Layang Layang, 2590 m, 1. bis 2.5.1987 (lg. Smetana, cMG); 1 Ex. Crocker Range, ca. 1000 m, 5.9.1988 (lg. Smetana, cMG); 1 Ex. Mount Kinabalu, Livagu River, 1.9.1988 (lg. Smetana, cMG); 2 Ex. Poring Hot Springs, 500 bis 600 m. 5-5. u. 7-6.1987 (lg. Burckhardt, cMG). DIAGNOSE: Dunkel rotbraun, die 4 gliederige Fühlerkeule länger als die Geißel, scharf abgesetzt, Halsschild mit basaler Querfurche. BESCHREIBUNG: Long. 1,10 bis 1,20 mm, lat. 0,60 mm. Dunkel rotbraun, bräunlich behaart. Kopf von oben betrachtet isodiametrisch rund mit vorgewölbten Augen, die Schläfen eineinhalbmal so lang wie der Augendurchmesser, abstehend behaart. Fühler zurückgelegt die Halsschildbasis nicht ganz erreichend, ihre Geißel etwas kürzer als die 4 gliederige Keule, die beiden ersten Glieder so lang wie breit, 3 bis 7 sehr klein, breiter als lang, 8 3- mal so breit wie 7, schwach, 9 und 10 etwas stärker quer, das Endglied gerundet- kegelförmig, nur wenig länger als breit. Halsschild konisch, an seiner Basis wenig breiter als der Kopf mit den Augen, seine Seiten struppig abstehend behaart, vor der Basis mit 2 großen Grübchen. Flügeldecken zusammen an der Basis etwas breiter als die Halsschildbasis, oval, wenig länger als Kopf und Halsschild zusammen, fein punktiert und schräg abstehend behaart, mit langer, von einer schrägen Humeralfalte begrenzten Basalimpression. Flügel ver- kümmert. 912 HERBERT FRANZ _Beine schlank und mäßig lang, Schenkel sehr schwach verdickt, Schienen gerade. Penis (Abb. 59) von oben betrachtet annähernd doppelt so lang wie breit, mit nach oben gebogenem, am Ende breit abgestutztem Apex. Basalöffnung dorsal gelegen mit breitem sklerotisierten Rahmen, Parameren leicht S-förmig gebogen, das Penisende fast erreichend, mit je 2 terminalen Tastborsten. Euconnus (Napochus) fuscus nov. spec. MATERIAL: Holotypus d, Kinabalu Nat. Park, Poring Hot Springs, 475 m, 23.8.1988 (Ig. Smetana, cMG); 1 Paratypus, ebenda 11.5.1987 (lg. Burckhardt u. Löbl, cF); Mount Kinabalu Nat. Park, Livagu River, 490 m, 18.5.1987 (lg. Smetana, cMG). DIAGNOSE: rotbraun stark glänzend, Halsschild mit basaler Querfurche und darin mit 4 Griibchen, Fliigeldecken mit langer, lateral von einer langen Humeralfalte begrenzter Basalimpression. BESCHREIBUNG: Long. 1,30 mm, lat. 0,65 mm. Dunkel rotbraun, die Extremitäten heller gefärbt. Kopf von oben betrachtet queroval mit seitlich stark vorgewölbten Augen und bärtig behaarten Schläfen. Halsschild konisch, an der Basis nur wenig breiter als der Kopf mit den Augen, mit breiter basaler Querfurche und darin mit 4 Grübchen, die Seiten abstehend behaart. Flügeldecken stark gewölbt und seitlich stark gerundet, glatt und glänzend, schütter, nach hinten gerichtet behaart, mit tiefer, lateral von einer Humeralfalte begrenzter Basal- impression. Beine kurz, Schenkel schwach verdickt, Schienen distalwärts etwas verbreitert. Penis (Abb. 60) von oben betrachtet kurzoval, mit annähernd trapezförmigem, scharf abgesetzten Apex. Seine Basalöffnung auf der Dorsalseite nahe zur Penismitte verschoben, von einem sehr breiten Sklerotinrahmen umgeben, Parameren an dessen distalem Rand inserierend, stark nach außen und dann wieder zur Mitte gebogen, das Penisende erreichend, mit je 2 terminalen Tastborsten. Euconnus (Napochus) funestus nov. spec. MATERIAL: Holotypus d (Penispräparat) und 4 Paratypen, Poring Hot Springs, 500 bis 550 m, 6. u. 12.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda 3 Paratypen (cF); 2 Paratypen, ebenda (lg. Smetana, cMG); 1 Paratypus, ebenda, 23.8.1988 (lg. Smetana, cMG); 1 Paratypus, Mount Kinabalu, Summit Pondok Ubah, 2050 m, 26.4.1987 (lg. Smetana, CMG). DIAGNOSE: Gekennzeichnet durch schwarze Körperfarbe, sehr lange Fühlerkeule und gedrungenen Fühlerbau. BESCHREIBUNG: Long. 1,50 bis 1,60 mm, lat. 0,65 mm, Schwarz, die Extremitäten hellbraun, weißlich behaart. Kopf von oben betrachtet gerundet-rautenförmig, mit den stark vorgewölbten Augen breiter als lang, oberseits flach, Schläfen und Hinterkopf dicht und abstehend behaart. Fühler zurückgelegt die Halsschildbasis erreichend, mit 4 gliederiger Keule, das 7. Glied aber schon breiter als das 6., die beiden ersten Glieder nicht ganz so lang wie breit, 3 bis 7 klein, 8 bis 10 doppelt breiter als 7, schwach quer, das 11. Glied querüber abgeschnürt, der distale Teil schmäler als der basale. Halsschild konisch, an der Basis kaum breiter als der Kopf mit den Augen, mit bisweilen undeutlicher basaler Querfurche, seitlich abstehend behaart. Fliigeldecken kurzoval, kaum länger als Kopf und Halsschild zusammen, gemeinsam breiter als die Halsschildbasis, seitlich sehr stark gerundet, lang und schräg nach hinten SCYDMAENIDAE VON SABAH 913 gerichtet behaart, mit tiefer, von einer schrägen Humeralfalte scharf begrenzter Basal- impression. Flügel entwickelt. Beine kurz, Schenkel schwach verdickt. Penis (Abb. 61) wie für die Untergattung Napochus typisch gebaut. Peniskörper von oben betrachtet gerundet-viereckig, doppelt so breit wie die davon scharf abgesetzte Apikalpartie, Basalöffnung dorsobasal gelegen mit schmalem sklerotisierten Rahmen, Parameren am distalen Rand der Basalöffnung angesetzt, schmal und dünnhäutig, distal der Mitte S-förmig gekrümmt, das Penisende erreichend, mit je 2 terminalen Tastborsten. Im Penisinneren liegt ein umfangreicher sklerotisierter Komplex, vor dessen Basis knapp hinter der Basalöffnung eine schmale Querleiste liegt. Von den beiden Enden dieser Querleiste ziehen Muskelstränge zu dem Sklerotinkomplex, von dem beiderseits ein sichelförmig nach hinten gekrümmter Zahn absteht. Hinter den beiden Zähnen steht am apikalen Ende des Sklerotinkomplexes auf beiden Seiten eine schmale Sklerotinschleife ab. Apikal ragt am Ende des Sklerotinkomplexes eine Spitze unter dem zungeförmigen Apex penis nach hinten vor. Euconnus ? (Napochus) sabahinanus nov. spec. MATERIAL: Holotypus ¢ (Penispräparat), Mount Kinabalu Nat. Park, Poring Hot Springs area Eastern Ridge Tr., 1000 m, 28.6.1988 (lg. Smetana, cMG); 1 Paratypus Poring Hot Springs, 500 bis 600 m, 9.5.1987 (Ig. Burckhardt u. Löbl, cMG); ebenda 2 Paratypen, 1030 m, (lg. Smetana, cF); 1 Paratypus, Mount Kinabalu at Livagu, 1803 m, 2.9.1988 (lg. Smetana, cMG); Crocker Range, 9-9- 1988 (lg. Smetana cMG). DIAGNOSE: Sehr klein, schwarz, Extremitäten rotbraun, gedrungen gebaut, Kopf klein, so lang wie breit, Halsschild isodiametrisch, Flügeldecken kurzoval. BESCHREIBUNG: Long. 1,00 bis 1,10 mm, Schwarz, Extremitäten rotbraun. Kopf von oben betrachtet rundlich, mit stark vorstehenden Augen und bärtig behaarten Schläfen. Fühler mit lockerer 4 gliederiger Keule, zurückgelegt die Halsschildbasis erreichend, ihre beiden ersten Glieder leicht gestreckt, 3 bis 7 stark quer, 8 3-mal so breit wie 7,9 und 10 noch etwas breiter, alle 3 viel breiter als lang, das Endglied fast so lang wie die beiden vorletzten zusammen, sein distaler Teil schmäler als der proximale, wie für Napochus kennzeichnend. Auch die vorletzten Glieder wie für Napochus typisch gebaut. Halsschild annähernd konisch, breiter als lang, mit 2 in einer Querfurche stehenden Basalgrübchen, dicht und struppig abstehend behaart. Flügeldecken kurzoval, schon an der Basis zusammen breiter als die Halsschildbasis, mit tiefer, lateral durch eine lange Humeralfalte begrenzter Basalimpression, lang, nach hinten gerichtet behaart. Beine kurz und schlank, Schenkel nur schwach verdickt. Penis (Abb. 62) im einzigen vorliegenden Präparat nur unvollständig erhalten. Die Apikalpartie zweispitzig, die Basalöffnung und eine Paramere verloren. Die erhaltene rechte Paramere ist leicht S-förmig gekrümmt und hat keine Tastborsten. Euconnus (Napochus) borneoi nov. spec. MATERIAL: Holotypus d, (Penispräparat), Sabah, Poring Hot Springs, 500 m, (1g. Burckhardt u. Löbl, cMG); ebenda, 1 Paratypus (Penispräparat) 9.5.1987 (Burckhardt u. Löbl, cF); 1 Paratypus, Mount Kinabalu, 2600 m, 23.5.1987 (lg Burckhardt u.Löbl, cMG); 1 Paratypus, Kinabalu Nat. Park, HQ Liongu River, ca. 1000 m, 5.9.1987 (lg. Smetana, cMG); Paratypus 9 Crocker Range, ca. 1000 m, 5.12.1988 (Ig. Smetana, cMG). DIAGNOSE: Nur im weiteren Sinn in das Subgenus Napochus gehörig. Durch rund- lichen Kopf mit großen vorgewölbten Augen, kurze Fühler mit breiter 4 gliederiger Keule 914 HERBERT FRANZ ABB. 72-77. 72: Euconnus fraudulentus nov. spec., Penis in Dorsolateralansicht. 73: Euconnus kinabalumontanus nov. spec., Penis in Dorsalansicht. 74: Euconnus parakinabalumontanus nov. spec., Penis in Dorsalansicht. 75: Euconnus minutipenis nov. spec. Penis in Dorsolateralansicht. 76: Euconnus glandulipenis nov. spec., Penis in Dorsalansicht. 77: Euconnus robusticeps nov. spec. Penis in Dorsalansicht. SCYDMAENIDAE VON SABAH 915 sowie konischen Halsschild mit 2 Basalgrübchen, jedoch ohne basale Querfurche gekenn- zeichnet. BESCHREIBUNG: Long. 1,15 bis 1,20 mm, lat. 0.,55 mm. Rotbraun, gelblich behaart. Kopf von oben betrachtet isodiametrisch-rundlich mit großen vorgewölbten Augen und dicht behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis erreichend, mit breiter 4 gliederiger Keule, diese reichlich so lang wie die Geißel, das Basalglied gestreckt, 2 bis 7 klein, Glied 8 bis 10 3mal so lang wie 7, breiter als lang, das Endglied sehr kurz eiförmig. Halsschild so lang wie breit, konisch, an der Basis so breit wie der Kopf mit den Augen, mit 2 großem Basalgrübchen und steif abstehend behaarten Seiten. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, etwas länger als Kopf und Halsschild zusammen, mit kleiner, von einer hoch emporgwölbten Humeralfalte begrenzter Basalimpression, fein punktiert und schräg abstehend behaart. Flügel entwickelt. Beine schlank, Schienen in der Längsmitte etwas verdickt. Penis (Abb. 63) von oben betrachtet an einen Hutpilz erinnernd, Peniskörper und Apex nicht scharf getrennt, ihm sitzt basal haubenförmig eine “Schale” auf, die den Peniskörper fast bis zur Längsmitte einhüllt und in der dorsobasal die Basalöffnung liegt. Diese besitzt keinen sklerotisierten Rahmen. Diese “Schale” steht über einem kleinen querovalen Sklerotinkörper, der hinter der Basalöffnung liegt, und über die dünnhäutigen Parameren mit dem Peniskörper in Verbindung. Dieser ist etwa in seiner Längsmitte von beiden Seiten eingeschnürt und erweitert sich dahinter wieder zu einem schwalben- schwanzförmigen Endteil. Im Inneren des Peniskörpers liegt nahe der Basis eine keulenförmige Blase, die sich distal schiffchenfòrmig erweitert. An sie schließt der Ductus ejaculatorius, ein dickes Rohr, das in einem weiten Kanal nicht ganz bis zur Penisspitze reicht. Von oben und hinten betrachtet links befindet sich ein breiter und flacher, schräg nach hinten und zur Mitte gerichteter Zahn. Die Parameren sind breit, erreichen das distale Drittel der Penislänge und liegen dem Peniskörper seitlich eng an. Jede tägt eine terminale Tastborste. BESTIMMUNGSABELLE DER Napochus-ARTEN AUS SABAH. Be Snenimiss-sliedenigerEühlerkeulesr AMENER IR IL TONO 2 A ntenimit4-cliederiperEiuhlerkeuleB® Sass. SR ee 3 2 Etwas größer, das 7. bis 10. Fühlerglied nur leicht gestreckt, Kopf mit den stark vor- gewölbten Augen so breit wie die Halsschildbasis ............... quinquearticulatus n. sp. — Etwas kleiner, das 7. bis 10. Fühlerglied wesentlich länger als breit, Kopf mit den wenig vorgewölbten Augen nicht so breit wie die Halbschildbasis ............. ventriculosus n. sp. SE elanyzsrob (lon el: 60 MM) e I LN set Meee E SONORO RT 4 = Körperlängesl SO, mmiund!daruntere a RUN E I SO CRUE ATEN RAR 5 4 11. Fühlerglied im distalen Teil abrupt verschmälert, Fühler zurückgelegt die Hals- sehildbasissuberragend nr E sear net oe teen eee ee burckhardtianus n. sp. — Distaler Teil des 11. Fühlergliedes nicht eingeschniirt und distal abrupt verschmälert, Fühler zurückgelegt die Halsschildbasis knapp erreichend ............. layangensis n. sp. Sam Korpenläansezsumsl, 30: ME Te Re RR 6 ze Köıperlängen]30immmund.darunterhe es IT IT 7 9837615410, Eühlerslied3;malisorbreitwielang. nenn valdeobscurus n. sp. — 8. bis 10. Fühlerglied höchstens doppelt so breit wie lang .................. funestus n. sp. imubdorpeniance les 0mm PenissAbby Ole: ee fuscus n. sp. = Körperlänge 20/mmrund darunter, Benis’anders gebauten RR RE 8 916 HERBERT FRANZ .8 Halsschild ohne basale Querfurche, nur mit Basalgübchen ................................ 9 —EHalsschild’mit/basaler.@uerfurche..........222.2222202 022 rene ERE eee eee 10 SERRKopBschmälenalssder/Halsschild RTRT kinabalui n. sp. zu Kopkso,breitiwie.der.Halsschild.. en. an. er re borneoi n. sp. lORKorperfarbe\schwarzg&i er me ee I eerie sabahinanus n. Sp. = MIKOrperfarbe rotbraun. nn... hierher 4 Arten die nur im männlichen Geschlecht aufgrund des Baues des männlichen Kopulationsapparates sicher unter- schieden; werden kOnnem 22.42.4204 MARNE RE ER Untergattung Napoconnus Franz Euconnus (Napoconnus) cephalotes nov. spec. MATERIAL: Holotypus d (Penispräparat) Mount Kinabalu Nat. Park, HG at Livagu River, 1600 m, 25.4.1987 (lg. Smetana, cMG); ebenda, 1500 m, 2.9.1988 (lg. Smetana, cMG); 24 Paratypen, Mount Kinabalu, 1450 bis 1550 m, 21. u. 23.5.1987 (lg. Burckhardt u. Löbl, cF); ebenda, 3 Paratypen (lg. Burckhardt u. Löbl 1987, cMG); 1 Paratypus ebenda, 1450 bis 1530 m (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Kopf besonders beim d groß, Schläfen und Hinterkopf außerordentlich dicht, pelzig behaart, Fühler sehr kurz, Körper auffällig gedrungen gebaut. BESCHREIBUNG: Long. 1,60 bis 1,80 mm, lat. 0,60 bis 0,70 mm. Rotbraun, bräunlich behaart. Kopf von oben betrachtet annähernd isodiametrisch-rundlich, sehr flach gewölbt, beim 6 mit den Augen nahezu so breit wie die Halsschildbasis, beim 9 wesentlich schmäler, die Stirn in beiden Geschlechtern spitz nach vorne vorspringend. Fühler zurückgelegt knapp die Halsschildmitte erreichend, zwischen dem 1. und 2. Glied deutlich abknickbar, das Basalglied zweimal, das 2. eineinhalbmal so lang wie breit, 3 bis 8 klein, distalwärts zunehmend quer, 9 und 10 mehr als doppelt so breit wie lang, 10 länger als 9, das gerundet-kegelförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild konisch, so lang wie an der Basis breit, ohne Basalgrübchen, kurz und dicht, abstehend behaart. Flügeldecken schon an der Basis zusammen deutlich breiter als die Halsschildbasis, seitlich stark gerundet erweitert, wenig länger als Kopf und Halsschild zusammen, ohne Basalimpression und ohne Schulterbeule. Flügel voll entwickelt. Beine ziemlich kurz, Schenkel mäßig verdickt, Schienen gerade. Penis (Abb. 65) langgestreckt, von oben betrachtet zum apikalen Ende leicht verbreitert, der Apex tief gespalten, zu beiden Seiten mit je 2 Zähnen endend. Am Grunde des Ausschnittes zwischen diesen tritt ein in 2 Spitzen gespaltener Vorsprung zutage. Unter und zwischen den Zähnen steht eine horizontale, stark sklerotisierte Platte hervor, die mediodistal in einem spitzen Zahn endet. Noch ventraler als diese Platte befinden sich zwei sichelförmig zur Mitte gebogene Zähne. Die Basalöffnung des Penis liegt dorsobasal, sie hat ein kleines Lumen und ist von einem sehr breiten Sklerotinrahmen umfaßt. An diesem entspringen die dünnen geraden Parameren, die das Penisende nicht erreichen. Jede trägt terminal 2 lange Tastborsten. Euconnus (Napoconnus) parallelipenis nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Mount Kinabalu Nat. Park, Poring Hot Springs, 530 m, 30.8.1988 (lg. Smetana, cMG). SCYDMAENIDAE VON SABAH 917 DIAGNOSE: Gekennzeichnet durch das Fehlen von Basalgrübchen am Halsschild, durch glatte, schwach behaarte Flügeldecken mit runder Basalimpression ohne Humeralfalte und stark verdickte Vorderschenkel sowie durch die Penisform. BESCHREIBUNG: Long. 1,40 mm, lat. 0,60 mm. Kastanienbraun, bräunlich behaart. Kopf etwas länger als mit den kleinen Augen breit, flach gewölbt, an den Schläfen und am Hinterkopf schütter behaart. Fühler zurückgelegt die Halsschildmitte knapp erreichend, ihre beiden ersten Glieder doppelt so lang wie breit, 3 bis 8 klein, 9 und 10 dreimal so breit wie 8, das Endglied um die Hälfte länger als breit. Halsschild so lang wie breit, ein wenig breiter als der Kopf mit den Augen, schütter behaart, ohne Basalgrübchen. Flügeldecken sehr fein punktiert, spärlich behaart, mit kleiner, runder Basalimpression. Vorderschenkel stärker verdickt als die der Mittel- und Hinterbeine, Schienen schlank und gerade. Penis (Abb. 65) sehr langgestreckt, parallelseitig, seine Basalöffnung mit sehr breitem, stark sklerotisierten Rahmen, mit sehr dünnen, fast penislangen Parameren mit je 2 terminalen Tastborsten, mit aus stark sklerotisierten, zangenförmigen Teilen bestehenden Apex und mit 2 aus dem terminal gelegenen Ostium herausragenden, basal verbreiterten, distal schmalen und parallelen Stäben. Euconnus (Napoconnus) valdepullus nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Mount Kinabalu, 1550 bis 1650 m 24.4.1987 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus, ebenda (cF). DIAGNOSE: Für die Untergattung ungewöhnlich klein. BESCHREIBUNG: Long 0,90 mm, lat. 0,35 mm. Schwarzbraun, Beine und Palpen bräunlich gelb, hellgrau behaart. Kopf von oben betrachtet länglich-rautenförmig, im Niveau der vor seiner Längsmitte stehenden Augen am breitesten und hier so breit wie der Vorderrand des Halsschildes, nach hinten gerichtet, lang behaart. Fühler zurückgelegt die Halsschildmitte erreichend, ihr Basalglied doppelt, das 2. eineinhalbmal so lang wie breit, 3 bis 8 sehr klein, 9 3 mal, 10 4-mal so breit wie 8, das Endglied gerundet- kegelförmig, so lang wie breit. Halsschild ein wenig länger als an der Basis breit, mit 2 Basalgrübchen, besonders an den Seiten dicht behaart. Flügeldecken stark gewölbt, an der Basis zusammen wenig breiter als die Halsschildbasis, mit runder und tiefer Basalimpression, fein behaart. Flügel voll entwickelt. Beine schlank und kurz. Penis (Abb. 66) leicht dorsalwärts gekrümmt, seine Basis lappenförmig dorsalwärts gebogen, Apex penis spitzwinkelig-dreieckig distalwärts vorspringend, Operculum haken- förmig nach oben gebogen. Vor dem Ostium penis liegt im Penisinneren ein kapuzen- förmiges Sklerotingebilde. Euconnus (Napoconnus) langananensis nov. spec. MATERIAL: Holotypus d (Penispräparat). Poring Hot Springs, 850 m, 14.8.1987 (1g. Burckhardt u. Löbl, CMG); 1 Paratypus d (Penispräparat), ebenda 30.4.1987 (lg. Burckhardt u. Löbl (CMG); 1 Paratypus (dé), Mount Kinabalu, 1500 m, 21.4.1987 (lg. Burckhardt u. Löbl, cF); 1 Paratypus 3 (Penispräparat) Mount Kinabalu, 1500 m, 21.7.1987 (lg. Burckhardt u. Löbl, cF); 1 Paratypus dé (Penispräparat), Crocker Range, 1650 m, 16.5.1987 cMG und 2 Paratypen, ebenda (cF). DIAGNOSE: Gekennzeichnet durch großen Kopf, dieser mit den Augen so breit wie die Halsschildbasis, so lang wie breit. 918 HERBERT FRANZ È BESCHREIBUNG: Long. 1,30 bis 1,40 mm, lat. 0,70 mm. Rotbraun, gelblich behaart. Kopf so lang wie breit, Schläfen und Hinterkopf sehr dicht, nach hinten gerichtet behaart. Fühler zurückgelegt die Halsschildbasis knapp erreichend, ihr Basalglied und das 2. doppelt so lang wie breit, 3 bis 8 sehr klein, 9 3 mal so breit wie 8, 10 noch etwas breiter, beide stark quer, das Endglied so lang wie breit. Halsschild knapp so lang wie an der Basis breit, mit 2 Basalgrübchen, dicht, nach hinten gerichtet behaart. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, feiner und schütterer als der Halsschild behaart, mit wenig hervortretender, lateral von einer kurzen Humeralfalte begrenzter Basalimpression. Flügel entwickelt. Beine kurz, Vorderschenkel stärker verdicht als die der beiden anderen Beinpaare. Penis (Abb. 67) weniger langgesteckt als bei den meisten anderen Arten der Untergattung, nach oben gebogen, der Apex 2-spitzig, Operculum ebenfalls in Form von 2 Doppel- spitzen nach hinten vorspringend. Parameren das Penisende nicht erreichend, mit je einer terminalen Tastborste versehen. Euconnus (Napoconnus) elongatior nov. spec. MATERIAL: Holotypus 4 und Paratypus 4 (Penispräparate), Crocker Range, 1350 m, km 60 Kota Kinabalu, Tambunan, 17.5.1987 (lg. Burckhardt u. Löbl, cMG); 4 Paratypen, Crocker Range, 1550 bis 1650 m, 16.5.1987 Ig. Burckhardt u. Löbl, cF); 5 Paratypen, Crocker Range, 17.5.1987 route Kota Kinabalu, Tambunan (lg. Burckhardt u. Löbl, cMG); 2 Paratypen, ebenda, 1550 bis 1650 m 16. bis 18.5.1987 (CMG); 2 Paratypen, Poring Hot Springs, 7. bis 12.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Dem nachstehend beschriebenen E. proceripenis sehr ähnlich, weicht von diesem vor allem durch schlankere Fühlerkeule, den Besitz von 4 Basalgrübchen des Halsschildes und durch anders geformten Penis ab. BESCHREIBUNG: Long. 1,30 mm, lat. 0,60 mm, dunkel rotbraun, bräunlichgrau behaart. Kopf von oben betrachtet gerundet-rautenförmig, im Niveau der stark vorgewölbten Augen am breitesten und mit den Augen breiter als lang, Hinterkopf und Schläfen abstehend behaart. Fühler zurückgelegt die Halsschildbasis fast erreichend, ihr Endglied spitz-eiförmig, so lang wie die beiden vorletzten zusammen. Halsschild an der Basis so breit wie lang, mit 4 Basalgrübchen. Flügeldecken sehr fein punktiert und behaart, vor der Basis mit kleiner Basalimpression. Flügel voll entwickelt. Beine schlank, Vorderschenkel kaum stärker verdickt als die der beiden anderen Beinpaare. Penis (Abb. 68) sehr langgestreckt, distalwärts verbreitert, seine Basalöffnung im rechten Winkel nach oben gebogen. Parameren dem Peniskörper eng anliegend, das Penisende fast erreichend, mit 2 terminalen Tastborsten. Apex penis zweispitzig. Euconnus (Napoconnus) tambunanus nov. spec. MATERIAL: Holotypus dé (Penispräparat), Crocker Range, 1000 m, (lg. Burckhardt u. Löbl, cMG); 4 Paratypen, Poring Hot Springs (lg. Burckhardt u. Löbl, cF); ebenda 6 Paratypen (CMG); 1 Paratypus, Mount Kinabalu, 1534 m, 29.5.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Fühlerkeule schlanker als bei E. proceripenis, Halsschild mit 2 Basal- grübchen, Flügeldecken abstehend behaart, Vorder- und Mittelschenkel keulenförmig verdickt. SCYDMAENIDAE VON SABAH 919 BESCHREIBUNG: Long. 1,20 bis 1,30 mm, lat. 0,60 mm, Rotbraun, weißlich behaart. Kopf von oben betrachtet so lang wie mit den großen, vorgewölbten Augen breit, lang, nach hinten gerichtet behaart. Fühler zurückgelegt fast die Halsschildbasis erreichend, das 1. Glied fast 3 mal so lang wie breit, das 11. Glied in einer scharfen Spitze endend, so lang wie die beiden vorletzten zusammen. Halsschild leicht gewölbt, seine Basis nur so breit wie der Kopf mit den Augen, vor der Basis mit 2 Grübchen. Flügeldecken zusammen nicht viel breiter als die Halsschildbasis, abstehend behaart, an der Basis mit einer länglichen, wenig tiefen Basalimpression. Penis (Abb. 69) dem des E. langananensis ähnlich, wie dieser in flachem Bogen dorsal- wärts gekrümmt. Basalöffnung dorsobasal gelegen, mit nur mäßig breitem Sklerotin- rahmen umgeben. Parameren dünn, dem Peniskörper eng anliegend, das Penisende nicht ganz erreichend, im Spitzenber ich mit 4 Tastborsten verschiedener Länge. Apex mit zwei stumpfen Spitzen, zwischen diesen mäßig tief ausgeschnitten. Unmittelbar vor dem apikal gelegenen Ostium penis liegt ein wenig differenzierter Sklerotikomplex. Euconnus (Napoconnus) proceripenis nov. spec. MATERIAL: Holotypus 4 (Penispräparat) und Paratypus ®, Crocker Range, 1600 m, 18.5.1987 (lg. Burckhardt u. Löbl, CMG); 1 Paratypus d Penispräparat, Mount Kinabalu, 1500 m, 21.5.1987 (lg. Burckhardt u. Löbl cF); Crocker Range, 1530 bis 1650 m und 1200 m, 63 km route Kota Kinabalu, 16.5.1987 (lg. Burckhardt u. Löbl cMG); 1 Paratypus, Poring Hot Springs, 900 bis 950 m, 12.5.1987 (lg Burckhardt u. Löbl, CMG) und ebenda 1 Paratypus d, 11.5.1987 (cF). DIAGNOSE: Gekennzeichnet durch breite Fühlerkeule, deren letztes Glied breiter als das vorletzte, am distalen Ende abgerundet, Halsschild mit 2 Basalgrübchen. BESCHREIBUNG: Long. 1,30 mm, lat. 0,60 mm. Dunkel rotbraun, bräunlichgrau be- haart. Kopf von oben betrachtet gerundet-rautenförmig, im Niveau der stark vorgewölbten Augen am breitesten, mit den Augen breiter als lang, Hinterkopf und Schläfen abstehend behaart. Fühler zurückgelegt die Halsschildbasis erreichend, ihr Endglied spitz-eiförmig, so lang wie die beiden vorletzten zusammen. Halsschild an der Basis so breit wie lang, mit 4 Basalgrübchen. Flügeldecken sehr fein punktiert und behaart, an der Basis mit kleiner Basalimpression. Flügel voll entwickelt. Beine schlank, Schenkel in der Dicke wenig verschieden. Penis (Abb. 70) sehr langgestreckt distalwärts schwach verbreitert, seine Basalöffnung im rechten Winkel nach oben gebogen, Parameren dem Peniskörper eng anliegend, das Penisende fast erreichend, mit terminaler Tastborste. Apex penis 2-spitzig. BESTIMMUNGSTABELLE DER Napoconnus-ARTEN AUS SABAH ee Schriklem Worperlansei0 90M en valdepullus n. sp. Se Grober Rorpergl ine 20 MUNITE 2 2 Kopf breit, mit den Augen beim d annähernd so breit wie die Halsschildbasis, Fühler zurückgelegt die Halsschildmitte nicht erreichend, Körper sehr gedrungen gebaut — Kopf schmäler, Fühler zurückgelegt die Halsschildmitte überragend .................... 4 3 Schläfen und Scheitel sehr dicht pelzig behaart, Halsschild ohne Basalgrübchen ARA TRITATO RE REN cephalotes n. sp. 920 HERBERT FRANZ -— Schläfen und Hinterkopf dicht, aber nicht pelzig behaart, Halsschild mit 2 Basal- RUM CHEM Maes. EN ee ee ER langananensis n. Sp. 4 Halsschild ohne Basalgrübchen, 8. Fühlerglied breiter als das 7 ............................. LAOS R IS e eroe d Sd. SWElalsschildsmitBasalgrubehen te. MR TTT 5 5 Halsschild mit 2 Basalgrübchen, Vorderschenkel mäßig verdickt ..................... ren LER TR OE RARE tambunanus n. Sp. — Halsschild mit 4 Basalgrübchen, Vorderschenkel stark verdickt. Hierher die 2 nur durch Unterschiede im Penisbau sicher unterscheidbaren Arten ............................ Euconnus-species incertae sedis Euconnus globicollis nov. spec. MATERIAL: Holotypus d (Penispräparat), Sabah, Poring Hot Springs, 6.5.1987 (1g. Burckhardt u. Löbl, CMG); 2 Paratypen ebenda, (CMG), und 1 Paratypus (cF). DIAGNOSE: Gekennzeichnet durch länglich-rautenförmigen Kopf, undeutlich abgesetzte 5-gliederige Fühlerkeule, schlanken konischen Halsschild mit 2 Basalgrübchen und kräftige Beine mit mäßig verdickten Schenkeln und gekrümmten Vorder- und Mittelschienen. BESCHREIBUNG: Long. 1,80 mm, lat. 0,70 bis 0,80 mm. Rotbraun, die Beine heller gefärbt, gelblich behaart. Kopf von oben betrachtet gerundet-rautenförmig und wenig länger als mit den großen, stark gewölbten Augen breit, die Schläfen doppelt so lang wie der Augendurchmesser, lang abstehend behaart. Fühler allmählich zur Spitze verdickt, mit undeutlich abgesetzter 5- gliederiger Keule, zurückgelegt die Halsschildbasis knapp erreichend, ihre beiden ersten Glieder leicht gestreckt, die folgenden isodiametrisch, vom 7. an gegen die Spitze zunehmend verdickt, das spitz-eiförmige Endglied so lang wie die beiden vorhergehenden zusammen. Halsschild so lang wie breit, fast konisch, seitlich abstehend behaart, mit 2 Basalgrübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, stark gewölbt und seitlich stark erweitert, nur mit Andeutung einer Basalimpression, ohne Humeralfalte, lang abstehend behaart. -Beine kräftig, Vorderschenkel stärker verdickt als die der Mittel- und Hinterbeine, Vorder- und Mittelschienen leicht medialwärts gekrümmt. Penis (Abb. 71) sehr eigenartig gebaut, Apex penis stark nach oben gekrümmt, ebenso auch die Parameren, Operculum dagegen in der Ebene der Ventralwand des Penis gelegen, in einer stark verschmälerten am Ende spatelförmigen Spitze endend. Aus dem Ostium penis ragt rechtsseits ein starker Stachel nach oben: Die Parameren erreichen das Penisende nicht, sie tragen je 2 terminale Tastborsten. Euconnus fraudulentus nov. spec. MATERIAL: Nur Holotypus d (Penispräparat), Mount Kinabalu, 1500 m, 25.4.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Dem E. parakinabalumontanus sehr ähnlich, von ihm aber durch nur 3- gliederige Fühlerkeule und ganz andere Penisform verschieden. Gekennzeichnet außerdem durch kleinen, rautenförmigen Kopf, kurze Fühler und kurze Beine mit medialwärts gekrümmten Schienen. SCYDMAENIDAE VON SABAH 921 Vio r ti Ù N 1 I N i ' ' 1 1 u % ' ABB. 78-83. 78: Euconnus paramerorum nov. spec., Penis in Dorsalansicht. 79: Euconnus circumlatus nov. spec., Penis in Dorsalansicht. 80: Euconnus filipenis nov. spec., Penis in Dorsalansicht. 81: Euchonnus stylifer nov. spec., Penis in Dorsalansicht. 82: Euconnus paraglobicollis nov. spec., Penis in Dorsalansicht. 83: Euconnus pondoki nov. spec., Penis in Dorsalansicht. 922 HERBERT FRANZ BESCHREIBUNG: Long. 2,00 mm, lat. 1,00 mm. Schwarz, Beine rotbraun, dicht bräun- lichgrau behaart. Kopf von oben betrachtet rautenförmig mit stark vorgewölbten Augen, so breit wie lang, oberseits nach hinten gerichtet, an den Schläfen seitlich abstehend, dicht behaart. Fühler kurz, zurückgelegt nicht einmal die Halsschildmitte erreichend, alle Glieder mit Ausnahme des ersten und letzten breiter als lang, das 2. breiter als das 7., 3 mal so breit wie lang, halb so breit wie das 9., dieses fast, das 10. exakt doppelt so breit wie lang, das eiförmige Endglied so lang wie die beiden vorletzten zusammen. Halsschild so lang wie breit zum Vorderrand viel stärker als zur Basis verengt, sehr dicht und kurz, aufgerichtet behaart, mit 2 Basalgrübchen. Flügeldecken zusammen schon an der Basis viel breiter als die Halsschildbasis, mit sehr kleiner, lateral von einer wenig vortretenden Humeralfalte begrenzter Basalimpression, dicht punktiert und dicht abstehend behaart. Flügel entwickelt. Beine kurz, Schenkel keulenförmig verdickt, Schienen medialwärts gekrümmt. Penis (Abb. 72) wie bei vielen anderen Euconnus-Arten geformt, mit oberseits flachem, ventral vorgewölbtem Peniskörper und von diesem scharf abgesetztem, dreieckigem Apex. Die Basalöffnung mit stark sklerotisiertem Rahmen und das Penisende nicht ganz erreichenden Parameren. Diese mit je 3 terminalen Tastborsten, leicht gebogen. Seiten des Apex penis schwach ausgeschweift. Aus dem Ostium penis ragen mehrere Sklerotinstäbe heraus, von denen 2 von oben und hinten besehen rechts liegende mit Widerhaken versehen sind. Euconnus kinabalumontanus nov. spec. MATERIAL: Holotypus d (Penispräparat), Crocker Range, 1600 m, 18.8.1987 (lg. Burckhardt u. Löbl, cMG; 1 Paratypus, Mount Kinabalu, 1500 m, 21.5.1987 (lg. Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch gestreckten, gerundet-rautenförmigen Kopf mit kleinen Augen, steif abstehend behaarten, isodiametrischen Halsschild mit 4 Basal- grübchen und stark medialwärts gekrümmte Vorderschienen. BESCHREIBUNG: Long. 1,90 mm, lat. 0,88 bis 1,00 mm, schwarz, die Beine rötlich- braun, braun behaart. Kopf von oben betrachtet lang-rautenförmig, die kleinen Augen an den Kopfseiten weit nach vorne gerücket, lang und dicht abstehend behaart. Fühler mit scharf abgesetzter, 4- gliederiger Keule, zurückgelegt die Halsschildmitte erreichend, ihre beiden ersten Glieder eineinhalbmal so lang wie breit, 3 bis 7 klein, isodiametrisch, 8 3-mal so breit wie 7, 9 und 10 noch etwas breiter, alle 3 stark quer, das Endglied sehr kurz eiförmig, fast so lang wie 9 und 10 zusammen. Halsschild breiter als der Kopf, so lang wie breit, beinahe konisch, mit 4 Basalgrübchen, dicht und abstehend behaart. Flügeldecken zusammen an der Basis so breit wie die Halsschildbasis, mit von einer langen Humeralfalte begrenzter Basalimpression, fein punktiert und abstehend behaart. Beine ziemlich lang und schlank, Schenkel schwach verdickt, Mittelschienen medio- distalwärts gebogen. Penis (Abb. 73) mit einem von oben betrachtet eineinhalbmal so langen wie breiten, parallelseitigen Peniskörper und einem von diesem scharf abgesetzten zungenförmigen Apex. Basalöffnung dorsal gelegen, mit sklerotisiertem Rahmen, Parameren gerade, die Basis des Apex penis etwas überragend, mit je einer terminalen Tastborste. Operculum weniger als halb so lang wie der Apex, sein Hinterrand halbkreisförmig, seine Basis flach bogenförmig begrenzt, stark sklerotisiert. SCYDMAENIDAE VON SABAH 923 Euconnus parakinabalumontanus nov. spec. MATERIAL: Holotypus d (Penispräparat), Mount Kinabalu, 1450 bis 1550 m, 23.5.1987 (Burckhardt u. Löbl, CMG); 1 Paratypus ©, 1450 m, HQ Liwago River, 7.8.1988 (lg. Smetana, cF). DIAGNOSE: Gekennzeichnet durch lange Behaarung der Oberseite, kurze Fühler mit 4-gliederiger Keule, Penis sehr ähnlich dem von E. kinabalumontanus. BESCHREIBUNG: Long. 1,90 bis 2,10 mm, lat. 0,80 mm. Dunkel rotbraun, graubraun behaart. Kopf von oben betrachtet länglich-rund mit kleinen vor seiner Länsmitte stehenden Augen, Hinterkopf und Schläfen sehr lang und dicht behaart. Fühler mit 4 gliederiger Keule, zurückgelegt nur die Halsschildmitte erreichend, ihr Basalglied und das 2. um ein Drittel länger als breit, 3 bis 7 klein, isodiametrisch, 8 mehr als doppelt so breit wie 7, wie auch 9 und 10 mehr als doppelt so breit wie lang, das gerundet-kegelförmige Endglied nicht ganz so lang wie 9 und 10 zusammen. Halsschild isodiametrisch, seitlich mäßig gerundet, zum Vorderrand nur wenig mehr als zur Basis verengt, sehr dicht, und abstehend behaart. Flügel voll entwickelt. Beine kurz, Schenkel mäßig verdickt. Penis (Abb. 74) von oben betrachtet aus einem distal verbreiterten Peniskörper und einem davon abgeschnürten, gerundet rechteckigen Apex bestehend. Parameren nur die Basis des Apex penis erreichend, am distalen Ende mediodistal gekrümmt, mit je 2 terminalen Tastborsten. Apex penis gerundet-viereckig, seine Umrandung stärker sklerotisiert. Im Penisinneren liegen vor der Basis des Apex, spiegelbildlich zur Sagittalebene, 4 Sklero- tinkörper. Euconnus minutipenis nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Kinabalu Nat. Park, Sumit Trail, Pondok Lowii, 2300 bis 2400 m, 28.4.1987 (lg. Smetana, CMG); 2 Paratypen, Crocker Range, 1550 bis 1659 m, 18.5.1987 (Ig. Burckhardt u. Löbl, cMG); 2 Paratypen, Poring Hot Springs, 550 bis 950 m, 9. bis 12.5.1987 (lg. Burckhardt u. Löbl, cF); ebenda, 1 Paratypus, 485 m, 20.8.1987 (lg. Smetana cMG); 2 Paratypen, Mount Kinabalu, 1550 bis 1700 m, 24. bis 27.5.1987 (lg. Burckhardt u. Löbl, cF); 1 Paratypus, Kinabalu Nat. Park, Mempasi Trail, 1600 m, 22.5.1987 (lg. Smetana, cMG); 1 Paratypus, Kinabalu Nat. Park, Sumit Trail, Pondok-Utah, 2050 m, 26.4.1987 (lg. Smetana, cMG); Crocker Range, ca. 1000 m, 5.9.1988 (Ig. Smetana, cMG). Dragnose: Äußerlich dem E. globicollis ähnlich, aber heller rotbraun, schütterer behaart, Vorderschienen kaum merklich medialwärts gebogen, Flügeldecken zusammen schon an der Basis wesentlich breiter als die Halsschildbasis, Fühler mit deutlich abgesetzter 4 gliederiger Keule, Penis ganz anders geformt. BESCHREIBUNG: Long. 1,70 bis 1,80 mm, lat. 0,60 bis 0,70 mm. Rotbraun, Palpen und Beine heller gefärbt, bräunlichgelb behaart. Kopf von oben betrachtet gerundet-rautenförmig, die Augen beim d größer und stärker vorgewölbt als beim 9, Hinterkopf beulenförmig über den Hals vorragend, Schläfen dicht, seitlich abstehend behaart. Fühler dick, beim 9 zurückgelegt die Halsschildbasis etwas, beim d um die beiden letzten Glieder überragend, ihr Basalglied dicker als die folgenden, das 2. beim d reichlich, beim 2 knapp doppelt so lang wie breit, die folgenden Glieder bis zum 7. kugelig, 8 bis 10 isodiametrisch, doppelt so breit wie 7, das kegelförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Halsschild beim d leicht gestreckt, beim @ so breit wie lang, mit 2 großen medialen und 2 kleinen lateralen Basalgrübchen, auf der Scheibe schütter, an den Seiten sehr dicht, abstehend behaart, zum Vorderrand sehr stark, zur Basis fast nicht verengt. 924 HERBERT FRANZ - Flügeldecken zusammen schon an der Basis breiter als die Halsschildbasis, beim d gestreckter als beim 2, mit flacher, aber lateral von einer Humeralfalte scharf begrenzter Basalimpression, fein punktiert und schräg abstehend behaart. Schenkel beim d stärker als beim © keulenförmig verdickt, Schienen mediodistal flach ausgerandet und mit Haarfilz bedeckt, beim d stärker als beim 9 mediodistal gekrümmt, Tarsen sehr zart. Penis (Abb. 75) sehr klein dorsalwärts gekrümmt, der Apex nicht vom Peniskörper abgesetzt, dreieckig, aber vor der Spitze breit abgerundet. Basalöffnung dorsobasal gelegen, mit einem sklerotisierten Rahmen umgeben, Parameren das Penisende nicht ganz erreichend und jede mit 4 terminalen Tastborsten versehen. Das spitzwinkelig-dreieckig ausgeschnittene Operculum erreicht den Hinterrand des Apex, es ist stärker sklerotisiert als der Peniskörper. Euconnus glandulipenis nov. spec. MATERIAL: Nur Holotypus d, Poring Hot Springs, 500 m, 6.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch schokoladebraune Färbung, lange, 4- bis 5-glie- derige Fühlerkeule, gerundet-rautenförmigen Kopf mit stark vorgewölbten Augen und nahezu konischen Halsschild. Im Habitus täuschend an Napoconnus erinnernd, aber durch die lange 4- bis 5-gliederige Fühlerkeule und durch den Penisbau als einer ganz anderen Verwandtschaftsgruppe zugehörig erwiesen. BESCHREIBUNG: Long. 1,50 mm, lat. 0,76 mm. Dunkel schokoladebraun, hellgrau behaart. Kopf von oben betrachtet gerundet-rautenförmig, mit großen, stark vorgewölbten Augen und beulenförmig gegen den Hals vortretendem Hinterkopf, lang grau behaart. Fühler zurückgelegt die Halsschildbasis überragend, ihre 4- bis 5-gliederige Keule länger als die Geißel, das Basalglied sehr kurz, das 2. zweieinhalbmal länger als breit, 3 bis 5 annähernd isodiametrisch, 6 und 7 klein, breiter als lang, 8 bis 10 so lang wie breit, das eiförmige Endglied fast so lang wie 9 und 10 zusammen. Halsschild etwas länger als breit, nahezu konisch, an der Basis nur wenig breiter als der Kopf mit dem Augen, stark gewölbt, mit 2 Basalgrübchen, schütter, anliegend behaart. Flügeldecken sehr kurz oval, hoch gewölbt, schon an der Basis zusammen viel breiter als die Halsschildbasis, mit lateral von einer Humeralfalte begrenzter Basalimpression, mit langer, fast anliegender Behaarung. Beine kurz. Schenkel schwach verdickt. Penis (Abb. 76) sehr gedrungen gebaut, der Peniskörper von oben betrachtet fast kreisrund, der Apex aus 2 spitzwinkelig-dreieckigen Teilen bestehend. Basalöffnung ohne sklerotisierten Rahmen, von den Parameren ist am einzigen Präparat nur eine erhalten, die nur die Basis des Apex penis erreicht. Sie ist an der Spitze verbreitert und mit mehreren Tastborsten besetzt. In der Mitte des Peniskörpers liegt ein rundlicher sklerotisierter Körper und distal davon ein doppelt- halbmondförmiges Sklerotinband. Euconnus robusticeps nov. spec. MATERIAL: Holotypus d (Penispräparat) und Paratypus, Mount Kinabalu Nat. Park, area Langanan Crk., 885 m, 22.8.1988 (lg. Smetana, cMG); 1 Paratypus, Poring Hot Springs, 500 m, 3.5.1987 (Ig. Burckhardt u. Löbl, cMG) und 1 Paratypus ebenda, 13.5.1987 (lg. Burckhardt u. Löbl, cF); 1 Paratypus, Crocker Range, km. 6 NE Kota Kinabalu Tambunan, 19.5.1987 (lg. Burckhardt u. Löbl, cMG). SCYDMAENIDAE VON SABAH 925 DIAGNOSE: Sehr ausgezeichnet durch gedrungenen Körperbau, großen runden Kopf und kugelig gewölbten Halsschild, sowie kurze Fühler und Beine. BESCHREIBUNG: Long. 1,80 bis 2,00 mm, lat. 0,60 mm. Schwarz, Beine rotbraun, Palpen bräunlichgelb, braun behaart. Kopf von oben betrachtet nahezu kreisrund, flach gewölbt, fast so lang und breit wie der Halsschild, an den Schläfen dicht und steif abstehend behaart, Augen klein, weit nach vorne gerückt. Fühler dick, zurückgelegt nur das vorderste Viertel der Halsschildlänge erreichend, die Geißelglieder kurz und gedrängt aneinanderschließend, die 4-gliederige Keule fast so lang wie die Geißel, Glied 8 um die Hälfte breiter als 7, 9 um mehr als die Hälfte breiter als 8, wie auch 10 stark quer. Das gerundet kegelförmige Endglied so lang wie die beiden vorletzten zusammen. Halsschild kugelig, gleichmäßig vom Vorderrand bis zur Basis gerundet, sehr dicht abstehend behaart, mit 2 großen medialen und 2 kleinen lateralen Basalgrübchen. Flügeldecken an der Basis zusammen kaum breiter als die Halsschildbasis, stark gewölbt und seitlich gleichmäßig gerundet, mit ziemlich tiefer Basalimpression, fein punktiert und nach hinten gerichtet behaart. Flügel entwickelt. Beine sehr kurz und dick, alle Schenkel stark verdickt, die Schienen in der Längsmitte breiter als an den beiden Enden, leicht medialwärts gekrümmt. Penis (Abb. 77) von oben betrachtet oval, der Apex zweistufig zur Spitze verschmälert, in einer scharfen Spitze endend. Basalöffnung groß, nur die distale Hälfte ihrer Umrahmung stark sklerotisiert. Parameren die Längsmitte des Apex ereichend, mit je 3 terminalen Tastborsten. Im distalen Viertel der Penislänge befindet sich ein stark sklerotisierter Komplex, der die ganze Penisbreite einnimmt. Euconnus paramerorum nov. spec. MATERIAL: Holotypus d (Penispräparat), Mount Kinabalu, 1750 m, 27.4.1987 (lg. Burckhardt u. Läbl, cMG); ebenda Paratypus d (Penispräparat) (lg. Burckhardt u. Löbl, cF); Paratypus, Poring Hot Springs, 550 bis 600 m, 9.5.1987 (1g. Burckhardt u. Löbl, cMG) und Paratypus above Poring Hot Springs, 570 m, 15, 8, 1988 (Ig. Smetana, cMG). DIAGNOSE: Sehr ausgezeichnet durch annähernd kugelförmigen Kopf, unscharf abgesetzte, 5-gliederige Fühlerkeule, beim d sehr stark verdickte Vorderschenkel und durch den Penisbau. BESCHREIBUNG: Long. 1,60 m, lat. 0,60 mm. Schwarz, Beine rotbraun, Palpen hell gelbbraun. Kopf von oben betrachtet ungefähr kugelig, Scheitel beulenförmig emporgewölbt, Augen groß, aber wenig gewölbt, allseits abstehend behaart. Fühler beim & mit undeutlich abge- grenzter 5-gliederiger, beim 2 mit 4-gliederiger Keule, zurückgelegt die Halsschildbasis knapp erreichend, ihre beiden ersten Glieder leicht gestreckt, 3 bis 6 kugelig, 7 beim d um die Hälfte breiter als 6, beim ? kaum größer als das vorhergehende, 8 bis 11 doppelt so breit wie 6, 8 kaum, 9 und 10 zunehmend breiter als lang, das gerundet-kegelförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild kugelig gewölbt, zum Vorderrand stärker als zur Basis verengt, so lang wie breit, mit 2 Basalgrübchen, abstehend behaart. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, kaum länger als Kopf und Halsschild zusammen; sehr fein punktiert und mäßig dicht, nach hinten gerichtet behaart. Basalimpression seicht, lateral von einer Humeralfalte begrenzt. Beine mittellang, Vorderschenkel etwas stärker verdickt als die der beiden anderen Beinpaare, Schienen schwach gebogen. 926 HERBERT FRANZ Penis (Abb. 78) von oben betrachtet birnförmig, mit großer, von einem skerotisierten Rahmen umgebener Basslöffnung. Apex nicht abgesetzt, Ostium penis terminal gelegen, Parameren sehr lang, distal der Mitte in weitem Bogen zur Sagittalebene gebogen. Euconnus circumlatus nov. spec. MATERIAL: Holotypus d (Penispräparat) Kinabalu Nat. Park, Poring Hot Springs, 520 m, 15.8.1988 (Ig. Smetana, CMG); 1 Paratypus, ebenda 9.5.1987 (lg. Smetana, cMG); 6 Paratypen, ebenda 500 bis 600, 7. bis 13.5.1987 (lg. Burckhardt u. Löbl, cMG); 4 Paratypen, Kinabalu Nat. Park, Sumit Trail Pondok Ubah, 2000 bis 2400 m, 26.4.1987 (lg. Smetana, CMG); HQ Liwagu River, 1500 bis 1550 m, 27.4.1987 (lg. Smetana, CMG); Mount Kinabalu, 1750 m 27.4.1987 (lg. Burckhardt u. Löbl, cMG); 4 Paratypen, ebenda, Mount Kinabalu, 2600 m, 2.5.1987 (1g. Burckhardt, u. Löbl, cF) und 3 Paratypen ebenda (cMG); 4 Paratypen, ebenda, 1550 m, (CMG). DIAGNOSE: Sehr ausgezeichnet durch von oben betrachtet fast kreisrunden Kopf, sehr scharf abgesetzte, 4-gliederige Fühlerkeule und durch den Penisbau. BESCHREIBUNG: Long. 1,50 bis 1,80 mm, lat. 0,60 bis 0,80 mm. Kastanienbraun, die Beine rotbraun, bräunlich behaart. Kopf von oben betrachtet fast kreisrund, mit großen, flach gewölbten Augen und bärtig abstehend behaarten Schläfen, Fühler zurückgelegt die Halsschildmitte wenig überragend, ihre beiden ersten Glieder um ein Viertel länger als breit, breiter als die folgenden, 3 bis 7 annähernd so lang wie breit, 8 3mal so breit wie 7, sehr schwach quer, 9 und 10 noch etwas breiter als das Endglied, dieses gerundet- kegelförmig, wenig länger als breit. Halsschild konisch, nicht breiter als der Kopf, dicht, abstehend behaart, mit 2 weit an die Seiten gerückten Basalgrübchen, Flügeldecken an der Basis zusammen nur wenig breiter als die Halsschildbasis, mit kleiner furchenförmig vertiefter Basalimpression, sehr fein punktiert und abstehend behaart. Flügel entwickelt. Beine lang, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 79) gedrungen gebaut, schwach sklerotisiert, von oben betrachtet gerundet- rechteckig, leicht nach oben gebogen, Umrahmung der Basalöffnung distal stark sklero- tisiert, von ihrem Basalrand ragt ein Sklerotinzapfen nach vorne. Parameren sehr kurz, nicht einmal die Längsmitte des Penis erreichend, mit einer terminalen Tastborste. Apex vom Peniskörper abgesetzt, eine breite und kurze, nach oben gerichtete Platte darstellend. In ihren Hinerecken steht beiderseits eine lange, nach hinten gerichtete Tastborste. Im Penisinneren steht sagittal eine tropfenförmige Apophyse, von der beiderseits ein Sklerotinbogen zur Seite und nach hinten zieht. Zwischen den beiden Bögen befinden sich in unregelmäßiger Anordnung kleine Sklerotinkörper. Im Apikalbereich zieht von oben und hinten betrachtet nach hinten rechts ein langer Sklerotinstab, an dessen linker Seite basal weitere kleine Skerotinkörper angeordnet sind. Euconnus filipenis nov. spec. MATERIAL: Holotypus 3 (Penispräparat) Mount Kinabalu, 1500 m, 25.4.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Gekennzeichnet durch geringe Größe, querovalen Kopf mit stark vorge- wölbten, großen Augen und schlanken Fühlern mit 4-gliederiger Keule, isodiametrischem Halsschild mit 2 durch eine Querfurche verbundenen Basalgrübchen. Penis sehr schlank. BESCHREIBUNG: Long. 1,20 mm, lat. 0,50 mm, hell rotbraun, wenig dicht hellbraun behaart. SCYDMAENIDAE VON SABAH 927 See ABB. 84-88. 84: Euconnus crockeranus nov. spec. Penis in Dorsalansicht. 85: Euconnus valdepilosus nov. spec., Penis in Dorsolateralansicht. 86: Euconnus tortricornis nov. spec., Penis in Dorsalansicht. 87: Euconnus borneoensis nov. spec. Penis in Dorsalansicht. 88: Euconnus crockeri nov. spec., Penis a) in Dorsal-, b) in Lateralansicht. 928 HERBERT FRANZ ‚Kopf mit den großen, vorgewölbten Augen queroval. Die Fühler mit lockerer, 4- gliederiger Keule, zurückgelegt die Halsschildbasis erreichend, ihr Basalglied und das 2. um die Hälfte länger als breit, 3 bis 7 annähernd isodiametrisch, 8 bis 10 mehr als doppelt so breit wie 7, und breiter als lang, das Endglied gerundet-kegelförmig, kürzer als 9 und 10 zusammen. Halsschild so lang wie breit, kaum breiter als der Kopf mit den Augen, seitlich gleich- mäßig gerundet, vor der Basis mit 2 durch eine Querfurche verbundenen Grübchen, seitlich abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit lateral durch eine Humeralfalte begrenzter Basalimpression, schütter behaart. Beine schlank, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 80) sehr schmal und langgestreckt, schwach sklerotisiert, seine Dorsalwand in der Längsmitte im Bogen begrenzt, distal von der Begrenzung ragen aus dem Penisinneren zahlreiche Stachel nach hinten. Das Operculum ist nahezu so lang wie der Peniskörper, sehr spitzwinkelig-dreieckig, seine äußerste Spitze aufgebogen. Euconnus stylifer nov. spec. MATERIAL: Holotypus d (Penispräparat), Poring Hot Springs, 500 m, 8.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda 2 Paratypen (cF); 1 Paratypus 2 Kinabalu Nat. Park, HQ Sillar, 1540 m, 14.8. bis 1.9.1988 (lg. Smetana, cMG). DIAGNOSE: Gekennzeichnet durch kastanienbraune Färbung, unscharf abgesetzte 5- gliederige Fühlerkeule, kugelig gewölbten Halsschild mit sehr dichter, senkrecht ab- stehender Behaarung und lang-strichförmig in einer Eindellung verlaufende Basal- impression. BESCHREIBUNG: Long. 1,50 bis 1,60 mm, lat. 0,70 bis 0,75 mm. Kastanienbraun, braungrau behaart. Kopf von oben betrachtet gerundet-rautenförmig, Schläfen sehr dicht, abstehend behaart, Augen klein, seitlich vorgewölbt. Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit unscharf abgesetzter 5- gliederiger Keule, ihre beiden ersten Glieder gestreckt, 3 bis 6 klein, isodiametrisch, 7 um die Hälfte breiter als 6, 8 bis 10 gleichbreit, doppelt so breit wie 7, alle 3 stark quer, das eiförmige Endglied kürzer als 9 und 10 zusammen. Halsschild fast so lang wie breit, sehr dicht und steif aufgerichtet behaart, mit 4 tiefen Basalgrübchen. Flügeldecken schon an der Basis zusammen viel breiter als die Halsschildbasis, fein punktiert und abstehend behaart, mit tiefer Basalimpression und schräger, hoch empor- gewölbter Humeralfalte. Flügel voll entwickelt. Beine mittellang, Vorderschenkel stärker verdickt als die der beiden anderen Beinpaare, Mittelschienen medialwärts gekrümmt. Flügel voll entwickelt. Penis (Abb. 81) langgestreckt, Peniskörper von oben betrachtet doppelt so lang wie breit, die Basalöffnung von einem breiten sklerotisierten Rahmen umgeben, die Parameren die Mitte des Peniskörpers wenig überragend, zur Spitze verbreitert, ohne Tastborsten. Aus dem Ostium penis ragt ein langer, ventralwärts gebogener Sklerotinstab nach hinten heraus. Er wurzelt hinter der Längsmitte des Peniskörpers und ist an der Basis ver- hältnismäßig breit. Er verschmälert sich distalwärts trichterförmig. Auf seiner von hinten und oben betrachtet rechten Seite liegen hintereinader 2 große ovale Sklerotinkörper und unter diesen ein schmales stark sklerotisiertes Operculum. Die Dorsalwand des Penis ist über das Operculum hinaus distalwärts verlängert und wellenförmig begrenzt, sie ist in SCYDMAENIDAE VON SABAH 929 dem Bereich vor dem Hinterrand mit zahlreichen Poren besetzt. Eine Abgrenzung der apikalen Partie gegen den Peniskörper ist nicht vorhanden. Euconnus paraglobicollis nov. spec. MATERIAL: Holotypus d (Penispräparat), Crocker Range, 1550 bis 1650 m, 16.6.1987 (lg. Burckhardt u. Löbl, cMG); 3 Paratypen, Poring Hot Springs, 500 m, (lg. Burckhardt u. Löbl, cMG); 1 d (Penispräparat) ebenda (cF). Diagnose: Äußerlich dem E. globicollis sehr ähnlich, von ihm durch scharf abge- setzte 4-gliederige Fühlerkeule und abweichenden Penisbau verschieden und mit ihm deshalb nicht nahe verwandt. BESCHREIBUNG: Long. 1,60 bis 1,65 mm, lat. 0,60 bis 0,70 mm. Rotbraun, bräunlich behaart. Kopf von oben betrachtet gerundet-rautenförmig, länger als breit, stark gewölbt, die kleinen Augen seitlich etwas vorragend. Schläfen steif abstehend behaart. Fühler zurückgelegt die Halsschildbasis etwas überragend, die 4-gliederige Keule etwas kürzer als die Geißel, die beiden ersten Glieder etwas länger als breit, 3 bis 7 isodiametrisch, 8 nicht ganz, 9 und 10 reichlich doppelt so breit wie 7, das eiförmige Endglied etwas kürzer als die beiden vorletzten zusammen. Halsschild schwach gerundet, fast konisch, an der Basis kaum merklich breiter als der Kopf mit den Augen, mit kleinen Basalgrübchen, an den Seiten abstehend behaart. Flügel voll entwickelt. Beine kräftig, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 82) von oben betrachtet eiförmig, der Apex schwach abgesetzt. Basalöffnung des Penis im einzigen vorhandenen Präparat bei Präparation zerstört, Parameren abge- brochen. Die von oben und hinten betrachtet linke in der vermutlich richtigen Position in der Abbildung dargestellt, distal leicht verbreitert, an der Spitze mit 2 längeren Tastborsten, davor auf beiden Seiten mit je 2 kleinen Börstchen. Im Penisinneren entspringt der Präputialsack distal aus einem breiten, lateral durch sklerotisierte Leisten begrenzten Raum. Er verengt sich trichterförmig zu einem Rohr, das bis zur Basis des Apex penis reicht und dessen distales Ende düsenförmig verengt ist. In dem breiten basalen Teil befindet sich sagittal ein sklerotisierter dreieckiger Körper, an dessen beiden Seiten 2 kleinere, rundliche angedeutet sind. Der Apex penis ist zweiteilig, die beiden Teile sind durch einen bogenförmigen Ausschitt getrennt. Euconnus pondoki nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Kinabalu Nat. Park, Sumit Trail Pondok Ubah, 2050 m, 26.4.1987 (lg. Smetana, CMG); Paratypus d (Penispräparat), Poring Hot Springs, 350 bis 600 m, 9.5.1987 (lg. Burckhardt u. Löbl, cF). DIAGNOSE: Zu den Arten mit rautenförmigem Kopf und konischem Halsschild gehörig. Unter diesen verhältnismäßig klein. BESCHREIBUNG: Long. 1,40 mm, lat. 0,60 mm. Rotbraun, bräunlich behaart. Kopf von oben betrachtet gerundet-rautenförmig, im Niveau der etwas vor seiner Längs- mitte stehenden Augen am breitesten, leicht gestreckt, die Schläfen und der Hinterkopf abstehend behaart. Fühler mit 4-gliederiger Keule, zurückgelegt die Halsschildbasis knapp erreichend, ihre beiden ersten Glieder leicht gestreckt, 3 bis 7 klein, breiter als lang, 8 doppelt so breit wie 7, isodiametrisch, 9 und 10 breiter als lang, das eiförmige Endglied kürzer als die beiden vorletzten zusammen. 930 HERBERT FRANZ _Halsschild konisch, etwas länger als breit, mit 2 Basalgrübchen, an den Seiten dicht behaart. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, fein punktiert und behaart, mit einer lateral von einer Humeralfalte begrenzten Basal- impression. Flügel voll entwickelt. Beine mäßig lang, Schenkel keulenförmig verdickt, Schienen gerade. Penis (Abb. 83) von oben betrachtet eiförmig, distal verschmälert, der Apex nicht abgesetzt, sein Hinterrand in der Mitte eingekerbt. Basalöffnung ohne sklerotisierte Umrahmung. Parameren das Hinterende des Penis nicht erreichend, vor der Spitze mit je 3 lateralen Tastborsten. Hinter der Basalöffnung befindet sich im Penisinneren ein U- förmiger Sklerotinkörper. Zwischen den basalwärts gerichteten Armen des U befindet sich ein kleiner, tropfenförmiger Sklerotinkörper. Distal schließen an das U 2 parallel zur Sagittalebene liegende Sklerotinstäbe an, die distal kugelig erweitert sind. Die Seiten des Penis sind zwischen der Basalöffnung und der Apikalregion stärker sklerotisiert als der Basal- und Apikalbereich. Vor dem Hinterrand des Penis stehen spiegelbildlich zur Sagittalebene 2 parallele Sklerotinstäbe. Euconnus crockeranus nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Crocker Range, 1200 m km 63 NE Kota Kinabalu Tambunan, 11.5.1987 (lg. Burckhardt u. Löbl, CMG). DIAGNOSE: Gekennzeichnet durch sehr kurze Fühler mit gedrungener 4-gliederiger Keule, schwarze Körperfarbe und rotbraune Extremitäten. BESCHREIBUNG: Long. 1,40 mm, lat. 0,60 mm. Schwarz, die Extremitäten rotbraun, bräunlich behaart. Kopf von oben betrachtet gerundet-rautenförmig, länger als breit, die Schläfen doppelt so lang wie der Augendurchmesser, abstehend behaart. Fühler zurückgelegt nicht einmal die Halsschildmitte erreichend, Glied 1 sehr kurz, 2 dick, isodiametrisch, 3 bis 7 sehr klein, breiter als lang, 8 3-mal so breit wie 7, 9 und 10 noch etwas breiter, alle 3 stark quer, das Endglied knapp so lang wie breit. Halsschild gleichmäßig zum Vorderrand und zur Basis verengt, so lang wie breit, seitlich dicht und struppig behaart, mit 2 Basalgrübchen. Flügeldecken kurzoval, schon an der Basis etwas breiter als die Halsschildbasis, nur mit Andeutung einer Basalimpression, fein punktiert und abstehend behaart. Beine kurz, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 84) von oben betrachtet oval, der Peniskörper in den basalen 2 Dritteln schwach sklerotisiert und auch die Basalöffnung nur am distalen Rand stärker versteift. Parameren schlank, das Penisende nicht erreichend, mit einer terminalen Tastborste. Das distale Drittel des Penis stark sklerotisiert, in der Längsmitte am Hinterrand im Bogen ausgeschnitten. In das distale Ende des Ausschnittes ragen von den Seiten 2 spiegel- bildlich zu einander stehende Sklerotinhaken vor. An sie schließt basal auf beiden Seiten der Rand des Apex penis an, der im Winkel an den Peniskörper grenzt. Hier liegt in der Sagittalebene ein dünnhäutiges Fenster der Peniswand, das laterodistal auf beiden Seiten von einem stark sklerotisierten, annähernd halbmondförmigen Gebilde begrenzt wird. Euconnus valdepilosus nov. spec. MATERIAL: Holotypus d (Penispräparat) Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus ebenda (cF) und 1 Paratypus (CMG). DIAGNOSE: Gekennzeichnet durch robuste Körperform, dichte und steif aufgerichtete Behaarung der ganzen Oberseite und schwach abgesetzte 4-bis 5-gliederige Fühlerkeule. 931 SCYDMAENIDAE VON SABAH 90 Y1Omm ABB. 89-94. 89: Euconnus sabahi nov. spec., Penis in Dorsalansicht. 90: Euconnus protectus nov. spec., Penis in Dorsalansicht. Euconnus crockericola nov. spec., Penis in Dorsalansicht. 92: Euconnus 91: excelsipenis nov. spec., Penis in Dorsalansicht. 93: Euconnus longitubus nov. spec., Penis in Dorsalansicht. 94: Euconnus indecorus nov. spec. Penis in Dorsalansicht. 932 HERBERT FRANZ } BESCHREIBUNG: Long. 2,30 bis 2,50 mm, lat. 0,30 bis 1,00 mm. Braunschwarz. Beine dunkel-rotbraun, Palpen heller, Behaarung schwarzbraun. Kopf von oben betrachtet rund, stark gewölbt, fast kugelig, allseits dicht und steif abstehend behaart, mit kleinen, hinter den Fühlerwurzeln stehenden Augen. Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit scharf abgesetzter, 4-bis 5- gliederiger Keule, diese sehr unscharf abgesetzt, das 2. Glied um ein Drittel länger als breit, 3 bis 7 leicht gestreckt, 8 bis 10 beim d nicht ganz doppelt so breit wie lang, beim 2 etwas schmäler, das eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild stark gewölbt, etwas breiter als lang, kaum breiter und länger als der Kopf, allseits dicht und abstehend behaart, mit von der Behaarung stark verdeckten Basal- grübchen. Flügeldecken stark gewölbt, schon an der Basis zusammen viel breiter als die Hals- schildbasis, mit von einer sehr kurzen Humeralfalte begrenzter Basalimpression, fein punktiert und lang, nach hinten gerichtet behaart. Flügel entwickelt. Beine mit stark verdickten Schenkeln und schwach gebogenen Schienen. Penis (Abb. 86) aus einem kurzovalen Peniskörper, einem spitzwinkelig-dreieckigen Apex und einem diesem an Länge gleichen Operculum bestehend. Das Operculum ist stark sklerotisiert und besitzt die Form eines H, wobei dessen basale Teile der Verankerung im Peniskörper dienen, während die distalen unter dem Apex nach hinten ragen. Basalöffnung des Penis groß, dorsobasal gelegen. Parameren am distalen Rand der Basalöffnung wurzelnd, das Penisende fast erreichend, terminal mit je 2 langen und 2 kürzeren Borsten versehen. Euconnus tortricornis nov. spec. MATERIAL: Holotypus ¢ (Penispräparat), Crocker Range, 1550 bis 1650 m (lg. Burckhardt u. Löbl, cMG); Paratypus 9 Crocker Range, 1270 m, km 60 route Kota Kinabalu-Tambunan, 17.5.1987 (1g Burckhardt u. Löbl, cF); ebenda & Paratypus, ebenda (CMG). DIAGNOSE: 6 gekennzeichnet durch die Fühlerbildung, mittlere Fühlerglieder exzen- trisch. BESCHREIBUNG: Long. 2,00 mm, lat. 0,90 mm. Schwarz, Palpen und Beine rotbraun, braun behaart. Kopf von oben betrachtet gerundet-rautenförmig isodiametrisch, die Augen in der Längsmitte des Kopfes stehend, stark vorgewölbt, die Schläfen eineinhalbmal so lang wie der Augendurchmesser, seitlich abstehend, der Hinterkopf nach hinten gerichtet behaart. Fühler zurückgelegt die Halsschildbasis erreichend, beim d die 3 ersten Glieder normal geformt, 1 und 2 gestreckt, 3 so lang wie breit, 4 dem 3. exzentrisch aufsitzend, ebenso 5 und 6, die Fühler dadurch in ihrem Bereich nach außen gedreht, 7 wieder in die Achse zurückgedreht, 8 bis 11 die normale Keule bildend, 8 bis 10 stark quer, das eiförmige Endglied so lang wie die beiden vorletzten zusammen. Beim ® sind die Fühler normal gebildet. Halsschild so lang wie breit, zum Vorderrand stärker verengt als zur Basis, dicht, an den Seiten struppig abstehend behaart, mit 2 Basalgrübchen. Flügeldecken sehr kurzoval, schon an der Basis zusammen viel breiter als die Halsschildbasis, fein punktiert und lang, abstehend behaart, mit breiter, lateral von einer Humeralfalte begrenzter Basalimpression. Beine kurz, Schenkel mäßig verdickt, Schienen medialwärts gekrümmt. Penis (Abb. 86) aus einem von oben betrachtet ovalen Peniskörper und einem lang- gestreckten schmalen Apex bestehend. Dieser ist dorsalwärts gebogen, die Spitze schmal SCYDMAENIDAE VON SABAH 933 abgestutzt. Parameren das Penisende fast erreichend, mit einer sehr kräftigen terminalen Borste versehen. Der Apex ist durch ein H-förmiges Sklerotingerüst versteift. Dieses ist in der basalen Häfte breiter, distalverschmälert. Zwischen und vor den basalen Ästen des H befindet sich ein großes querovales, dünnhäutiges Fenster. Euconnus borneoensis nov. spec. MATERIAL: Holotypus d (Penispräparat) Mount Kinabalu, 1750 m, 27.4.1987 (lg. Burckhardt u. Löbl, CMG); 1 Paratypus, ebenda (cF); 1 Paratypus, Poring Hot Springs, 550 bis 600 m, 9.5.1987 (lg Burckhardt u. Löbl, cMG); 1 Paratypus, Poring Hot Springs, area Eastern Ridge Trail; 1000 m, 29.9.1988 (Ig. Smetana, CMG). DIAGNOSE: Gekennzeichnet durch großen runden Kopf, kurze Fühler mit 4- gliederiger Keule, konischen Halsschild sowie geringe Größe. BESCHREIBUNG: Long. 1,40 mm, lat. 0,60 mm. Rotbraun, gelblich behaart. Kopf von oben betrachtet kreisrund, flach gewölbt, mit bärtig behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis nicht erreichend, mit scharf abgesetzter, 4-gliederiger Keule, die beiden ersten Glieder gestreckt, 3 bis 7 sehr klein, 8 bis 10 sehr stark quer, mehr als 3-mal so breit wie 7, das Endglied gerundet-kegelförmig, nicht ganz so lang wie breit. Halsschild annähernd konisch, nicht breiter als der Kopf mit den Augen, seitlich struppig behaart, mit 2 großen Basalgrübchen. Flügeldecken kurzoval, schon an der Basis zusammen breiter als die Halsschildbasis, mit lateral von einer kurzen Humeralfalte begrenzter Basalimpression, fein und wenig auf- fällig, jedoch aufgerichtet behaart. Beine kurz und schlank. Penis (Abb. 87) von oben betrachter wenig länger als breit, in den basalen drei Vierteln distalwärts verbreitert, dann stufenförmig verschmälert, der Apex kurz, zum Hinterrand abgestuft. Zu beiden Seiten der am weitesten distal stehenden Stufe steht eine käftige Borste. Basalöffnung ohne sklerotisierten Rahmen, Parameren dünn, nur das distale Drittel der Penislänge erreichend, mit einer terminalen Tastborste. Im Penisinneren befindet sich ein in der Anlage 5-eckiger Sklerotinkörper, der wie ein vekehrtes U distalwärts offen ist. Euconnus crockeri nov. spec. MATERIAL: Holotypus d und Paratypus d (Penispräparate), Crocker Range, 1550 bis 1650 m, 16.5.1987 (Ig. Burckhardt u. Löbl, cMG); Paratypus 1 d (Penispräparat) Crocker Range, 1200 m, km 63 Kota Kinabalu-Tambunan, 19.5.1987, (CMG); ebenda, 1 Paratypus (cF). DIAGNOSE: Gekennzeichnet durch unscharf abgesetzte 5-gliederige Keule der nur 10- gliederigen Fühler, gerundet-rautenförmigen Kopf, konischen Halsschild und bedeutende Größe. BESCHREIBUNG: Long. 2,20 bis 2,40 mm, lat. 0,80 bis 0,90 mm. Dunkel rotbraun, bräunlich behaart. Kopf von oben betrachtet annähernd kreisrund, der Augendurchmesser nur so groß wie der des 1. Fühlergliedes, Schläfen abstehend behaart. Fühler nur 10-gliederig, das Endglied aber distal leicht abgeschnürt, ähnlich wie bei Napochus, zurückgelegt die Halsschildbasis erreichend, mit unscharf abgesetzter 5- gliederiger Keule, das Basalglied etwas dicker als 2, dieses um ein Viertel länger als breit, 3 bis 5 sehr klein, 6 etwas größer, 7 um ein Drittel breiter als 6, 8 bis 10 noch breiter als 7, deutlich breiter als lang, das gerundet-kegelförmige Endglied fast so lang wie 9 und 10 zusammen. 934 HERBERT FRANZ Halsschild so breit wie lang, fast konisch, mit 2 großen Basalgrübchen, seitlich struppig abstehend behaart. Flügeldecken langoval, an der Basis zusammen nur wenig breiter als die Halsschildbasis, mit breiter, lateral von einer schrägen Humeralfalte begrenzter Basalimpression, schräg abstehend behaart. Beine kräftig und ziemlich lang, Vorderschenkel stärker verdickt als die der beiden anderen Beinpaare, Vorder- und Mittelschienen leicht medialwärts gekrümmt. Penis (Abb. 88 a, b) von oben betrachtet eiförmig, mit schwach abgesetztem Apex, bei seitlicher Betrachtung nach oben gekrümmt. Unter dem Apex liegt noch stärker gekrümmt als dieser und in der Ruhelage mit der Spitze diesem genähert, von oben und hinten betrachtet links der schmale Ductus ejaculatorius, der nur in Lateralsicht als stark sklerotisiertes Rohr erkennbar ist. Die Basalöffnung des Penis besitzt nur distal einen stark sklerotisierten Rahmen, an dem die S-förmig gekrümmten Parameren inserieren. Sie sind mit je 3 terminalen Tastborsten bewehrt. Euconnus sabahi nov. spec. MATERIAL! Holotypus 3 (Penispräparat), Mount Kinabalu, 1580 m 27.4.1987 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus 9, Poring Hot Springs, 6.5.1987 (lg. Bruckhardt u. Löbl, cF); ebenda 1 Paratypus, 500 m, 11.5.1987 (Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch gerundet-rautenförmigen Kopf, scharf abgesetzte 4- gliederige Fühlerkeule, konischen Halsschild mit 2 Basalgrübchen und durch die Penisform. BESCHREIBUNG: Long. 1,30 mm, lat. 0,65 mm, rotbraun, lang gelblich behaart. Kopf von oben betrachtet gerundet-rautenförmig, mit den Augen nicht ganz so breit wie lang, die Schläfen eineinhalbmal so lang wie der Augendurchmesser, dicht abstehend behaart. Fühler zurückgelegt die Halsschildbasis knapp erreichend, ihre beiden ersten Glieder gestreckt, 3 bis 7 klein, isodiametrisch, 8 bis 10 3-mal so breit wie 7, viel breiter als lang, das Endglied gerundet-kegelförmig, so lang wie breit. Halsschild länger als breit, fast konisch, mit 2 Basalgrübchen, nach hinten gerichtet behaart. Flügeldecken zusammen schon an der Basis breiter als der Halsschild, mit kurzer, lateral von einer kurzen Humeralfalten begrenzter Basalimpression, lang, nach hinten gerichtet behaart. Flügel voll entwickelt. Beine schlank, Vorderschienen schwach medialwärts gekrümmt. Penis (Abb. 89) sehr gedrungen gebaut, der Apex gerundet-dreieckig, vom Peniskörper nicht scharf gesondert. Operculum ein wenig länger als der Apex, aus 2 schmalen Sklerotinkörpern bestehend, nach vorne lateralwärts gebogen. Basalöffnung des Penis unmittelbar hinter dem Basalrand des Penis gelegen, nur am distalen Rand stark sklerotisiert, Parameren hier lateral wurzelnd, mit einer terminalen Tastborste versehen. Im Penisinneren liegt distal der Mitte ein großer sklerotisierter Trichter, dessen Mündung in den Spalt zwischen den beiden Teilen des Operculums führt. Von dem von oben und hinten betrachtet rechten Rand des Trichters entspringt ein langer distalwärts verbreiterter und nach rechts ausbiegender Sklerotinstab, der über den Rand des Operculums hinausragt. Euconnus protectus nov. spec. MATERIAL: Holotypus d Kinabalu-Nat. Park, HQ. Silau-Silau Trail, 1540 m, 14.8. bis 1.9.1988 (lg. Smetana, CMG); 2 Paratypen, 2 2 Mount Kinabalu, 1450 his 1550 m, 25.4. und 23.5.1987 (Ig. Burckhardt u. Löbl. cMG); ebenda, 1 Ex. (cF); Poring Hot Springs, 500 m, 7.4.1987, 12.5.1987; (Burckhardt u. Löbl, cF). SCYDMAENIDAE VON SABAH 935 DIAGNOSE: Gekennzeichnet durch verhältnismäßig schlanke Fühler mit lockerer 4- gliederiger Keule und rotbraune Färbung. BESCHREIBUNG: Long. 1,40 mm, lat. 0,60 mm. Rotbraun, gelblich behaart. Kopf gerundet-rautenförmig, die Schläfen doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis erreichend, mit lockerer 4-gliederiger Keule, das 7. Glied intermediär zwischen Geißel und Keule, Glied 2 gestreckt, 3 bis 6 klein, isodiametrisch, 8 bis 10 breiter als lang, das Endglied spitz eiförmig, länger als breit. Halsschild konisch, länger als breit, an der Basis etwas breiter als der Kopf mit den Augen, an den Seiten abstehend behaart, vor der Basis mit 2 Grübchen. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, ohne deutliche Basalimpression, schräg nach hinten abstehend behaart. Flügel entwickelt. Beine ohne besondere Merkmale. Penis (Abb. 90) von oben betrachtet aus einem gerundet- viereckigen Peniskörper und einem davon abgesetzten Apex bestehend. Dieser ist distalwärts verschmälert, zum Ende aber wieder etwas verbreitert. Vor ihm liegt zum Teil im Inneren des Peniskörpers ein länglicher Sklerotinkörper, der zur Basis verbreitert ist und distal im Bereich des Apex penis einen über die ganze Breite reichenden Bogenausschitt besitzt. Die Parameren erreichen das Penisende fast und tragen im Spitzenbereich lateral je 3 Tastborsten. Euconnus crockericola nov. spec. MATERIAL: Holotypus d (Penispräparat) und 4 Paratypen, Crocker Range, 1200 m, 63 km Kota Kinabalu-Tambunan, 19.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 2 Paratypen (cF). DIAGNOSE: Gekennzeichnet durch relativ bedeutende Größe, gerundet-rautenförmigen Kopf, kräftige Fühler mit unscharf abgesetzter, 4-bis 5-gliederiger Keule, kugeligen, relativ kleinen Halsschild, und sehr stark bauchig erweiterte Flügeldecken. BESCHREIBUNG: Long. 2,20 bis 2,40 mm, lat. 0,90 bis 1,00 mm. Dunkel rotbraun, dicht und aufgerichtet bräunlichgelb behaart. Kopf von oben betrachtet gerundet-rautenförmig, stark, der Scheitel und Hinterkopf beulenförmig gewölbt. Länge der Schläfen gleich dem Zweieinhalbfachen des Augen- durchmessers, allenthalben an den Schläfen besonders grob und steif behaart. Fühler zurückgelegt die Halsschildbasis erreichend, das 2. Glied um die Hälfte länger als breit, 3 leicht gestreckt, 4 bis 7 kugelig, 8 und die folgenden doppelt so breit wie 6, alle 3 schwach quer, das eiförmige Endglied fast so lang wie die beiden vorletzten zusammen. Halsschild kugelig, kaum breiter als der Kopf, so breit oder etwas breiter als lang, dicht und abstehend behaart, mit 4 Basalgrübchen. Flügeldecken schon an der Basis zusammen viel breiter als die Halsschildbasis, mit tiefer, lateral von einer langen Humeralfalte begrenzter Basalimpression, sehr lang und dicht, abstehend behaart. Beine sehr kräftig, Schenkel keulenförmig verdickt, Schienen medialwärts gekrümmt. Penis (Abb. 91) in dem einzigen vorliegenden Präparat sehr stark geschrumpft, aus einem ziemlich kurzovalen Peniskörper und einem langen spitzwinkelig-dreieckigen, leicht nach oben gebogenen Apex bestehend. Unter diesem liegt ein schmal-zungenförmiges Operculum, das stärker sklerotisiert ist als der Apex. Von den beiden Parameren ist im Präparat nur eine erhalten. Sie ist in der basalen zwei Dritteln sehr breit, zur Spitze stark verschmälert und an dieser mit einer sehr starken und steif medialwärts gesichtenen und 2 weiteren terminalen Borsten versehen. 936 HERBERT FRANZ Euconnus excelsipenis nov. spec. MATERIAL: Holotypus d (Penispräparat) Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG); 1 Paratypus vom selben Fundort (cF). DIAGNOSE: Gekennzeichnet durch scharf abgesetzte 4-gliedrige Fühlerkeule, gerundet-rautenförmigen Kopf, namentlich aber durch den außerordentlich komplizierten Bau des männlichen Kopulationsapparates. BESCHREIBUNG: Long. 1,70 mm, lat. 0,60 mm. Schwarz, die Extremitäten rotbraun, gelblichbraun behaart. Kopf von oben betrachtet gerundet-rautenförmig, der Hinterkopf beulenförmig empor- gewölbt. Fühler mit scharf abgesetzter 4-gliederiger Keule, zurückgelegt die Hals- schildbasis erreichend, ihr Basalglied kurz, das 2. leicht gestreckt, 3 bis 7 klein, breiter als lang, 8 doppelt so breit wie 7, wie auch 9 und 10 breiter als lang, das Endglied gerundet- kegelförmig, etwas länger als breit. Halsschild rund, etwas breiter als lang, zum Vorderrand stärker als zur Basis verengt, viel breiter als der Kopf, dicht und abstehend behaart, mit 4 Basalgrübchen. Flügeldecken schon an der Basis zusammen etwas breiter als die Halsschildbasis, sehr fein punktiert, stark glänzend, schwach behaart. Flügel voll entwickelt. Beine kurz. Schenkel keulenförmig verdickt. Penis (Abb. 92) reich differenziert und sehr eigenartig gebaut, von oben betrachtet kurzoval, seine Basis mit 2 Höckern versehen, zwischen diesen mit einem spitzbogen- förmigen Ausschnitt. Apex ebenfalls mit 2 dicken, an der Spitze distal ausgerandeten Sklerotinstäben. Deren Basis ist medial rechtwinklig erweitert, aber nicht verbunden, sondern durch einen tiefen sagittalen Spalt getrennt. Am basalen Ende des Spaltes mündet ein kurzes, dickes Rohr, das in einem weiten Trichter in der Penismitte entspringt. In diesen Trichter ragt von vorne ein großer tropfenförmiger Sklerotinkörper, zu dessen beiden Seiten sich basal zwei kleine weitere Sklerotinkörper befinden. Unmittelbar vor ihnen liegt die Basalöffnung des Penis, die nur am Hinterrand einen schwach sklerotisierten Rahmen besitzt. An diesem entspringen die beiden Parameren, die S-förmig gekrümmt bis an das apikale Ende des Penis reichen. Vor ihrer Spitze sind sie gegabelt. Sie entsenden lateral einen sehr kurzen Ast, der gerade abgeschnitten ist und dort 3 Tastborsten trägt. Der Hauptast ist ab der Stelle der Gabelung verschmälert und trägt eine terminale Tastborste. Im Penisinneren befindet sich beiderseits des beschriebenen Trichters eine flügelförmige, an ihrem lateralen Rand schmal sklerotisierte Fläche. Euconnus longitubus nov. spec. MATERIAL: Holotypus 4 (Penispräparat), Mount Kinabalu, 1750 m, 27.4.1953 (lg. Burckhardt u. Löbl, cMG); ebenda 3 Paratypen, 1550 bis 1600 m, 16. u. 18.4.1987 (CMG); 3 Paratypen (1 Penispräparat) Crocker Range, 63 km Kinabalu-Tambunan, 17.-18.5.1987 (lg. Burckhardt u. Löbl, cF); ebenda, 14 Paratypus, ca. 1000 m, 5.9.1988 (lg. Smetana, cMG); 19 Paratypus, Mount Kinabalu, 2600 m, 1.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch verhältnismäßig bedeutende Körpergröße, kleinen konischen Halsschild, und breite, stark gewölbte Flügeldecken. BESCHREIBUNG: Long. 1,60 mm, lat. 0,70 mm. Dunkel rotbraun, bräunlich behaart. Kopf beim & von oben betrachtet gerundet- rautenförmig, die Schläfen nicht ganz doppelt so lang wie der Augendurchmesser, steif abstehend behaart, beim 2 größer, von oben betrachtet rund, Fühler zurückgelegt die Halsschildbasis mit dem Endglied überragend, ihr 3. bis 7. Glied isodiametrisch, ebenso 8 bis 10, diese aber doppelt so breit wie 7, das Englied eiförmig, fast so lang wie die beiden vorletzten zusammen. SCYDMAENIDAE VON SABAH 937 ! : \ \ \ 1 \ \ \ \ \ A N Br nu 5 ABB. 95-101. 95: Euconnus pilosiceps nov. spec., Penis in Dorsolateralansicht. 96: Euconnus smetanai nov. spec., Penis in Dorsalansicht. 97: Euconnus silauensis nov. spec., Penis in Lateralansicht. 98: Euconnus acuticornis nov. spec., Penis in Dorsolateralansicht. 99: Euconnus livagoensis nov. spec. Apikalpartie des Penis in Lateralansicht. 100: Euconnus poringensis nov. spec., Penis a) in Dorsal- b) in Lateralansicht. 101: Euconnus longeacuminatus nov. spec., Penis in Dorsalansicht. 938 HERBERT FRANZ Halsschild klein, so lang wie breit, konisch, an der Basis etwas breiter als der Kopf, mit 4 Basalgrübchen, seitlich abstehend behaart. Flügeldecken zusammen schon an der Basis wesentlich breiter als die Halsschildbasis, mit breiter, lateral von einer Humeralfalte begrenzter Basalimpression, schräg abstehend behaart. Beine kräftig, Schenkel mäßig verdickt, Schienen mäßig medialwärts gekrümmt. Penis (Abb. 99) im Bauplan an E. excelsipenis erinnerend, der Ductus ejaculatorius über den Apex weit hinausragend, dessen Hinterrand in der Mitte nicht tief eingeschitten sondern nur im flachen Bogen breit ausgerandet. Basalöffnung des Penis groß, mit breitem Sklerotinrahmen. Parameren das Penisende so weit überragend wie der Ductus ejacula- torius, vor der Spitze lateral breit ausgerandet, lateral und medial beiderseits mit je 3 Tastborsten. Der Ductus ejaculatorius ist basal trichterförmig erweitert, in dem Trichter ruht eine lang-herzförmige Blase neben der von oben und hinten betrachtet links am Trichterrand ein runder Sklerotinkörper steht. Euconnus indecorus nov. spec. MATERIAL: Holotypus d (Penispräparat), Poring Hot Springs, Langanan River, 850 m, 14.5.1987 (Ig. Burckhardt u. Löbl, cMG); 3 Paratypen, Mount Kinabalu, 1430 bis 1520 m, 25.4.1987 (lg. Burckhardt u. Löbl, cMG); ebenda 2 Paratypen (cF). DIAGNOSE: Gekennzeichnet durch geringe Größe, braunchwarze Färbung, kompakte, 4-gliederige Fühlerkeule und gedrungene Gestalt. BESCHREIBUNG: Long. 1,15 bis 1,25 mm, lat. 0,45 bis 0,50 mm. Braunschwarz, braungrau behaart. Kopf von oben betrachtet rund, flach gewölbt, die Schläfen abstehend behaart, Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit scharf abgesetzter, 4-gliederiger Keule, ihre beiden ersten Glieder leicht gestreckt, die folgenden klein, nur ein Drittel so breit wie 8 und die folgenden. Halsschild konisch, so lang wie breit, an der Basis etwas breiter als der Kopf mit den Augen, mit 2 Basalgrübchen, seitlich abstehend behaart. Flügeldecken kurzoval, schon an der Basis zusammen breiter als die Halsschildbasis, mit breiter, lateral von einer langen Humeralfalte begrenzter Basalimpression, nach hinten gerichtet behaart. Beine kurz, Schenkel schwach verdickt, Schienen gerade. Penis (Abb 94) sehr klein, sein Apex 2-spitzig, dünnhäutig, im Inneren des Peniskörpers jedoch mit stark sklerotisierten Differenzierungen. Im basalen Penisteil steht ein lang- tropfenförmiges Gebilde, hinter dem ein langgestreckter Sklerotinkörper folgt, der sich distal verjüngt und an beiden Seiten von einer sklerotisierten Schale umfaßt ist, die distal im rechen Winkel umbiegt und im Bereich des Apex penis seitlich mit einer scharfen Spitze ins Freie tritt. Sie tritt dort mit dem distalen Ende der Parameren in Verbindung, deren Basis ebenso wie die Basalöffnung des Penis im Präparat nicht erhalten ist. Der Apex endet in 2 abgerundeten Spitzen, zwischen denen der Hinterrand im Bogen ausgeschnitten ist. Euconnus pilosiceps nov. spec. MATERIAL: Holotypus & (Penispräparat), Mount Kinabalu, 1450 bis 1550 m, 23.5.1987 (lg. Burckhardt u. Löbl. CMG); 1% wahrscheinlich dieser Art unter Layang Layang, 2595 m, 2.5.1987 (lg. Smetana, cF). SCYDMAENIDAE VON SABAH 939 DIAGNOSE: Sehr ausgezeichnet durch sehr dicht aufgerichtet behaarten, von oben betrachtet runden Kopf und kurze Fühler mit 4-gliederiger Keule. BESCHREIBUNG: Long. 1,90 mm, lat. 0,80 mm. Braunschwarz, die Beine rotbraun, braunschwarz behaart. Kopf von oben betrachtet rund, der Hinterkopf schwach beulenförmig vorgewölbt, sehr dicht aufgerichtet behaart, mit kleinen, an den Kopfseiten stehenden Augen. Fühler zurückgelegt die Halschildbasis nicht erreichend, mit scharf abgesetzter, 4-gliederiger Keule, die beiden ersten Glieder gestreckt, 3 bis 7 sehr klein, breiter als lang, 8 bis 10 stark quer, 3-mal so breit wie 7, das Endglied gerundet-kegelförmig, so lang wie 9 und 10 zusammen. Bei dem vermutlich zugehörigen 2 die Fühler länger mit lockerer 4- gliederiger Keule, sonst aber der Kopf genau so geformt und behaart. Halsschild so lang wie breit, seitlich schwach gerundet, verhältnismäßig klein, allseits sehr dicht behaart, unter der Behaarung glänzend, ohne Basalgrübchen. Flügeldecken kurzoval, stark gewölbt, schon an der Basis zusammen viel breiter als die Halsschildbasis, mit breiter, von einer langen Humeralfalte begrenzter Basalimpression, dicht und aufgerichtet behaart, fein punktiert. Flügel voll entwickelt. Beine kurz, Schienen leicht medialwärts gekrümmt, beim vermutlich zugehörigen 9 die Mittelschienen mediodistal mit einem feinen Dorn. Penis(Abb. 95) mehr als doppelt so lang wie breit, schwach sklerotisiert, mit unscharf abgesetzter Apikalpartie, diese zweilappig, die Lappen unvollständig getrennt. Basal- öffnung groß, mit sehr schmalem, sklerotisierten Rahmen, Paramere dünn, nur die Basis des Apex penis erreichend, mit je einer terminalen Tastborste. Im Peniskörper ist vor dessen distalem Ende ein unregelmäßig geformter Sklerotinkörper vorhanden. Anmerkung: In Papua habe ich 2 Euconnus-? gefunden, das eine bei der Karawari-Lodge, das andere in der Bismarck-Range, die dem E. pilosiceps äußerlich außerordentlich ähnlich sind. Der Kopf ist bei ihnen allerdings deutlich länger als breit. Euconnus smetanai nov. spec. MATERIAL: Kinabalu Nat. Park, Poring Hot Springs, area Eastern Ridge Tr. 790 m, 17.8.1988 (lg. Smetana, cMG); 1 Paratypus ebenda, 510 m, 30.8.1988 (lg. Smetana, CMG); ebenda 3 Paratypen, 1450 in 1550 m, 29.4. bis 22.5.1987 (1g. Burckhardt u. Löbl, cF). DIAGNOSE: Gekennzeichnet durch nur 3-gliederige Fühlerkeule, gerundet-rauten- förmigen, länglichen Kopf und konischen Halsschild. BESCHREIBUNG: Long. 1,20 mm, lat. 0,50 mm. Dunkel rotbraun, braun behaart. Kopf von oben betrachtet gerundet-rautenförmig, die Schläfen doppelt so lang wie der Augendurchmesser, abstehend braun behaart. Fühler mit 3-gliederiger Keule, sehr breit, fast so lang wie die Geißel, ihre beiden ersten Glieder gestreckt, 3 bis 8 sehr klein, 9 dreimal, 10 und 11 4-mal so breit wie 8, 9 und 10 sehr stark quer, das Endglied kegel- förmig, so lang wie breit. Halsschild konisch, so lang wie breit, an der Basis breiter als der Kopf mit den Augen, seitlich abstehend behaart. Flügeldecken zusammen schon an der Basis breiter als die Halsschildbasis, mit kleiner Basalimpression und langer, abstehender Behaarung. Beine kurz. Penis (Abb. 96) oval, sein Apex vom Peniskörper nicht deutlich abgesetzt, Parameren das Penisende fast erreichend, mit einer terminalen Tastborste. Im Penisinneren befindet sich hinter der Mitte des Peniskörpers ein sklerotisierter Komplex, dessen vorderer Teil nach hinten gerichtet spitzwinkelig-dreieckig ist und abgerundete Ecken besitzt, während der 940 HERBERT FRANZ distale Teil zungenförmig ist und am Hinterrand in der Mitte leicht eingekerbt ist. Vor der Penisspitze befinden sich 2 zwei zueinander und nach hinten konvergierende Stäbe. Euconnus silauensis nov. spec. MATERIAL: Holotypus 4 (Penispräparat) Mount Kinabalu Nat. Park, HQ Silau-Silau. Tr. 1550 m, 14.8.1988 (Ig. Smetana, cMG); ebenda 22 (? diese Art) Mount Kinabalu, 1500 m, 29.4.1987 (1g. Burckhardt CMG); ebenda 29 (? diese Art), 21. bis 25.4.1987 (lg. Burckhardt u. Löbl, cF); Mount Kinabalu, 1500 bis 1550 m, 23-28.4.1987 (lg. Burckhardt u. Löbl, cMG, ? diese Art). DIAGNOSE: Kleine Art mit kurzen Fühlern und mit gedrungen gebauter 4-gliederiger Keule. BESCHREIBUNG: Long. 1,20 bis 1,30 mm, lat. 0,50 bis 0,55 mm. Dunkel rotbraun, bräuniglich behaart. Kopf, von oben betrachtet gerundet-rautenförmig, mit großen Augen und bärtig behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit kompakter 4- gliederiger Keule und dünner Geißel, ihre beiden ersten Glieder kaum gestreckt, die folgenden klein, breiter als lang, 8 bis 10 3-mal so breit wie 7 das Endglied kurz, schmäler als die vorhergehenden. Halsschild konisch, so lang wie breit, an der Basis etwas breiter als der Kopf mit den Augen, seitlich struppig behaart, mit 2 Basalgrübchen. Flügeldecken kurzoval, schon an der Basis zusammen breiter als die Halsschildbasis, fein punktiert und ziemlich lang behaart, mit unscheinbarer Basalimpression. Flügel voll entwickelt. Beine kurz, ohne besondere Merkmale. Penis (Abb. 97) aus einem annähernd isodiametrischen Peniskörper und einer wenig kürzeren zungenförmigen Apikalpartie bestehend. Parameren nur von halber Penislänge. Vor der Basis des Apex penis liegen im Penisinneren Sklerotindifferenzierungen. Euconnus acuticornis spec. nov. MATERIAL: Holotypus d (Penispräparat) Crocker Range 1200 m, km 63 NE Kota Kinabalu- Tambunan, 19.7.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch querovalen Kopf, dieser in seiner Länge fast derjenigen der 3 vorletzten Fühlerglieder gleich. Das Endglied der Fühler sehr spitz und lang, Halsschild klein, mit 2 sehr großen Basalgrübchen. BESCHREIBUNG: Long. 1,40 mm, lat. 0,70 mm. Rotbraun, gelblich behaart. Kopf von oben betrachtet queroval, Augen stark vorgewölbt, Schläfen doppelt so lang wie der Augendurchmesser, ohne steif abstehende Behaarung, Fühler zurückgelegt die Hals- schildbasis etwas überragend, ihre beiden ersten Glieder doppelt so lang wie breit, 3 sehr klein, 4 bis 6 zunehmend größer, 7 kugelig um die Häfte breiter als 6, 8 nicht ganz doppelt so breit wie 7, 9 und 10 noch etwas breiter als 8, zunehmend breiter als lang, das in einer scharfen Spitze endende Englied fast so lang wie 8 bis 10 zusammen. Halsschild isodiametrisch, nicht breiter als der Kopf mit den Augen, mit 2 großen Basalgrübchen, an den Seiten steif abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit von einer Humeralfalte scharf begrenzter Basalimpression, fein punktiert und lang, abstehend behaart. Flügel entwickelt. Beine mittellang, Vorderschenkel etwas stärker verdickt als die der beiden anderen Beinpaare. SCYDMAENIDAE VON SABAH 941 Penis (Abb. 98) aus einem von oben betrachtet gerundet- länglich rechteckigen Penis- körper und einer nur wenig kürzeren Apikalpartie bestehend. Basalöffnung des Penis von einem breiten, stark sklerotisierten Rahmen umgeben. Parameren die Basis des Apex penis etwas überragend mit je 2 terminalen Tastborsten. Apex langgestreckt im basalen Viertel verengt, das 2. und 3. Viertel parallelseitig, das letzte dreieckig verschmälert. Operculum zungenförmig, kürzer und schmäler als der Apex, aber stärker sklerotisiert. Im Inneren des Peniskörpers befinden sich parallele sklerotisierte Falten der Peniswand, unter denen ein von oben und hinten betrachtet links stehender winkelig gebogener Stachel auffällt. Vor ihm steht ein spitzwinkelig dreieckiger nach links außen gerichteter Zahn. Euconnus livagoensis nov. spec. MATERIAL: Holotypus d (Penispräparat), Kinabalu Nat. Park, HQ at Livago, 1505 m 2.9.1988 (1g. Smetana, cMG); ebenda 1 Paratypus (1g. Smetana, cMG); 1 Paratypus, Mount Kinabalu (cF). DIAGNOSE: Eine gedrungen gebaute Art mit gerundet-rautenformigem Kopf mit abstehend behaarten Schläfen, konischem Halsschild und kurzovalen Flügeldecken. BESCHREIBUNG: Long. 1,10 mm, lat. 0,50 mm. Rotbraun, die Extremitäten hell gelb- braun, braun behaart. Kopf von oben betrachtet gerundet-rautenförmig, die dicht und steif abstehend behaarten Schläfen eineinhalbmal so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis erreichend, mit breiter, 4-gliederiger Keule, ihre beiden ersten Glieder dicker als die folgenden, 2 leicht gestreckt, 3 bis 7 klein, 8 mehr als 3-mal so breit wie 7, 9 und 10 noch etwas breiter, alle stark quer, das Endglied halbkugelig, ebenfalls breiter als lang. Halsschild konisch, so lang wie breit, an den Seiten struppig behaart, mit 2 Basalgrübchen. Flügeldecken zusammen wesentlich breiter als die Halsschildbasis, nur so lang wie Kopf und Halsschild zusammen, mit nach hinten verflachter Basalimpression, fein punktiert und nach hinten gerichtet behaart. Beine kurz und schlank, Schenkel nur schwach verdickt. Penis (Abb. 99) ganz immatur, so daß nur einige morphologische Details des apikalen Teiles desselben erkennbar sind. Der Apex ist zungenförmig, zu seinen beiden Seiten sind die Ende den Parameren erkennbar. Sie tragen eine Mehrzahl langer, wellig gebogener Tasthaare. Ein sehr langes biegt zunächst nach hinten und dann hinter der Spitze des Apex penis von oben und hinten besehen nach rechts. Es endet in einer sehr feinen Spitze in der Nähe der Borsten der rechten Paramere. Die angeführten Merkmale werden ausreichen, um die Art im männlichen Geschlecht wiedererkennen zu können, da sie bei keiner anderen Euconnus-Art aus Sabah bisher in einer annähernd ähnlichen Weise bekannt sind. Euconnus poringensis nov. spec. MATERIAL: Holotypus & (Penispräparat) und 1 Paratypus, Poring Hot Springs 8.5.1987 (lg. Burckhardt u. Löbl, CMG); ebenda 1 4 Paratypus (Penispräparat) (cF). DIAGNOSE: Kopf relativ klein, gerundet-rautenförmig mit dicht abstehend behaarten Schläfen. Fühler zurückgelegt die Halsschildbasis knapp erreichend, mit scharf abgesetzter 4-gliederiger Keule, ihre beiden ersten Glieder gestreckt, 3 bis 7 kleinkugelig, 8 eineinhalbmal, 9 und 10 doppelt so breit wie 7, isodiametrisch, das Endglied spitz eiförmig, ein wenig kürzer als 9 und 10 zusammen. Halsschild konisch, kaum merklich länger als breit, an der Basis nur wenig breiter als der Kopf mit den Augen, seitlich struppig, abstehend behaart, mit 2 großen Basalgrübchen. 942 HERBERT FRANZ Flügeldecken zusammen schon an der Basis wesentlich breiter als die Halsschildbasis, dicht punktiert, lang und wenig dicht behaart. Beine ziemlich kurz und schlank, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 100 a, b) von oben betrachtet aus einem ovalen Peniskörper und einem davon abgesetzten, zungenförmigen Apex bestehend. Basalöffnung groß, Parameren an dem einzigen vorhandenen Penispräparat nicht erkennbar. Hinter der Basalöffnung befindet sich im Penisinneren eine gerundet-langrechteckige Platte, die bis in den Apikalbereich zurückreicht und an der Basis schwach, am Hinterrand aber tief im Bogen ausgeschnitten ist. Hinter ihn befindet sich bis weit in die apikale Region hineinreichend eine zweite, etwas längere Platte, die an der Basis klammerartig ausgeschnitten und am distalen Ende stumpfwinkelig-dreieckig begrenzt ist. Daran schliesst sich unter dem Apex penis ein aus 2 parallelen Stäben bestehendes Operculum an, das nicht ganz bis zum Penisende reicht. In dem Raum zwischen den beiden Platten sicht man bei Betrachtung von oben einen kleinen annähernd querovalen Sklerotinkörper. Dieser erweist sich bei lateraler Betrachtung als die Spitze eines großen Zahnes, der von der Ventralwand des Penis nach oben ragt. Die distale Platte biegt bei seitlicher Betrachtung ventralwärts zur Ventralwand des Penis und findet in den beiden Stäben des Operculums ihre Forsetzung. Euconnus longeacuminatus nov. spec. MATERIAL: Holotypus d (Penispräparat) Mount Kinabalu, 1500 m, 25.4.1987 (lg. Burckhardt u. Löbl, cMG); Paratypus d (Penispräparat) Crocker Range 1200 m, km 63 von Kota Kinabalu nach Tambunan, 19.5.1987 (lg. Burckhardt u. Löbl, cF); ebenda, 6 Parytypus, Penispräparat (CMG). Dragnose: Äußerlich dem E. kinabalui ähnlich, aber wesentlich weniger robust, im Bau des männlichen Kopulationsapparates sehr von ihm verschieden. BESCHREIBUNG: Long. 1,90 mm, lat. 0,80 mm. Schwarzbraun, Beine und Palpen rotbraun, braun behaart. Kopf von oben betrachtet rautenförmig, mit stark vorgewölbten Augen, die Schläfen 3-mal so lang wie der Augendurchmesser, dicht und steif, auch die Kopfoberseite lang und steif nach hinten gerichtet behaart. Fühler zurückgelegt die Halsschildbasis erreichend, mit unscharf abgesetzter, 5-gliederiger Keule, ihr 2. Glied um ein Viertel länger als breit, 3 und 4 leicht gestreckt, 5 und 6 kugelig, 7 ebenso aber etwas größer, 8 größer als 7, 9 und 10 schwach quer, das spitz-eiförmige Endglied etwas kürzer als die beiden vorletzten zusammen. Halsschild gestreckt etwas breiter als der Kopf mit den Augen, zum Vorderrand stärker als zur Basis verengt, dicht und abstehend behaart, mit 4 Basalgrübchen. Flügeldecken an der Basis zusammen etwas breiter als die Halsschildbasis, mit breiter, lateral von einer Humeralfalte scharf begrenzter Basalimpression, sehr fein rugos skulptiert (80-fache Vergrößerung), lang, nach hinten gerichtet behaart. Flügel voll entwickelt. Beine ziemlich schlank, Schenkel mäßig verdickt, Mittelschienen des d medioterminal mit einem feinen Dorn. Penis (Abb. 101) sehr langgestreckt, mit einem langen, in einer scharfen Spitze endenden Apex. Operculum breit und viel kürzer als der Apex. Basalöffnung des Penis mit einem breiten sklerotisierten Rahmen. Parameren breit, nur die Basis des Apex penis erreichend, ohne Tastborsten. Apex penis basal mit einem langen sagittalen Dorn, zu beiden Seiten desselben stark sklerotisiert. Davor stehen im Penisinneren mehrere unregelmäßig geformte Sklerotinkörper. SCYDMAENIDAE VON SABAH 943 Euconnus cuneipenis nov. spec. MATERIAL: Holotypus d (Penispräparat), Crocker Range, 1550 bis 1650 m, 16.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch länglichen, gerundet-rautenförmigen Kopf, kleinen, konischen Halsschild mit 4 Basalgrübchen und sehr kurzovale Flügeldecken. BESCHREIBUNG: Long. 1,80 mm, lat. 0,90 mm. Rotbraun, bräunlichgelb behaart. Kopf von oben betrachtet länglich gerundet-rautenförmig, mit langen, steif abstehend behaarten Schläfen und kleinen, vorgewölbten Augen. Fühler zurückgelegt die Halsschildbasis erreichend, mit unscharf abgesetzter 4-gliederiger Keule, ihr Basalglied dicker als die folgenden, das 2. leicht gestreckt, 3 bis 7 kugelig, 7 schon etwas größer als 6, 8 um die Hälfte größer als 7, 9 und 10 mehr als doppelt so breit wie 8, stark quer, das kegelförmige Endglied nicht ganz so lang wie die beiden vorletzten zusammen. Die Fühler erweisen sich bei genauer Betrachtung als schwach asymmetrisch. Die Glieder vom 7. bis 9. sind medial länger als lateral. Die Glieder 8, 9 und 10 sind überdies seitlich erweitert und auf der Erweiterung mit einem Büschel dickerer Borsten versehen. Medial spingt der distale Rand des 8. und namentlich des 9. und 10. Gliedes mit einer scharfen Spitze über die Basis des nächstfolgenden Gliedes vor. Halsschild fast konisch, an seiner Basis nur wenig breiter als der Kopf mit den Augen, stark gewölbt, seitlich abstehend behaart, mit 6 Basalgrübchen. Flügeldecken sehr kurzoval, schon an der Basis zusammen viel breiter als die Halsschildbasis, mit tiefer, lateral von einer Humeralfalte begrenzter Basalimpression. An der Naht hinter dem Schildchen über beide Flügeldecken mit einer flachen, länglichen Eintiefung, sehr deutlich punktiert, stark glänzend, ziemlich schütter behaart. Beine kräftig, Vorderschenkel etwas stärker verdickt als die der beiden anderen Beinpaare, Schienen leicht medialwärts gekrümmt. Penis (Abb. 102) von oben betrachtet keilförmig, mit kleiner, runder Basalöffnung ohne stärker sklerotisierten Rahmen und fast die Penisspitze erreichenden Parameren mit je 2 terminalen Tastborsten. Ductus ejaculatorius gerade, sagittal verlaufend. Er entspringt in einer kleinen Kugel, die seine Basis trichterförmig umfaßt. Seine ganze Basalregion ist von einer Membran kugelförmig umfaßt. Diese Membran ist basal von einem geraden Rohr durchbrochen, das basal in die Kugel mündet und hinter der Basalöffnung in einem fast die ganze Penisbreite einnehmenden Sklerotinbogen entspringt. Die Spitze des Ductus ejaculatorius überragt in der Ruhelage die Spitze des Apex ein wenig. Euconnus ubahanus nov. spec. MATERIAL: Holotypus d (Penispräparat), Kinabalu Nat. Park, Sumit Trail Pondok, Ubah, 2050 m, 26.4.1989 (lg. Smetana, cMG). DIAGNOSE: Eine der vielen Arten des Gebietes, die durch einen rautenförmigen Kopf, durch einen kleinen annähernd konischen Halsschild und kurzovale Flügeldecken ausgezeichnet sind. Sehr ausgezeichnet durch den Bau des männlichen Kopulations- apparates, der von oben betrachtet fast genau gerundet-quadratisch ist und dem ein Apex und ein Operculum fehlen. BESCHREIBUNG: Long. 1,40 mm, lat. 0,70 mm. Dunkel rotbraun, braun behaart. Kopf gerundet-rautenförmig, so lang wie mit den Augen breit, die Schläfen 3-mal so lang wie der Augendurchmesser, steif abstehend behaart. Fühler mit scharf abgesetzter 4- 944 HERBERT FRANZ 108 ABB. 102-108. 102: Euconnus cuneipenis nov. spec., Penis in Dorsalansicht. 103: Euconnus ubahanus nov. spec., Penis in Dorsalansicht. 104: Euconnus masculinus nov. spec., Penis in Dorsalansicht. 105: Euconnus ovulipenis nov. spec., Penis in Dorsalansicht. 106: Euconnus bicornipenis nov. spec., Penis in Dorsalansicht. 107: Euconnus fontium nov. spec., Penis in Dorsolateralansicht. 108: Protoscydmus sabahensis nov. spec., Penis in Dorsalansicht. gliederiger Keule, zurückgelegt die Halsschildbasis nicht ganz erreichend, ihr Basalglied dicker als die folgenden, doppelt so lang wie breit, 2 leicht gestreckt, 3 bis 7 iso- diametrisch, klein, 8 3-mal so breit wie 7, 9 und 10 noch etwas breiter, alle 3 sehr stark quer, das Endglied gerundet-kegelförmig, breiter als lang. Halsschild annähernd konisch, an der Basis etwas breiter als der Kopf mit den Augen, seitlich dicht abstehend behaart, vor der Basis mit 2 nahe an den Seitenrand gerückten Grübchen. Flügeldecken kurzoval, schon an der Basis zusammen viel breiter als der Halsschild, mit tiefer, grübchenförmiger Basalimpression, fein punktiert und lang abstehend behaart. Flügel verkümmert. Beine ziemlich kurz, Schenkel schwach verdickt. SCYDMAENIDAE VON SABAH 945 Penis (Abb. 103) von oben betrachtet gerundet-viereckig, wenig länger als breit, ohne Apex und Operculum, mit kleiner Basalöffnung mit nur am Hinterrand sklerotisiertem Rahmen. Parameren dünnhäutig, nur halb so lang wie der Penis, mit je einer terminalen Tastborste. Hinter der Basalöffnung befindet sich im Penisinneren ein lang tropfen- förmiger Sklerotinkörper, der auf beiden Seiten von einem Sklerotinbogen umfaßt ist. Distal von dem tropfenförmigen Körper befindet sich ein viel kleinerer querovaler und dahinter ein sagittal verlaufendes Rohr, das am Hinterrand des Penis endet, ohne daß dort ein geräumiger Ostium penis vorhanden wäre. Am Hinterrand des Penis befindet sich auf jeder Seite eine Borste. Die Borsten sind einander zugewendet. Euconnus masculinus nov. spec. Material: Nur Holotypus 4 (Penispräparat), Kinabalu Nat. Park, HQ 1500 m, 25. bis 30.4.1987 (1g. Smetana, cMG). DIAGNOSE: Kopf rautenförmig, Halsschild konisch, Fühlerkeule locker 4-gliederig, Fühler zurückgelegt die Halsschildbasis erreichend. BESCHREIBUNG: Long. 1,00 mm, lat. 0,50 mm. Rotbraun, dicht grau behaart. Kopf von oben betrachtet gerundet-rautenförmig, mit den großen, vorgewölbten Augen etwas breiter als lang, sehr flach gewölbt. Fühler mit scharf abgesetzter, lockerer, 4- gliederiger Keule, ihre beiden ersten Glieder eineinhalbmal so lang wie breit, 3 bis 7 etwas breiter als lang, 8 bis 10 sehr schwach quer, das eiförmige Endglied fast so lang wie die beiden vorhergehenden zusammen. Halsschild konisch, etwas länger als breit, an der Basis breiter als der Kopf mit den Augen, ohne Basalgrübchen, an den Seiten dichter behaart als auf der Scheibe. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, wenig länger als zusammen breit, mit kleiner, von einer kurzen Humeralfalte begrenzter Basal- impression, fein punktiert und anliegend behaart. Beine kurz, Schenkel schwach verdickt, Schienen gerade. Penis (Abb. 104) aussergewöhnlich gebaut, sodaß daran zu denken ist, die Art einem eigenen Subgenus zuzuordnen. Peniskörper von oben betrachtet sehr kurzoval, am Hinterrand in seiner ganzen Breite abgestutzt, an ihn schließt der gerundet stumpf- winkelig-dreieckige Apex an, der nicht ganz so breit ist wie der Peniskörper. Aus diesem tritt an der von oben und hinten besehen rechten Seite der Ductus ejaculatorius nach hinten aus. Er entspringt im Inneren des Penis schmal trichterförmig, zieht außerhalb des Apexrandes nach hinten um den Hinterrand des Apex herum bis zu dessen linkem Rand, wobei er sich allmählich verschmäkert. Bei seinem Austritt aus dem Peniskörper steht von ihm ein großer sichelförmiger Zahn ab, der zur Mitte des Penis gerichtet ist. Die Basalöffnung des Penis liegt dorsobasal und besitzt keinen stärker sklerotisierten Rahmen. Von ihren Seiten entspringen die ebenfalls nur schwach sklerotisierten Parameren, die das Penisende erreichen. Sie tragen vor ihrer Spitze lateral eine lange Borste und terminal 2 klöppelförmige Forsätze. Die Basalöffnung des Penis befindet sich in einer Art “Haube”, die den Peniskörper von 3 Seiten umhüllt. Im Penisinneren befindet sich vor dem Ursprung des Ductus ejaculatorius ein sklerotisierter Komplex, der links aus dem Ostium penis einen breiten Lappen nach hinten entsendet. Euconnus ovulipenis nov. spec. MATERIAL: Nur Holotypus d (Penispräparat) Crocker Range, 1200 m, 63 km Straße Kota Kinabalu-Tambunan, 19.5.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Sehr ausgezeichnet durch die sehr lange Fühlerkeule, annähernd konischen Halsschild, sowie durch die Penisform. 946 HERBERT FRANZ BESCHREIBUNG: Long. 1,20 mm, lat. 0,70 mm. Dunkel rotbraun, gelblich behaart. Kopf von oben betrachtet rundlich, flach gewölbt, mit den flach gewölbten Augen nur wenig breiter als lang, die gerundet zur Basis konvergierenden Schläfen knapp doppelt so lang wie der Augendurchmesser, wenig auffallend abstehend behaart, die Behaarung der Kopfoberseite lang, nach hinten gerichtet, wenig dicht. Fühler zurückgelegt die Halsschildbasis knapp überragend, ihr Basalglied und das 2. etwas gestreckt, 3 bis 7 sehr stark quer, 8 doppelt so breit wie 7, auch 9 und 10 leicht gestreckt, 10 breiter als 9, das eiförmige Endglied um die Hälfte länger als 10, die Keule mehr als doppelt so lang wie die Geißel. Halsschild nahezu konisch, zur Basis aber schwach verengt, diese mit 2 durch eine Querfurche verbundenen Grübchen, obserseits schütter, seitlich struppig abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis mit breiter, lateral von einer langen Humeralfalte begrenzter Basalimpression, fein punktiert und schräg abstehend behaart. Flügel voll entwickelt. Beine kurz, ohne besondere Merkmale. Penis (Abb. 105) von oben betrachtet kurz eiförmig, der Apex vom Peniskörper nicht abgesetzt. Die Basalöffnung mit breitem, stark sklerotisiertem Rahmen, Parameren breit, die Penisspitze nicht ganz erreichend. Operculum spitzwinkelig-dreieckig, das Penisende nicht erreichend. Im Penisinneren liegen distal der Basalöffnung 2 an der Basis mit- einander verbundene sagittal parallel zueinander apikalwärts verlaufende Sklerotinzapfen, die im distalen Viertel der Penislänge enden. Euconnus bicornipenis nov. spec. MATERIAL: Nur Holotypus d (Penispräparat) Sabah, Mount Kinabalu, 1750 m, 27.4.1987 (lg. Burckhardt u. Löbl, cMG). DIAGNOSE: Gekennzeichnet durch zurückgelegt die Halsschildbasis erreichende Fühler mit 4-gliederiger Keule, sehr stark querem 8. bis 10. Fühlerglied und sehr großem Endglied, dessen distaler Teil schwach vom basalen abgesetzt und verschmälert ist. Das einzige vorliegende Exemplar ist stark beschädigt, es fehlen ihm u. a. beide Flügeldecken. Penis sehr eigenartig gebaut, sodaß die Erkennung der Art durch die Penismerkmale allein schon ausreichend gesichert ist. BESCHREIBUNG: Long. ca. 1,20 mm, lat. ca. 0,70 mm. Dunkel rotbraun. Kopf von oben betrachtet gerundet-rautenförmig, die Schläfen doppelt so lang wie der Augendurchmesser. Fühler zurückgelegt die Halsschildbasis überragend, ihre beiden efsten Glieder leicht gestreckt, 3 bis 7 isodiametrisch bis schwach quer, 7 schon etwas breiter als 6, 8 mehr als doppelt so breit wie 7, stark, 9 und 10 noch stärker quer, das Endglied so breit wie 10, nur sehr wenig kürzer als 9 und 10 zusammen, sein distaler Teil etwas schmäler als der basale, seine Spitze breit abgerundet. Halsschild konisch, so breit wie lang, an der Basis breiter als der Kopf mit den Augen, so weit erkennbar nur mit 2 Basalgrübchen. Flügeldecken verloren gegangen, Flügel voll entwickelt. Beine ziemlich kurz, Schienen schwach medialwärts gekrümmt. Penis (Abb. 106) von oben betrachtet annähernd doppelt so lang wie breit, distalwärts etwas verschmälert, der Apex nicht scharf abgesetzt. Die Basalöffnung nur an ihrem distalen Rand mit sklerotisierter Umrahmung, Parameren am einzigen Präparat nicht vorhanden. Ostium penis dorsoapikal gelegen, aus ihm ragt distalwärts eine stark skleroti- sierte, horizontale Platte heraus. Diese ist vor ihrem apikalen Ende spatelförmig verbreitert SCYDMAENIDAE VON SABAH 947 und dahinter wieder eckig verschmälert. Der Hinterrand ist 2-spitzig. Die Platte ist auf ihren beiden Seiten im Penisinneren mit einem langen Sklerotinstab verankert, die beiden Stäbe sehen von oben betrachtet wie 2 Stierhörner aus. Die Penisseiten reichen seitlich der horizontalen Platte stachelförmig verschmälert bis zur Penisspitze. Euconnus fontium nov. spec. MATERIAL: Holotypus & (Penispräparat), Sabah, Poring Hot Springs, 500-600 m, 9.5.1987 (lg. Burckhardt u. Löbl, cMG); ebenda, 2 Paratypen, 500 bis 620 m, 16.5.1987 (cF); ebenda 3 Paratypen (cMG); 1 Paratypus, Crocker Range, 1000 m, 5.9.1988 (Ig. Smetana, cMG). DIAGNOSE: Sehr klein, sehr gedrungen gebaut, Fühler sehr kurz. BESCHREIBUNG: Long. 0,90 mm, lat. 0,60 mm. Schwarz, Beine größtenteils rotbraun, dunkel behaart. Kopf von oben betrachtet nahezu kreisrund, dicht und abstehend behaart. Fühler zurückgelegt die Halsschildbasis nicht erreichend, ihre Keule sehr scharf abgesetzt, die Geißel dünn, ohne das Basalglied so lang wie die Keule, das 2. Glied um die Hälfte länger als breit, wie auch das erste viel dicker als die folgenden, Glied 8 doppelt, 9 und 10 3-mal so breit wie lang. Halsschild konisch, an der Basis so breit wie der Kopf, auch an den Seiten nur schütter behaart, stark glänzend, vor der Basis mit 2 großen Grübchen. Flügeldecken sehr kurz oval, nur so lang wie Kopf und Halsschild zusammen, schon an der Basis zusammen viel breiter als die Halsschildbasis, ohne Basalimpression und ohne Humeralfalte, schütter, abstehend behaart. Flügel verkümmert. Beine sehr kurz. Penis (Abb. 107) dünnhäutig, nur die Umrahmung der Basalöffnung, der Apex und 2 nahe beieinander liegende Schleifen im Penisinneren stärker sklerotisiert. Basalöffnung gerundet-viereckig, Apex dreieckig, Ostium terminal gelegen. BESTIMMUNGSTABELLE DER Euconnus-ARTEN INCERTAE SEDIS VON-SABAH simigasymmetnischenrkühlergliedern et". ee 2 = © WMC NO ENT RUNS 11e eee ne le us Pareles. + 2 Mittlere Fühlerglieder exzentrisch, Körpergröße 2,0 mm, schwarzbraun, dicht und abstehendibehaantarr = ne eure ha een tortricornis n. Sp. — Glieder der Fühlerkeule beim d schwach asymmetrisch .................................. 3 3. 8., 9. und 10. Fühlerglied des d medial erweitert und mit einem Borstenbiindel ver- SCREME Sp One SEA A cairn en AT Ne Ne PRES er Eühlerkeulerdrsliedenier Re creas celta aya siete see ten hessen cuneipenis n. sp. — Fühlerkeule 3-gliederig, ihre Glieder medial beim d erweitert ........... smetanai n. sp. 4 Fühler scheinbar 10-gliederig, Glied 3 sehr klein, schwer sichtbar, Keule 5-gliederig DISTORTO EEE Mina lola crockeri n. Sp. = Eühlexdeutlichy leghedener RM een nes eee sect te ee 5 5 Die ganze Körperoberseite lang abstehend dunkel behaart, Körperlänge 1,70 mm und darüber a rec hehe re ee ER Le i Lera 6 — Körper meist ohne lang abstehende dunkle Behaarung, Größe unter 1,60 mm ......... 9 6 Fühler kurz, zurückgelegt die Halsschildbasis nicht erreichend ........................... U —e Eühler lang zurückgeles@die Halsschildbasisüberagend nennen: 9 7 Kopf, Halsschild und Flügeldecken annähernd gleich breit, 8. und 11. Fühlerglied schmälewals/das Sundial Oren Arne Ne INCA een rotundiceps n. Sp. 948 HERBERT FRANZ — Kopf und Halsschild deutlich schmäler als die Flügeldecken ............................. 8 8 Körper schütter behaart, der Untergrund unter der Behaarung allenthalben sichtbar SIGN VARIANTI CNS lee en NEE circumlatus n. Sp. — Körper sehr dicht, emporgerichtet behaart, Untergrund unter der Behaarung nur an beszenzten'Stellen sichtbar! "ira... RI STI pilosiceps n. sp. 9m Körperlänge.über 1, 00:mm 1... ent ER OE 10 = Korperlängerunter 160mm}... MEN sans nee IRR ee Cee Eee ER REE Ree 14 I0BKorpenläaneeubenl: 901mm Erna nen TO crockericola n. sp. ie Konpenlänse unter (SOM init 11 MÉHaAISSChId'KONISCh 007.010 URLS en 2 a Ne PT 12 — Halsschild seitlich gerundet, Fühler zurückgelegt die Halsschildbasis erreichend IR RS IT O nd rei SEE i RURAL OR ac u co couacooccc 13 12 Fühler mit scharf abgesetzter 4-gliederiger Keule ........................ minutipenis n. sp. “gunlermits-pliederiserKenle sc e REI globicollis n. sp. 13 8. Fühlerglied deutlich kleiner und schmäler als die folgenden, Kopf stärker gewölbt undıdiehtenbehaatte rn. een errechnet fraudulentus n. sp. — 8. Fühlerglied kaum kleiner als die folgenden, Kopf flacher und weniger dicht behaart Sa BE De CARRE A A LIRE AES I parakinabalui montanus n. Sp. 14% Körperlänge 1,40 bis/1,60:mmi....... re an ee Mu nade aetna EGE ERP eee 22 WE K6rperlänge unter KAO!MM vi. Ri E E 15 IS@RilemerAntenivomi0:90bistl:00mm'Korperange nn. ....1.0 Peer 16 — Etwas größere Arten von 1,20 bis 1,40 mm Körperlänge ...............................… 17 16 Fühler zurückgelegt die Halsschildbasis erreichend, die 4-gliederige Keule locker, die Glieder 8 bis 10 nur schwach quer, Körperfarbe rotbraun ................ masculinus n. sp. — Fühler zurückgelegt die Halsschildbasis nicht erreichend, die 4-gliederige Keule sehr kompakt, die Glieder 9 und 10 mehr als 3-mal so breit wie lang, Körperfarbe schwarz OPA E cae nn a tee A Sonate P ahs ERE SO I CES CR fontium n. sp. Körperlänge um 1,20 mm... ee Rene RE 18 = Körperlänge 1,30.bis 1,40'mm ..... 2... une see. acces STORTO 19 18 Körper schlank, hellbraun gefärbt, Fühler schlank ..........................- filipenis n. sp. u KOrpenrobuster schwarzbraun, Eühlerrdieker =... cn. livagoensis n. Sp. 19 Fühlerkeule fast doppelt so lang wie die GeiBel, die Keulenglieder alle gestreckt LED ce ME BER RARE SA NO RASE EN OS ER RER RG LL ovulipenis n. sp. — Fihlerkeule nicht oder nur wenig länge als die Keule, ihre Glieder nicht stark gestreckt CROATO NG SR TR cd 0000 0 0 oo 20 20 Fühlerkeule 5-gliederige, die Fühler zurückgelegt die Halsschildbasis ereichend OO SSR INNE TA IR RE I Nea protectus n. Sp. — Eühlerkeule 4 eltederig, Fühlermeist kirzert e 21 200 und OP Eühlerslied3-mal’soi/breit wie langer nenne indecorus n. Sp. Mundi ONFihlergledh6chstens schwach quer "CREME I 22) SR PME CORRE SEEN RES I SORA ROUES RES uo silauensis n. Sp. 22 Kopf stark quer, letztes Fühlerglied sehr spitz eiförmig länger als die beiden vorletzten ZUSAMMEN: cm re ee scuticornis n. SP. andere Merkimalskombinatronem TO MN RE RER RE Hierher alle anderen Arten, die nur aufgrund der Genitalmerkmale sicher bestimmt werden kônnen. Gattung Protoscydmus nov. gen. Die Gattung gehört zu den Scydmaeniden mit dem primitivsten Bau des männlichen Kopulationsapparates. Der Penis besitzt keine scharf begrenzte Basalöffnung. Die Para- SCYDMAENIDAE VON SABAH 949 meren haften am Peniskörper ohne Verbindung mit sklerotisierten Ansatzstellen, im Penisinneren fehlen irgendwelche sklerotisierte Organe, der Apex penis ist nicht vom Peniskörper abgesetzt, ein Operculum fehlt ebenso wie ein scharf umgrenztes Ostium. Das neue Genus gehört in die Verwandtschaft der ebenfalls primitiven Genera Alloraphes, Stenichnaphes und Parastenichnaphes, der Genitalapparat ist aber noch weniger entwickelt. Die Gattung ist auf die einzige bisher bekannte Art Protoscydmus sabahensis nov. spec. begründet. Als Merkmale von generischem Rang sind zu bewerten: Die sehr geringe Größe und die angegebenen Merkmale des männlichen Genitalapparates. Protoscydmus sabahensis nov. spec. MATERIAL: Holotypus d (Penispräparat) Kinabalu Nat. Park, Poring Hot Springs area below Langanan Fall, 800 m, 12.5.1987 (lg. Smetana, cMG); 2 Paratypen ebanda, 520 m 13.5.1987 (lg. Smetana, CF); 1 Paratypus, Eastern Ridge Tr. 790 m, 17.8.1988 (lg. Smetana, cMG); 1 Paratypus, Kinabalu Nat. Park, 1100, 24.5.1897 (1g. Smetana, cMG). DIAGNOSE: Gekennzeichnet durch sehr geringe Größe, konischen Halsschild und 4- gliederige Fühlerkeule. BESCHREIBUNG: Long. 0,80 bis 0,90 mm, lat. 0,40 bis 0,42 mm. Braunschwarz, Fühlergeißel, Beine und Palpen rotbraun, gelblich behaart. Kopf rundlich, der Hinterkopf etwas beulenförmig emporgewölbt, Schläfen fein behaart, Fühler zurückgelegt nur die Halsschildmitte erreichend, mit scharf abgesetzter, 4- gliederiger Keule, ihr 2. Glied eineinhalbmal so lang wie breit, 3 bis 7 sehr klein, 8 bis 10 3-mal so breit wie 7, breiter als lang, 11 schmäler als 10. Halsschild konisch, isodiametrisch bis leicht gestreckt, mit 2 Basalgrübchen, an den Seiten schütter, abstehend behaart. Flügeldecken schon an der Basis zusammen breiter als die Halsschildbasis, mit kleiner Basalimpression, abstehend behaart. Flügel entwickelt. Beine kurz, ohne besondere Merkmale. Penis (Abb. 108) mit schlanken Parameren ohne Tastborsten. Die übrigen Merkmale wurden schon in der Gattungsdiagnose aufgezählt. KATALOG DER BESPROCHENEN ARTEN Gattung Scydmaenus Latreille Untergattung Scydmaenus s. str. novaehollandiae Lhoste minangkabauensis Blattny pseudovestitoides nov. spec. vestitoides Reitter kinabaluensis nov. spec. trapeziceps nov. spec. bukitulari nov. spec. crockerensis nov. spec. borneoensis nov. spec. borneoi nov. spec. fraternus nov. spec. Untergattung Agathelor Schaufuss deplanatus Schaufuss 950 HERBERT FRANZ Untergattung Armatoscydmaenus Franz brevitarsis Schaufuss trapezicollis Lhoste laticeps nov. spec. Untergattung Androscydmaenus nov. subg. densepunctatus nov. spec. Untergattung Mimoscydmaenus Franz crockeri nov. spec. complexipenis nov. spec. Untergattung Mascarensia Franz dissimilis nov. spec. Untergattung Eustemmoides Franz punctatus nov. spec. alessmetandi nov. spec. silvicola nov. spec. cuneipenis Nov. spec. thermarum nov. spec. furcatus nov. spec. burckhardtloebli nov. spec. filipenis nov. spec. parafilipenis nov. spec. allofilipenis nov. spec. bidentipenis nov. spec. frater nov. spec. poringensis nov. spec. livagui nov. spec. sabahensis nov. spec. sabahi nov. spec. allosabahensis nov. spec. parasabahensis nov. spec. sabahanus nov. spec. Gattung Loeblites Franz sabahensis nov. spec. Gattung Horaeomorphus Schaufuss loeblianus nov. spec. sabahensis nov. spec. punctatissimus nov. spec. Gattung Syndicus Motschulsky kinabalui nov. spec. Gattung Syndicomorphus nov. gen. magnus NOV. spec. Gattung Borneosabahia nov. gen. mirifica nov. spec. Gattung Euconnus Thomson Untergattung Euconnus s. str. pseudosukhotanus nov. spec. allosukhotanus nov. spec. kinabaluanus nov spec. simillimus nov. spec. latus nov. spec. SCYDMAENIDAE VON SABAH paenetypicus nov. spec. paeneglaber nov. spec. smetanaensis NOV. spec. apicefurcatus nov. spec. Untergattung Borneoconnus nov. sub. laticlava nov. spec. sabahanus nov. spec. eremita nov. spec. Untergattung Napochus Reitter kinabalui nov. spec. mirus NOV. Spec. allomirus nov. spec. layangensis nov. spec. quinquearticulatus nov. spec. ventriculosus nov. spec. burckhardtianus nov. spec. valdeobscurus nov. spec. parakinabalui nov. spec. fuscus nov. spec. funestus nov. spec. sabahinanus nov. spec. borneoi nov. spec. paramirus nov. spec. Untergattung Napoconnus Franz cephalotes nov. spec. parallelipenis nov. spec. valdepullus nov. spec. langananensis nov. spec. elongatior nov. spec. tambunanus nov. spec. proceripenis nov. spec. Euconnus spec. incertae sedis globicollis nov. spec. fraudulentus nov. spec. kinabalumontanus nov. spec. parakinabalumontanus nov. spec. minutipenis nov. spec. glandulipenis nov. spec. robusticeps nov. spec. paramerorum nov. spec. circumlatus nov. spec. filipenis nov. spec. stylifer nov. spec. paraglobicollis nov. spec. pondoki nov. spec. crockeranus nov. spec. valdepilosus nov. spec. tortricornis NOV. spec. borneoensis nov. spec. crockeri nov. spec. 951 952 HERBERT FRANZ sabahi nov. spec. protectus nov. spec. crockericola nov. spec. excelsipenis nov. spec. longitubus nov. spec. indecorus nov. spec. pilosiceps nov. spec. smetanai nov. spec. silauensis nov. spec. acuticornis nov. spec. livagoensis nov. spec. poringensis nov. spec. longeacuminatus nov. spec. cuneipenis NOV. spec. ubahanus nov. spec. masculinus nov. spec. ovulipenis nov. spec. bicornipenis nov. spec. fontium nov. spec. Gattung Protoscydmus nov. gen. sabahensis nov. spec. PHYLOGENETISCH- BIOGEOGRAPHISCHE ERGEBNISSE Die Bearbeitung der Scydmaenidenfauna von Sabah hat zur Abrundung meiner seit 20 Jahren schrittweise durchgeführten Studien der Scydmaenidenfauna Südostasiens geführt. Sie hat gezeigt, dass der Raum des Nat. Parks des Mount Kinabalu als eines der Entwicklungszentren der Bodenfauna von SO-Asien anzusprechen ist, einer Fauna, die von Borneo über Indonesien bis Thailand reicht. Im Raum des Kinabalu-Nat. parks sind nun 114 Scydmaeniden-Arten nachgewiesen, ausserordentlich viele, wenn man vergleichsweise berücksichtigt, dass Burckhardt und Löbl im Zuge ihrer Aufsammlungen in Thailand, in einer etwa 1000 Exemplare umfassenden Scydmaenidenausbeute nur 47 Arten von Scydmaeniden gefunden haben (FRANZ 1987, 1989a). Die Verwandtschaft der Scyd- maenidenfauna von Thailand zu der von Sabah hat sich allerdings als sehr eng erwiesen, was aus dem Umstand hervorgeht, dass die Gattung Loeblites Franz und die Untergattung Mimoscydmaenus Franz der Gattung Scydmaenus diesen beiden Gebieten als Endemismen gemeinsam sind. In die gleiche Richtung weist auch die nahe Verwandtschaft der Euconnus- Fauna von Thailand und Sabah, vor allem der Arten der Untergattung Euconnus s.str. Aufschlussreich ist auch ein Vergleich der Scydmaenidenfauna von Sabah mit derjenigen von Malaysia, Singapore und Sumatra (SCHAUFUSS 1884, BLATTNY 1926, 1935, FRANZ 1970, 1984 und 1989b), der ergibt dass auch diese Gebiete dem südostasiatischen Faunenbereich angehören. Dagegen gehört die Fauna von Sri Lanka, von wo 198 Scydmaenidenarten bekannt sind (BESUCHET 1971, FRANZ 1971, 1981) einem anderen Faunenbereich an, dem südindischen (vgl. FRANZ im Druck) und letzten Endes der Fauna des alten Gondwana-Kontientes mit naher Verandtschaft zu Madagaskar (FRANZ 1981). Die Fauna von Papua (FRANZ in Druck) ist australisch und somit einem ganz anderen Faunenbereich angehörig. SCYDMAENIDAE VON SABAH 953 LITERATUR BESUCHET, CL. 1971. Les Clidicus de Ceylan (Col. Scydmaenidae). Mitt. Schweiz. ent. Ges. 43: 249-257. BLATTNY, C. 1926. Fauna Sumatrensis (Bitrag Nr. 24) Scydmaenidae (Col.). Suppl. Entom. 14: 1-8, Taf. 1. — 1935. Fauna Sumatrensis (Bijdrage Nr. 76) Scydmaenidae, Col.). Pars IL. Tijdschr. Entom. 78: 240-248. FRANZ, H. 1970. Zur Kenntnis der Scydmaeniden-Fauna von Singapore, Malakka und Indonesien. Beitr. Entom. 20: 535-578. = 1971. Revision der Gattung Syndicus Motsch. (Coleopt. Scydmydmaenidae). Kol. Rdsch. 49: 11-28. — 1975. Zweiter Beizrag zur Kenntnis der Scydmaenidenfauna Südostasiens. Sber. Österr. Akad. Wiss., Math. nat. Kl. Abt. I, 183: 51-107. = 1981. Coleoptera: Die Scydmaenidae Sri Lankas (mit Ausnahme der Genera Cephennium s.lat., Clidicus und Syndicus). Entom. scand. suppl. 111: 125-274. — 1983. Scydmaeniden des Ungarischen Naturwissenschaftlichen Museums in Budapest aus Südostasien: Sri Lanka, Thailand und Vietnam. Folia entom. Hungar. 44: 175-187. - 1984. Beitrag zur Kenntnis der Scydmaeniden von Sumatra und benachbarter Gebiete. Sber. Osterr. Akad. Wiss., Math. Nat. KI. Abt. I, 193: 89-142. - 1985. Neue und ungenügend bekannte Scydmaneniden (Coleoptera) aus Taiwan, Fukien, und Thailand. Mitt. Münch. ent. Ges. 74: 91-128. = 1986. Ein neues Genus und ein neues Subgenus der Scydmaenidae (Coleoptera) aus Siidost- Asien. Revue suisse Zool. 93: 965-970. - 1989a.4. Beitrag zur Scydmaenidenfauna von Thailand (Die Scydmaenidenausbeute von D.H. Burckhardt u. I. Löbl). Revue suisse Zool. 96: 33-80. — 1989b Neue Scydmaenidenarten (Coleoptera) aus Malaysia. Ztschr. Arbeitsgem. Österr. Entom. 41: 84-88. ~ 1992. Scydmaeniden aus Malaysia und Sulawesi (Coleoptera: Scydmaenidae). Kol. Rdschau 62:69-76. LHOSTE, J. 1938-1939. Etude de quelques Scydmaenidae des rives de l'Océan Indien. Arb. morph. taxon. Ent. Berlin 5: 109-127, 6: 1-10 und 281-287. REITTER, R. 1883. Beitrag zur Pselaphiden- und Scydmaeniden-Fauna von Java und Borneo. Verh. zool. bot. Ges. Wien 33: 283-302. SCHAUFUSS, L.W. 1884. Die Scydmaeniden Nord- und Ostasiens, der Sundainseln und New Guineas im Museo Civico di Storia Naturale zu Genua. Annali Mus. civ. Stor. nat. Giacomo Doria. (2), 1, (XXI): 393-420. — 1889. Neue Scydmaeniden im Museum Ludwig Salvator. Berl. ent. Z. 33: 1-42. via Uhr LAST REVUE SUISSE DE ZOOLOGIE Volume 99 — Fascicule 4 ZooLoGIA 92 “Systematics and phylogeny: theoretical, morphological, biochemical and biogeographical aspects” Geneve, 3-4 April 1992. (Annual Conference of the Swiss Zoological Society) ................. KELLER, Albert. Note sur une étude comparative des jarres primaires de trois espèces d'Equidae: Equus asinus, E. przewalskii et E. caballus ConDE, Bruno. Un Palpigrade énigmatique de Thaïlande avec une brève reyuerdesserandesidivisions dellordrer... rn. engen MULLER, Sonia et Claude WEBER. Les dents des sous-familles Hyposto- minae et Ancistrinae (Pisces, Siluriformes, Loricariidae) et leur valeur CAXONOIMIQUE serene ee NR BOLETZKY, Sigurd v. Evolutionary aspects of development, life style, and reproductive mode in incirrate octopods (Mollusca, Cephalopoda) .... BASSET, Yves and Daniel BURCKHARDT. Abundance, species richness, host utilization and host specificity of insect folivores from a woodland site, with particular reference to host architecture ........................ GERY, J. and Volker MAHNERT. Notes sur quelques Brycon des bassins de l'Amazone, du Parana-Paraguay et du Sud-Est brésilien (Pisces, @haraciformessCharacidae) re NE ne MENDES, Luis F. Some new data on the Nicoletiidae (Insecta: Zygentoma) KonmWEurope AN ASIA MINOR... ee ee SARTORI, Michel. Mayflies from Israel (Insecta; Ephemeroptera) I.- Hepta- geniidae, Ephemerellidae, Leptophlebiidae & Palingeniidae ........... FRANZ, Herbert. Monographie der Scydmaenidae (Coleoptera) von Sabah (NOÉ one) Re Re. thee ei a lh eRe Pages 727-734 733-139 741-746 747-754 755-770 771-791 793-819 821-834 835-858 859-953 REVUE SUISSE DE ZOOLOGIE Volume 99 — Number 4 ZOOLOGIA 92 “Systematics and phylogeny: theoretical, morphological, bio- chemical and biogeographical aspects” Genève, 3-4 April 1992. (Annual Conference of the Swiss Zoological Society) .................... KELLER, Albert. Note on a comparative study of the guard hair of three © Equidae species: Equus asinus, E. przewalskii and E. caballus ............ CONDE, Bruno. An enigmatic Palpigrade from Thailand, with a short review Cite main divasions.of the Order... ee MULLER, Sonia and Claude WEBER. The teeth of the subfamilies Hyposto- minae and Ancistrinae (Pisces, Siluriformes, Loricariidae) and their taxonomicalbavalue; sit-start SITE BOLETZKY Sigund, v. Evolutionary aspects of development, life style, and reproductive mode in incirrate octopods (Mollusca, Cephalopoda) .... BASSET, Yves and Daniel BURCKHARDT. Abundance, species richness, host utilization and host specificity of insect folivores from a woodland site, With particular reference to hostrarchitecture "Pr Re GERY, J. and Volker MAHNERT. Notes on some Brycon (Pisces, Characi- formes, Characidae) from the Amazon, Parana-Paraguay and South- Eastern» Brazillsystems-.......s0.r.e ee TALI TI MENDES, Luis F. Some new data on the Nicoletiidae (Insecta: Zygentoma) fromsEuropesand. Asia Minora.----..-0--s as eme ne OUTRE SARTORI, Michel. Mayflies from Israel (Insecta; Ephemeroptera) I. Hepta- geniidae, Ephemerellidae, Leptophlebiidae & Palingeniidae ............ FRANZ, Herbert. 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