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MAM BID BER, OA tee PPT Rho ALII AREA CAPE ETAT eae RS TETE CCE ET ERP AS PEER TES EE RTC ET 3 pada AR SEES erben aOR oA en cho A DM GOR LA ay Ze a ete BEER BEE FR ER Lath wine FEN rs em en Geben re sr COPIES AIS SAMA Len CPC CAO EA IR RASTA RELA RSM SIE BM pi eae nes di AA DE a EEE BIETEN nd MR oh A ARSA A = deine nik ST TE to she eA atk: DEA nes VO hi nun GR IRRE En Vin (tA hh 6100822 ALINA SS AREA He na habe FE BEI CRUE Sar en 27 Dessins Zar dieta, ra EEE TE 4 na age 288,0 è CRETE NE ey DLE RER) Par eras A Msi eek ieee Eee Re rm “sos wre Die BEE ee ER entrate air TEST tg TAPER matt yd SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE REVUE SUISSE DE ZOOLOGIE TOME 107 — FASCICULE 3 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève MANUEL RUEDI Chargé de recherche au Muséum d'histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d'histoire naturelle de Genève Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des Instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L'ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230.— (en francs suisses) Les demandes d'abonnement doivent étre adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse ANNALES de la SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genéve tome 107 fascicule 3 2000 g kl GENEVE SEPTEMBRE 2000 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE SUISSE Fr. 225.— REVUE SUISSE DE ZOOLOGIE TOME 107 — FASCICULE 3 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève MANUEL RUEDI Chargé de recherche au Muséum d’histoire naturelle de Genève CHARLES LIENHARD Charge de recherche au Muséum d’histoire naturelle de Geneve Comité de lecture I] est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Geneve et de représentants des Instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 N A EN Internet: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L'ABONNEMENT: (en francs suisses) Les demandes d'abonnement doivent étre adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse UNION POSTALE Fr. 230.— REVUE SUISSE DE ZOOLOGIE 107 (3): 419-559; septembre 2000 The genus Dianous Leach in China (Coleoptera, Staphylinidae) 261. Contribution to the knowledge of Steninae Volker PUTHZ c/o Limnologische Fluss-Station des Max-Planck-Instituts für Limnologie Damenweg 1, D-36110 Schlitz, Germany. The genus Dianous Leach in China (Coleoptera, Staphylinidae). 261. Contribution to the knowledge of Steninae. - A revision of the species of Dianous Leach of China, including 4 new synonyms and descriptions of 36 new taxa, 10 of them from Taiwan (= T), 2 from Laos, | from Vietnam and 1 from the Philippine Islands, is presented: Dianous adonis sp. n., D. alcy- oneus sp. n. (Laos), D. angulifer sp. n. (T), D. ater sp. n. (T), D. atro- coeruleus sp. n., D. atrocyaneus sp. n. (T), D. atroviolaceus sp. n., D. biguttulus sp. n., D. bilunatus Puthz, 1994 syn. n. = D. taiwanensis Puthz, 1971, D. bioculatus sp. n., D. brevicornis sp. n., D. cebuensis sp. n. (Phi- lippine Islands), D. coeruleovestitus sp. n., D. coeruleotinctus sp. n., D. cupreogutta sp. n. (T), D. cyaneocupreus Sp. n. (T), D. dubiosus sp. n., D. electrigutta Sp. n. (T), D. fauveli sp. n. (Vietnam), D. gemmosus sp. n., D. gonggamontis Sp. n., D. grandistigma Sp. n., D. haraldi sp. n. (Laos), D. inaequalis laevior ssp. n. (T), D. karen Rougemont, 1981 spec. propr. (not syn. of D. vietnamensis Puthz), D. lobatipes sp. n., D. mendax sp. n. (T), D. miripes Rougemont, 1985 syn. n. = D. latitarsis L. Benick, 1942, D. ocelli- fer sp. n., D. puberulus L. Benick, 1942 syn. n. = D. punctiventris Champion, 1919, D. rougemontianus sp. n., D. rugipennis sp. n., D. rugo- sipennis sp. n., D. schoenmanni sp. n., D. senex sp. n., D. septentrionalis Naomi, 1988 syn. n. = D. japonicus Sawada, 1960, D. sucinigutta Sp. n. (T), D. suciniguttatus sp. n., D. sucininotatus sp. n. (T), D. variegatus sp. n., D. yangae sp. n.. Some morphological characters are discussed, and the genus is tentatively grouped. 24 first records for various species and countries, distribution tables, and a key to species are presented. Key-words: Coleoptera — Staphylinidae — Dianous — South-East Asia - China — taxonomy — morphology — distribution. INTRODUCTION The genus Dianous Leach, with (now) 200 taxa, is mainly distributed in the mountainous areas of the Oriental region. When I first revised the genus (Puthz, 1980, 1981) 17 species were known from China. Recent collections have increased species Manuscript accepted 29.03.2000 420 VOLKER PUTHZ number considerably, a tendency which is still in progress. In this revision 78 species, 32 of them new to science, are reported from China. This study was started in 1992 and exclusively focussed on the Taiwanese species, but later on it was extended to the whole of China, including also neighbouring countries. The constant influx of new material required a permanent extension. Besides, an unexpected impediment delayed completion of the manuscript: in 1993 and 1994 Zheng published two articles on Chinese Dianous in Chinese with short English summaries, but reprints and some paratypical material for comparison were not sent to me until 1998! Consequently, I had to change the manuscript several times. Fortu- nately, most of the species described by Zheng were found among the material at hand, but original material of four taxa (D. acutus, D. alternans, D. bashanensis, D. emei- ensis) is still missing. Although the revision of the Chinese Dianous is still at its beginning, I think it reasonable and necessary to fix the present state of knowledge, on which future investigations may be based. General remarks on the grouping of the species are added, and the present knowledge of their distribution is given in tables, including numerous first records. PART I: GENERAL REMARKS In 1981, I proposed two groups of the genus: group I, including all those species which were at first thought to belong to the genus Stenus Latreille (middle of frons concave, medial portion not elevated, eyes very large, without or with very short temples); group II, with species in which the frons has two distinct lateral furrows and a distinctly elevated medial portion (eyes mostly smaller, distinct temples present). This grouping works in most cases. Only few species do not exactly fit the group definitions: D. corticicola Puthz, D. puthzi Lundgren, D. naicus Puthz, D. smetanai Puthz have more or less elevated medial portions of the frons and/or distinct temples. The latter two seem to be intermediate between both groups. Species of group I, about 30 % of all Dianous, represent a close complex (large, Stenus-like eyes, tarsal structure simple, structure of sternum 9 and of the aedeagus rather uniform, sculpture of the exosceleton also rather uniform). The following are the taxa of group I (distribution only given for species which are not listed in table 1): amicus Puthz: Borneo carinipennis (Bernhauer) araeocerus (L. Benick): Borneo cebuensis sp. n.: Philippine Islands aurichalceus (Champion) concretus Puthz: Borneo baliensis Rougemont: Bali corticicola (Puthz) benicki Puthz: Philippine Islands cupreoaeneus (Champion) bhotius Rougemont cyaneovirens (Cameron) bhutanensis Rougemont dajak Puthz: Borneo boops Puthz: Borneo davaomontium (Puthz): Philippine Islands bracteatus (Champion) flavicoxatus (L. Benick): Sumatra burckhardti Puthz: Borneo fluctivagus Puthz THE GENUS DIANOUS IN CHINA 421 haraldi sp. n. nokrekensis Puthz hirsutus Rougemont philippinus Puthz: Philippine Islands hygrobius (L. Benick): Java pykaranus (Cameron) inconspicuus Rougemont reformator Rougemont iridicolor (Scheerpeltz) rougemonti Puthz: Borneo jaechi Puthz: Ceram saxicola (Puthz): Philippine Islands Javanicola Puthz: Java schillhammeri Puthz: Sumatra kinabalumontis Puthz: Borneo semicoeruleus (Cameron): Malaysia lahu Rougemont shan Rougemont lasti Puthz siberutensis Puthz: Siberut lividus (L. Benick): Philippine Islands striatellus (L. Benick): Java, Bali loebli Rougemont sulcipennis Puthz loeblianus Puthz : Borneo tiomanensis Puthz: Tioman martensi Rougemont tonkinensis (Puthz) meo Rougemont uncinipenis Puthz nagamontium Puthz viridicupreus Rougemont nepalensis Rougemont viriditinctus (Champion) niger Rougemont wittmeri Rougemont nigrovirens (Fauvel) yao Rougemont nilgiriensis Puthz About 70% of the Dianous belong to group II with a variety of morphological types and probably also monophyletic complexes, which are here tentatively distinguished. Some of the morphological characters which are used in defining species complexes need some explanation and comments, respectively: (a) The tarsal shoe (figs 1-6): this very peculiar character has been described by Rougemont (1985: p. 136 nota 2) who states that there are several species which have such “soles” and that this character is more or less developed in different species. The tarsal shoe consists of densely set, long setae, which in some species are modified into willow-like leaves (figs 3, 5). These setae are found on all tarsal segments or on the distal ones. The tarsal shoe can be best seen in tarsi, which are free of mounting glue, and should be observed in ventral aspect of the protarsi and in lateral aspect of the last tarsal segment of meso- and of metatarsi (as in figs 1, 4). In species which have a weak tarsal shoe one finds a thin setaceous sole which is longer than half the length of tarsal segment 5. For functional analysis I refer to my friend Rougemont: “It may be assumed that the structure is functional, increasing the tarsus’ resistance to the surface tension of water, and possibly also increasing the animal’s bouyancy by trapping air between the sole, the setae and the tarsus”. (b) Structure of tarsal segment 4: most species have “simple tarsi” without any lobes. In few species tarsal segment 4 is asymmetrically lobed (fig. 7), in other very deeply bilobed (figs 8, 9). Intermediate forms occur between these three character states (figs 10-12, 139-142, 219, 220). In some species segment 3, too, is bilobed. For grouping the species, this character is of much more restricted value than in the genus Stenus Latreille. 422 VOLKER PUTHZ (c) Female sternum 8: The outline of the female sternum 8 is an important taxonomic character and requires description. Usually this sternum has an evenly rounded posterior margin, but in some groups there are emarginations of different depth (figs 47, 48, 76, 77, 79, 80, and variability: figs 61-67), in others the medioposterior portion is elongated and more or less projecting (e. g. figs 88, 93-98, 193-199). (d) Sternum 9: The outline of this sternum, including apicolateral brush (length, strength), is also an important character, which should be described. Posterior margin of this sternum can be acute laterally (with a distinctly proment “tooth”: figs 50, 90, 95, 99, 100, 116, 118-120, 135, 136, 138, 150, 204-208, 237), serrate (figs 75, 78, 91, 96, 117, 229-233) or smooth (without cuticular fringes: figs 42, 43, 53). This character 1s of restricted value for species grouping, since there are intermediate forms. In the males of numerous species, sternum 9 has peculiar cuticular structures apicoventrally: the most usual type is represented by apically spinulose arcuate cuticular laminae (= asacl; figs 13-18) which can be seen even by higher light-microscopical magnifications. Additionally, there are areas with glandular openings (figs 16, 18). Similar cuticular characters are widely distributed in arthropods (e. g. in Isopoda (Eisenbeis & Wichard, 1985: 125), Plecoptera (Kapoor, 1997), Trichoptera (Wichard er al., 1995: 201). In Dianous, they are specific not only for single species but also for species complexes. Number, density and distribution of these cuticular structures might be also of taxonomic value and should be studied carefully. Narrow cuticular protuberances on sternum 9 and at the actual posterior margin of that sternum render it fimbriate (fig. 14). In Dianous gemmosus sp. n. there is a field of fine cuticular spines apicomedially (des: (e) Aedeagus: Compared to Stenus Latreille, the aedeagi of Dianous are much more uniform both in outline and in internal structure. Nevertheless some “types” can be distinguished: (1) Species which have two groups of parameral setae (groups which differ in their setal strength and groups which are placed on different portions of the parameres; this is an easy observable character: figs 82, 83). (2) Species with an entire outline of the anterior median lobe, which in most cases is narrowed triangularly (e. g. figs 38-40, 74, 177-181, 209-214) . (3) Species which have an excavated anterior portion of the median lobe (figs 19, 121-130, 143, 144). (4) Species with a basally case- like internal sac (figs 39, 40, 44-46, 69) or with a distally nightcap-like internal sac (fig. 113) or with membranous structures densely set with chitinous tufts (fig. 70). (f) It should be noted that in some species the outline of tergum 10 differs between: male and female; in those cases, the female tergum is distinctly triangular, not or much less so in the males (figs 71-73, 200, 201). (g) Pubescence: Numerous species are characterized by special arrangement of the pubescence, especially that of the elytra, forming whirls or other patterns (fig. 20). Some species have very long pubescence, which can be erect (figs 21, 22) or curved and/ or more or less recumbent (figs 23, 24). (h) Sculpture of the exosceleton: Besides a punctation of differing strength and density, numerous species have areas in which the sculpture is rugose, coalescent or vorticose (figs 25-28). The microsculpture may be more or less distinct and was found to be of diagnostic value. THE GENUS DIANOUS IN CHINA 423 Proposed species complexes of group II (? = one of both sexes missing or has to be verified by more material): (1) calceatus-complex: mostly relatively small eyes, temples at least 1/2 the length of eye, tarsi (mostly) with a distinct tarsal shoe (setae willow-like broadened in * figs 3, 5) (missing in #), segment | of metatarsi (mostly) much longer than the combined length of segments 2-4, segment 4 simple or asymmetrical. Sternum 9 (mostly) serrate or smooth apicolaterally, lacking apically spinulose arcuate cuticular laminae. Male: median lobe narrowed conically (broadly rounded in D. gregarius), apex rounded, internal structures membranous with tufts and a case-like structure basally, setae of parameres uniform, arranged equally. Female: Sternum 8 broadly rounded or (mostly) emarginate apically. Sculpture of exosceleton mostly fine and dense (not coarse and rugose). Coloration variable: unicolorous, metallic and/ or with yellowish spots on elytra. adonis sp. n. gregarius Rougemont * arachnipes Puthz inaequalis Champion alternans Zheng inaequalis laevior ssp. n. annandalei Bernhauer kabakovi Puthz # assamensis Cameron * latitarsis L.Benick bifoveifrons Champion malayanus Cameron # brevitarsis Puthz margaretae Rougemont caeruleonotatus Champion * moritai Naomi * calceatus Puthz # nigrocyaneus Puthz # camelus Puthz obliquenotatus Champion cameronianus Jarrige obscuroguttatus Cameron chetri Rougemont ? pallitarsis L. Benick coeruleomicans Puthz ponticus Fagel coeruleotinctus sp. n. punctiventris Champion coeruleovestitus sp. n. # robustus Cameron consors Cameron scabricollis Champion convexifrons Puthz schoenmanni sp. n. cyanogaster Champion * sichuanensis Puthz elegantulus Zheng siwalikensis Cameron fauveli sp. n. srivichaii Rougemont flavoguttatus Puthz tumidifrons Puthz * gracilipes Champion versicolor Cameron (2) yunnanensis-complex: similar to the calceatus-complex, tarsal shoe thin, segment | of metatarsi as long as or slightly less long than the combined length of segments 2-4, sternite 9 rounded apicolaterally. Male: internal structures of aedeagus different: no case-like structure basally, but with spinulose, membranous or strongly sclerotized tube (figs 70, 74). Female: sternum 8 emarginate apically. acuminifer Puthz uniformis Zheng angulifer sp. n. yunnanensis Puthz 424 VOLKER PUTHZ (3) lobigerus-complex: large eyes, temples less than 1/2 the length of eye, without tarsal shoe, tarsal segment 4 very deeply bilobed (figs 8, 9), sternum 9 with a long tooth apicolaterally (one exception: rounded-serrate in D. spiniventris), no spinu- lose arcuate cuticular laminae apicomedially. Male: Apical portion of median lobe broadly lanceolate or narrowly triangular with ventral pubescence, internal sac with a moderately broad, long, sclerotized tube (some species have an strongly sclerotized expulsion mechanism distally: *), parameres with one group of uniform setae. Female: sternum 8 broadly rounded or with a rounded projection apicomedially. Sculpture mostly coarse, well-defined or rugose. Coloration: black or metallic, some species have elytral spots. azureus Champion lobigerus Champion * bimaculatus Cameron luteoguttatus Champion cribrarius Champion * siamensis Rougemont cruentatus L. Benick spiniventris Puthz * flavoculatus Puthz * strabo Puthz * hainanensis Puthz (4) aereus-andrewesi-complex: eyes large, temples less than 1/2 the length of eyes, without tarsal shoe, tarsal segment 4 simple, sternum 9 serrate apicolaterally (2 exceptions: *), with spinulose arcuate cuticular laminae or spines. Male: median lobe narrowed conically, apex not emarginate, apical portion densely set with long setae and/ or ventral ridges (+), internal sac with coarse tufts basally, parameres with two groups of setae of different strength. Female: sternum 8 roundly prominent apicomedially. Sculpture of the exosceleton variable, mostly with rugose areas. Coloration: metallic, no yellowish spots on elytra. + aeneus Cameron + minor Champion + aereus Champion +? radiatus Champion + andrewesi Cameron + rougemontianus Sp. n. ? brevicornis sp. n. + ruginosus Zheng + championi Cameron ? rugipennis Sp. n. ? cupreoviolaceus Puthz ? rugosipennis sp. n. ? femoralis Cameron + vietnamensis Puthz + frater Cameron + viridipennis Cameron + gracilis Puthz + * violaceus Puthz + karen Rougemont (5) ocellatus-complex: eyes large, temples less than 1/2 the length of eye, without tarsal shoe, tarsal segment 4 (mostly) shortly bilobed, sternum 9 serrate or acute apicolaterally with spinulose arcuate cuticular laminae apicomedially. Male: Median lobe more or less triangularly narrowed, rounded apically, densely pubescent apicoventrally (figs 85, 177-181), internal sac with a nightcap-like distal portion, para- meres with one group of uniform setae. Female: sternum 8 rounded, mostly projecting apicomedially. Sculpture variable, moderately coarse, punctation well-defined or rugose. Coloration: most species have elytral spots. THE GENUS DIANOUS IN CHINA 425 alcyoneus sp. n. ocellatus Cameron atrocoeruleus Sp. n. (incl. acutus Zheng) distigma Champion ocellifer sp. n. emarginatus Zheng oculatipennis Puthz grandistigma Sp. n. variegatus Sp. n. luteolunatus Puthz verticosus Eppelsheim luteostigmaticus Rougemont ? psilopterus L. Benick (6) chinensis-complex: large eyes, temples less than 1/2 the length of eye, without tarsal shoe, tarsal segment 4 simple to moderately bilobed (figs 9-11, 139-142), sternum 9 serrate or acute apicolaterally with spinulose arcuate cuticular laminae. Male: apical portion of median lobe broad, emarginate, internal sac with a nightcap-like distal portion (fig. 113), parameres with two groups of setae of different strength. Female: sternum 8 roundly prominent apicomedially. Sculpture of the exosceleton variable. Coloration: metallic, mostly with yellowish spots on elytra. aequalis Zheng hammondi Rougemont atrocyaneus Sp. n. hummeli Bernhauer atroviolaceus Sp. n. senex Sp. n. banghaasi Bernhauer socius Zheng bashanensis Zheng subtortuosus Champion bioculatus sp. n. subvorticosus Champion chinensis Bernhauer taiwanensis Puthz cyaneocupreus Sp. n. tortuosus Champion dubiosus Sp. n. tortus Cameron emeiensis Zheng yangae sp. n. gemmosus Sp. n. yoshidai Naomi gonggamontis sp. n. (7) coerulescens-complex: eyes large, temples less than 1/2 the length of eye, without tarsal shoe, tarsal segment 4 simple or bilobed (*), sternum 9 rounded-serrate, or acute apicolaterally, spinulose arcuate cuticular laminae present apicomedially. Male: Apical portion of median lobe more or less triangularly narrowed, internal sac distally simple or pointed (#, figs 209, 218, 220), parameres with one group of uniform setae. Female: sternum 8 rounded or slightly projecting apicomedially. Sculpture: punctation mostly well-defined. Coloration: unicolorous, metallic and/ or with elytral spots. Some species (see below) are close to group I of the genus by their general habitus, one of them (D. puthzi) has thin tarsal shoes and resemble D. malayanus Cameron, but the internal structure of the aedeagus and the 8" sternum of the female do not fit the definition of the calceatus- complex. # amamiensis Sawada * cameroni Champion ater sp. n. chalybaeus Le Conte 426 VOLKER PUTHZ coerulescens Gyllenhal morimotoi Naomi (with ssp. anatolicus Korge and ssp. nitidulus Le Conte elegans Khnzorian) shibatai Sawada cupreogutta Puthz #* sucinigutta sp. n. cupreostigma Puthz #* suciniguttatus sp. n. #* electrigutta sp. n. #* sucininotatus sp. n. # freyi L. Benick * sulcatipennis Puthz gongen Watanabe viridicatus Naomi iwakisanus Watanabe Japonicus Sawada near Dianous group I: # klapperichi L. Benick naicus Puthz * lobatipes sp. n. puthzi Lundgren mendax sp. n. smetanai Puthz Concerning the ecology of Dianous-species (Puthz, 1981: 91 f.), there is only little supplementary information: G. M. de Rougemont published some substantial observations in 1985; e. g. concerning Dianous robustus Cameron: “was found clinging to and moving over a vertical rock face down which ran a steady film of water, some of the insects ‘up to their knees’ in the water, while 2 exx. were seen totally submerged.” New records confirm the published data according to which several Dianous species occur together in the same habitat (see also Puthz, 1988: 633), some- times in large numbers (figs 29, 30). Most Dianous are steneceous and belong to the bryomadicolous fauna (Vaillant, 1955). There are, however, some exceptions: D. corticicola Puthz (Vietnam) was found by sifting bark of tree, D. concretus Puthz (Borneo), remarkable species by the concrescent elytra, by sifting dead leaves and dry mosses in open forest (Puthz, 1988: 638), and some of the new described Taiwanese species (D. sucininotatus, D. electri- gutta, D. ater) were found in soaking wet moss growing on rotten wood around small ponds in a coniferous forest, by sifting moss and debris on a forest seapage, by sifting of wet moss on large fallen trees in an Abies-forest and in tufts of Carex at swampy edge of a small lake. Since there are no records from exclusively dry habitats, it can be assumed that species, which do not live in or near torrents, on rocks in running waters, at sprays of waterfalls etc. nevertheless need a wet or humid environment as is provided by mosses and humus in humid forests. — The genus occurs from lower elevations to nearly the alpine zone: a Taiwan locality below of Yushan main Peak (3650 m) is the highest hitherto known locality (see below: D. sucinigutta sp. n.). The distribution of species is shown in three tables: (1) a table which lists all the Dianous species of the Oriental region except those which are restricted to Malaysia and the Sunda-Archipelago, (2) a table showing the distribution of all species known from China including neighbouring countries, (3) a distribution table of the Taiwanese species according to Taiwanese provinces. Table | is divided into 13 columns for the respective countries; India is listed in three columns, since there is a (relictal) special Dianous-fauna found in the South THE GENUS DIANOUS IN CHINA 427 Indian mountains and some differences between the Assam-fauna and that of the rest of northwest India. Aside from records given in the taxonomic section of this article, the following first records are included: Dianous aereus Champion: Burma: 23 d d, 28 E Piyin Oo Lwin, 1070 m, 22°03.523° N 96°31.95 Schillhammer (20) (NHMW, cP). Dianous bimaculatus Cameron: Nepal: 1 d: env. Shivalaya, banks of Kimti Khola. 2. V. 1993, A. Kleeberg (cP). Dianous cribrarius Champion: Nepal: 2 d d, 2 9 2, Annapurna Reg., env. Biretanthi, 1200 m, Bhurungdi Khola, 4. X. 1992, A. Weigel (Naturhistorisches Museum Erfurt, cP). Laos: | 9: Prov. Lg. Nam Tha ca. 5 km S Muang Sing, 650 m, Huay Giuom river, 10. VI. 1996, Schillhammer (21) (NHMW). Dianous hammondi Rougemont: N. India: 1 4,2 9 ©, Khasi Hills (IRSnB, MHNG). Dianous hirsutus Rougemont: Vietnam: 1 © (cf. det.), 40 km NW An Khe Buon Luoi, 620- 750 m, 14°10° N, 108°30° E, 28. IIL.- 12. IV. 1995, Pacholatko & Dembicky (NHMW). Dianous kabakovi Puthz: Laos: 6 d d, 5 © 2, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./ 13. VI. 1996, Schillhammer (25) (NHMW, cP); 1 4, Bolikhamsai prov., Ban- Nape- Kaow- Nua- Pass, 18. IV. — 1. V. 1998, Jendek & Sausa (coll. Hromädka). Dianous lobigerus Champion: Nepal: 1 d, env. Shivalaya, banks of Kimti Khola, 2. V. 1993, A. Kleeberg (cKleeberg). Dianous minor Champion: Laos: 8 d 4,7 2 2, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./13. VI. 1996, Schillhammer (25) (NHMW, cP); 3 64,2 9 9: Prov. Lg. Nam Tha, ca. 30 km NW Lg. Nam Tha, 800 m, 16./18. VI. 1996, Schillhammer (28, 30) (NHMW, cP). Dianous obliquenotatus Champion: Thailand: 1 2, Sob Pong, 24. XII. 1989, H. Malicky (cP); 1 2, Doi Inthanon, Bang Khun Klang, 98°32‘E, 18°32‘N, 900 m, 6. I. 1990, Chantaramongkoi & Malicky (MHNG). Dianous scabricollis Champion: Laos: 1 4, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./13. VI. 1996, Schillhammer (25) (NHMW): This specimen is distinguished from N. Indian specimens (types) by the in general dark Prussian- blue colour, elytral band sky- blueish, posteriolateral quarters of elytra strongly aeneous; aedeagus similar to that of the type series. ? 2: Myanmar, Mandalay Division, 8 km 6:EX6Pwe Kauk Wit, 195 xe 1.998; Dianous siamensis Rougemont: Vietnam: 1 9, 12 km N Dalat, Lang Bian, 28.- 30. IV. 1994, Pacholatko & Dembicky (NHMW); 1 2, 16 km N Dalat- Ankroat, 1400 m, 12°05‘N, 108°24° E, 15. IV. 1995, Pacholatka & Dembicky (NHMW). Dianous siwalikensis Cameron: Vietnam: 1 E, Tonkin: Kao- Bang (IRScNB). Dianous srivichaii Rougemont: Laos: 17 88,15 2 2, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./13. VI. 1996, Schillhammer (25) (NHMW, cP). Burma: 1 6,7 9 9: Myanmar, Mandalay Division, 8 km E Piyin Oo Lwin, 1070 m, 22°03.523° N 96°31.956°E, Pwe Kauk Wf., 19. X. 1998, Schillhammer (20) (NHMW, cP). Dianous strabo Puthz: Thailand: 1 d, Phetchanban Prov., Nam Nao N.P., 24. XI. 1995, Zettel (22) (NHMW). Dianous yunnanensis Puthz: Nepal: 1 d, 2 9 ©, Sindhupalchowk District, Dolalghat, Sunkoski, 7. XI. 1993, Moog & al. (NHMW, cP). Table 2 lists the occurence of Dianous species in 26 main administrative units of China (mostly provinces, indicated by letters) and in 8 adjacent countries (number). The following 6 biogeographic units are recognized (separated by #): Biogeographic unit 1 (1) N. INDIA, Kashmir; (2) NEPAL, BUTHAN; (A) Sichuan; (3) VIETNAM, LAOS, BURMA, THAILAND; (B) Yunnan; (C) Guizhou 428 VOLKER PUTHZ Biogeographic unit 2 (D) Guangxi; (E) Guangdong + Hongkong + Hainan; (F) Fujian; (G) Taiwan; (4) Riuku-Islands & other Japanese Islands Biogeographic unit 3 (H) Hunan; (I) Hubei; (J) Jiangxi; (K) Zhejiang + Shanghai Biogeographic unit 4 (L) Jiangsu; (M) Anhui; (N) Henan; (O) Shaanxi; (P) Shanxi; (Q) Shandong; (R) Hebei + Beijing + Tianjin; (S) Liaoning Biogeographic unit 5 (5) JAPAN (main country); (6) KOREA; (T) Jilin; (U) Heilongjiang; (7) Russia E of 120°E Biogeographic unit 6 (V) Tibet; (W) Quinghai; (X) Gansu + Ninxia Hui; (Y) Xinjiang Uighur; (Z) Nei Mongol (Inner Mongolia); (8) MONGOLIA There are two well-founded proposals for dividing China into major biogeo- graphical regions, that of Takhtajan (1986) and that of Jäch & Li (1995). Since China has an extremely complicated topography and orography the limits of distinguished regions in both proposals are more or less arbitrary and do not work exactly to classify each individual locality. Therefore, in my table 1 prefer to use the exactly defined political borders of the provinces (or administrative units) and arranging these units according to the aforementioned biogeographical concepts as much as possible. Except for the South China Region and for Sichuan (which should belong to three different zoogeographical regions), most of the other proposed regions in Jäch & Li fit to some degree the complexes of administrative units used in my table. This table will be used also in a series of papers in preparation on the Chinese Stenus-fauna and consequently takes into consideration the adjacent Palaearctic areas, from which no Dianous has hitherto been reported. The genus Dianous is of Northern Oriental origin and has its distribution centre in the mountainous areas between the Palaearctic and the Oriental regions south of 31°northern latitude as can be seen from map | (fig. 31 a). Few species occur either in Europe, in the Caucasus and in the Altai mountains (D. coerulescens with subspecies) or in North America (D. chalybeaus Lec. and D. nitidulus Lec.). Today there is a considerable distance between the Altai population of D. coerulescens and its main range in Europe and the Caucasus, and also between the East Asian Dianous and those of North America (compare map 1 in Puthz, 1981). Since all my hypotheses to explaining these disjunctions are highly speculative, I only present the facts, new records might bring light into the matter. Regarding group- and species- diversity, the distribution centre of Dianous seems a major centre both of speciation and of the preservation of ancient forms. A comparison of map | (fig. 31 a) with map 3 in my first revision of the genus in 1981 shows a considerable increase of knowledge concerning species numbers of different countries and areas, especially for China. Furthermore the distribution area of THE GENUS DIANOUS IN CHINA 429 group II now also includes Sumatra and Java, the South Japanese islands and Japan. It may be noted that group II does not transgress Wallace’s line (sensu Huxley). Dianous jaechi Puthz from Ceram extends the range of the species of group I eastward to Lydekker’s line (no Dianous has been reported from New Guinea!). Map 2 (fig. 31 b) shows the distribution of Dianous in China according to administrative units of that country. Most species are known from Southern Sichuan and from Yunnan, areas which are close to Assam and to Nepal/ Northwest India and which belong to the centre of evolution of the genus. The number of species decreases from southern to northern China and from the west to the east, showing that Dianous is mainly an Oriental element. Six Dianous are reported from Hainan, 5 of them endemic, 1 of them (D. punctiventris) is widespread in the central range of the genus and also known from Vietnam, Yunnan and Burma. The Hainan species belong to three different complexes of group II of the genus, which indicates that at least three different ancestors lived in the larger SE-Indochinese area, from which Hainan now 1s separated as an island. Close relatives have been recorded from Vietnam, which supports close relationships between Hainan and the mainland fauna. I would also exspect a species of group I of the genus, but as in the much better known Taiwan none has hitherto been found. Apart from 1 species (D. luteoguttatus), the remaining 14 Dianous from Taiwan are endemic (not counting D. dubiosus, for which the record needs confirmation). Most of the species occur in the warm temperate mixed forest zone (800-2000 m) and in the cool temperate coniferous forest zone (1900-2800 m), some (D. electrigutta, D. sucininotatus, D. sucinigutta) also or mainly in the cold temperate coniferous forest zone (2600-3000 m) and/ or in the subalpine coniferous forest zone (3000-3600 m). D. sucinigutta has been found from 1660 m up to 3650 m. The Taiwanese species belong to five different complexes of group II of the genus and have mostly strong affinities to the mailand fauna or to the fauna of the southern Japanese islands. This phenomenon is shown in the following closely related species: D. arachnipes — D. moritai (Amami- Oshima); D. sucinigutta — D. freyi (Fujian), D. amamiensis (Amami, Nansei); D. inaequalis laevior — D. inaequalis (Sichuan, Yunnan); D. atrocyaneus — D. yoshidai (Iriomote); D. taiwanensis — D. banghaasi (Fujian). Of special interest 1s D. luteo- guttatus Champion because of its wide range (Western Himalaya/ Nepal — Taiwan) and the broad disjunction between these areas. This species is an example of the strong connection between the old Himalayan fauna and the fauna of Taiwan. Although many Taiwanese Dianous occur at high elevations, no Palaearctic element was found, as e. g. in the genus Stenus Latreille; this is explained by the Oriental origin of Dianous. However, it is worth noting that the most common Taiwanese species, D. taiwanensis, reported from 1600 m up to 2600 m, is closely related to several Northeastern Chinese species which also occur at high elevations (e. g. D. gonggamontis) and which in their general habitus resemble the Dianous of the northern hemisphere. - Two complexes of group II of the genus have not been reported from Taiwan (aereus-andrewesi- complex; ocellatus-complex); the former, however, is known from Hainan. This indicates a different faunal history and supports published data that the Taiwanese fauna is of greater age and has fewer Oriental elements than the fauna of Hainan. The high species 430 VOLKER PUTHZ number of Taiwan in relation to the adjacent mainland provinces of China seems to be striking. I would exspect many more species to occur in Fujian and Zhejiang than are now known; future collecting may be hoped to fill the gap. In the descriptions the following abbreviations are used: acam: antecoxal area of metasternum asacl : apically spinulose arcuate cuticular laminae adE : average distance between eyes als : apicolateral setae of parameres ams : apicomedial setae of parameres Ad: allotype EI: holotype IB: eye length el: length of elytra IR: length of pronotum IS: length of suture ie: length of temples PM: proportional measurements PTE paratype(s) WEI: greatest width of elytra wH: width of head WP : width of pronotum BMNH: British Museum, Natural History, London (now: The Natural History Museum, London) FMCh: Field Museum of Natural History, Chicago CR: coll. Puthz Che coll. de Rougemont CS: coll. Smetana IRSCNB: Institut Royal des Sciences Naturelles de Belgique, Bruxelles MHNG: Muséum d'histoire naturelle, Genève MM: Museo civico di Storia naturale, Milano MNHNP: Muséum National d’ Histoire Naturelle, Paris NHMB: Naturhistorisches Museum Basel NHMW: Naturhistorisches Museum Wien USNM: United States National Museum, now known as National Museum of Natural History, Washington, D.C. ZMB: Museum fiir Naturkunde (der Humboldt-Universitàt) Berlin My thanks are due to all colleagues and institutions which provided their material for study. Above all, I am deeply indebted to Dr. Ales Smetana who contri- buted a major portion of the material, especially from Taiwan, and who revised the major portion of my English descriptions and key. Warm thanks are also due to my friend Guillaume de Rougemont, who generously left his material from China at my THE GENUS DIANOUS IN CHINA 43] disposal, and to Andreas Pütz and Michael Schülke who provided their Chinese material for study. I thank Dr. O. Betz (Kiel) for the REM-photographs and last but not least Volker Assing for correcting the English text of the introduction and of the general remarks. PART II: TAXONOMY AND FAUNISTICS Dianous yao Rougemont Dianous yao Rougemont, 1981a: 330 ff. figs.; 1981b: 359; 1983: 18 Distribution. This species, previously known from Burma and Thailand, occurs also in China: Guizhou: 6 specimens Huangguoshu, X. 1986, G. M. de Rougemont (Rougemont i. L., CR). New to China. Comments. The 2% recorded by L. Benick (1942) as “Stenus cf. assamensis Cameron” from Kambaiti, 2000 m, 1./6. VI. 1934, Malaise (Museum Stockholm) belongs to D. yao. Dianous cebuensis sp. n. Type Material. Holotype (2): Philippine Islands: Cebu: S Badian Matutinao, Kawasan Falls, 2-50 m, 23.- 24. II. 1997, H. Zettel (116) (coll. Zettel in NHMW). PM of HT: wH: 38.5; aE: 22; wP: 28.4; IP: 32; wEl: 37; lEl: 41; IS: 34. This new species belongs to group I of the genus and resembles closely D. saxicola (Puthz) and D. tonkinensis (Puthz). It is remarkable by the sparsely punctate frons. Strongly shining, black with dark bluish-olivaceous reflection, frons moderately coarsely, sparsely punctate, punctation of the pronotum coarse, dense, slightly confluent, elytral punctation (except on sides and near posterior margin) very coarse and transversely confluent, abdomen very finely and sparsely punctate. Pubescence of the fore parts indistinct, distinct and recumbent on the abdomen. Antennae reddish brown, club infuscate, maxillary palpi reddish yellow, segment 3 infuscate, legs reddish yellow, kees narrowly blackish, tarsi slightly infuscate. Clypeus and labrum dark- metallic, sparsely pubescent. Length: 4.0 - 4.5 mm (fore parts: 2.4 mm). Male: unknown. Female: Sternite 8 somewhat triangularly projecting apically (about as in fig. 198). Valvifer serrate apically, apical brush thin. Tergite 10 subtriangular. Head broader than elytra, frons moderately deeply impressed in anterior half, punctation moderately coarse and sparse, diameter of punctures about as large as basal cross section of antennal segment 3, interstices at least as large as diameters of punctures, mostly somewhat larger. Antennae slender, the last segment extends beyond the posterior margin of the pronotum when reflexed, penultimate segments nearly 1.5 x as long as broad. Shape of pronotum as in the related species; punctation coarse and dense, slightly confluent transversely on about middle, diameter of punctures about as large as apical cross section of antennal segment 2, interstices nearly half as wide as diameters of punctures. Elytra subquadrate, longer than broad, punctation very coarse 432 VOLKER PUTHZ and very dense (except on the declining side portions and a narrow area posterio(and) - laterally, where the punctures become extinct), forming long transverse rugae. Abdo- men as in the related species; punctation of paratergites very fine and extremely sparse, punctation of tergites very fine and sparse, interstices often twice as large as diameters of punctures. Legs very slender, metatarsi 3/4 as long as metatibiae, segment 1 dis- tinctly longer than segments 2-4 combined, much longer than segment 5. The whole body lacks groundsculpture. | Comments. Dianous cebuensis is easy to identify by the moderately coarse and sparse punctation of the frons. In my key to the species of group I it has to be placed at couplet # 26 (25): From D. tonkinensis (Puthz) it is distinguished by the much finer and sparser punctation of the frons, nearly extinct punctation of the outer border- areas of the elytra and by the much sparser abdominal punctation; from D. saxicola (Puthz) by the same characters. Dianous haraldi sp. n. Type materal. Holotype (3): N Laos: Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./ 13. VI. 1996, Harald Schillhammer (25) (NHMW). PM of HT: wH: 32; aE: 28; wP:25.5; IP: 26; WEI: 34.5; IEl: 37; IS: 30. This new species belongs also to group I of the genus, where it is the sister species of D. meo Rougemont, to which it is very similar so that a full description is not necessary. Moderately shining, black with dark purple reflection, frons moderately finely and very densely punctate, punctation of the pronotum coarse, very dense, strongly confluent, elytral punctation yet coarser, arranged into long rugae, punctation of the abdomen very fine and very dense. Pubescence of the fore parts indistinct, that of the abdomen dense and recumbent. Antennae black with some metallic reflection. Maxillary palpi with the first segment and bases of segments 2 and 3 reddish yellow, rest dark brown. About basal third of the femora reddish yellow, rest black with some metallic reflection, tibiae blackish brown, tarsi dark brown. Clypeus and labrum black with metallic reflection, sparsely pubescent. Length: 3.4 — 3.9 mm (fore parts: 2.1 mm). Male: Sternite 8 with a triangular notch in about posterior fifth. Sternite 9 with a small tooth apicolaterally, apicolateral brush moderately strong. Tergite 10 rounded. Aedeagus with a triangularly narrowed median lobe (fig. 240) which has a moderately strongly sclerotized expulsion mechanism as also has D. meo. Female: unknown. Head, antennae and pronotum almost similar to those of D. meo (punctation of the frons yet finer than that of the pronotum; on the pronotum the punctures are only well-defined on the actual middle, on the rest of surface punctures are confluent into long transverse rugae). Sculpture of elytra generally similar to that of D. meo, but coarser and stronger on anterior half, rugae nearly as broad as antennal segment 2. Abdomen similar to that of D. meo, finely and very densely punctate including the paratergites. Legs as in D. meo. THE GENUS DIANOUS IN CHINA 433 Comments. Dianous haraldi — which I warmly dedicate to my friend Harald Schillhammer, Vienna, who collected the holotype — has to be placed in my key (1981) at couplet # 63 (64) [spec. 6 Rougemont = D. meo Rougemont]. It is distinguished from D. meo by the reddish yellow bases of femora, by the coarser elytral sculpture and by the sexual characters (in D. meo the apical portion of the median lobe has a different outline (fig. 239) and the apical notch of sternite 8 is less deep); from D. tiomanensis Puthz it is distinguished by the coarser elytral sculpture, the less extended reddish yellow coloration of the bases of femora and by the sexual characters. Dianous arachnipes Puthz Dianous arachnipes Puthz, 1971a: 90f.; Shibata, 1979: 7 f. figs; Puthz, 1984a: 101; Naomi, 1997: 2 fig. Distribution. This species 1s widely distributed in Taiwan (see map in Shibata): Taipei Hsien: 3 9 9: Nanshi Str. (seapage area) 2 km S. Wulai, 22. III. 1991, H. G. Nelson (FMCh, cP). Ilan Hsien. Taoyüan Hsien. Taichung Hsien. 1 ©: Pahsienshan, 11. VII. 1932, Eaki (Kyushu University). Nantou Hsien. Chiayi Hsien. Kaohsiung Hsien. Hualien Hsien: 1 9; Urai, 1.X. 1921, Esaki (Kyushu University). Dianous latitarsis L. Benick Dianous latitarsis L. Benick, 1942: 42 ff. figs; Rougemont, 1983a: 11 f.; Zheng, 1993: 199. Dianous miripes Rougemont, 1985: 134 f. figs, syn. nov. Material examined: Holotype (4) from Burma; Sichuan: 1 d: Emei Shan, X. 1986, G. M. de Rougemont (cP), 21 more specimens from Emei Shan in cR. (in litt.). Zheng (1993) also records this species from Emei Shan. Comments. This species is easily to identify by its “pseudotetramerous” tarsi with segment 4 having a tube-like bristle shoe enveloping segment 5 except claws. This character distinguishes it easily from D. gracilipes Champion. Slight differences noted in the description of D. miripes belong to the range of variability, as confirmed by my friend de Rougemont (in litt.). Dianous versicolor Cameron Dianous versicolor Cemaron, 1915: 533 f.; 1930: 411 f. fig.; Puthz, 1981a: 114 f. fig.; Rouge- mont, 1985: 136 f.; Puthz, 1990: 123 Material examined: 1 d: Yunnan: 100 km W. Baoshan, Gaoligongshan Natural Reserve, 14.- 21. VI. 1993, E. Jendek & R.Sausa (NHMW); Sichuan: 18 exx. Emei Shan, X. 1986, G. M. de Rougemont (t. Rougemont, cR); 2 2 2: Emei Shan, 1530 m, 22. VI. 1994, torrent on slate with large crystalline rocks, H. Schillhammer (NHMW, cP). Comments. This species is very remarkable because of its asymmetrical, distinctly bilobed fourth tarsal segment with strong bristle shoe (figs 4-7). The female sternite 8 has only a minute medio apical excision (fig. 32). Dianous versicolor was previously known from India, Nepal, and Bhutan; new for China. Dianous chetri Rougemont Dianous chetri Rougemont, 1980: 172 ff. figs; 1985: 139 434 VOLKER PUTHZ Material examined: Yunnan: 1 2: Daju, Tiger George, 27°21’ N, 100°19’ E, 24.- 28. VI. 1992, E. Jendek (NHMW): 2 2 £ : 100 km W Baoshan, Gaoligongshan Natural Reserve, 14.- 21. VI. 1993, E. Jendek & O. Sause (NHMW). Sichuan: 1 d: Emei Shan, X. 1986, G. M. de Rougemont (cR). 1 d: NE India: Meghalaya, W Garo Hills, Nokrekek N. P., ca. 1100 m, 25°29.6° N, 90°19.5’ E, 9.-17. V. 1996, Jendek & Sausa (HNMW):; 1 2: W. Almora, Kumaon, H. G. Champion (cP): new for India. 1 d: N. Laos: Prov. Lg. Nam Tha, ca. 30 km NW Lg. Nam Tha, 800 m, 16./18. VI. 1996, Schillhammer (NHMW): new for Laos! PM of the male from Sichuan: wH: 59; adE: 39; IE: 20; IT: 15; wP: 45; IP: 43; Wir Sale lew 31541085 Male: In general similar to that of D. alternans Zheng. Metasternum broadly impressed medially, shining area broader (as broad as long), each side of it with about 10 moderately fine punctures. Impunctate, but microsculptured, middle area of sternites 3-6 broader, about half as broad as sternites, sternite 6 simple (not impressed) medially, with a few, very sparse, moderately fine punctures. Sternites 7 to 9 same as those of D. alternans. Aedeagus similar to that of the compared species, but median lobe relatively broader anteriorly, apex with five lateral setae, internal sac less strongly sclerotized and somewhat different in shape (fig. 46), parameres with 15 or 16 setae, two proximate setae widely separate from the following ones. Comments. The male specimen from Emei Shan, found together with D. alter- nans Zheng, differs in some apparently minor respects from typical specimens of D. chetri: Head and elytra at shoulders are bluish (as it is seen in specimens from N. India, Kumaon), and the punctation of pronotum is nearly obsolete anteriorly and posteriorly (much less distinct than in Nepalese specimens). The median lobe is somewhat broader anteriorly, but has also 5 apicolateral setae, parameres have fewer and evenly set setae (paratype from Sundarijal Cascades, Nepal, has 19/17 setae). Because the internal sac is identical with that of D. chetri, I believe that the differences fall within the variability range of the species. The Yunnan specimens resemble very much those from Nepal. New to China. Dianous alternans Zheng 1993 Dianous alternans Zheng, 1993: 36: 201 ff. figs Material examined: 4 d d, 3 2 2: Sichuan: Emei Shan, 7. X. 1985/ X. 1986, G. M. de Rougemont (cR, cP); 4 dd, 5 © ©: ibidem 1530 m, 22. VI. 1994, torrent on slate with large crystalline rocks, H. Schillhammer (NHMW, cP). This species is the sister species of D. chetri Rougemont; it also has ,,tarsal shoes“. Moderately shining, variegate metallic: head blue, pronotum bluish anteriorly, posteriorly, and laterally, brassy medially, or totally brassy; elytra (fig. 34) aeneous and/or with coppery tint on some areas, each with a large, remarkable bright bluish band extending from base through shoulders and then directed inwards; abdomen dark blue. Head, elytra, and abdomen finely and very densely punctate, pronotum with somewhat coarse punctation anteriorly, posteriorly, and laterally, but very finely and sparsely punctate in middle, punctation even becoming obsolete there. Antennae dark blue metallic, narrow bases of segments brownish, club more or less dark brown. Maxillary palpi dark, blue metallic, base of segments reddish. Legs dark blue metallic, THE GENUS DIANOUS IN CHINA 435 bases of trochanters reddish yellow. Clypeus and labrum bright bluish, densely set with long setae, lateral pubescence of clypeus particularly dense. Segment 4 of metatarsi simple, with a narrow bristle shoe. Length: 6.5 — 8.5 mm (fore parts : 3.7 — 4.2 mm). PM of one male from Emei Shan: wH: 63; adE: 41; IE: 23; IT: 15; wP: 45; IP: 45; wEl: 68; IEI: 76; IS: 65. Male: Metasternum broadly convex, slightly impressed posteriomedially, with a shining triangular area in middle slightly longer than broad, each side of this area sparsely set with about 10-16 moderately fine punctures, shining area densely, isodiametrically microsculptured, acam impunctate but shallowly microsculptured, sides of metasternum abruptly, very densely and finely punctate and pubescent. Sternites 3-5 with broad, impunctate shining area distinctly less broad than half the sternite (e. g. 22: 55 on sternite 4), medially, area finely microsculptured, Sternite 6 broadly impressed medially, sparsely set with moderately fine punctures, Sternite 7 with a broad and distinct medial impression more densely and somewhat more coarsely punctate than sides, densely set with yellowish pubescence posteriorly, posterior margin shallowly emarginate. Sternite 8 with a broad, obtusely-angled notch in about posterior tenth. Sternite 9 (fig. 50) with a broad and distinct tooth apicolaterally, sparsely set with numerous very short setae and few longer setae ventrally, apicolateral brush strong and dense, curved inwards distally, nearly as long as the entire sternite. Tergite 10 broadly rounded. Aedeagus (fig. 38) with a triangularly narrowed median lobe, its apical portion about twice as broad as parameres at broadest point, broadly rounded apically, with 2-3 distinct setae on each side. Internal sac strongly sclerotized (outline: fig. 45). Parameres distinctly longer than median lobe, with 12-16 strong setae. Female: Ventral characters resembling those of male, but sternite 6 simple and impunctate medially, sternite 7 very shallowly impressed medio-apically, apical margin very shallowly emarginate. Sternite 8 with a distinct medio-apical notch in triangularly produced medio-apical portion (figs 56- 58). Valvifers more or less rounded apically. Tergite 10 moderately broadly rounded, with a slight, concave median impression. Head as that of D. chetri, with shallow longitudinal furrows on anterior half of frons, punctation fine, even, very dense. Antennae slender, by about 3 outer segments extending beyond posterior margin of pronotum when reflexed, penultimate segments nearly twice as long as broad. Pronotum with deep impressions and lateral tubercles, slightly deeper and more elevated than those of D. chetri, posterior impressions coarsely and moderately densely punctate, rest of pronotum moderately finely to very finely punctate, punctation in middle very sparse, microsculpture dense, isodiametrical, variegate and deep (see below). Elytra with curved bands of whitish pubescence and an area of whitish pubescence medio-laterally; punctation about as fine as that on head, very dense on bluish bands, less dense on anterio-medial portion, whith interstices slightly larger than half diameter of punctures. Abdomen with broad, slightly upwards directed finely and very densely punctate paratergites, paratergites of tergite 4 about as broad as metatibia at apex, tergite 7 with a broad palisade fringe apically, punctation throughout fine and very dense, that of tergites 8 and 10 abruptly coarser and less 436 VOLKER PUTHZ dense. Legs slender, metatarsi 3/5 as long as metatibiae, first segment about as long as the fifth, distinctly longer than segments 2-4 combined, fourth segment simple with a narrow „shoe“ of long setae 3/5 length of claw segment. Pronotum more or less deeply reticulate, rest of body with more or less faint groundsculpture. Variability: One male paratype differs from the other paratypes by smaller size: fore parts: 3.5 mm, total length: 6.0- 7.0 mm, by the median lobe slightly narrower anteriorly and bearing only two setae at each side, and by the lower number of paramere setae (13/12). Specimens captured by Schillhammer have much deeper reticulation than the series captured by de Rougemont, especially on pronotum, also the pronotum is totally dull with less distinctly pierced punctation, while in the holotype the pronotum is distinctly shining because of much shallower reticulation. At the first these specimens seem to belong to a different species, but I can not find any additional differences, e. g. in the ventral punctation or in the genitalia, to support this concept. Comments. Dianous alternans Zheng differs from D. chetri Rougemont, which has been captured at the same place, by the sharply delimited blue elytral band (in the middle of the elytra of D. chetri are two more or less greenish blue areas that may be variably confluent), by more brilliant (less dull) pronotum, and by the sexual characters. In D. alternans head and shoulders are bluish, while in most specimens of D. chetri they are greenish-olivaceous, but there are also specimens of D. chetri with bluish head and bluish shoulders. Dianous coeruleovestitus Sp. n. Type Material. Holotype (dé): Hongkong: New Territories, Great Falls, Kadoorie Experimental Farm, CL 2209, 23. XI. 1985, J. T. & D. A. Polhemus (USNM); 8 88,5 9 2: Tai Mo Shan, 28. IV. 1996, G. M. de Rougemont (cR, cP). This new species resembles at first glance D. inaequalis Champion, it is the sister species of D. coeruleomicans Puthz. Moderately shining, green/blue metallic, head, broad band on elytra (fig. 33), and abdomen bluish, pronotum blackish blue, elytra on medial basal half and on posterior fifth greenish-olivaceous with some brassy tint. Head, elytra, and abdomen very finely and very densely punctate and pubescent, pronotum moderately finely to very finely and sparsely punctate, interstices nearly smooth, brilliant. Antennae dark blue-violet, club brownish. Palpi dark brown. Legs dark blue-violet, bases of tro- chanters reddish yellow, tarsi blackish brown. Clypeus and labrum blue, densely pubescent. Legs as those of compared species: segment 4 of metatarsi simple, with narrow bristle-shoes. Length: 6.0 — 6.2 mm (fore parts: 3.0 mm). PM of HT? wH: 44,5: adE: 30; IE: 17; IT: 9: wP: 34,5: IP: 35: WEI ileal Eley [SAS Male : Metasternum convex, shallowly impressed posteriomedially, with a long, triangular shining area in middle about 1/3 narrower than long, densely and shallowly microsculptured, and with about 10 moderately fine punctures on each half; sides of metasternum abruptly, very finely and very densely punctate and pubescent, acam almost impunctate. Sternites 3- 5 with an impunctate, shining median area about 1/3 as THE GENUS DIANOUS IN CHINA 437 broad as sternite and very shallowly microsculptured, sternite 6 medially somewhat more coarsely and less densely punctate than laterally, posterior margin very broadly, shallowly emarginate; sternite 7 with moderately deep, very finely and very densely punctate and pubescent impression posteriomedially, posterior margin moderately shallowly emarginate. Sternite 8 with a broad emargination in about posterior ninth. Sternite 9 with a broad, blunt tooth (fig. 41) apicolaterally, ventrally with sparse short setae, apicolateral brush strong and dense, about as long as whole sternite. Tergite 10 broadly rounded. Aedeagus (fig. 39) with median lobe triangularly narrowed, apex slightly broader than parameres at tip, with 4 moderately strong setae laterally, internal sac resembling that of D. chetri. Parameres longer than median lobe, with 11 or 12 setae apically (no setae at base). Female: Ventral characters similar to those of male. Sternite 8 with a shallow notch apicomedially (about as in fig. 76). Valvifer broadly rounded apically, with a strong brush, 1.5 x as long as the valvifer. Tergite 10 broadly rounded. Head and antennae in general as in D. chetri. Pronotum about as in D. alternans, but brilliant, groundsculpture almost obsolete, punctation laterally and posteriorly somewhat coarser, anteriorly as coarse as on head, becoming finer mediad, almost obsolete on middle; interstices in lateral impressions and posteriorly nearly as large as diameter of punctures, much wider medially. Elytra (fig. 33) in general as in D. alternans, but bluish areas larger and less distinctly separate from greenish areas (several violet-coppery transition zones present), surface less deeply impressed, punc- tation and pubescence very similar. Abdomen about as in D. alternans, but paratergites slightly narrower, those of tergite 4 about as broad as metatibia at middle. Comments. Dianous coeruleovestitus can be distinguished from both D. chetri Rougemont and D. alternans Zheng by the smaller size, by the coloration of the elytra, by the structures (degree of shininess) of the pronotum, and by the sexual characters, from D. coeruleomicans Puthz by different metallic reflection and the aedeagus. Etymology: coeruleus (Latin)= blue; vestitus (Latin) = clothed : Clothed with blue. Dianous coeruleotinctus sp. n. Type Material. Holotype (4) and allotype (9): NW- Hunan: Wulingyuan, N. Dayong Sioxiyu, 400 m, 31. X. 1993, large river in broad and shallow metal bed (NHMW). Paratypes: 4 d d,3 2 2: same data as holotype (NHMW, cP). This new species is allied to D. malayanus Cameron; it resembles several blue metallic species, e. g. D. acuminifer Puthz. Bright blue metallic, moderately shining, head, elytra, and abdomen finely, or moderately finely, and very densely punctate, pronotum finely to moderately coarsely and more or less sparsely punctate, shiny. Pubescence whitish yellow, dense, recum- bent, elytra with distinct tortuous bands of pubescence. Antennae, palpi, and legs blue metallic, antennal club brownish. Clypeus and labrum blue metallic, densely pubescent. Fourth tarsal segment simple with a sparse and short bristle shoe. Length: 5.2 — 6.5 mm (fore parts: 3.0 — 3.3 mm). PI\Miothil WEIS23 32 2dE2 IFJE17 17292 WRE3721P3753WEES6S31E1 265: IS: 54. 438 VOLKER PUTHZ Male: Metasternum broadly convex, triangular medial area flat, moderately finely, regularly, and very densely punctate, narrow interstices with fine groundsculp- ture, with medial, narrow, impunctate shining area about as broad as last antennal segment; acam finely and very densely punctate throughout; declining lateral areas of metasternum about as coarsely and densely punctate as medial area. Abdominal sternites 3-6 very densely and finely punctate throughout, sternite 7 posteriomedially with a shallow but distinct impression slightly more densely punctate and pubescent than sides, posterior margin shallowly emarginate. Sternite 8 with a broad emargination in about posterior fourteenth. Sternite 9 short and broad, posterior margin nearly blunt, finely crenulate (fig. 53), apicolateral brush very strong and dense, about 1.5 x as long as sternite. Tergite 10 very broadly rounded. Aedeagus (fig. 40) with median lobe triangularly narrowed, apically slightly broader than parameres at apex, apex rounded, internal structures resembling those of D. inaequalis Champion. Parameres slightly longer than median lobe, with about 12 evenly set long setae. Female: Ventral characters similar to those of male, but sternite 7 not impressed. Sternite 8 with two oval areas densely set with short setae, apically more or less deeply excavate (figs 52, 55). Valvifer rounded, apical brush very strong, 1.5 x as long as valvifer. Tergite 10 obtusely angulate apically. Head similar to that of D. acuminifer. Antennae slender, with at least last 3 segments extending beyond posterior margin of pronotum when reflexed, penultimate segments about 1.5 x as long as broad. Pronotum with deep lateral and oblique impressions, middle area appearing somewhat bumplike elevated, punctation variable, coarse and dense laterally and on basal impressions, with punctures possibly nearly as large as middle cross section of third antennal segment and interstices smaller than half diameters of punctures; distinctly less coarse and less dense on anterior portion of pronotum, with diameters of punctures about as large as one middle eye facet and interstices as large as diameters of punctures or larger; punctation in very middle even finer and sparser, but so irregular that a distinctly separate shining area can not be identified. Elytra subquadrate, sutural impression deep and long, humeral impressions shallow, punctation about as coarse as that on head but extremely dense, rough on posterior half. Abdomen broadly margined, paratergites distinctly directed upwards, those of segment 4 nearly as broad as metatibia at apex, punctation of paratergites fine and very dense, still denser than punctation of tergites which appears very dense throughout, except slightly sparser on middle portions of tergites; tergite 7 with a broad palisade fringe apically (the insect is fully winged); tergite 8 with punctation much coarser and less dense than that of tergite 7; tergite 10 with a few very fine punctures. Legs slender, metatarsi about 3/5 as long as metatibiae, first segment distinctly longer than following 3 segments combined (37: 33), somewhat longer than last segment (37: 35), fourth segment simple, bristle shoe thin. Interstices of frontal punctation smooth, pronotum densely but very shallowly reticulated (shiny !), abdomen with shallow ground sculpture. Comments. Dianous coeruleotinctus is distinguished from D. malayanus Cameron by its bright blue metallic lustre (D. malayanus Cameron has a more blue- violet shine), coarser pronotal punctation, coarser punctation of elytra, and the sexual THE GENUS DIANOUS IN CHINA 439 characters, from D. acuminifer Puthz and very similar species by much coarser punctate and shiny pronotum. Etymology: coeruleus (Latin) = bright blue, tinctus (Latin) = dipped/colored. Dianous inaequalis Champion Dianous inaequalis Champion, 1919: 45; Cameron, , 1930: 415 f. fig.; Puthz, 1971a: 91 fig.: Puthz, 1981a: 99, 122, fig. 42; Rougemont, 1983a: 2; Puthz, 1984a: 104 (see below) Dianous caeruleoguttatus Cameron, 1927: 8 Material examined : Yunnan: 1 4 : Kunming, 5. X. 1985, G. M. de Rougemont (CR): 1 9: Dali, X. 1986, G. M. de Rougemont (CR); 3 9 2: Lugu Lake — Luo Shui, 27°45’N, 100°45’E, 8.- 9. VII. 1992, E. Jendek (NHMW, cP); 1 d: ca. 100 km NW Lijiang, Hengduanshan, 2200 m, Jiduan — Weixi, 1. VII. 1994, H. Schillhammer (10) (NHMW). Sichuan: 1 ¢: Emei Shan, X. 1986, G. M. de Rougemont (cR). This is one of the most widely distributed species of group II. It has already been recorded from China (Sichuan: Puthz, 1971a). Male: Metasternum broadly convex, flat medially, middle very slightly impressed posteriorly, anterior process not separate from rest of metasternum by any kind of impression or transverse incision; sides finely and very densely punctate and shortly pubescent; middle with a broad, triangular shining area: moderately finely and sparsely punctate posteriorly, impunctate in very middle; triangular area about 1/3 as broad as entire width of metasternum posteriorly, its microsculpture deep, very dense, about isodiametrical. Ventral aspect of abdomen: fig. 116, Cameron, 1930. Sternites 3-5 medially with a moderately broad, impunctate, finely chagrined area, about 1/4 as broad as respective sternite, sternite 6 medially punctate, punctation sparser than that on sides (no distinctly differentiated impunctate area in middle). Rest of abdomen as in the Taiwanese subspecies (see below). Ae deagus (about as in fig. 69), median lobe slightly longer but also less long as parameres, parameres (mostly ?) with less than 20 long setae. Female: Metasternum and sternites 3- 5 about as in male. Sternite 6 medially with a narrow impunctate 1/8 or 1/9 of width of sternite, impunctate area distinctly narrower than that of sternite 5. Rest of abdomen as in Taiwanese subspecies, sternite 8 (figs 59, 60). Comments. The ventral characters in all specimens listed above are same as in the Indian specimens. The bluish areas of the elytra are slightly larger than those of the typical specimens, but are distinctly less large than in the Taiwanese subspecies. One female from Luo Shui has also bluish color on the humeral portion, but the blue color is not quite the same as that in the middle of elytra: some violet or brassy tint amongst the blue is apparent. At present, the Chinese material is not sufficient to allow the decision whether the slight differences between the Chinese and Indian/Nepalese specimens are clinal differences or belong to the general variability. Dianous inaequalis laevior ssp. n. Dianous inaequalis; Puthz, 1984a: 104 Type Material. Holotype (3) and allotype (2): Taiwan: Nantou Hsien: Shanlinchi, 1650 440 VOLKER PUTHZ m, 19. V. 1991, on bear moits rocks directly in a creek (see photo 2 in Smetana, 1995), A. Smetana (MHNG). Paratypes: 59 8 4,53 © 2: same data as HT (cP, cR, cS); 1 d: cataract nr Choshui Hast., rte. 151 0.7 km S. Hsitou (11 km SSE Luku) 6.1V.1991, H. G. Nelson (FMCh). Ilan Hsien: 1 4: Taipingshan, 1820 m, 15. VII. 1993, same habitat as HT, A. Smetana (cS). Kaohsiung Hsien: 5 4 4,2 2 2: Peinantashan trail, 2065 m, 6. VII. 1993, on wet vertical wall with running water, A. Smetana (cP, cS). Chiayi Hsien: 3 d d: Alishan, 2400 m, 3.-9. VII. 1972, T.C.Maa (Bishop Museum, cP). Hualien Hsien: 2 & 4,2 2 2: Taroko N. P., Nanhushi Hut, 2200 m, 8. V. 1990, on wet large rocks with thin layer of soaking fallen leaves in a creek, A. Smetana (cP, cS). This Dianous represents the Taiwanese subspecies of the variable Dianous inaequalis, which is one of the most wide-spread central-oriental Dianous. The Taiwa- nese specimens resemble very closely in nearly all respects the mainland nominate form, but show some differences in the coloration, and distinct differences in the ventral punctation. Length: 6.0 — 7.5 mm (for parts 3.4 — 3.8 mm. Male: Metasternum broadly convex, flat medially, posteriomedially very slightly impressed, anterior process separate from rest of metasternum by a more or less deep, narrow transverse incision; sides of metasternum finely and very densely punctate and shortly pubescent; middle of metasternum with a very broad, triangular, sharply delimited impunctate area (only a few fine punctures on sides of shining area), some- what broader than half of entire width of metasternum, with dense, moderately deep, sometimes slightly transverse microsculpture. Abdominal sternites 3-5 medially with a broad, finely chagreened impunctate shining area, at least 1/3 as broad as respective sternite; sternite 6 medially also with that remarkable impunctate area, 1/4 to 2/5 as broad as the sternite; sternite 7 broadly and shallowly impressed posteriomedially, impression very densely and moderately finely punctate, and with long, yellowish, brushlike pubescence, posterior margin broadly emarginate. Sternite 8 with a broad medio-apical notch in about posterior 10" to 11°. Sternite 9 (fig. 68), with a more or less blunt tooth latero-apically. Tergite 10 very broadly rounded at posterior margin. Aedeagus (fig. 69) with median lobe triangularly narrowed, internal sac densely set with spines; parameres extending well beyond apex of median lobe, slender, with many (20- 25) long setae. Female: Metasternum about same as in male, anterior process sometimes sepa- rate from rest of metasternum by a transverse impression. Abdominal sternites 3-6 medially with a broad, shining impunctate area, resembling that of male, sternite 6 about same sternite 5; sternite 7 densely and moderately finely punctate medially, more densely so near broadly and shallowly emarginate posterior margin; yellowish pubes- cence very dense posteriorly, long, brushlike convergent, distinctly extending beyond posterior margin of sternite; sternite 8 with a more or less deep apical notch (variability: figs 61-67). Valvifer rounded apically, apical brush 1.5 — 2 x as long as valvifer. Tergite 10 broadly, triangularly narrowed, rounded apically. Shining, head and abdomen blue to blue-black, sometimes with slight brassy or coppery reflex, pronotum dark-blue with variegate tints (olive, coppery, bronze), elytra golden- to brassy-metallic with a large and broad, curved blue-metallic band, and with some coppery tint in narrow transition zone between golden/brassy and blue-metallic THE GENUS DIANOUS IN CHINA 44] areas. Punctation of head and abdomen fine and very dense, except tergite 8 moderately coarsely and sparsely punctate; pronotum moderately coarsely and moderately densely (varying) punctate; upper golden/brassy areas of elytra very finely, more or less densely punctate (see below), less finely and extremely densely punctate laterally and on bluish areas. Pubescence short and dustlike-dense, on elytra brushed in different directions: a distinctly separate whitish area of pubescence laterally slightly before the middle, a narrow curved band of whitish pubescence on each elytron beginning behind shoulder (about 1/4 of elytral length from anterior margin) turning toward suture at about middle of elytron and then turning again backwards in sutural quarter, ending about 1/4 of elytral length before posterior margin of elytra. Antennae, palpi, and legs bluish-black, bases of trochanters brownish. Head and abdomen with shallow microsculpture, pro- notum with deep and dense, nearly isodiametrical microsculpture, elytra between punc- tures mainly smooth (see below). Legs slender; metatarsus 3/5 as long as metatibia, segment | somewhat longer than the following 3 segments combined, about as long as last segment, segment 4 simple with a distinct, but narrow bristle-shoe about 2/3 as long as claw-segment. Comments. Dianous inaequalis laevior is distinguished from D. inaequalis inaequalis by the elytral coloration. Bluish areas are distinctly larger: a broad and long area extends from shoulders somewhat behind middle, at its narrowest portion the transverse bluish band is mostly 1/3 — 1/4 (in the nominate form 1/6 — 1/5) as wide as the length of elytra. The elytra are bluish behind shoulders in all specimens; the brassy/golden anterior area of the elytra is distinctly separate from the lateral bluish portion in most specimens; the bluish portion behind shoulders may be 1/5 — 1/4 as wide as one elytron (in dorsal aspect); in a few specimens the lateral bluish portion may become only 1/10 as wide as one elytron, but the brassy/golden anterior coloration does not extend to the declining lateral portion behind shoulders, as it does in D. inaequalis inaequalis. The elytral punctation is also somewhat variable: interstices of the punc- tation on the basal brassy/golden area are about equal to diameters of punctures or slightly larger in most specimens, but the punctures in one female PT are slightly larger with interstices slightly larger than half the diameter of punctures (the microsculpture is also more distinct in this female). Etymology: comparative of Latin laevis = smooth; „more shining“. Dianous elegantulus Zheng 1993 Dianous elegantulus Zheng, 1993: 203 figs. Material examined: 3 dd, 3 9 £ Sichuan: Emei Shan, 7. X. 1985, G. M. de Rougemont (cR, cP); 1 2: Sichuan: Yachow, 2200 ft., 15. I. 1929, D. C. Graham (USNM). NW-Hunan: 9 dd, 12 9 9: Wulingyuan, N. Dayong Suoxiyu, 400 m, 31. X. 1993, large river in broad and shallow metal bed, H. Schillhammer (NHMW, cP); 1 3, 1 E 2: ca. 60 m W Ya’an, 1500 m and 1600 m, 4 km W Xingou Village, Ji & Wang, 12./13. VI.1996 (CWBS 232: stream, ca. 2-3 m wide, tributary to Tiangüan He, cold and fast flowing through secondary forest, shaded; 233: upstream of loc. 232, small stream, ca. 1- 2 m wide, cold and fast flowing through secondary forest, large stones, leaf packs) (NHMW); 1 6, 1 2- PT: Emei Shan, 710- 1120 m, 23.- 24. VII. 1983, Lu Ying & Gou Zheng-yue [HT and other PT in the Department of Biology, Sichuan Teacher’s College, Nanchong]. 442 VOLKER PUTHZ This Dianous is one of the most beautiful species of the genus. It resembles at first glance D. ponticus Fagel externally, but it is more closely related to D. inaequalis Champion and resembling species; its sister species is D. adonis sp. n. (see below). Moderately shining, variegate metallic: Head, abdomen, and legs blue, pro- notum dark blue with violet or greenish tint, elytra tricolorous (fig. 36): central, sharply delimited, cross bright blue (dull), outer quarters brilliant brassy-golden-coppery. Head, elytra, and abdomen finely and very densely punctate and pubescent, pronotum moderately coarsely and moderately densely to moderately sparsely punctate. Antennae and palpi blue metallic, antennal club brownish. Bases of trochanters brownish, tarsi blackish, becoming dark brown toward apex. Clypeus and labrum blue, densely pubescent. Tarsi simple, 4th segment with a distinct bristle- shoe. Length: 7.5 — 9.5 mm (fore parts: 4.3 — 4.7 mm). PM of a male from Emei Shan: wH: 68; adE: 44; IE: 21,5; IT: 17; wP: 49; IP: 46; wEl: 82; IEl: 90; IS: 77. Male: Metasternum convex with a shining, plain or slightly impressed triangular median area 2/3 as broad as long, distinctly but shallowly microsculptured, impunctate medially, with 10-12 moderately coarse punctures on each side; acam impunctate, with nearly obsolete groundsculpture; sides of metasternum abruptly, very finely and densely punctate and pubescent. Abdominal sternites 3-6 medially with impunctate (but microsculptured) area about 1/5 — 1/6 as broad as respective sternite, sternite 7 posterio- medially more or less (very shallowly) impressed and moderately finely, very densely punctate and pubescent, posterior margin very shallowly emarginate. Sternite 8 with a very broad, obtusely angulate excision in about posterior fourteenth. Sternite 9 (fig. 43) obtuse apically, irregularly crenulate, with sparse short setae, apicolateral brush strong and dense, about 1.5 x as long as the entire sternite. Aedeagus (fig. 44) with median lobe triangularly narrowed toward broadly rounded apical portion about twice as broad as parameres apically, with 4 small lateral setae; internal sac moderately strongly sclerotized, resembling that of related species. Parameres longer than median lobe, with about 18 moderately strong setae, a few of them situated in basal half. Female: Ventral characters about as those of male. Sternite 8 apically with a bifid projection with variable medial excision (figs 51, 54). Valvifer broadly rounded- serrate apically, with a very long and strong brush. Tergite 10 triangularly narrowed. Head about as in D. ponticus, with long lateral furrows, comparatively small eyes and strongly rounded temples (temple-suture can be seen from above). Antennae slender, about 3 outer segments extending beyond the posterior margin of pronotum when reflexed, penultimate segments 1.5 — 1.8 x as long as broad. Pronotum with deep oblique impressions in posterior half, a constriction near anterior margin, a lateral impression near base and two less deep impressions medially near posterior margin; punctation of basal impressions moderately coarse and dense, diameters of punctures about as large as cross section of one tarsal claw, interstices slightly less large than diameters of punctures; anterior punctation nearly as coarse as, medial and lateral punctation distinctly finer and sparser than that of basal impressions, finest punctures finer than those on frons, interstices at least as large as to 2-3 times larger than diameters of punctures (see also below). Elytra (fig. 36) with punctation on bright blue THE GENUS DIANOUS IN CHINA 443 dull central cross extremely dense, that of brassy-golden-coppery areas less dense, shining interstices no larger than diameters of punctures. Abdomen very broad with very broad, somewhat upward directed paratergites, paratergites of tergite 4 at least as broad as metatibia at apex. Legs slender, metatarsi 2/3 as long as metatibae, first segment slightly longer than the 3 following combined, about as long as the last segment. Head with nearly obsolete microsculpture, pronotum distinctly but not deeply, isodiametrically microsculptured, central portions of bluish area of elytra densely and deeply reticulated, outer quarters smooth, abdomen shallowly to very shallowly reti- culated. Variability: Punctation of the middle of pronotum may be nearly as coarse and dense as that on anterior portions. Comments. Dianous elegantulus can be distinguished at first glance from all other Dianous by its very beautiful coloration, from D. adonis by the stronger elytral punctation, different elytral color, the ventral and the sexual characters. Dianous adonis sp. n. Type material. Holotype (4), allotype (9) and 1 ?- paratype: Sichuan: Ganzi pref., Daxue shan, 5 km E Kangching, 102°00’E, 30°03’N, river valley, ca. 3000 m, 20.-23.V.1977, A. Piitz (ZMB, coll. Piitz, cP). This new species resembles strongly D. elegantulus Zheng and is its sister species. Moderatey shining, variegate metallic: Head, pronotum, abdomen and legs dark blue-metallic, elytra bicolorous (fig. 37), main surface including central area shining blue, smaller areas near base and on outer posterior quarter brilliant brassy-golden. Head finely and very densely punctate, diameters of punctures as large as medial eye facets, interstices mostly smaller than half diameters of punctures. Pronotum modera- tely finely and sparsely punctate (central area) to coarsely and densely punctate (near posterior margin). Elytral punctation slightly finer and slightly less dense than that on frons. Abdomen finely and densely punctate. Pubescence remarkable, except on pro- notum short and dense throughout, elytral pubescence variegate in color and arrange- ment: curved fields of golden shining setae-bands contrast with the darker main pubescence. Antennae and maxillary palpi dark blue metallic, antennal club brownish. Clypeus and labrum blue metallic, moderately densely pubescent. Tarsi simple, segment 4 with a distinct bristle shoe, first segment as long or slightly longer than the last segment. Length: 8.0 — 9.5 mm (fore parts: 4.2 — 4.4 mm). BMEORENESWELZ64 Stade: MAMIE 2 ETES NP 494122462 Wels aE: IS: 66. Male: Metasternum convex with a broad, plain or slightly impressed shining median area, which is shallowly reticulate with 8-10 fine punctures on each side; punctation of the sides of metasternum different, sharply delimited: finely and densely punctate and pubescent, acam impunctate. Sternites 3-6 with a broad, impunctate and finely reticulate shining median area (that of sternite 4 somewhat narrower than half width of sternite: 25:57), sides finely and densely punctate and pubescent. Sternite 7 444 VOLKER PUTHZ impunctate and densely reticulate medially, moderately finely to finely punctate and long yellowish pubescent posteromedially. Sternite 8 with a broad obtusely-angled moderately shallow apical emargination (length of sternite: depth of emargination = 100: 6). Sternite 9 as in D. elegantulus, apical brush strong, distinctly longer than the sternite. Tergite 10 broadly rounded. Ae deagus nearly as in D. elegantulus, the median lobe less broad anteriorly, parameres with 15-16 setae. Female: Ventral characters similar to the male, sternite 8 with a bifid projection as in D. elegantulus. Valvifer and tergite 10 as in the compared species. Variability: The anterior brassy area of elytra is occasionally narrower than in 0339, Comments. D. adonis is distinguished from D. elegantulus by the much finer and less dense punctation of the bluish elytral centre, by the differently colored elytra, by the narrower pronotum, by the broader shining median areas of sternites and by the sexual characters. Etymology: Adonis (Latin): in Greek mythology a very handsome young man loved by Aphrodite. Dianous schoenmanni sp. n. Type material. Holotype (4): Sichuan: Guan Xian Co., Balang Shan, 3 km W, stream, 0.5 — 1.0 m wide, fast flowing, cold, flowing throigh meadows, partly shaded by bushes, small crystalline gravel and sand, ca. 1950 m, |. VII. 1998, Schònmann, Ji, Wang (CWBS 343) (NHMW). This new species also resembles D. elegantulus Zheng and is closely related to it. Moderately shining, metallic blue with small brassy to coppery areas on elytra (fig. 35). Head finely and very densely punctate, diameters of punctures as large as medial eye facets, interstices mostly smaller than half diameter of punctures. Pronotum finely and moderately sparsely punctate anteriorly and posteriorly, very sparsely on central area. Elytral punctation nearly as fine as that on frons. Abdomen finely and densely punctate. Except on pronotum the pubescence is distinct, particularly conspi- cuous on the elytra, where a narrow transverse band can be seen medially, which after a short interruption seems to be continued on declining sides laterally, where a round area of shining pubescence exists; also on posterior angles the pubescence is more conspicuous than on the rest of the surface. Antennae, maxillary palpi and legs dark blue-metallic. Clypeus and labrum blue-metallic, moderately densely pubescent. Tarsi simple, segment 4 with a distinct bristle-shoe, three quarters of the length of the claw segment; segment | slightly longer than the following 3 segments combined (18: 16), slightly shorter than the last segment (18: 19). Pronotum and abdomen with shallow ground sculpture. Length: 6.5 — 7.5 mm (fore parts: 4,1 mm). PM of HT: wH: 61; adE: 42: JE: 21: IT: 18; wP: 43.5; JP: 43.52 mE 136851 BE 74; IS: 65. Male: Metasternum with a broad, triangular, flat shining median area which is shallowly reticulate and has some 4-6 moderately fine punctures laterally, acam THE GENUS DIANOUS IN CHINA 445 smooth. Sternites 3-6 with moderately broad impunctate median areas, nearly one third as broad as the respective sternites. Sternite 7 moderately sparsely and slightly more coarsely punctate medially than laterally, densely pubescent near posterior margin, which is very shallowly emarginate. Sternit 8 with a broad obtusely- angled emargi- nation in about posterior 12' (8: 94). Sternite 9 similar to that of D. elegantulus (see fig. 43) but more prominent apicolaterally (fig. 42), posterior margin smooth medially, apicolateral brush strong, distinctly longer than the sternite. Tergite 10 broadly rounded. Aedeagus nearly as in D. elegantulus (see fig. 44), apical portion of median lobe about as broad as in the compared species, parameres less long, slightly longer than the median lobe with 19 setae. Female: unknown. Comments. Dianous schoenmanni may be distinguished from D. elegantulus Zheng by the longer, much finer punctate pronotum, by the narrower elytra with much smaller brassy areas and by the median areas of sternites, which are much broader impunctate. It is distinguished from D. adonis sp. n. by the minor length, sparser medial punctation of the pronotum, less large brassy areas on elytra, narrower impunctate median areas of sternites and broader apical portion of the median lobe. Etymology: This new species is named in honour of Dr. H. Schönmann, Vienna, who collected the new insect. Dianous punctiventris Champion Dianous punctiventris Champion, 1919: 48; Cameron, 1930: 420 fig.; Puthz, 1981a: 123 fig. Dianous puberulus L.Benick, 1942: 39 ff. figs, syn. nov.; Puthz, 1981a: 123 fig. Material examined: India: 1 d, 1 9: W. Almora, Champion (types; BMNH); 1 d: Chakrata district: Bangar, 6000 feet, 1. V. 1921, Cameron (cP); 1 4,2 9 2: Dehra Dun, 1940-45, Liesenfeldt (FMCh, cP); BURMA: 1 4,6 9 9: Washaung, 200 m, 16. III. 1934, R.Malaise (syntypes of D. puberulus; FMCh, Museum Stockholm, cP); 12 d d, 12 2 2: Shingbwiang, 10. III. 1914, L. C. Kuitert (Snow Ent.Mus., Kansas; cP). Laos: 1 3: Prov. Lg Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12.- 13. VI. 1996, Schillhammer (25) (NHMW). Vietnam: 2 85,2 22: Bao Lac (coll.Fauvel, IrScNB); 18, 4 9 9: Cao Bang, 1. XI. 1893, Dr. Billet (coll.Fauvel; IrScNB, cP). China: Yunnan: 1 à, 1 2: Kingtung, 1200 m, 10. III. 1957, Panfilov (Museum Leningrad, cP); 4 EE: Ruli, 4. II. 1993, G. M. de Rougemont (cR, cP). Hainan: 2 2 2: 3 km NE Maoyang; Wuzhi Shan River slowly flowing through wide, deforested valley, only few riffles, banks with sand, grass and mud, 100 m, 19. I. 1996, Ji & Wang (198) (NHMW, cP); 1 2:2 km S Mao’an, 250 m, 20. I. 1996, Jäch (199) (NHMW). Comments. For a long time I doubted the status of both taxa. After having studied more than 50 specimens from various localities, amongst them several males with everted aedeagi, I am convinced that both taxa are conspecific. As I noted (1980a: 238), the aedeagus-figure published by Jarrige 1950 for D. cameronianus Jarrige (taken from an unknown specimen, not from the HT) shows the everted aedeagus of D. puncti- ventris. In D. punctiventris (normal state) the median lobe is narrowly, triangularly narrowed (fig. 44, Puthz, 1981a). As known from related species (e. g. figs 61-66), in females the shape of the posterior portion of sternite 8 is variable, it can be more or less deeply excised. 446 VOLKER PUTHZ Dianous fauveli sp. n. Type material. Holotype (¢): Vietnam: Cao Bang, Dr. Billet (IRScNB). PM of HT: wH: 55; adE: 37; IE: 19; IT: 10: WP 45,5: IP? SAVE Eom leas. 19263. This new species is closely allied and similar to D. yunnanensis but differs markedly in some respects. It was found amongst the historical material in the Fauvel- collection. Blackish blue metallic, rather dull, head moderately finely and densely punctate, pronotum slightly more finely and densely punctate (also medially), punctation of elytra about as coarse as that on head, but very dense, extremely densely and rough on posterior half, abdomen finely and very densely punctate. Pubescence whitish yellow, dense, recumbent, elytra with tortuous bands of pubescence. Antennae, palpi, and legs black with blue metallic tint. Clypeus and labrum blue metallic, densely pubescent. Fourth tarsal segment simple, with sparse and short bristle shoe. Length: 7.0 — 8.5 mm (fore parts: 4.0 mm). Male: Trochanters with very small, inconspicuous, blunt tooth. Metasternum broadly convex, medial area very broad, nearly as broad as long, narrowly and shallowly impressed medially, with linear impunctate area, punctation of rest of median area moderately fine (less regular in size of punctures than in D. uniformis) and very dense, interstices distinctly reticulate; acam finely and densely punctate throughout, not only medially. Abdominal sternites 3-6 finely and densely punctate throughout; sternite 7 with a shallow posteriomedian impression densely punctate and set with long pubescence, posterior margin distinctly but shallowly emarginate. Sternite 8 with a shallow round emargination in about posterior thirteenth. Sternite 9 broadly rounded and crenulated apically, very shallowly emarginate, apicolateral brush strong and dense, slightly longer than sternite. Tergite 10 broadly rounded. Ae deagus relatively short and broad, much shorter that those of D. acuminifer and of D. yunnanensis, apical portion (fig. 49) triangularly narrowed, apically as broad as parameres at apex, internal structures (everted in HT) resembling those of D. yunnanensis: a long and broad, strongly sclerotized tube and some weak membranes set with indistinct elements. Parameres slightly longer than median lobe, with about 20- 22 lang setae on anterior half. Female: unknown. Comments. Dianous fauveli resembles in nearly all respects D. yunnanensis, but differs by the distinctly different sexual characters. The holotype is an old, somewhat damaged specimen (antennae and legs partly missing). Etymology: Dedicated to the great staphylinidologist Albert Fauvel (1840-1921) in which’s collection this specimen was found as „Dianous billeti sp. n.“. Dianous acuminifer Puthz Dianous acuminifer Puthz, 1984b: 104 fig. Material examined. Taiwan: Nantou Hsien: 2 & g, 1 2: Shanlinchi 1650 m, 16. V. 1990, on wet large rocks in a creek (see photo 2 in Smetana, 1995), A. Smetana (cS, cP); 11 dd, 9 THE GENUS DIANOUS IN CHINA 447 2 2: Sun Lin Sea, rte. 151, 16 km SSE Luku, 6. IV. 1991, H. G. Nelson (FMCh, cP); 1 2: tr. Chen Yu Lau, 2 km above Lona, 37 km S Shuili (6.3 km ex jet. rte. 21), 7. IV. 1991, H. G. Nelson (FMCh). Taitung Hsien: Tzepeng, 1.-II. 1964, T. C. Maa (HT; Bishop Museum). Hualien Hsien: 1 © : Taroko National Park, Chungyantienshi (River) Waterfall, 2300 m, 10. V. 1990, wet moss and debris in a spray zone of a waterfall, A. Smetana (cS). 1 2: Nantou Hsien: Sun Lin Sea, rte. 151, 16 km SSE Luku, 6. IV. 1991, H. G. Nelson (FMCh) (cf. det.). PM of a male from Shanlinchi: wH: 56; adE: 39; IE: 18; IT: 11; wP: 44,5; IP: 45; wie ples El 7531S: 65: Male: Trochanters simple. Medial area of metasternum flat, narrowly impressed and impunctate posteriomedially, remainder of medial area finely and very densely punctate, interstices much narrower than half diameters of punctures; with shallow microsculpture; acam with a slightly finer and slightly less dense punctation, punctation of sides of metasternum about as coarse as, but slightly less dense and shallower, than that on medial area. Sternites 3-6 finely and densely punctate throughout, interstices smooth, sternite middle indistinctly impressed posteriomedially, more densely punctate and pubescent medially, posterior margin distinctly emarginate. Sternite 8 with a broad emargination in about posterior sixth. Sternite 9 broadly rounded and crenulated apically (fig. 78), apicolateral brush strong and dense, distinctly longer than the sternite. Tergite 10 as in fig. 72, less triangularly pointed than in female (see fig. 71). Aedeagus long with median lobe triangularly narrowed (fig. 74), apical portion at least as broad as parameres anteriorly, internal sac consisting of a long and broad, strongly sclerotized tube and of membranes with strong spines basally and less strong spines apically. Parameres somewhat longer than median-lobe, with about 30 evenly set setae. Comments. The male is here described for the first time. Dianous acuminifer is in external characters quite similar to several related species and may be positively identified only by its sexual characters. The metallic color is variable in specimens from Sun Lin Sea: some areas are bright bluish, others brassy to dark-green, no specimen is totally bluish. One female of this series differs from the other 9 by shape of sternite 8 (fig. 80) and by weak brush of valvifer, but the punctation of tergite 8 and the metallic tint fit well the other specimens, so this single female is regarded as a variation of D. acuminifer. Dianous angulifer sp. n. Type Material. Holotype (2): Taiwan: Chiayi Hsien: Alishan, 2180 m, Sister Ponds, forest litter, 26. IV. 1990, L. LeSage (LL 24 a) (MHNG). This new species is the sister species of D. acuminifer Puthz which it resembles in nearly all respects, but it differs mainly by the distinctly different shape of the terminal abdominal sterna. Dark blue metallic, rather dull, entire body moderately finely and very densely punctate and densely set with golden, recumbent pubescence, elytra with remarkable tortuous bands of pubescence. Antennae, palpi, and legs black with blue metallic tint, antennal club and tarsi blackish brown. Clypeus and labrum bright blue metallic, densely pubescent. Segment 4 of metatarsi simple, with a short bristle shoe. 448 VOLKER PUTHZ Length: 6.5 — 7.5 mm (fore parts: 3.7 mm). PM of Hl: wHe 54: adE: 37; IE: 17; VE? 10; wR: 42> IP: AS WEINMOMERNTES [S302 Male: Unknown. Female: Ventral characters closely resembling those of D. acuminifer (see above), but punctation of sternite 8 somewhat coarser and distinctly denser, interstices distinctly less large than diameter of punctures, posterior margin shallowly emarginate (fig. 76). Valvifer rounded apically, apical brush much weaker than that of D. acumi- nifer, 2/3 as long as valvifer. Tergite 10 (fig. 73) triangularly narrowed, apex rounded. In most other respects quite similar to D. acuminifer, but interstices between punctation of head smooth, interstices of pronotal punctation nearly smooth, broad abdominal paratergites directed slightly more (distinctly) upwards, the punctation of tergite 8 about as coarse as that on elytra and dense, interstices distinctly smaller than half diameters of punctures (in all D. acuminifer except one specimen punctation of tergite 8 is distinctly less coarse and less dense, interstices as large as half diameters of punctures or larger). Comments. It may be argued that the described characters fit into the variation range of D. acuminifer. But I do not think so, mainly because of the entirely different apical brush of the sternum 9 (valvifer). This character distinguishes D. angulifer positively also from D. uniformis Zheng and D. fauveli sp. n. 1 9: Nantou Hsien: Sun Lin Sea, rte. 151, 16 km SSE Luku, 6. IV. 1991, H. G. Nelson (FMCh) is with some doubt regarded as belonging to D. angulifer (see Comments under D. acuminifer). Dianous angulifer is also very near to D. yunnanensis Puthz but differs from it by the lack of spine at metatrochanters. Etymology: angulus (Latin) = angle; Dianous who has an (apical) angle. Dianous uniformis Zheng Dianous uniformis Zheng, 1993: 203 f. figs. Material examined: 8 d d, 2 © 9: Sichuan: Emei Shan, 7.X.1985, G. M. de Rougemont (CR, cP); 2 6d, 2 2 2-PT: Emei Shan, 23-24. VII.1983, Gou Zheng-yue & Lue Ying (cP). [HT and other PT gd d, 2 9 9-PT: Emei Shan, 23-24.VII.1983, Gou Zeng-yue & Lu Ying (cP). [HT and other PT deposited in the Dept. Biol., Sichuan Teachers College] This species is very similar, and closely related, to D. acuminifer and D. angulifer. Blackish blue metallic (sometimes with olivaceus shine), rather dull, head moderately finely and densely punctate, pronotum (except for narrow medial area) slightly more finely and densely punctate, elytra somewhat more coarsely and very densely punctate (punctation less dense on anterior half, extremely dense, nearly confluent posteriomedially), abdomen finely and densely punctate, somewhat less den- sely so medially. Pubescence whitish yellow, very distinct, recumbent, especially remarkable on elytra with tortuous bands of pubescence. Antennae, palpi, and legs dark blue metallic. Clypeus and labrum blue metallic, densely pubescent. Fourth metatarsal segment simple, with moderately sparse bristle shoe. Length: 7.5 — 9.5 mm (fore parts: 4.3 mm). THE GENUS DIANOUS IN CHINA 449 PM of one male: wH: 61; adE: 41; IE: 21; IT: 11; wP: 46; IP: 47,5; wEl: 79; IEl: 85:18:73. Male: Trochanters simple. Medial area of metasternum flat, slightly impressed posteriomedially with narrow, impunctate shining area medially about as broad as second segment of maxillary palpi, lateral areas finely and densely punctate, interstices nearly half as large as half diameters of punctures, shallowly microsculptured, acam more finely and less densely punctate. Punctation on middle of sternites 3-6 about as fine and dense as that on lateral portions, sternite 7 simple, very finely and very densely punctate and pubescent posteriomedially, posterior margin shallowly emarginate. Sternite 8 with a broad and shallow emargination in about posterior tenth. Sternite 9 rounded and crenulated apically (fig. 75), apicolateral brush strong and dense, slightly longer than sternite. Tergite 10 very broadly rounded at posterior margin. Ae deagus in general as that of D. acuminifer, but apical portion of median lobe (fig. 70) narrower, narrower than parameres at apex, internal sac consisting of broad, strongly sclerotized tube and numerous membranes densely set with short, strong spines. Parameres slightly longer than median lobe, with about 22 long and strong setae on apical quarter. Female: Ventral characters as those of male, but sternite 7 without apical emargination. Sternite 8 with distinct excision apically (fig. 79). Valvifer rounded api- cally, with strong brush about 1/3 longer than valvifer. Tergite 10 as that of D. angulifer (see fig. 73). Punctation and pubescence very similar to that of D. acuminifer, but punctation of fore parts not as dense, interstices of frontal punctation repeatedly as large as diameters of punctures, punctation of pronotum less crowded, punctation of anterior half of elytra slightly less dense, interstices often half as large as half diameters of punctures, rarely larger, punctation near posterior margin of elytra less rough. Comments. Dianous uniformis may be easily distinguished from D. acuminifer and D. angulifer by its blackish blue metallic habitus, by the slightly less dense punctation, and by the sexual characters. It differs from D. yunnanensis Puthz by the broader head, slightly longer pronotum, larger size, and the sexual characters (the aedeagus of D. yunnanensis is similar externally, but has a totally different internal sac with a very broad internal tube narrowed posteriorly, and only very weakly sclerotized membranes set with indistinct toothlike elements; also parameres have strong setae at least on anterior half). Dianous luteoguttatus Champion Dianous luteoguttatus Champion, 1919: 98 f.; Puthz, 1980b: 6, fig.; Rougemont, 1987a: 49 Material examined: Taiwan: Nantou Hsien: | 2: Shanlinchi, 1650 m, 19. V. 1991, on wet rocks in a creek (see photo 2 in Smetana, 1995), A. Smetana (cS); 1 9: ibidem, 10. V. 1990. fauchage dans forét primaire, L. LeSage (LL 90-4) (coll. LeSage). Comments. This species was until now known from N. India and Nepal: n e w to China. The elytral spot is somewhat larger in the Taiwanese females than in the Nepalese male (from Induwa Khola Valley), with diameter nearly as large as the length of the first segment of metatarsus, while in the Nepalese male it is only as large as the 450 VOLKER PUTHZ length of the eighth antennal segment. In the Taiwanese females the sternites 3-6 are more sparsely punctate than in the male described below, with the interstices of the medial punctation distinctly larger (up to twice as large) than diameters of punctures. The male sexual characters have not been described yet, therefore their description based on a male from Nepal follows: Male: Metasternum convex, median area plain, with a narrow, medial, line-like incised longitudinal impression, lateral punctation rather fine and moderately dense, diameters of punctures smaller than basal cross section of third antennal segment, interstices shallowly microsculptured, larger than diameters of punctures but less large than two punctures combined, acam very finely and densely punctate and pubescent; declining lateral areas of metasternum finely and densely punctate, interstices shallowly reticulated. Medial punctation of sternites 3-6 slightly coarser and somewhat sparser than lateral punctation, interstices mostly nearly as large as diameters of punctures; sternite 7 much more densely punctate and pubescent medially than laterally, posterior margin slightly emarginate; sternite 8 with a broad emargination in about posterior eighth; sternite 9 with a distinct tooth apicolaterally (fig. 90), some sclerotized fringes and asacl apicomedially, apicolateral brush moderately strong, 3/4 as long as sternite. Tergite 10 broadly rounded. Aedeagus (fig. 87) with long, anteriorly narrow, lanceolate median lobe, apical portion about twice as broad as parameres at apex; internal sac consisting of a broad, strongly sclerotized tubal piece and several mem- branes set with differently shaped spines. Parameres slender, with 11 or 12 long setae only on anterior half. Female: Ventrally similar to those of male, but sternite 8 broadly rounded apically (no projection). Valvifer (fig. 1, Puthz, 1980b), apical brush somewhat shorter than the valvifer. Tergite 10 rounded apically. Dianous rougemontianus sp. n. Dianous vietnamensis; Zheng, 1994: 479 Type material. Holotype (male) and allotype (female): Sichuan: Emei Shan, X. 1986, G. M. de Rougemont (cR in BMNH). Paratypes: 4 d 4,8 9 © same data as holotype (cR, cP); 1 9: ibidem 600-1050 m, 5.-19. V. 1989, L. Bocäk (NHMB); 2 G d: ibidem 1530 m, 22. VI. 1994, H. Schillhammer (4a) (NHMW): 2 G d: Da Bai Shan, IX. 1980, Zhou Ping (cP, cZheng). The specimens recorded as ,.D. viernamensis“ by Zheng should be regarded as belonging to this new species. This new species is closely allied to D. viernamensis Puthz and has been confounded with him. It also resembles very much D. ruginosus Zheng. Blackish green metallic, with olivaceous and/or brassy tint, moderately shining, head moderately coarsely and densely punctate, pronotum coarsely and very densely, slightly confluently punctate, elytral sculpture consisting of mostly long rugae, directed on anterior half from suture posteriolaterad to about middle and from there posterio- mediad on posterior half; abdomen very finely and very densely punctate. Pubescence of head and abdomen distinct, short, recumbent. Antennae, palpi, and legs black with metallic tint, three outer segments lighter, tarsi + brownish. Clypeus and labrum greenish-blue, moderately sparsely pubescent. Fourth tarsal segment simple, lacking bristle shoe. THE GENUS DIANOUS IN CHINA 45] Length: 4.3 — 5.8 mm (fore parts: 2.7 — 2.9 mm). PM of the HT: wH: 42; adE: 25.5; IE: 18; IT: 5; wP: 33.5; IP: 34.5; wEl: 48; lEl: 51; IS: 44. Male: Metasternum broadly and shallowly impressed, deeply and densely reticulate, but not as dull as e. g. that of D. aereus, impunctate medially, lateral areas of triangular medial area with about 18-20 moderately coarse punctures as large as basal cross section of third palpal segment, acam very finely and sparsely punctate, interstices with very shallow groundsculpture, shiny. Sternites 3-5 very sparsely, finely punctate medially, but without distinct separate smooth area; sternite 6 with a distinct and broad posteriomedial impression, medial punctation about as dense as that on sides, posterior margin broadly emarginate; sternite 7 with a distinct medial impression, sides of impression distinctly elevated and toothlike prominent posteriorly, distinctly extending beyond semicircularly emarginate posterior margin of sternite; sternite 8 with a deep notch in posterior third (length of sternite: depth of notch = 83: 28); sternite 9 (fig. 99), with sclerotized fringes and asacl apicomedially, apicolateral brush strong but distinctly shorter than sternite. Tergite 10 broadly rounded. Aedeagus (fig. 83), anterior portion of median lobe with long and distinct ventrolateral ridges (missing in D. vietna- mensis, but present in D. karen and in D. aereus), strongly and densely pubescent; parameres with 18 very strong and long equally set amS and with additional dorsal series of about 12 shorter setae. Female: Ventral characters similar to those of male, but sternites 6 and 7 not impressed; sternite 8 (fig. 97). Valvifer subtriangular apically, apical brush shorter than valvifer. Tergite 10 triangular. In most respects as D. ruginosus Zheng, but larger, punctation of head some- what coarser, median portion of frons less strongly elevated, antennae longer; extending by 3.5 segments (in D. ruginosus by 3 segments) beyond posterior margin of the pronotum when reflexed, pronotum less broad, slightly coarser but less confluently punctate, elytra narrower (longer than broad), punctures anterio-medially mostly well- defined (in D. ruginosus mostly confluent). Metatarsi about 4/5 as long as metatibiae, segment | slightly shorter than three following segments combined, distinctly longer than last segment. Entire body distinctly, but shallowly reticulated. Comments. Dianous rougemontianus differs from D. vietnamensis Puthz and D. karen Rougemont by the greenish-olivaceous tint of the fore parts (blackish-blue in the compared species), by the broader head, longer antennae and longer elytra, by the coarser but somewhat less rugose punctation of the fore parts, and by the the sexual characters. It differs from D. aereus Champion by the less elevated medial portion of the frons, by the coarser punctation of the head, the longer antennae, and by the sexual characters. Etymology: This new species is warmly dedicated to my dear friend Guillaume de Rougemont (London), who made substantial contributions to the knowledge of the genus Dianous and who provided his Chinese Dianous for this study. 452 VOLKER PUTHZ Dianous vietnamensis Puthz Dianous vietnamensis Puthz, 1980a: 234 f. fig. Material examined: 2 d 5: Hainan: 30 km E Maoyang, above Wuzhi Shan Resort, stream ca. 5 m wide, densely shaded, flowing partly through primary forest, 700-800 m, 18. I. 1996, Jäch (194) (NHMW): 1 © (cf. det.): Xishuangbanna, 20. I. 1993, G. M. de Rougemont (cR). 1 d, 1 9: N Laos: 28 dd, 36 9 9, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12./13. VI. 1996, Schillhammer (25) (NHMW, cP); 4 d d, 7 2 2, Prov. Lg. Nam Tha, ca. 30 km NW Lg. Nam Tha, 800 m, 16./18. VI. 1996, Schillhammer (28, 30) (NHMW, cP): 1 6, Bolikhamsai Prov., Ban Nape-Kaew-Nua-Pass, 600 m, 18°22.3° N, 105° 9.1 E, 18. IV. - 1. V. 1998, Jendek & Sausa (cHromädka). Described from Vietnam: new to China and Laos. The identification of the Yunnan-female has to be assured by the respective male. Dianous karen Rougemont spec. propr. Dianous karen Rougemont, 1981a: 357 ff. figs.: 1983a: 12: 1986: 265 f. The description of D. rougemontianus made it necessary to revise closely related taxa. The improvement of our knowledge of the genus has considerably increased in the last years, allowing the usage of characters that were formerly neglected, misvalued, or not really understood, e. g. arrangement and number of the setae on parameres, sculp- ture of the metasternum, shape of tergite 10, etc. Using this knowledge, I believe now that D. karen, which has been synonymized by its author with D. vietnamensis in 1986, is a good species. It is distinguished from D. vietnamensis by the number and arran- gement of the parameral setae, by the sculpture of the metasternum, and by the gene- rally less strongly pronounced ventral characters of the terminal sternites in the male. To make the comparison of the two species possible, these are the PM of a Thailand male: wH: 37; adE: 20; IE: 15; IT: 3; wP: 29; IP: 28.5; wEl: 45; IEl: 46.5; IS: 40. Length of body: 3.7 — 4.5 mm (fore parts: 2.3 — 2.4 mm). Male (from Doi Suthep, 1000 m, 14. I. 1990, H. Malicky, cP): Medial area of metasternum broadly flattened somewhat impressed posteriomedially, entire medial area deeply and very densely reticulated, dull with faint shine, reticulation becoming slightly less dense toward middle coxae, but there is no distinct bipartition in sculpture as in D. vietnamensis (metasternum shallowly reticulate, shiny and moderately coarsely and moderately sparsely punctate anterio-medially, but extremely densely and deeply reticulate and quite dull, with punctation indistinct posteriomedially), punctation of medial area moderately fine to moderately coarse and moderately sparse, acam finely and moderately densely punctate, interstices distinctly reticulated but shiny. Sternites 3- 6 medially slightly more coarsely and somewhat more sparsely punctate than laterally; sternite 7 distinctly impressed posteriomedially, posterior sides of impression carinate, tooth-like extending beyond posterior margin of sternite, which is distinctly emarginate between teeth (impression and teeth somewhat stronger in D. vietnamensis). Sternite 8 with a deep notch in about posterior quarter; sternite 9 (fig. 96), with fine sclerotized fringes and asacl apicomedially, apicolateral brush strong, somewhat shorter than sternite. Tergite 10 broadly rounded. Ae deagus (fig. 82); proximal half of each THE GENUS DIANOUS IN CHINA 453 paramere without setae, distal half with 9 (published fig.ure of the holotype) — 11 (Doi Suthep) strong setae, dorsal series of each paramere setae with 7 setae in my Doi Suthep specimen (holotype: ?, not figured; aedeagus lost: Rougemont, 1981). The species lives also in Laos: 1 d Laos c.: Bolikhamsai Prov., Ban-Nape-Kaow-Nua-Pass, 18. IV.- 1. V. 1998, Jendek & Sousa (cHromädka): new to Laos. Comments. Since the material is still sparse, the variability ranges of D. vier- namensis and D. karen cannot be positively assessed; the number of setae is probably variable in a limited space, I think it is reasonable to consider the differences in the arrangement of parameral setae as a specific character. Dianous brevicornis sp. n. Type material. Holotype (female) and 1 ?-paratype: Yunnan: 50 km N. Lijiang, Yulongshan Natural Reserve, (NHMW, c P). Very similar to D. aereus Cameron, and different only as follows. Greenish-aeneous metallic, rather shiny, head moderately finely and densely punctate, pronotum moderately coarsely, irregularly and confluently punctate, elytral sculpture consisting of long rugae directed on anterior half from suture posteriomediad, on posterior half from middle backwards toward the suture; abdominal punctation very fine and very dense. Antennae, palpi, and legs black with greenish metallic tint, anten- nal club and tarsi indistinctly paler. Clypeus and labrum greenish, sparsely pubescent. Pubescence of pronotum and elytra indistinct, that of head and abdomen distinct, short, dense, recumbent. Fourth segment of metatarsus simple, with a few longer setae, lacking bristle shoe. Length: 3.3 — 4.1 mm (fore parts: 2.2 mm). BMEOCETIWER 32 Sade O11E: 145: EA WP TS AP DT os WERADZIEN: AD Sal S735. Male: Unknown. Female: Metasternum strongly convex, medial area very broadly flat, shallowly impressed and impunctate posteriomedially, lateral portions of medial area each with 18-20 moderately fine punctures, interstices densely but moderately shallowly reti- culate, strongly shiny (thus distinctly different from D. aereus Cameron and D. ruginosus Zheng), acam nearly impunctate, ground sculpture even shallower than that of middle of metasternum. Sternites 3-6 finely and very sparsely punctate medially, interstices at least twice as large as diameters of punctures; sternite 7 more densely punctate and pubescent medially than on sides, not impressed, posterior margin very slightly emarginate. Sternite 8 about as in fig. 97. Valvifer obliquely truncate apically, apical brush thin, nearly half as long as valvifer. Tergite 10 moderately triangularly pointed. Antennae distinctly shorter than those of D. aereus, extending beyond posterior margin of pronotum by 1-5 segments when reflexed, outer segments nearly as broad as long. Punctation on middle of abdominal tergites somewhat less dense, interstices on middle of tergite 6 distinctly larger than very fine punctures (in D. aereus less large than diameters of less fine punctures). 454 VOLKER PUTHZ Comments. Dianous brevicornis differs from D. aereus Cameron by the dis- tinctly shorter antennae and by the ventral characters described above, particularly by the the median area of metasternum, which, in D. aereus, is completely dull because of the extremely dense and deep reticulation. Etymology: brevicornis (Lat.) = with short antennae. Dianous ruginosus Zheng Dianous ruginosus Zheng, 1993: 204 figs. Material examined: 1 ¢: Sichuan: Emei Mt., 500 m, 4.-20. V. 1989 (NHMB); 2 9 2: Mt. Emei, 9 2-PT: Emei Shan, 1500 m, 30. VII. 1982, Gao Xin-fen (cP) [HT and more PT in the Dept.Biol., Sichuan Teachers College. | This species also closely resembles D. aereus Cameron. Blackish-green metallic with olivaceous tint, moderately shining, head mode- rately finely and densely punctate, pronotum moderately coarsely and very densely, confluently punctate, elytral sculpture consisting mostly of long rugae, directed on anterior half from suture posteriolaterad to about middle and from there posteriomediad on posterior half; abdomen very finely and very densely punctate. Pubescence of head and abdomen distinct, short, recumbent. Antennae, palpi, and legs black with metallic tint, three outer antennal segments paler, sometimes light brown, tarsi more or less brownish. Clypeus and labrum blackish with metallic tint, moderately sparsely pubescent. Fourth segment of metatarsi simple, lacking bristle shoe. Length: 3.8 — 4.7 mm (fore parts: 2.3 — 2.4 mm). PM of one male: wH: 38; adE: 24; IE: 17; IT: 4; wP: 30.5; IP: 31; wEl: 44; IEl: 44; IS: 38. Male: Medial area of metasternum broadly flat, densely and deeply reticulate, with a faint shine (not quite dull as in D. aereus), middle shallowly impressed, punctation about equally distributed, moderately coarse, diameters of punctures about as large as basal cross section of third palpal segment, each half of medial area with about 20 punctures, acam with a few finer, shallow punctures, interstices with shall- ower reticulation than that on medial area of metasternum, declining lateral areas of metasternum finely and densely punctate, interstices with very shallow ground- sculpture, shiny. Medial punctation of sternites 3-6 as fine as, but slightly sparser, than that on sides, interstices larger than diameters of punctures, but less large than two punctures combined; sternite 6 with a broad and very shallow emargination posteriorly; sternite 7 with a distinct impression posteriomedially, sides of impression elevate and carinated posteriorly, somewhat extending beyond posterior margin of sternite, puncta- tion of impression denser than lateral punctation, posterior margin distinctly emar- ginate. Sternite 8 with a broad and deep emargination in about posterior quarter; sternite 9 acute apicolaterally (fig. 100), with short sclerotized fringes apicomedially, apico- lateral brush yellowish brown, weak and moderately short, about two thirds as long as sternite. Tergite 10 broadly rounded. Ae deagus as in fig. 81, median lobe broadly rounded anteriorly, with long setae ventrally, internal sac with broad tube, and with strongly spinose membranes proximally, and finely crenulate membranes distally. THE GENUS DIANOUS IN CHINA 455 Parameres much longer than median lobe, each with about 20 evenly set moderately strong setae. Female: Ventral characters similar to those of male; sternite 7 slightly more densely punctate and pubescent posteriomedially than laterally, no impression, no apical emargination. Sternite 8 as in fig. 98. Valvifer acute apically, apical brush short, indistinct, distinctly shorter than valvifer. Tergite 10 triangularly narrowed. In most respects as D. aereus Cameron, but differs by the last 3 antennal segments lighter than the rest of antennae, and by the ventral characters: in D. aereus the median area of the metasternum is completely dull, the apical brush of the sternum 9 is brownish and longer, and the aedeagus is clearly different Comments. I originally considered Dianous ruginosus Zheng to represent a subspecies of the Himalayan D. aereus Cameron. — One female is a more robust specimen and mainly black (PM: wH: 39; adE: 25; IE: 18; IT: 4; wP: 32.5; IP: 33; wEI: 44; JEl: 44; IS: 36), sternites 3-6 are somewhat more sparsely punctate, interstices medially two times as large as diameters of punctures Dianous rugipennis sp. n. Type material. Holotype (2): NW- Hunan: Wulingyuan, N. Dayong Sioxiyu, 400 m, 31. X. 1993 (NHMW). This new species resembles both D. championi Cameron and D. cupreogutta sp. n., it is the sister species of D. rugosipennis Sp. n. Black with dark blue metallic tint, shining, fore parts coarsely and densely punctate, sculpture of almost entire elytra strongly rugose, forming long confluent rugae. Abdomen finely and moderately sparsely punctate. Pubescence of fore parts moderately distinct, abdomen with rather dense and recumbent pubescence. Antennae black, club dark brown. Maxillary palpi with first segment brown, segments 2 and 3 dark brown. Legs black, trochanters and tarsi blackish brown. Clypeus and labrum black with faint bluish tint, sparsely pubescent. Segment 4 of metatarsi simple. Length: 4.3 — 5.3 mm (fore parts: 2.6 mm). Poni wh "42 ade 3 IE 2 I 17875: we: 2971 32. wels43 971 EI 348318: 38. Male: Unknown. Female: Midline of middle portion of metasternum narrowly impunctate, sides moderately densely set with coarse punctures, acam with some very fine punctures. Abdominal sternites 3-6 finely and moderately sparsely punctate medially, sternite 7 finely and densely punctate and pubescent posteriomedially, posterior margin simple. Sternite 8 (fig. 93). Valvifer (fig. 92) apically acute, apical brush moderately distinct, less than half as long as valvifer. Tergite 10 rounded. Head distinctly but slightly narrower than elytra, longitudinal furrows deep and very long, median portion of frons about as broad as and as strongly elevated as each of the side portions; punctation coarse and dense, diameters of punctures slightly less than basal cross section of third antennal segment, interstices mostly narrower than half diameter of punctures, but becoming as large as, to somewhat larger on a small area 456 VOLKER PUTHZ posteriorly near medial eye margins (not on middle of frons). Antennae moderately slender, when reflexed, extending only by last segment beyond posterior margin of pronotum, outer segments nearly 1.5 x as long as broad. Pronotum distinctly constricted anteriorly and posteriorly, impressions about same as in D. cupreogutta, thus less deep than in D. championi; punctation coarse and dense, rarely slightly confluent, diameters of punctures as large as, or slightly larger, than basal cross section of third antennal segment, interstices smaller than half diameter of punctures, except posteriomedially, on small impunctate area about as broad as two and as long as three punctures combined. Elytra subquadrate, sutural and humeral impressions distinct; punctation coarser than that on pronotum, forming long oblique rugae directed laterad in anterior half, mediad in posterior half; some vorticose sculpture present on posterior half but no distinct rosette and no traces of coppery or any other similar spot. Abdomen relatively convex, paratergites narrower than those of compared species, appreciably declining (sides of sternites visible in dorsal aspect), paratergites of segment 4 about as broad as metatibia at base, punctation of paratergites very fine, dense anteriorly, becoming sparser posteriad; punctation of tergites 3-7 very fine and moderately sparse, diameters of punctures slightly less than one inner eye facet, interstices at least 1.5 x as wide as diameters of punctures, somewhat larger on middle of tergites. Legs moderately slender, metatarsi about 5/8 as long as metatibiae; first segment slightly shorter than 3 following segments combined (25: 27), distinctly shorter than last segment (25: 30), segment 4 simple. No microsculpture, except tergite 7 faintly reticulated, tergite 8 with shallow microsculpture, tergite 10 nearly smooth. Comments. Dianous rugipennis is distinguished from D. cupreogutta by its unicolorous elytra, the denser punctation of the head and pronotum, by the more strongly rugose sculpture of the anterior portion of the elytra, by the narrower, decli- ning paratergites, and by the simple tarsi. It differs from the very similar D. championi (specimens of which are not fully winged) by the denser punctation of the head and the pronotum, by the shorter antennae and legs, the not acute rugae on elytra (rugae are narrowly rounded above), the narrower, declining paratergites, the golden (not silver) pubescence, and by the simple fourth tarsal segment. From D. rugosipennis sp. n. it 1s distinguished by smaller elytra, less coarse sculpture, longer antennae and smooth interstices of the punctation of tergite 3-6 (see above). Etymology: ruga (Latin) = fold or ruga, -pennis (Latin) = on elytra: Dianous with rugae on the elytra. Dianous rugosipennis sp. n. Type material. Holotype (d ): Fujian: Chong’an Wuyi Shan, 1 km W Wuyi Gong Village (= Shangian, ca. 10 km S Chong’an City), residual pools in dry riverbed in steep valley, crystalline rock, 250 m, 15./ 18. I. 1997, Ji & Wang (CWBS 240) (NHMW). Paratypes: 1 d, 4 2 ©: Anhui: Huang Shan 60 km NNW Huang Shan City [=Tunxi], stream (beneath cable car), ca. 3- 5 m wide, almost completely dried out, surrounded by primary forest, very big granitic rocks and sand, short sections with running water (20-30 cm wide), small residual pools with decaying leaves, near Tang Kou, 900-1000 m [figs 17, 20 in Jäch & Li, 1998], 31. X. 1997, Schönmann (CWBS 292) (NHMW, cP). THE GENUS DIANOUS IN CHINA 457 This new species resembles D. radiatus Champion in general habitus, it is very similar to its sister species D. rugipennis sp. n.. Moderately shining, black with more or less distinct dark-greenish metallic tint. Head coarsely and densely punctate, pronotum very coarsely and slightly confluently punctate, elytra very coarsely punctate, sculpture confluent, abdomen finely and mode- rately densely punctate. Pubescence of fore parts less distinct, that of abdomen distinct and recumbent. Antennae, maxillary palpi, and legs blackish with faint metallic tint, last antennal segments slightly infuscate. Clypeus and labrum dark-metallic, sparsely pubescent. Segment 4 of metatarsi simple. Length: 4.8 — 5.8 mm (fore parts: 2.8- 3.0 mm). BNKoORHIESWwER AA ade: 2631ER 211: SVP M3 SAP: SSVELSO0SMERZG: IS: 46. Male: Metasternum broadly convex, median area strongly shining with very faint microsculpture, punctation moderately fine and sparse, diameters of punctures smaller than basal cross section of second segment of maxillary palpi, interstices twice and more as wide as diameters of punctures, posterior middle narrowly impunctate, acam almost smooth, with a few very fine punctures. Medial punctation of sternites fine and sparse. Sternite 7 shallowly impressed and densely punctate and pubescent postero- medially, posterior margin shallowly emarginate. Sternite 8 with a semicircular emargi- nation in about posterior eighth. Sternite 9 acute apicolaterally (fig. 95), apical brush about half as long as the sternite, with a small area set with asacl apicomedially. Tergite 10 rounded. Aedeagus (fig. 84, 86) with slender median lobe triangularly pointed apically, apical portion densely set with moderately long setae ventrally, internal sac strongly sclerotized. Parameres slender with 15-16/ 24-25 equally distributed setae. Female: Sternite 8 (fig. 94), stronger triangular apically than in D. rugipennis. Valvifer about as in D. rugipennis, apical brush stronger, nearly as long as stronger sclerotized portion of the valvifer. Tergite 10 rounded. Head distinctly narrower than elytra, longitudinal furrows of frons deep and very long, median portion nearly as broad as each of the side portions, flat anterior, distinctly elevated posterior; punctation coarse and dense, diameters of punctures variable, large punctures about as large as basal cross section of third antennal segment, interstices mostly smaller than half diameters of punctures. Antennae short, when reflexed exten- ding to about posterior margin of the pronotum, penultimate segments about one third longer than broad. Pronotum with the sides strongly convex in anterior half, strongly concave posteriorly; a deep transverse impression near anterior margin, deep lateral impressions in about middle, two distinctly elevated portions posteriorly behind the lateral impressions; punctation very coarse and dense, distinctly rugose near posterior margin, diameters of punctures about as large as medial cross section of third antennal segment, interstices mostly smaller than half diameters of punctures, some exceptions posteromedially, where interstices can become as large as punctures. Elytra subqua- drate, posterior margin deeply emarginate, sutural impression distinct in anterior half, a shallow impression in lateral middle where internal rugae end in a point; sculpture very coarse and dense with long rugae, which are directed posterolaterad in anterior half of elytra, anterolaterad in posterior half; rugae about as wide as first segment of middle 458 VOLKER PUTHZ tarsi. Abdomen moderately convex, paratergites moderately broad, in sagittal level, those of segment 4 about as braod as metatibiae at broadest point of its anterior third, punctation of paratergites fine and very dense anteriorly, distinctly sparser posteriorly; punctation of tergites 3-7 fine and moderately dense, interstices about as large or slightly larger than diameters of punctures; punctation of tergites 8 and 10 fine and sparse, distinctly sparser than that of tergite 7. Legs slender, metatarsi two thirds as long as metatibiae, first segment slightly shorter than 3 following segments combined (30: 32), about as long as segment 5, segment 4 simple. Fore body with very shallow traces of microsculpture, microsculpture of abdomen slightly more distinct (for identi- fication in a key this species should run amongst those which are lacking micro- sculpture). Variability: The specimens from Anhui are black with faint greenish tint; the aedeagus is almost identical to that of the HT, but parameres have 24-25 setae. Comments. Dianous rugosipennis is distinguished from D. radiatus Champion by coarser and denser punctation of head and pronotum, by the shorter antennae, and by the darker greenish shine, from D. championi Cameron also by coarser and denser punctation of head and pronotum and shorter antennae, and by the distinct greenish shine. From both species by the aedeagus, especially by the equal distribution of paramere setae. From the very similar D. rugipennis it is distinguished by the much larger elytra (WH: wEl in D. rugipennis = 0.97, in D. rugosipennis = 0.84- 0.90), coarser sculpture of the fore parts, shorter antennae and faint but distinct microsculpture on abdomen. Etymology: rugosipennis (Latin) = having rugosely sculptured elytra. Dianous atrocyaneus sp. n. Type material. Holotype (4 ): Taiwan: Taoyuan Hsien: Takuanshan Forest, 1600 m, 17. IV. 1990, on wet rocks in a creek, A. Smetana (cS). This new species belongs to the group of D. chinensis Bernhauer and is the sister species of D. yoshidai Naomi from Iriomote Island near Taiwan. In general habitus (proportions and sculpture), it closely resembles D. banghaasi Bernhauer. Moderately shining, fore parts completely black dorsally, entire abdomen and fore parts bright blue metallic ventrally. Punctation of fore parts coarse and dense, slightly confluent on some areas of elytra, abdomen finely and densely punctate. Pubescence of fore parts whitish, long, erect, especially conspicuous at sides of elytra, abdomen with dense, moderately recumbent golden pubescence. Antennae black, club brownish. Palpi dark brown. Legs black with dark blue metallic tint, trochanters dark brown. Clypeus and labrum dark blue metallic with long and moderately dense pubescence. Fourth segment of metatarsi slightly but distinctly bilobed. Length: 5.2 — 6.2 mm (fore parts: 3.0 mm). PM of HT: wH: 41,5: adE: 25: IE: 18; lle: 6: wP: 34.5: IP? 36:52 wEIED» Sale e 55; IS: 49. Male: Metasternum broadly convex, flat medially, narrow medial area (as wide as 4-5 punctures combined) impunctate, punctation coarse and rather dense, becoming THE GENUS DIANOUS IN CHINA 459 much finer and sparser toward declining lateral portions of metasternum, acam almost smooth. Sternites 3-5 medially finely and somewhat sparser punctate than on the sides; sternite 6 very shallowly impressed in posterior middle, punctation as on the sides, sternite 7 with long, distinct medial impression finely and densely punctate posteriorly and very densely pubescent at posterior border, posterior margin shallowly emarginate. Sternite 8 with a deep, broad, rounded emargination in about posterior quarter. Sternite 9 with a distinct prominent tooth apicolaterally (fig. 116), apicolateral brush about 2/3 as long as sternite. Tergite 10 broadly rounded. Ae deagus (fig. 113) of the „chinensis-type“, parameres with 8 + 10/12 setae, apex of median lobe with about 26 short setae ventrolaterally. Female: Unknown. Head similar to that of D. banghaasi, punctation of frons coarse and dense, diameters of punctures slightly less than cross section of third antennal segment, interstices mostly smaller than half diameter of punctures, somewhat larger, medially and posteriolaterally, rarely as large as or larger than diameters of punctures. Antennae slender, similar to those of D. banghaasi. Pronotum similar to that of D. banghaasi, punctation coarser than on head, diameters of punctures may become as large as middle cross section of third antennal segment, punctation transversely confluent near posterior margin. Elytra very large, subquadrate, sutural impression long and distinct, humeral impression shallow; punctation as coarse as or somewhat coarser than in D. banghaasi, somewhat confluent in middle of each elytron, diameters of largest punctures about as large as cross section of second antennal segment, interstices distinctly smaller than half diameter of punctures, distinctly larger posteriomedially on each elytron. Abdomen similar to that of D. banghaasi, with very broad paratergites, and fine and dense punctation. Legs slender, metatarsi at least 2/3 as long as metatibiae, first segment much shorter than the 3 following segments combined (36: 47), about as long as last segment, segment 4 shortly bilobed (most distinct in anterior tarsi). Elytra with very faint traces of microsculpture, interstices of abdominal punctation, except on tergite 8, nearly smooth. Comments. Dianous atrocyaneus may be distinguished from all closely related species by the coloration. It differs from D. yoshidai Naomi (which has a dark coppery dot in the middle of the elytra just behind middle, a character that was not indicated in the original diagnosis but was taken from the holotype) by its larger size and by larger elytra. The hitherto undescribed male sexual characters of D. yoshidai are very similar to those of D. atrocyaneus, but sternite 8 has a deeper apical notch (in about posterior third), the median lobe has some more small setae ventro-laterally (27-30), parameres with less setae (7- 8). Etymology: ater (Latin) = black, cyaneus (Latin) = bright blue. Refers to the coloration. Dianous gemmosus sp. n. Type material. Holotype (6) and allotype (9): Yunnan: 100 km W Kunming, Diaolin Natural Reserve, 22. V.-2. VI. 1993 (NHMW, cP). Paratypes: 7 dd, 5 99: Yunnan: Yipinglang, 1800 m, 25°05’N, 101°53’E, 8.-10.VI.1993, Bolm (NHMB, cP); 1 2: ibidem 2000 m, 25°05’N, 101°55’E, V. Kuban (NHMB). 460 VOLKER PUTHZ This new species belongs to the chinensis-complex and is the sister species of D. hammondi Rougemont which it resembles closely. It is distinguished from him e. g. by a narrow triangular golden shining area in posterior middle of elytra. Rather shining, black with bluish and some aeneous tint, elytra each with conspicuous golden-coppery area on posterior half near suture. Head moderately coar- sely and densely punctate, pronotum coarsely and mostly densely punctate, somewhat rugose near anterior margin and in posterior impressions, elytra each coarsely sculptu- red, with long, mostly oblique rugae, which form a + regular oval rosette on posterior two thirds (figs 27, 28), abdomen, except tergites 8 and 10, very finely and densely punctate. Pubescence of fore parts moderately distinct, that of abdomen distinct and recumbent. Antennae blackish with slight metallic tint, base of individual segments narrowly brown, club dark brown. First segment of maxillary palpi brown, second and third segments blackish brown, bases of segments lighter. Legs black with some metal- lic tint, trochanters brownish, tarsi blackish brown. Clypeus and labrum blue metallic, moderately densely pubescent. Segment 4 of metatarsi distinctly bilobed (fig. 114). Length: 5.3 — 6.5 mm (fore parts: 3.0 — 3.2 mm). PM of HT: wH:45; adE: 28; IE: 19: 11: 8: wP: 33,5; IP737;5; WE 532121393: ISO Male: Metasternum broadly convex, broad triangular median area dull, extre- mely densely reticulated, medially impunctate, laterally moderately sparsely set with numerous moderately fine punctures, declining lateral areas of metasternum finely and densely punctate and pubescent, interstices smooth and shiny, acam finely and densely punctate and pubescent. Punctation of sternites 3-5 in middle slightly less fine and somewhat sparser than that on sides, sternite 6 with distinct impression in posterior half, punctation about same as that on previous sternites, posterior margin shallowly emarginate in similar way as that of sternite 5; sternite 7 with a long and narrow median impression, posterior half of impression bordered laterally by sharp keel toothlike prominent posteriorly, anterior half of impression very finely and very sparsely punc- tate, punctation of posterior half moderately fine and very dense, posterior margin broadly emarginate. Sternite 8 with a round emargination in about posterior fifth. Sternite 9 serrate apically (fig. 117), apicolateral brush strong and long, as long as sternite. Tergite 10 broadly rounded. Aedeagus (fig. 123) with median lobe broad, deeply, about V-like emarginate apically, apical portion with about 30 short setae ventro- laterally. Parameres with two groups of setae: 9 or 10 long and strong ventral setae at apex, and about 7 shorter setae dorsally in anterior half. Female: Ventral characters resembling those of male, but medial area of meta- sternum very shallowly reticulate and moderately shiny, lateral portions of medial area moderately coarsely and moderately densely punctate; sternite 6 simple, sternite 7 with a very shallow impression posteriomedially, moderately finely and very densely punc- tate, posterior margin very shallowly emarginate. Sternite 8 (fig. 115) triangularly rounded apically. Valvifer acute apically, apical brush longer than entire valvifer. Tergite 10 rounded, densely set with short and strong setae apically. THE GENUS DIANOUS IN CHINA 461 Head in general similar to that of the previous species; diameters of punctures on frons about as large as basal cross section of third palpal segment (thus distinctly smaller than basal cross section of third antennal segment), interstices mostly smaller than half diameters of punctures, distinctly larger posteriolaterally and on elevated medial portion, where they may exceed diameters of neighbouring punctures. Antennae very long, at least extending by last four segments well beyond posterior margin of the pronotum when reflexed, penultimate segments about twice as long as broad. Pronotum in general as that of the previous species; punctation distinctly coarser than on head, diameters of punctures about as large as basal cross section of third antennal segment, interstices mostly slightly smaller than half diameters of punctures laterally, larger centrally, middle with distinct impunctate area about as long as fourth antennal segment and as broad as length of tenth antennal segment; punctation on anterior transverse impression slightly confluent, punctation of the posterior impressions (near posterior margin) distinctly confluent. Elytra subquadrate, sutural and humeral impressions distinct; in anterior third with punctation about as coarse as that on middle of pronotum, but punctures obliquely extended; posterior two thirds with mainly long oblique rugae directed from middle of lateral margin posteriomediad toward suture and forming an obliquely-oval, more or less distinct rosette on lateral half (fig. 28); despite deep rugae and very narrow interstices surface of elytra distinctly shiny, not as dull as e. g. in D. frater Cameron. Abdomen with broad, slightly upwards directed paratergites, those of segment 4 about as broad as middle cross section of metatiba, punctation of paratergites very fine and very dense throughout; tergite 7 with a broad palisade fringe apically (the species is fully winged); punctation very fine and dense, diameters of punctures smaller than medial eye facets, interstices about as large as (laterally) or somewhat larger (medially) than punctures; punctation of tergite 8 coarser but also dense; tergite 10 finely and sparsely punctate. Legs slender, metatarsi 2/3 as long as the metatibiae, first segment distinctly shorter than following 3 segments combined (43: 54), distinctly longer than last segment (43: 34), segment 4 distinctly, moderately deeply bilobed (fig. 114). Fore parts lack microsculpture, abdomen with traces of very shallow groundsculpture, tergite 8 with shallow reticulation, tergite 10 rather smooth. Comments. Dianous gemmosus is distinguished from D. hammondi Rougemont by the less regular punctation of the frons, by the coarser punctation of pronotum which has no median longitudinal furrow, by the golden elytral area, the less dense abdominal punctation, and by the sexual characters (in D. hammondi, the anterior emargination of the median lobe is broadly rounded and there are only about 20 short ventral setae on each side of the apex of the median lobe). Etymology: gemma (Latin)= precious stone, jewel; Dianous set with precious stone or jewel. Dianous chinensis Bernhauer Dianous chinensis Bernhauer, 1916: 28; Puthz, 1971a: 89 figs. Material examined: Shandong: 4 d d, 4 9 ©: Kiautschau [Chiao-chou Wan] (types) (FMCh, ZMB, cP). Jiangxi: 1 © „Kiangsi“ without further indication (MM). Hunan: 7 6 6, 1 9: Wulingyuan, N. Dayong Sioxiyu, 400 m, 30. X. 1993, H. Schillhammer (5) (NHMW, cP). 462 VOLKER PUTHZ Shaanxi: 1 d: „Tsinling [Ching-ling Shan], Hsin-nganfu-Peiho (= Si-an-fu, Wie-ho, see Filchner, 1925: 69 ff.; = Xi'an) (ZMB). Comments. This species seems to be rare. It may be easily recognized by the dense and very regular punctation and by the distinct reticulation throughout. Elytra: fig. 102. Sternite 9 of male (figs 118, 119). Aedeagus (figs 121, 122), ratio of breadth of median lobe at bottom of apical emargination to depth of apical emargination = 1.9- 2.3; parameres with 7-9 alS and 8-13 amS, apical lobes of median lobe with 29-34 short setae. This species is also very close to D. aequalis Zheng but larger, reticulated, the median lobe less deeply emarginate, in the female apicomedial projection of sternite 8 is less narrow and less long. In the NHMW there are two females from Fujian, which differ from typical D. chinensis by dark blue color (lacking greenish tint), one of them has a very large elytral spot (larger than the length of antennal segment 3), the other a distinctly smaller elytral spot. Since I have no males from the respective localities these two specimens should be regarded as ,,chinensis abw.“ 1 9: Fujian: Longyan 2 km S Dachi, 750 m, Ceyan Shan, stream ca. 1-2 m wide, rocky steps with waterfalls, pools with crystalline sand, coarse granitic gravel, in the upper section surrounded by broadleaf forest, 29. I. 1997, Ji & Wang (CWBS 262); 1 2: Longyan Xiachi (20 km W Longyan), Meihua Shan, 650 m, mountain stream, 3-5 m wide, flowing through densely forested (broadleaf trees) steep valley descending from Meihua Shan, small waterfalls (figs 14, 16 in Jach & Li, 1998), 30. I. 1997, Ji & Wang (CWBS 263). Dianous yangae sp. n. Dianous chinensis; Zheng, 1993: 198 Type material. Holotype (male) and allotype (female): Sichuan: Emei Shan, 7. X. 1985, G. M. de Rougemont (cR in BMNH). Paratypes: (4) © 2, (4) 9 9: same data as HT (cR, cP); 2 dd: Suifu, 1.-10. XI. 1929, D. C. Graham (USNM); 1 d: Sichuan: Quingfeng Shan NW Chengdu, 600 m, Flussufer, 30.55 N, 103.30 E, 4. VI. 1997, M. Schülke (cSchiilke); 25 à, 2 2 2: Emei Shan, 630-710 m, 23. VII. 1983 („chinensis“ det. Zheng)(cP). This new species is the sister species of D. chinensis, from which it may be distinguished at first glance by the much smaller elytral spots. Moderately shining, Prussian-blue metallic, elytra each with a moderately large orange spot (fig. 103). Head and pronotum moderately coarsely and densely punctate, elytral punctation coarse and dense, regular, abdomen finely and very densely punctate. Pubescence remarkably dense, whitish yellow, long and semierect. Antennae, maxillary palpi, and legs blackish blue. Clypeus and labrum blue, densely pubescent. Fourth tarsal segment distinctly, moderately deeply bilobed. Length: 6.7-7.5 mm (fore parts: 3.3 — 3.6 mm). PM of HT: wH: 45; adE: 29; IE: 18; ITe: 6; wP: 37.5; IP: 40; wEl: 64; lEl: 64; ISSSSÌ Male: Medial area of metasternum flattened, slightly impressed posterio- medially, punctation moderately coarse and very dense, diameters of punctures nearly as large as basal cross section of third segment of maxiallary palpi, interstices mostly smaller than half diameters of punctures, densely reticulated; narrowly impunctate THE GENUS DIANOUS IN CHINA 463 posteriomedially; acam finely and densely punctate, interstices nearly smooth; decli- ning lateral areas of metasternum about as coarsely and densely punctate as medial area. Sternites 3-6 moderately finely and very densely punctate throughout; sternite 7 with a broad and shallow posterio-medial impression more densely punctate and pubescent than sides, posterior margin broadly emarginate. Sternite 8 with a broad and deep notch in nearly posterior quarter. Sternite 9 acute apicolaterally (fig. 120), with dense sclerotized fringes and asacl apicomedially, apicolateral brush strong, nearly as long as sternite. Tergite 10 blunt or very shallowly emarginate apically. Aedeagus (figs 124, 125) similar to that of D. banghaasi, ratio of breadth of median lobe at bottom of apical emargination : depth of apical emargination = 1.5 — 1.9; parameres with 11 alS and 12-14 amS; apical lobes of median lobe with 22- 26 ventral setae. Female: Ventral characters similar to those of male, but sternite 7 not impressed, very finely and densely punctate and pubescent posteriomedially. Sternite 8 (fig. 153). Valvifer acute, apical brush about as long as valvifer. Tergite 10 distinctly, but moderately triangular. Punctation generally remarkably regular, that of head and pronotum equally coarse and dense, diameters of punctures about as large as basal cross section of second segment of maxillary palpi, distinctly smaller than basal cross section of third antennal segment, interstices mostly smaller than half diameters of punctures, larger posteriorly near inner magins of eyes and on midline of pronotum (but no distinctly separate impunctate areas). Antennae long, extending beyond posterior margin of pronotum by about 4 segments when reflexed, outer segments nearly 1.5 x as long as broad. Pronotum with distinct, but shallow oblique lateral impressions on posterior half. Elytra coarsely, very densely, regularly punctate, all individual punctures well-defined, dia- meters of punctures as large as basal cross section of third antennal segment, interstices distinctly smaller than half diameters of punctures, shallowly reticulated, punctation becoming somewhat finer and sparser anteriorly and posteriorly. Abdomen with very broad, upwards directed, finely and very densely punctate paratergites; punctation of tergites fine and very dense, diameters of punctures nearly as large as medial eye facets, interstices mostly as large as half diameters of punctures, middle of each tergite hardly more sparsely punctate; punctation of tergites 8 and 10 slightly coarser than that of the previous tergites. Legs slender, metatarsi as long as metatibiae, first segment distinctly shorter than three following segments combined, slightly longer than last segment, fourth segment distinctly bilobed. Entire dorsal surface densely, more or less deeply reticulated, resulting in somewhat fatty lustre. Comments. Dianous yangae differs from D. chinensis by the much smaller elytral spots (compare fig. 102), by the narrower apical lobes of the median lobe, by the different ratio of anterior median lobe (2.3 in one male syntype of D. chinensis), and by the larger size. The specimens from Emei Shan quoted by Zheng as „chinensis Bernhauer“ belong to D. yangae as could be verified by specimens from the quoted series. Etymology: Dedicated to Miss Yang Ping, who helped in collecting. 464 VOLKER PUTHZ Dianous senex sp. n. Type material. Holotype (3) and allotype (2): China: W-Sichuan: Daxue Shan, Hailuogo Glacier Park, Camp 1, 2100 m, 29.36.00 N, 102.03.35 E, 27.- 31. V. 1997, M. Schiilke (ZMB); Paratypes: 2 d d: same data as HT (cP, cSchülke); 2 d d: ibidem 102°04‘ E, 29°36‘ N, river valley ca. 3 km above Camp I, 2100 m, 28./ 31. V. 1997, A. Pütz, D. Wrase (coll. Pütz, coll. Schiilke). This new species belongs to the group of D. chinensis and resembles many species of that group, especially D. vangae. It is remarkable by moderately long greyish pubescence of the fore parts. Moderately shining, black with (Prussian-)blue metallic reflection, elytra each with a moderately large, oval, orange spot. Punctation of fore parts coarse and dense, very slightly confluent between elytral spot and suture, abdominal punctation fine and dense. Woolly pubescence of the fore parts greyish, moderately long, half erect, longest setae of pronotum about as long as antennal segment 4, abdominal pubescence yellowish, dense, recumbent. Antennae, maxillary palpi, and legs black with blue metallic tint, trochanters black/metallic. Clypeus and labrum blue metallic with long and dense pubescence. Sternum 9 with distinct apicolateral tooth (fig. 135). Fourth tarsal segment shortly bilobed, distinct also at metatarsi. Length: 5.2 — 6.4 mm (fore parts: 3.0 — 3.1 mm). PMUotAt: wH: 45; adE: 29: IE? 18: life: 7: wP: 352 IRe 372 whe S4lEiS>- IS: 49. Male: Metasternum moderately convex, medial area coarsely and moderately densely punctate except on actual middle which is impunctate, interstices distinctly reticulated, acam with some fine punctures, punctation of sternites moderately coarse and moderately dense, interstices nearly smooth. Sternite 7 with a broad impression apicomedially which is very densely punctate and pubescent, posterior margin shallowly emarginate. Sternite 8 with a broad, rounded emargination in about posterior quarter. Sternite 9 (fig. 135), densely set with sclerotized fringes and asacl apico- medially, apical brush strong, about as long as the sternite. Tergite 10 broadly rounded and slightly concave apicomedially. Ae deagus (fig. 126) about as in the related species, with a strongly sclerotized, nightcap-like internal sac, parameres with 8/10 alS and 12/16 amS, apex of median lobe with about 17-22 short setae ventrolaterally. Female: Ventral characters similar to those of male, sternite 8 rounded (probably triangular apicomedially; incomplete in the allotype). Apical brush of valvifer nearly as long as valvifer. Tergite 10 triangularly rounded. Pubescence of fore parts greyish and woolly on head and pronotum. Head and pronotum less densely punctate than elytra, in the middle of pronotum an impunctate area about as long as antennal segment 4 and about as broad as three punctures combined. Diameter of oval elytral spot less long than antennal segment 3, about as long as antennal segments 6 + 7 combined, punctation of elytra coarse and very dense, except on a small elevated area between humeral and sutural impressions in anterior third, where interstices may become half as large as diameters of punctures, and except on posteriolateral quarter; punctation of elytral spots dense, between spot and suture very dense or slightly confluent longitudinally. Punctation of abdomen fine and dense, THE GENUS DIANOUS IN CHINA 465 punctures slightly smaller than medial eye facets, interstices also in middle of tergites at most as large as diameters of punctures, punctation of tergite 8 coarser, punctures about as large as medial eye facets; punctation of paratergites fine and extremely dense. Reticulation at places faint, somewhere distinct, at places missing. Comments. Dianous senex differs from D. yangae by smaller size, smaller elytra, less dense and coarser punctate metasternum, by the elytral spot which is comparatively larger and (longitudinally-) oval, and by the slightly coarser punctation of pronotum’s dic, from D. mendax, D. emeiensis, D. hummeli, and D. dubiosus by distinct apicolateral tooth of sternum 9, from D. atroviolaceus, D. banghaasi, D. bashanensis, D. bioculatus, D. gonggamontis, D. socius and D. taiwanensis by the greyish pubescence of the fore parts, and from some of them furthermore by semierect pubescence, from D. chinensis by the shorter, woolly pubescence of pronotum, from all by the sexual characters. This new species is very close to D. yangae sp. n. and might represent a subspecies of it, which is restricted to high mountains. Etymology: senex (Lat.) = aged man. Refers to its greyish pubescence. Dianous banghaasi Bernhauer, 1916 Dianous Bang-Haasi Bernhauer, 1916: 27 f.; L. Benick, 1940: 575 figs. Dianous banghaasi; Puthz, 1970: 32; Puthz, 1971a: 89 fig.; Zheng, 1993: 198 Dianous pilosus Champion, 1919: 54 f. Material examined: Shandong: 5 d d, 2 2 2: Kiautschau [Chiao-chou Wan] (including types of D. banghaasi) (FMCh, MM, cP). Zhejiang: 3 6 4,4 9 9: Ningpo [Yin-hsien] (including 1 type of D. pilosus) (BMNH, IRScNB, ZMB, cP); 1 6: Da-zeh near Tychau, Walker (syntype of D. pilosus) (BMNH). Jiangxi: 1 5 (without precise locality) (syntype of D. banghaasi) (FMCh); 1 d: Kuling near Kiukiang [Jiujiang], H. F. Loomis, 18. X. 1919 (USNM); 8 66,3 22: Tienmuschan, Reitter (FMCh, NHMW, ZMB, cP). Hunan: 1d, 1 9: Shaoshan, 18. X. 1966, P. M. Hammond (BMNH); 7 6d, 2 9 2: SW Huitong, env. Guangping, 300 m, 2. XI. 1993, H. Schillhammer (7), 3 d 4,3 © 2: ibidem 350 m, 4.X1.1993, H. Schillhammer (10 a); 1 2: ibidem 350 m, 8. XI. 1993, L. Li (16) (NHMW); 7 88,4 £ 2: Wulingyuan, N. Dayong Zangjiajie, 450 m, 30. X. 1993, H. Schillhammer (4), 1 2: Wulingyuan, N. Dayong Sioxiyu, 400 m, 31. X. 1993, H. Schillhammer (5) (NHMW, cP). Guizhou: 1 d,5 ® 2: no further indication (NHMW); 5 9 9: „Kouy-tcheou, région de Pin-fa, 1906, J. Cavalerie (IRScNB, MNHNP); 2 dé 4: Huangguoshu, X. 1986, G. M. de Rougemont (cR, cP), 31 exx. Longong, Huaxi, and Huangguoshu, X. 1986, G. M. de Rougemont (t. Rougemont, cR); 4 dd, 8 9 2: Bije Pref., Minxi Co., Dou Lou Lake, 1200-1300 m, 30. VII. 1997, M. Wang (NHMW, cP). Guangxi: 1 d: Dist. Lipu, 120 km S Guilin, Berge bei Siuren, 350 m, 12. XI. 1993, Schönmann & Schillhammer (19)(NHMW). Fujian: 13 4 4,21 9 2: Chungan, Bohea Hill, 16. II., 24. IV., 30. IV., 3. V. 1940, 1 d: Shaowu, Shuipeichieh, 13. I. 1940, 1 dg: Shaowu, Kaishow, 4. I. 1944, 2 2 2: Shaowu, Ku Hsien kai, 6. VI. 1943, 16. III. 1944, 1 d, 19: Changting, Hotien, 15. IV. 1941, 1 d: Changting, Ninling, 24. IV. 1941, 1 6: Yungan, 31. III. 1941: all leg. T. C. Maa (Bishop Museum, cP); 1 d: Shaowu, 500 m, 29. X. 1937, J. Klapperich (cP). Guangdong: 10 gd d, 139 2: „Canton, lebend in den Kisten“, Mell (ZMB, cP). Sichuan: 16 (abw.): N Ya’an, Baoxing County, ca. 5 km SW Baoxing, 900 m, river (Xichuan He), ca. 5 m wide, with pebbles and cobbles, cold, fast flowing through forest, Ji & Wang, 11. VI. 1996 (CWBS 228) (NHMW); 1 & (abw.): Emei Shan, 7. X. 1985, G. M. de Rougemont (cR). Zheng records numerous specimens from Emei Shan, of which he sent mee 6.2 9 2115; Vie 152 VII and 237. Vile 1983): Comments. This is the most common Dianous of China, easy to identify by its remarkably erect pubescence and the large elytral spots (fig. 104). The extension of the elytral spots seems to be variable. In most specimens examined (including types) the 466 VOLKER PUTHZ large, somewhat transverse spot on each elytron extends toward the declining lateral areas and is seemingly (in dorsal view) fused with lateral margin but is (in lateral view) actually separate from the lateral margin by a distance about equal to about the length of the fourth antennal segment. In one of the two males from Guizhou: Huangguoshu, the elytral spot is somewhat smaller and thus in dorsal view separate from darker elytral sides (the diameter of the spot is about as large as the length of the third antennal segment; about as large as the length of the third and fourth antennal segments combined in typical specimens of D. banghaasi). Some specimens from Sichuan: Emei Shan have the spots broader than long but narrower than in the typical specimens, with the length slightly larger than the length of the third antennal segment. Dissection of the genitalia and careful study of the ventral characters confirmed that these are only somewhat aberrant specimens. The labels are marked with „abw.“(= aberrant). Dianous banghaasi probably reaches the distributional limits in Sichuan and Guizhou and is represented there by peripheric populations which to some degree differ from the populations in the centre of the distribution areal (as it is known in D. coerulescens Gyllenhal, see Puthz, 1980b: 7 ff.). The aedeagus (fig. 127) is also variable in the number of the setae: the parameres have 8-11 alS and 9-11 amS, the apicolateral lobes of the median lobe have 18-24 small ventral setae; the number of setae varies both in one specimen and in specimens from the same series. A good working distinguishing character is the ratio of breadth of median lobe at bottom of apical emargination: depth of emargination, which varies in the material examined between 1.2-1.4 (this ratio differs from that of the related species, e. g. D. taiwanensis Puthz, D. aequalis Zheng, D. bioculatus sp. n.). Sternite 8 of the female has a distinct triangular projection apically (fig. 131). Dianous aequalis Zheng Dianous aequalis Zheng, 1993: 199 f. fig. Material examined: | S-PT: Sichuan: Emei Shan, 710- 1120 m, Gou Zheng-yue (cP) [HT, AT and 4 PT in the Dept.Biol., Sichuan Teachers College, Nanchong]. Guangdong: 2 dd, I 2: „China, Canton, lebend in den Kisten“, Mell (ZMB, cP). This species is closely related to D. banghaasi, it also has conspicuous erect pubescence. The description is taken from a male labelled „Canton“ (see below). Moderately shining, black with violet (-bluish) tint, each elytron with a large orange spot (diameter about as large as length of third antennal segment, or even larger). Head moderately coarsely and moderately densely punctate, pronotum coarsely and densely, near posterior border somewhat coalescently, punctate, elytral punctation coarse, mostly well-defined, except sometimes slightly confluent near medial border of elytral spot, abdomen finely and moderately densely punctate. Pubescence yellowish, long, dense, erect. Antennae brown, club lighter, maxillary palpi brownish, legs blackish brown, tarsi lighter. Clypeus and labrum bluish, densely pubescent. Fourth segment of metatarsi distinctly but shortly bilobed. Length: 4.2 — 5.2 mm (fore parts: 2.5 — 2.7 mm). PM of d from „Canton“: wH: 38; adE: 23; IE: 17; ITe: 4; wP: 31; IP: 33; wEl: 49; JE]: 49; IS: 42. THE GENUS DIANOUS IN CHINA 467 Male: Metasternum broadly convex, medial area slightly impressed and impunc- tate posteriorly (impunctate area triangular, about as long as fifth segment of metatarsus, and as broad as length of second metatarsal segment posteriorly), punc- tation moderately coarse, regular, dense, diameters of punctures about as large as basal cross section of third segment of maxillary palpi, interstices slightly larger than half diameters of punctures, nearly smooth, acam with some fine and scattered punctures, shiny. Sternites 3-6 somwhat more sparsely punctate medially than laterally; sternite 7 with very shallow posteriomedial impression, punctation and pubescence of impression fine and very dense, posterior margin simple. Sternite 8 with round notch in about posterior quarter. Sternite 9 with acute tooth apicolaterally (fig. 136), densely set with long sclerotized fringes and asacl on an oval field apicomedially, apicolateral brush strong, as long as sternite. Tergite 10 broadly rounded. Ae deagus similar to those of related species, apical emargination of median lobe deep (fig. 128), ratio of breadth of median lobe at botom of apical emargination: depth of apical emargination = 1.3 — 1.7; parameres slightly longer than median lobe, with 5-7 (8) alS and 8- 9 (10) amS; apical lobes of median lobe with 13-19 small ventral setae. Female: Ventral characters similar to those of male. Sternite 8 (fig. 132) with a long triangular projection. Valvifer acute apically, apical brush about as long as valvifer. Tergite 10 rounded. Dianous aequalis is in many respects very close to D. banghaasi, but the outer antennal segments are longer, nearly twice as long as broad, the elytral sculpture denser, intermediate between D. banghaasi and D. chinensis with posterio-lateral quarter coarsely and densely punctate, punctures as coarse as on anterior half of elytra, interstices smaller than half diameters of punctures (see below), elytral spots (in dorsal view) distinctly separate from the blackish-blue lateral areas. Variability: In the PT from Emei Shan the posteriolateral punctation of elytra is less coarse and less dense than in the Canton-d, but distinctly more dense than in D. banghaasi; the parameres have 7/8 alS and 10 amS. Comments. Dianous aequalis Zheng is distinguished from D. banghaasi by smaller size, by the characters given above, and by the sexual characters, from D. chinensis also by smaller size, by the smooth interstices of punctation, and by the aedeagus with narrower apical lobes of the median lobe and deeper apical emargination. R. E. Mell (1878- 1970) lived a long time in Canton; possibly the material has been collected there and shipped alive to the Berlin museum; pers. comm. Dr. Uhlig. Dianous dubiosus sp. n. Type material. Holotype ( ): Guizhou: Huangguoshu, X. 1986, G. M. de Rougemont (coll. Rougemont in BMNH). 1 2 paratype: Guangxi: Da Ming Shan, 1000 m, 13. V. 1997, J. Fellowes (cP). This Dianous resembles in almost all characters very strongly D. emeiensis Zheng but has a distinct aedeagus and a more deeply (distinctly) bilobed segment 4 of the metatarsi (fig. 141). Moderately shining, blackish-blue-metallic (Prussian blue), elytra each with a moderately large organge spot (about as in fig. 105 or slightly smaller). Punctation of 468 VOLKER PUTHZ head coarse and very dense, interstices mostly smaller than half diameters of punctures, deeply reticulated, punctation of pronotum as coarse as in D. emeiensis but slightly denser, elytral sculpture very coarse, and dense, slightly transversely coalescent on lateral portions of anterior half, also coalescent around anterior and sutural border of orange spots, abdomen moderately finely but more densely punctate than in D. emei- ensis. Pubescence, antennae, palpi, and legs as in D. emeiensis, segment 4 of metatarsi distinctly, but shortly bilobed (fig. 141). Length: 4.5-5.7 (fore parts: 2.7-2.8 mm). PM of HT: wH: 40.5; adE: 24; wP: 33: IP: 34,5; wEl: 52; IEL: 55; IS: 47. Male: Metasternum strongly convex, medial area not flattened, with midline narrowly impunctate, sides moderately finely and densely punctate, punctures smaller than basal cross section of palpal segment 3, interstices smaller than diameters of punctures, very densely, moderately shallowly reticulated, shiny; acam very finely, sparsely punctate, interstices smooth. Medial punctation of sternites 3-6 moderately fine, slightly sparser than lateral punctation, sternite 7 with posterior broad, shallow impression more densely punctate than sides, posterior margin shallowly emarginate. Sternite 8 with a broad and deep notch in about posterior quarter to fifth. Sternite 9 as in fig. 137, irregularly serrate apically, without apicomedial fringes, apicolateral brush strong, distinctly shorter than sternite. Tergite 10 very broadly rounded, slightly concave posteriomedially. Aedeagus as in fig. 130, resembling that of D. bashanensis Zheng (fig. 144), apical portion of median lobe less deeply emarginate, with about 16- 17 apicoventral short setae on each side; parameres longer than in D. bashanensis and stronger apically, with 7 alS and 11 amS. Female: Metasternum nearly as in the male, but no distinct microsculpture throughout. Sternite 8 pointed apicomedially. Valvifer oblique-triangular posteriorly, apical brush nearly as long as valvifer. Tergite 10 rounded. Variability: The elytral spot of the ©-paratype is smaller than that of the holotype. Comments. Dianous dubiosus is distinguished by the aedeagus and the distinctly bilobed segment 4 of metatarsus. It is more coarsely sculptured and more densely reticulated than D. emeiensis. It can be distinguished from D. bashanensis Zheng by the larger elytra, the stronger metallic tint, and by the elytral spot situated in about posterior half of elytra. | 2: Taiwan: Nantou Hsien: Sun Lin Sea, rte. 151, 16 km SSE Luku, 6.1V.1991, H.G.Nelson (FMCh): cf. det.: This specimen resembles closely the type of D. dubiosus but elytral sculpture is even coarser, abdominal punctation slightly less dense. Etymology: dubiosus = dubious; it refers to the difficulty to distinguish this species from D. emeiensis by the external characters and to the lack of the type material for comparison. Dianous cyaneocupreus sp. n. Type material. Holotype (JS): Taiwan: Taipei Hsien: South Fork Ching Mei Str., 3 km SE Ching Mei (19 km SE Teipei), 26. II. 1991, H. G. Nelson (FMCh). THE GENUS DIANOUS IN CHINA 469 This new species resembles many of the bluish species with elytral spots, it belongs to the group of D. chinensis Bernhauer. Moderately shining, black with distinct bluish tint, elytra each with a large cupreous spot. Punctation of fore parts coarse and dense, somewhat confluent on some areas of elytra, abdomen very finely and sparsely punctate. Pubescence of fore parts brownish, very long, erect, especially remarkable on clypeus and on pronotum; abdomen with dense, long, recumbent, golden pubescence. Antennae, maxillary palpi, and legs black with some dark bluish metallic tint, trochanters dark brown. Clypeus and labrum blue metallic with very long pubescence. Fourth segment of pro- and mesotarsi distinctly but slightly biolobed, that of metatarsi almost simple (fig. 142). Length: 5.0 — 6.0 mm (fore parts: 3.0 mm). PM of HT: wH: 44; adE 27; IE: 19; IT: 6; wP: 35; 1P:38; wEl: 55; IEI: 57; 1S: 49. Male: Metasternum, acam and ventral abdomen about as in D. atrocyaneus, punctation of sternites very fine, medially slightly sparser than laterally. Sternite 8 with a deep, broad, rounded emargination in about posterior quarter. Outline of sternite 9 about as in D. atrocyaneus, with a broad area densely set with long sclerotized fringes and minute asacl apicomedially (fig. 138), apicolateral brush strong, 5/6 as long as the sternite. Tergite 10 broadly rounded. Ae deagus about as in D. atrocyaneus (see fig. 113), parameres with 6 + 12 setae, apex of median lobe with about 23-27 short setae ventrolaterally. Female: Unknown. Elytra with a large cupreous spot in posterior half, diameter slightly smaller than length of antennal segment 3; elytral punctation even coarser and mostly denser than on pronotum, in anterior half transversely confluent, between spot and suture longitu- dinally confluent, very dense and slightly confluent on the spot. Comments. Dianous cyaneocupreus differs from D. cupreostigma Puthz by much longer pubescence and finer and sparser abdominal punctation, from D. yoshidai Naomi by coarser punctation of frons and elytra, longer pubescence and much finer and sparser abdominal punctation, from D. taiwanensis Puthz by color, longer pubescence of pronotum and very dense punctation of paratergites, from D. dubiosus sp. n. by very fine and very sparse abdominal punctation and smooth interstices of punctation of nearly the whole insect, from all these species by the aedeagus. Etymology: cyaneus (Latin) = bright blue, cupreus (Latin) = cupreous. Refers to the coloration. Dianous hummeli Bernhauer, 1936 Dianous Hummeli Bernhauer, 1936: 3; Puthz, 1971a: 90f. figs. Dianous szechuanus Bernhauer, 1936, l.c. : 3 f. Material examined. Sichuan: Holotype (male) of D. hummeli and holotype (female) of D. szechuanus: Kia-ling-ho, Youen-kou (FMCh). Male: Metasternum broadly convex, medial area flattened, with midline narrowly impunctate, sides of medial area moderately coarsely, densely punctate, diameters of punctures slightly smaller than basal cross section of palpal segment 3, 470 VOLKER PUTHZ interstices slightly larger than half diameters of punctures, distinctly reticulated, moderately shiny, acam finely and sparsely punctate, interstices nearly smooth. Ster- nites 3-6 finely punctate (punctures distinctly smaller than one eye facet), there are fewer punctures medially than laterally, sternite 7 slightly flattened medially, but not impressed, very densely punctate and pubescent, posterior margin simple. Sternite 8 with a broad and very deep notch in about posterior third to fourth. Sternite 9 as in fig. 152, posterior margin with medial sclerotized fringes, apicolateral brush strong, about as long as sternite. Tergite 10 broadly rounded. Aedeagus (fig. 143) with median lobe broad and broadly emarginate at apex; apical lobes each with about 20 short ventral setae; parameres not much longer than median lobe, each with 6 alS and 10-11, strong and moderately long, amS. Female: Ventral characters similar to that of male. Sternite 8 (fig. 134) Tergite 10 subtriangular. Comments. The punctation of head and pronotum is comparatively sparse and not as coarse as in the following two species: the diameters of punctures on frons are nearly as large as basal cross section of palpal segment 3, interstices are mostly larger than half diameters of punctures, but can be also as large as diameters of punctures. The punctation of the pronotum is somewhat coarser than that on frons, but it is relatively sparse in the middle, where interstices become much wider than diameters of punctures; the impunctate area in the middle of the pronotum is about as broad as the length of the antennal segment 11: the medial punctures of pronotum are slightly smaller than the basal cross section of antennal segment 3, but the punctation becomes much coarser and much denser toward posterior margin, where slight coalescence can be seen; the interstices of pronotal and frontal punctation are nearly smooth, without appreciable reticulation. Dianous emeiensis Zheng, 1993 sp. propr. Dianous hummeli emeiensis Zheng, 1993: 201 figs. Material examined: 4 4 4,4 2 © Guizhou: Huaxi, X. 1986, G. M. de Rougemont (coll. Rougemont, coll. Puthz). This Dianous closely resembles D. hummeli Bernhauer and is its sister species. Main differences, that are of specific rank, are in male genitalia. Moderately shining, blackish- blue- metallic (Prussian blue), elytra each with a moderately large orange spot (fig. 105). Punctation of head moderately coarse and dense, that of pronotum conspicuously coarser, very dense and somewhat coalescent, punctation of elytra about as coarse as on pronotum, coalescent around orange spots, abdomen moderately finely, densely punctate. Pubescence whitish, conspicuously dense, long and semi-erect. Antennae, palpi, and legs black with blue metallic tint, trochanters with brownish tint, narrow bases of middle antennal segments reddish. Clypeus and labrum black with blue metallic tint, moderately densely pubescent. Segment 4 of metatarsi very shortly (indistinctly) bilobed (fig. 140). Length: 5.0-6.5 mm (fore parts: 2.8-3.0 mm). PM of one male: wH: 40; adE: 25.5; wP: 32; IP: 34; wEl: 54; IEl: 54; IS: 46. THE GENUS DIANOUS IN CHINA 471 Male: Ventral characters very close to those of D. hummeli, but punctation of sternites less fine, punctures on middle of sternites about as coarse as one eye facet. Sternite 8 with a broad notch in about posterior quarter to fifth (less deep than in D. hummeli). Sternite 9 as in fig. 149, middle of posterior margin with numerous sclerotized fringes and a field of asacl apicomedially, apicolateral brush strong, a little longer than sternite. Tergite 10 broadly rounded. Ae deagus as in figs 145, 147, 148, median lobe broad, apical portion less broadened than in D. hummeli, apical margin variable, not emarginate; apical lobes each with 20-26 short ventral setae; parameres longer than in D. hummeli, number of setae variable: 6-10 alS and 11-19 amS, setae very strong and somewhat longer than in D. hummeli. Female: Ventral characters similar to those of male, sternite 8 as in fig. 146, apicomedial projection narrower than in D. hummeli. Tergite 10 subtriangular. Comments. Dianous emeiensis is more robust than D. hummeli, punctation of the fore parts, especially the head and pronotum, is coarser and denser. The vorticose sculpture near the elytral spots is more distinct, and the punctation of the abdomen is slightly coarser than in D. hummeli. Dianous bashanensis Zheng, 1994: 480 f. figs. Material examined: NW Hunan: 1 © Wulingyuan N. Dayong, Zangjiajie, 650 m, 29. X. 1993, Schillhammer; 2 8 d: Wulingyuan N. Dayong, Sioxiyu, 450 m, 31. X. 1983, Schönmann (NHMW, cP). PM of a male from NW Hunan: wH: 40; adE: 25; wP: 32; IP 32.5; wEl: 46; IEl: 46; IS: 38. Male: Metasternum with an impunctate and strongly shining triangular area medially, posteriorly this area is about as broad as the length of segment 2 of the maxillary palpi; sides of shiny area moderately finely and moderately densely punctate, diameters of punctures somewhat smaller than basal cross section of palpal segment 3, interstices about as large as diameters of punctures, smooth; toward the declining sides follows a narrow strip of very fine and very dense punctation laterally, declining sides moderately finely and sparsely punctate, acam with very few fine punctures. Sternites 3-5 medially finely and distinctly sparser punctate than laterally, sternite 6 broadly flattened, medial punctation slightly coarser than lateral one. Sternite 7 with a shallow longitudinal impression, sparsely punctate anteriorly, densely punctate and pubescent near posterior border, where it is shallowly emarginate. Sternite 8 with a broad rounded notch in about posterior quarter to fifth. Sternite 9 (fig. 150), with numerous sclerotized fringes and dense asacl apicomedially, apicolateral brush moderately strong, as long as the sternite. Tergite 10 broadly rounded. Aedeagus (fig. 144), apical portion of median lobe broad and deeply emerginate, each side ventrally with 14-16 small setae; parameres slender, distinctly longer than the median lobe, with about 7 alS and about 10 amS. Female: Sternite 8 (fig. 133). Valvifer acute apically. Tergite 10 subtriangular. Comments. This species was described from Sichuan: Da Bai Mountains. It is very close to D. socius Zheng, from which it is distinguished best by its sexual characters. 472 VOLKER PUTHZ Dianous atroviolaceus sp. n. Type material. Holotype (4) and allotype (2): NW Yunnan: ca. 10 km SW Lijiang, 2700 m, 5. VII. 1994, Li (14) (NHMW). Paratypes: 3 dé 4 same data as HT; 1 6,2 2 2: Yunnan: 100 km W Kunming, Diaolin Natural Reserve, 22. V.-2. VI. 1993, E. Jendek & O. Sausa; 1 2: Lugu Lake — Luo Shui, 27°45’ N, 100°45’ E, 8.- 9. VII.1992, E. Jendek (NHMW, cP). This new species is very closely related to D. taiwanensis, from which it differs easily by the dark violet tint. In general habitus the micropterous form resembles strongly D. taiwanensis, the macropterous form D. chinensis Bernhauer. Variably winged, moderately shining, black with dark violet tint, elytra each with variably large orange spot. Punctation of head moderately coarse and moderately dense, that of pronotum somewhat coarser, mostly dense, slightly confluent near posterior margin, punctation of elytra coarse, dense, punctures obliquely extended, coalescent between spots and suture (also somewhat coalescent at lateral border of orange spot), abdomen very finely and moderately sparsely punctate. Pubescence blackish, dense, that of fore parts half erect, longest setae nearly as long as second antennal segment. Antennae, maxillary palpi, and legs blackish with dark violet to dark coppery tint. Clypeus and labrum dark violet, moderately densely pubescent. Fourth segment of metatarsi distinctly, shortly bilobed. Length: 4.7 — 6.0 mm (fore parts: 2.4 — 3.0 mm). PM of the (incompletely winged) HT and of the (fully winged) d- PT from Diaolin Nat.Res.: wH: 40 (42,5); adE: 26.5 (28); IE: 16.5 (17); ITe: 7 (7); wP: 31 (36); IPS GO) ES 68) 1ELAS 69) 1S 57610); Male: Medial area of metasternum broadly flattened, triangularly impunctate posteriomedially, side portions moderately coarsely and moderately densely punctate, diameters of punctures about as large as basal cross section of third segment of maxillary palpi, interstices as large as, or slightly larger, than diameters of punctures, densely reticulated but shiny; acam very finely and sparsely punctate, nearly smooth; declining lateral areas of metasternum finely and moderately sparsely (near medial area) to moderately coarsely and densely (laterally) punctate. Medial punctation of abdominal sternites 3-6 somewhat sparser than lateral punctation; sternite 7 distinctly impressed posteriomedially, punctation of impression moderately fine and dense, posterior margin distinctly, shallowly emarginate. Sternite 8 with a deep notch in about posterior quarter. Sternite 9 (about as in fig. 135), numerous sclerotized fringes and dense asacl apicomedially, apicolateral brush strong, as long as the sternite. Tergite broadly rounded. Aedeagus (fig. 154) of the D. banghaasi-type, ratio of breadth of the median lobe at bottom of apical emargination to depth of apical emargination = 1.50 — 1.60; parameres each with 7-9 alS and 9-12 amS; apical lobes of median lobe with 28-30 small setae. Female: Ventral characters similar to those of male, but sternite 7 not impressed. Sternite 8 (fig. 164). Valvifer triangular apically, apical brush strong, as long as valvifer. Tergite 10 subtriangular. In nearly all respects about same as the variable D. taiwanensis but differs as follows: color dark violet, not black or black with greenish blue tint, elytral sculpture denser, with punctures very dense and slightly longitudinally confluent also near lateral THE GENUS DIANOUS IN CHINA 473 border of orange spots (not at all so in D. taiwanensis), and ventral punctation less coarse and less dense. Comments. D. atroviolaceus differs from D. bioculatus by the slightly confluent punctation near the lateral borders of the elytral spots, and by the male sexual characters. The males of D. atroviolaceus from 2700 m are much narrower and have smaller elytral spots (nearly as large as the length of the third antennal segment) than the male from Diaolin Natural Reserve (diameter of its elytral spot distinctly larger thn length of 3' antennal segment), but in the females from the respective localities the size of the spot varies. Etymology: ater (Lat.) = black, violaceus (Lat.) = violet. Dianous gonggamontis sp. n. Type material. Holotype (4) and allotype (9): Sichuan: Gongga Shan, above Camp 3, 3050 m, on silty soil near a creek, 22. VII. 1994, A. Smetana (NHMG). Paratypes: 7 88,4 9 9: same data as HT; 1 d : ibidem, 3300 m, gravel bank of a creek, 23. VII. 1994, A. Smetana (cS, cP); 1 4,3 £ 2: Gonggashan: Hailuogou, 1900-2900 m, 29°36’ N, 102°06’ E, 6. VII. 1994, D. Kral & J. Farkac (coll. Hromadka, cP); 1 4: W- Sichuan: Daxue Shan, Hailuoguo Glacier Park, Camp 2, 2550- 2700 m, 29.35.16 N, 102.01.53 E, 30./ 31. V. 1997, M. Schülke (cSchiilke); 2 3 d: ibidem 2650 m, 30. V. 1997, A. Pütz (coll. Pütz); 1 4: ibidem, 2500-2700 m, 30./ 31. V. 1997, Wrase (coll. Schülke). Material „cf. det.“: Sichuan: 1 9: Guan Xian Co., Balang Shan 15 km SW Wolong, ca. 2300 m, mountain stream, 5 — 10 m wide (river bed up to 50 m wide), crystalline gravel and sand, very fast flowing, very cold, 1. VIII. 1998, Schönmann, Ji, Wang (CWBS 342) (NHMW); 1 d, 1 2: Wanglang, 15. VII. 1988 (cP). This new species closely resembles D. banghaasi Bernhauer, also in its sexual characters, but the elytral spot is smaller and the pubescence of the fore parts much shorter, and semierect. Blackish-blue-metallic (Prussian blue), shining, elytra each with a large orange spot, which is mostly somewhat less large than the length of antennal segment 3 (exceptions !) and distinctly separate from the elytral side. Fore parts coarsely and very densely punctate, punctation coalescent between orange spot and suture (short rugae), abdominal punctation fine and moderately dense, moderately sparse in middle of tergites, where interstices are distinctly or much larger than diameters of punctures; punctation of tergite 8 distinctly, but not conspicuously, coarser than that of tergite 7. Pubescence blackish, dense, short, semierect, setae of pronotum not as long as antennal segment 2. Antennae, palpi, and legs blackish, with blue-metallic tint. Clypeus and labrum dark blue-metallic, moderately densely pubescent. Segment 4 of metatarsi simple (lobation about as in D. emeiensis, see fig. 140). PM of HT: wH: 40; adE: 24; wP: 31.5; IP: 32; wEl: 45.5; 1El: 48; IS: 40. Male: Metasternum broadly flattened, narrowly impunctate posteriomedially, punctation on disc moderately coarse and moderately sparse, diameters of punctures about as large as basal cross section of palpal segment 3, interstices mostly distinctly larger than diameters of punctures, may become twice as large as, shallowly reticulated, shiny; declining sides finely and sparsely punctate (no distinct strip of denser punctation between fine lateral and coarse medial punctation), acam nearly smooth. Medial punctation of sternites 3-5 slightly coarser and slightly sparser than lateral 474 VOLKER PUTHZ punctation, sternite 6 flattened posteriomedially, somewhat denser punctate than on the sides, posterior margin slightly emarginate. Sternite 7 shallowly impressed posterio- medially, impression finely and densely punctate and pubescent, posterior margin shallowly emarginate. Sternite 8 with a deep notch in about posterior quarter. Sternite 9 about as in D. banghaasi, with numerous sclerotized fringes and fine asacl apico- medially and a broad tooth apicolaterally. Tergite 10 broadly rounded. Aedeagus about as in D. banghaasi, median lobe broadened apically, deeply emarginate apico- medially (index 1.30-1.40), parameres with 8-10 alS and 9-13 amS. Female: Ventral characters similar to those of male, except last sternites; sternite 7 in posterior middle finely and densely punctate and pubescent, sternite 8 subtrian- gular, about as in D. banghaasi. Tergite 10 moderately narrowly rounded, not distinctly subtriangular. Comments. Some specimens from Sichuan differ from the type series by slightly different sculpture and are therefore not labelled as paratypes. Dianous gonggamontis sp. n. is distinguished by its short, semierect pubescence and sparse punctation of the apical quarter of elytra. Dianous taiwanensis Puthz Dianous taiwanensis Puthz, 197 1a: 88 f. fig.; Puthz, 1984a: 101 Dianous bilunatus Puthz, 1984a: 101 f. figs syn. nov. Material examined: holotype (9): Taiheizan, 6. V. 1932, L. Gressitt (USNM). Taipei Hsien: 1d: Pihu, 49 km E. of Taipei, on Highway 9 (to Yilan), 18. II. 1972, T. C. Maa (Bishop Museum). Ilan Hsien: 9 4 4,99 2: Taipingshan, 1880 m, 14. VII. 1993, in soaking wet moss on a vertical wall in a creek; 7 6d, 4 9 9: ibidem 1820 m, 15. VII. 1993, in wet moss on large rocks directly in a creek. Taoyuan Hsien: 6 d d: Takuanshan Forest, 1600 m, 17. IV. 1990, on wet rocks in a creek. Nantou Hsien: 2 dd (abw.): Shanlinchi, 1650 m, 16. V. 1990, same habitat as previous; 2 646, 1 9: ibidem 19. V. 1991 all leg. A. Smetana (cS, cP). Chiayi Hsien: 1 d, 2 9 2: Alishan, 2400 m, 3.- 9. VII. 1972, T. C. Maa (types of D. bilunatus) (Bishop Museum, cP); 1 d: ibidem 2300 m, 9. IV. 1965, Y. Hirashima (Kyushu Univ.); 1 d: ibidem, 17. V. 1968, Y. Watanabe (cP); 1 dg: Alishan, Sister Ponds, 2180 m, 26. IV. 1990, soaking wet moss on rotting wood around small ponds, A. Smetana (cS). Kaohsiung Hsien: 1 4,2 9 9: Creek 2 km E Tien Chih, Hwy. 20, 2400 m, in soaking wet moss on a forest sepage, 22. VII. 1993; 10 88,14 9 9: Creek 4 km E Yakou, 2600 m, 23. VII. 1993, in soaking wet moss on large rocks in a creek, A. Smetana. Pingtung Hsien: 2 © 9: Peitawushan Kuai-Ku Hut, 2120 m, 27. IV. 1992, wet moss on a large fallen tree. Hualien Hsien: 7 4 4,5 © 9: Taroko N.P., Nanhushi Hut, 2220 m, 8. V. 1990, in wet leaf litter on large rocks along a creek; 3 dd, 2 © ©: Taroko N.P., Chungyantienshi (River) Waterfall, 2300 m, 10. V. 1990, in soaking wet moss on a vertical wall near a creek, all leg. A. Smetana (cS, cP). Comments. This species is widely distributed in Taiwan and considerably variable. Main differences given in my descriptions of both taxa (D. taiwanensis and D. bilunatus) prooved to fall within the variability range: (a) proportions of the individual tagmata vary in size (figs 107-112) but no specimen is as fully winged as e. g. D. banghaasi, winges are in different degrees shortened, (b) the size of the elytral spot varies in the same series (figs 108, 112), (c) the rugoseness of the elytral sculpture is variable: most (somewhat backwards extended) punctures are well-defined, or (in other specimens from the same locality) confluent longitudinally, but they are distinctly separate at lateral borders of spots, (d) the reticulation varies: interstices of the fore THE GENUS DIANOUS IN CHINA 475 parts are smooth in some specimens, shallowly reticulate in others, (e) punctation of the broad median area of the metasternum is slightly variable: the median portion is impunctate, the side portions are moderately coarsely and sparsely punctate; interstices vary from about as large as diameters of punctures to twice as large, reticulation of the surface of interstices varies from very shallow to deep, (g) most specimens are black, some + bluish (-green). Aedeagus in general as in D. banghaasi and related species (figs 157-160), the number of setae on parameres varies between 8-11 alS and 8-13 amS and between 19-24 ventrally on apical lobes of the median lobe; the ratio width of median lobe at base of apical emargination: depth of emargination varies from 1.27 (T 60) — 1.82 (T 24). The two males from Shanlinchi (T 60) differ from other specimens by the larger elytra (fig. 107), the deep reticulation of the entire body, and by the deep apical emargination of the median lobe (ratio: 1.27), but I cannot find any other characters which would justify a separate status of these extreme specimens. Dianous taiwanensis is very close to D. bioculatus sp. n., but it may be easily distinguished by the more oblique shoulders, the shorter pubescence, and by the aedeagus. Dianous bioculatus sp. n. Type material. Holotype (¢) and allotype (2): Yunnan: 100 km W. Kunming, Diaolin Natural Reserve, 22. V.- 2. VI. 1993, E. Jendek & O. Sausa (NHMW). Paratypes: 11 dd; 19 2 2: same data as holotype; 2 6 d, 10 2 2: Yunnan: Yipinglang, 1800 m, 25°05‘N, 101°53;, 8.- 10. VI. 1993, Bolm (NHMB, cP, cZheng). This new species is allied to D. taiwanensis which it closely resembles. Moderately shining, black with very faint dark blue tint, elytra each with a moderately large orange spot (about as in fig. 107). Punctation of head moderately coarse and dense, that of pronotum somewhat coarser, also dense, punctation of elytra coarse and dense, more or less coalescent around orange spots, abdomen finely and moderately sparsely punctate. Pubescence conspicuous, blackish, dense, long, erect. Antennae, maxillary palpi, and legs black with some metallic tint, trochanters brownish, narrow bases of intermediate antennal segments reddish. Clypeus and labrum black with blue/violet metallic tint, moderately densely pubescent. Fourth segment of metatarsi distinctly but shortly bilobed. Length: 4.5 — 6.0 mm (fore parts: 2.4 — 2.9 mm). PMrotiiht=wH 405 7adke255 1b 217.5: TE 76 32WE EI SIEB EMERATIIEE 49; IS: 41. Male: Medial area of metasternum triangularly flattened, shallowly impressed and impunctate but shallowly reticulated posteriomedially, lateral punctation modera- tely coarse and moderately dense, diameters of punctures about as large as basal cross section of third palpal segment, interstices somewhat larger than diameters of punc- tures, acam with some 2 or 3 very fine punctures, smooth; declining lateral portions of metasternum finely and densely punctate, punctation becoming coarser and denser toward lateral borders. Medial punctation of sternites 3-5 slightly less dense than lateral 476 VOLKER PUTHZ punctation; sternite 6 with shallow impression posteriomedially with punctation and pubescence denser than that of sides, posterior margin very shallowly, broadly emar- ginate; sternite 7 with distinct finely and densely punctate and pubescent median impression in about posterior two thirds, posterior margin shallowly emarginate. Ster- nite 8 with round notch in about posterior fourth. Sternite 9 with acute tooth apicola- terally (fig. 162), numerous sclerotized fringes and dense asacl apicomedially, apico- lateral brush strong, slightly shorter than sternite. Tergite 10 broadly rounded. A e d e a- g u s similar to that of D. banghaasi, but apical lobes of median lobe shorter and broader (fig. 155), ratio of breadth of median lobe at bottom of apical emargination: depth of apical emargination = 2.8-4.0; parameres each with about 7-8 alS and 8- llamS; apical lobes of median lobe with about 27 short ventral setae. Female: Ventral characters similar to those of male, but sternites 6 and 7 simple, sternite 7 posteriomedially more densely punctate and pubescent than on sides. Sternite 8 with a considerable triangular projection (fig. 165). Valvifer obliquely acute apically, apical brush about as long as valvifer. Tergite 10 broadly rounded, not triangular. Head small, much narrower than elytra, longitudinal furrows of frons distinct but not as deep posteriorly as those of D. distigma and closely related species, medial and lateral portions broadly elevated, distinctly extending beyond level of inner eye margins, punctation moderately coarse and dense, diameters of punctures nearly as large as basal cross section of third antennal segment, some interstices about as large as half diameters of punctures, some (especially posteriomedially and near posterior inner eye margins) larger, becoming as large as diameters of punctures, but not forming distinctly separate shining areas. Antennae very long, extending beyond posterior mar- gin of pronotum when reflexed by nearly four segments. Pronotum with deep oblique impressions on posterior half and with transverse impression near anterior margin; punctation coarse and dense (except on actual middle), diameters of punctures as large as basal cross section of third antennal segment or slightly larger, interstices mostly distinctly smaller than half diameters of punctures, sculpture sometimes more or less coalescent e. g. posteriomedially; a distinct impunctate shining area, about 2-3 punc- tures broad and 4-6 punctures long, or shallow longitudinal impression in middle. Elytra subquadrate, shoulders prominent, sculpture coarser and denser than that of pronotum, largest punctures nearly as large as apical cross section of third antennal segment, punctation becoming somewhat confluent and somewhat tortuous around elytral spots, but rugae are short, not forming long channels. Abdomen with broad, upwards directed, finely and very densely punctate paratergites, punctation of tergites even finer and sparser, punctures nearly as large as medial eye facets, interstices about twice as large as diameters of punctures, punctation of tergites 8 and 10 slightly coarser. Legs slender, metatarsi 5/7 as long as metatibiae, first segment distinctly shorter than following three segments combined, somewhat longer than last segment, fourth seg- ment distinctly, but shortly bilobed. Faint reticulation on tergites 7-10, rest of surface lacking any groundsculpture. Comments. Dianous bioculatus differs from D. taiwanensis by the rectangular shoulders, by the slightly longer pubescence, and by the the aedeagus. THE GENUS DIANOUS IN CHINA 477 Etymology: bi- oculatus (Lat.) = with two eyes. Dianous socius Zheng Dianous socius Zheng, 1993: 200 f. figs. Material examined: 1 dé: Sichuan: Mt. Emei, 600-1050 m, 5.- 19. V. 1989, L. Bocak (NHMB); 1 2: Emei Shan, 7. X. 1985, G. M. de Rougemont (CR); 1d: Quingcheng Shan 65 km NW Chengdu, 103.33 E, 30.53 N, 8 km W Talping, 800-1000 m, 18. V./ 3.- 4. VI. 1997, A. Pütz (coll. Pütz); 2 ¢ 4 -PT: Emei Shan, 710-1120 m, Gou Zheng-yue & Lu Ying (cP). [HT, AT and 4 paratypes from Emei Shan in the Dept.Biol, Sichuan Teacher’s College, Nanchong. | PM of the first & listed above: wH: 42; adE: 27; IE: 17.5; ITe: 7; wP: 34.5; IP: Sop wiles 2 EE 53:5; 15: 47. This species is allied to D. banghaasi Bernhauer, which it closely resembles in general habitus except for much smaller elytral. spots. Shining black with faint dark-blue metallic tint, elytra each with a moderately large orange spot (diameter about as large as length of first metatarsal segment). Head moderately coarsely and moderately densely punctate, pronotum coarsely and densely punctate, punctation of elytra even coarser than that of pronotum, slightly coalescent at anterio-medial border of orange spots, abdomen finely and moderately sparsely punctate. Pubescence remarkably long, dense and erect, even slightly longer than in D. banghaasi. Antennae, maxillary palpi, and legs black with some metallic tint, narrow bases of intermediate antennal segments reddish, club brownish, trochanters brownish, tarsi sometimes brownish. Clypeus and labrum bluish, densely pubescent. Fourth segment of metatarsi distinctly but shortly bilobed. Length: 4.7 — 6.2 mm (fore parts: 2.9 — 3.0 mm). Male: Metasternum broadly convex, shallowly impressed posteriomedially, impunctate and smooth on a small triangular area, sides of medial area moderately coarsely and densely punctate, diameters of punctures about as large as basal cross section of second segment of maxillary palpi, smooth interstices mostly about as large as diameters of punctures, acam with a few very fine, scattered punctures. Medial punctation of sternites 3-6 fine and sparse, smooth interstices twice as large, or even larger, than diameters of punctures; sternite 7 shallowly impressed posteriomedially, finely and moderately densely punctate and pubescent, posterior margin nearly imper- ceptibly emarginate. Sternite 8 with a broad and deep notch in apical quarter. Sternite 9 with a very strong tooth apicolaterally (fig. 163), apicolateral brush strong, slightly shorter than sternite. Tergite 10 broadly rounded. Ae deagus in general similar to those of related species but apical lobes of the median lobe very broad, apical emargination relatively shallow (fig. 156, ratio of breadth of median lobe at bottom of apical emargination : depth of apical emargination = 3.1-3,3); parameres with 8 alS and 11-13 amS; apical lobes of median lobe with about 18-21 short ventral setae. Female: Ventral characters similar to those of male, but sternite 8 with a remarkable, long triangular projection apically (fig. 167). Valvifer triangularly acute apically, apical brush about as long as sternite. Tergite 10 rounded. In most respects as D. banghaasi, that is with small head, shallow posterior longitudinal furrows of frons, moderately coarse and dense punctation of frons, very 478 VOLKER PUTHZ long antennae extending beyond posterior margin of pronotum by four segments when reflexed, relatively shallow pronotal impressions. Different by the punctation of the pronotum being slightly coarser and somewhat confluent posteriomedially (less regular, less distinctly well-defined than in D. banghaasi), and by the presence of a distinct but small impunctate shining area on actual middle of the pronotum. Elytral sculpture very similar to that of D. banghaasi, but distinctly confluent at anterior and medial borders of the orange spots, which are much smaller than in D. banghaasi. No microsculpture, except for faint traces on tergites 8 and 10. Comments. Dianous socius Zheng is easily distinguished from D. banghaasi Bernhauer by its much smaller elytral spots, generally blackish coloration throughout, and the sexual characters especially the very long apicolateral tooth of sternum 9. These characters also distinctly distinguish D. socius from one male of D. banghaasi from Emei Shan, which has smaller elytral spots than usual (see above). Dianous ocellatus Cameron, 1930 Dianous ocellatus Cameron, 1930: 430 f. fig.; Puthz, 1980b: 10 fig. ? Dianous acutus Zheng, 1994: 479 f. figs. Material examined: Yunnan: 8 dd, 11 9 9: Xishuangbanna, 20./ 24. I. 1993, G. M. de Rougemont; 14: Ruili, 4. II. 1993, G. M. de Rougemont (cR, cP). 4 dd, 2 22: Thailand: Chiang Mai Prov., Doi Suthep, 750- 800 m, 2. XI. 1995, Zettel (4) (NHMW, cP). Comments. These specimens differ from Assam specimens (types) in having a larger elytral spot, but sexual characters are almost the same (see figs 177, 182, 183, 196, 202, 203) (number and strength of apicomedial sclerotized fringes of sternite 9 are somewhat different). Dianous ocellatus was described from India (Assam) and is also known from Vietnam. New to China and Thailand. Zheng described Dianous acutus based on specimens from Sichuan. Regarding the figures given by Zheng, this taxon seems to be very close to D. ocellatus. Verification by the compa- rison of the types is necessary. Unfortunately, my attempt to get the types for study was not successful. Dianous ocellifer sp. n. Type material. Holotype (male) and allotype (female) and 1 ?- PT: Yunnan: Ruili, 4. I. 1993, G. M. de Rougemont (cR in BMNH, cP). 1 4,2 9 ?- PTT: N Laos: Prov. Lg Nam Tha, ca. 20 km SE Muang Sing, 950 m, 12.- 13. VI. 1996, Schillhammer (25)(NHMW, cP). 1 d: Laos C.: Bolikhamsai Prov., Ban-Nape-Kaow-Nua-Pass, 18. IV.- 1. V. 1998, Jendek & Sausa (cHromadka). This species closely resembles D. ocellatus Cameron. Moderately shining with blue metallic and/or coppery tint, each elytron with a large orange spot (resembling that of D. grandistigma). Head moderately finely and densely punctate, punctation of pronotum moderately coarse and dense, punctation on basal third of elytra about as coarse as that on pronotum, rest of elytra with dense, oblique sculpture of long rugae, abdomen very finely and densely punctate. Pubescence distinct, whitish yellow, that of fore parts semierect, that of abdomen very distinct, recumbent. Antennae blackish, bases of individual segments remarkably reddish, club THE GENUS DIANOUS IN CHINA 479 brownish. Palpi blackish. Trochanters reddish, legs black with metallic tint, tarsi some- times blackish brown. Clypeus and labrum blue metallic, densely pubescent. Fourth tarsal segment deeply bilobed. Length: 4.8- 5.8 mm (fore parts: 2.9 mm). PMrofithe AL =wE: 4S"adE 27 1B 19312 55WP 333: 1B23533WwEl23321E1733; IS: 45. Male: Medial area of metasternum moderately coarsely and densely punctate, interstices reticulated, moderately shiny. Sternites 3-6 slightly more sparsely punctate medially than laterally, sternite 6 with a very shallow impression posteriomedially, sternite 7 with a distinct medial impression on posterior half, punctation and pubes- cence dense, posterior margin with distinct emargination; sternite 8 with a deep subtriangular excision in about posterior third; sternite 9 with a blunt tooth apico- laterally (fig. 207), fields of fine striae apicomedially, apicolateral brush moderately strong, as long as sternite. Tergite 10 broadly rounded. Aedeagus (figs 180, 189) broad, apical portion of median lobe broadly lanceolate, densely set with strong and very long setae ventro-laterally, internal structures similar to those of D. distigma. Parameres somewhat exceeding median lobe, evenly set with about 22 long setae. Female: Ventral characters similar to those of male. Sternite 7 with more finely and more densely punctate posteriomedially. Sternite 8 (fig. 198). Valvifer with a strong apical brush, as long as the valvifer. Tergite 10 slightly subtriangular, not as rounded as in D. ocellatus Cameron. Head with strongly elevated medial and lateral portions of frons, diameters of punctures nearly as large as basal cross section of third palpal segment, interstices on half of the frons about as large as half diameters of punctures, on other half of frons (particularly posteriorly) about as large as diameters of punctures, rarely larger, no distinct shining area. Antennae very long, by about four segments extending beyond posterior margin of pronotum when reflexed. Pronotum with deep oblique impressions on posterior half and with transverse impression near anterior margin, rest of surface comparatively even (e. g. no median impression); punctation regular, moderately coarse, diameters of punctures somewhat smaller than basal cross section of third antennal segment (but distinctly larger than those on frons), interstices mostly smaller than half diameters of punctures, with a small shining area on very middle. Elytra similar to those of D. ocellatus, but orange spot larger than in the typical form (as large as in the Yunnan-populations), no vortex of rugae on posterior quarter. Abdomen similar to that of D. ocellatus. Legs moderately slender, metatarsi 5/7 as long as metatibiae, first segment distinctly shorter than following three segments combined, somewhat longer than last segment, third segment broad, shortly bilobed, fourth segment deeply bilobed. Groundsculpture of fore parts nearly obsolete, that of abdomen distinct. Comments. Dianous ocellifer may be distinguished from D. ocellatus Cameron by the male sexual characters: in D. ocellatus, the posteriomedial impression of sternite 7 is much shallower and the posterior emargination is much shallower, the notch of sternite 8 is found in posterior quarter, the apicolateral brush of sternite 9 is nearly as long as the sternite, the aedeagus is much larger (compare that of D. ocellifer, fig. 177), 480 VOLKER PUTHZ the apical portion of the median lobe (fig. 189) is not lanceolate and is differently setose. The two species are so close in the sculpture of the exosceleton that good distinguishing characters cannot be given at the moment (punctation of elytra in posteriolateral quarter seems to be less vorticose, punctures more distinctly well- defined); the elytral spot is larger than in the type specimens from Assam, as large as in Yunnan specimens. Etymology: ocellifer (Latin) = bearing little eyes. Dianous grandistigma sp. n. Type material. Holotype (male) and allotype (female): Yunnan: Xishuangbanna, 24. I. 1993, G. M. de Rougemont (cR in BMNH). Paratypes: 15 8 4,6 £ 9: Xishuangbanna, 20. and 24. I. 1993, G. M. de Rougemont (cR, cP). This new species closely resembles D. distigma Champion and D. variegatus sp. n.. The female has very striking characters on valvifer. Moderately shining with faint blue metallic tint, elytra each with large orange spot posteriomedially (fig. 171). Head moderately finely to moderately coarsely and densely punctate; pronotum coarsely and very densely punctate, rugose anterio- and posteriomedially, medial area more or less impunctate, shining; elytra about as coarsely punctate as pronotum, densely rugose on posterior half, vorticose at spots; abdomen very finely and densely punctate. Pubescence of head and elytra whitish, moderately long, semierect, that of abdomen very dense recumbent. Antennae blackish brown, club brownish, palpi dark brown, trochanters reddish, legs black with metallic tint, tarsi sometimes blackish brown. Clypeus and labrum blue metallic, very densely pubescent. Fourth tarsal segment deeply bilobed. Length: 5.0 — 6.5 mm (fore parts: 2.8 — 3.0 mm). PMiof Hl? wH:45:5;:adE:28; IE? 19:172\6;) wP 36 MP-STMERSSAIERSSE 15747. Male: Medial area of metasternum moderately broadly flat, with a narrow medial impression, punctation of medial area moderately coarse and dense, punctures nearly as large as basal cross section of second palpal segment, interstices one to two times as large as diameters of punctures, densely isodiametrically reticulated and shiny, acam very finely and moderately densely punctate and pubescent, interstices with very shallow groundsculpture; declining lateral areas of metasternum finely and moderately sparsely punctate, interstices nearly smooth. Sternites 3-6 slightly more sparsely punctate medially than laterally, interstices one and one half to two times as large as diameters of punctures, shining, groundsculpture nearly extinct; sternite 6 very shal- lowly impressed posteriomedially, sternite 7 with a distinct medial impression on posterior three quarters, punctation moderately fine and very dense, pubescence long, dense, convergent, posterior margin distinctly but shallowly emarginate. Sternite 8 with a deep subtriangular notch in about posterior quarter; sternite 9 with a distinct tooth apicolaterally as in fig. 205, distinct sclerotized fringes and fields of asacl apico- medially, apicolateral brush moderately strong, slightly shorter than sternite. Tergite 10 broadly rounded. Aedeagus (figs 178, 186), median lobe triangularly narrowed and strongly curved upwards, dorsally with a longitudinal ridge on each side, apical THE GENUS DIANOUS IN CHINA 48] portion densely set with moderately long setae inserted mainly dorso-laterally, internal structures similar to those of D. distigma. Parameres distinctly longer than median lobe, evenly set with (about) 20-22 long setae. Female: Punctation of metasternum as in male, but interstices very shallowly reticulated, shiny. Punctation of sternites as in male, sternite 7 finely and very densely punctate and pubescent posteriomedially. Sternite 8 as in fig. 194. Valvifer (fig. 192) narrowly rounded apically, dorsal side smooth and shiny, ventral side densely set with moderately long, very strong, modified setae (comblike, fig.191). Tergite 10 slightly subtriangular. In most respects very similar to D. distigma Champion, but the elytral spot considerably larger, elytra shorter, and the male’s sexual characters different: the flattened median area of the metasternum is distinctly less broad and its posterior middle punctate, the aedeagus narrower, its apical portion shorter (see figs 181, 187). Valvifer totally different. Comments. This species is the first one observed with such conspicuous charac- ters on valvifers; these characters make it easy, to distinguish it from other species similar in exosceletal habitus. — One male in the Zoological Museum in St. Petersburg from Vietnam: Mountains near Tuan Giao, 800 m, 28. XI. 1962, G. Kabakov has also larger elytral spots than those of the Himalayan specimens of D. distigma, and with the aedeagus strongly resembling that of D. grandistigma. This specimen (not at hand) may belong actually to D. grandistigma. Etymology: grandis (Lat.) = large, stigma (Lat.) = a mark. Dianous variegatus sp. n. Type material. Holotype (à ), allotype (2), and 1 © - paratype: Yunnan: Ruili, 4. II. 1993, G. M. de Rougemont (cR in BMNH, cP). This new species is the sister species of D. grandistigma sp. n. based e. g. on the modified setae on the valvifer. It also closely resembles D. verticosus Eppelsheim. Variegate metallic (blue-violet-coppery), rather shining, elytra each with a large orange spot posteriomedially (about as large as length of antennal segment 4). Head moderately finely and moderately densely punctate, pronotum coarsely and very densely punctate (except centre), somewhat coalescent anteriorly and posteriorly; punctation of elytra about as coarse as that on pronotum, but vorticose on posterior half; abdomen very finely and densely punctate. Pubescence, antennae, palpi, and legs same as in D. grandistigma. Length: 5.5-6.5 mm (fore parts: 3.0- 3.1 mm). PM of the holotype: wH: 48; adE: 29; wP: 37,5; IP: 39.5; wEl: 57,5; IEl: 57.5; IS: 50. Male: Ventral characters similar to those of D. grandistigma, but punctation of metasternal disc distinctly finer and slightly sparser, reticulation shallower (interstices strongly glistening), diameter of punctures distinctly smaller than basal cross section of palpal segment 2, interstices often more than two times as large as diameters of punctures. Interstices between the punctures on sternites with very faint but distinct 482 VOLKER PUTHZ reticulation. Sternite 8 with a broad and deep triangular excision in about posterior quarter (length of sternite: depth of excision = 92: 21). Sternite 9 (fig. 204). Tergite 10 broadly rounded. Aedeagus (fig. 184) in general as in D. grandistigma, apical portion of median lobe broader, no dorsal ridge at apical portion. Female: Punctation of metasternum finer and sparser, more strongly shining than that of D. grandistigma. Sternite 8 (fig. 193), less sharply triangular posteriorly. Valvifers about as in D. grandistigma, also with long and strong modified setae; setae not as densely set as in D. grandistigma, somewhat narrower and much more acute on central portion (modified setae at posterior border of valvifer rounded apically as in D. grandistigma). Tergite 10 subtriangular (fig. 201). Comments. D. variegatus is more shining than D. grandistigma and more brightly variegate-metallic. It is also distinguished by its sexual characters, although they are close to those of D. grandistigma. Since nothing is known about variability of that species, the differences may fall within the variability of D. grandistigma, or be of subspecific rank.- The new species also resembles externally the variable D. verticosus Eppelsheim, but the elytral spot is larger, the apical portion of the median lobe is narrower (compare fig. 185), sternite 8 of the female is less triangular (fig. 195) and the valvifer is different (in D. verticosus all the setae are pointed). Dianous emarginatus Zheng, 1993 Dianous emarginatus Zheng, 1993: 201 figs. Material examined: 1 d, 1 2: Yunnan: Heishui, 35 km N Lijiang, 27°13’ N, 150°19’ E, 1.-19. VII. 1993, E. Jendek (NHMW, cP); 4 d 4,3 9 2: ibidem 18. VI.- 4. VII. 1993, S. Becvar (NHMG, cP); 1 2: Lugu Lake — Luo Shui, 27°45’N, 100°45’ E, 8.- 9. 7. 1992, E. Jendek (NHMW). 1 G-PT: Sichuan: Emei Shan, 1600 m, 30. VII. 1982, Tian Hua (cP) [HT and 3 PT in the Dept.Biol., Sichuan Teacher’s College, Nanchong]. — 1 ® (cf. det.): Burma: Carin Asciuii Cheba, 800- 1400 m, XII. 1887, L. Fea (coll. Fauvel, IRScNB). This species is allied and closely similar to D. distigma Champion. Moderately shining, black with faint blue metallic tint, elytra each with a moderately large orange spot posteriomedially (compare fig. 172). Head moderately finely and densely punctate, pronotum coarsely and densely, near anterior and posterior borders rugosely, punctate, elytra punctate about as coarsely as pronotum, each with close oblique rugae on posterior half, abdomen very finely and densely punctate. Pubescence, except on pronotum, distinct, especially on abdomen, whitish, semierect on fore parts, recumbent on abdomen. Antennae blackish brown, club brownish, palpi dark brown, trochanters reddish, legs blackish with metallic tint, tarsi occasionally dark brown. Clypeus and labrum black with bluish tint, densely set with short setae. Fourth tarsal segment narrowly and deeply bilobed. Length: 4.5 — 5.5 mm (fore parts: 2.6 mm). PM of a Yiinnan- d: wH:41; adE:26; IE: 18; IT: 5; wP: 32: IP: 31; wEl: 48; IE: 49.5; IS: 42. Male: Medial area of metasternum broadly flat, with a short impression posteriomedially, impunctate around impression, otherwise moderately finely and den- sely punctate; interstices distinctly reticulated, diameters of punctures nearly as large as THE GENUS DIANOUS IN CHINA 483 basal cross section of second palpal segment, interstices nearly as large as half diameters of punctures, acam finely and moderately sparsely punctate, reticulation of interstices even shallower than that of medial area; declining lateral areas of meta- sternum finely and moderately densely punctate, interstices very shallowly reticulated, shining. Sternites 3-6 slightly more sparsely punctate medially than laterally, interstices nearly twice as large as diameters of punctures, sternite 7 with a distinct but shallow, finely and very densely punctate and pubescent median impression in posterior two fifths, posterior margin shallowly emarginate; sternite 8 with a deep subtriangular notch in about posterior fifth; sternite 9 with a short, blunt tooth apicolaterally (fig. 206), fields of fine striae apicomedially, apicolateral brush short, about half as long as sternite. Tergite 10 broadly rounded. Aedeagus (figs 179, 188), median lobe broader than that of D. distigma (figs 181, 187), apical portion densely set with very long setae ventro-laterally, internal structures consisting of expulsion bands, a strongly sclerotized tube-like internal piece, and a complex of densely spinose membranes. Parameres slightly exceeding apex of median lobe, evenly set with about 16- 22 long setae (much longer in D. distigma). Female: Ventral characters similar to those of male, interstices of medial punctation of metasternum slightly larger, nearly as large as diameters of punctures, sternite 7 simple, apical emargination nearly imperceptible. Sternite 8 (fig. 197). Valvifer with a distinct tooth apically, apical brush 2/3 as long as valvifer. Tergite 10 evenly rounded apically. In most respects very similar to D. distigma: Head with strongly elevated medial and lateral portions of frons, diameters of punctures nearly as large as basal cross section of third palpal segment, interstices mostly smaller than half diameters of punctures, small impunctate area posteriorly near inner eye margin, medial portion of frons slightly less densely punctate than rest of frons, but interstices rarely as large or slightly larger than diameters of neighbouring punctures, no distinct shining area. Antennae very slender, by 3-4 segments extending beyond posterior margin of pro- notum when reflexed. Punctation of pronotum slightly less dense than that of D. di- stigma, diameters of punctures as large as basal cross section of third antennal segment, interstices mostly distinct, nearly as large as half diameters of punctures, larger in very middle, which is shiny and slightly impressed; punctation transversely confluent near anterior and especially near posterior border. Elytra less large than those of D. distigma, punctation on anterior half mostly well-defined, interstices regularly larger than diameters of punctures anteriomedially; posterior half with dense, obliquely-rugose sculpture, rugae somewhat convex at orange spot, but not distinctly vorticose. Abdomen and legs as those of D. distigma, metatarsi about 2/3 as long as metatibiae, first segment distincty shorter than three following segments combined, slightly longer than last segment, third segment shortly bilobed, fourth segment deeply bilobed. Entire body with very shallow reticulation. Comments. Dianous emarginatus Zheng differs from D. distigma Champion by its narrower body and by the sexual characters; it differs from D. grandistigma sp. n. by the much smaller elytral spot, by the shorter antennae, and by the sexual characters. 484 VOLKER PUTHZ Dianous luteolunatus Puthz Dianous luteolunatus Puthz, 1980b: 3 figs. Material examined: SE Guanxi: 1 9: District Yulin, Liuwan Mountains SW Yulin, Kuishan 300 m, 17. XI. 1993, H. Schillhammer (22) (NHMW). Laos: 17 88,12 2 2, Prov. Lg. Nam Tha, ca. 20 km SE Muang Sing, 12./13. VI. 1996, Schillhammer (25) (NHMW, cP); 2 GG, 4 2 9, Prov. Lg. Nam Tha, ca. 30 km NW Lg. Nam Tha, 800 m, 16.-18. VI. 1996, Schillhammer (28, 30) (NHMW); 1 ©, Bolikhamsai prov., Ban- Nape- Kaow- Nua- Pass, 18. IV.- 1. V. 198, Jendek & Sausa (cHromadka). Comments. This species was described from a single male from Vietnam (Mountains NO Thai Nguyen), new to China and Laos. Dianous atrocoeruleus sp. n. Type material. Holotype (male) and allotype (female): Yunnan: 100 km W.Kunming, Diaolin Natural Reserve, 22. V.-2. VI. 1993, E. Jendek & O. Sausa (NHMW). Paratypes: 2 dd, 2 2 2: same data as holotype. This new species is closely related to the orange spotted species D. luteolunatus Puthz and D. luteostigmaticus Rougemont, but differs from both easily by the uni- colorous surface of the elytra. Black with Prussian-blue metallic tint, rather dull, head moderately finely and densely punctate, pronotum coarsely and mostly densely punctate, elytra coarsely sculptured, each with long rugae directed on anterior half from suture posterio-laterad toward middle of elytron and from there posterior-mediad toward suture to form rugose cross with its center in middle of combined elytra; punctation on anterio-lateral areas of elytra mostly separate, on posterio-lateral quarters mostly shortly confluent the fore parts moderately erect, that of abdomen recumbent. Antennae, maxillary palpi and legs black with blue metallic reflection, bases of trochanters reddish, last antennal segments may be slightly brownish. Clypeus and labrum blackish-blue metallic, densely pubescent. Tarsal 4 segment simple with some weak shoe-bristles. Length: 4.8 — 6.0 mm (fore parts: 2.8 — 3.1 mm). PM of'the HT: wH: 40; adE: 26.5; IE: 17: 17: 6; wP: 33: IP? 342 whi ssalE iow: IS: 48. Male: Metasternum strongly convex, narrow triangular medial area flat, extre- mely densely and deeply reticulated but with a faint reflex, impunctate in middle, laterally with about 10 moderately fine punctures, acam with a few moderately fine punctures, densely but much more shallowly reticulated and slighty more shining than medial area; declining lateral portions of metasternum finely and very densely punctate and pubescent, interstices shiny with shallow ground sculpture. Medial punctation of sternites 3 and 4 somewhat coarser and slightly sparser than lateral punctation; medial punctation of sternites 5 and 6 nearly same as that on sides, posterior margins of ster- nites 3 and 4 shallowly emarginate; sternite 7 distinctly but shallowly impressed poste- riomedially, finely and very densely punctate and pubescent, posterior margin with a distinct emargination. 8" sternite with a round and deep emargination nearly in posterior third; sternite 9 broad, apicolaterally with more or less distinct tooth apico- laterally (fig. 101), with strong sclerotized fringes and fields of fine asacl apico- THE GENUS DIANOUS IN CHINA 485 medially, apicolateral brush strong and dense, nearly as long as sternite. Tergite very broadly rounded. Aedeagus (fig. 85), anterior portion of median lobe broadly lanceolate, densely set with long ventral setae anteriolaterally, internal structures consisting of a stronger sclerotized, broad tube and densely spinose membranes. Para- meres somewhat longer than median lobe, densely set with 29 moderately strong, long setae. Female: Ventral characters similar to those of male, but sternite 7 simple, except for very shallow apical emargination; sternite 8 (fig. 88). Valvifer more or less obliquely truncate apically, apical brush about as long as valvifer. Tergite 10 slightly triangularly narrowed, apex rounded. Punctation of head moderately fine, diameters of punctures nearly as large as basal cross section of third palpal segment (thus much smaller than basal cross section of third antennal segment), interstices mostly smaller than half diameters of punctures, slightly larger on elevated median portion and posteriolaterally, but no distinct shining areas. Antennae moderately slender, by about two segments extending beyond posterior margin of the pronotum when reflexed, outer segments slightly longer than broad. Pronotum with a deep, transverse impression near anterior margin and deep, oblique lateral impressions on posterior half; punctation deeply pierced, mostly dense, dia- meters of punctures as large as basal cross section of third antennal segment, interstices smaller than half diameters of punctures, much larger posteriomedially on a shining, but finely reticulated area, of which the width possibly as large as length of antennal segment eighth; punctation somewhat confluent near anterior margin and in posterior impressions near base. Elytra strongly rugose, rugae may form more or less distinct vortex latero-medially. Abdomen with broad, distinctly upwards directed, paratergites, those of segment four about as broad as metatibiae in apical third, punctation of paratergites fine and very dense, nearly as fine as that of tergites. Tergite 7 with a broad apical membranous fringe (winges fully developed). Punctation of tergites 8 and 10 twice as coarse and much sparser than that of tergite 7. Legs moderately slender, metatarsi 2/3 as long as metatibiae, first segment distinctly shorter than three following segments combined (33: 38), longer than last segment (33: 28), fourth segment simple. Entire body densely, more or less deeply reticulated. Comments. Dianous atrocoeruleus. is similar in general appearance to D. luteo- lunatus Puthz and D. luteostigmaticus Rougemont, but differs by the lack of the orange elytral spots and by the different sexual characters.- The allotype has some attached mites. Etymology: ater (Latin) = black, coeruleus (Latin) = blue: blackish blue Dianous. Dianous alcyoneus sp. n. Type material. Holotype (4) and allotype (9) and 1 d, 2 9 - paratypes: N Laos: Prov. Lg. Nam Tha: ca. 30 km SE Muang Sing, 800 m, 16./ 18. VI. 1996, H. Schillhammer (28, 30) (NHMW, cP). This new species is the sister species of D. atrocoeruleus sp. n.. 486 VOLKER PUTHZ Moderately shining, black with Prussian-blue reflection, elytra with a silver- bluish spot posteriolaterally, frons moderately coarsely and densely punctate, pronotum coarsely, densely and somewhat confluently punctate, elytral sculpture coarse, rugose, vorticose, abdomen finely and very densely punctate; pubescence of the fore parts indistinct, on abdomen short, dense, recumbent. Antennae, maxillary palpi and legs black with some metallic reflection. Clypeus and labrum blackish blue metallic, mode- rately densely pubescent. Length: 4.7 — 5.6 mm (fore parts: 2.6 — 2.8 mm). PM of HT: wH: 40; aE: 24; IE: 17; IT: 6; wP: 34; IP: 33; wEl: 55; IEI: 58; IS: 49. Male: Metasternum strongly convex, narrow triangular median area flat, extremely deeply and densely reticulate, completely dull, with a few moderately fine punctures, acam finely and densely punctate throughout with shallow microsculpture (much more shining than median area of metasternum). Sternit 3 with a triangular impunctate area medially, sternite 4-5 slightly coarser and slightly less densely punctate medially than laterally, sternite 6 densely punctate medially, sternite 7 very densely punctate and pubescent posteriomedially (no distinct impression), posterior margin very shallowly emarginate. Sternite 8 with a broad notch in about posterior sixth. Sternite 9 serrate apically (fig. 241) with a small area of asacl posteriomedially, apical brush strong, as long as the sternite. Tergite 10 broadly rounded. Ae deagus resembling that of D. atrocoeruleus (fig. 85; also very similar to that of D. luteostigmaticus Rougemont) but the apical portion of the median lobe different (fig. 243); internal structures similar to those of the compared species, parameres with 21 setae. Female: Ventral characters as in the male, but sternite 7 simple. Sternite 8 broadly rounded and slightly bent upwards posteriorly. Tergite 8 with a broad excision apically (fig. 242). Valvifer serrate apically, brush very strong. Tergite 10 triangular. Comments. In nearly all respects similar to D. atrocoeruleus but the sexual characters different: sternite 8 of the male is much less emarginate, aedeagus slightly but distinctly different, tergite 8 of the female has an apical excision. In my key (1981) the new species should be placed at # 150 (151): it is distinguished from D. vietnamensis Puthz by the larger size, by the shorter antennae with broader penultimate segments and by the sexual characters; in the key of this article it might be placed either at # 204 or at # 214 (it is distinguished from D. subvorticosus by lacking dense pubescence of the fore body). Etymology: Alcyoneus (Latin) = resembling a kingfisher by its silver-bluish elytral spot. Dianous oculatipennis Puthz Dianous oculatipennis Puthz, 1980b: 3 ff. figs. Material examined: Yunnan: 2 d d, 2 9 9: Xishuangbanna, 20./ 24. I. 1994, G. M. de Rougemont (cR, cP). Male: Metasternum flattened medially, narrowly impressed posteriomedially, punctation of disc moderately coarse and sparse, diameters of punctures nearly as large as basal cross section of palpal segment 2, interstices two to three times as large as THE GENUS DIANOUS IN CHINA 487 diameters of punctures, densely and deeply reticulated, weakly glistening (neither shining nor dull), acam very finely and sparsely punctate, interstices nearly smooth. Sternites 3-6 medially as fine and sparse punctate as on the sides, punctures much smaller than eye facets, interstices twice as large as punctures, groundsculpture nearly extinct. Sternite 5 and 6 very shallowly emarginate at posterior margin, sternite 7 with shallow but distinct impression posteriomedially, punctation and pubescence of impression fine and very dense, posterior margin shallowly emarginate. Sternite 8 with a broad, triangular notch nearly in posterior third. Sternite 9 (fig. 208). Tergite 10 broadly rounded. Aedeagus (fig. 190). Female: Punctation of metasternum as in male, but interstices narrower (although distinctly larger than diameters of punctures), and shallowly reticulated, shining. Sternites as in male, sternite 7 with medial punctation and pubescence denser than that on sides. Sternite 8 triangular (fig. 199). Valvifer irregularly serrate apically, apical brush strong, nearly as long as valvifer. Tergite 10 rounded, not subtriangular. Comments. This species was described from a single male from Yunnan (Siamonyang, 25 km N Ch’e-li). Dianous lobatipes sp. n. Material: Holotype (3): Anhui: Huang Shan, 60 km NNW Tunxi nr. Tang Kou, 900- 1000 m, stream (beneath cable car), ca. 3-5 m wide, almost completely dried out, surrounded by primary forest, very big granitic rocks and sand, short sections with running water (figs 17, 20 in Jäch & Li, 1998), 31. X. 1997, M. Wang (CWBS 292) (NHMW). Moderately shining, head blue-metallic with olive tint, rest of insect olivegreen- metallic, head moderately finely, densely punctate, pronotum and elytra coarsely and mostly confluently punctate, abdomen very finely and very densely punctate; pubes- cence short, recumbent, conspicuous on abdomen. Antennae, maxillary palpi and legs dark green-metallic, last segments of antennae and last tarsal segments slightly brow- nish. Clypeus and labrum blue-metallic, moderately densely pubescent. Fourth tarsal segment very deeply bilobed. Length: 5.0- 5.8 mm (fore parts: 2,7 mm). Male: Metasternum broadly convex, triangularly impressed posteriomedially, median portion shining, almost smooth, about 10 moderately fine punctures on each side, acam smooth. Sternites 3-6 almost impunctate medially with faint microsculpture, shining median area about 1/4 as broad as the respective sternite, not sharply delimited laterally, transition zone between impunctate and densely punctate areas sparsely and moderately finely punctate. Sternite 7 finely and densely punctate with dense pubes- cence posteriomedially, posterior margin very shallowly emarginate. Sternite 8 with a broad, rounded notch in about posterior 6th. Sternite 9 apically serrate (fig. 238), apicolateral brush strong, distinctly somewhat shorter than length of sternite. Tergite 10 broadly rounded. Outline of aedeagus resembling that of D. rugosipennis (see figs 84, 86), but internal structure totally different: no strongly sclerotized basal portion of internal sac but a broad and long membranous sac densely set with tufts: stronger sclerotized lateral portions of the apical portion narrower than in the compared species with few short setae (fig. 224); parameres with 15- 16 long setae in apical half. 488 VOLKER PUTHZ Female: unknown. Head slightly broader than elytra (45: 43,5), eyes large (IE: 21; IT: 5), frons broad (adE 27) with distinct, complete longitudinal furrows, median portion about as broad as each of the side portions, slightly but distinctly convex, elevated to about level of inner eye margins; punctation moderately fine and dense, diameters of punctures nearly as large as basal cross section of segment 3 of the maxillary palpi, interstices mostly distinctly smaller than diameter of punctures, larger at few places. Antennae very long, last four segments extend beyond the posterior margin of the pronotum when reflexed, penultimate segments 1.5 x as long as broad. Pronotum slightly longer than broad (36: 33,5), sides convex anteriorly, concave in posterior third, a distinct round impression on each side of the middle; punctation coarse, coarser anteriorly and posteriorly, less coarse on middle, very dense and rugose anteriorly and posteriorly, dense laterally or moderately dense on actual middle, largest punctures about as large as medial cross section of antennal segment 3, smallest punctures (on the centre) about as large as basal cross section of antennal segment 8, interstices mostly much narrower than half diameter of punctures, as large or somewhat larger than punctures on medial centre. Elytra rectangular, slightly narrower than head (43,5: 45), distinctly longer than broad (48: 43,5), no distinct impressions; sculpture coarse, dense and rugose, rugae slightly vorticose on posterior half, about as broad as medial cross section of antennal segment 3, interstices smaller than half width of rugae; declining sides of elytra coarsely or moderately corarsely punctate, shining interstices as large as diameter of punctures or larger. Abdomen with broad, upwards directed paratergites, basal furrows of tergites very deep, tergite 7 with a broad membranous fringe apically (the insect 1s fully winged); punctation very fine and very dense, tergite 8 moderately finely and moderately densely punctate, interstices as large as diameter of punctures or larger, tergite 10 with some moderately fine punctures. Legs slender, metatarsi longer than half metatibiae (35: 58), first segment slightly shorter than the following 3 segments combined (14: 16), slightly longer than the last segment (14: 12); segment 4 narrowly and deeply bilobed, lobes as long as half of the claw segment. Faint microsculpture on frons, abdominal microsculpture distinct on tergite 8-10. Comments. This new species is remarkable by the deeply bilobed fourth tarsal segment, narrow general facies and apically serrate tergite 10. There are several species which have resembling aedeagi (D. /uteostigmaticus Rougemont, D. rugosipennis sp. n.), but the internal structures are different from that of the new species. So, at present, I do not Know its sister- species. Tentatively the new species should be placed into the complex of Dianous coerulescens. Etymology: lobus (Latin) = lobe, pes (Latin) = foot. Dianous cupreogutta sp. n. Type material. Holotype (d ): Taiwan: Nantou Hsien: Shanlinchi, 1650 m, 19. V. 1991, A. Smetana, on wet rocks in a creek (see photo 2 in Smetana, 1995), A. Smetana (MHNG); allotype (9 ): same, 16. V. 1990, A. Smetana (cS). Paratypes. 1 d, 1 9: same data as HT and AT (cS, cP). Kaohsiung Hsien: 1 4: road from Tengchih to Chuyenshan, 1400 m, 25. IV. 1990, A. Smetana (cS). THE GENUS DIANOUS IN CHINA 489 This new species is remarkable by the dark, brilliant general habitus and by the coarse, rugose sculpture of the elytra, which have a small coppery spot on the posterior half. It resembles D. championi Cameron but is closer related to D. cameroni Champion. It also resembles in general appearance the west Palaearctic Stenus asphal- tinus Erichson. Brilliant black with slight blue metallic tint (head anteriorly, pronotum laterally, tip of abdomen), elytra on posterior half with a small, sometimes indistinct, coppery spot. Punctation of head and pronotum moderately coarse, moderately dense (frons) to moderately sparse (pronotum), elytra coarsely, rugosely sculptured with long vorticose rugae on posterior half, abdomen very finely and sparsely punctate except last 3 tergites. Pubescence rather long, moderately recumbent, that of abdomen quite distinct. Antennae black, club somewhat blackish brown. First segment of maxillary palpi light brown, second segment brown, third segment dark brown. Legs black, trochanters brownish, tibiae and tarsi with brownish tint. Clypeus and labrum dark blue metallic, sparsely pubescent. Segment 4 of metatarsi short, slightly bilobed. Length: 3.8 — 4.8 mm (fore parts: 2.3 — 2.5 mm). PM of Ale wheal adE 22 %IE 19:11: 455 wP230.IP23lawEl: 4142: 15237. Male: Middle of metasternum plain, moderately impressed posteriorly, brilliant, impunctate medially, with 8-9 moderately coarse punctures on each side, acam impunctate. Abdominal sternites 3-6 medially finely and very sparsely punctate, punc- tation slightly sparser than that on sides, sternite 7 shallowly impressed posterio- medially, finely and very densely punctate and pubescent, posterior margin shallowly emarginate. Sternite 8 with a broad emargination in about posterior eighths to ninth. Sternite 9 (fig. 228) with a broad tooth apicolaterally, with short sclerotized fringes apicomedially, densely set with rather long setae ventrally, some fields of fine, short striae apicodorsally, apicolateral brush somewhat shorter than sternite. Tergite 10 broadly rounded. Aedeagus (fig. 214) of the „coerulescens-type“, median lobe triangularly narrowed, apical portion with numerous long setae, most of them dorsally. Parameres slender, with about 12-16 long setae. Female: Ventral characters almost as those of male, but apical impression and emargination of sternite 7 shallower. Sternite 8 rounded, obtusely angulated produced medially (about as in fig. 215). Valvifer apically moderately acute. Tergite 10 narrowly rounded. Head about as broad as elytra in male (may be slightly narrower or slightly broader), slightly, but distinctly, narrower in female (e. g. 41.5: 45.5); longitudinal furrows deep and long, middle portion nearly as broad as each of lateral portions, strongly elevated, as high as elevated lateral portions of frons; punctation moderately coarse, moderately dense, diameters of punctures about as large as basal cross section of third antennal segment, interstices mostly as large or slightly larger than diameters of punctures, narrower in longitudinal furrows. Antennae moderately slender, about one and half of the outer segments extending well beyond posterior margin of pronotum when reflexed, penultimate segments about 1.5 x as long as broad. Pronotum with distinct oblique impressions posteriolaterally and with transverse impression near 490 VOLKER PUTHZ anterior margin; posterior third distinctly constricted, with conspicuous bump posterio- laterally; punctation about as coarse or somewhat coarser than that on head, moderately dense laterally, sparse to very sparse medially, interstices 3 or more times wider than diameters of punctures in very middle. Elytra subquadrate, sutural and humeral impressions shallow; punctation on anterior half mostly well- defined (occasionally confluent), on posterior half strongly vorticose forming a more or less distinct rosette around small coppery spot situated directly behind longitudinal middle, diameter of coppery spot about as large as length of antennal segment 5 or 6; the cavity of largest furrows can receive second antennal segment. Abdomen with broad, slightly upward directed paratergites, those of tergite 4 as broad as metatibia at apex, punctation of paratergites dense anteriorly, becoming very sparse posteriad; punctation of tergites 3- 7 very fine and sparse, diameters of punctures smaller than medial eye facets, interstices medially at least three times as large as diameters of punctures; Tergite 7 with a broad palisade fringe apically (the species is fully winged), punctation of tergites 8 and 10 coarser and denser. Legs moderately slender, metatarsi about 3/4 as long as metatibiae, first segment distinctly shorter than 3 following combined (e. g. 32: 35), very slightly longer than last segment, segment 4 slightly broader than segment 3, distinctly but shortly bilobed. Most parts of this insect are brilliant lacking microsculpture, faint groundsculpture can be seen on tergites 6 and 7, distinct reticulation on the last 3 tergites. Comments. | suppose that this species is more variable in the size of the elytra (in relation to that of the head) than described above, as it is the case in D. championi. Dianous cupreogutta may be distinguished from D. championi Cameron by smaller size, the coppery elytral spot, the relatively broader paratergites, by the more sparsely punctate pronotum lacking longitudinal median impression, and by the sexual characters. It differs from D. cameroni Champion, which has a very resembling aedeagus, by the much sparser punctation, the brilliant lustre, by the smaller size, and by the shorter tarsi. Since the degree of bilobation of segment 4 of the metatarsi is not so clear in this species, in keys it should be quoted in both colums: with simple and with bilobed tarsi. Etymology: cupreus (Latin) = coppery, gutta (Latin) = drop: Dianous with coppery drops. Dianous ater sp. n. Type material. Holotype (d) and allotype (9): Taiwan: Taoyuan Hsien, Takuanshan Forest, 1600 m, on wet rocks in a small mountain stream, 17. IV. 1990, A. Smetana ( NHMG). Paratypes: 93 d d, 120 2 2: same data as holotype (cP, cR, cS). Ilan Hsien: 6 d d, 14 9 2: Chyr Duan 1050 m, 18. IV. 1990, same habitat as HT, A. Smetana (cP, cS); 1 4,2 2 2: ibidem, 1100 m, 18. IV. 1990, swampy edge of a small lake, in tufts of Carex, A. Smetana (cS). This species is closely related to D. amamiensis Sawada, which it resembles in general habitus and sculpture, but it is easily distinguished by the blackish legs. At first glance, it seems to represent a species of Stenus near the Palaearctic S. intricatus Er. Moderately shining, upper surface black with very faint dark-green-metallic tint (distinct only on anterior portion of head and on last 3 abdominal segments, more or THE GENUS DIANOUS IN CHINA 49] less distinct also on posteriolateral rugose portion of elytra, ventral side distinctly greenish-metallic. Punctation of fore parts coarse and very dense, coalescent on parts of pronotum and on posterior half of elytra, abdomen finely and very densely punctate. Pubescence of fore parts short and little distinct, abdomen densely pubescent. Antennae blackish with faint metallic tint, penultimate segments brownish. Maxillary palpi with segment | and narrow base of segment 2 brownish, rest black with metallic tint. Legs blackish with metallic tint, femora dark brown in about basal half, trochanters reddish brown. Labrum and clypeus black with dark-green tint, moderately densely pubescent. Segment 4 of metatarsi distinctly, moderately broadly bilobed. Length: 4.7- 5.7 mm (fore parts: 2.8- 3.0 mm). Male: Metasternum broadly convex, very slightly impressed posteriomedially; punctation of flat triangular middle moderately coarse and moderately dense, much coarser than that on declining lateral portions, interstices mostly as large as punctures, with more or less deep, isodiametrical groundsculpture, posteriomedial area narrowly impunctate, acam with some fine punctures. Abdominal sternites 3-6 moderately finely and densely punctate, interstices with more or less distinct groundsculpture; sternite 7 shallowly impressed posteriomedially, impression finely and very densely punctate and pubescent, posterior margin shallowly emarginate. Sternite 8 with a broad triangular excision in about posterior eighth. Sternite 9 (fig. 229) blunt and serrate apicolaterally, apical brush about 2/3 length of sternite. Tergite 10 broadly rounded at posterior margin. Ae deagus (fig. 209) with apical portion of median lobe triangularly narrowed and curved toward ventral side. Female: Ventral characters about same as in male, apical impression of sternite 7 very shallow. Sternite 8 (fig. 215) with rounded projection posteriomedially. Valvifer apically rounded-serrate. Tergite 10 broadly rounded. Length: 4.7- 5.7 mm (fore parts: 2.8- 3.0 mm). Head much narrower than elytra (40.5: 50), eyes large (IE: 20; IT: 4); frons broad (adE: 23) with two distinct longitudinal furrows, medial portion about as broad as each of side portions, distinctly elevated, extending well beyond the level of inner eye margins; sides portions distinctly elevated posteriorly; punctation coarse and dense, diameter of punctures about as large as cross section of basal third of antennal segment 3, interstices distinctly smaller than half diameter of punctures, larger on middle portion and on elevated side portions, with interstices becoming as large as (about) one neighbouring puncture. Antennae moderately long, extending slightly more than by last segment beyond posterior margin of pronotum when reflexed, penultimate segments 1.5 x as long as broad. Pronotum slightly longer than broad (34: 31), broadest at about middle, moderately convex in anterior two thirds, distinctly constricted in posterior third; with moderately deep impression laterally somewhat behind middle, transverse impression near anterior margin, and sharp, very narrow, short, and more or less distinct medial impression at about middle. Punctation coarser than that on head and more or less coalescent, distinctly coalescent on posterior half, diameters of punctures about as large as middle cross section of antennal segment 3, interstices always much smaller than half diameter of punctures. Elytra subquadrate, much broader than head (50: 40.5), slightly longer than broad (53: 50), shoulders prominent, sides shallowly 492 VOLKER PUTHZ rounded, posterior margin deeply emarginate (IS: 44); sutural impression distinct, humeral impression less distinct, shallow. Punctation as coarse or slightly coarser than that on pronotum, very dense and mostly distinctly coalescent on following areas: anteriorly near suture (short transverse rugae in most specimens), on inner third of posterior half there are oblique rugae directed posteriad from elytral middle toward suture, and in lateral two thirds directed straight toward posterior margin or somewhat curved, but not forming a distinct (or complete) rosette. Abdomen with broad para- tergites, those of segment 4 as broad as metatibia at its broadest portion, slightly directed upwards and very finely and densely punctate. Basal impression of basal tergites deep, tergite 7 with distinct palisade fringe apically (wings fully developed). Punctation fine and very dense throughout, punctures about as large as medial eye facets, interstices smaller than diameters of punctures, sternite 8 with punctation dis- tinctly coarser and slightly less dense than that of tergite 7, tergite 10 moderately finely and moderately densely punctate. Legs slender, metatarsi (fig. 220) nearly 4/5 as long as metatibiae, segment | distinctly shorter than following 3 segments combined, much longer than last segment; segment 4 distinctly, moderately broadly bilobed (fig. 220). Entire surface with more or less deep ground sculpture. Variability: In general the punctation of frons is very dense, the very small impunctate areas are completely reduced in numerous specimens. The coalescent areas of elytra are of different sizes: the rugae are apparent only on the middle portion of the posterior half of elytra and are short and very slightly curved (nearly straight) in some specimens, while in others the rugose areas are much more distinct; however there is not One specimen amongst the large type series which completely lacks rugae. One paratype is much deeper microsculptured than the rest of the material. A few (less mature ?) specimens have the legs brownish. Comments. Dianous ater may be easily distinguished from S. amamiensis Sawada (and from the closely related spotted species, as e. g. D. freyi L.Benick) by dark body with dark-greenish-metallic tint, and by blackish legs. Etymology: ater (Lat.) = black. Dianous sucinigutta sp. n. Type material. Holotype (4) and allotype (9): Taiwan: Chiai Hsien: Alishan, Sister Ponds, 2180 m, 18. IV. 1990, in soaking wet moss growing on rotten wood around small ponds in a coniferous forest, A. Smetana (NHMG). Paratypes: 11 d d, 12 9 9: same data as HT. Ilan Hsien: 16: Taipingshan, 1880 m, 14. VII. 1993, in wet moss on large rocks at the base of a vertical wall in a creek. Taoyuen Hsien: 1 8,2 2 2: Takuanshan Forest, 1660 m, 17. IV. 1990; 1 &: same, 1600 m, in moss on rocks in a small creek. Taichung Hsien: 2 4 4, 1 9: Hsiaohsueh Shan, 2650- 2750 m, 1. V. 1990, among vegetation on a small seepage; 1 9: Anmashan, 2120 m, 1.V.1990, in wet moss on a fallen tree in a forest seapage; 1 2: Anmashan, Creek, 2136 m, 12. V. 1992, same habitat as previous. Nantou Hsien: 13 88,7 2 9: Houhuanshan Kuenyang, 3050 m, 27. IV. 1990, in soaking wet moss on a vertical wall; 3 6d, 1 2: same, 4. V. 1991, same as previous; 1 d,2 9 2: same; 10 46,7 99 Houhuanshan, 3175 m, 15. V. 1990, in soaking wet moss at edges of a small creek; 5 dd, 1 9: Maifeng, 2130 m, 3. V. 1991, among vegetation on wet vertical wall; 3 d d: Yushan N.P., Mun- Li Cliff, 2700 m, 13. V. 1991; 5 8 d: same, 18.V.1991; 1 2: SW Slope below Yushan Mountain Peak, 3650 m, 14. V. 1991, debris on wet forest floor; 2 d d: Nankaoshan trail, Yuenhai Hut, 2350 m, 4. V. 1992, sifting lush vegetation along wet edges of a small creek; 2 d d: same, THE GENUS DIANOUS IN CHINA 493 Tenchi Hut, 2900 m, 5. V. 1992, in wet moss on fallen trees along a creek; 1 d, 2 29: Nankaoshan, 1,5 km SW Tenchi Hut, 2830 m, 6. V. 1992, sifting of fallen leaves and moss at the base of a vertical wall (cf. photo fig. 16 in Bousquet & Smetana 1996); 1 ¢: Tenchi Hut, 2895 m, 7. V. 1992, same as previous. Chiai Hsien: 3 d d: Yushan N.P., Ta- Ta Ghia, 2750 m, 27. IV. 1990, in wet moss on fallen trees; 1 9: Alishan, 2300 m, 9. V. 1965, Y. Hirashima (Ent.Lab., Kyushu Univ.). Kaohsiung Hsien: 1 6: Kuanshan, Kuanoshing Hut, 2950 m, 18. IV. 1992; 1 d, 2 2: Creek 2 km E. Tien Chih, Hwy 20, 2400 m, 22. VII. 1993, in soaking wet moss on a seapage near a creek; 1 d: Creek 4 km E. Yakou, 2600 m, 23. VII. 1993, in soaking wet moss on rocks directly in a creek. Hualien Hsien: 1 d, 2 9 2: Taroko N.P., Chungyantianshi (River) waterfall, 2300 m, 10. V. 1990, soaking wet moss and other debris in a spray of a waterfall; 1 d: same, 2280 m, 10. V. 1990; 1 &: Taroko N.P., Nanhushi Hut, 2220 m, 12. V. 1990, wet moss on rocks in a creek, except | all leg. A. Smetana (cS, cP, cR). This new species and the following two are closely related to the complex of D. freyi L. Benick, D. klapperichi L. Benick, and D. amamiensis Sawada, but may be easily distinguished from them by the very broad head and short wings. Shortwinged, shining, black, abdomen with very faint aeneous tint, elytra with a slightly elevated, small orange spot (fig. 175). Punctation of fore parts coarse and dense, more or less coalescent on pronotum, distinctly coalescent on elytra; abdominal punctation moderately fine and moderately sparse (variable: s. b.). Pubescence of fore parts indistinct, that of abdomen sparse. First antennal segment dark brown, segment 2 brown, segments 3-5 (6) light brown to reddish brown, remaining segments slightly infuscate. Maxillary palpi reddish brown, third segment infuscate toward apex. Legs reddish brown to brownish, posterior half of femora (often) slightly infuscate (espe- cially on dorsal side), apical portion of femora narrowly dark brown, tarsi distinctly darker (mostly dark brown) than tibiae. Clypeus and labrum black, sparsely pubescent. Segment 4 of metatarsi moderately deeply and narrowly bilobed. Length: 4.8- 6.4 mm (fore parts: 2.8- 3.1 mm). Male: Metasternum broadly convex, narrowly and slightly impressed posterio- medially, punctation on middle portion coarse and dense, diameters of punctures about as large as basal cross section of segment 2 of the maxillary palpi, interstices mostly as large as half diameter of punctures; narrow triangular area impunctate posteriomedially; acam finely and moderately sparsely punctate. Punctation of basal abdominal sternites moderately fine and moderately dense, about same as that on sides of sternites; sternite 6 posteriomedially with slightly finer and denser punctation than that on sides, posterior margin slightly emarginate; sternite 7 with distinct but not deep medial impression in posterior 2/3, its punctation fine and very dense, posterior margin shallowly emarginate. Sternite 8 with broad, subtriangular notch in about posterior fifth. Sternite 9 (fig. 233), rounded-serrate apicolaterally, apical brush slender, nearly half as long as sternite. Tergite 10 very broadly rounded posteriorly. Aedeagus (figs 210, 211, 218) with median lobe with subtriangular apical portion, its apical „knob“ more or less pro- nounced and slightly curved upwards, parameres much longer than median lobe, each with about 15 long setae. Female: Ventral characters about same as in male, but sternite 7 posterio- medially finely and very densely punctate and pubescent. Sternite 8 slightly, obtusely- rounded, no distinct median projection. Valvifer apically rounded-serrate. Tergite 10 (about) semicircularly rounded at posterior margin. 494 VOLKER PUTHZ Head very large, distinctly broader than elytra (48: 45), eyes large (IE 22), temples moderately long (IT 8), frons broad (adE 27) with two deep longitudinal furrows, medial portion narrower than side portions, distinctly elevated to about level of inner eye margins, lateral portions elevated posteriorly, forming impunctate swelling diameter of which sometimes about as large as length of antennal segment 2. Punctation coarse and dense, diameters of punctures as large as basal to middle cross section of antennal segment 3, interstices mostly smaller than half diameter of punctures, possibly (mostly about as large as diameter of punctures) in middle of frons and on lateral swelling, latter in most cases more distinct than impunctate median area. Antennae moderately slender, extending by last or last two segments beyond posterior margin of pronotum when reflexed, penultimate segments 1,5 x to twice as long as broad. Pro- notum little longer than broad (37: 34), broadest at about middle, sides convex in anterior 2/3, moderately concave posteriorly; with deep, oblique impression directed from sides posteriomediad in posterior half, with a transverse impression near anterior margin, and with short, narrow, and less distinct longitudinal impression just behind middle in most specimens. Punctation coarse, dense and irregular, more or less coalescent anteriorly and posteriorly, diameters of punctures becoming as large as apical cross section of antennal segment 3, interstices mostly distinctly smaller than half diameter of punctures, rarely becoming as large as diameters of punctures. Elytra (fig. 175) narrower than head (45: 48), slightly longer than broad (47,5: 45), shoulders prominent, sides feebly rounded, posterior margin moderately deeply emarginate, with distinct humeral impression. Punctation coarse to very coarse, irregular, coalescent, dense but less dense than on pronotum, interstices often as large as half diameter of punctures, in some areas larger; long and somewhat curved rugae around the slightly elevated orange spot in most specimens present, orange spot coarsely punctate but not rugose. Interstices may become larger near posterior border of elytra, sometimes also near suture, and/or near middle of anterior half of elytra. Abdomen broadly marginate, paratergites of segment 4 as broad as metatibiae at middle (d ) or at apex (9), modera- tely finely and moderately sparsely punctate; basal furrows of first tergites deep, tergite 7 with a broad palisade fringe apically (the insect is winged, but wing length is dis- tinctly reduced in comparison to macropterous species). Punctation variable: modera- tely fine to fine and moderately dense to moderately sparse, interstices on first tergites distinctly smaller than diameters of punctures or distinctly larger than diameters of punctures. In general abdominal punctation on most tergites, especially posterior ones, is moderately sparse with interstices distinctly larger than punctures. Punctures of tergite 7 about as large as one inner eye facet, interstices up to 2 x as large as diameter of punctures. Punctation of tergite 8 slightly coarser and denser, tergite 10 with few fine scattered punctures. Legs slender, metatarsi about 5/7 as long as metatibiae, segment 1 slightly longer than two following segments combined, distinctly but not much longer than last segment; segment 4 moderately deeply and narrowly bilobed (about as in fig. 219). Entire dorsal surface distinctly microsculptured in most specimens, with microsculpture of fore parts shallow, distinct only on a few areas; abdomen mostly more distinctly, although sometimes very faintly, microsculptured, occasionally nearly smooth. THE GENUS DIANOUS IN CHINA 495 Variability: The elytra are slightly broader than head in a few females. The degree of the elevation of the median portion of frons is variable; it is distinctly less elevated than the level of inner eye margins in some specimens, in others it distinctly extends beyond that level. Also the antennae vary somewhat in their length (s. a). The size of the elytral spot normally varies between the two sizes shown in fig. 175 (about half of the specimens have larger spots, about as long as the first segment of mesotarsi, in about the other half the spots are about as long as the second antennal segment); one specimen has larger spots (about as long as third antennal segment), in a few specimens the elytral spot is reduced to the size of two or one punctures and is therefore very indistinct. Middle of the pronotum has rarely a more or less distinct shining area. The most remarkable variability is found in the abdominal punctation, which varies consi- derably even in specimens from the same locality. Comments. This species occurs mostly in seepages or other quite wet habitats in coniferous, or mixed broadleaved + coniferous forests; it seems to avoid creeks or other running water (where Dianous species live usually). The specimens were taken by sifting moss, leaf litter, and other debris, sometimes moss on large fallen trees; all these habitats are not the typical Dianous habitats as we know them. The locality below of Yushan main Peak (3650 m) is the highest hitherto known locality for the genus. Dianous sucinigutta may be distinguished from the other two related Taiwanese species, D. sucininotatus Sp. n. and D. electrigutta sp. n. by the smaller size of its elytra spot, from D. sucininotatus also by the in general shorter elytra and less broad head; from D. electrigutta by the different outline of the elytra (rounded, not trapeziform), by the oblique rugae in posterior sutural third, and by the mostly somewhat sparser and finer abdominal punctation. Etymology: The name combines the Latin word for amber and for drop = Dianous with amber drops. Dianous sucininotatus sp. n. Type material. Holotype (3) and allotype (9): Taiwan: Pingtung Hsien: Peitawushan above Kuai- Ku Hut, 2750 m, 29. IV. 1992, sifting of moss and debris on wet habitats along a creek, A. Smetana (MHNG, cS). Paratypes: 2 dd, 2 2 2: same data as HT; 2 9 9: same, 22. IV. 1991, sifting moss and debris on a forest seapage, A. Smetana (cS, cP). In nearly all respects very closely resembling D. sucinigutta and is probably its sister species. Shortwinged, shining, black with faint aeneous tint, elytra with a slightly elevated large orange spot (fig. 174). Punctation of fore parts coarse and dense, somewhat coalescent around elytral spot, abdominal punctation moderately fine and moderately dense. Pubescence of fore parts indistinct, that of abdomen short and sparse. First antennal segment dark brown, second segment brown, segments 3-5 (or 6) light brown, remaining segments more or less infuscate. Maxillary palpi reddish brown, third segment infuscate at apex. Legs reddish brown, apices of femora narrowly darkened, tarsi distinctly infuscate. Clypeus and labrum black, sparsely pubescent. Fourth segment of metatarsi moderately deeply and narrowly bilobed. 496 VOLKER PUTHZ Length: 5.0-6.3 mm (fore parts: 3.0- 3.2 mm). Male: Ventral characters about same as those of D. sucinigutta, median puncta- tion of metasternum slightly less coarse, sternite 6 lacking special characters, impression of sternite 7 very shallow, sternite 8 with a broad, subtriangular notch in about posterior fifth. Sternite 9 rounded-serrate apically, apical brush slender, distinctly shorter than half sternite. Tergite 10 very broadly rounded posteriorly. A e d e agus as in fig. 212, not significantly different from that of D. sucinigutta. Female: As in D. sucinigutta. Head, antennae and pronotum about same as those of D. sucinigutta, but sculp- ture of pronotum less coarse, not as dense, and nearly completely separate (sometimes slightly confluent near posterior margin). Elytra with a very large spot, which can be enlarged inwards at its anterior portion, where it becomes darker/brownish and less distinct. Elytral sculpture less confluent than that of D. sucinigutta, more regular, some oblique rugae running from the spot backwards and inwards, more rugae sometimes present around elytral spot; shiny interstices of punctation often as large as half diameter of punctures. Abdomen and legs as those of D. sucinigutta, metatarsi as in fig. 219. Microsculpture of fore parts mostly indistinct, that of abdomen dense and distinct. Comments. Dianous sucininotatus may be distinguished from D. sucinigutta by the much larger elytral spots, (lighter colored legs), and by the less coarse and less rugose sculpture of the fore parts; from D. electrigutta sp. n. by the longer, rounded elytra with larger spots and by the less coarse and less dense punctation of the entire body. Dianous sucininotatus 1s probably the sister species of D. sucinigutta. It seems to represent this Dianous-type in most southern mountain ranges in southern Taiwan. Etymology: sucini-notatus (Latin) = marked with amber. Dianous electrigutta sp. n. Type material. Holotype (4) and allotype (9): Taiwan: Nantou Hsien: Yushan N.P., 2 km W Pai- Yun Hut, 3350 m, 16. V. 1991, sifting of wet moss on large fallen trees in an Abies- forest, A. Smetana (MHNG). Paratypes: 7 6 4,5 9 2: same data as holotype; 3 d d, 1 9: same but 3528 m, 15. V. 1991, sifting of wet moss, grasses, and bamboo debris along a trickle in a coniferous forest (T 83); 9 dd, 5 9 9: same but 3375 m, 17. V. 1991, same habitat as HT, all leg. A. Smetana (cS, cP). Closely related and similar to D. sucinigutta and D. sucininotatus, but different by sculpture and by the elytral spot. Shortwinged, moderately shining, black with feeble aeneous tint, elytra each with a large, oblong orange spot (fig. 176). Fore parts very coarsely and densely punctate, punctation slightly coalescent on elytra, abdominal punctation moderately coarse and dense. Pubescence of fore parts indistinct, abdomen with short and sparse pubescence. Antennae with two segments blackish to dark brown, third segment (and sometimes a following segments) reddish brown, segments 6 to 8 mostly dark brown, club lighter brown. Maxillary palpi reddish brown, last segment infuscate. Legs reddish brown, apices of femora narrowly infuscate, tarsi remarkably darker than tibiae, nearly THE GENUS DIANOUS IN CHINA 497 blackish. Clypeus and labrum black, sparsely pubescent. Segment 4 of metatarsi moderately deeply and narrowly bilobed. Length: 5.0- 6.0 mm (fore parts: 2.7- 3.0 mm). PMFof Hl? wEl: 47.5: adE: 272 1E A205 AAS: we7355. 12364 wel 445 1Bi: 46; IS: 38. Male: Ventral characters about same as those of D. sucinigutta, but metasternal punctation coarser, sternite 6 simple, sternite 7 with a very shallow impression in posterior half, sternite 8 with a broad, subtriangular notch in posterior fifth. Sternite 9 and tergite 10 as in D. suciniguttasp.n. Aedeagus (fig. 213) very similar to that of D. sucinigutta, apical tip of median lobe seems to be slightly more set off. Female: Ventral characters about same as those of D. sucinigutta. Head in general as that of D. sucinigutta, punctation coarse and very dense, narrow interstices distinctly reticulate, impunctate areas reduced, indistinct. Antennae shorter than in most specimens of D. sucinigutta extending beyond posterior margin of pronotum by more than last segment when reflexed, penultimate segments about 1.5 x as long as broad. Punctation of pronotum as coarse as in D. sucinigutta but denser, interstices generally much smaller than half diameters of punctures, often only few punctures are confluent. Elytra slightly shorter, more divergent, elytral spot oval. Punctation slightly coarser than that on pronotum, mostly somewhat extended longitu- dinally, more or less distinctly coalescent at lateral border of elytral spot and/or between elytral spot and suture. Abdominal punctation in general dense, as dense as in most densely punctured specimens of D. sucinigutta, ground sculpture very dense and deep. Variability: Coalescently punctured areas of pronotum and elytra vary in size; most of the punctation may be clearly separate in some specimens. Comments. D. electrigutta may be easily distinguished from D. sucinigutta and from D. sucininotatus by its trapezoid elytra, by the size and shape of the elytral spot, and by the very dense but relatively rare coalescence of the punctation. Etymology: electrum (Latin) = amber, gutta (Latin) = drop; electrigutta = with amber drops Dianous suciniguttatus Sp. n. Type material. Holotype (é ) and allotype (9 ): Sichuan: Emei Shan, 7. X. 1985, G. M. de Rougemont (cR in BMNH, cP). Paratypes: 2 & 4,2 9 9: Quingcheng Shan 65 km NW Chengdu, 8 km W Taiping, 800-1000 m, 103°33’E, 30°53’N, 18.V./ 3.-4. VI. 1997, A. Pütz (coll. Pütz, cP): 1 4,3 2: ibidem, 18. V. 1997, Wrase (coll Schülke, cP). Closely resembling D. sucinigutta but easily distinguished by the much narrower head and by the shining surface, especially that of abdomen. Shining, black with faint aeneous-coppery tint, elytra with a slightly elevated orange spot (fig. 173). Punctation of fore parts coarse and dense, coalescent posterio- medially on pronotum and around elytral spots, abdominal punctation fine to modera- tely fine and moderately dense, distinctly less dense on middle of tergites; pubescence of fore parts indistinct, abdominal pubescence dense, long, recumbent. First antennal segment dark brown, remaining segments light brown to yellowish brown. Maxillary 498 VOLKER PUTHZ palpi reddish brown, third segment infuscate. Legs light reddish brown, apical portion of femora distinctly infuscate, bases of tibiae darker than apical two thirds, tarsi as light as tibiae apically. Clypeus and labrum black, moderately densely pubescent. Segment 4 of metatarsi moderately deeply and narrowly bilobed. Length: 5.0 — 6.4 mm (fore parts 3.0 mm). PMrof Ai:wH: 462 adE: 25; IE: 21-5: IT: 6: we? 3412437 wE S22 IS: 44. Male: Ventral characters about same as those of D. sucinigutta but punctation mostly coarser, sternite 7 with posteriomedian impression slightly more distinct. Sternite 8 with a broad, triangular notch in about posterior fifth. Sternite 9 (fig. 237) with a prominent, blunt tooth apicolaterally, apical brush slender, about 1/3 as long as sternite. Tergite 10 broadly rounded. Ae de agus (fig. 221) with median lobe with a subtriangular, apically broadly rounded apical portion. Parameres slender, with about 20 long setae. Female: Ventral characters resembling those of male. Sternite 8 apically roun- ded, slightly produced in middle. Valvifers acute apically. Tergite 10 broadly rounded. Head remarkably narrower than that of D. sucinigutta. Impressions of pronotum as those of D. sucinigutta but the punctation less coarse and more equal in size, diameter of punctures about as large as medial cross section of 3" antennal segment, interstices smaller than half the diameter of punctures. Elytra subquadrate with coarse and deep punctation and long rugae around elytral spot. Abdomen similar to that of D. sucinigutta but more convex (parts of sternites are visible in dorsal aspect), punctation of middle portions of tergites distinctly less dense than that of the sides. Entire dorsal surface lacks microsculpture, except for only traces on tergite 8. Comments. D. suciniguttatus is distinguished from D. amamiensis Sawada by the broader head, the much deeper longitudinal furrow of frons, by the deeper impressions of pronotum, the coarser and less dense punctation of the entire body, by the declining, finely and moderately sparsely punctate paratergites, and by the aede- agus. In addition from D. freyi L. Benick by the light color of the legs, which may be found only in immature specimens of D. freyi. Etymology: The name combines the Latin word for amber and for dropped = amber-dropped Dianous. Dianous freyi L. Benick Dianous Freyi L. Benick, 1940: 593 ff.; L. Benick, 1942: 76-79. figs; Puthz, 1971a: 88; Zheng, 1993: 199. Material examined. Zhejiang: T’ein-mu Shan (holotype) (NHMB); 1 d, 1 9: Ning-po (FMCh); Jiangxi: 18 dd, 19 9 2: Kuling near Kiu Kiang, 18. X. 1919, H. F. Loomis (USNM, cP). Fujian: 4 88,2 9 9: Shaowu, 1937, J. Klapperich (FMCh, cP); 1 2: Kuatun, IV. 1938, J. Klapperich (DEI); 10 dd, 10 2 2: Chungan, Bohea Hills, IV. 1940, IV. 1941, T. C. Maa (Bishop Museum, cP): 1 2: Yungan, 11. IT. 1940, T. C. Maa (Bishop Museum); 1 9: Changting: Niuling, 24. IV. 1941, T. C. Maa (Bishop Museum). Anhui: 1 2: Huang Shan, 30 km NW Tunxi, 5 km E Nantang, 350-400 m, 28. X. 1997, M. Wang (NHMW). Hunan: 1 9: Wulingyuan, N.Dayong Zangjiajie, 650 m, 29. X. 1993, H. Schillhammer (1) (NHMW); 1 d: Wulingyuan, N.Dayong Sioxiyu, 400 m, 31. X. 1993, H. Schillhammer (5) (NHMW); 1 d, 5 22: env. THE GENUS DIANOUS IN CHINA 499 Guidong, 26°04’ N,113°56’ E, 26.-31. V. 1994 (NHMW, cP). Guizhou: 1 d, 2 29: Huangguoshu, X. 1986, G. M. de Rougemont (cR). Yunnan: 2 d d: Yulongshan, 3000-3500 m, 27°06’ N, 100°15’ E, Ganhaizi pass, 18.-23. VII. 1990, V. Kuban (NHMB); 12 dd, 12 22: Leigongshan, Xijiang, 1200- 1900 m, 28. V. — 2. VI. 1997, Bolm (NHMB, cP). Sichuan: 1 2: Xichuan, X. 1986, G. M. de Rougemont (cR); 80 d 8,55 9 9: Mt. Emei, 500-1200 m, 4.-18. V. 1989, S. & J. Kolibec; 55 d d, 68 9 2: ibidem 600-1050 m, 5.-19. V. 1989, L. Bocäk, 56 d d 43 2 2: ibidem 4.-20. V. 1989, V. Kuban (NHMB, cP); 25 exx. ibidem, X. 1985, X. 1986, G. M de Rougemont (cR); 1 d, 2 9 9: ibidem, above Fuhu Monastery, 700-1000 m, stream, ca. 2 m wide, slowly flowing through deep gorge, shaded, with gravel, boulders and sand, Ji & Wang, 15. VI. 1996 (CWBS 238) (NHMW): ] 3: ca. 16 km N Ya’an City, 3 km N Shangli, 950 m, small stream, ca. 0.5 m wide, Ji & Wang, 9. VI. 1996 (CWBS 225) (NHMW). Comments. This is a variable species very closely allied to D. klapperichi (see below). Dianous klapperichi L. Benick Dianous Klapperichi L. Benick, 1942: 76 ff. figs. Material examined. Fujian: 2 dd, 1 9: Shaowu, 500 m, XI. 1937, J. Klapperich (syntypes) (FMCh). Sichuan: 2 d d, 3 2 2: Mt. Emei, 500 m, 4.-20. V. 1989, V. Kuban (NHMB, cP); 1 d ibidem, 7. X. 1985, G. M. de Rougemont (cR); 1 d: ibidem 1530 m, 22. VI. 1994, Schillhammer (4 a) (NHMW); 2 dd: ca. 16 fa N Ya’an City, 3 km N. Shangli, 950 m, small stream, ca. 0.5 m wide, Ji & Wang, 9. VI. 1996 (CWBS 225) (NHMW, cP); 1 d: ca. 14 km N Ya’an Vity, rd. To Shangli, 800 m, river, ca. 5-8 m wide, with large rocks, some of these partly moss-covered, flowing through secondary forest, Ji & Wang, 8. VI. 1996 (CWBS 221) (NHMW). Comments. This species is very close to the variable D. freyi and possibly only an extreme, robust form of this species. I was repeatedly in doubt about the status of this taxon, and today I am still not sure about it, but I propose to regard it tentatively as a distinct species. It differs from D. freyi by the coloration of the femora (basal half reddish, apical half, rather sharply delimited, blackish as compared to only basal third more or less reddish, mostly darker, in D. freyi, but this cannot be seen in immature specimens) and by the male sexual characters: In D. klapperichi the sternites 5 and 6 are shallowly emarginate posteriorly (in D. freyi simple), the apical portion of the median lobe is shorter than in D. freyi (figs 226, comp. 222, 223, 225), but D. freyi is also variable in this character (fig. 12 b in Benick, 1942 is an artefact, the apex of the median lobe has been broken (split) in this specimen). Dianous klapperichi is more robust than most specimens of D. freyi and has smaller elytral spots (diameter about as large as length of fourth antennal segment). Both species, D. freyi and D. klapperichi differ easily from similar species by their apically serrate sternum 9 (figs 230, 231). Dianous cupreostigma Puthz Dianous cupreostigma Puthz, 1984a: 102 ff. figs. Material examined: 1 4: Taiwan: Nantou Hsien: Shanlinchi, 1650 m, 16. V. 1990, on wet rocks in a creek (see photo 2 in Smetana, 1995), A. Smetana, (cS). Comments. This species was described from Chito Experimental Forest, 1150 m (holotype in the B. P. Bishop Museum, Honolulu, paratype in cP). The specimen. from Shanlinchi differs from the original specimens in having a distinct orange spot on the elytra (not a coppery mark); the spot is distinctly separate from the darker ground color 500 VOLKER PUTHZ of the elytra and has a narrow coppery transition zone at periphery. In all other respects, the specimen agrees well with the original specimens. Dianous mendax sp. n. Type material. Holotype (male) and allotype (female): Taiwan: Nantou Hsien, Shan- linchi, 1650 m, 19. V. 1991, on wet rocks in a creek (see photo 2 in Smetana, 1995), A. Smetana (MHNG, cS). Paratypes: Taoyuan Hsien: 1 9: Takuanshan Forest, 1600 m, 17. IV. 1990, same habitat as HT, A. Smetana (cP). Hualien Hsien: 1 9: Rakao National Park, Chungyantienshi (River) Waterfall, 2300 m, 10. V. 1990, wet moss and debris in spray zone of a waterfall, A. Smetana (cS). This new species is the sister species of D. cupreostigma Puthz, which it closely resembles. Moderately shining, black with bluish and some aeneous tint, elytra each with a rather large orange spot (slightly larger than in fig. 172), distinctly delimited but with coppery transition zone at periphery. Head moderately coarsely and densely punctate, pronotum coarsely and very densely punctate, punctures somewhat coalescent near anterior and posterior margins, punctation of basal third of elytra same as that on pronotum, sculpture of posterior two thirds becoming increasingly more confluent around not confluently punctate orange spots, but not forming long rugae as e. g. in D. grandistigma, abdominal punctation fine and dense. Pubescence whitish, that of the fore parts dense and semierect, that of abdomen very dense and recumbent. Antennae, palpi and legs blackish with some metallic tint, antennal club, palpi, and tarsi some- times slightly lighter. Clypeus and labrum blackish blue, moderately densely pubescent. Fourth tarsal segment distinctly bilobed (fig. 9). Length: 4.8 — 6.2 mm (fore parts: 2.0 — 3.0 mm). PM of HT: wH: 45; adE: 26; IE} 21; I: 5; wP2 325; IP? 3353 wEEBl JE ES: IS: 45.5. Male: Medial area of metasternum moderately broadly flat, shallowly impressed and impunctate posteriomedially, brilliant, sides of medial area coarsely and moderately densely punctate, interstices nearly smooth, brilliant, diameters of punctures about as large as basal cross section of third antennal segment, interstices about as large or slightly larger than diameters of punctures, acam impunctate, brilliant. Sternites 3-6 somewhat less dense punctate medially than laterally, sternite 7 slightly flattened posteriomedially, finely and very densely punctate and pubescent, posterior margin very shallowly emarginate; sternite 8 with a broad emargination in about posterior sixth; sternite 9 rounded-serrate apically (fig. 232), apical brush moderately strong, somewhat shorter than the sternite. Tergite 10 broadly rounded. Aedeagus (fig. 217), median lobe broad and + rounded anteriorly, with many long setae ventro- laterally, internal structures consisting of small expulsion bands, a strong tube, and densely spinose membranes. Parameres somewhat longer than the median lobe, evenly set with 17- 20 long setae. Female: Ventral characters similar to those of male. Sternite 8 (fig. 216), less prolonged apically than in D. cupreostigma (fig. 227). Valvifer obliquely rounded- serrate apically, apical brush as long as valvifer. Tergite 10 moderately narrowly rounded (not triangular). THE GENUS DIANOUS IN CHINA 501 Head broad, frons with long and deep longitudinal furrows, medial portion distinctly elevated, but not as much as the side portions, punctation coarse and mostly dense, diameters of punctures as large as basal cross section of third antennal segment, interstices mostly smaller than half diameters of punctures, distinctly larger posteriorly on side portions and on elevated medial portion, but rarely as large, or slightly larger, than diameters of punctures (therefore no distinctly separate shining areas). Antennae moderately slender, extending beyond posterior margin of pronotum by two segments when reflexed. Pronotum with usual oblique impressions on posterior half and with transverse impression near anterior margin; punctation coarse and very dense, diameters of punctures about as large as median cross section of third antennal segment, inter- stices much smaller than half diameters of punctures, distinctly larger posteriomedially near a small impunctate area (as large as one or two neighbouring punctures). Elytra similar to those of D. cupreostigma, but spots much larger. Abdomen as in D. cupreo- stigma with very broad, upwards directed, finely and very densely punctate para- tergites; punctation of tergites fine and dense, less dense medially with interstices exceeding diameters of punctures; tergite 8 somewhat more coarsely and densely punctate than sternite 7, tergite 10 moderately sparsely punctate. Legs slender, metatarsi as long as metatibiae, first segment slightly shorter than three following segments combined, slightly longer than last segment, fourth segment moderately deeply bilobed (fig. 9). Entire surface more or less shallowly reticulated (reticulation may be nearly obsolete on fore parts). Comments. Dianous mendax may be distinguished from the very similar D. cupreostigma Puthz by the larger elytral spots (at least 1.5 x as large as those of D. cupreostigma), by the more bilobed fourth tarsal segment (fig. 9, compare fig. 12), and by the sexual characters. Etymology: mendax (Latin) = false; this species ,,imitates®* D. cupreostigma. PART HI: KEY TO THE DIANOUS-SPECIES OF CHINA AND JAPAN 1(4) Frons concave, medial portion not elevated (group I, Puthz, 1981a), temples very short. [Note: In D. latitarsis L. Benick the frons is broadly concave but its medial portion is slightly to distinctly elevated. This large species is easily recognized by its long temples at least half as long as the eyes: see # 10]; sternite 9/valvifer apically serrate, apicolateral brush short, less than half as long as sternite/valvifer 2 (3) Head about as broad as, or broader than elytra, lateral portions of frons even, not declining, medial portion impressed. Interstices of punctation of head and pronotum narrow but distinct, strongly shining. Male: apical notch of sternite 8 triangular with straight sides. Aedeagus (fig. 16, Euh SI TO ES 00mm See ee ey ed ae tele D. tonkinensis (Puthz) Vietnam, Thailand, Indonesia (Borneo, Sulawesi), China: Yunnan: | à: Xishuangbanna, 22. I. 1993, G. M. de Rougemont (cR):ne w to China. 3(2) Head distinctly narrower than elytra, entire frons (including lateral portions) concave, interstices of punctation of head and pronotum 502 VOLKER PUTHZ extremely narrow (punctation somewhat confluent), head and pronotum nearly dull. Male: Apical notch of sternite 8 triangular, with sinuate sides. Aedeagus as in fig. 2 A, Rougemont (198 1a). SATTE AR RO AR, ase) eae D. yao Rougemont Burma, Thailand, China: Guizhou 4(1) Medial portion of frons distinctly elevated, frons with two distinct longitudinal furrows, or broadly elevated with shallow, longitudinal impressions (group II, Puthz, 198 1a) 5 (44) First segment of metatarsus distinctly longer than following three segments combined (species doubtful in this character are treated in both couplets). Fourth segment of metatarsus symmetrical or asymmetrical First segment of metatarsus longer than remaining segments combined Tarsi pseudotetramerous: tarsal segment four with tube-like bristle shoe, enveloping segment five except claws 8 (11) Frons with broad and shallow but distinct impression posteriomedially 9 (10) Head as broad as or broader than elytra (fig. 1, Shibata, 1979), eyes smaller (IT: IE > 0.75). Punctation of frons less dense, interstices on posteriolateral portions as large as, or slightly larger, than diameters of punctures. Elytra with distinct impressions. Male: aedeagus (fig. 2, Shibata, 1979), a button-shaped apex of median lobe projection dorsally (about as in fig., 20, Puthz, 1981). Female: 8th sternite broadly rounded. 70 SS ARR ER NE PAR ST RL EE NE Sr AP ANR EE D. arachnipes Puthz China: Taiwan 10 (9) Head distinctly narrower than elytra, eyes larger (IT: IE < 0.75). Punc- tation of frons denser, interstices on posteriolateral portions distinctly smaller than diameters of punctures. Elytra nearly even. Male: aedeagus (fig. 19, Puthz, 1981), anterior portion of median lobe triangularly narrowed, apex less button shaped and slightly curved dorsally. Female: Sthisterniterounded 752 9-5 MM: 2. 2er er D. latitarsis L. Benick India: Sikkim; Kashmir; Nepal; Thailand; China: Sichuan Frons not impressed posteriomedially, broadly convex, oval impression only around ocelli. Pronotum impunctate. Larger species. Head broader than elytra, eyes relatively small (IT: IE = 0.80). Elytra nearly even, no distinct impressions. Male: unknown. Female: 8th sternite broadly rounded 0 8 0m LEI ae ee D. moritai Naomi Japan: Amami Islands 12 (7) Tarsi distinctly 5- segmented, tarsal shoe present only on ventral third of segment 5. Pronotum distinctly punctate. Smaller species. Head broader or nearly as broad as elytra, eyes relatively large (ITe: IE = 0.6). Elytra nearly even. Male: sternite 8 with a broad and shallow emargination in about posterior fifteenth. Sternite 9 (fig. 6, Puthz, 1997), bluntly pro- duced apicolaterally. Tergite 10 broadly rounded. Aedeagus (fig. 7, Puthz, 1997), median lobe triangularly narrowed, apex narrowly roun- ded, parameres slightly longer than median lobe with about 16 long 6 (13 7 (12 TZ RED 11 (8 i 15 (24) ISO) 17 (18) 18 (17) 19 (20) 20 (19) 21217) 2205) THE GENUS DIANOUS IN CHINA 503 setae. Female: sternite 8 very broadly rounded, slightly emarginated apicomedially. Tergite 10 broadly rounded. CSS NN RE MA EN a PIERRE D. nigrocyaneus Puthz China: Hainan First segment of metatarsus distinctly shorter than remaining segments combined, segment four with more or less distinct bristle shoe Elytra bi- or tricolorous (specimen to be observed under various directions of light; e. g. a bluish area in middle of a greenish area is sometimes immediately apparent) Longitudinal furrows of frons distinct posteriorly Elytra blue with 4 separate, variegate-metallic areas (figs 35- 37) Outer quarters of elytra brilliant brassy-golden-coppery, centre bright blue (fig. 36). Impunctate shining median areas of sternites less broad than of the width of sternite. Male: sternum 9 (fig. 43), tergite 10 broadly rounded, aedeagus (fig. 44). Female: sternite 8 (figs 48, 51, 54), tersited Oitmanculars/'S:=)9.5 amma, Mi PN ne D. elegantulus Zheng China: Sichuan, Hunan Elytra blue with 4 brassy-golden-coppery areas on lateral quarters, which are separate from the sides by metallic blue (figs 35, 37). Impunctate shining median areas of sternites broader than of the width of sternite Larger: 8.0 -9.5 mm, brassy areas on elytra larger (fig. 37), impunctate median areas of sternites nearly half as broad as width of sternite. Male: aedeagus about as in D. elegantulus, median lobe slightly less broad anteriorlyAklemalerasıın Dielesaniulus ae isan sn D. adonis sp. n. China: Sichuan Smaller: 6.5 -7.5 mm, brassy areas on elytra smaller (fig. 35), impunc- tate median areas of sternites nearly 1/3 as broad as width of sternite. Male: aedeagus similar to that of D. elegantulus, but parameres shorter, slightly longer than the median lobe. Female: unknown. D. schoenmanni sp. n. Elytra differently colored Bluish areas of elytra smaller, transverse bluish band in middle of elytra at its narrowest point 1/6-1/5 as wide as length of elytra; brassy/golden anterior coloration of elytra extending laterally toward the sloping portion of elytra behind shoulders. Sternites 3-5 medially with a moderately broad impunctate area, about as broad as respective sternite. Male: as subspecies, see below. Female: sternite 8 (figs 59, 60). COS im ee lem D. inaequalis inaequalis Champion India: U.P., Sikkim, Assam; Nepal; China: Yunnan, Sichuan Bluish areas of elytra larger, transverse bluish band in middle of elytra at its narrowest point 1/3 - 1/4 as wide as length of elytra; brassy/golden anterior coloration of elytra distinctly separate from lateral bluish portion. Sternites 3-5 medially with broader impunctate area (1/4-2/5 as 504 24 (14) 23182) 26 (31) 27 (28) 28 (27) 29 (30) 30 (29 — VOLKER PUTHZ broad as the respective sternite). Male: aedeagus (fig. 69). Female: sternite 8 as in figs 61-67. 6.0 - 7.5 mm........ D. inaequalis laevior ssp. n. China: Taiwan Longitudinal furrows of frons becoming obsolete posteriorly Tarsı simple, fourth segment of protarsus broad but not bilobed, bristle shoe distinct but not remarkably strong Pronotum distinctly punctate, diameters of punctures, especially near posterior margin, at least as large as one medial eye facet. Blue areas of elytra larger, distinctly separate Larger species: 6.5 - 8.5 mm. Bluish areas of elytra (fig. 34). Pronotum deeply reticulated, moderately shining or dull. Tergite 10 rounded. Male: aedeagus (fig. 38). Female: sternite 8 (figs 56-58)... D. alternans Zheng China: Sichuan Smaller species, bluish areas of elytra as in fig. 33. Pronotum nearly without groundsculpture, brilliant. Two very closely related species Elytra with brassy- greenish reflection on anterior sutural half (fig. 33). Male: sternite 9 (fig. 41). Tergite 10 rounded. Aedeagus (fig. 39), median lobe broader, parameres with only 11-12 setae apically. Female: sternite 8 shallowly emarginated apicomedially (about as in fig. 76). NYO ERRORI E AR D. coeruleovestitus sp. n. China: Hongkong Elytra with violet-coppery reflection on anterior half. Male: sternite 9 (fig. 8, Puthz, 1997) resembling fig. 41. Tergite 10 rounded. Aedeagus (fig. 9, Puthz, 1997), median lobe narrower, parameres with 19-20 setae distributed along entire length of each paramere. Female: unknown. SORTA oa alate de D. coeruleomicans Puthz China: Hainan 31 (26) Pronotum impunctate or indistinctly punctate, diameters of punctures smaller than one medial eye facet. Bluish areas of elytra smaller, less distinct, and less distinctly separate. Tergite 10 rounded. Male: aedeagus about as in fig. 40, apical portion of median lobe broader than in previous species. Female: sternite 8 (fig. 8, Puthz, 1981a) about as in 23964 40-81 RE I Ro. D. chetri Rougemont India: U.P.; Nepal; Bhutan; Laos; China: Yunnan, Sichuan Tarsi asymmetrically bilobed (fig. 7), fourth segment of protarsus bilobed in about apical quarter, and with a very strong bristle shoe. Male: aedeagus as in fig. 28, Puthz,1981a, with median lobe broadly rounded anteriorly. Female: sternite 8 (fig. 32) GO: Sema Bee AREAS re Lace PET PAS D. versicolor Cameron India; Nepal; Bhutan; China: Yunnan, Sichuan Elytra unicolorous, sometimes with various metallic tint, without distinctly delimited areas contrasting in color from main color of elytra Elytra with coarse and dense vorticose sculpture, punctures/ rugae as large as basal cross section of antennal segment 3 or larger. Male: 35 (34) 36 (39) STAGES \ 383.67) 39 (36) 40 (43) 41 (42) 42 (41) 43 (40) 44 (5) 45 (72) THE GENUS DIANOUS IN CHINA 505 sternite 9 (fig. 151). Aedeagus (fig. 27, Puthz, 1981a), parameres spoonlike broadened apically. Female: sternite 8 about as in fig. 198. DONC NII ces RA III A: D. andrewesi Cameron India, Nepal, China: Yunnan: 1 d: Xishuangbanna, 22. I. 1993, G. M. de Rougemont (cR), new to China. Elytra (moderately) finely punctate, not coarsely vorticose, punctures much smaller than basal cross section of antennal segment 3 Punctation of pronotum slightly finer than that of head and elytra, dense, interstices mostly about as large as half diameters of punctures Metatrochanter in both sexes with distinct prominent tooth.Male: tergite 10 very broadly rounded or blunt apically, aedeagus (fig. 1, Puthz, 1980a). Female: sternite 8 (fig. 5, Puthz, 1980a, apical notch deeper than in fig. 76). Tergite 10 triangular, apical point less prominent as that oDirangulifers6:0 249 an Are. ma cee D. yunnanensis Puthz China: Yunnan; India: Assam Metatrochanter simple Punctation of pronotum, at least partially, distinctly coarser than that of head and elytra, mostly less dense Smaller species, less than 6.5 mm long Elytral punctation very fine and extremely dense, elytra quite dull, punctures smaller than one medial eye facet; pronotum with deep im- pressions and four conspicuous calli, punctation mostly dense, inter- stices mostly smaller than diameters of punctures. Male: tergite 10 rounded, aedeagus as in fig. IA (Rougemont, 1981b) median lobe triangularly narrowed, apical portion twice as long as broad. Female: sternite 8 (fig. 47), tergite 10 obtusely angled. GI) I (OE OI PRE EL NA D. srivichaii Rougemont Burma; Thailand; Vietnam; Laos; China: Yunnan: | d: Xishuang- banna, 22. I. 1993, G. M. de Rougemont (cR), Fujian Elytral punctation less fine and less crowded, elytra somewhat shiny, punctures somewhat larger than one medial eye facet; pronotum with Shallowsimpresstons 920 SME RR D. coeruleotinctus sp. n. China: Hunan Larger species exceeding 6.5 mm. Male: tergite 10 broadly rounded. Aedeagus as in fig. 44, Puthz (1981a) median lobe narrowly trian- gularly narrowed. Female: sternite 8 with a distinct apical excision as in figs 10, 11, Puthz (1981a) about as in fig. 48; tergite 10 broadly fran cular: OE ARI RE Ronee D. punctiventris Champion India; Burma; Laos; Vietnam; China: Yunnan, Hainan First segment of metatarsus as long as, or shorter than following three segments combined. Fourth segment of metatarsus more or less symmetrical Head broader than elytra or about as broad as elytra, definitely not narrower than elytra (this character might be deceptive; in some cases measurements should be taken) 506 46 (67) 47 (52) 48 (51) 49 (50) 50 (49) 51 (48) VOLKER PUTHZ Legs black or preponderantly dark (dark brown to black) Fourth segment of metatarsus conspicuously, deeply bilobed (fig. 8). Larger species with yellowish spotted elytra. Sternum 9/valvifer with very distinct apicolateral teeth/tooth (e. g. fig. 90) Punctation of pronotum and elytra coarse, very dense and rugose punctate, interstices microsculptured. Elytral spot much smaller. Male: aedeagus (fig. 87), median lobe with a narrow triangular apical portion, which is much longer than broad, parameres shorter than median lobe. Female: sternite 8 broadly rounded, valvifer (fig. 1, Puthz, 1980b), with apremanrkableitooth ©1022 mms yeast ea eee D. luteoguttatus Champion India; Nepal; China: Taiwan Punctation of pronotum and elytra moderately coarse, moderately dense, distinctly well-defined, interstices smooth. Elytral spot much larger (fig. 168). Male: aedeagus (fig. 15, Puthz, 1997), medianlobe with a broad triangular apical portion, which is broader than long; parameres longer than median lobe. Female: similar to previous species MORES OMS MINNIS IE RSS ISO PINTO D. hainanensis Puthz Smaller species, elytra lack yellowish spots. Male: sternite 9 serrate apically (fig. 238). Aedeagus with a broad lanceolate apical portion of the median lobe (fig. 224). Female: unknown. 5.0 - 5.8 mm. D. lobatipes sp. n. China: Anhui Fourth segment of metatarsus not, or only slightly bilobed. Fore body without microsculpture, dark blue-violet or brilliant-black with a coppery spot on each elytron Dark blue-violet, punctation of pronotum coarse and dense, confluent anteriorly and posteriorly. Paratergites narrower, those of segment 4 as broad as antennal segment 10. Male: sternite 8 with a triangular excision in about posterior quarter. Sternite 9 (fig. 2, Puthz, 1997) with a distinct tooth apicolaterally. Aedeagus (fig. 1, Puthz, 1997), median lobe triangular apically, with ventrolateral ridges, and dense, long setae on apical portion, parameres about as long as median lobe with about 10-12 setae. Female: unknown. 3.7 - 4.7 mm............ D. violaceus Puthz China: Hainan Brilliant-black, elytra each with a small coppery spot on posterior half, punctation punctures. Paratergites broader, those of segment 4 broader than antennal segment 10. Male: sternite 9 (fig. 228); Tergite 10 broadly rounded. Aedeagus (fig. 214). Female: sternite 8 about as in fig.215; tergite 10 narrowly rounded. 3.8 - 4.8 mm...... D. cupreogutta sp. n. China: Taiwan Fore body with miscrosculpture (at least partially), coloration different, mostly dark bronze-green Paratergites narrow and directed downwards, paratergites of segment 4 about as broad as first antennal segment, with punctation fine an mode- 96137) 59 (60) 60 (59) 61 (62) 62 (61) 63 (64) 64 (63) 65 (66) 66 (65) 67 (46) THE GENUS DIANOUS IN CHINA 507 rately sparse. Fore parts as in fig. 2, Sawada, 1960. SORA ee ae D. shibatai Sawada Japan: Honshu Paratergites broader and mostly directed upwards, sometimes sagittal, paratergites of segment 4 at least twice as broad as third antennal segment (Five taxa described from Japan, which are very closely related and may belong to one variable species) couplets 51-56 in this key are mostly taken from Naomi, 1988 and Watanabe, 1984 Larger species, more than 4.5 mm long, when abdomen extended. Male: aedeagus as in figs 6, 7, Watanabe, 1984. ATA Ro E E II e D. iwakisanus Watanabe Japan: Honshu Smaller species, not exceeding 4.5 mm in length, when abdomen extended Black with dark bluish reflection. Male: median lobe of aedeagus parallel-sided in apical quarter (figs 3, 4, Watanabe, 1984). Shape of bodyzshisällies3.9 242mm en er D. gongen Watanabe Japan: Honshu Black with bronze-green reflection. Male: median lobe of aedeagus more or less narrowed apically Metallic reflection very distinct. Male: median lobe about as in D. Japonicus (see below), parameres slightly shorter (fig. 2 B, Naomi, OS Amatori casina ssi LR D. viridicatus Naomi Japan: Honshu Almost black, with feeble metallic reflection Broader; pronotum covered with subrugose punctures on base. Male: parameres of aedeagus longer, with denser and longer setae apically (fig. 1 C, Naomi, 1988). Shape of body: fig. 1, Sawada, 1960. SERE ho iaia D. japonicus Sawada Japan: Honshu. Through the kindness of Drs. Sawada and Naomi I studied the holotype of D. japonicus and two paratypes of D. septentrionalis Naomi, together with some males in my collection. Sculptural differences between both taxa (as noted by Naomi), as well as slight differences in the shape of the anterior portion of the median lobe, fall within the range of variability. Dianous septentrionalis should be regarded as a synonym of D. japonicus. syn. nov. ! Narrower; pronotum densely covered with round punctures on base. Male: parameres of aedeagus shorter, with sparser and shorter setae apically (fig. 1 B, Naomi, 1988). 3.9 mm. ............. D. morimotoi Naomi Japan: Kyushu Legs reddish or paler 508 VOLKER PUTHZ 68 (69) Elytra trapeziform, elytral spot large, oval (fig. 176). Male: sternite 9 (fig. 236). Aedeagus (fig. 213). Female: sternite 8 slightly, obtusely COUNACAP0AISO IMMERSE MERE PE PR eee D. electrigutta sp. n. China: Taiwan 69 (68) Elytra subquadrate, elytral spot less oval 70(71) Elytral spot smaller as in fig. 175; head narrower; elytral sculpture more confluent. Male: sternite 9 (figs 233, 234). Aedeagus (figs 210, 211). Female: sternite 8 slightly, obtusely rounded. ASSO: Asim = EEE Oe IE ene ee D. sucinigutta sp. n. China: Taiwan 71(70) Elytral spot larger, as in fig. 174; head broader; elytral sculpture more regular, less confluent. Sexual characters resembling those of the previous species (see figs 212, 235). 5.0 - 6.3 mm... D. sucininotatus sp. n. China: Taiwan 72 (45) Head distinctly narrower than elytra 73 (76) Legs predominantly reddish 74(75) Smaller species. Head much narrower than elytra. Paratergites broader, distinctly directed upwards, very densely punctate. Male: aedeagus as in fig. 1 A, Naomi, 1988, about as in figs 222, 223. Shape of body: fig. 3, Sawada, 1960. 4.0 - 5.0 mm. . D. amamiensis Sawada Japan: Amami Oshima Is. 75 (74) Larger species. Head distinctly, but not remarkably narrower than elytra. Paratergites narrower, directed downwards, moderately sparsely punctate. Male: sternite 9 (fig. 237). Aedeagus (fig. 221). Female: sternite 8 rounded, slightly produced in middle. China: Sichuan 76 (73) Legs black or predominantly dark, often with metallic tint 77 (78) Longitudinal furrows of frons obsolete posteriorly, frons evenly convex posteriorly: see # 32: D. versicolor 78 (77) Longitudinal furrows of frons complete, extending from antennal tubercles to neck 79 (158) Fourth segment of metatarsus distinctly bilobed (figs 8-12, 114, 219, 220x). Species doubtful in this character are treated in both couplets. (Note: Lobation of metatarsi is less strong than that of anterior tarsi). 80 (101) Elytra unicolorous or variegate metallic, without orange spot 81 (88) Punctation of elytra distinctly well- defined (sometimes a few punc- tures may be confluent; if dark coppery- violet : see # 93: D. cupreo- violaceus Puthz) 82 (83) Fourth segment of metatarsus deeply bilobed, lobes longer than half segment 5. Abdominal punctation sparser, interstices on tergite 4 at least twice as large as diameters of punctures. Male: Sternites 3-6 with an acute projection apicolaterally; tergite 10 broadly rounded. Aedeagus as in fig. 2, Puthz, 1980a. Female: Sternite 8 triangular; tergite 10 truncate- triangular. 83 (82) 84 (85) 85 (84) 86 (87) 87 (86) 88 (81) 89 (90) 90 (89) 9102) 92 (91) 93 (94) THE GENUS DIANOUS IN CHINA 509 SIA IT. A. EEE o PR noi. D. spiniventris Puthz China: Yunnan Fourth segment of metatarsus less deeply bilobed, lobes shorter than half segment 5. Abdominal punctation denser, interstices on tergite 4 less than twice as large as diameters of punctures Eyes larger, temples about 1/5 as long as eyes. Abdominal punctation very dense, interstices on tergite 4 smaller than diameters of PUNCtUTES HAS) Sm ee ee (D. ater sp. n.) see # 89 Eyes smaller, temples about 1/3 as long as eyes. Abdominal puncta- tion less dense, interstices on tergite 4 larger than diameters of punctures Larger species. Elytra entirely black. Male: sternite 8 with apical notch in about posterior quarter; sternite (fig. 116). Aedeagus (fig. 113), parameres with 8 + 10 (-12) setae. SIZE 2 VON) Vata NO RINO AR D. atrocyaneus sp. n. China: Taiwan Smaller species. Elytra with a cupreous spot on posterior half. Male very similar to that of D. atrocyaneus, sternite 8 with deep notch in about apical third, aedeagus about same as in D. atrocyaneus, but parameres with 7+ 7 (- 8) setae. 4.5 - 5.2 mm.......... D. yoshidai Naomi Japan: Iriomote Is. Punctation of elytra (most of it) distinctly confluent Elytra entirely black, abdominal punctation very dense, interstices on tergite 4 smaller than diameters of punctures. Tergite 10 broadly rounded. Male: sternite 9 (fig. 229). Aedeagus (fig. 209). Female: SlemmiteyS*(hies.21'5) 74 WS 76M PER ARE RE D. ater sp. n. China: Taiwan Elytra with more or less distinct metallic tint Pronotum broader than long. Elytra with bluish tint and a more or less distinct, skyblue spot on posterior half. Male: sternite 9 serrate apically; tergite 10 broadly rounded. Aedeagus as in figs 24-26 (Puthz, 1981a), apical portion of median lobe enlarged, ore or less emarginated medially. Female: sternite 8 about as in fig. 93. Tergite 10 narrowly rounded. 5.5 - 7.0 mm........... D. subvorticosus Champion India; China: Yunnan Pronotum longer than broad Dark with strong coppery-violet reflection. Punctation of elytra well- defined on sutural third, more or less confluent on lateral two thirds. Paratergites not directed upwards, narrower, those of segment four as broad as antennal segment 2. Male: sternite 8 with a subtriangular notch in about posterior third. Sternite 9 (fig. 5, Puthz, 1997) with a broad tooth apicolaterally. Tergite 10 broadly rounded. Aedeagus (fig. 3, Puthz, 1997) with a lancolate apex of median lobe which is curved dorsally and densely set with granules and some longer setae, 510 94 (93) 95 (96) 96 (95) 97 (100) 98 (99) 99 (98) 100 (97) VOLKER PUTHZ parameres longer than median lobe, densely set with long setae. Female: sternite 8 (fig. 4, Puthz, 1997), narrowly rounded apico- medially. Tergite 10 semicircularly rounded. EUTIN EN. rec ee to erano RE PES D. cupreoviolaceus Puthz China: Hainan Differently colored. Punctation of elytra confluent, including sutural third. Paratergites directed upwards, broader, those of segment 4 twice as broad as antennal segment 2 Metatarsal segment four very shortly bilobed, lobes much shorter than 1/3 segment 5 (fig. 12). Pronotum with strong and homogeneous punctation lacking deep impressions. Elytra bluish with a coppery spot on posterior half. Male: sternite 9 serrate apically; tergite 10 broadly rounded. Aedeagus as in fig. 4 (Puthz, 1984a), apical portion of median lobe triangular, densely set with long ventro-lateral setae. Female: sternite (fig. 227); tergite 10 narrowly rounded. A WO) > DOV TOT NUL ARA PAIA AC A, eS SA ı D. cupreostigma Puthz China: Taiwan Metatarsal segment four deeply trilobed, lobes as long as 1/3 of segment 5 (fig. 114, see also fig. 16, Puthz, 1998). Pronotum with coarse, variegate dense punctation and deep impressions Medial portion of frons strongly elevated, extending well beyond the level of medial eye margins Abdominal punctation very dense, interstices smaller than punctures, tergite 8 finely and densely punctate with fatty reflex, interstices about as large as diameters of punctures, deeply reticulate; interstices of ventral abdominal punctation distinctly reticulate. Elytra blackish- blue with a coppery reflex (or an orange spot) behind middle. Male: apical portion of median lobe rounded, fig. 185). Female: sternite 8 (fig. 195). Tergite 10 subtriangular (fig. 200). SIA AO RI a se ee N D. verticosus Eppelsheim India, Burma, Vietnam, China: Yunnan, ? Sichuan Abdominal punctation less dense, interstices as large as punctures, tergite 8 less finely and less densely punctate, more brilliant- shining, interstices larger than diameters of punctures, less deeply reticulate: interstices of ventral abdominal punctation smooth. Elytra dark blue with a golden- coppery area on posterior half near suture. Tergite 10 rounded. Male: sternite 9 (fig. 117). Aedeagus (fig. 123). Female: sternite 8 (fig. 115). Tergite 10 rounded. 5.3 - 6.5 mm... D. gemmosus sp. n. China: Yunnan Medial portion of frons less elevated, not extending to the level of medial eye margins. Elytra bronze-black-bluish with a bluish or coppery reflex on the rosette. Tergite 10 rounded. Male: sternite 9 serrate apicolaterally. Aedeagus with the median lobe triangularly THE GENUS DIANOUS IN CHINA 511 narrowed (fig. 30, Puthz, 1981). Female: sternite 8 triangular as in the PLEVAOUS SPECIES PS Oma an er D. cameroni Champion Pakistan (Punjab), India, Nepal, China: Yunnan 101 (80) Elytra with distinct orange spots 102 (123) Diameter of elytral spot about as large as length of third antennal segment or larger 103 (106) Pubescence of fore parts conspicuously long and erect, single setae as long as fourth antennal segment 104 (105) Smaller: 4.2- 5.2 mm. Coloration mainly blackish-violet. Interstices of punctation smooth. Tergite 10 rounded. Male: sternite 9 (fig. 136). Aedeagus (fig. 128), apical emargination of median lobe deeper, apical lobes narrower. Female: sternite 8 (fig. 132). AD ATI AR MeL ap Stew anne D. aequalis Zheng China: Sichuan, Guangdong (?) 105 (104) Larger: 4.6 - 6.3 mm. Coloration markedly bluish (-green). Interstices of punctation more or less distinctly reticulated. Tergite 10 rounded. Male: sternite 9 (figs 118, 119). Aedeagus (figs 121, 122), apical emargination of median lobe less deep, apical lobes broader. Female: Stemmite Sta boutas nie al sn ser fee. erates D. chinensis Bernhauer China: Shandong, Shaanxi, Anhui, Hunan, Jiangxi, ? Fujian 106 (103) Pubescence of fore parts less long, less conspicuous, half erect or recumbent, longest setae distinctly shorter than fourth antennal seg- ment 107 (114) Sculpture of elytra very distinctly confluent/rugose around, and partly on, orange spot. Fully winged species 108 (109) Elytral spot larger, diameter as large as length of antennal segments 3 and 4 combined; abdominal punctation sparser, interstices in middle of tergites twice as large as diameters of punctures. Male: sternite 9 (fig. 208, see also fig. 6, Puthz, 1980b), with a distinct tooth apico- laterally; tergite 10 broadly rounded. Aedeagus (fig. 190, see also fig. 5, Puthz, 1980b), apical portion of median lobe broadly trian- gular, densely set with long ventrolateral setae. Female: sternite 8 (fig. 199), tergite 10 rounded apically. 5.0 - 6.0 mm. . D. oculatipennis Puthz China: Yunnan 109 (108) Elytral spot smaller, diameter distinctly smaller than length of anten- nal segments 3 and 4 combined; abdominal punctation denser, inter- stices in middle of tergites mostly at most as large as diameters of punctures 110 (111) Elytral sculpture around orange spot and on its periphery strongly vorticose, in centre of spot some (more or less) punctures well- defined. Male: sternite 9 (fig. 205) with a distinct tooth apicolaterally: tergite 10 broadly rounded. Aedeagus (fig. 178). Female: valvifers (fig. 192) densely set with moderately long, very strong setae on its posteniorhalt (tieaslOl) 23.0265 mm kee oe ee ae D. grandistigma sp. n. China: Yunnan; Vietnam (?) 512 VOLKER PUTHZ 111 (110) Elytral sculpture, including (complete) sculpture of orange spots, densely confluent on posterior half, long and narrow rugae pass obliquely backwards. Two very similar species 112 (113) Male: apical notch of sternite 8 in about posterior third. Aedeagus as in fig. 179, apical portion of median lobe broader (fig. 189). Female: sternite 8 (fig. 198), apical projection shorter; tergite 10 slightly SUbtran gUlart4 899 SENTE RR ae D. ocellifer sp. n. China: Yunnan; Laos 113 (112) Male: apical notch of sternite 8 in about posterior quarter. Sternite 9 (figs 202, 203). Aedeagus (figs 177, 182, 183), median lobe narrower. Female: sternite 8 (fig.196), apical projection longer; tergite 10 apically rounded. CO AIAR A eee ate D. ocellatus Cameron (incl. acutus Zheng) India: Assam; Thailand; Vietnam; China: Yunnan; Sichuan 114 (107) Sculpture of elytra more or less confluent only on area between orange spot and suture. Fully winged or brachypterous species 115 (116) Segment four of metatarsus broadly and deeply bilobed. Elytra (fig.170), orange spots extending towards declining sides. Male: sternite 9 as in fig. 13 (Puthz, l.c.). Aedeagus as in fig. 11 (Puthz, Key)yEemalesunknown43:036 SIM FE EEE D. strabo Puthz China: Yunnan; Thailand 116 (115) Segment four of metatarsus very slightly bilobed. Elytral spots distinctly separate from declining sides (as in figs 105-112) 117 (118) Abdominal punctation less fine and denser, punctures of tergite 5 as large as one punctures; punctation of paratergites very dense, all interstices distinctly smaller than half diameters of punctures. Male: sternite 9 (fig. 137); tergite 10 very broadly rounded, slightly concave in posterior middle. Aedeagus (fig. 130). Female: sternite 8 pointed apicomediallys4:5; 3557: mA bb. Sane ET D. dubiosus sp. n. China: Guizhou, Guangxi, ? Taiwan 118 (117) Abdominal punctation finer and less dense, punctures of tergite 5 smaller than one eye facet, interstices in middle of tergites much larger than diameter of punctures; punctation of paratergites less dense, interstices at least somewhere as large as half diameters of punctures, or larger 119 (122) Punctures near lateral border of elytral spot distinctly well-defined 120 (121) Black, or with dark blue (-green) metallic tint. Punctation of posterio- lateral quarter of elytra very coarse and dense, interstices mostly much smaller than diameters of punctures: see # 135 D. taiwanensis Puthz 121 (120) (Blackish) blue metallic (Prussian blue). Punctation of posterior- lateral quarter of elytra moderately coarse and moderately dense to moderately sparse, interstices often as large as diameters of punc- tures, or larger: see # 178 D. gonggamontis sp. n. THE GENUS DIANOUS IN CHINA 513 122 (119) Punctures near lateral border of elytral spot very dense and slightly longitudinally confluent. Black with dark violet tint. See # 136 D. atroviolaceus Sp. n. 123 (102) Diameter of elytral spot smaller than length of third antennal segment 124 (137) Elytral sculpture simple, punctures well-defined, not confluent or vorticose (rarely, a few punctures coalescent) 125 (128) Pubescence of fore parts conspicuously long and erect, single setae as long as fourth antennal segment (similar to fig. 22) 126 (127) Black with faint bluish tint. Abdominal punctation much sparser, interstices on groundsculpture. Tergite 10 rounded. Male: sternite 9 (fig. 163), with a very long apicolateral tooth. Aedeagus (fig. 156). Female: sternite 8 (fig. 167); valvifer triangularly acute apically. AST) OM Ne Re EAN RS a D. socius Zheng China: Sichuan 127 (126) Distinctly bluish. Abdominal punctation much denser, interstices on middle of tergite 4 about as large as diameters of punctures, mode- rately shiny with distinct reticulation. 4.8 - 6.0 mm. See # 105 D. chinensis Bernhauer 128 (125) Pubescence of fore parts shorter, half erect, single setae about as long as second antennal segment, if longer, then distinctly curved (see fig. 24) 129 (132) Pubescence of fore parts greyish 130 (131) Larger species with smaller elytral spot (fig. 103). Abdominal punc- tation dense, interstices on middle of tergites smaller than half diameters of punctures, reticulate. Male: sternite 9 (fig. 120); tergite 10 blunt or very shallowly emarginate apically. Aedeagus (figs 124, 125). Female: sternite 8 (fig. 153); tergite 10 moderately triangular. COESIONE e OE D. yangae sp. n. China: Sichuan 131 (130) Smaller species with larger, longitudinally-oval spot. Abdominal punctation less dense, interstices on middle of tergites about as large as diameters of punctures, smooth or indistinctly reticulate. Male: sternite 9 (fig. 135); tergite 10 as above. Aedeagus (fig. 126). Female: sternite 8 (?), tergite 10 as above. 5.2 - 6.4 mm............ D. senex sp. n. China: Sichuan 132 (129) Pubescence of fore parts blackish 133 (134) Pubescence of fore parts longer, longer than second antennal segment, more erect. Elytra subquadrate.Tergite 10 rounded. Male: sternite 9 (fig. 162). Aedeagus (fig. 155). Female: sternite 8 (fig. 165). LOTTERIA EIERN RER: D. bioculatus sp. n. China: Yunnan 134 (133) Pubescence of fore parts somewhat shorter, about as long as second antennal segment, less erect 514 VOLKER PUTHZ 135 (136) Black or with dark blue (-green) metallic tint. Punctures near lateral margin of elytral spot distinctly well-defined. Medial area of meta- sternum broadly-triangularly impunctate in middle, side portions of medial area each with at most 20 coarse punctures. Medial punctation of sternites moderately fine. Male: sternite 9 (fig. 161); tergite 10 broadly rounded. Aedeagus (figs 157-160). Female: sternite 8 (fig. 166); tergite 10 subtriangular. General outline and size of elytral spot Vanable (nes KU ee ERA ee D. taiwanensis Puthz China: Taiwan 136 (135) Black with dark violet tint. Punctures near lateral margin of elytral 137 (124 — 138 (145) 139 (140) 140 (139) 141 (142) spot very dense triangularly impunctate, side portions of medial area with distinctly more than 20 more or less coarse punctures. Medial punctation of sternites fine. Male: sternite 9 (fig. 89); tergite 10 broadly rounded. Aedeagus (fig. 154). Female: sternite 8 (fig. 164); tergite 10 subtriangular. General outline and size of elytral spot very vanables4526.0. mm wanes ne Nee D. atroviolaceus sp. n. China: Yunnan Elytral sculpture at least partly confluent, mostly confluent between orange spot and suture Both sternite 9 and valvifer rounded- serrate apically (figs 230-232). (If decision is doubtful, try also opposite couplet) Head broader (bEl: DH < 115). Entire body with distinct bluish tint. Male: sternite 9 (fig. 232); tergite 10 broadly rounded. Aedeagus (fig. 217). Female: sternite 8 (fig. 216); tergite 10 narrowly rounded. 18:02, mm ae es Rh el D. mendax sp. n. China: Taiwan Head narrower (bEl: BH > 120) Body blackish-blue-metallic (Prussian blue). Legs entirely black. Pronotum with distinct pubescence. Fore body very densely punctate andineticulatezlessishinin SE en chiens D. dubiosus sp. n. (see # 117) 142 (141) Body blackish. Legs at least at parts brownish or lighter. Pronotum 143 (144) 144 (143) 145 (138) lacking pubescence. Fore body less densely punctate, without or with faint microsculpture, more shining. Two closely related taxa, belon- ging possibly to one variable species Basal half of femora reddish, rather distinctly delimited from blackish apical half. Male: shorter.4.5-5.5mm.......... D. klapperichi L. Benick China: Fujian, Sichuan Basal third of femora + reddish, mostly darker, indistinctly delimited from darker apical portion. Male: sternite 9 (fig. 230). Aedeagus (figs 222, 223, also 225), apical portion of median lobe longer. 4.5 - 5.5 1 i ae Pema. ID.) jen erica China: Zhejiang, Anhui, Jiangxi, Hunan, Fujian, Guizhou, Yunnan, Sichuan Both sternite 9 and valvifer with a more or less distinct tooth apicolaterally (figs 152, 161, 204, 206) THE GENUS DIANOUS IN CHINA 515 146 (151) Elytral sculpture more rugose, rugae present also on orange spot 147 (148) Pronotum broader than long. Smaller: 4.5-5.5 mm. Black,with faint (blue-) metallic tint. Male: notch of sternite 8 in about posterior fifth. Aedeagus (figs 179, 188). Female: sternite 8 (fig. 197); valvifer with apical brush; tergite 10 rounded. 4.5 - 5.5 mm....... D. emarginatus Zheng China: Sichuan, Yunnan; ? Burma 148 (147) Pronotum longer than broad. Larger: 5.5- 6.5 mm 149 (150) Variegate metallic (blue-violet-coppery). Male: notch of sternite 8 in about posterior strong, apically rounded setae, about as in fig. 191; tergite 10 subtriangular (fig. 201). 5.5 - 6.5 mm....... D. variegatus sp. n. China: Yunnan 150 (149) Elytra blackish-blue with a coppery reflex or an orange spot behind middle. Male: notch of sternite 8 in about posterior fourth. Apical portion of median lobe broader, rounded (fig. 185). Female: sternite 8 more triangular (fig. 195), valvifer with apically pointed setae; tergite 10 subtriangular (fig. 200) 5.5 - 6.5 mm......... D. verticosus Eppelsheim India, Burma, Vietnam, China: Yunnan, ? Sichuan 151 (146) Elytral sculpture less rugose, no rugae on orange spot, rugae only around orange spot or between spot and suture. Three very similar species 152 (157) Black, with more or less distinct, Prussian-blue or bluish-green, metallic tint 153 (154) Black and/or metallic. Punctation of posteriolateral quarter of elytra very coarse and dense, interstices mostly much smaller than dia- meters of punctures. Tergite 10 rounded. Male: sternite 9 (fig. 161), mostly with a distinct tooth apicolaterally. Aedeagus (figs 157-160). Female: sternite 8 (fig. 166). 4.3 - 5.7 mm. ..... D. taiwanensis Puthz (pars) China: Taiwan 154 (153) Distinctly blue metallic. Punctation of posteriolateral quarter of elytra moderately coarse and moderately dense to moderately sparse, interstices often as large as diameters of punctures, or larger 155 (156) Head much narrower (wEl: wH > 1.30), elytra quadrate with rectan- gular shoulders. Pubescence of pronotum whitish. Punctation of head moderately coarse and moderately sparse. Male: sternite 9 (fig. 152; see also fig. 7, Puthz, 1971a), serrate apically, with toothlike pro- jection laterally; tergite 10 rounded. Aedeagus (fig. 143), apical portion of the median lobe more or less truncate and broadly emar- ginate. Female: sternite 8 (fig. 134); tergite 10 subtriangular. AN = SED NINE N REES RE NR ee EN D. hummeli Bernhauer China: Sichuan 156 (155) Head broader (bEl: bH < 1.25), elytra slightly trapeziform with somewhat oblique shoulders. Pubescence of pronotum brownish. Punctation of head coarse and very dense. See # 178 D. gongga- montis Sp. n. 516 VOLKER PUTHZ 157 (152) Black, with bluish-violet metallic tint. Brachypterous or fully WATTS SOR SESH BO RR TM anes OLS Ce Nee D. atroviolaceus sp. n. 158 (79) Fourth segment of metatarsus simple 159 (186) Elytra with distinct orange spots 160 (163) Elytral spot very large, diameter distinctly larger than length of third antennal segment (e.g. fig. 104) 161 (162) Pubescence conspicuously long, erect, setae of pronotum distinctly longer than antennal segment 2 (figs 23, 24). Elytral spot very large, extending mostly to sides of elytra in dorsal view (fig. 104). Tergite 10 rounded. Male: sternite 9 (about as in fig. 135). Aedeagus (fig. 127). Female: sternite 8 (fig. 131) 4.8 - 6.0 mm... . D. banghaasi Bernhauer China: Shandong, Shaanxi, Anhui, Jiangxi, Hunan, Guizhou, Guang- dong, Guangxi, Fujian, Sichuan 162 (161) Pubescence much shorter, semierect, setae of pronotum shorter than antennal segment 2. Elytral spot smaller, slightly larger than length of antennal segment 3, not extending to sides of elytra in dorsal view. SCTE Oey N ne BRUT AU Etang SUE EN D. gonggamontis Sp. n. 163 (160) Elytral spot smaller, diameter as wide as, or distinctly smaller, than length of third antennal segment 164 (183) Abdominal punctation fine, or moderately fine, moderately dense to moderately sparse, interstices on middle of tergites distinctly larger than diameters of punctures; punctation of tergite 8 distinctly, but not conspicuously, at most twice as coarse as that on tergite 7 165 (180) Pubescence of pronotum darker, brownish 166 (175) Pubescence of pronotum erect 167 (170) Pubescence of fore parts longer, longer than antennal segment 2, more erect 168 (169) Punctation on posteriolateral quarter of elytra much denser, inter- stices much smaller than diameters of punctures. Head broader (bH: bE = 0.86). Male: sternite 8 (fig. 162). Aedeagus (fig. 155), apical portion of median lobe broader excavated, with more apicoventral setae (27). Female: sternite 8 (fig. 165), less strongly triangular. SEC mn. Shunt sree aie totaal A ee D. bioculatus sp. n. China: Yunnan 169 (168) Punctation on posteriolateral quarter of elytra much sparser, interstices often as large as or larger than diameters of punctures. Head narrower (bH: bE = 0.81). Male: sternite 9 (fig. 163). Aedeagus (fig. 156), apical portion of median lobe less broadly excavated, with less apicoventral setae (18-19). Female: sternite 8 (fig. 167), stronger triangular. 416.2 mm. tee MEER Ee D. socius Zheng China: Sichuan 170 (167) Pubescence of fore parts somewhat shorter, about as long as antennal segment 2, or shorter, less erect THE GENUS DIANOUS IN CHINA 517 171 (172) Black with dark violet tint. Punctures near lateral margin of elytral spot very dense and slightly confluent longitudinally. Male: sternite 9 (about as in fig. 150). Aedeagus (fig. 154). Female: sternite 8 (fig. 164). General outline and size of elytral spot very variable. 81-26: 0: 1a RL SER FEN PR EU CE ar D. atroviolaceus sp. n. China: Yunnan 172 (171) Black, or with dark blue (-green) metallic tint. Punctures near lateral margin of elytral spot distinctly well-defined 173 (174) Abdominal punctation less fine and denser, diameters of punctures nearly as large as one eye facet, interstices in middle of tergites slightly larger than diameters of punctures; punctation of paratergites very dense, interstices smaller than half diameters of punctures. Male: sternite 9 (fig. 150). Aedeagus (fig. 144), median lobe nearly parallel-sided, anterior margin less deeply emarginated. Female: ster- nite 8 (fig. 133), slightly less triangular. 4.0 - 5.0 mm. D. bashanensis Zheng China: Sichuan, Hunan 174 (173) Abdominal punctation finer and sparser, punctures distinctly smaller than one eye facet, interstices in middle of tergites at least twice as large as diameters of punctures; punctation of paratergites less dense, interstices often larger than diameters of punctures. Male: sternite 9 (fig. 161). Aedeagus (figs 157-160), median lobe concave laterally, anterior margin deeply emarginated. Female: sternite 8 (fig. 166), a little more triangular. General outline and size of elytral spot very vamable (tes 107-1143 ES mme tener D. taiwanensis Puthz (pars) China: Taiwan 175 (166) Pubescence of pronotum semierect 176 (179) Punctation of paratergites very dense, interstices much smaller than diameters of punctures 177 (178) Punctation of the fore parts very coarse and very dense. SEHR Ii SER BREI IM D. dubiosus sp. n. 178 (177) Punctation of the fore parts less coarse and less dense. Male and female: About as in D. banghaasi (compare figs 127, 131, 135). ADS 0 NAT se tee eet DIO EN A EUR D. gonggamontis sp. n. China: Sichuan 179 (176) Punctation of paratergites less dense, interstices often as large as diameter of punctures, or larger see # 174 D. taiwanensis Puthz 180 (165) Pubescence of pronotum whitish and semi-erect 181 (182) Punctation of fore parts less coarse and less dense, diameters of punctures on frons nearly as large as basal cross section of palpal segment 3, interstices mostly larger than half diameters of punctures, often as large or larger than diameters of punctures. Male: sternite 9 (fig. 152). Aedeagus (fig. 143), median lobe distinctly emarginate apically. Female: sternite 8 (fig. 134). 4.5 - 5.3 mm. . D. hummeli Bernhauer Sichuan (NO) 518 VOLKER PUTHZ 182 (181) Punctation of the fore parts coarser and denser, diameters of punc- tures on frons slightly larger than basal cross section of palpal segment 3, interstices mostly as large as, or slightly larger, than diameters of punctures. Male: sternite 9 (fig. 149).Aedeagus (figs 145, 147, 148), median lobe not emarginate apically. Female: sternite ) AS TM Wiese WS4h ANS CHO) ie So SE D. emeiensis Zheng Sichuan, Guizhou 183 (164) Abdominal punctation very dense and very fine, interstices in middle of tergites distinctly smaller than diameters of punctures, punctation on tergite 8 conspicuously coarser, at least twice as coarse as that of tergite 7 184 (185) Punctation of fore parts coarser, punctures on pronotum about as large as basal cross section of third antennal segment. Three or four incomplete oblique rugae between orange spot of elytra and suture. Male: sternite 9 with a broad, blunt tooth apicolaterally. Aedeagus as in fig. 2, Rougemont, 1986, median lobe broadly lanceolate. Female: tergite 10 subtriangular.4.5-5.8mm...... D. luteostigmaticus Rougemont China: Hongkong 185 (184) Punctation of fore parts less coarse, punctures of pronotum about as large as basal cross section of second palpal segment. About four rather complete, oblique rugae between orange spot of elytra and suture. Male: sternite 9 with a short tooth apicolaterally (fig. 4, Puthz, 1980b). Aedeagus (fig. 3, Puthz, 1980b), with apical portion of median lobe triangular. Female: tergite 10 triangular. AN OES SORIA ash sted iA ys cone aso TONO « à D. luteolunatus Puthz Vietnam; Laos; China: Guangxi [Specimens recorded from Thailand probably belong to a closely related new species. | 186 (159) Elytra variegate, without orange spots 187 (188) Punctation of pronotum very fine and sparse, punctures distinctly smaller than medial eye facets. Fourth tarsal segment with conspi- cuous bristle shoe (figs 1-3). Tergite 10 broadly rounded. Male: ster- nite 8 (fig. 9, Puthz, 1990), with a broad tooth apicolaterally. Aede- agus (fig. 6, Puthz, 1990), with median lobe broadly triangular apically. Female: sternite 8 broadly rounded, apical middle truncate or slightly emarginate medio-apically. 7.5 - 9.0 mm. . D. sichuanensis Puthz China: Sichuan 188 (187) Punctation of pronotum coarser and denser, punctures at least as large as medial eye facets. Fourth tarsal segment without bristle shoe, or only with rather thin, sparse bristles 189 (216) Punctation of pronotum as coarse as, or distinctly coarser, than that of frons 190 (215) Elytral punctation distinctly confluent, at least on some areas 191 (208) THE GENUS DIANOUS IN CHINA 519 Elytra unicolorous, without distinct, differentiated area of different colour (except an indistinctly delimited silver-bluish area on posterior half) 192 (195) Punctation of paratergites variegate, very dense anteriorly, distinctly sparser posteriorly 193 (194) Head much narrower than elytra (WH: wEl = 0.84- 0.90), antennae 194 (193) 195 (192) 196 (203) 197 (202) 198 (199) 199 (198) 200 (201) 201 (200) shorter, when reflexed extending towards posterior margin of pronotum, abdomen with faint microsculpture throughout. Male: sternite 8 with a broad, semicircular emargination in posterior 8th. (figs 84, 86). Female: sternite 8 (fig. 94). SANS IN ee RE ns SA em Mel D. rugosipennis sp. n. China: Fujian, Anhui Head broader, nearly as boad as elytra (WH: wEl = 0.97), antennae longer, when reflexed last segment extending beyond posterior mar- gin of pronotum, abdomen, except tergite 7-10, without micro- sculpture. Male unknown. Female: sternite 8 (fig. 93). SoS LUS reti re eek a Ho at D. rugipennis sp. n. China: Hunan Punctation of paratergites uniform, very dense Species with greenish-metallic shine Head broader (bEI: DH < 1.22), with larger eyes. Metasternum deeply reticulated Frons broader, medial portion posteriorly as elevated as middle. Male: sternite 9 (fig. 99), tergite 10 broadly rounded. Aedeagus (fig. 83). Female: sternite 8 (fig. 97; tergite 10 triangular. AS Ar OMI IN pepsi seta ee ae Snir D. rougemontianus sp. n. China: Sichuan Frons narrower, medial portion more elevated posteriorly than middle Antennae entirely dark. Male: medial area of metasternum completely dull. Sternite 9 (fig. 91). Aedeagus (fig. 36, Puthz, 1981a), parameres broadened apically, with about 8-12 very strong and very long medial and 8-9 fine and short dorsomedial setae. Female: sternite 8 about as in fig. 93, broader triangular apically; tergite 10 triangular. AN (Rey SES TANNIN N ae PAL cL DENE D. aereus Champion India, Nepal, Bhutan, Burma, China: Yunnan: 1 d: Ruth, 4.11.1993, G.M. de Rougemont (cR):new to China. Three outer antennal segments lighter than rest of antennae. Male: medial area of metasternum densely and deeply reticulated, but with a faint shine. Sternite 9 (fig. 100). Aedeagus (in fig. 81), parameres slender apically, with about 20 equally set, moderately strong and moderately long setae. Female: sternite 8 (fig. 98), narrower trian- gular apically; tergite 10 triangular. 3.8 - 4.7 mm...... D. ruginosus Zheng China: Sichuan 520 VOLKER PUTHZ 202 (197) Head narrower (bEl: DH > 1.26), with smaller eyes. Metasternum shallowly reticulated, strongly shining. Male unknown. Female: sternite 8 about as in fig. 97; tergite 10 triangular. BEAR RE SERIE D. brevicornis sp. n. China: Yunnan 203 (196) Species predominantly bluish (blackish blue, greenish blue), elytra with a silver-bluish shining area posteriolaterally 204 (205) Punctation of frons much finer than that of pronotum. Blackish blue. Male: sternite 9 with a (more or less) distinct tooth apicolaterally (fig. 101); tergite 10 broadly rounded. Aedeagus (fig. 85). Female: sternite 8 (fig. 88); tergite 10 slightly triangular. AO OT MR Licks Wis ACRE Sch ENI N D. atrocoeruleus sp. n. China: Yunnan 205 (204) Punctation of frons about as coarse as that of pronotum, or only slightly finer 206 (207) Larger species: 5.5 - 7.0 mm. Elytral sculpture finer, with completely bright (silvery) bluish shine. See #91......... D. subvorticosus Champion 207 (206) Smaller species: 3.7 - 5.0 mm. Elytral sculpture coarser, coarse rugae not forming complete vortex on posterior half. Male: sternite 9 about as in fig. 99. Aedeagus (fig. 4, Puthz, 1980 a) about as in fig. 83, median lobe broadly rounded apically (not triangular), parameres with strong setae also in basal half. Female: tergite 10 triangular. [see also D. karen Rougemont in part I of this article]... . D. vietnamensis Puthz Vietnam; Laos; China: Hainan, ? Yunnan 208 (191) Elytra with a differentiated area of different metallic color 209 (212) Pubescence of pronotum very long, erect, longest setae about as long as antennal segment 4 210 (211) Elytral punctation coarser, largest punctures as large as apical cross section of antennal segment 3, abdominal punctation very fine and very sparse, elytra with a distinct cupreous spot, diameter nearly as large as the length of antennal segment 3. Male: sternite 9 (fig. 138). Aedeagus (fig. 129). Female unknown. HINUNTER DIRT ee D. cyaneocupreus sp. n. China: Taiwan 211 (210) Elytral punctation less coarse, largest punctures about as large as medial cross section of antennal segment 3, abdominal punctation fine and moderately dense, elytra with indistinet cupreous spot, diameter about as large as antennal segment 11. See #87. D. yoshidai Naomi 212 (209) Pubescence of pronotum much shorter, curved distally, longest setae about as long as antennnal segment 10 213 (214) Pronotal punctation coarse, diameters of punctures as large as widest cross section of antennal segment 3, interstices smooth, elytra each Withicupreousispots See 9er ren re ee D. cupreostigma sp. n. THE GENUS DIANOUS IN CHINA 52] 214 (213) Pronotal punctation moderately fine, diameters of punctures smaller than basal cross section of antennal segment 3, interstices reticulate. SEHE... ee RN. O D. subvorticosus Champion 215 (190) Elytral punctation well-defined (rarely a few punctures slightly COAIESCENL) ASCE A: Born RA ee D. atrocyaneus sp. n. 216 (189) Punctation of pronotum less coarse than punctation of frons. Four very similar species 217 (218) Metatrochanter with distinct tooth in both sexes. See # 37 D. yunnanensis Puthz 218 (217) Metatrochanter simple 219 (222) Punctation of tergite 8 less coarse and less dense, interstices as large as half diameters of punctures or even larger, mostly with shallow reticulation. Apicolateral brush of sternum 9 strong, longer than the sternum. Female: apical emargination of sternite 8 at least as deep as wide (figs 77, 79, 80) 220 (221) Punctation slightly less dense. Blackish blue. Male: sternite 9 (fig. 75); tergite 10 broadly rounded. Aedeagus (fig. 70). Female: sternite 8 (fig. 79); tergite 10 triangular, narrowly rounded apically (about as TR) IS NN E AR RE im. OT, D. uniformis Zheng China: Sichuan 221 (220) Punctation somewhat denser. Dark blue metallic. Male: sternite 9 (fig. 78); tergite 10 (fig. 71). Aedeagus (fig. 74). Female: sternite 8 (fig. ZN atersite 10e 72) 60 ER D. acuminifer Puthz 222 (219) Punctation of tergite 8 coarser and very dense, as coarse as on elytra, interstices smaller than half diameters of punctures, deeply reticu- lated. Apicolateral brush of valvifer weak, shorter than valvifer. Female: sternite 8 (fig. 76), apical emargination much less deep; tergite 10 (fig. 73) less acute. Male: unknown. CSM IM n een ane D. angulifer sp. n. China: Taiwan 522 VOLKER PUTHZ TABLE | Geographical distribution of the Southeast-Asian Dianous based on a political map; India is listed in 3 columns: SInd = mountainous areas of southern India (Nilgiris etc.). NWInd = northern India excluding Assam (Himalayan areas) and Assam (see also figs 31 a, b). (Afgh.) = the species concerned is also known from Afghanistan. DIANOUS PASSENT ENS ZUBE Desa uil 1 | D D TD) D acuminifer Puthz ?acutus Zheng adonis sp. n. aeneus Cameron x aequalis Zheng x aereus Champion Xeno Xe NX x alcyoneus sp. n. x alternans Zheng x amamiensis Sawada x andrewesi Cameron EIER: x angulifer sp. n. x annandalei Bernhauer Re KG eX arachnipes Puthz x assamensis Cameron x ater sp. n. atrocoeruleus Sp. n. atrocyaneus Sp. n. atroviolaceus Sp. n. aurichalceus (Champion) x x azureus Champion (Afgh.) x banghaasi Bernhauer x bashanensis Zheng X bhotius Rougemont x bhutanensis Rougemont X bifoveifrons Champion RX bimaculatus Cameron KIN EX bioculatus sp. n. x bracteatus (Champion) x x brevicornis sp. n. x brevitarsis Puthz x caeruleonotatus Champion (Afgh.) x calceatus Puthz x camelus Puthz x cameroni Champion E Sax cameronianus Jarrige x carinipennis (Bernhauer) X championi Cameron x x chetri Rougemont Ru ex chinensis Bernhauer coeruleomicans Puthz coeruleotinctus Sp. n. coeruleovestitus Sp. n. consors Cameron x pi dI PI PI pd pi pi x» DIANOUS convexifrons Puthz corticicola (Puthz) cribrarius Champion cruentatus L. Benick cupreoaeneus (Champion) cupreogutta sp. n. cupreostigma Puthz cupreoviolaceus Puthz cyaneocupreus Sp. n. cyaneovirens (Cameron) cyanogaster Champion distigma Champion dubiosus sp. n. electrigutta sp. n. elegantulus Zheng emarginatus Zheng emeiensis Zheng femoralis Cameron flavoculatus Puthz flavoguttatus Puthz fluctivagus Puthz frater Cameron freyi L. Benick gemmosus Sp. n. gongen Watanabe gonggamontis Sp. n. gracilipes Champion gracilis Puthz grandistigma sp. n. gregarius Rougemont hainanensis Puthz hammondi Rougemont haraldi sp. n. hirsutus Rougemont hummeli Bernhauer inaequalis Cameron inaequalis laevior ssp. n. inconspicuus Rougemont irididolor (Scheerpeltz) iwakisanus Watanabe Japonicus Sawada kabakovi Puthz karen Rougemont klapperichi L. Benick lahu Rougemont lasti Puthz latitarsis L. Benick lobatipes sp. n. (Afgh.) DER D rg THE GENUS DIANOUS IN CHINA ans aN ee Sr OS}, ON >» Se Sb BIS Fe 0) — Ze Sep OMS ie m mi mx» Mn DI DO DI »” 5 DT D & 524 VOLKER PUTHZ DIANOUS ln mem gi ans nu N W (ay {=} LIST AZIZ | mm. DS POSE sex D @ eo A puren 5 pm © = lobigerus Champion loebli Rougemont luteoguttatus Champion luteolunatus Puthz luteostigmaticus Rougemont margaretae Rougemont martensi Rougemont mendax Sp. n. meo Rougemont minor Champion morimotoi Naomi moritai Naomi nagamontium Puthz nepalensis Rougemont niger Rougemont nigrocyaneus Puthz nigrovirens (Fauvel) nilgiriensis Puthz nokrekensis Puthz obliquenotatus Champion obscuroguttatus Cameron ocellatus Cameron ocellifer sp. n. oculatipennis Puthz pallitarsis L. Benick psilopterus L. Benick punctiventris Champion pykaranus (Cameron) radiatus Champion reformator Rougemont robustus Cameron rougemontianus Sp. n. ruginosus Zheng rugipennis Sp. n. rugosipennis Sp. n. scabricollis Champion schoenmanni sp. n. senex Sp. n. shan Rougemont shibatai Sawada siamensis Rougemont sichuanensis Puthz siwalikensis Cameron socius Zheng spiniventris Puthz srivichaii Rougemont strabo Puthz subtortuosus Champion x pi PI PI * * M pi pi THE GENUS DIANOUS IN CHINA 525 DIANOUS an mn — pae Set a {= clog) SISI »oen SES} Es een DESIO eS] Dre "y ans subvorticosus Champion Kx x sucinigutta sp. n. x suciniguttatus sp. n. x sucininotatus sp. n. x sulcatipennis Puthz x sulcipennis Puthz x taiwanensis Puthz x tonkinensis (Puthz) X X X tortuosus Champion KHK tortus Cameron X tumidifrons Puthz x uncinipenis Puthz x uniformis Zheng variegatus sp. n. versicolor Cameron x x verticosus Eppelsheim Xx x x vietnamensis Puthz x x violaceus Puthz viridicatus Naomi x viridicupreus Rougemont x viridipennis Cameron x viriditinctus (Champion) x x wittmeri Rougemont X yangae sp. n. X yao Rougemont EX x yoshidai Naomi x yunnanensis Puthz x X m M M di Di OK TABLE 2 Distribution of Dianous-species in China and neighbouring countries; for abbreviations see the introduction (p. 427). No species is known from biogeographic unit 6 which is therefore not unit | unit 2 unit 3 unit 4 À units A3BC#DEFG4#HIJK#LMNOPQR#SS5S6TU7 D | DIANOUS 1 acuminifer Puthz x ? acutus Zheng x adonis sp. n. x aequalis Zheng x x aereus Champion X X X X alternans Zheng x amamiensis Sawada x andrewesi Cameron xx x angulifer sp. n. x arachnipes Puthz ater sp. n. x atrocoeruleus sp. n. x atrocyaneus sp. n. x atroviolaceus sp. n. x banghaasi Bernhauer x NAME xX xX x x x bashanensis Zheng x x bioculatus sp. n. x x 526 VOLKER PUTHZ Biogeographic unit | unit 2 unit 3 unit 4 unit 5 DIANOUS 12A3BC#DEFG4#HIJK#LMNOPQR#S56TU7 brevicornis sp. n. x chetri Rougemont SIG SE chinensis Bernhauer ? Xa eX x x x coeruleomicans Puthz x coeruleotinctus sp. n. coeruleovestitus sp. n. x cupreogutta sp. n. cupreostigma Puthz x cupreoviolaceus Puthz x cyaneocupreus sp. n. x dubiosus sp. n. x a electrigutta sp. n. x elegantulus Zheng emarginatus Zheng emeiensis Zheng freyi L. Benick ) gemmosus Sp. N. x gonggamontis sp. n. x gongen Watanabe x grandistigma Sp. n. EX hainanensis Puthz x hummeli Bernhauer X inaequalis Champion DE SC inaequalis laevior ssp. n. x iwakisanus Watanabe x Japonicus Sawada x karen Rougemont > 02) klapperichi L. Benick x latitarsis L. Benick IGIENICI DELI lobatipes sp. n. x luteoguttatus Champion XX x luteolunatus Puthz x x luteostigmaticus Rougem. x mendax sp. n. x morimotoi Naomi x moritai Naomi x nigrocyaneus Puthz x ocellatus Cameron x x ocellifer sp. n. x oculatipennis Puthz punctiventris Champion x x rougemontianus sp. N. x ruginosus Zheng x rugipennis sp. n. x rugosipennis Sp. n. x x schoenmanni sp. n. x senex sp. n. x shibatal Sawada x sichuanensis Puthz x socius Zheng x spiniventris Puthz srivichaii Rougemont x strabo Puthz x subvorticosus Champion x sucinigutta sp. n. x suciniguttatus SP. n. x sucininotatus SP. n. x taiwanensis Puthz x tonkinensis (Puthz) EX uniformis Zheng x variegatus Sp. n. versicolor Cameron XIX verticosus Eppelsheim Dx viernamensis Puthz x violaceus Puthz x viridicatus Naomi x yangae sp. n. x yao Rougemont x x yoshidai Naomi x yunnanensis Puthz XX x x bal x lai 2 m ms x MK 2 ” Pas x De Scie oe ISIDE OX x X THE GENUS DIANOUS IN CHINA 527 TABLE 3 Distribution of Dianous in Taiwan according to districts (hsien) (see also map 3, fig. 31 c). I: Taipeh, 2: Ilan, 3: Hsinchu, 4: Taoyuan, 5: Miaoli, 6: Taichung, 7: Changhua, 8: Nantou, 9: Chiayi, 10: Yunlin, 11: Tainan, 12: Kaohsiung, 13: Pingtung, 14: Taitung, 15: Hualien. (N°) Dianous in Districts of Taiwan l BR SCT RS Oe NOS 12468) 14109465 acuminifer Puthz x Hi X angulifer sp. n. x arachnipes Puthz EX x x XX x x ater sp. n. x atrocyaneus sp. n. X cupreogutta sp. n. x x cupreostigma Puthz x cyaneocupreus sp. n. x dubiosus sp. n. ? electrigutta sp. n. inaequalis laevior ssp. n. x luteoguttatus Champion mendax sp. n. x sucinigutta sp. n. X x x sucininotatus sp. n. taiwanensis Puthz XIX x XX x Pi KK pi 16 species Sona UC 2 2 CS Pai D |» > 528 VOLKER PUTHZ Fics 1-6 Figs 1-3: Tarsal shoe of Dianous sichuanensis Puthz (2-PT): Apex of metatarsus, lateral aspect (1), ventral aspect (2), detail of latter (3). — Figs 4-6: Metatarsus of Dianous gracilipes Champion (d, Naga Hills): Lateral aspect of apex (4), ventral aspect of segments 2-4 with willow-leave- like setae (5), ventral aspect of segment 5 between claws (6). THE GENUS DIANOUS IN CHINA 529 NM 7 8 A A & 10 11 12 Fics 7- 12 Outline of tarsal segment 4: Dianous versicolor Cameron (Emei Shan, metatarsus of d, 7), D. azureus Champion (Tanakpur, metatarsus of à, 8), D. mendax sp. n. (2-PT, metatarsus, 9), D. banghaasi Bernhauer (Emei Shan, protarsus and metatarsus of 2, 10, 11), D. cupreostigma Puthz (2-PT, metatarsus, 12).- Scale = 0.1 mm. 530 VOLKER PUTHZ Fics 13-16 Figs 13, 14: Dianous taiwanensis Puthz: Male sternite 9, dorsal aspect, with apically spinulose arcuate cuticular laminae. — Figs 15, 16: Dianous taiwanensis Puthz: Male sternite 9, dorsal aspect, apically spinulose arcuate cuticular laminae and glandular openings. THE GENUS DIANOUS IN CHINA Fics 17-20 Figs 17, 18: Dianous taiwanensis Puthz: Male sternite 9 (T 152), dorsal aspect, apically spinulose arcuate cuticular laminae and glandular openings. — Figs 19, 20: Dianous taiwanensis Puthz (T 162): Apical portion of the median lobe of the aedeagus (19), Dianous elegantulus Zheng (Wulingyuan): Left elytron with dense pubescence in characteristic arrangement (20). 332 VOLKER PUTHZ Fics 21-24 Figs 21, 22: Dianous banghaasi Bernhauer (env. Guangping): Pronotum (dorsal and tangential aspect) with long, erect pubescence (compare figs 23, 24). — Figs 23, 24: Dianous gonggamontis sp.n. (PT): Pronotum (dorsal and tangential aspect) with moderately long, curved/ recumbent pubescence (compare figs 21, 22). 533 THE GENUS DIANOUS IN CHINA Fics 25-28 , 26: Dianous ocellatus Cameron (Xishuangbanna): Le ft elytron with rugose-coalescent Habitus s oO 28: Dianous gemmosus sp.n. (PT): Figs 25 howing coarse vorticose 5 sculpture. — Figs 27 sculpture on the elytra. 534 VOLKER PUTHZ Fics 29, 30 Large numbers of Dianous (f. e. D. freyi L. Benick) in small humid places at the Emei Shan, Sichuan (photograph by Zheng Fa-ke). THE GENUS DIANOUS IN CHINA PN, Ries 3ira The genus Dianous Leach in Southeast Asia. Numbers indicate the number of species recorded from the respective countries, islands or provinces (Yunnan and Sichuan): 37 from NW-India and from Nepal, 32 from Assam, 30 from Sichuan, 27 from Yunnan, 17 from China (excluding Sichuan, Yunnan, Hainan and Taiwan), 16 from Taiwan, 15 from Thailand, 14 from Vietnam, 13 from Borneo, 11 from Laos, 10 from Bhutan, 7 from Burma, 6 from Hainan, 5 from Madras and from Honshu, 4 from Punjab, Sumatra and from Java, 3 from Afghanistan and from Malaysia, 2 from Afghanistan, Bali, Luzon, Mindanao, Ryukyu, | from Siberut, Timor, Sulawesi, Ceram, Negros, Basilan, Cebu, Mindoro, Kyushu, Shikoku. The black line delimites the distribution of group I of the genus, the dotted line that of group II. The open circle shows an isolated record from the Altai mountains. LL = Lydekker’s line. 536 VOLKER PUTHZ Fic. 31 b The genus Dianous Leach in China and neighbouring countries: numbers indicate the number of species recorded from the individual provinces of China and the surrounding countries (see also table 2). THE GENUS DIANOUS IN CHINA Fic. 31 c 537 The genus Dianous Leach in Taiwan: numbers indicate the number of species recorded from the individual districts (see also table 3). 538 VOLKER PUTHZ 32 Fics 32-37 Sternite 8 of the female (32) and coloration of the elytra (33-37, see descriptions): Dianous versicolor Cameron (Emei Shan, 32), D. coeruleovestitus sp.n. (PT, 33), D. alternans Zheng (Emei Shan, 34), D. schoenmanni sp.n. (HT, 35), D. elegantulus Zheng (Sioxiyou, 36), D. adonis sp.n. (HT, 37). THE GENUS DIANOUS IN CHINA 539 Fics 38-44 Ventral aspect of the aedeagus (38-40, 44) and ¢ sternite 9 (41-43): Dianous alternans Zheng (Emei Shan, 38), D. coeruleovestitus sp.n. (HT, 39, 41), D. coeruleotinctus sp.n. (PT, 40), D. schoenmanni sp. n. (HT, 42), D. elegantulus Zheng (Emei Shan, 43, 44). Scale = 0.1 mm (40 = 39, 41; 42 = 43, 44). 540 VOLKER PUTHZ 47 46 48 50 51 52 \ 774 53 54 55 y LS 56 57 58 Fics 45-58 Stronger sclerotized portion of the internal sac (45, 46), outline of the 9 sternite 8 (total: 47; posterior portion: 48, 51, 52, 54-58), outline of the anterior portion of the aedeagus (49), posterior portion the ¢ sternite 9 (50, 53): Dianous alternans Zheng (Emei Shan, 45, 50, 56-58), D. chetri Rougemont (Emei Shan, 46), D. srivichaii Rougemont (Tailand: Mae Yao, 47), D. elegantulus Zheng (Emei Shan, 48, 51, 54), D. fauveli sp.n. (HT, 49), D. coeruleotinctus sp.n. (PT, 52, 53, 55). Scale = 0.1 mm (45= 46, 49; 47= 48, 51, 52, 54-58; 50= 53). THE GENUS DIANOUS IN CHINA 54] 60 62 | ai i 64 | a) Al 66 68 KK Fics 59-69 Outline of the posterior portion of the 9 sternite 8 (59-67), outline of the posterior portion of the 3 sternite 9 (68), ventral aspect of the aedeagus (69): Dianous inaequalis Champion (Nepal: Basantapur: 59; Yunnan: Lugu Lake: 60), D. inaequalis laevior ssp. n. (PTT: Taiwan loc. 87: 61- 69). Scale = 0.1 mm (59 = 60-67; 69 = 68). 542 VOLKER PUTHZ Fics 70-80 Anterior portion of the aedeagus, ventral aspect (70, 74), tergite 10 (71, 72, 73), outline of the posterior portion of the ? sternite 8 (76, 77. 79, 80), outline of the posteror portion of the & sternite 9 (75, 78): Dianous uniformis Zheng (Emei Shan, 70, 75, 79), D. acuminifer Puthz (3 Taiwan T 50: 71; © Taiwan: Sun Lin Sea: 72; T 50: 77; T 60: 74, 78; Sun Lin Sea: 80), D. angulifer sp.n. (2 HT: 73, 76). Scale = 0,1 mm (70= 74, 75, 78; 71 = 72, 73, 76, 77, 79, 80). THE GENUS DIANOUS IN CHINA 543 — FIGs 81-87 Ventral aspect of the aedeagus and the anterior portion of the aedeagus, respectively: Dianous ruginosus sp. n. (HT, 81), D. karen Rougemont (Thailand: Doi Suthep, 82), D. rougemontianus sp. n. (PT, 83), D. rugosipennis sp. n. (HT, 84, 86), D. atrocoeruleus sp. n. (PT, 85), D. luteo- guttatus Champion (Nepal: Induwa Khola Valley, 87). Scale = 0;1 mm (81 = 86; 82 = 83: 85 = 87). 544 VOLKER PUTHZ 88 91 92 vi 93 94 v = 97 98 101 Fics 88-101 Outline of the © sternite 8 (88, 93, 94, 97, 98), outline of the posterior portion of the ¢ sternite 9 (89-91, 95, 96, 99-101), outline of the valvifer (92): Dianous atrocoeruleus sp.n. (PT, 88, 101), D. atroviolaceus sp.n. (PT, 89), D. luteoguttatus Champion (Nepal: Induwa Khola Valley, 90), D. aereus Champion (Ruili, 91), D. rugipennis sp.n. (HT, 92, 93), D. rugosipennis sp. n. (PT, 94, 95), D. karen Rougemont (Thailand: Doi Suthep, 96), D. rougemontianus sp. n. (PTT, 97, 99), D. ruginosus Zheng (Emei Shan 98, 100). Scale = 0.1 mm (88 = 93, 94, 97, 98: 89 = 90, 91, 95, 96, 99-101). THE GENUS DIANOUS IN CHINA 545 102 104 106 107 108 109 110 112 Fics 102-112 Elytra of various Dianous species (including the variability of D. taiwanensis): D. chinensis Bernhauer (PT, 102), D. yangae sp. n. (PT, 103), D. banghaasi Bernhauer (Emei Shan, 104), D. emeiensis Zheng (Huaxi, 105), D. bimaculatus Cameron (Nepal: env. Shivalaya, 106), D. taiwa- nensis Puthz (T 60: 107; T 87: 108, 112; T 48: 109; T 54: 110; T 152: 111). a VOLKER PUTHZ 113 114 = 121 117 122 ‘ Rainy tp Ÿ 120 154 os Fics 113-125 Ventral aspect of the aedeagus (apical portion: 113, complete: 123, outline of the apical portion of the median lobe: 121, 122, 124), mesotarsi (114). outline of the © sternite 8 (115), outline of the posterior portion of the ¢ sternite 9 (116- 120). Dianous atrocyaneus sp. n. (HT, 113, 116), D. gemmosus sp. n. (PT: 114, 115; HT: 117, 123), D. chinensis Bernhauer (PT: 118, 121; Wulingyuan: 119, 122), D. yangae sp. n. (PTT, 120, 124, 125). Scale = 0.1 mm (113 = 121- 125; 116 = 117- 120). THE GENUS DIANOUS IN CHINA 547 132 : ©. 1 Fics 126-142 Outline of the anterior portion of the median lobe or aedeagus (126-130), outline of the © sternite 8 (131-134), outline of the posterior portion of the 4 sternite 9 (135-138), segments 4 and 5 of the metatarsi (139-142): Dianous senex sp. n. (PT, 126, 135), D. banghaasi Bernhauer (PT, 127: „Kanton“, 135), D. aequalis Zheng (,Kanton“, 128, 132, 136), D. cyaneocupreus sp.n. (HT, 129, hummeli Bernhauer (HT of D. szechuanus, 134, HT of D. hummeli, 139), D. emeiensis Zheng (Huaxi, 140). Scale = 0.1 mm (126 = 127-129; 130 = 135-138; 131 = 132-134; 139 = 140-142. 548 VOLKER PUTHZ 150 149 av 151 153 152 Fics 143-153 Apical portion of the aedeagus without internal structures (143-145, 147, 148), Outline of the 9 sternite 8 (146; posterior portion: 153), outline of the posterior portion of the d sternite 9 (149- 152: Dianous hummeli Bernhauer (HT, 143, 152), D. bashanensis Zheng (Wulingyuan, 144, 150), D. emeiensis Zheng (Huaxi, 145-149), D. andrewesi Cameron (Naga Hills, 151), D. yangae sp. n. (PT, 153). Scale = 0.1 mm (143 = 144, 145, 147-152; 146 = 153). THE GENUS DIANOUS IN CHINA 549 154 155 158 159 164 165 166 167 Fics 154-167 Outline of the anterior portion of the median lobe (154-160), outline of the posterior portion of the d sternite 9 (161-163), outline of the 9 sternite 8 (164-167): Dianous atroviolaceus sp.n. (PTT, 154, 164), D. bioculatus sp.n. (PT, 155, 162, 165), D. socius Zheng (Eimei Shan, 156, 163, 167), D. taiwanensis Puthz (T 60: 157; HT: 158; T 24: 159: Alishan: 160; T 50: 161, 166). Scale = 0-1 mm (154 = 155-160; 161 = 162, 163; 164 = 165-167). ° 550 VOLKER PUTHZ 168 169 173 72 175 176 Fics 168-176 Elytra of Dianous hainanensis Puthz (HT, 168), D. flavoculatus Puthz (PT, 169), D. strabo Puthz (PT, 170), S. grandistigma sp. n. (PT, 171), D. distigma Champion (PT, 172), D. suciniguttatus sp.n. (HT, 173), D. sucininotatus sp.n. (left and right elytron taken from 2 PTT, T 107, 174), D. sucinigutta sp.n. (left and right elytron taken from 2 PTT, T 24, 175), D. electrigutta sp.n. (PT, 176). THE GENUS DIANOUS IN CHINA 55] Fics 177-191 Figs 177-181: Ventral aspect of the aedeagus, internal sac and setae partly ommitted: Dianous ocellatus Cameron (PT, 177), D. grandistigma sp. n. (HT, 178), D. emarginatus Zheng (Heishui, 179), D. ocellifer sp. n. (HT, 180), D. distigma Champion (HT, 181). Scale = 0.1 mm. — Figs 182- 191: Outline of the apical portion of the median lobe (ventral aspect; 182-190), posterior portion of the valvifer (191): Dianous ocellatus Cameron (PT: 182; Ruli: 183), D. variegatus sp. n. (HT, 184), D. verticosus Eppelsheim (setae ommitted, Burma: Irawaddi, 185), D. grandistigma sp.n. (HT, 186; PT, 191), D. distigma Champion (setae ommitted, HT, 187), D. emarginatus Zheng (Heishui, 188), D. ocellifer sp. n. (HT, 189), D. oculatipennis Puthz (Xishuangbanna, 190). Scale = 0.1 mm (182 = 183-190). 552 VOLKER PUTHZ 216 Fics 192-217 Figs 192-208: Valvifer (192), outline of the 2 sternite 8 (193-199), outline of the © tergite 10 (200- 201), outline of the posterior portion of the ¢ sternite 9 (202-208): Dianous grandistigma sp. n. (PT, 192, 194; HT, 205), D. variegatus sp.n. (PT, 193, 201; HT, 204), D. verticosus Eppelsheim (HT, 195, 200), D. ocellatus Cameron (Xishuangbanna, 196, 202; Ruli, 203), D. emarginatus Zheng (Heishui, 197, 206), D. ocellifer sp. n. (PT, 198, 207), D. oculatipennis sp.n. (Xishuangbanna, 199, 208). Scale = 0.1 mm (193 = 194- 201; 202 = 203-208). — Figs 209-217: Ventral aspect of the aedeagus and the anterior portion of the median lobe, respectively (209- 214, 217), outline of the 2 sternite 8 (215, 216): Dianous ater sp. n. (PT, 209, 215), D. sucinigutta sp. n. (PTT, T 24, T 24, 210, 211), D. sucininotatus sp. n. (PT, 212), D. electrigutta sp. n. (PT, 213), D. cupreogutta sp. n. (PT, 214), D. mendax sp. n. (PT, 216; HT, 217). Scale = 0.1 mm (209 = 210- 213; 214 = 217; 215 = 216). THE GENUS DIANOUS IN CHINA 553 225 226 227 Fics 218-227 Figs 218-221: Anterior portion of the aedeagus (218, 221), metatarsus (219, 220): Dianous sucinigutta sp. n. (PT, T 29, 218), D. sucininotatus sp. n. (PT, 219), D. ater sp. n. (PT, 220), D. suciniguttatus sp. n. (HT, 221). Scale = 0.1 mm (218 = 221; 219 = 220). — Figs 222-227: Outline of the anterior portion of the median lobe (222-226), outline of the © sternite 8 (227): Dianous freyi L. Benick (Emei Shan: 222; Wulingyuan: 223; Kuling: 225); D. lobatipes sp.n. (HT, 224), D. klapperichi L. Benick (Emei Shan: 226); D. cupreostigma sp. n. (PT, 227). Scale = 0.1 mm O22 =223-226). 554 VOLKER PUTHZ 237 ) Coe f / 238 241 240 242 Fics 228-243 Figs 228-238: Outline of the posterior portion of the d sternite 9 (233-238 without asacl): Dianous cupreogutta sp. n. (PT, 228), D. ater sp. n. (PT, 229), D. freyi L. Benick (Emei Shan, 230), D. klapperichi L. Benick (Emei Shan, 231), D. mendax sp. n. (HT, 232), D. sucinigutta sp. n. (PT, T 24: 233; T 13: 234), D. sucininotatus sp. n. (PT, 235), D. electrigutta sp. n. (PT, 236), D. suciniguttatus sp. n. (HT, 237), D. lobatipes sp. n. (HT, 238). 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INDEX New names in bold letters, synonyms in italics acuminifer Puthz 423, 446 alternans Zheng 423, 434 acutus Zheng 425, 478 amamiensis Sawada 425, 429 adonis sp.n. 423, 443 amicus Puthz 420 aeneus Cameron 424 anatolicus Korge 426 aequalis Zheng 425, 466 andrewesi Cameron 424, 505 aereus Champion 424, 427, 453,519 angulifer sp. n. 423, 447 alcyoneus sp. n. 425, 485 annandalei Bernhauer 423 558 VOLKER PUTHZ arachnipes Puthz 423, 429, 433 dajak Puthz 420 araeocerus (L. Benick) 420 davaomontium (Puthz) 420 assamensis Cameron 423 distigma Champion 425 ater sp.n. 425, 490 dubiosus sp. n. 425, 467 atroeruleus sp. n. 425, 484 electrigutta sp.n. 426, 429, 496 atrocyaneus sp.n. 425, 429, 458 elegans Khnzorian 426 atroviolaceus sp.n. 425, 472 elegantulus Zheng 423, 441 aurichalceus (Champion) 420 emarginatus Zheng 425, 482 azureus Champion 424 emeiensis Zheng 425, 470 baliensis Rougemont 420 fauveli sp. n. 423, 446 banghaasi Bernhauer 425, 429, 465 femoralis Cameron 424 bashanensis Zheng 425, 471 flavicoxatus (L. Benick) 420 benicki Puthz 420 flavoculatus Puthz 424 bhotius Rougemont 420 flavoguttatus Puthz 423 bhutanensis Rougemont 420 fluctivagus Puthz 420 bifoveifrons Champion 423 frater Cameron 424 bilunatus Puthz 474 freyi L. Benick 426, 429, 498 bimaculatus Cameron 424, 427 gemmosus sp.n. 425, 459 bioculatus sp. n. 425, 475 gongen Watanabe 426 boops Puthz 420 gonggamontis sp.n. 425, 473 bracteatus (Champion) 420 gracilipes Champion 423 brevicornis sp.n. 424, 453 gracilis Puthz 424 brevitarsis Puthz 423 grandistigma sp. n. 425, 480 burckhardti Puthz 420 gregarius Rougemont 423 caeruleoguttatus Cameron 439 hainanensis Puthz 424 caeruleonotatus Champion 423 hammondi Rougemont 425, 427 calceatus Puthz 423 haraldi sp. n. 421, 432 camelus Puthz 423 hirsutus Rougemont 421, 427 cameroni Champion 425 hummeli Bernhauer 425, 469 cameronianus Jarrige 423 hygrobius (L. Benick) 421 carinipennis (Bernhauer) 420 inaequalis Champion 423, 429, 439 cebuensis sp. n. 420, 431 inaequalis laevior ssp. n. 423, 429, 439 chalybaeus LeConte 425, 428 inconspicuus Rougemont 421 championi Cameron 424 iridicolor (Scheerpeltz) 421 chetri Rougemont 423, 433, 436 iwakisanus Watanabe 426 chinensis Bernhauer 425, 461 jaechi Puthz 421, 429 coeruleomicans Puthz 423 japonicus Sawada 426, 507 coeruleotinctus sp. n. 423, 437 javanicola Puthz 421 coeruleovestitus sp. n. 423, 436 kabakovi Puthz 423, 427 coerulescens Gyllenhal 426, 428 karen Rougemont 424, 452 coerulescens anatolicus Korge 426 kinabalumontis Puthz 421 coerulescens elegans Khnzorian 426 klapperichi L. Benick 426, 499 concretus Puthz 420, 426 lahu Rougemont 421 consors Cameron 423 lasti Puthz 421 convexifrons Puthz 423 latitarsis L. Benick 423, 433 corticicola (Puthz) 420, 426 lividus (L. Benick) 421 cribrarius Champion 424, 427 lobatipes sp. n. 426, 487 cruentatus L. Benick 424 lobigerus Champion 424, 427 cupreoaeneus (Champion) 420 loebli Rougemont 421 cupreogutta sp. n. 426, 488 loeblianus Puthz 421 cupreostigma Puthz 426, 499 luteoguttatus Champion 424, 429, 449 cupreoviolaceus Puthz 424 luteolunatus Puthz 425, 484 cyaneocupreus sp. n. 425, 468 luteostigmaticus Rougemont 425 cyaneovirens (Cameron) 420 malayanus Cameron 423, 438 cyanogaster Champion 423 margaretae Rougemont 423 THE GENUS DIANOUS IN CHINA 559 martensi Rougemont 421 mendax sp. n. 426, 500 meo Rougemont 421,433 minor Champion 424, 427 miripes Rougemont 433 morimotoi Naomi 426 moritai Naomi 423, 429 nagamontium Puthz 421 naicus Puthz 420, 426 nepalensis Rougemont 421 niger Rougemont 421 nigrocyaneus Puthz 423 nigrovirens (Fauvel) 421 nilgiriensis Puthz 421 nitidulus LeConte 426, 428 nokrekensis Puthz 421 obliquenotatus Champion 423, 427 obscuroguttatus Cameron 423 ocellatus Cameron 425, 478 ocellifer sp. n. 425, 478 oculatipennis Puthz 425, 486 pallitarsis L. Benick 423 philippinus Puthz 421 pilosus Champion 465 ponticus Fagel 423 psilopterus L. Benick 425 puberulus L. Benick 445 punctiventris Champion 423, 429, 445 puthzi Lundgren 420, 426 pykaranus (Cameron) 421 radiatus Champion 424 reformator Rougemont 421 robustus Cameron 423, 426 rougemonti Puthz 421 rougemontianus sp.n. 424, 450 ruginosus Zheng 424, 454 rugipennis sp.n. 424, 455 rugosipennis sp.n. 424, 456 saxicola (Puthz) 421 scabricollis Champion 423, 427 schillhammeri Puthz 421 schoenmanni sp. n. 423, 444 semicoeruleus (Cameron) 421 senex sp.n. 425, 464 septentrionalis Naomi 507 shan Rougemont 421 shibatai Sawada 426 siamensis Rougemont 424, 427 siberutensis Puthz 421 sichuanensis Puthz 423 siwalikensis Cameron 423, 427 smetanai Puthz 420, 426 socius Zheng 425, 477 spiniventris Puthz 424 srivichaii Rougemont 423, 427, 505 strabo Puthz 424, 427 striatellus (L. Benick) 412 subtortuosus Champion 425 subvorticosus Champion 425 sucinigutta sp.n. 426, 429, 492 suciniguttatus sp. n. 426, 429, 497 sucininotatus sp.n. 426, 495 sulcatipennis Puthz 426 sulcipennis Puthz 421 szechuanus Bernhauer 469 taiwanensis Puthz 425, 429, 474 tiomanensis Puthz 421 tonkinensis (Puthz) 421, 501 tortuosus Champion 425 tortus Cameron 425 tumidifrons Puthz 423 uniformis Zheng 423, 448 uncinipenis Puthz 421 variegatus sp.n. 425, 481 versicolor Cameron 423, 433 verticosus Eppelsheim 425, 482 vietnamensis Puthz 424, 452 violaceus Puthz 424 viridicatus Naomi 426 viridicupreus Rougemont 421 viridipennis Cameron 424 viriditinctus Champion 421 wittmeri Rougemont 421 yangae sp.n. 425, 462 yao Rougemont 421,431 yoshidai Naomi 425, 429, 459 yunnanensis Puthz 423, 427 shit] #41} at | Er pat pipi a iger “ici ays at tre Ar ack ni inf BE, © 4 eee | ve a Na i REVUE SUISSE DE ZOOLOGIE 107 (3): 561-577; septembre 2000 Contribution to the systematics of the genus Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar J.-L. GATTOLLIAT & M. SARTORI Museum of Zoology, P.O. Box 448, CH-1000 Lausanne 17, Switzerland; e-mail: Jean-Luc Gattolliat @ie-zea.unil.ch Contribution to the systematics of the genus Dabulamanzia (Ephemer- optera: Baetidae) in Madagascar. - At the present time, the Afro- Malagasy genus Dabulamanzia is represented at Madagascar by two species. Three new species D. gladius, D. gigantea and D. concolarata are described at the larval stage. D. gladius and D. gigantea present a high degree of specialisation of the mandibles and labrum that clearly distinguish them from any other species of Dabulamanzia. D. concolorata also owns characteristic mouthparts, especially the labrum. A fourth species is described at the imaginal stage, but unamed as long as its larva remains unknown. The relative position of these new species is discussed. Key-words: Ephemeroptera - Baetidae - Dabulamanzia - new species - Madagascar. INTRODUCTION The genus Dabulamanzia has been recently erected for some species previously assigned to the tarsale group of Afroptilum Gillies (Lugo-Ortiz & McCafferty, 1996b). This genus is well defined at the larval stage by the following apomorphies: bulbous segment 3 of the labial palp and a small proximal arc of setae on the tibiae; at the imaginal stage, hindwing with a single hooked spur and a well-developed apophysis at the base of the first segment of the gonopod are the main characters. Some of these characters suggest that Dabulamanzia is related to the Cloeodes complex, even if the imaginal stage differs greatly by the number of intercalary veins, the presence of hindwing and the shape of the gonopods (Lugo-Ortiz & McCafferty, 1996a) . The first species of Dabulamanzia from Madagascar has been recently described (Lugo-Ortiz & McCafferty, 1997c). Its name, D. improvida indicates the unexpected discovery of the genus in Madagascar. Since then, an other species has been found in this Island (Gattolliat et al., 1999). Herein we describe three new species and we discuss their position within the genus. The holotypes and some of the paratypes are housed in the Museum of Zoology, Lausanne, Switzerland. Other paratypes are deposited in the Museum National d'Histoire Naturelle, Paris. Manuscript accepted 11.04.2000 562 J.-L. GATTOLLIAT & M. SARTORI Dabulamanzia gladius Gattolliat sp. n. Holotype. Larva, P0861, Madagascar, Rianila Bas., unnamed riv., Loc. road to Lakato, Long. 48°21'48" E, Lat. 19°02'40" S, Alt.1050m, 8.4.1999. J.-L. Gattolliat and N. Raberiaka. Paratypes. Two larvae same data as holotype. Two larvae, P0693, same locality as holotype, 22.4.1997. J.-L. Gattolliat, C. Rochat and N. Raberiaka. Three larvae, P0862, Mada- gascar, Rianila Bas., tributary riv. to Sahatandra Riv., Loc.near Ambalafotsy, road to Lakato. Long. 48°21'51" E, Lat. 19°02'22" S, Alt.1050m, 8.4.1999. J.-L. Gattolliat and N. Raberiaka. Other material. One female larva (91b), Madagascar, Manampanihy Bas., tributary riv. of Manampanihy Riv., Loc. Fenoevo, Long. 46°53'39" E, Lat. 24°41'00" S, Alt. 72m, 15.4.1992. J.-M. Elouard. One larva, P0814, Madagascar, Antongombato Bas., Makis Riv., Loc. 100m downstream of the Great Wasterfall, Long. 49°10'14" E, Lat. 12°29'17" S, Alt. 675m, 22.3.1999. J.-L. Gattolliat and Z. Rabeantoandro. LARVA Maximal length (no mature specimens): Body 6.7 mm. Cerci and terminal filament broken. Head. Coloration almost uniformly light yellow, except brown between the eyes without vermiform marking on vertex and frons. Antennae pale light yellow. Eyes and ocelli black. Labrum (Fig. la) rectangular, with distal margin almost straight, with two kinds of setae, one row of feathered setae (Fig. Ib) and one row of multifid setae (Fig. Ic); dorsally with a continuous arc of about 20 long setae, abundant setae in the proximal half; without setae ventrally. Hypopharynx as in figure 2, lingua with minute thin setae, superlinguae well developed and clearly separated of the lingua. Right mandible (Fig. 3) with two sets of incisors, the outer formed only by one single well-developed and laterally reinforced tooth and the inner one reduced to a single small tooth; prostheca long and thin, without apical teeth; length of the tuft of setae between prostheca and mola reducing toward the mola; tuft of small setae near the mola well-developed; tuft of setae at the apex of the mola reduced to 2 or 3 setae; basal half without setae dorsally. Left mandible (Fig. 4) with incisors fused in a single tooth; prostheca with 4 teeth, the apical one much more developed; length of the tuft of setae between prostheca and mola reducing toward the mola; tuft of setae at the apex of the mola reduced to 3 setae; basal half without setae dorsally. Maxillae (Fig. 5) with 4 teeth, the distal one opposed to the three others; 2 rows of setae, the first one formed by abundant small setae and the second by long stout setae ending with 4 twice as long as the others, without pectinate or spine-like setae in the middle of the range: 6 to 7 setae at the base of the galea roughly arranged in a row; 1 single small seta perpendicularly to the margin of the galea: palp 2-segmented as long as the galealacinia, first segment 1.4 time shorter than the second; second segment ending with a small rounded protuberance; thin setae on the external margin of the first and second segments, especially numerous at the apex of the second. Labium (Fig. 6) with glossae subequal in length to paraglossae, and more slender than them: apical half of glossae with stout setae, long setae randomly distributed on the basal half of the ventral side; paraglossae apically rounded, with 2 rows of simple setae; one simple long seta on the margin of the paraglossae. Labial palp THREE NEW MALAGASY SPECIES OF DABULAMANZIA 563 Fics 1 to 6. Larval structures of Dabulamanzia gladius : la : labrum (left : ventral; right : dorsal). lb : multifid seta of the labrum. Ic : feathered seta of the labrum. 2 : hypopharynx. 3 : right mandible. 4 : left mandible. 5 : left maxilla. 6 : labium. 564 J.-L. GATTOLLIAT & M. SARTORI 3-segmented; first segment stout, 1.3 time smaller than the second and third combined; second segment enlarged at the apex, row of about 4 setae; third segment very broad, truncated and incurved at the apex, almost completely covered with setae. Thorax. Coloration light brown. Hindwing pad present. Legs yellow, except the apex of femora and tibia light brown. Forelegs (Fig. 7a) with coxa with few setae. Femora dorsally with a row of 12 long setae, only 2 of them in the distal half, apical half with thin short setae; 6 submarginal blunt setae; femoral patch of 4 spatulated setae; numerous short setae on the ventral margin. Tibiae dorsally with only short and thin setae, small subproximal arc of short setae; apex dorsally with a single long curved seta; ventral margin with abundant setae; tibio-patellar suture absent. Tarsi dorsally with only short and thin setae, subproximal arc of setae absent; ventral margin with a row of stout setae; tarsal claws (Fig. 7b) with a single row of 3 subequal teeth, subapical pair of setae absent. Second and third legs similar to foreleg, except setae of the ventral margin less abundant and tibio-patellar suture present. Abdomen. Coloration of the terga uniformly light brown, except terga 5 and 6 brown, darker proximally and laterally. Sterna yellow except 5 and 6 brown. Asymme- trical gills (Fig. 8a) on abdominal segments 1 to 7; dark tracheation weil developed, serrated with thin setae apically and posteriorly (Figs 8b and 8c). Paraproct (Fig. 9) unusually elongated, with about 25 pointed marginal spines, increasing in length at the apex; surface covered with more than 35 scale bases; setae insertion randomly distributed more abundant near the apex; postero-lateral extension with numerous minute spines along the margin; about 10 scale bases close together. Cerci and median caudal filament dark brown. Male and female imagoes unknown. Dabulamanzia gigantea Gattolliat sp. n. Holotype. One female subimago with larval skin, (167a), Madagascar, Manampatrana Bas., Sahanivoraky Riv., Loc. tributary riv. of lantara Riv., Long. 47°00'41" E, Lat. 22°13'33" S, Alt. 1400m, 19.11.1993. J.-M. Elouard, F.-M. Gibon. LARVA Head. Labrum (Fig. 10) sub-rectangular, with a smooth anteromedial emargi- nation; distal margin bordered with two kinds of setae, one row of feathered setae (as in Fig. 1b) and one row of multifid setae (as in Fig. Ic); dorsally with a continuous row of about 15 long setae, abundant setae and insertion of setae in the proximal half; ventral face with a single small seta laterally and a row of thin setae medially. Hypopharynx similar to figure 2. Right mandible (Fig. 11a) with two sets of incisors; prostheca long, thin and unforked, without apical teeth (Fig. 11b); length of the tuft of setae between prostheca and mola reducing toward the mola; tuft of small setae near the mola; tuft of setae at the apex of the mola reduced to 2 or 3 setae; basal half without setae dorsally. Left mandible (Fig. 12a) with incisors fused; prostheca with 4 teeth and the apical much more developed (Fig. 12b); tuft of setae between prostheca and mola present; tuft of setae at the apex of the mola reduced to 3 setae; basal half without setae dorsally. THREE NEW MALAGASY SPECIES OF DABULAMANZIA 565 Ws EN à NYA Fics 7 to 9. Larval structures of Dabulamanzia gladius : Ta : left foreleg. 7b : tarsal claw. 8a : fourth gill. 8b : anterior margin of the fourth gill. 8c : posterior margin of the fourth gill. 9 : paraproct. Maxillae (Fig. 13) with 4 teeth, the distal one distinct from the three others; one row of small setae with, in the middle three stouter setae and apically five setae twice as long as the others; row of 7 setae at the base of the galea; 1 single small seta perpendicularly to the margin of the galea on a well-marked apophysis; palp 2- segmented longer than galealacinia, first segment 1.4 time shorter than the second; thin setae on the inner margin of the first and second segments, especially numerous at the apex of the second; micropores on the inner margin of the first segment. Labium (Fig. 14) with glossae subequal in length to the paraglossae; apical half of glossae with stout setae, patch of five setae on the basal half of the ventral side; paraglossae apically rounded, with 2 rows of simple setae. Thin setae on the lateral side of the mentum. Labial palp 3-segmented; first segment stout, 1.1 smaller than the second and third combined; second segment much larger at the apex than at the base, dorsally with a row of about 4 setae ending with three smaller setae; third segment broad, apex truncated and substraight, ventrally almost completely covered with setae, much larger apico-laterally. Thorax. Hindwing pad present. Forelegs (Fig. 15a) with coxa covered with few spines and an arc of micropores. Femora with a dorsal row of at least 20 long setae, 566 J.-L. GATTOLLIAT & M. SARTORI 11b 12b Fics 10 to 14. Larval structures of Dabulamanzia gigantea : 10 : labrum (left : ventral; right : dorsal). lla : right mandible. 11b : right prostheca. 12a : left mandible. 12b : left prostheca. 13 : left maxilla. 14 : labium. THREE NEW MALAGASY SPECIES OF DABULAMANZIA 567 especially numerous in the proximal half, submarginal setae absent, apical half of dorsal margin with thin short setae; femoral patch of at least 10 spatulated setae; numerous short setae on the ventral and lateral margins; arc of fine setae absent. Tibiae dorsally with only short and thin setae, small subproximal arc of short setae visible on the both sides; apex dorsally with a single long curved setae (Fig. 15b); ventral margin with abundant setae, apex with 3 long and acute setae; tibio-patellar suture absent. Tarsi dorsally with only short and thin setae, subproximal arc of setae absent, apex with a patch of thin setae; ventral margin with a row of short and stout setae; tarsal claws (Fig. 15c) with a single row of 5 teeth, the apical two much smaller, subapical pair of setae absent. Second and third legs similar to foreleg, except setae of the ventral margin less abundant and tibio-patellar suture present. Abdomen. Elongated and asymmetrical gills (Fig. 16) on abdominal segments | to 7, dark tracheation well-developed, serrated with thin setae apically and posteriorly, anterior and posterior margin well-sclerotized. Male and female imagoes unknown. FEMALE SUBIMAGO Forewing length 8.8 mm. Pterostigma with 5 vertical cross-veins. One inter- calary vein between longitudinal veins except apically, two transverse veins between the subcostal and first radial veins (fig. 17). Hindwing length 1.7 mm. Two longitudinal veins well-marked, joined at the base. Two incomplete and less marked veins. Single spur weakly developed (Fig. 18). Dabulamanzia concolorata Gattolliat sp. n. Holotype. Larva female (818a), Madagascar, Antongombato Bas., Makis Riv., Loc. Camp base WWF, Sacred Waterfall, Long. 49°10'09" E, Lat. 12°31'40" S, Alt.1075m, 23.3.1999. J.-L. Gattolliat and Z. Rabeantoandro. Paratypes. Four larvae, same data as holotype. Four larvae, PO810, Madagascar, Antongombato Bas., Makis Riv., Loc. Camp base, 500m downstream of P0818, Long. 49°10'21" E, Lat. 12°31'27" S, Alt.1030m, 21.3.1999. J.-L. Gattolliat and Z. Rabeantoandro. Two larvae, P0814, Madagascar, Antongombato Bas., Makis Riv., Loc. 100m downstream of the Great Waterfall, Long. 49°10'14" E, Lat. 12°29'17" S, Alt.675m, 22.3.1999. J-L Gattolliat and Z. Rabeantoandro. Two larvae, P0822, same locality as P0814, 24.3.1999. J.-L. Gattolliat and Z. Rabeantoandro. LARVA Maximal length : Body 7.2 mm. Cerci 2.5 mm. Terminal filament subequal to the cerci. Head. Coloration almost uniformly light yellow, except brown between the eyes without vermiform marking on vertex and frons. Antennae pale light yellow, except scapus and pedicellus light brown. Eyes and ocelli black. Labrum (Fig. 19) narrow, rounded with a narrow anteromedial emargination; distal margin bordered with simple fine setae; without other setae ventrally; dorsally with an arc of five long setae and a submedial setae, few setae in the proximal half. Hypopharynx as in figure 20, lingua 568 J.-L. GATTOLLIAT & M. SARTORI Fics 15 to 16. Larval structures of Dabulamanzia gigantea : 15a : left foreleg. 15b : apex of the foretibia. 15c : tarsal claw. 16 : fourth gill (damaged apically). covered with numerous setae, superlinguae poorly developed. Right mandible (Fig. 21a) with two sets of incisors, slender and turned backwards; prostheca (Fig. 21b) short and stout, with about seven stout setae; the tuft of setae between prostheca and mola quite short; tuft of small setae near the mola well-developed; tuft of setae at the apex of the mola reduced to 2 stout setae; basal half with setae dorsally. Left mandible (Fig. 22), incisors fused to a group of five teeth; prostheca with 4 teeth, the apical one much more developed and a comb-shaped structure; length of the tuft of setae between prostheca and mola reducing toward the mola; tuft of setae at the apex of the mola reduced to 3 setae; basal half with setae dorsally. Maxillae (Fig. 23) with 4 teeth, the distal one opposed to the three others; 2 rows of setae formed by abundant small setae and long stout setae ending with 3 much longer setae, without pectinated or spine-like setae in the middle of the range; 6 to 7 short setae at the base of the galea arranged in a row; a couple of small setae perpendicularly to the margin of the galea; palp 2- segmented, longer than galealacinia, first segment 1.5 time shorter than the second; few thin setae on the external margin of the first and second segments. Labium (Fig. 24) with glossae subequal to paraglossae; apical half of glossae with stout setae, row of setae subparallel to the inner margin; paraglossae apically rounded, with 2 rows of simple setae. Labial palp 3-segmented; first segment stout, 1.25 smaller than the second and third combined; second segment moderately enlarged at the apex, row of about 6 setae; third segment apically rounded, as broad as the apex of the second. THREE NEW MALAGASY SPECIES OF DABULAMANZIA 569 N MA 1 \ 1 7 - TT TI 4 AL \K Y UVA VIE \\ We 3 Il \V\ Fics 17 to 18. Female subimaginal structures of Dabulamanzia gigantea : 17 : forewing. 18 : hindwing. Thorax. Coloration light brown. Hindwing pad present. Legs yellow, except the dorsal margin of the whole leg and the apex of femora light brown. Forelegs (Fig. 25a), coxa with few setae. Femora with dorsal row of 13 long and broad setae, only 2 or 3 of them in the distal half, no submarginal seta, apical half of dorsal margin with thin setae; femoral patch of 2 spatulated setae; apex crenated with thin setae (Fig. 25b); short setae on the ventral margin. Tibiae dorsally with only thin setae, small subproximal arc of long and thin setae (Fig. 25b); apex dorsally with a single long curved seta (Fig. 25c); ventral margin with few short setae; tibio-patellar suture absent. Tarsi dorsally with only thin setae, subproximal arc of setae absent; ventral margin with a row of short setae; tarsal claws (Fig. 25d) with a single row of 3 short and 2 long teeth, subapical pair of setae absent. Second and third legs similar to foreleg, except setae of the ventral margin less abundant, tibio-patellar-suture present and the subproximal arc of setae longer. Abdomen. Coloration of the terga almost uniformly light brown, except a yellow spot on segments 2 to 9, sometimes surrounded with brown, distal margin darker on segments 4 to 9, segments 3 to 6 with a brown mark laterally (Fig. 26). Sterna yellow except sternite 9 and paraproct light brown. Asymmetrical gills (Fig. 27) on abdominal segments | to 7; dark tracheation well developed, apically and posteriorly serrated with thin setae; anterior and to a less extent posterior margin sclerotized. Paraproct (Fig. 28), with about 16 pointed marginal spines; surface covered with more than 80 scale bases; 570 J.-L. GATTOLLIAT & M. SARTORI Fics 19 to 24. Larval structures of Dabulamanzia concolorata : 19 : labrum (left : ventral; right : dorsal). 20 : hypopharynx. 21a : right mandible. 21b : right prostheca. 22 : left mandible. 23 : left maxilla. 24 : labium. THREE NEW MALAGASY SPECIES OF DABULAMANZIA 57] Fics 25 to 28. Larval structures of Dabulamanzia concolorata : 25a : left foreleg. 25b : apex of the forefemur. 25c : apex of the foretibia. 25d : tarsal claw. 26 : abdomen (dorsal view). 27 : fourth gill. 28 : paraproct. 572 J.-L. GATTOLLIAT & M. SARTORI setae insertion randomly distributed more abundant near the apex; postero-lateral extension with about 35 minute spines along the margin; about 30 scale bases. Cerci and median caudal filament uniformly brown. Male and female imagoes unknown. Dabulamanzia sp. A Material examined. Five male imagoes, two males subimagoes, P0658, Madagascar, Rianila Bas., unnamed Riv., Loc. Road to Lakato, Long. 48°21'48" E, Lat. 19°02'40" S, Alt. 1050m, 26.11.1996. J. Legrand and D. Ramdriamasimanana. One male imago, P0792, same locality P0658, 18.11.1998. J. Legrand and N. Raberiaka. One male imago, P0292, Madagascar, Sakanila Bas., Lakato Riv., Loc. Road to Lakato, Long. 48°25'08" E, Lat. 19°08'33" S, Alt. 690m, 20.3.1995. J.-M. Elouard. Twenty two male imagoes and subimagoes, P0764, same locality as P0292, 17.10.1998. J. Legrand and N. Raberiaka. One male imago, P0771, Madagascar, Sakanila Bas., Lakato Riv., Loc. Road to Lakato, Long. 48°21'38" E, Lat. 19°03'07" S, Alt. 1050m, 26.10.1998. J. Legrand and N. Raberiaka. Six male imagoes, P0794, Madagascar, Rianila Bas., Sahatandra Riv., Loc Road to Lakato. Long. 48°21'43" E, Lat. 19°02'40" S, Alt. 1050m, 19.11.1998. J. Legrand and N. Raberiaka. MALE IMAGO Maximal length : Body 7.3 mm; caudal filaments 18 mm. Head. Yellowish brown without marks (Figs 29 and 30). Turbinate, subcylin- drical eyes deep orange (becoming honey-brown after preservation and storage in alcohol). Stout carina between the antennae. Thorax. Prothorax light brown with dark brown pattern, meso and metathorax uniformly light brown. Forewing length 6.8 mm. Surface hyaline except pterostigma light grey, with 5 to 7 vertical cross-veins not reaching the Sc vein. One intercalary vein between longitudinal veins except between subcostal and first radial veins (fig. 31a). Costal margin serrated (Fig. 31b). Hindwing length 1.1 mm. Surface hyaline. Two well- marked longitudinal veins, joined at the base. Third vein free, incomplete and less marked than two others. Single stout spur covered with small teeth (Fig. 32). Legs yellowish brown, except the apex of tibiae and of each article of the tarsi dark brown. Abdomen pale cream, with a brown narrow transverse line in the distal part of each segment. Genitalia formed by three-segmented gonopods, the limit between the first and the second not visible. Length of articles 1 and 2 0.50 mm. Article 3 0.14 mm. Second segment long, clearly enlarged subapically, with the inner margin covered with small teeth, base of the first segment with a stout apophysis without a brush of setae. Third segment elongated, with the inner margin incurved (Fig. 33). Female imago unknown. Larva unknown. MALE SUBIMAGO Similar to the male imago. THREE NEW MALAGASY SPECIES OF DABULAMANZIA 573 = - 31b Fics 29 to 33. Male imaginal structures of Dabulamanzia sp. A : 29 : head (lateral view). 30 : head (dorsal view). 3la : forewing. 31b : costal margin of the forewing. 32 : hindwing. 33 : genitalia. 574 J.-L. GATTOLLIAT & M. SARTORI AFFINITIES The presence of a subproximal arc of setae on the tibiae of D. gladius sp. n., D. concolorata sp. n. and D. gigantea sp. n. (Figs 7a, 15a and 25a), tibio-patellar suture present only on the second and third legs and serrated costal margin of forewing (Fig. 31b) suggest that these species belong to the Cloeodes complex (Lugo-Ortiz & McCafferty, 1996a; Lugo-Ortiz & McCafferty, 1996b). The presence of teeth on the tarsal claws (Figs 7b, 15c and 25d), the presence of hindwings, the absence of arc of setae on the tarsi clearly demonstrate that these three species do not belong to Cloeodes (Waltz & McCafferty, 1987b; Waltz & McCafferty, 1987a) nor to Maliqua (Lugo- Ortiz & McCafferty, 1997a). The absence of a thumb-like distomedial process on labial palp segment 2 (Figs 6, 14 and 24) and the two subapical not extremely enlarged teeth of the tarsal claw (Figs 7b, 15c and 25d) distinguish these three species from the genus Crassabwa (Lugo-Ortiz & McCafferty, 1996a). At the opposite, the shape of the labial palp (Figs 6, 14 and 24), especially of the second and third segments, the single row of subequal teeth on the tarsal claws (Figs 7b, 15c and 25d), the shape, the venation and the serration of the gills (Figs 8a, 8b, 8c, 16 and 27) are apomorphic features of the genus Dabulamanzia (Lugo-Ortiz & McCafferty, 1996b) . D. sp. A presents the following features which clearly argue that this species belongs to the genus Dabulamanzia: hindwing with a single hooked spur (Fig. 32), forewing with single intercalary veins (Fig. 3la) and serrated costal margin (Fig. 31b) and male genital forceps with a well-developed apophysis on the first segment (Gies33): Larvae of D. gladius and D. gigantea are quite close. The great size of D. gigantea, the shape of the apex of the third segment of the labial palp (Figs 6 and 14), the shape of the labrum (Figs la and 10), the shape of the legs (Figs 7a and 15a) and the number of tarsal teeth (Figs 7b and 15c) allow us to separate these two species. They are distinguished from African and all the other Malagasy larvae of Dabula- manzia by the shape of the labrum, the shape of the third segment of the labial palp, the right prostheca and the fused teeth of the mandibles (Wuillot & Gillies, 1993; Lugo- Ortiz & McCafferty, 1996b; Lugo-Ortiz & McCafferty, 1997c; Gattolliat er al., 1999). They appear much closer to D. tarsale (Gillies): the shape of the labrum and of the third segment of the labial palp are very similar. However, the right prostheca is different, bifid in D. tarsale (Gillies, 1990) and reduce to a single bristle-like in D. gladius and D. gigantea (Figs 3 and 11b). Moreover the incisors of the mandibles of D. gladius and D. gigantea are very characteristic. D. concolorata differs clearly from all other species of Dabulamanzia by the shape of the labrum (Fig. 19). It is closely related to D. duci Gattolliat & Elouard and D. improvida Lugo-Ortiz & McCafferty (Lugo-Ortiz & McCafferty, 1997c; Gattolliat et al., 1999). However, it differs from them in the coloration of the abdomen (Fig. 26), the shape of third segment of the labial palp (Fig. 24), the right mandible with incisors slender and turned backwards (Figs 21a and 22) and the maxillae with an unusual couple of small setae perpendicularly to the margin of the galea (instead of a single seta in most of the other species and genera of Baetidae) (Fig. 23). THREE NEW MALAGASY SPECIES OF DABULAMANZIA 57 Nn D. sp. A differs from the African species by the shape of the apophysis of the first segment of the gonopods (Fig. 33). It appears more closely related to the Malagasy species D. duci. The following features allow us to distinguish the two species: second segment of the gonopod clearly enlarged subapically and apophysis of the first segment without a brush of setae, inner margin of the third segment incurved (Fig. 31). As this species 1s only known at the imaginal and subimaginal stages, we refrain to name it, even if the above-mentioned features distinguish it from all the other Malagasy species. The great size of D. sp. A clearly indicates that it could not be the imago of D. gladius, even if theses two species are present in the same area. DISCUSSION The genus Dabulamanzia is actually known by 11 species, 6 in mainland Africa and 5 in Madagascar. However, we cannot conclude that this genus presents the same diversity in both areas. The number of species is greatly underestimated in Africa for two main reasons. First, the baetid fauna of most African regions is poorly known (McCafferty & de Moor, 1995) and secondly, even if Dabulamanzia 1s widespread and very common, it is seldom caught by light-traps (Gillies, 1990). This could be due to the peculiar mating behaviour of the males: they wait in the morning for the females in an horizontal flight a few centimetres above flat stones or rocks (Gattolliat er al., 1999): The status of the genus Dabulamanzia is not clear in Madagascar. Two of the five species, D. improvida and D. duci are extremely close. Moreover, a third species, Nesydemius polhemusorum Lugo-Ortiz & McCafferty, appears to be also very close to them. These authors claim that the presence of thin setae on the dorsal margin of the legs and the relative size of the subapical teeth of the tarsal claws are apomorphies, distinguishing the monospecific genus Nesydemius from others species of Dabula- manzia (Lugo-Ortiz & McCafferty, 1998). A single specimen of Nesydemius polhemu- sorum was mounted; consequently the intraspecific variation among and between populations cannot be correctly estimated. The tiny differences observed between single specimens of N. polhemusorum and D. improvida need to be strengthened and the validity of the genus Nesydemius confirmed. The mouthparts of D. gladius and D. gigantea show clear adaptation for scraping the tops of stones. They share this kind of specialisation with the genus Xyrodromeus Lugo-Ortiz and McCafferty that also shows the same adaptations: teeth of the mandibles fused and extremely developed, labrum almost straight (Lugo-Ortiz & McCafferty, 1997b). Similar adaptations are also present in other genera such as Baetis (Müller-Liebenau, 1969). However, they must be strictly considered as a convergence due to the same foraging behaviour. The shape of the labial palp, the maxillae, the subproximal arc of setae on the tibiae and especially the single row of teeth on the tarsal claw clearly distinguish these three genera and demonstrate that they do not belong to the same complex of genera. Since the remaining species lack apomorphic features and D. gladius and D. gigantea share with the other Dabulamanzia species a number of apomorphies, it is not justified to erect a new genus for these two species. 576 J.-L. GATTOLLIAT & M. SARTORI KEY TO THE MALAGASY SPECIES OF DABULAMANZIA IL Distal margin of the labrum almost straight; mandibles with fused incisors; third segment of the labial palp apically truncated................ 2 - Distal margin of the labrum with a clear anteromedial emargination, mandibles with incisors not fused; third segment of the labial palp apicallysroundediorpomtedì iv... IE 8 2. Third segment of the labial palp apically concave; labrum almost quadrangular; femora with a dorsal row of about 12 setae.......... D. gladius - Third segment of the labial palp apically straight; labrum more rounded; femora with a dorsal row of at least 20-setae............4.- .-0% D. gigantea 3% Tergae almost uniform light brown; labrum narrow; third segment of the labialpalpnot@breader thanithe second... un 02.9 D. concolorata - At least tergae 3 to 6 brown with 3 lighter median spots, labrum rounded not narrow, third segment of the labial palp broader than the second......... 4 4 Third segment of the labial palp rounded, second segment of the maxillary palp slender and pointed= 2%... A O 2.0 ve ee D. improvida - Third segment of the labial palp slighty pointed, second segment of the maxillaryspalprasıbroad as the second. VERRE aa yee cee D. duci ACKNOWLEDGEMENTS We thank the whole team of the LRSAE (Laboratoire de Recherche sur les Systemes Aquatiques et leur Environnement, Antananrivo, Madagascar), especially its director Dr J.-M. Elouard for logistical assistance and donation of specimens, and Sophie Rabeantoandro for great help during our field work in Madagascar. We also want to thank Dr M.T. Gillies, Lewes England, for useful comments on this paper. A travel grant was provided to JLG by the Swiss Academy of Sciences. This paper is the contribution n° 39 to the series "Aquatic Biodiversity of Madagascar". REFERENCES GATTOLLIAT, J.-L., SARTORI, M. & ELOUARD, J.-M. 1999. Aquatic Biodiversity from Madagascar 12: three new species of Baetidae (Ephemeroptera) from the Réserve Naturelle Intégrale d'Andohahela. Fieldiana ns 94: 115-124. GiLLIES, M. T. 1990. A revision of the African species of Centroptilum Eaton (Baetidae, Ephemeroptera). Aquatic Insects 12(2): 97-128. LUGO-ORTIZ, C. R. & MCCAFFERTY, W. P. 1996a. Crassabwa: a new genus of small minnow mayflies (Ephemeroptera: Baetidae) from Africa. Annales de Limnologie 32(4): 235-240. LUGO-ORTIZ, C. R. & MCCAFFERTY, W. P. 1996b. The composition of Dabulamanzia, a new genus of Afrotropical Baetidae (Ephemeroptera), with description of two new species. Bulletin de la Société d'Histoire Naturelle de Toulouse 132: 7-13. LUGO-ORTIZ, C. R. & MCCAFFERTY, W. P. 1997a. Maliqua: a new genus of Baetidae (Ephemer- optera) for a species previously assigned to Afroptilum. Entomological News 108: 367-371. LUGO-ORTIZ, C. R. & MCCAFFERTY, W. P. 1997b. New Afrotropical genus of Baetidae (Insecta: Ephemeroptera) with bladelike mandibles. Bulletin de la Société d'Histoire Naturelle de Toulouse 133: 41-46. THREE NEW MALAGASY SPECIES OF DABULAMANZIA Sia LuGo-ORTIZ, C. R. & MCCAFFERTY, W. P. 1997c. New species and first reports of the genera Cheleocloeon, Dabulamanzia and Mutelocloeon (Insecta : Ephemeroptera) from Mada- gascar. Bulletin de la Société d'Histoire Naturelle de Toulouse 133: 47-53. LuGo-ORTIZ, C. R. & MCCAFFERTY, W. P. 1998. Phylogeny and biogeography of Nesydemius, n. gen., and related Afrotropical genera (Insecta : Ephemeroptera : Baetidae). Bulletin de la Société d'Histoire Naturelle de Toulouse 134: 7-12. MCCAFFERTY, W. P. & DE Moor, F. C. 1995. South African Ephemeroptera : Problem and priorities. (pp. 463-476). In: Ciborowski C. L.D & J. H. H. (eds.). Current directions in research in Ephemeroptera. Canadian Scholars' Press, Toronto. MULLER-LIEBENAU, I. 1969. Revision der Europäischen Arten der Gattung Baetis Leach, 1815. Gewässer und Abwässer 53/54: 1-214. WALTZ, R. D. & MCCAFFERTY, W. P. 1987a. Generic revision of Cloeodes and description of two new genera (Ephemeroptera: Baetidae). Proceedings of the Entomological Society of Washington 89(1): 177-184. WALTZ, R. D. & MCCAFFERTY, W. P. 1987b. Revision of the genus Cloeodes Traver (Ephemer- optera: Baetidae). Annals of the Entomological Society of America 80(2): 191-207. WUILLOT, J. & GILLIES, M. T. 1993. New species of Afroptilum (Baetidae, Ephemeroptera) from West Africa. Revue d'Hydrobiologie Tropicale 26: 269-277. i Mi toner Fara ER REVUE SUISSE DE ZOOLOGIE 107 (3): 579-589; septembre 2000 Das Mannchen von Zelotes zellensis Grimm (Araneae: Gnaphosidae) Christoph MUSTER & Konrad THALER Institut fiir Zoologie und Limnologie der Universitit, Technikerstraße 25, A-6020 Innsbruck, Austria. The male of Zelotes zellensis Grimm (Araneae: Gnaphosidae). — The previously unknown male of Zelotes zellensis Grimm, 1982 is described from the Bavarian Alps (Germany). Characters to distinguish this species from the closely related Z. aeneus (Simon, 1878) are indicated and illus- trated. For both species notes on habitat, maturity period and distribution are given. Key-words: Araneae - Gnaphosidae - Zelotes - Alps. EINLEITUNG Die Gnaphosidae Mitteleuropas bergen, obwohl von Grimm (1985) gründlich revidiert, noch ungelöste taxonomische Probleme, besonders ım alpinen Raum. Dort finden sich die vier Arten, deren Männchen Grimm unbekannt blieben: Berlandina nubivaga (Sımon, 1878) und Gnaphosa alpica Simon, 1878 in den Westalpen, sowie die erst rezent von Grimm (1982) beschriebenen Zelotes devotus und Z. zellensis. Die Männchen von G. alpica und von Z. devotus wurden inzwischen identifiziert (Ovtsha- renko et al., 1992; Thaler, 1989a). Von Z. zellensis waren bisher nur das Holotypus- Weibchen und ein subadultes Exemplar aus den Kitzbühler Alpen (Salzburg) bekannt. Einzelne Männchen der Art aus den Bayerischen Alpen haben bereits vor Jahren J. Wunderlich vorgelegen (in litt.). Sie wurden als Z. aeneus (Simon, 1878) nahestehend erkannt, eine nähere taxonomische Zuordnung war damals jedoch nicht möglich. Das Auffinden einer größeren Serie von Exemplaren beider Geschlechter am Geigelstein (Chiemgauer Alpen) im Jahr 1999 erlaubt es nun, das Männchen von Z. zellensis zu beschreiben sowie Verbreitung und Habitat dieser Art zu schildern. Abkürzungen: CM = Goll. € Muster, ETh’= Coll IR Thaler MHNG = Muséum d'histoire naturelle Geneve, NMW = Naturhistorisches Museum Wien, ZMH = Zoologisches Museum Hamburg. Mafiangaben sind in mm. Zelotes zellensis Grimm, 1982 Z. zellensis Grimm, 1982: 177-178, Beschreibung des 2; Grimm (1985: 262, 2); Platnick (1989: 496, 1993: 671); Heimer & Nentwig (1991: 450, 2). Manuskript angenommen 25.04.2000 580 CHRISTOPH MUSTER & KONRAD THALER ABB. 1-4: Zelotes zellensis Grimm (1, 2), Z. aeneus (Simon) (3, 4). & -Taster von ventral (1, 3) und retrolateral (2, 4). Abkiirzungen: E Embolus, K Konduktor, Na Nebenarm des End- apparates, TA Terminalapophyse des Tegulum. Maßstab 0,1 mm. MANNCHEN VON ZELOTES ZELLENSIS 58] ABB. 5-9: Zelotes zellensis Grimm (5, 7, 8, 9), Z. aeneus (Simon) (6). d -Taster, Endapparat von ventral (5, 6), Embolus von dorsal (7), Epigyne von ventral (8), Vulva von dorsal (9). Maßstab 0,1 mm. Material: DEUTSCHLAND: BAYERN, Karwendel, Soiernspitze, 1780 m, 14 (10.08.- 10.09.1998); Chiemgauer Alpen, Geigelstein, 1650 — 1800 m, 12 (25.09.1998-18.05.1999), 23, 12 (30.06.-29.7.1999), 23 (29.07.-21.08.1999), 18, 49 (21.08.-11.09.1999), 23, 69 (11.09.-17.10.1999); OSTERREICH: SALZBURG, Tennengebirge, Samer Alm, 1520 m, Id (07.07.-04.08.1999); Kitzbühler Alpen, Schmittenhöhe bei Zell am See, 1400-1800 m, 1° Holotypus (ZMH, 05.07.1979, leg. Grimm). Mit Ausnahme des Holotypus wurden alle Ex. in Barberfallen gefangen, leg. Muster. Deponierung: Ex. vom Geigelstein 18, 22 MHNG, 16, 19 NMW, 18, 12 ZMH, 18,1% CTh, übrige Ex. CM. Männchen: Maße (n = 8): Gesamtlänge 4,5 (4,0 — 5,0), Prosomalänge 1,85 (1,7 — 1,9), Prosomabreite 1,4 (1,3 — 1,5), Sternumlänge 1,1 (1,0 — 1,2), Cymbiumlänge 0,65 (0,62 — 0,67), Bein I: Fe 1,33 (1,23 — 1,37), Pt 0,74 (0,67 — 0,80), Ti 1,06 (0,98 — 1,25), Mt 0,80 (0,75 — 0,87), Ta 0,78 (0,72 — 0,80). Cheliceren: vorderer Klauenfurchenrand mit 3-4, hinterer mit 1-2 Zähnchen. Beinbewehrung: Mt I und II mit 2 Paar ventraler Stacheln, je eines im basalen und im mittleren Drittel des Beingliedes; gelegentlich nur 2 oder 3 Stacheln vorhanden. Opisthosoma: mit großem, dunkelbraunem Scutum, ca. 40% der Abdomenlänge ein- nehmend. 582 CHRISTOPH MUSTER & KONRAD THALER ABB. 10-15: Zelotes zellensis Grimm (10, 12, 14), Z. aeneus (Simon) (11, 13, 15). d-Taster von schräg ventral (10, 11), Endapparat von schräg ventral (12, 13), Spitze des Nebenarmes (14, 15). MANNCHEN VON ZELOTES ZELLENSIS 583 Pedipalpus: Abb. 1, 2 , 10. Tibia-Apophyse gerade, Tegulum und Retinaculum ohne Besonderheiten. Endapparat: Abb. 5, prolateral-distal mit Höcker (X), retro- lateral-proximal gerundet (Y); Nebenarm (Na in Abb. 1) lang dornförmig, schwach gebogen; Konduktor (K in Abb. 2) groß, polsterförmig, von ventral deutlich sichtbar und Embolus teilweise überdeckend; Embolus (E in Abb. 2) lang, S-förmig, prolateral mit ganzrandigem, lamellösem Saum; Spermophor vor Eintritt in den Embolus blasenartig erweitert. Nebenarm des Endapparates wie bei einigen anderen Arten der Gattung (Miller, 1967: 256) von einem feinen Kanal durchzogen, der an der Erweiterung des Spermophor vorbei in den Embolus führt und dort nicht weiter zu verfolgen war. Der Nebenarm erscheint demnach als ein ”2. Embolus” (Grimm, 1985: 188). Die biolo- gische Bedeutung dieses ”’rätselhaften Organs” (Miller, 1967: 256) ist nach wie vor unbekannt. Nach Grimm (1985: 18/19) weist der Nebenarm im distalen Bereich eine Skleritfalte auf; Abb. 14, 15 zeigen die komplizierten Mündungsstrukturen. Weibchen: Mit den Beschreibungen von Grimm (1982, 1985) gut überein- stimmend, Maße (n = 11): Gesamtlänge 5,1 (4,5 — 5,6), Prosomalänge 1,9 (1,8 — 2,1), Prosomabreite 1,5 (1,3 — 1,6), Sternumlänge 1,2 (1,0 — 1,2), Bein I: Fe 1,24 (0,95 — 1,40), Pt 0,75 (0,63 — 0,80), Ti 0,98 (0,92 - 1,05), Mt 0,80 (0,75 — 0,87), Ta 0,75 (0,72 — 0,78); Epigyne und Vulva Abb. 8, 9. In Tab. 1 wird das Mittelfeld der Epigyne in Anlehnung an Chyzer & Kulczynski (1897: 195, Fußnote 1), Schenkel (1925) und Lohmander (1944) als "Areola” bezeichnet. VERWANDSCHAFTSBEZIEHUNGEN Die Zugehörigkeit von Z. zellensis zur Z. subterraneus-Gruppe (Grimm, 1982) wird durch die Merkmale des d-Tasters bestätigt: Embolus relativ kurz, distal retrolateral entspringend, Endapparat mit Nebenarm (embolar projection) (Platnick & Shabad, 1983; Grimm, 1985). Größte Ähnlichkeit besteht zu Z. aeneus (Abb. 1, 2, 10 vs. 3, 4, 11), insbesondere da bei beiden Arten der Nebenarm den Rand des Cymbiums erreicht. Unterschiede zeigen sich im Detail (Abb. 5, 12 vs. 6, 13): bei Z. aeneus ist der Embolus kürzer, nicht so stark gewunden, der Innensaum mit unregel- mäßiger Kontur, nach Job (1969) “in eine feine...Spitze ausgezogen”, der Endapparat erscheint retrolateral-proximal beilförmig (Y in Abb. 6). Der Spermophor von Z. aeneus ist im Endapparat vor der Einmündung in den Embolus nicht blasenartig erweitert, der Konduktor ist kleiner, das Tegulum distal-medial zu einer Spitze ausgezogen ("terminal apophysis”, Platnick & Shabad, 1983, TA in Abb. 3). Zudem unterscheiden sich beide Arten deutlich in ihren Dimensionen, besonders in der Länge des Cymbiums (Abb. 16, 17). Zelotes aeneus ist in beiden Geschlechtern die größere Art, die Variabilität ist im Gegensatz zu Z. zellensis beträchtlich. Die wesentlichen Unterschiede zwischen den beiden Arten sind in Tab. | zusammengefaßt. 584 CHRISTOPH MUSTER & KONRAD THALER + Z. aeneus (Innsbruck- Martinswand) m Z. aeneus (Otztal- Forchet) A Z. aeneus (Südtirol) | A Z. zellensis (Chiemgauer Alpen - Geigelstein) Cymbiumlange in mm D Z. zellensis (Karwendel - Soiernspitze) 1,50 2,00 2,50 3,00 3,50 Prosomalänge in mm ABB. 16: Relation zwischen Cymbium- und Prosomalänge bei adulten Männchen von Zelotes zellensis und Z. aeneus. + Z. aeneus (Innsbruck - Martinswand) | IZ. aeneus (Otztal-Forchet) | AZ. aeneus | (Südtirol) Länge Tibia | in mm | AZ. zellensis (Chiemgauer Alpen - Geigelstein) 1,3 2,0 2,9 3,0 Prosomalänge in mm ABB. 17: Relation zwischen Länge der Tibia I und Prosomalänge bei adulten Weibchen von Zelotes zellensis und Z. aeneus. MANNCHEN VON ZELOTES ZELLENSIS 585 Tas. 1: Differenzierung von Zelotes aeneus und Z. zellensis Merkmal 3d: Cymbiumlänge 3: Embolus 3d: Tegulum 3: Opisthosoma 2: Länge Tibia I 9: Epigyne 3d, 9: Bestachelung des Metatarsus I Lebensraum Gesamtverbreitung aeneus 0,78 — 1,1 mm kürzer, Innensaum unregelmäßig (Abb. 6, 13) distal-medial mit Spitze (TA, Abb. 3, 4) Scutum klein (< 1/3 Abdomen-Länge) 1,1 - 1,5 mm Areola fast quadratisch, Vorderecken der Epigyne weit vor der Areola (Grimm, 1985: Abb. 310) ventral 1 Paar Fels- und Steppenheide unter 1300 m Westeuropa, dispers in zellensis 0,62 — 0,67 mm länger, stärker gewunden, Innensaum ganzrandig (Abb. 5, 7,12) distal-medial gerundet (Abb. 1, 2) Scutum größer (> 1/3 Abdomen-Länge) 0,9- 1,1 mm Areola herzförmig, Vorderecken der Areola genähert (Abb. 8) ventral meist 2 Paare Geröllfelder, Felsrasen und Zwergstrauchheide der Subalpinstufe in 1400 — 1800 m mittlere Nordalpen (Abb. 18) Mitteleuropa (Grimm, 1985: 298) EINNISCHUNG UND VERBREITUNG Die beiden Vergleichsarten sind geographisch und vor allem ökologisch von- einander isoliert. Zelotes zellensis ist anscheinend auf die subalpine Höhenstufe der Ostalpen beschränkt, Z. aeneus in Mitteleuropa dispers an Xerothermstandorten in Höhenlagen unter 1300 m verbreitet. Alle Individuen von Z. zellensis wurden an südexponierten Hängen im Bereich der Krummholzzone zwischen 1400 und 1800 m gefangen. Vorzugslebensraum sind Kalkschutthalden im Mosaik mit Legföhren und Zwergstrauchbeständen. Die höchsten Fangzahlen am Geigelstein wurden in Barberfallen erzielt, die direkt im vegetationsfreien Blockschutt installiert waren. Das Weibchen aus dem Tennengebirge stammt von einem ausgedehnten Borstgrasrasen über Werfener Schiefer. Die Hauptaktivitätszeit von Z. zellensis liegt wie bei Z. aeneus (siehe Stein- berger, 1990) in den Herbstmonaten. Adulte Individuen wurden von Juli bis Oktober gefangen, mit einem Aktivitätsmaximum im September. Bei diesen Arten fehlt somit eine Aktivitätsspitze im Frühjahr, wie sie andere in subalpinen Lebensräumen am Nordabfall der mittleren Ostalpen vorkommende Zelotes-Arten nach der Überwin- terung aufweisen: Z. apricorum (L. Koch, 1876), Z. clivicola (L. Koch, 1870), Z. petrensis (C. L. Koch, 1839), Z. similis (Kulczynski, 1887). 586 CHRISTOPH MUSTER & KONRAD THALER * Zelotes zellensis | | % Zelotes aeneus E22] > 1000 m über NN ABB. 18: Verbreitung von Zelotes zellensis (Gesamtareal) und Z. aeneus in den mittleren Ostalpen. Fundorte von Z. aeneus nach Noflatscher (1991), Thaler (1995, 1997). Fragezeichen markieren die vermutlichen Fundorte von Z. zellensis im Wimbachgries (Korge, 1977) und am Attersee (Pfeiffer, 1901). Nach aktuellem Wissensstand ist Z. zellensis ein Endemit der Ostalpen. Die Art wurde bisher nur nördlich des Alpenhauptkammes gefunden, an wenigen Stand- orten des Alpennordrandes zwischen Isar und Tennengebirge (Abb. 18). Wir erwarten, dass sie in diesem Gebiet in ihrem Vorzugshabitat regelmäßig anzutreffen sein wird. Auch die Funde von "Zelotes aeneus” im Nationalpark Berchtesgaden (Korge, 1977) und in den Bayerischen Alpen (ohne nähere Angaben, Platen in Blick & Scheidler, 1991) dürften mit hoher Wahrscheinlichkeit auf Z. zellensis zu beziehen sein. Korge sammelte seine Exemplare ja an ”Schotterflächen mit primären Schutt- besiedlern...bei noch recht geringem Bedeckungsgrad” in 1400 m Höhe. Unklar bleibt die Arealgrenze im Süden und Osten. Der Locus typicus am Ostrand der Kitzbühler Alpen ist der südlichste Fundort. In den Zentralalpen scheint die Art weitgehend zu fehlen, wie Fallenfänge in den Untersuchungsräumen Obergurgl (Puntscher, 1980) und Hohe Tauern (Thaler, 1989b; Relys, 1996) nahelegen. Auch aus den Nord- ostalpen ist den Verfassern keine unmittelbar auf Z. zellensis zu beziehende Angabe bekannt (siehe Wiehle & Franz 1954). Vielleicht hat aber schon Kulczynski (1915: 916) ein Männchen von Z. zellensis gesehen. Es handelt sich um das bereits von Pfeiffer (1901, sub Prosthesima setifera) genannte Exemplar von ”Z. aeneus” aus Oberösterreich. Der Fundort Weyeregg am Attersee liegt in der Nähe des Tennen- gebirges und weit vom nächsten aeneus-Vorkommen entfernt. Dieser Verdacht wird durch die folgende Überlegung gestützt: dieses d soll sich nämlich in Details von einem von Kulczynski (1887, Abb. 63, sub P. sarda) aus Südtirol abgebildeten Exemplar unterscheiden. Diese Abbildung dürfte aber den “echten” Z. aeneus dar- stellen, zumindest ist die mediale Spitze des Tegulum deutlich erkennbar. An MANNCHEN VON ZELOTES ZELLENSIS 587 Wärmestandorten Südtirols wurde Z. aeneus im Sinne der rezenten Autoren in- zwischen konstant und abundant nachgewiesen (Noflatscher, 1988, 1991). Möglicherweise kommt Z. zellensis auch in den Karpaten vor. Miller & Svaton (1978) bilden mit gewissem Vorbehalt ein ”Z. aeneus” - 8 aus der Slowakei ab, das viele Merkmale von Z. zellensis aufweist: Innensaum des Embolus ganzrandig, Konduktor groß, Vorderrand des Tegulum ohne Spitze. Leider ist das Exemplar in der Sammlung Miller nicht mehr vorhanden (Kürka, 1994). Dagegen ist Z. aeneus eher als atlantisches Faunenelement anzusprechen, mit deutlichem Verbreitungsschwerpunkt in Westeuropa (Grimm, 1985: 298) und dispersen Vorkommen in Mitteleuropa. Die Fundorte in Nordtirol (Thaler, 1997) lassen eine Besiedlung von Südtirol aus über den Reschenpaß vermuten. Periphere Vorkommen bestehen im Raum Innsbruck und bei Seefeld (Murphy & Murphy, 1984). Weiter östlich, wie in der Steiermark (Kropf & Horak, 1996) und weiten Teilen Kärntens (Komposch & Steinberger, 1999), scheint die Art dagegen groß- räumig zu fehlen. Die Nennung vom Magdalensberg in Kärnten ist auf Z. larreillei (Simon, 1878) zu beziehen (Steinberger, 1988 und in litt.). Das kürzlich festgestellte Vorkommen am Dobratsch bei Villach (Komposch, 1997) könnte mit einem entlang des Alpensüdrandes zu vermutenden Teilareal in Verbindung stehen. Auch der Fundort bei Wiener Neustadt (Malicky, 1972) sollte aus dem Vorland erreicht worden sein. DANK Wir danken Herrn Dr. H. Dastych (Hamburg) für die Ausleihe des Typus- materials von Z. zellensis, Herrn Dr. K. H. Steinberger (Innsbruck) für Vergleichs- material von Z. aeneus und den Herrn K. Eller und K. Schatz (Innsbruck) für Hilfe bei den Rasteraufnahmen. Die Arbeit ist Teil des Dissertationsvorhabens von C. M. im FB Biologie der Universität Hamburg. Der Erstautor dankt für finanzielle Unter- stützung der Graduiertenförderung Hamburg, dem DAAD und dem Evangelischen Studienwerk Haus Villigst e.V. LITERATUR BLICK, T. & SCHEIDLER, M. 1991. Kommentierte Artenliste der Spinnen Bayerns (Araneae). Arachnologische Mitteilungen 1: 27-80. CHYZER, C. & KULCZYNSKI, L. 1897. Araneae Hungariae. 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Wagner, Innsbruck, 664 pp. Sr hae En - di GS REVUE SUISSE DE ZOOLOGIE 107 (3): 591-599; septembre 2000 Note sur les musaraignes (Soricidae, Mammalia) du Parc National du Souss-Massa, Maroc Peter VOGEL!, Mark LAWRENCE? & Ali AGHNAJ? | Institut d'écologie, Université de Lausanne, CH-1015 Lausanne, Suisse. 2 International Foundation for Conservation and Developement of Wildlife, B.P. 116, Inezgane, Maroc (current address: National Avian Research Center, P.O. Box 45553, Abu Dhabi, UAE). 3 Direction du Parc National Souss-Massa, B.P. 107, Inezgane, Maroc. Shrews (Soricidae, Mammalia) of the National Park Souss-Massa, Morocco. - With 420 trap-nights we sampled shrews in different habitats of the Souss-Massa National Park. Crocidura viaria (n=11) was found in 5 habitats (cultivated land, wetland, beach dunes with Chenopodia, sand dunes with patchy vegetation and also in the Euphorbia association). No shrews were captured in Eucalyptus forest with undergrowth. Southward from Essaouira, C. viaria is obviously the most frequent species in dense or patchy lowland vegetation. Two other species (C. whitakeri and C. tarfayaensis) which occur in sympatry in the Agadir region were not captured. They live probably not in syntopy with C. viaria and their ecological requirements remain to be studied. Key-words: Soricidae - Crocidura viaria - shrews - Souss-Massa National Park. INTRODUCTION Le Parc National du Souss-Massa (PNSM), fondé en 1991, a pour but principal de protéger la plus grande population d’Ibis chauve (Geronticus eremita). Elle compte entre 250 et 300 individus, représentant plus des deux tiers de la population mondiale (Brindley er al., 1995). Au vu d’une espece aussi menacée, la protection de cette région se justifie largement. Compte tenu de la richesse ornithologique rencontrée aux embou- chures des rivieres Souss et Massa, de la diversité des milieux rencontrés dans ce parc (Anonyme, rapport des Eaux et Foréts) et la diversité faunistique due a la rencontre d’elements paléarctiques et afrotropicaux (Benhamza, 1995), cette région mérite d’être mieux connue sur le plan scientifique. Concernant les mammiferes, Aulagnier & Thévenot (1986a) ont publie un aperçu basé sur une collection étendue de données diverses. Une étude plus ciblée sur les musaraignes de la région du Oued Massa par Aulagnier & Hermas (1989) a été basée essentiellement sur l’analyse de pelotes de déjection de rapaces nocturnes, Manuscrit accepte le 08.05.2000 592 P. VOGEL, M. LAWRENCE & A. AGHNAJ mettant en évidence la présence de trois espèces a fréquence pratiquement égale (Crocidura viaria, C. whitakeri et C. tarfayaensis) et d’une espèce très rare (Crocidura lusitania). Plus récemment, Benhamza (1995) a repris le sujet de la faune mammalienne du PNSM, réunissant une vaste base de données par observations directes, analyse de traces, de feces, de parties squelettiques et en ce qui concerne les mircromammiferes, par piégeages effectués sur une période de deux ans. Si ce travail a permis de mieux cerner les exigences écologiques de certaines especes (Carnivores, Ongulés, Rongeurs), le résultat concernant les Soricidae est étonnant: sur 58 micromammiferes capturés par trappes, aucune musaraigne n'y a été prise. Ces données divergentes obtenues par des méthodes peu comparables nous ont incité a rechercher une explication. En 1986, l’un de nous a eu l’occasion de faire un essai de piegeage dans un champ cultivé de Sidi R’bat ayant comme résultat la capture de 5 musaraignes (Vogel et al., 1988). Ceci a donné l’idée d’appliquer la même technique pour vérifier |’ hypo- these d’une régression Eventuelle, évaluer la situation actuelle de ces insectivores et comparer les peuplements des musaraignes des différents milieux du parc. MATERIEL ET METHODES Les milieux typiques du parc sont, selon Benhamza (1995), au nombre de 7, soit les dunes fixées (I), dunes non fixées (II), milieu steppique (III), reboisement a Euca- lyptus (IV), formation a Euphorbes cactoides (V), cultures (VI), zones humides a végétation spécialisées (VII). Tous ces milieux ont été prospectés, sauf le milieu steppique, la végétations semblait trop faible pour convenir aux musaraignes. En ce qui concerne le milieu cultivé, nous nous référons à Vogel et al. (1988). Apres l’inspection d’un milieu, une ou deux lignes de 30 trappes Longworth® (Penlon, Abingdon, GB) ont été posées, explorant ainsi des sous-unites du même biotope. L’espacement moyen des trappes était d’environ 7 m (3 à 15 m), l’empla- cement ayant été si possible choisi sous un strate de végétation dense pour maximiser la capture des musaraignes. Les piégeages, totalisant 420 pieges-nuits, ont eu lieu en juillet/aoüt 1998. Les trappes, dont la boite de nid est garnie de foin pour absorber l’urine, ont été appatées de viande ou de sardines, de flocons d’avoine, de graines de tournesol et d’un raisin. Les individus capturés ont été pesés et mesurés sans anesthésie et relachés sur place. Une analyse plus poussée était prévue en cas de probleme de détermination. Celle-ci s’est basée sur les travaux d’Aulagner & Thévenot (1986b), Benhamza (1995) et l’expérience personnelle (Vogel & Maddalena, 1987; Vogel et al., 1988). RESULTATS Au total, 19 micromammiferes ont été capturés, soit 11 musaraignes et 8 ron- geurs. Ceci suggére que dans les milieux prospectés, la faune des micromammiferes semble étre dominée par les musaraignes et non pas par les rongeurs. Toutes les musaraignes appartenaient à la même espèce Crociura viaria (Fig. 1). MUSARAIGNES DU PARC NATIONAL DU SOUSS-MASSA 593 TABLEAU | Répartition des 19 micromammiferes capturés dans le Parc National Souss- Massa en fonction du milieu, avec indication du nombre de trappes-nuits (T-N). * = captures de 1986. Habitat T-N Crocidura Mus Mastomys Gerbillus Gerbillus viaria spretus erythroleucus campestris hoogstraali Dunes a végétation 120 | I (D Dunes à Chenop. 120 2 5 (ID) Steppe È (ID) Forét a Eucalyptus 30 (IV) Euphorbes cactoides 60 2 (V) Champs cultivés* 60 5 2 (VD Zone humide (VII) 90 6 l | Les captures, regroupées en fonction du milieu, sont résumées au Tableau 1. Il s’en suit que le milieu le plus riche en musaraignes (6 individus sur 90 trappes-nuits) correspond a la végétation du lit du ruisseau (Fig. 2), dominé par les plantes halophiles (Salicornia sp.), des joncs (Juncus acutus) et des Tamarisques (Tamarix gallica). Dans les autres milieux, le taux de captures était plus faible avec environ 2 musaraignes sur 60 trappes. En ce qui concerne les dunes, tout dépend de la couverture du sol: dans les dunes les plus proches de la mer et isolées, avec couverture de Chenopodium murale, aucune musaraigne n'a été capturée. C’est en fait l'habitat typique de Gerbillus hoogstraali. Ceci n’est pas seulement démontré par les captures assez nombreuses de cette espèce, mais encore par les quantités importantes de traces en forme de cercle autour de la dune et rayonnant entre les dunes. Néanmoins, dans une parcelle de Chenopodium tres étendue d’environ 30 m de long en bordure des dunes fixes (Fig. 3), un couple de C. viaria a été capturé. Dans les dunes partiellement boisées (Acacia cyanophylla), cette musaraigne était également présente. Mais dans ce type de milieu, elle n’a jamais été prise dans les reboisements a Eucalyptus. Dans des dunes stabilisées et une forêt d’Eucalyptus avec sous-bois longeant l’embouchure de l’Oued Souss, aucun micromammifere n’a été pris. Finalement, les lignes posées dans un milieu a Euphorbes cactoides sur un sol rocheux au-dessus de Sidi R’bat (Fig. 4) ont mis en évidence la présence de C. viaria (2 individus sur 60 trappes). Rappelons que les cultures ont permis en juillet 1986 un taux de captures élevé. 594 P. VOGEL, M. LAWRENCE & A. AGHNAJ FIG. 1 Crocidura viaria, mäle adulte. La queue de cette espece est visiblement un lieu de stockage de graisse. (=) Milieu humide, occasionnellement inondé, vers l’embouchure de la rivière Massa. Il s’agit d’un biotope particulièrement riche. MUSARAIGNES DU PARC NATIONAL DU SOUSS-MASSA 59 Un FIG. 4 Milieu à Euphorbes cactoïdes. 596 P. VOGEL, M. LAWRENCE & A. AGHNAJ DISCUSSION La systématique des musaraignes du Maroc a posé quelques problèmes, notamment à la suite du travail de Vesmanis & Vesmanis (1979). Celui-ci, basé sur le matériel de la Smithonian Institution, faisait miroiter l’existence de 13 espèces du genre Crocidura. Hutterer (1986) ne retenait que 5 espèces valides, interprétation confirmée par Aulagnier & Thévenot (1986b). Quatre de ces espèces vivent dans la région d’ Agadir (Aulagnier & Hermas, 1989). La plus grande espèce, décrite a l’origine sous le nom de Crocidura bolivari Morales Agacino, 1934, présente une formule chromoso- mique qui les rapproche aux crocidures géantes (Crocidura olivieri) d’ Afrique tropicale (Vogel et al., 1988). Selon analyse morphologique de Hutterer (1986), C. bolivari est synonyme de C. viaria (I. Geoffroy Saint-Hilaire, 1834). Cette interpretation a été confirmée par Maddalena (1990) qui n’a pas trouvé des differences alléliques impor- tantes par rapport a des populations de C. viaria situées au sud du Sahara (Burkina Faso). Si les unités spécifiques sont actuellement établies avec une assez grande certi- tude, deux problemes importants restent a résoudre. Le premier concerne l’origine phylogénétique de deux especes mal connues: C. whitakeri et C. tarfayaensis. Le deuxieme concerne les connaissances écologiques de toutes ces especes. Nos résultats aident à mieux comprendre l’écologie de C. viaria, seule espèce capturée. Un point intéressant concerne le rapport numérique entre musaraignes et ron- geurs. Notre première tentative de capture en 1986 a révélé la présence de C. viaria dans des terrains cultivés du village Sidi R’bat (Vogel er al., 1988). L'absence de captures de musaraignes par Benhamza (1995) dans le périmètre du parc laissait penser soit à un recul de ce groupe dû à l’utilisation de pesticides dans les terrains cultivés, soit à un effet provoqué par des changements climatiques. Les résultats présentés ici Suggerent une autre explication, soit celle d’un biais dû aux trappes utilisées. Les trappes Longworth sont connues pour leur efficacité pour capturer les micromammi- feres de petite taille. L’entrée de 4.5 x 4 cm du tunnel d’acces est attirant pour les musaraignes et le système de déclenchement est très sensible s’il est muni d’une rampe. Même Suncus etruscus avec 1.8 g peut déclencher le mécanisme, mais une fois prise, cette musaraigne minuscule arrive à s’échapper par le trou de passage du ressort. En revanche, ce systeme avec boite de nid métallique totalement fermée est moins attrayant pour les rongeurs désertiques et ne permet pas de capturer d’ animaux dépas- sant 80 g. Les trappes-cages grillagées de Manufrance utilisées par Benhamza (1995) semblent plus attractives pour les grands rongeurs, mais inefficaces pour les plus petites espèces: le déclenchement se fait par traction sur l’appät et le grillage d’une maille de 2 x | cm laisse passer toutes les espèces de musaraignes, expliquant le résultat négatif avec ce type de trappe. Concernant les exigences écologiques de C. viaria, cette espèce se trouve dans la région du parc dans presque tous les milieux, à condition que la végétation soit assez dense pour lui procurer une protection contre les prédateurs. Le milieu à végétation d’halophytes semble être son terrain de prédilection, mais des terres agricoles à couver- ture dense comme un champ de luzerne conviennent également. La couverture à elle MUSARAIGNES DU PARC NATIONAL DU SOUSS-MASSA 597 seule n’est pas suffisante puisque dans la forêt d’ Eucalyptus avec sous-bois, aucune prise n’a été faite. Ce milieu particulier semble inhospitalier, étant donné qu'il n’héberge pas une riche faune d’invertebres. Il n’est pas étonnant que les musaraignes soient quasiment absentes des dunes dont la couverture est très clairsemée. Ce milieu peut se présenter plus favorablement lors d’une année de pullulation de rongeurs; leurs galeries nombreuses peuvent alors étre exploitées par les musaraignes. La présence de C. viaria dans le milieu a Euphorbes cactoides était plutot inattendue, puisque la végé- tation se présente sous forme de touffes discontinues. Mais ce méme milieu héberge selon Benhamza (1995) une population d’Elephantulus rozeti, un autre micro- mammifere a régime insectivore. Ce dernier étant caractérisé par un rythme d’activite diurne (Seguignes, 1983), il éviterait une éventuelle compétition avec C. viaria qui est strictement nocturne (Tettamanti, 1997). La question se pose sur la valeur d’un sondage sur une période si courte. Malgré un taux de capture relativement faible (4.5%. des trappes occupées), le total de nos captures correspond a un tiers des prises effectuées par Benhamza (1995) en deux ans. La représentativité relativement bonne de notre campagne est confirmée par l’échan- tillon des rongeurs. Malgré le petit nombre d'individus, il correspond assez bien aux résultats de Benhamza (1995) pour les espèces de petite taille (présence de Gerbillus hoogstraali, G. campestris, Mastomys erythroleucus et Mus spretus), qui ont été trou- vés dans les milieux correspondant a leur préférence écologique (Tab. 1). Face a la richesse spécifique des rongeurs sur un échantillon de seulement huit individus, l’uniformité monospécifique pour les musaraignes est étonnante. Les données obtenues par l’analyse de pelotes de la région (Aulagnier & Hermas, 1989) ne montrent qu’une légère dominance de C. viaria. Or, ce travail n’a pas séparée les données selon leur origine. Une comparaison du nombre minimal d’individus provenant des deux villages de l’embouchure de l'Oued Massa (Sidi R’bat et Sidi Ouassay) avec les individus provenant du barrage Youssef ben Tachine, localité a 30 km en amont du PNSM et située dans les montagnes, est très intéressante (S. Aulagnier, comm. pers.) : sur les 19 Crocidures provenant de la cöte C. viaria domine avec 63%, suivie de C. tarfayaensis avec 21% et C. whitakeri avec 16%. Au barrage en revanche, sur les 26 Crocidures (non inclues deux C. lusitania), C. tarfayaensis dominait avec 43%, suivie de C. viaria avec 31% et C. whitakeri avec 27%. Les deux petites espèces semblent dès lors plutöt exploiter des regions plus sechardes et rocailleuses. Si les donnees bibliographiques s’accordent pour définir l’exigence écologique de C. trafayaensis en l’associant à un milieu rocheux (Aulagnier & Thévenot, 1986; Hutterer, 1986), la situation est moins claire pour C. whitakeri. Selon Heim de Balsac (1968) et Rzebik- Kowalska (1988), elle vivrait aussi en milieu rocheux, mais selon d’autres, elle serait associée aux dunes, comme le suggère l’échantillon retiré de pelotes analysées par Hutterer (1986) en provenance de Moulay Bousselham (93 C. whitakeri, 48 C. russula). Concernant l’oued Massa, cette espece a été en effet capturée une fois dans les mémes dunes fixées où notre effort est resté infructueux. Il s’agit d’une femelle portante (Fig. 5), capturée le 4.10.1989 (M. Ruedi, comm. pers.). En guise de conclusion, dans la région du sud marocain, y compris d’ Essaouira a Sidi Ifni (P. Vogel obs. pers.), C. viaria, dont l’origine est afrotropicale, est une musa- 598 P. VOGEL, M. LAWRENCE & A. AGHNAJ FIG. 5 Musaraigne capturée, photographiée et déterminée comme C. whitakeri par M. Ruedi. II s’agit d’une femelle capturée a Sidi R’bat qui a donné naissance à deux petits, avant de s’évader. Il semble que ce soit le seul individu vivant photographié. Le blanc du ventre et du dessus des pattes ainsi que le pelage grivelé sont des critères distinctifs (Aulagnier & Thévenot, 1986a). En revanche, la queue en réalité courte (27 à 39 mm pour une longueur de tête et corps de 53 à 64 mm) apparaît sur cette photo plutôt longue. raigne omniprésente dans tous les milieux à végétation dense (oueds et milieux cultivés). Toutefois, sa répartition est limitée aux basses altitudes. Plus au nord, en tous cas de Casablanca a Tanger, elle est remplacée par C. russula, d’origine paléarctique. Cette dernière supporte mieux le froid (Sparti, 1990). Comme montré ailleurs (Vogel & Maddalena, 1987), C. russula vit depuis les milieux halophiles en bordure d’estuaires (Mulay Bousselham) jusqu'à 2°800 m dans le Haut Atlas (Oukaimeden). Remarquons que ce sondage n’a pas permis de préciser l'écologie de C. whita- keri et de C. tarfayaensis qui, selon les analyses de pelotes, existent dans le périmètre du PNSM. Malheureusement, la chouette chevéche (Athene noctua) que nous avons rencontrée a plusieurs reprises semble spécialisée dans la chasse aux insectes et aux reptiles (obs. pers.), les micromammiferes ne jouant qu'un rôle secondaire dans leur régime. La chouette effraie (Tyro alba) a en revanche un régime alimentaire repre- sentant mieux la faune des petits mammiferes. Selon le faible nombre de pelotes a disposition de Benhamza (1995), ce rapace nocturne semble très rare. L'installation de nichoirs pour la chouette effraie à des endroits bien protégés de toute perturbation pourrait fournir un matériel d’étude à long terme. Ceci serait non seulement intéressant sur un plan scientifique, mais également sur un plan didactique, soit pour | instruction des élèves, étudiants et touristes qui visitent le nouveau centre d’accueil placé au cœur du parc. MUSARAIGNES DU PARC NATIONAL DU SOUSS-MASSA 599 REMERCIEMENTS Nous remercions M. Mohamed Ribi, Directeur du Parc, pour nous avoir associé au programme de recherche du PNSM, ainsi que M. Paul McCormick, Directeur de la IFCDW, pour le soutien concernant cette étude. S. Aulagnier a mis a disposition des données non publiées et M. Ruedi sa prise de vue unique de C. whitakeri. BIBLIOGRAPHIE Anonyme. Parc National de Souss-Massa, Plan d'aménagement et de gestion. Administration des Eaux et Foréts et de la conservation des sols. Pp. 11-22. AULAGNIER, S. & THEVENOT, M. 1986a. Note sur les Mammifères de la région de l'embouchure de l’Oued Massa. Bulletin de l'Institut Scientifique, Université Mohammed V, Rabat 10: 193-199. AULAGNIER, S. & THEVENOT, M. 1986b. Catalogue des Mammifères sauvages du Maroc.- Travaux de l'Institut scientifique cherifien publiés par la Société des Sciences naturelles du Maroc, Série zoologie 41: 1-164. AULAGNIER, S. & HERMAS, J. 1989. Etude biométrique des Crocidures (Soricidae, Insectivora) de la région de Massa (Souss, Maroc). Donana, Acta Vertebrata 16: 89-104. BENHAMZA, J. 1995. Mammiferes du Parc National du Souss-Massa (Agadir) : Composition et répartition cartographiée. These, Université Ibnou Zohr, Agadir, 144 pp. BRINDLEY, E., DIMMICK, C., BOXDEN, C., RiBi, M., HOFFMANN, D. & DEL Nevo, A. 1995. The bald Ibis: a species on the brink? RSPB Conservation Review 9: 76-79. HEIM DE BALSAC, H. 1968. Les Soricidae dans le milieu désertique saharien. Bonner Zoologische Beitrdge 3/4: 181-188. HUTTERER, R. 1986. The species of Crocidura (Soricidae) in Morocco. Mammalia 50: 521-534. MADDALENA, T. 1990. Systématique, évolution et biogéographie des Musaraignes Afrotropicales et Paléarctiques de la sous-famille des Crocidurinae: une approche génétique. Thése de doctorat, Université de Lausanne, 172 pp. RZEBIK-KOWALSKA, B. 1988. Studies on the genus Crocidura (Insectivora, Mammalia) in Algeria. Acta Zoologia Cracoviensia 31: 167-192. SEGUIGNES, M. 1983. La torpeur chez Elephantulus rozeti (Insectivora, Macroscelididae). Mam- malia 47: 87-91. SPARTI, A. 1990. Comparative temperature regulation of African and European shrews. Compa- rative Biochemistry and Physiology, Part A, Comparative physiology 97: 391-397. TETTAMANTI, C. 1997. Etude de la température corporelle chez trois musaraignes du genre Crocidura (Insectivora, Mammalia). Travail de diplome, Université de Lausanne. VESMANIS, LE. & VESMANIS, A. 1979. Beitrag zur Kenntnis der Crociduren-Fauna Marokkos (Mammalia Insectivora, Soricidae). 1. Die Wimperspitzmäuse aus der Sammlung der Smithonian Institution. Zoologische Abhandlungen 36/23: 11-80. VOGEL, P. & MADDALENA, T. 1987. Note sur la répartition altitudinale et la fréquence de la Musaraigne musette (Crocidura russula yebalensis) au Maroc. Mammalia 51: 465-467. VOGEL, P., MADDALENA, T. & AULAGNIER, S. 1988. Le caryotype de Crocidura bolivari Morales Agacino, 1934 (Mammalia, Soricidae). Revue suisse de Zoologie 95: 779-783. ie eae 17 REVUE SUISSE DE ZOOLOGIE 107 (3): 601-656; septembre 2000 A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People’s Republic of China, II Ivan LÖBL Museum d’histoire naturelle, case postale 6434, CH-1211 Geneve 6, Switzerland. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People’s Republic of China, II. - The present paper is the second part of a review of the scaphidiines of the People’s Republic of China. It treats the members of Scaphisoma Leach, which is the most speciose genus of the subfamily. Within the collections examined 54 species are recognized, from which the following 30 are described as new: S. inversum, S. emei- cum, S. serosum, S. subtile, S. aciculare, S. latro, S. signum, S. wolong, S. styloides, S. apertum, S. pseudantennatum, S. oppositum, S. acclivum, S. linum, S. serpens, S. parasolutum, S. pseudosolutum, S. vexator, S. pseudovarium, S. heishuense, S. paravarium, S. fibrosum, S. incisum, S. migrator, S. mutator, S. suspiciosum, S. irruptum, S. volitatum, S. vesti- gator, and S. dumosum. Scaphisoma lautum Löbl and Scaphisoma sinense Pic are placed in synonymy of Scaphisoma haemorrhoidale Reitter. A_key to the species of Scaphisoma of the People’s Republic of China is provided. Key-words: Coleoptera - Staphylinidae - Scaphidiinae - China - syste- matics. INTRODUCTION Scaphidiinae are mycophagous beetles feeding on a variety of fungi (e. g., Nuss, 1975; Lawrence & Newton, 1980; Newton, 1984). They are common on Polyporaceae, but also feed and develop on Hydnaceae, Clavariaceae, Agaricales, Heterobasidiomycetes, and slime moulds. They tunnel in carps and gills of the gilled and other mushrooms, build characteristic retreats and produce silk-lined pupal chambers (Ashe, 1984; Leschen, 1988 and 1994; Hanley, 1996). Scaphidiines are usually associated with dead wood or other decaying vegetational matter. Many species are cryptic but may be easily found in sieved forest litter. Their highest diversity is in moist, warm temperate, subtropical, and tropical forested areas. Such habitats have been largely destroyed in mainland China (see Jäch & Li, 1995), or are difficult to survey. Nevertheless, substantial new collections of scaphidiines were gathered in the course of the past decade. I have recorded 78 species in 12 genera in the first part of my review of the scaphidiines of the People’s Republic of China (Löbl, 1999). Most of these species were recorded for the first time from China, and Manuscript accepted 22.03.2000 602 IVAN LÖBL a 18 were new to science. The present study treates Scaphisoma Leach which is the most speciose genus of the subfamily. At present 54 species are recognized, 30 of which are new and described below. The total number of scaphidiines is raised from 31 species described or recorded from mainland China prior to my 1999 paper, to 122 species. Yet it is difficult to estimate how far this number is representative and to extrapolate data pertaining to species diversity of scaphidiines of China. While members of almost all genera likely to occur in the country have been found, numerous additional species in most genera may be expected. The poor knowledge of the Chinese scaphidiines may be illustrated by the two records (each for one species) from the Northeastern Region of China, whereas from the neighboring Russian Far East 19 species are known (Löbl, 1993). No scaphidiines are recorded from Inner Mongolia, Tibet, and the Northwestern areas of China although at least two common species may be expected to occur there: Scaphisoma inopinatum Löbl which is distri- buted throughout large parts of the Eurasian continent and known from Mongolia and the Russian Far East (Löbl, 1970), and Scaphium quadraticolle Solsky, widely distri- buted in Central Asia, northern Pakistan and Kashmir inlcuded. Curiously, scaphi- diines are unknown from Hainan and only two are from Guangdong, while at least 11 species occur in Hong Kong. Most material comes from Yunnan (72 species in total, 32 of them Scaphisoma) and Sichuan (with 33 species, 13 of them Scaphisoma), where the highest number of presumed narrow endemics also occur. Seven species are known from Fujian, all from a collection made in 1946 at «Kuatun» (= Guadun Village, Wuyi Shan, North Fujian; see Smetana, 1996); from other provinces usually less than 5 species are represented in the material studied. An other notable feature of the Chinese Scaphisoma ıs the relatively high number. of species of the Palaearctic S. subalpinum and the Oriental S. rouyeri groups, and the low number of species of the Oriental S. unicolor group. MATERIAL AND METHODS Reference is made only to material coming from the People’s Republic of China, referred below as to China. Most is from recent collections and housed in the Muséum d’histoire naturelle, Geneva (MHNG). Other collections which include specimens used in the present study are: MNHN Muséum National d’ Histoire Naturelle, Paris MMUE The Manchester Museum, Manchester NHMB Naturhistorisches Museum, Basel NMPC. National Museum, Praha PCAP Private collection A. Pütz, Eisenhüttenstadt PCMS Private collection M. Schülke, Berlin SMNS _ Staatliches Museum für Naturkunde, Stuttgart SNMB Slovak National Museum, Bratislava ZMB Zoologisches Museum, Berlin The methods are as given in my preceding publications: the length of the body is measured from the anterior pronotal margin to the inner apical angle of the elytra; SCAPHIDIINAE OF CHINA, II 603 the lateral keels of the pronotum and elytra, and the plane of the inner and outer apical angles of the elytra is observed in dorsal view. The presence, but not the absence, of the antecoxal puncture row of the metasternum is quoted. The length ratio of the antennal segments is measured from dry specimens, at high (100 to 200x) magni- fication. This ratio is not given in species of the S. haemorrhoidale group in which it does not provide diagnostic characters. The genitalia of all males were dissected and mounted in Canada balsam on slides. The length of the aedeagi is taken from genitalia mounted on slides. The previously described species are listed alphabetically, the new species are ranged according their relationships. Nevertheless, some males and numerous females from various collections could not be assigned to any defined species. KEY TO THE SCAPHISOMA OF CHINA I Elytra each with sutural stria extended along base to form basal stria....... 2. - Bilytraswithounbasal SA EE ia ne te RUE 18 2 Elytra with basal'striae entre joinmedtollateral sta eut ee es: 3 - Elytra with basal striae extending to midwidth of base, or to humeral areamnotyoine duto lateral striae eet mn en. e 5 3 Elyisareachswithidarkgdisealispofa ee ze ee er notatum Löbl - Eiyaradlackinsidiseal spot e Mier re Me e - 4 Antennal segment 5 much shorter than segments 3 and 4 combined ROERO I IO E MR I ei styloides sp. n. Antennal segment 5 about as long as segments 3 and 4 combined. serpens sp. n. 5 Antennal segment 5 much longer than segments 3 and 4 combined........ 6 - Antennal segment 5 about as long as, or distinctly shorter than, serments>rand 4 combined eso. septic. u O 7 6 Antennal segment 5 about 3 times as long as segment 4. . . pseudorufum Löbl - Antennal segment 5 twice as long as segment 4............. serosum Sp. N. U Metacoxallinesparallel ar 22 2 N RS SPA en 8 - Metacoxalilineyareuatelozsubansulatene mica cr ot elise cee ee 9 8 Antennal’segments’> and Gequally lone. 2... 2.2.22... aciculare sp. n. - Antennal segment 5 much shorter than segment6........... uniforme Löbl 9 Antennal segment 6 longer than segments 4 and 5 combined............ 10 - Antennal segment 6 shorter than segments 4 and 5 combined ............ 12 10 Aedeagus gradually narrowed apically, basal bulb much shorter than distal process; flagellum robust, angulate in basal portion (Fig. 27) RENE AE CES EAC a A rela cae nae Fe NR Lae ER AR ER hee, Ruy A oppositum sp.n. - Aedeagus abruptly narrowed apically, basal bulb much longer than distaliprocess:iflacellumino tan sulate e Rete e ere e oe Il Il Aedeagal apex truncate, flagellum very thin (Fig. 25) . pseudantennatum sp. n. - Aedeagal apex hook-like, flagellum robust (Figs 29, 30)..... acclivum sp. n. 12 Abdomen with microsculpture consisting of transverse striae . . . latro sp. n. 604 IVAN LOBL Abdomen lacking microsculpture, or with punctulate microsculpture..... 18 Mesocoxalllıne’subparallel'.... Rie Ro ee eee 14 Mesocoxaliline areuate or'subangulater... RE 15 Punctation on mediobasal portion of first abdominal sternite much more coarse and dense than that on apical or lateral portions of same sternite. Internal sac of aedeagus lacking spinose or squamose structures . subtile sp. n. Punctation of first abdominal sternite entirely very fine and sparse. Internal sac of aedeagus with spinose structures (Fig. 23)..... apertum Sp. n. Large species, body 2.1 to 2.3 mm long. Internal sac of aedeagus with flaselum Bee BAGO, Ge Bui. ARE A RR AREE linum sp. n. Small species, body 1.2 to 1.7 mm long. Internal sac of aedeagus with Or Without flagellum 225300 u. RR RR 16 Mesocoxal line subangulate, mesocoxal area as long as, or longer than, metacoxal area. Aedeagus asymmetrical.............. brunneonotatum Pic Mesocoxal line arcuate, mesocoxal area shorter than metacoxal area. Medeagus symmetrical). IN ORI IE 17 Internal sac of aedagus with sclerotized flagellum............ wolong Sp. n. Internal sac of aedeagus without flagellum or rods ............ portevini Pic Minute species, body length below 1 mm ......... minutissimum Champion Parserspecies body.lensthiexceding»Iimm?.. RR 19 Antennal segment 6 as long as, or longer than, segments 4 and 5 combined 2 m. ARE NOT IU TR NP AR N EEE 20 Antennal segment 6 shorter than segments 4 and 5 combined............ 21 Addomen with punctulate microsculpture. Aedeagus asymmetrical Mien DER LE RER ARE A Ted dia ahs ral ter conde a A Ge falciferum Lobl Abdomen with microsculpture consisting of striae. Aedeagus symma- (TENCE My tad e LE ce NER NORE a Japonicum Löbl Elytra with sutural striae short, starting posterior level of tip of pronotal lobe; distant from basal’marsın. . 22... u. RE eee 22 Elytra with sutural striae longer, starting at each side of pronotal lobe, CLOSE REO bere FT o ET ee er ER elytral margin 23 Abdomen with microsculpture consisting of striae. Body large, length HL SS) IVT OWS ei ee le Ps EN ee inline Man oily cn pseudosolutum sp. n. Abdomen lacking microsculpture. Body smaller, length 1.3-1.5 mm I I nn ORSO RAI laevigatum Löbl Elytra with adsutural areas conspicuously widened toward base, sutural Stnlae-aneulateng nn ee eu O ICE dohertyi Pic Elytra with adsutural areas not particularly widened anteriorly, sutural stniaenot ansulate antemorly ... 2.2 inh ke he oo ONORE 24 Prohypomera distinctly microsculptured CCR, RE oo eee 25 Prohypomera lacking microsculpture.-= >... SONS SS 26 Anterior two thirds of elytra coarsely punctate, posterior third of elytra very finely punctate, appearing impunctate................ rufescens (Pic) 29 30 31 SCAPHIDIINAE OF CHINA, II 605 Entire elytra with fairly coarse punctation............ pseudodelictum Löbl Antennal segment 3 conspicuously long, about as long as two thirds of SEINE CRE ANNE ER ARR ER I DANA PN parasolutum Sp. n. Antennal segment 3 short, usually about as long as one third, not longer thamhalft: ose nent AH Wr La PERL Re 27] Exposed abdominal segments lacking microsculpture, or with punc- tulatesmieroseulptures# nl RE a ee URL Re A ee O. 28 All exposed abdominal segments with microsculpture consisting of transverse _ striae, or first sternite lacking microsculpture, remainder Of abdomen with distinct striate microsculpture NN 31 Mesocoxal line substriangular, mesocoxal area longer than metacoxal ANG AR D PIE RENT Mask SEE ah ER EEE maindroni Achard Mesocoxal line arcuate, mesocoxal area as long as or shorter than Metacoxalrarearz ur, HM Sn DEEE N NOSTRO I AE eet rae 29 Elytra each with dark, transverse, central and apical fasciae . . . invertum sp. n. Elytrallackinsidarksiasciae We 22:2 DANA, EE NL PRES nr ER DI IE ONE i o 30 Aedeagus asymmetrical, with parameres extremely narrow I MOR UN PEN LEA O RARE He RATA cts ri DETAIL dar unicolor Achard Aedeagus symmetrical, with parameres normally wide .......... segne Löbl Pronotum light, with dark median fascia extended from base to anterior LEO MOT EER adh baal RIEN RE EAN ÇA ANT Er EUR LENS vexator sp. n. Rronotum withoutidark median fasciatin qm. nen ciel ae eee ores 32 Pronotum dark brown or black on large central part of disc, light ochreous along lateral margins; elytra each with black basal area and very dark brown or black subapical spot ................ paravarium sp. n. ColounspattermiGueheEre mtyecencvonc: ences REN Sed ck a MPa ee LIEB a 33 Elytra and pronotum uniformy ochreous or dark brown, or elytra slightly:darkened: subapicallyar... wen ziert nee AE 34 Elytra bicolorous, often with distinctive colour patter, not darkened subapicallyaneı. 8 Aus EME DER RI Re 38 Aedeagus with basal bulb partly overlapping distal process ............. 33 Aedeagus with basal bulb not overlapping distal process ............... 36 Parameres with distinct membranous lobe in middle portion . . . . pressum Löbl Parameres without distinct membranous lobe (Fig. 61)........ incisum Sp. n. Parameres expanded proximally to form large basal lobes entirely exposed in dorsal view; apical portion of parameres widened in dorsal VICE (RIOT NER EIN IN Ia BE signum sp. n. Parameres not expanded proximally to form lobes and usually not widenedkapicallys an ro IR Ce CE coh RR NA RAR. 37 Antennal segment 6 wider than segment 5. Body dark reddish-brown. Adsuturalfareas oßelytrardistinetly zaisedn m en. emeicum SP. n. Antennal segments 5 and 6 equally wide. Body light ochreous or light reddish-brown. Adsutural areas of elytra flat ................ nakanei Löbl 606 40 41 47 IVAN LOBL Elytra each with darkened basal area usually forming triangular pattern, and often with additional dark central or subapical spot ................ 39 Elytra each with apical sixth to apical third light, remainder of surface uniformly reddish-brown to black, as pronotum 72... 1.2.2. Gases 42 Elytra usually with darkened central area, prothorax light ochreous. Parameres of aedeagus arcuate, each with large, wide membranous lobe RO de ne ele cause nie eine COUDE LES atronotatum Pic Elytral not darkened in center: prothorax dark. Parameres usually with HAROWMSMANAOBES RENE TA 12222... See oe a STE 40 Elytra with inner apical angles prominent and acute. Elytral punctation CONSPICUOUSIVICORESCHRE AU. Gi ae EP RI fibrosum Sp. n. Elytra with apical angles rounded, not prominent. Elytral punctation not onspieuouslylcoarser.. wh. Ga ll 2.2 LI REVO ISTE 4] Parameres of aedeagus lobed in middle, arcuate in apical half (Fig. 46) IRPINO CRONO eat nichts, AV IO Pree EAUX SARE pseudovarium sp. n. Parameres of aedeagus lacking lobes, arcuate in apical fourth (Fig. 49) VR ISTAT, STAVO Den ARTERIE SIE heishuiense sp. n. Aedeagus with distal process completely overlapped by basal bulb....... 43 Aedeagus with distal process not, or partly overlapped by basal bulb...... 45 Aedeagus with parameres arcuate in dorsal view ...................... 44 Aedeagus with parameres sinuate in dorsal view (Fig. 71) .. suspiciosum sp. n. Aedeagus with parameres widened posterior middle, with wide apical portion; internal sac with subapical narrow rod, lacking central rod (RSS) Reta O Sake ee LENS TAT PER irrumptum sp. n. Aedeagus with parameres equally wide in apical half; internal sac with long, narrow, centralirod (Figs, 83, 85)... 2... LEE vestigator sp. n. Internal sac of aedeagus with one or two sclerotized rod/s .............. 46 Imtermallsac of aedeagus lacking rods . Eee eee eee 47 Parameres of aedeagus curved, wide, in apical portion much wider than al namowest point near middle (Fig. S2)E 2... SERE volitatum sp. n. Parameres of aedeagus with parameres narrow, almost straight, in apical portion ashwide ASIA EAP EEE cederholmi Löbl Internal sac of aedeagus with several, long, strongly sclerotized, Straight or slightly oblique, apical:denticles tie eee a= eee 48 Internal sac of aedeagus without apical denticles, or with cluster of numerous; short, curved; apical denticles 52... i Zen eee 50 Internal sac of aedeagus with subapical, triangular, sclerotized plate (Rie RS Sed u. el cutee st cates akt au bee dumosum sp. n. Internal sac of aedeagus lacking subapical, sclerotized plate............. 49 Denticles present throughout entire width of apical portion of internal SACHE PR ee ee HORA E SIE ROOT migrator Sp. n. Denticles present in medio-apical portion of internal sac only . . morosum Löbl Internal sac of aedeagus with two subapical clusters of strongly sclero- ved curved denticles i. SE geminatum Löbl SCAPHIDIINAE OF CHINA, II 607 - Internal sac of aedeagus without subapical clusters of strongly sclero- GIZECUGEMEICIES rarest RE ae. ENA VICI EN NL Tot, SI DI Basal portion of internal sac of aedeagus lacking median row of denticles . 52 - Basal portion of internal sac of aedeagus with long, median row of Stronsilvesclerotizedidenticles 2 CU ah. NANTERRE ee NE 52 Punctation on large basal portion of elytra very fine, contrasting with punctation on elytral center. Internal sac without basal spicules . innotatum Pic - Punctation on most of basal area of elytra about as coarse as that on elytral center. Internal sac with proximal spicules ...... fortipatum Champion 53 Internal sac of aedeagus with pair of lateral lobes; paramers weakly arcuate. Elytra often with reddish, humeral spot...... haemorrhoidale Reitter - Internal sac of aedeagus lacking lateral lobes; parameres strongly arcuate (Fig. 68). Elytra without humeral spot.............. mutator Sp. n. NEW RECORDS AND COMMENTS Scaphisoma atronotatum Pic, 1920 Redescription, aedeagus: Löbl, 1970: 152, Figs 5, 6; Löbl, 1990b: Figs 67, 68. Materiel examined. China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont, 2 (MHNG); Gaoligong Mts, 2200-2500m, 24°57’N, 98°45’E, 8-16.v.1995, O. Semela, 1 (NHMB) and same data but 1500-2500m, 17-24.v. 1995, 4 (NHMB, MHNG): Yunnan, Henduan Mts, Meili, 2700m, 28°06° N 98°54’E, 5-8.v11.1996, O. Semela, 1 (MHNG): Yunnan, Kumming, western hills, 9.vii.1990, L. & M. Bocak, 1 (NHMB); Yunnan, Yipinglang, 1800m, 25°05’N, 101°53’E, 8-10.vi.1993, Bolm, 1 (NHMB); same but 1800-2000m, 25°04’, 17- 20.v1.1994, V. Kuban, 2 (NHMB). Distribution. Nepal; Burma; Thailand; China: Yunnan. Scaphisoma brunneonotatum Pic, 1923 Aedeagus: Löbl, 1980, Figs 27, 28; diagnostic characters: Löbl, 1982: 105. Material examined. China, Hong Kong, Aberdeen, 10.viii.1996, J. Fellowes, 16 (MHNG); Hong Kong, Saikung, Tai Mong Tsai, 1.vii1.1996, J. Fellowes, 24 (MHNG); Hong Kong, iil, v, 1x-xil. 1996, G. de Rougemont, 36 (MHNG); Hong Kong, N.T., Tai Po Kau, iv.1992, G. de Rougemont, 11 (MHNG); Guangxi, Guilin, 22.1x.1995,-G. de Rougemont, 12 (MHNG). Distribution. Japan: Ryukyu’s; Taiwan; Thailand; Vietnam; China: Guangxi, Hong Kong. Scaphisoma cederholmi Lobl, 1971 Diagnostic characters, aedeagus: Löbl, 1986a: 193, Fig. 74. Material examined. China, Hong Kong, v.1996, G. de Rougemont, 2 (MHNG). Distribution. Sri Lanka; India; Thailand; China: Hong Kong. Scaphisoma dohertyi Pic, 1915 Diagnostic characters, aedeagus: Löbl, 1981b: 107, Fig. 5. 608 IVAN LOBL Material examined. China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont, 2 (MHNG); Yunnan, Mengyang Nat. Res., ca 500m, 9.1x.1994, S. A. Kurbatov, 3 (MHNG). Distribution. India; Indonesia; Thailand; Malaysia; Vietnam; China: Yunnan. Scaphisoma falciferum Löbl, 1986 Description, aedeagus: Löbl, 1986b: 359, Figs 14, 15. Material examined. China, Yunnan, Gaoligong Mts, 25°22°N, 98°49’E, 17-24.v.1995, O. Semela, 1 (MHNG). Distribution. Pakistan; India; Nepal; China: Yunnan. Scaphisoma forcipatum Champion, 1927 Diagnostic characters, aedeagus: Löbl, 1986a: 198, Figs 88-91. Material examined. China, Yunnan, 25°22°N, 98°49’E, Gaoligong Mts, 1500-1800m, 17-24.v.1995, O. Semela, 1 (NHMB). Distribution. Pakistan; India; Nepal; China: Yunnan. Scaphisoma geminatum Löbl, 1986 Description, aedeagus: Löbl, 1986a: 212, Figs 112-114. Material examined. China, Guangdong, Qi Mu Zhang, 5.iv.1997, leaf litter, J. Fellowes, 1 (MHNG). Distribution. India; China: Guangdong. Comments. This specimen has the two central clusters of robust denticles in the internal sac of the aedeagus oriented apically, and notably longer and those in the specimens from India. Besides, it differs slightly in the shape of the apical portion of the parameres and in the structure of the basal portion of the internal sac. Scaphisoma haemorrhoidale Reitter, 1877 Redescription, aedeagus: Löbl, 1970: 791, Figs 76, 77. Type material examined. Scaphisoma sinense Pic, 1920: 5; lectotype d (MNHM), paralectotype 2 (SNMB), from Nankin - syn. n. Scaphisoma lautum Löbl, 1965b: 30; holotype d from Fujian, Kuatun (MNHM) - syn. n. Additional material examined. China, Beijing, Xiaolongmen, 1200m, 1.v11.1993, G. de Rougemont, | (MHNG); Beijing, airport, 50m, 17.v.1997, M. Schiilke, 1 (PCMS); Jiangsu, Nanjjing, 17.viii.1994, G. de Rougemont, 2 (MHNG); Jiangsu, Nanjing Agric. Univ., vii.1991, R.J.Cooter, 1 (MMUE); "Nankin", 1 (MHNG); Hubei, Chengde, 15.1x.1995, G. de Rougemont, 1 (MHNG); Luoyang, viii.1982, G. de Rougemont, 4 (MHNG). Distribution. Japan; Korea; Russian Far East; China: "Manchuria", Beijing, Hubei, Jiangsu, Liaoning, Fujian, Yunnan. Comments. The variability of some characters was underestimated by Löbl (1965b). In addition, the lectotype of S. sinense and the holotype of S. lautum have the internal sacs of the aedeagi extruded, making observation of the diagnostic structures difficult. It is notable that almost all specimens from China have the subhumeral area of the elytra reddish, in addition to the light apical area. A similar colour pattern is rarely present in specimens from Japan. SCAPHIDIINAE OF CHINA, II 609 Scaphisoma innotatum Pic, 1926 Diagnostic characters, aedeagus: Löbl, 1881b: 110, Fig.8; redescription, aedeagus: Löbl, 1986a: 2201, Figs 92-94. Material examined. China, Yunnan, Ruili, 4.11.1993, G. de Rougemont, 4 (MHNG). Distribution. India; Nepal; Thailand; Vietnam; China: Yunnan. Scaphisoma japonicum Löbl, 1965 Redescription, aedeagus: Löbl, 1970: 779, Figs 62, 63. Material examined. China, Liaoning, Mt. Shi Fang Ding, Dandong City, 25.v111.1996, J. Li, 9 (MHNG). Distribution. Far East Russia; Japan; China: Liaoning. Scaphisoma laevigatum Löbl, 1970 Description: Löbl, 1970: 755, Figs 29, 30; diagnostic characters, aedeagus: Löbl, 1980: 103, Fig. 15. Material examined. China, North Yunnan, 30km N of Lijiang, 3000m, 3.v11.1990, L. & M. Bocäk, 2 (MHNG NHMB); Yunnan, Jizushan, 2500-3100m, 25°58’N, 100°21°E, 30.v.- 3.v1.1993, V. Kuban, | (MHNG) and Bolm, | (NHMB). Distribution. Far East Russia; Japan; Taiwan; China: Yunnan. Scaphisoma maindroni Achard, 1920 Diagnostic characters, aedeagus: Löbl, 1979: 102, Fig. 24. Material examined. China, Hong Kong, iii-iv. 1996, G. de Rougemont, 4 (MHNG); Guizhou, Huaxi, x.1986, G. de Rougemont, 4 (MHNG); Yunnan, Shi Lin, 8.x.1985, G. de Rougemont, 1 (MHNG). Distribution. Pakistan; India; Nepal; Thailand; Vietnam; China: Guizhou, Hong Kong, Yunnan. Scaphisoma minutissimum Champion, 1927 Redescription, aedeagus: Löbl, 1992: 538, Fig. 126. Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 10.1x.94, litter and 12.1x.1994, rotten wood, S. A. Kurbatov, 2 (MHNG). Comments. Both specimens are females very similar in external characters to specimens of S. minutissimum from North India. Their indentification is tentative in absence of male. Scaphisoma morosum Löbl, 1990 Description, aedeagus: Löbl, 1990b: 589, Figs 129-131. Material examined. China, Yunnan, Ruili, 4.11.1993, G. de Rougemont, | (MHNG). Distribution. Burma; Thailand; China: Yunnan. 610 IVAN LOBL Scaphisoma nakanei Löbl, 1980 Description, aedeagus: Löbl, 1980: 107, Figs 22, 23. Material examined. China, Jiangsu, Nanjing, 17.viii.1994, G. de Rougemont, 4 (MHNG); Jiangsu, Nanjing, Zijishan, 8.v.1996, J. Cooter, 2 (MHNG); Hong Kong, iii, x- x11.1996, G. de Rougemont, 7 (MHNG). Distribution. Taiwan; China: Jiangsu, Hong Kong. Scaphisoma notatum Löbl, 1986 Description, aedeagus: Löbl, 1986a: 154, Figs 16, 17. Material examined. China, Yunnan, Gaoligong Mts, 2200-2500m, 24°57°N, 98°45’E, 8-16.v.1995, O. Semela, 2 (NHMB, MHNG); West Sichuan, Ganzi Tibet Aut. Pref., Luding Co. W Erlangshan-Pass, 2600m, 7km SSE Luding, 29°51’N, 102°15’E, 20.vi.1999, M. Schülke, 2 (PCMS); West Sichuan, Ya’an Pref., Shimian Co., Shimian-Ganluo, 27km south Shimian, 2450m, 29°02’N 102°31’48° E, springfedswamp, 8.v11.1999, A. Pütz, 2 (PCAP). Distribution. Pakistan; North India; Nepal; China: Sichuan, Yunnan. Scaphisoma portevini Pic, 1920 Redescription, aedeagus: Löbl, 1965b: 29, Figs 2a,b. Materiel examined. China, Yunnan, Lijiang, 26°53’N, 100°18’E, 1800m, 23.vi.- 21.vii.1992, S. Becvar, 8 (MHNG); Yunnan, Dali, 1600-2000m, 5-8.vii.1990, L. & M. Bocak, 1 (NHMB); Yunnan, Jizu Shan, 2500-2700m, 25°58’N, 100°21°E, 6-10.v11.1994, V. Kuban, 13 (NHMB, MHNG); Yunnan, Yipinglang, 1800-2000m, 25°04’N, 101°53°E, 17-20.vi.1994, V. Kuban, 3 (NHMB, MHNG): (NHMB, MHNG); Yunnan, Shi Lin, 8.x.1985, G. de Rougemont, 2 (MHNG); Yunnan, Ruili, 111.1993, G. de Rougemont, 1 (MHNG); Sichuan, Mt. Emei, 1700m, 22.1x.1994, S. A. Kurbatov, 1 (MHNG); South Sichuan, nr. Xichang, 1600m, 28.vii.1996, litter, S. A. Kurbatov, | (MHNG); NE Guangxi, 10km south Longsheng, ca 1000m, 15.v1.1995, S. A. Kurbatov, | (MHNG. Distribution. Korea; Japan; China: Guangxi, Sichuan, Yunnan. Comments. This species is very similar to S. rufum Achard for which it was mistaken a few times (see comments in Löbl, 1986a). As the shape of the internal sac of S. portevini is diagnostic only dissected males are recorded above. Scaphisoma pressum Löbl, 1990 Descripotion, aedeagus: Löbl, 1990: 571, Figs. 92-95. Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 11.1x.1994, S. A. Kurbatov, 4 (MHNG). Distribution. Thailand: China: Yunnan. Scaphisoma pseudodelictum Löbl, 1986 Description, aedeagus: Löbl, 1986a: 180, Figs 54, 55. Material examined. China, Yunnan, Ruili, 4.11.1993, G. de Rougemont, 1 (MHNG); Xishuangbanna, 24.1.1993, G. de Rougemont, 2 (MHNG); Yunnan, Mengyang Nat. Res., ca 500m, 9.1x.1994, S. A. Kurbatov, 4 (MHNG). Distribution. India; Thailand; China: Yunnan. SCAPHIDIINAE OF CHINA, II 611 Scaphisoma pseudorufum Löbl, 1986 Description, aedeagus: Löbl, 1986a: 143, Fig. 3. Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 9-14.1x.1994, S. A. Kurbatov, 7 (MHNG); Yunnan, Gaoligong, 1500-2500m, 25°22°N, 98°49'E, 17- 24.v.1995, O. Semela, 3 (NHMB, MHNG); Northeast Guangxi, 10km south Longsheng, ca 1000m, 15.v1.1995, S. A. Kurbatov, 1 (MHNG). Distribution. North India; Nepal; China: Guangxi, Yunnan. Scaphisoma rufescens (Pic, 1920) Redescription, aedeagus: Löbl, 198 1a: 157, Fig. 4. Material examined. China, Yunnan, Ruili, 4.11.1993, G. de Rougemont, 1 (MHNG); Xishuangbanna, 24.1.1993, G. de Rougemont, 2 (MHNG); Mengyang Nat. Res., ca 500m, 9.1x.1994, S. A. Kurbatov, 3 (MHNG). Distribution. East Malaysia; Singapore; Thailand; Vietnam; China: Yunnan. Scaphisoma segne Löbl, 1990 Description, aedeagus: Löbl, 1990b: 568, Figs 87-89. Material examined. China, Sichuan, Mt. Emei, 1500 and 1800m, 21 and 24.1x.1994, S. A. Kurbatov, 4 (MHNG); Sichuan, Mt. Emei, Wannian, 1050m, 19-30.111.1999, W. Schawaller, 1 (SMNS); Zhejiang, Lin’an County, W Tianmu Shan N.R., ca 500m, 16.v.1996, J. Cooter, 1 (MHNG); Yunnan, Yipinglang, 25°04’N, 101°55°E, 1800-2000m, 17-20.v1.1996, V. Kuban, 6 (NHMB, MHNG). Distribution. Thailand; China: Sichuan, Yunnan, Zhejiang. Comment. Four males are present in the collections examined, and all were dissected. Their aedeagi are very similar to that of holotype but they lack the large sclerified denticles in the internal sac. Scaphisoma unicolor Achard, 1923 Redescription, aedeagus: Löbl, 1970: 772, Figs 50, 51; aedeagus: Löbl, 1980: Fig. 26. Material examined. China, South Yunnan, Mengyang Nat. Res., ca 500m, 11.1x.1994, S. A. Kurbatov, 1 (MHNG). Distribution. India; Nepal; Thailand; Japan; Russian Far Easta; Taiwan; China: Yunnan. Scaphisoma uniforme Löbl, 1986 Description, aedeagus: Löbl, 1986a: 157, Fig. 20. Material examined. China, Yunnan, Weibaoshan, 25°12°N 100°24’E, 2000-3000m, 29- 30.vi.1992, V. Kuban, 16 (MHNG). Distribution. North India; Nepal; China: Yunnan. Comments. This species is variable in the colour of the body and elytral punctation. The shape of the basal portion of the flagellum is diagnostic. 612 | IVAN LOBL DESCRIPTIONS OF NEW SPECIES Scaphisoma invertum sp. n. Figs 1, 2 Holotype d : China, Northeast Guangxi, 10km south Longsheng, ca 1000m, 15.vi.1995, forest litter, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 2 4,7 2; South Yunnan, Mengyang Nat. Res., ca 500m, 14.1x.1994, in rotten wood, S. A. Kurbatov, 2 dé (MHNG); Yunnan, Maguan, 23°04°N, 104° 25’E, 1500-1600m, 25-26.vi.1994, V. Kuban, 1 ¢ (NHMB). Description. Length 1.3-1.5 mm. Body ochreous. Elytra usually dark along base, in middle and near apices. Central and subapical dark area usually distinct and forming transverse fasciae. Surface between dark areas lighter than pronotum. Apical abdominal segments and appendages light ochreous or yellowish. Length ratio of antennal segments 3° to Il as:follows: 3: 9: 11: 12: 16: 13: 15: 5317 Golotype): segment 4 narrow, 4 times as long as wide; segments 5 and 6 almost equally wide, distinctly wider than segment 4, each about 4 times as long as wide; segments 7 and 8 each about 4 times as long as wide; segment 11 about 3 times as long as wide. Pronotum with lateral margins almost evenly rounded; lateral keels entirely concealed or exposed in basal half; punctation sparse and extremely fine. Tip of scutellum exposed. Elytra each with lateral margin slightly arcuate near base and apices, oblique in middle; apical margin slightly rounded; inner apical angles at same level as outer apical angles; sutural margin not or slightly raised; adsutural areas flat; sutural striae almost parallel, near base slightly curved, not extended along basal margin; discal punctation fairly dense and very fine, more distinct than that on pronotum. Prohypo- mera smooth. Mesepimeral ridge slightly longer than interval to mesocoxa. Meta- sternum flat in middle portion, very finely and sparsely punctate, lacking median impressions and without microsculpture. Mesocoxal linea arcuate, finely punctate; mesocoxal area 0.05 mm long. Metepisternum in same plane as metasternum, flat, moderately narrowed anteriorly, with suture straight, rounded only at angles. Abdo- minal punctation even, very fine and sparse. First exposed sternite lacking micro- sculpture. Metacoxal lines arcuate, rather coarsely punctate; metacoxal areas 0.08 mm long. Apical abdominal tergites and sternites with punctulate microsculpture. Male sexual characters. Segments 1 to 3 of protarsi widened. Aedeagus (Figs 1, 2) 0.42-46 mm long. Median lobe symmetrical, weakly sclerotized, with large basal bulb. Ventral tubercles large, prominent. Distal process of median lobe short, obliquely inflexed, gradually narrowed, with apex wide in dorsal view, abruptly narrowed and with acute tip in lateral view. Internal sac tubular, long, widened api- cally, looped basally, lacking sclerites, covered by dense, minute, squamose or denti- cular structures. Parameres moderately bent in dorsal and lateral views, almost evenly wide. Comments. This species has the aedeagus structuraly similar to that in S. invisum Löbl, S. discretum Löbl, and S. binaluanum Pic. As the aedeagi in these species lack obvious derived characters, their relationships remain uncertain. Scaphi- soma inversum may be redily distinguished from S. discretum and S. invisum by the colour pattern of the elytra. In addition, S. invisum is characterized by the antennal SCAPHIDIINAE OF CHINA, II 613 segment 5 longer than segment 6, the mesocoxal areas relatively very large, longer than the metacoxal areas and the first abdominal sternite microsculptured. The new species differs notably from S. discretum by the antennal segment 4 much longer and the antennal segment 6 much shorter, compared with segment 5, and by the abdominal segments lacking striate microsculpture. It may be distinguished readily from S. binaluanum by the lack of striate micrsosculpture on abdominal segments. Scaphisoma emeicum sp. n. Figs 3, 4 Holotype d : China, Sichuan, Mt. Emei, 1700m, 22.1x.1994, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 1d, 29; same data but 23.ix., 1d; same data but 1500m, 21.IX, 26; same data but 1800m, 24.ix., 1d (all MHNG). Description. Length 1.4-1.5 mm. Body uniformly very dark brown or almost blackish. Apex of abdomen and appendages ochreous, tarsi and antennae lighter than tibiae and femora. Length ratio of antennal segments 3 to 11 as follows: 3: 9: 11: 11: 15: 12: 14: 14: 16 (holotype); segments 4 and 5 each about 4 times as long as wide; segments 6, 7, 8 and 11 each about 3 times as long as wide. Pronotum with lateral margins evenly arcuate; lateral keels not visible; punctation dense and very fine, consisting of minute, very shallow and not well delimited punctures. Scutellum completely covered by pronotal lobe, or minute point of scutellum exposed. Elytra each with slightly arcuate lateral margin; lateral keel not visible; apical margin truncate; inner apical angle lying at, or somewhat posterior to level of outer angle; sutural margin not raised; adsutural area flat near base, raised in posterior two thirds; sutural striae diverging in posterior third, parallel from apical third to level of pronotal lobe, slightly curved along pronotal lobe, not extended along basal margin. Prohy- pomera smooth. Mesepimeral ridge longer than interval to mesocoxa. Metasternum without microsculpture, flat in middle and without medio-apical impressions. Meta- sternal punctation fine and fairly dense, very fine on latero-anterior area. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.05 mm long. Metepisternum flat, in same plane as metasternum, strongly narrowed anteriorly, with suture straight. Abdo- men with microsculpture consisting of transverse striae and/or short waves. Punc- tation on pygidium extremely fine. First exposed sternite with punctation dense and fairly coarse in middle, sparse and very fine on lateral portions. Punctation on following sternites obsolete. Metacoxal line strongly arcuate, finely punctate; meta- coxal area 0.07 mm long. Sexual characters of male. Segments | to 3 of protarsi widened. Aedeagus (Figs 3, 4) 0.43-0.47 mm long. Median lobe symmetrical, with basal bulb fairly large and moderately sclerotized. Ventral tubercles large, slightly prominent. Distal process of median lobe slightly shorter than basal bulb, inflexed, curved, gradually narrowed apically, with tip obtuse in dorsal view, acute in lateral view. Dorsal valve single, membranous. Parameres moderately inflexed, wide, gradually narrowed toward apical third, slightly sinuate in dorsal view, straight in lateral view. Internal sac with denti- culate and spinose structures. Rows of large and stronger sclerotized denticles distinct in lateral view, in ventro-apical part of internal sac. 614 IVAN LOBL Comments. This species is likely related to S. vagans Löbl and S. iriomotense Lobl, with which it shares most of the aedeagal characters. The new species may be easily distinguished from S. sagans by the metepisternal suture which is convex, and the elytral punctation which is notably coarser in the latter species. Scaphisoma iriomotense may be easily distinguished by the antennaae with segments 3 to 5 about as long as segment 6. Scaphisoma serosum sp. n. Figs 5, 6 Holotype dé: China, Central Sichuan, Wolong Nat. Res., 1000m, 24.v.1994, rotten wood, S. A. Kurbatov (MHNG). Paratypes: same data as the holotype, 18 (MHNG); same data but 1700m, 19.v.1994, litter, 1d (MHNG). Description. Length 1.7 mm. Body dark reddish-brown. Elytra each with wide, light ochreous, subapical fascia. Femora and tibiae reddish, lighter than thorax. Apical abdominal segments, tarsi and antennae light, ochreous. Length ratio of antennal segments 3 to 11 as: 5: 10: 20: 19: 25: 20: 25: 25: 27 (holotype); segments 4 and 5 each about 4 times as long as wide; segments 6, 7 and 8 each about 3 times as long as wide; segment 11 about 3.5 times as long as wide. Pronotum with evenly arcuate lateral margins; lateral keels concealed: punctation dense and fine, punctures much smaller than puncture intervals. Tip of scutellum exposed. Elytra each with modera- tely rounded lateral margins; lateral keel hardly visible near base and near apex, not visible in middle portion (dorsal view); apical margin slightly arcuate; inner apical angle rounded, not prominent, situated at same level as outer apical angles; sutural margin not raised; adsutural area flat: sutural stria parallel to suture, curved near base and extended laterally to reach outer fourth of basal width, not joined to lateral stria; discal punctation fine and dense, slightly less fine than that on pronotum, with punc- ture intervals 2 to 5 times larger than puncture diameters. Pygidium with micro- sculpture consisting of very short striae. Prohypomera smooth, impunctate and lacking microsculpture. Mesepimeral ridge longer than interval to mesocoxa. Meta- sternum without microsculpture; medio-apical portion flattened, with fine and dense punctation, lateral and medio-anterior portions of metasternum with punctation very fine and scattered. Metepisternum narrowed anteriorly, flat, lying on slightly lower plane than metasternum, suture slightly sinuate. Mesocoxal line arcuate, finely punc- tate; mesocoxal area 0.04 mm long. Exposed abdominal sternites with microsculpture consisting of short striae, partly forming meshes. Abdominal punctation very fine and Sparse on most surface, relatively coarse on median portion of first sternite. Meta- xocal line arcuate, rather coarsely punctate; metacoxal area 0.09-0.10 mm long. Male sexual characters. Segments | to 3 of protarsi widened. Aedeagus (Figs 5, 6) 0.32-0.33 mm long, symmetrical. Median lobe with basal bulb fairly small, moderately sclerotized, ventral tubercles large, prominent. Distal process of median Fics 1 to 6 Aedeagi in Scaphisoma, dorsal and lateral views: 1 and 2, S. invertum sp. n.; 3 and 4, S. emeicum Sp. n.: 5 and 6, S. serosum sp. n. Scale bars = 0.1 mm. 615 SCAPHIDIINAE OF CHINA, II Ce SN 616 IVAN LOBL lobe narrow near bulb, strongly arcuate and inflexed in lateral view, slightly widened toward tip in dorsal view, with tip acute. Parameres almost evenly wide, slightly sinuate in dorsal view, slightly arcuate in lateral view. Internal sac simple, tubular, with apical spinose structures. Comments. This species resembles S. flavofasciatum Löbl in the conspicuous colour pattern but it may be easily distinguished by the relative length of the antennal segments 5 and 6, the length of the metacoxal area, and the abdominal micro- sculpture. Scaphisoma flavofasciatum has the segment 6 significantly longer than the segment 5, the first exposed abdominal sternite lacks microsculpture, and the metacoxal areas are much shorter than in S. serosum. The male characters are unknown from S. flavovasciatum. Although the aedeagus of S. serosum is similar to that of S. curtipenne (Pic), these species are probably not closely related. These two species differ drastically in exoskeletal characters (for S. curtipenne see Löbl, 1973), and their aedeagi lack obvious derived characters. Scaphisoma signum sp. n. Figs 7, 8 Holotype d: China, Central Sichuan, Wolong Nat. Res., 1500m, 22.v.1994, S. A. Kurbatov (MHNG). Description. Length 1.5 mm. Body reddish-brown, subapical portion of elytra darkened, apices of elytra, abdominal apex, femora and tibiae lighter than thorax, antennae and tarsi yellowish. Length ratio of antennal segments 3 to 11 as follows: 5: 11: 15: 15: 20: 15: 20: 20: 25; segment 4, 5 and 11 each about 4 times as long as wide; segments 6, 7 and 8 each about 3 times as long as wide. Pronotum with lateral margins evenly arcuate; lateral keels not visible; punctation sparse and very fine, consisting of minute but well delimited punctures. Tip of scutellum exposed. Elytra each with lateral margin rounded in basal third, oblique in apical two thirds; lateral keel entirely exposed in dorsal view; apical margin arcuate; inner apical angle rounded, not prominent, on same level as outer apical angles; sutural margin not raised; adsutural area flat; sutural stria parallel to suture, curved along pronotal lobe, not extended along basal margin; punctation near base as fine as that on pronotum, on most surface less fine, consisting of fine, not well delimited punctures. Prohypomera and mesepisterna smooth. Mesepi- meral ridge as long as interval to mesocoxa. Metasternum flattened medio-apically, with punctation sparse and very fine on most of surface, dense near inner margin of metacoxae. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.03 mm long. Metepisternum flat, moderately narrowed anteriorly, lying in same plane as meta- sternum, suture straight except near angles. Abdominal tergites and sternites with microsculpture consisting of transverse striae, punctation sparse and very fine. Metacoxal line arcuate, finely punctate; metacoxal area 0.09 mm long. Male sexual characters. Segment | to 3 of protarsi slightly widened. Aedeagus (Figs 7, 8) 0.44 mm long. Median lobe fairly sclerotized, symmetrical. Basal bulb large, with robust, not prominent ventral tubercle. Distal process of medin lobe gradually narrowed, with tip acute, strongly inflexed and curved, and dorsal valve single, sclerotized. Parameres with base very wide, narrowed toward curved apical portion, bearing large membranous lobe. Internal sac complex, with two median rows SCAPHIDIINAE OF CHINA, II 617 of large denticles and group of large, subapical denticles; membranes very finely denticulate apically, and finely spinose and incurved proximally. Comments. This new species is very similar to S. taiwanum Löbl in both exoskeletal and genital characters. It may be distinguished from the latter species by its aedeagus with parameres bearing large lobes and by the internal sac with rows of large denticles. Scaphisoma aciculare sp. n. Figs 9 to 11 Holotype d: China, Yunnan, Gaoligong Mts, 2200-2500m, 24°57’N, 98°45’E, 8-16.v.1995, S. Beévat (MHNG). Description. Length 2.10 mm. Body uniformly very dark reddish-brown, apex of abdomen, antennae and tarsi lighter. Length ratio of antennal segments 3 to 11 as follows: 7: 15: 19: 19: 25: 20: 23: 24: 28; segments 4, 5 and 8 each about 4 times as long as wide; segments 6, 7 and 11 each 3 times as long as wide. Pronotum with evenly arcuate lateral margins; lateral keels concealed; discal punctation extremely fine, fairly dense. Tip of scutellum exposed. Elytra each with lateral margin slightly rounded; lateral keel visible near base only; apical margin truncate, inner apical angle rounded, not prominent, at same level as outer apical angles; adsutural area flat anteriorly, slightly raised in apical two thirds; sutural stria curved near base and extended to outer third of basal width, forming basal stria not joined to lateral stria; discal punctation fairly dense and fine, consisting of punctures not well delimited, mostly much smaller than puncture intervals; punctation near base finer than that on discal center, similar to that on pronotum. Prohypomera smooth. Mesepimeral ridge about as long as interval to mesocoxa. Metasternum without microsculpture, very finely and sparsely punctate, convex in middle, with two medio-apical impressions. Mesocoxal line arcuate, distinctly punctate; mesocoxal areas 0.06 mm long. Metepis- terna flat, moderately narrowed anteriorly, lying on lower plane than metasternum; suture straight. Abdominal punctation sparse and very fine. First exposed sternite without microsculpture, following sternites with punctulate microsculpture. Meta- xocal line parallel to coxa, finely punctate; metacoxal area 0.03 mm long. Male sexual characters. Segments | to 3 of protibiae and segments | and 2 of mesotibiae moderately widened. Aedeagus (Figs 9 to 11) 0.69 mm long. Median lobe symmetrical. Basal bulb rather strongly sclerotized, relatively small. Ventral tubercles large, prominent. Distal process of median lobe about as long as basal bulb, gradually narrowed apically, with subapical denticle on dorsal side; tip acute in lateral view. Parameres with subbasal notch, expanded base, abruptly curved subapically in dorsal view. Inner side of parameres weakly sclerotized in apical half, sinuate in lateral view. Internal sac with long rod overlapped by rows of spines. Comments. This species possesses an aedeagus similar to that of S. simpli- cipenis Löbl, and is likely closely related to that species. However it may be easily distinguished from S. simplicipenis by the much large basal bulb, the less prominent ventral tubercles, and the presence of conspicuous, sclerotized spines in the internal sac. In external characters S. aciculare differs notably from S. simplicipenis by the sutural striae of elytra extended to form basal striae. 618 IVAN LOBL Scaphisoma latro sp. n. Figs 12, 13 Holotype d : China, Western Hubei, Shennonglia Nat. Res., 2000-2200m, 3-8.vi.1995, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 15,22 (MHNG). Description. Length 1.70-1.75 mm. Body dark reddish-brown to black, apical third of elytra yellowish, apex of abdomen ochreous. Femora and tibiae reddish- brown, lighter than thorax, antennae and tarsi yellowish. Length ratio of antennal segments 3 to 11 as follows: 6: 10: 16: 16: 22: 16: 20: 21: 25 (holotype); segment 4 about 3 times as long as wide; segment 5 about 4 times as long as wide; segment 6 about 3.5 times as long as wide; segment 7 about 3 times as long as wide; segment 8 about 2.5 times as long as wide; segment Il almost 3.5 times as long as wide. Pronotum evenly arcuate laterally; lateral keels concealed; discal punctation very fine and fairly sparse, diameters of punctures much smaller than puncture intervals. Tip of scutellum exposed. Elytra each with lateral margin arcuate in basal third, straight in posterior two thirds; apical margin slightly rounded; inner apical angle rounded, not prominent, at same level as outer apical angle; sutural margin not raised; adsutural area flat in anterior half, slightly raised in posterior half; sutural striae parallel in anterior half, converging toward apex, curved along base to form basal striae reaching outer third of basal width; discal punctation fine, fairly dense, less fine than that of pronotum but near elytral base about as fine as on pronotum. Prohypomera smooth. Mesepimeral ridge twice as long as interval to mesocoxa. Metasternum without microsculpture, with very fine and sparse punctation, flattened in middle, and with very shallow, medio-apical impressions; midline impunctate. Mesocoxal line arcuate, very finely punctate; mesocoxal area 0.06 mm long. Metepisterna flat, gradually narrowed anteriorly, lying at same plane as metasternum, with suture almost straight. Exposed abdominal segments with microsculpture consisting of transverse striae; abdominal punctation very fine and sparse, on mediobasal portion of first sternite more distinct than on remainder of sternal surface; microsculpture absent from latero- basal portion of first sternite. Metacoxal line strongly arcuate and very finely punctate; metacoxal area 0.10 mm long. Sexual characters of male. Segments | to 3 of protarsi widened. Aedeagus (Figs 12, 13) 0.59-0.6lmm long. Median lobe symmetrical. Basal bulb large, moderately sclerotized. Ventral tubercles large, somewhat prominent. Distal process of median lobe short, curved and inflexed, with tip wide in dorsal view, acute in lateral view. Parameres almost evenly narrow, slightly curved in dorsal view, slightly sinuate in lateral view. Internal sac with long membranous vesicle covered by squamose structures and with rows of strongly sclerotized, apical denticles. Comments. This species possesses the aedeagus similar to that of S. hiekei Löbl althought the latter species has the apical portion of the median lobe more strongly inflexed, and the parameres evenly narrow. Scaphisoma hiekei may be easily distinguished from S. latro by the elytra lacking basal striae and the antennal seg- ments 3 and 4 equally short. SCAPHIDIINAE OF CHINA, II 619 IS SSA leve 7-5 oe * rer CÈ x. Reel: 3 RISK ODE O io SAQA. Na, > DES ee a Ss SS 12 Fics 7 to 12 Aedeagi in Scaphisoma, dorsal and lateral views; 7 and 8, S. signum sp. n., scale bar = 0.1 mm: 9 to 11, S. aciculare sp. n.: 9 and 11, aedeagus, scale bar = 0.2 mm; 10, internal sac, scale bar = 0.1 mm; 12, S. latro sp. n., lateral view. 620 IVAN LÖBL Scaphisoma subtile sp. n. Figs 14, 15 Holotype d: China, South Sichuan, south Xichang, Lunji, 2300-2500m, 16-24.vii.1996, S. A. Kurbatov (MHNG). Description. Length 2.15 mm. Body blackish-brown, elytra lighter toward apex. Abdomen dark reddish-brown, with apex lighter. Femora blackish, tibiae reddish- brown, antennae and tarsi lighter than tibiae. Length ratio of antennal segments 3 to 11 as follows: 7: 12: 17: 20: 23: 19: 23: 22: 30; segments 4, 8 and 11 each about 3 times as long as wide; segment 5 about 3.5 times as long as wide; segment 6 about 4 times as long as wide; segment 7 about 2.5 times as long as wide. Pronotum with lateral margins evenly arcuate; lateral keels entirely visible; punctation dense and fairly fine, most puncture intervals distinctly larger than puncture diameters. Exposed point of scutellum minute. Elytra each with lateral margin strongly arcuate in basal half, slightly rounded in apical half; lateral keel exposed posterior middle area; apical margin truncate; inner apical angle rounded, not prominent, lying slightly posterior level of outer apical angles; sutural margin not raised; adsutural area flat in basal third, slightly raised in posterior basal third; sutural stria parallel to suture, curved near base and extended along basal margin to outer third of basal width, not joined to lateral stria; discal punctation fine and fairly sparse, punctures not well delimited, mostly as large as, or smaller than, pronotal punctures, much smaller than puncture intervals. Pygidium with distinct punctulate microsculpture; punctation fine near base, becoming very fine toward apex. Propygidium with punctation fairly coarse and dense. Prohypo- mera smooth, lacking microsculpture, with few extremely fine punctures. Mesepimeral ridge as long as interval to mesocoxa. Metasternum without microsculpture; medio- apical area flattened and with dense and fairly coarse punctation; medio-anterior and lateral areas of metasternum with very fine and sparse punctation. Mesocoxal line almost parallel to mesocoxa, finely punctate; mesocoxal area 0.04 mm long. Metepis- ternum flat, narrowed anteriorly, with almost straight suture, almost in same plane as metasternum. First exposed abdominal sternite lacking microsculpture; with punctation fairly coarse and dense on mediobasal area, very fine and sparse on medio-apical and lateral areas. Metacoxal line arcuate, finely punctate; metacoxal area 0.08 mm long. Following sternites with punctation very fine and sparse, and with distinct punctulate microsculpture. Male sexual characters. Segments | to 3 of protarsi slightly widened. Aedeagus (Figs 14, 15) 0.70 mm long. Median lobe symmetrical, with relatively small and weakly sclerotized basal bulb. Ventral tubercles large, prominent. Distal process of median lobe slender, longer than basal bulb, moderately inflexed, narrowed in apical portion; tip obtuse in dorsal view, slightly curved and acute in lateral view. Parameres almost evenly wide in apical two thirds, widened toward base, with margins sinuate in dorsal view, arcuate in lateral view. Internal sac with long, strongly sclerotized, almost straight rod; membranes lacking squamose or spinose structures. Comments. This species exhibits an aedeagus structuraly similar to that of S. punctaticolle Löbl, althought the ventral tubercles of the median lobe are much larger and prominent. Scaphisoma punctaticolle may be distinguished easily from S. subtile by the relatively coarsely punctate metasternum and the conspicuously microsculptured first abdominal sternite. SCAPHIDIINAE OF CHINA, II 621 Scaphisoma wolong sp. n. Figs 16, 17 Holotype d: China, Sichuan, Wolong Nat. Res. 1700m, 19.v.1994, S. A. Kurbatov (MHNG). Paratype 9: same data as holotype (MHNG). Description. Length 1.65 mm. Body uniformy very dark reddish-brown. Femora and tibiae slightly lighter than body. Antennae and tarsi ochreous or yellowish. Length ratio of antennal segments 3 to 11 as follows: 3: 6: 8: 11: 14: 10: 15: 14: 20 (holotype); segment 4 narrow, less than twice as long as wide, segment 5 wider than segment 4, almost 3 times as long as wide; segment 6 much wider than segment 5, about twice as long as wide; segments 7 and 11 each about 2.5 times as long as wide; segment 8 about twice as long as wide. Pronotum evenly arcuate laterally; lateral keels completely concealed; punctation very fine and dense, punctures relatively well delimited, much smaller than puncture intervals. Tip of scutellum exposed. Elytra each with lateral margin strongly arcuate anteriorly, oblique in apical portion; lateral keel not visible; apical margin distinctly rounded; inner apical angle rounded, not prominent, lying posterior to level of outer angle; sutural margin flat anteriorly, slightly raised in posterior half; adsutural area mostly flat, not or hardly oblique in posterior half; sutural striae parallel, curved along base and extended to humeral areas, forming basal striae not joined to lateral striae; discal punctation sparse and very fine on basal half, similar to that on pronotum, slightly less fine on apical half. Pygidium with punctation extremely fine and with punctulate microsculpture. Prohypomera smooth, impunctate. Mesepimeral ridge as long as interval to mesocoxa. Metasternum convex in middle, without medio-apical impressions and without microsculpture, with punctation very fine and sparse laterally and on center, dense and fairly coarse on medio-apical area. Mesocoxal line arcuate, rather coarsely punctate, mesocoxal area 0.06 mm long. Metepisternum flat, in same plane as metasternum, narrowed anteriorly, with straight suture. First abdominal sternite with punctation sparse and very fine laterally, dense and rather coarse medially. Metacoxal line arcuate, rather coarsely punctate; metacoxal area 0.06 mm long. Apical sternites with punctulate microsculpture. Male sexual characters. Protarsi not widened. Aedeagus (Figs 16, 17) 0.35 mm long. Median lobe fairly sclerotized, symmetrical. Basal bulb relatively small, with ventral tubercles distinct, not prominent. Distal process curved and moderately inflexed ventrally, gradually narrowed in dorsal view, with blunt tip. Dorsal valves membranous. Parameres slightly bent apically in dorsal view, slightly arcuate in lateral view. Internal sac bearing simple long flagellum; membranese very finely spinose. Flagellar base widened and bifid. Comments. This species is a member of the S. subalpinum group. It is similar to 5. inquietum Löbl, from which it differs in having the elytral and pronotal punc- tation more dense, the antennal segment 4 longer, and the internal sac bearing a simple rod, lacking denticulate and or quamose structures on membranes. The latter feature separates S. wolong from all other members of the S. subalpinum group. Scaphisoma styloides sp. n. Figs 18 to 20 Holotype d : China, West Hubei, Shennongjia Nat. Res., 2000-2200m, 3-8.vi.1995, S. A. Kurbatov (MHNG). Paratype 9 : sama data as holotype (MHNG). 622 IVAN LOBL Description. Length 1.95-2.30 mm. Body uniformly dark reddish-brown, apex of abdomen, tarsi and antennae lighter. Length ratio of antennal segments 3 to 11 as follows: 6: 10: 12: 15: 22: 16: 19: 18: 20 (holotype); segment 4 narrow, about 3 times as long as wide; segment 5 about 2.5 times as long as wide; segment 6 slightly more than 2 times as long as wide; segment 7 almost 3 times as long as wide; segment 8 and 11 each about 2 times as long as wide. Pronotum evenly arcuate laterally; lateral keels concealed; punctation very fine and sparse, puncture diameters much smaller than intervals. Tip of scutellum exposed. Elytra each with lateral margin arcuate in basal half, oblique in apical half; lateral keel not visible in dorsal view; apical margin truncate; inner apical angle rounded, not prominent, on same level as outer apical angle; sutural margin not raised; adsutural area flat; sutural stria parallel to suture, curved at base and extended to form basal stria joined to lateral stria; discal punc- tation fine, fairly dense, intervals between punctures mostly 2 to 4 times as large as puncture diameters. Pygidium very finely punctate, with punctulate microsculpture. Prohypomera smooth. Mesepisterna, lateral portions of metasternum, metepisterna and lateral portions of first exposed sternite with very fine and sparse punctation. Mesepimeral ridge about 1.5 times as long as interval to mesocoxa. Metasternum without microsculpture, convex in middle, with two medio-apical impressions densely punctate. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.06 mm long. Metepisterna flat, strongly narrowed anteriorly, in same plane as metasternum, with straight suture. Lateral portions of first exposed sternite and entire following sternites covered by punctulate microsculpture (conspicuous in holotype, hardly visible in paratype). Metacoxal line arcuate, finely punctate; metacoxal area 0.07 mm long. Sexual characters of male. Segments | to 3 of protarsi distinctly widened. Aedeagus (Figs 18 to 20) 0.55 mm long. Median lobe symmetrical, with basal bulb large, moderately sclerotized. Ventral tubercles small, slightly prominent. Distal pro- cess of median lobe much shorter than basal bulb, inflexed ventrally, gradually narrowed apically. Internal sac with flagellum simple, slender, gradually narrowed api- cally, slightly curved, and with large basal vesicular area covered by squamose structures. Parameres almost evenly wide in apical two third, straight in middle, curved in apical portion. Comments. This species is a membre of the S. subalpinum group. It is likely closely related to S. uniforme Löbl with which it shares most aedeagal characters. It differs from the latter species by the shape of the flagellum of the internal sac which is widened and hook-like in S. uniforme, the curved apices of parameres, and by the elytra with entire basal stria. Fics 13 to 18 Aedeagi in Scaphisoma, dorsal and lateral views; 13, S. latro sp. n., dorsal view, scale bar = 0.1 mm; 14 and 15, S. subtile sp. n., scale bar = 0.2 mm; 16 and 17, S. wolong sp. n., scale bar = 0.1 mm; 18, S. styloides sp. n., scale bar = 0.1 mm. 623 SCAPHIDIINAE OF CHINA, II 624 IVAN LOBL Scaphisoma apertum sp. n. Figs 21 to 23 Holotype 6: China, Yunnan, Jizu Shan, 2500-2700m, 25°58’ N 100°21’E, 6-10.vii. 1994, V. Kuban (MHNG). Paratypes: Yunnan, Gaoligong Mts, 2200-2500m, 25°57’N 98°45’E, 8-16.v.1995, O. Semela, 14,29 (NHMB, MHNG). Description. Length 2.1-2.2 mm. Body uniformly black, femora and tibiae dark reddish-brown, tarsi and antennae ochreous. Length ratio of antennal segments 3 to 11 as follows: 6: 14: 15: 21: 27: 23: 28: 26: 32 (holotype); segment 4 narrow, 3.5 times as long as wide; segments 5 to 8 each about 3 times as long as wide; segment 11 about 4 times as long as wide. Pronotum with lateral margins rounded in anterior half, oblique in basal half; lateral keels not visible; punctation dense and coarse, punctures not well delimited, mostly smaller than puncture intervals. Tip of scutellum exposed. Elytra each with lateral margin distinctly arcuate in basal third, slightly arcuate in apical two thirds; lateral keel visible near base and in apical third; apical margin arcuate; inner apical angle rounded, not prominent, on same level as outer apical angle; sutural margin not raised; adsutural area flat in basal third, slightly raised posterior basal third; sutural striae parallel, curved along base to form basal striae extended to outer third of basal width; discal punctation finer and less dense than pronotal punctation, puncture intervals mostly 2 to 4 times as large as puncture diameters. Pygidium and propygidium very finely punctate and with almost obsolete, punctulate microsculpture. Prohypomera smooth. Mesepimeral ridge slightly shorter than interval to mesocoxa. Metasternum without microsculpture, very finely and sparsely punctate, but with dense punctation on flattened medio-apical portion. Meso- coxal line subparallel, with coarse and very dense punctures; mesocoxal area 0.05 mm long. Mesepisterna flat, strongly narrowed anteriorly, lying below plan of meta- sternum; suture slightly sinuate. Abdominal sternites extremely finely punctate. First exposed sternite without microsculpture, following sternites with almost obsolete punctulate microsculpture. Metaxocal line arcuate, with dense and coarse punctures; metacoxal area 0.06 mm long. Sexual characters of male. Segments | - 3 of protarsi widened. Aedeagus (Figs 21 to 23) 0.72-0.73 mm long. Median lobe symmetrical, with basal bulb large, weakly sclerotized. Ventral tubercles well developed, slightly prominent. Distal process of median lobe shorter than basal bulb, inflexed, tapering. Dorsal valves membranous. Inner sac with flagellum long, strongly sclerotized, almost evenly thin, hook-like basally. Membranes with denticular and squamose structures, denticles partly arranged in rows. Comments. This is an additional member of the S. subalpinum group. It shares with S. castaneipenne Reitter, S. galloisi Achard and S. adnexum Löbl an aedeagus with long, thin flagellum, the internal sac with rows of denticular structures, and the parameres almost evenly narrow. Scaphisoma apertum may be distinguished from these species by the hook-like basis of the flagellum. In addition, it may be easily separated from S. castaneipenne and S. galloisi by the finer elytral punctation and finer abdominal microsculpture, and from S. adnexum by the larger size of the body and the more coarse pronotal punctation. SCAPHIDIINAE OF CHINA, II 625 The two examined males have the internal sac extruded. The flagellum is as illustrated, strongly bent in the holotype, stretched apically in the paratype. Scaphisoma pseudantennatum sp. n. Figs 24 to 26 Holotype ¢: China, Central Sichuan, Wolong Nat. Res., 1500m, 21.v.1994, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 46, 42 (MHNG); Sichuan, Liziping env. near Shimian, 200km south west Xa’an, 27.iv.-3.vii.1991, Z. Kejval, 19 (NHMB). Description. Length 2.6-2.7 mm. Body black, apex of abdomen very dark brown, tibiae reddish-brown, tarsi and antennae lighter than tibiae, ochreous or light brown. Length ratio of antennal segments 3 to 11 as follows: 8: 12: 15: 32: 35: 28: 30: 31: 38 (holotype); segments 4 and 5 each 3 times as long as wide; segment 6 conspi- cuously subparallel, wider than segment 5, almost 5 times as long as wide; segment 7 about 3.5 times as long as wide; segment 8 and 11 each about 4 times as long as wide (holotype). Thorax lacking microsculpture. Pronotum with almost evenly arcuate lateral margins, lateral keels not or barely visible; discal punctation distinct, punctures well delimited, puncture diameters smaller than puncture intervals. Tip of scutellum exposed. Elytra each with lateral margin arcuate anteriorly, oblique in middle portion; lateral keel partly visible in dorsal view; apical margin straight; inner apical angle rounded, not prominent, about on same level as outer apical angle; sutural margin not raised; adsutural area flat; sutural stria shallow, parallel to sutural margin, curved anteriorly and extended laterally along basal margin to humeral area, not joined with lateral stria; discal punctation even, similar to that on pronotum, as coarse as, or slightly coarser than, pronotal punctation. Mesepimeral ridge about as long as interval to mesocoxa. Metasternum flattened medio-apically; metasternal punctation very fine and very sparse on lateral and mediobasal areas, fairly dense and coarse on medio- apical area. Mesocoxal line arcuate, coarsely punctate; mesocoxal area 0.07-0.08 mm long. Metepisterna flat, strongly narrowed anteriorly, on same plane as metasternum; suture slightly sinuate. Exposed tergites with microsculpture consisting of punctures. Punctation on propygidium finer, that on pygidium much finer than elytral punctation. First exposed abdominal sternite lacking microsculpture; with punctation fairly dense and distinct on mediobasal area, very fine and very sparse on medio-apical and lateral surfaces. Following sternites with distinct punctulate microsculpture, normal punc- tation obsolete. Metacoxal line arcuate, coarsely puncate; metacoxal area 0.09-0.10 long. Male sexual characters. Segments | to 3 of protarsi distinctly widened. Aedea- gus (Figs 24 to 26) 1.15-1.20 mm long. Median lobe and parameres symmetrical, fairly strongly sclerotized. Basal bulb large. Ventral tubercles not prominent. Distal process of median lobe gradually inflexed and tapering; dorsal valves overlapping, weekly sclerotized. Internal sac with long flagellum curved basally and joined to short, hook-like sclerite. Membranes bearing complex squamose structures. Para- meres slightly narrowed apically and slightly curved in lateral view, sinuate or dis- tinctly curved and narrowed apically in dorsal view. 626 IVAN LOBL Comments. This new species is a member of the S. subalpinum group. It is similar to S. antennatum (Achard) with which it shares the antennal segment 6 long compared with segments 4 and 5. It may be distinguished readily from S. antennatum by the shorter antenal segment 11 and by the aedeagal characters. The aedeagus is smaller in S. antennatum, with the distal process of the median lobe shorter and not inflexed, the parameres slightly sinuate and the shape of the flagellum distinctive. Scaphisoma oppositum sp. n. Figs 27, 28 Holotype d: China, Yunnan prov., Gaoligongshan, 90 km west Baoshan, 26-26.v.1995, S. Beévat (MHNG). Paratype 2: same data as holotype (MHNG). Description. Length 2.55 mm. Very similar to S. pseudantennatum in most diagnostic characters. Antennae slightly distinctive, length ratio of antennal segments 3 to 11 as follows: 7: 10: 15: 31: 34: 28: 30: 30: 40 (holotype); segment 4 about 2.5 times as long as wide; segment 5 about 3 times as long as wide; segment 6 about 5 times as long as wide; segments 7 and 11 each 4 times as long as wide. Elytra with discal punctation finer and more sparse than that on pronotum or in S. pseudanten- natum. Microsculpture on propygidium and on lateral portions of first exposed abdominal sternite absent or hardly visible. Mesocoxal area 0.09 mm long. Metacoxal area 0.10 mm long. Male sexual characters. Segments | to 3 of protarsi widened. Aedeagus (Figs 27, 28) 1.10 mm long. Median lobe and parameres symmetrical, strongly sclerotized. Median lobe gradually narrowed apically, with basal bulb relatively small, shorter than distal process. Tip of median lobe acute and strongly curved. Ventral tubercles not prominent. Parameres arcuate, narrowed from base to middle third, almost evenly wide in posterior two thirds. Internal sac with flagellum sinuate and bifid proximally, and with additional, straight, subbasal rod. Basal portion of flagellum fairly thick, left flagellar rod narrow. Membranes of internal sac with squamose structures in middle. Comments. This species is an other member of the S. subalpinum group and is similar to S. antennatum and S. pseudantennatum. It differs from them notably by the more finely punctate elytra, the narrower aedeagus, and the very distinctive, bifid flagellum bearing an additional rod. Scaphisoma acclivum sp. n. Figs 29, 30 Holotype d: China, South Sichuan, Ya’an Prefecture, Shimian Co., Xiaoxiang Ling, lateral valley above Nanya Cun near Caluo, 11 km south Shimian, ca 1250m, 7.vii.1999, litter, bark, fungi, M. Schülke (ZMB). Paratypes: same data as holotype, 2 2 (MHNG, PCMS). Description. Length 2.6-2.9 mm. In external characters very similar to S. antennatum, S. pseudantennatum and S. oppositum. It differs by following characters. Fics 19 to 24 Aedeagi in Scaphisoma, dorsal and lateral view; 19 and 20, S. styloides sp. n.; 21 to 23, S. apertum sp. n., apical portion of median lobe with partly extruded internal sac; 24, S. pseud- antennatum sp. n., internal sac. Scale bars = 0.1 mm. 627 SCAPHIDIINAE OF CHINA, II as “CT EEZELS x WY = SID N (an min 3 x NE 628 IVAN LOBL Length ratio of antennal segments 3 to 11 as follows: 6: 10: 15: 28: 30: 24: 29: 25: 34 (holotype); segment 4 about 2.5 times as long as wide; segments 5 about 3 times as long as wide; segment 6 about 5 times as long as wide, parallel-sided; segment 7 almost 4 times as long as wide: segments 8 and 11 each 4 times as long as wide. Pronotal punctation dense and coarse, punctures well delimited, slightly smaller or as large as intervals. Elytral punctation less dense, near base finer, on center of disc slightly coarser than pronotal punctation. Metepisternum lying below plane of metasternum. Mesocoxal area 0.09 mm long, metacoxal area 0.10 mm long. Micro- sculpture on lateral portions of first exposed abdominal sternite distinct. Male sexual characters. Segments 1 to 3 of the protarsi and mesotarsi strongly widened. Aedeagus (Figs 29, 30) 1.55 mm. Median lobe strongly sclerotized, symme- trical, except for asymmetrical apex. Basal bulb fairly large, about as long as distal process, with ventral tubercles distinct, not prominent. Distal process moderately inflexed, rather oblique, asymmetrically narrowed toward apex; tip blunt in dorsal view; apex expanded ventrally and hook-like in lateral view. Parameres with large base, narrowed toward middle, with apical half almost equally wide and arcuate in dorsal view; sinuate in lateral view. Internal sac with flagellum robust, widened and curved in basal portion, almost straight in posterior basal portion, becoming gradually thiner toward apex. Proximal end of flagellum bearing small, oblique apophysis. Additional basal rod short, curved. Membranes very densely spinose in middle part of internal sac. Comments. This species may be readily distinguised from other members of the S. subalpinum group by the shape of the distal process of the median lobe, in particular by its hook-like apex. It is also characterized by the shape of the flagellum. The aedeagal characters of S. acclivum indicate close relationships to S. oppositum which possesses also a flagellum bearing a proximal apophysis and an additional basal rod. However, the shape of the median lobe, parameres and flagellum are distinct these two species. Scaphisoma linum sp. n. Figs 31432 Holotype d : China, Sichuan, Liziping env. near Shimien, 200 km southwest Ya’an, 27.vi.-3.vii.1991, Z. Kejval (MHNG). Paratypes: same data as holotype, 1 4,2 2 (MHNG); Sichuan, Daxue Shan, Gongga Shan, 102°04°E 29°36°N, Hailuogou Glacier Park, river valley about Ikm above Camp |, 2100m, 28-31.v.1997, A. Pütz, 3 d, 12 (MHNG, PCAP); same data but env. Camp II, 2650m, sifted, 30.v.1997, 1 & (MHNG): same data but above Camp III, 3000m, 1 4 (MHNG). Description. Length 2.15-2.30 mm. In external characters very similar to S. pseudantennatum, differs from the latter by: length ratio of antennal segments 3 to 11 as follows: 6: 10: 16: 18: 23: 17: 22: 21: 26 (holotype); antennal segment 4 about 3 times as long as wide; segments 5 and 6 each 4 times as long as wide; segments 7, 8 Fics 25 to 30 Aedeagi in Scaphisoma, dorsal and lateral views: 25 and 26, S. pseudantennatum sp. n., scale bar = 0.3 mm; 27 and 28, S. oppositum sp. n., scale bar = 0.3 mm: 29 and 30, S. acclivum sp. n., scale bar = 0.4 mm. 629 SCAPHIDIINAE OF CHINA, II 630 IVAN LOBL and 11 each about 3 times as long as wide; pronotum and elytra with almost even, fine punctation; inner apical angle of elytra lying posteriorly to outer apical angles; mesocoxal and metacoxal lines finely punctate; exposed abdominal tergites very finely punctate. Sexual characters of male. Segments | to 3 of protarsi widened. Aedeagus (Figs 31, 32) 0.65 - 0.70 mm long, symmetrical. Basal bulb moderately sclerotized, fairly large. Ventral tubercles large, prominent. Distal process of median lobe about as long as basal bulb, tapering, curved and inflexed ventrally. Parameres slightly curved, with large base, gradually narrowed toward middle, evenly wide in apical half (dorsal and ventral views). Internal sac with flagellum very long, almost evenly narrow, strongly curved basally and joined to long, membranous, basal vesicle. Squamose structures absent from internal sac. Comments. This species is also member of the S. subalpinum group. It may easily be distinguished by the distinctive shape of the flagellum. It has the ventral tubercles of the median lobe prominent, as in S. adjacens Löbl and S. serpens sp.n. Scaphisoma serpens sp. n. Figs 33, 34 Holotype d : China, Shaanxi, Qin Lin Shan, 10756’E, 33°45’N, route km 93 south of Zhouzhi, 108 km southwest of Xian, 1650m, mountain forest, sifted, 1-2.1x.1995, M. Schiilke (ZMB). Description. Length 1.9 mm. Body very dark reddish-brown, apex of abdomen lighter. Femora and tibiae ochreous. Tarsi and antennae light brown to yellowish. Length ratio of antennal segments 3 to 11 as follows: 5: 7: 13: 15: 20: 15: 19: 19: 23; segment 4 short, about 1.5 times as long as wide; segments 5 and 6 each about 2.5 times as long as wide; segments 7, 8 and 11 each about twice as long as wide. Pronotum with arcuate lateral margins; lateral keels concealed; punctation fine and dense, punctures not well delimited, very shallow, partly about as large as puncture intervals but mostly much smaller than latter. Tip of scutellum exposed. Elytra each with lateral margin slightly arcuate, lateral keel visible only near base; apical margin truncate; inner apical angle rounded, not prominent, lying posteriorly to outer angle; sutural margin not raised; sutural striae parallel, curved near base and extended laterally to form basal striae, approximate to base laterally and joined with lateral striae; discal punctation irregular, very fine to fairly coarse, dense; punctures usually larger than those on pronotum, smaller or about as large as puncture intervals. Exposed abdominal segments with punctulate microsculpture. Pygidium very finely and sparsely punctate, propygidium more coarsely punctate near base. Prohypomera impunctate, with punctulate microsculpture. Mesepimeral ridge about as long as interval to mesocoxa. Metasternum slightly convex in middle, impressed medio- apically; microsculpture absent, punctation very fine and sparse on most of surface, coarse and dense medio-apically. Mesocoxal line arcuate and finely punctate; mesocoxal area 0.05 mm long. Metepisternum flat, strongly narrowed anteriorly, with almost straight suture, almost on same level as metasternum. Exposed abdominal sternites extremely finely punctate. Metacoxal line arcuate, finely punctate; metacoxal area 0.07 mm long. SCAPHIDIINAE OF CHINA, II 63] Sexual characters of male. Segments | to 3 of protarsi widened. Aedeagus (Figs 33, 34) 0.75 mm long. Median lobe symmetrical, basal bulb large, weakly sclerotized. Ventral tubercles large, slightly produced. Distal process of median lobe gradually narrowed toward tip (dorsal view), inflexed obliquely and curved at apex: tip acute in lateral view. Parameres slender, slightly sinuate, curved in apical portion. Internal sac with flagellum wide, flat and almost straight in basal portion, bearing small but distinct, acute denticle on right side. Flagellum narrowed and strongly arcuate posteriorly to level of denticle and evenly narrow in apical half; membranes with fine squamose and, in apical portion, spinose structures. Comments. This species is an additional member of the S. subalpinum group. It has the aedeagus similar to that of S. adjacens Löbl to which it is obviously closely related. It may be distinguished from S. adjacens by the prohypomera and distinctly microsculptured first exposed abdominal sternite, and by the distinctive shape of the flagellum. ios) Scaphisoma parasolutum sp. n. Figs 35, 36 Holotype d: China, South Yunnan, Mengyang Nat. Res., ca 500m, 14.1x.1994, S. A. Kurbatov (MHNG). Description. Length 1.55 mm. Body dark reddish-brown, elytra slightly darker than pronotum. Ventral side of body and appendages ochreous. Length ratio of antennal segments 3 to 11 as follows: 10: 14: 23: 19: 25: 20: 25: 25: 26; segment 3 conspicuously elongate; segment 4 very narrow, about 5 times as long as wide; segment 5 slightly wider than segment 4, about 6 times as long as wide; segments 6, 7, 8 and 11 each about 5 times as long as wide. Pronotum with lateral margins almost oblique in basal half, convex in apical half; lateral keels visible; discal punctation dense and fine, punctures well delimited. Apical portion of scutellum exposed. Elytra each with lateral margin evenly arcuate; lateral keel entirely visible in dorsal view; apical margin rounded; inner apical angle rounded, not prominent, on same level as outer apical angle; sutural margin not raised; adsutural area flat; sutural striae parallel to suture in anterior half, slightly converging from middle to apex, reaching level of scutellum, not curved externaly; discal punctation dense and coarse, punctures mostly well delimited and smaller than intervals. Prohypomera smooth. Mesepisterna very finely punctate. Mesepimeral ridge 1.5 times as long as interval to mesocoxa. Meta- sternum flattened medio-apically; area between coxae with microsculpture consisting of transverse striae, lateral portions of metasternum without microsculpture; meta- sternal punctation sparse and very fine; antecoxal puncture row dense, fairly coarse. Mesocoxal line arcuate, coarsely punctate; mesocoxal area 0.04 mm long. Metepi- sterna slightly convex, strongly narrowed anteriorly, in same plane as metasternum, with arcuate suture. Abdominal tergites and sternites with microsculpture consisting of transverse striae; punctation sparse and very fine. Metacoxal line arcuate, coarsely punctate; metacoxal area 0.08 mm long. Sexual characters of male. Protarsi with segments | to 3 moderately widened. Aedeagus (Figs 35, 36) 0.88 mm long. Median lobe symmetrical. Basal bulb large, weakly sclerotized. Ventral tubercles small, somewhat produced. Distal process of 632 IVAN LOBL median lobe tapering, slightly inflexed. Parameres asymmetrical, in same plane as median lobe, each bearing oblique, membranous lobe. Internal sac with sclerotized rod gradually narrowed apically and membranes covered by squamose structures. Comments. This species is certainly closely related with S. solutum Löbl which shares the conspicuously elongate 3rd antennal segment and the structurally very similar aedeagus. Scaphisoma parasolutum may be distinguished from S. solutum by the presence of the metasternal antecoxal puncture row and by the internal sac of the aedeagus bearing a rod gradually narrowed. Scaphisoma pseudosolutum sp. n. Figs 37 to 39, 41 Holotype d: China, West Hubei, Shennongjia Nat. Res., 2000-2200m, 5.v1.1995, S. A. Kurbatov (MHNG) Description. Length 1.9 mm. Body very dark reddish-brown, apex of elytra, apical abdominal segments and legs lighter, antennae yellowish. Length ratio of antennal segments 3 to 11 as follows: 5: 12: 20: 17: 21: 15: 21: 20: 24; segments 4 and 5 about 4 times as long as wide; segments 6, 7, 8 and 11 each about 3 times as long as wide. Pronotum with lateral margins evenly arcuate; lateral keels visible in dorsal view; discal punctation dense and fine, consisting of well delimited punctures. Apex of scutellum exposed. Elytra each with lateral margin almost straight, rounded only near base and apex; apical margin truncate; inner apical angle on same level as outer apical angles, not prominent; adsutural area flat in basal fifth, raised between basal fifth and apex; sutural stria gradually, weakly diverging from apex to base, ending behind pronotal lobe, not curved externally; discal punctation dense and fairly fine, coarser than that on pronotum, consisting of punctures well delimited, smaller than puncture intervals. Prohypomera smooth. Mesepisterna extremely finely punctate, without microsculpture. Mesepimeral ridge almost twice as long as interval to mesocoxa. Metasternum with very fine microsculpture consisting of very short waves and striae. Medio-apical portion of metasternum impressed. Metasternal punc- tation fine and dense in middle, becoming very sparse laterally; antecoxal puncture row very dense, consisting of small punctures. Mesocoxal line subtriangular, dis- tinctly punctate; mesocoxal area 0.05 mm long. Metepisterna flat, strongly narrowed anteriorly, on same plane as metasternum; suture slightly sinuate. Abdominal tergites and sternites with distinct microsculpture consisting of transverse striae; punctation very fine and sparse. Metacoxal line strongly convex, finely punctate; metacoxal area 0.09 mm long. Male sexual characters. Protarsi with segments | and 2 strongly widened, segment 3 slightly widened. Mesotarsi with segment 1 strongly widened, segment 2 slightly widened. Aedeagus (Figs 37 to 39, 41) 0.98 mm long. Median lobe symme- trical, with basal bulb weakly sclerotized. Ventral tubercles small, not produced. Fics 31 to 36 Aedeagi in Scaphisoma, dorsal and lateral views; 31 and 32, S. linum sp. n., scale bar = 0.3 mm; 33 and 34, S. serpens sp. n., scale bar = 0.2 mm; 35 and 36, S. parasolutum sp. n., aedeagus in dorsal view, scale bar = 0.2 mm (35); internal sac, scale bar = 0.1 mm (36). 633 SCAPHIDIINAE OF CHINA, II ce CC eCECT (Sa 634 IVAN LOBL Distal process tapering and moderately inflexed, slightly sinuate near tip in lateral view. Dorsal valve simple, lamellar, weakly sclerotized. Parameres wide, hardly inflexed, bent apically, with large, subbasal, membranous lobe. Internal sac narrow in middle, widened apically, bulbous basally; bearing spinose and squamose structures in basal portion, squamose structures in middle and apical portions. Comments. This species appears to be allied to S. gentile Löbl, S. flexuosum Löbl, S. nigrum Löbl, S. solutum Löbl, and S. parasolutum sp. n. which share aedea- gal characters, in particular the shape of the median lobe and parameres. Scaphisoma pseudosolutum differs notably from S. gentile by the narrow apical portion of the parameres and much smaller squamose structures of the internal sac, from S. flexuosum by the wide middle portion of the internal sac, from S. nigrum and S. solutum by the parameral lobe situated subbasally, and from the latter species also by the internal sac lacking a sclerotized rod. These species may be easily distinguished from S. pseudosolutum by their smaller size (the body length in S. flexuosum is 1.35 mm, in S. nigrum and S. parasulutum 1.50 - 1.55 mm, and in S. solutum 1.3 mm). They may be readily separated also by the length ratio of the antennal segments, especially by the short segment 4 in S. nigrum, the relatively long segment 3 in S. solutum and S. parasolutum, and the segment 5 much longer than the segment 6 in S. flexuosum. Scaphisoma vexator sp. n. Figs 40, 42 Holotype d: China, Yunnan, Gaoligong Mts, 1500-2500m, 25°22’N, 98°49’E, 17- 24.v.1995, S. Beëvär (MHNG). Description. Length 2.0 mm. Most of body, and appendages ochreous. Pro- notum dark brown to black in mesal and basal portions. Elytra each with entire lateral portion, apex and base dark brown to black; subapical, transverse stripe lighter than remaining ochreous surface. Dark lateral area extended to elytral center, and dark basal area extended along suture to form triangle reaching to mid-length of suture. Mesosternum, metasternum and episterna brown. Length ratio of antennal segments 3 to 11 as follows: 4: 13: 22: 24: 26: 21: 24: 23: 26; segments 4 to 8 and 11 each about 4 times as long as wide. Pronotum with lateral margins arcuate; lateral keels entirely distinct; punctation fine and dense, well delimited and visible on dark areas, rather indistinct on ochreous areas. Elytra each with lateral margin arcuate; lateral keel entirely visible; apical margin rounded; inner apical angle lying posteriorly to level of outer apical angle, not prominent; sutural margin not raised; adsutural area flat; sutural stria parallel in anterior half, converging apically, curved near pronotal lobe, not extended along basal margin; discal punctation fairly fine and dense, punctures not well delimited, mostly larger than those on pronotum, and mostly distinctly Fics 37 to 42 Aedeagi in Scaphisoma; 37 to 39 and 41, S. pseudosolutum sp. n., dorsal and lateral views, scale bar = 0.2 mm (37, 38), paramere in ventral view, scale bar = 0.2 mm (39), internal sac in dorsal view, scale bar = 0.1 (41); 40 and 42, S. vexator sp. n., aedeagus in dorsal view, scale bar = 0.3 mm (40), paramere in ventral view, scale bar = 0.2 mm (42). 635 SCAPHIDIINAE OF CHINA, II < ID) N À 41 Maren FE ONTO TE TEN E Er > aie fee e CAD = <> AS SEI (PRAIA INI r SC UT = x Lee 6 ites DI EEE RG Sc ZA N Gao ce € rs BELG Gilt ar HE); “€ ERICH DS 4 636 IVAN LOBL smaller than puncture intervals but some punctures about as large as puncture intervals. Prohypomera smooth. Mesepimeral ridge longer than interval to mesocoxa. Metasternum slightly convex in middle, lacking impressions; with microsculpture consisting of transverse striae becoming obsolete laterally, punctation sparse and very fine; antecoxal puncture row present but relatively inconspicuous. Mesocoxal line arcuate, very finely punctate; mesocoxal area 0.08 mm long. Metepisterna flat, moderately narrowed anteriorly, lying on lower plane than metasternum; suture straight. Abdomen very finely and sparsely punctate, with microsculpture consisting of tranverse striae. Metacoxal line arcuate, finely punctate; metacoxal area 0.10 mm long. Sexual characters of male. Segments 1 and 2 of protarsi and segment | of mesotarsi strongly widened, about as wide as apex of tibiae. Segment 3 of protarsi and segments 2 and 3 of mesotarsi widened, narrower than preceding segment. Aedeagus (Figs 40, 42) 1.12 mm long. Median lobe asymmetrical. Basal bulb large, weakly sclerotized. Ventral tubercles small, not prominent. Distal process of median lobe consisting of strongly sclerotized dorsal and ventral arms. Dorsal arm partly overlapped by basal bulb, curved, tapering and inflexed. Ventral arm longer than dorsal arm, straight, parallel-sided in anterior middle, tapering in posterior middle. Parameres wide, curved, with weakly sclerotized inner lobes. Internal sac complex, with two large, strongly sclerotized teeth, one narrow, long rod, and bearing squa- mose and denticulate membranous structures. Comments. This species is closely related with S. velox Löbl. The aedeagi in both species are very similar (the length of the aedeagus of S. velor is not 0.55 mm as given by lapsus in the description, but 1.10 mm). Scaphisoma velox has the dorsal arm of the median lobe inconspicuous, the teeth-like sclerites of the internal sac longer, and the internal sac bearing an apical groups of denticles and a long basal rod which is absent from S. vexator. These two species may be easily distinguished by their colour pattern (the body is almost uniformly ochreous in S. velox). Scaphisoma pseudovarium sp. n. Figs 43 to 46 Holotype d: China, Yunnan, Gaoligong Mts, 2200-2500m, 24°57’N, 98°45’E, 8- 16.v.1995, S. Beévai (MHNG). Paratypes: same data as holotype, 3 4,5 2 (MHNG). Description. Length 1.9-2.0 mm. Body with distinetive although variable colour pattern. Head and pronotum reddish-brown to blackish, lateral portions of pronotum sometimes lighter than pronotal center. Elytra ochreous or yellowish, with entire base and small subapical spot dark brown to black. Dark basal zone expanded apically along suture to cover at least anterior half of adsutural area, sometimes expan- ded to form large, tringular zone joined to subapical spots. Prohypomera ochreous. Mesosternum, metasternum, episterna and metepimera, and basal two or three exposed abdominal sternites dark brown to black. Following sternites and exposed abdominal tergites ochreous to yellowish. Appendages ochreous. Length ratio of antennal seg- ments 3 to 11 as follows: 5: 15: 20: 20: 21: 19: 21: 20: 25 (holotype); segment 4 about 6 times as long as wide; segments 5 and 6 each about 7 times as long as wide; segment SCAPHIDIINAE OF CHINA, II 637 7 almost 4 times as long as wide; segments 8 and 11 each almost 5 times as long as wide. Pronotum with evenly arcuate lateral margins; lateral keels visible near base; punctation dense and fine or very fine, less fine near base than at center, punctures not well delimited, usually smaller than puncture intervals. Apex of scutellum exposed. Elytra each with lateral margin arcuate in basal half, oblique in apical half; lateral keel entirely visible in dorsal view; apical margin truncate; inner apical angle lying posteriorly to level of outer apical angle, not prominent; sutural margin not raised; adsutural area flat; sutural stria converging to apex, parallel to suture in anterior half, curved along basal lobe of pronotum, not extended laterally along basal margin; discal punctation dense and coarse, puncture intervals mostly about as large as, or up to twice as large as puncture diameters. Propypomera smooth. Mesepimeral ridge about 1.5 times as long as interval to mesocoxa. Metasternum lacking microsculpture, flattened in middle, without medio-apical impressions; punctation fine, dense on center, sparse laterally; antecoxal puncture row indicated by irregular row of punc- tures slightly larger than other metasternal punctures. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.05-0.06 mm long. Metepisterna flat, moderately narrowed anteriorly, lying below plane of metasternum, with suture rounded near angles, straight in middle. Exposed abdominal segments very finely punctate, with micro- sculpture consisting of transverse striae. Metacoxal line strongly arcuate, finely punctate, metacoxal area 0.11-0.13 mm long. Sexual characters of male. Segments 1 to 3 of protarsi and 1 and 2 of mesotarsi distinctly widened. Aedeagus (Figs 43 to 46) 1.02-1.04 mm long. Median lobe symme- trical. Basal bulb weakly sclerotized, slightly expanded apically, overlapping base of distal process. Ventral tubercles prominent. Distal process of median lobe inflexed, with ventral arm tapering and acute at apex, denticulate on dorsal side, bearing single denticle in middle of ventral side. Dorsal process partly fissured to forme two joined rods strongly sclerotized along fisure. Parameres symmetrical, sinuate, with wide base, lobed and membranous at middle. Internal sac with two central tooth-like sclerites and membranes bearing squamose and spinose structures. Comments. This species is very similar to S. varium Löbl with which it shares the colour pattern and most of the diagnostic characters. It may easily be distin- guished from S. varium by the ventrally denticulate distal process of the median lobe, the irregular inner lobes of the parameres and the internal sac bearing two fairly long, central, sclerotized teeth. The aedeagus of S. pseudovarium is structurally very similar to that of S. sasagoense Löbl and S. pinnigerum Löbl. Scaphisoma sasagoense has the parameres narrowed subapically, and distinctly widened in apical portion, lacking a membranous lobe, and the internal sac with clusters of strongly sclerotized basal and apical denticles. In addition, S. sasagoense may be distinguished by the uniformly reddish-brown body. In S. pinnigerum, the metasternal antecoxal puncture row is distinct, the parameral lobe lies more apically, and the internal sac of the aedeagus bears a proximal striate structure which is absent from S. pseudovarium and S. sasagoense. 638 IVAN LÖBL Scaphisoma heishuiense sp. n. Figs 47 to 50 Holotype d : China, Yunnan, Heishui, 35km north Lijiang, 27°13’N, 100°19’E, 18.vi.- 4.v11.1993, S. Beévaf (MHNG). Description. Length 1.9 mm. In external characters very similar to S. pseudo- varium, including the colour pattern. Body ochreous, with pronotum dark brown except near lateral margins and on anterior third, elytra with dark brown basal area expanded along suture to form not well delimited triangular spot, and with slightly darkened subapical spot, the latter not reaching to sutural stria. Mesosternum and metasternum with episterna and epimera dark brown. Appendages ochreous. Antennae almost as in S. pseudovarium; length ratio of antennal segments 3 to 11 as follows: 5: 12: 19: 17: 20: 16: 20: 19: 24. Pronotal punctation more sparse, with punctures mostly much smaller than puncture intervals. Metasternum with antecoxal puncture row obsolete. Mesocoxal areea 0.05 mm long. Metepisterna slightly narrowed anteriorly, with straight suture. Metacoxal area 0.11 mm long. Punctation on mediobasal area of first exposed sternite relatively coarse. Aedeagus (Figs 47 to 50) 0.90 mm long. Ventro-apical process of median lobe narrower than in S. pseudovarium, sinuate, more densely denticulate on dorsal surface, lacking ventral denticle. Parameres barely sinuate, bent apically, with inner lobe narrow. Spinose structures of internal sac long and strongly sclerotized, teeth-like sclerites larger than those in S. pseudovarium. Scaphisoma paravarium sp. n. Figs 51 to 55 Holotype d: China, Yunnan, Gaoligong Mts, 2200-2500m, 24°57’N, 98°45’E, 8-16.v.1995, S. Beévat (MHNG). Paratypes: same data as holotype, 2 2 (NHMB, MHNG) Description. Length 1.75-1.95 mm. In external characters very similar to S. pseudovarıum, with almost the same colour pattern. Antennae similar to those in preceding species; length ratio of antennal segments 3 to 11 as follows: 5: 12: 18: 18: 21: 18: 20: 19: 24 (holotype). Pronotum broadly lighter laterally and anteriorly; elytra with dark apical spot not reaching to sutural stria. Lateral keels of pronotum concealed in dorsal view, those of elytra entirely visible or concealed in middle. Elytra each with sutural margin raised; punctation very fine near base, consisting of fairly large but very shallow and not clearly delimited punctures on posterior two thirds of disc. Sexual characters of male. Protarsi with segments | and 2 strongly widened, segment 3 slightly widened. Mesotarsi with segments | and 2 slightly widened. Aede- agus (Figs. 51 to 55) 0.90 mm long. Median lobe symmetrical, with basal bulb weakly sclerotized, slightly expanded apically, lacking ventral tubercles. Distal process of Fics 43 to 50 Aedeagi in Scaphisoma; 43 to 46, S. pseudovarium sp. n., aedeagus in dorsal and lateral views, scale bar = 0.2 mm (43, 44), apical portion of median lobe, in lateral view, scale bar = 0.1 mm (45), paramere in ventral view, scale bar = 0.2 mm (46); 47 to 50, S. heishuense sp. n.. aedeagus in dorsal and lateral views, scale bar = 0.2 mm (47, 48), paramere in ventral view. scale bar = 0.2 mm (49), apical portion of median lobe, in lateral view, scale bar = 0.1 mm (50). 639 SCAPHIDIINAE OF CHINA, II 640 IVAN LÖBL median lobe almost in same plane as basal bulb, tapering, denticulate dorso-apically. Dorsal valves apparently joined medianly, strongly sclerotized along inner margin. Parameres sinuate, almost evenly narrow, enlarged base excepted, with short, narrow, mebranous lobe in posterior half. Internal sac lacking rods or robust teeth-like sclerites. Membranes of basal half of internal sac bearing denticulate structures over- lapped laterally by squamose structures forming oblique, irregular rows. Denticular structures in apical portion of internal sac variable in size, more distinct than those in basal portion. Comments. This species may be distinguished from the very similar S. varium, S. pseudovarium and S. heishuiense by the finer elytral punctation. It is well charac- terized by the shape of the parameres and the structures of the internal sac. Scaphisoma fibrosum sp. n. Figs 56, 57 Holotype d : China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont (MHNG). Description. Length 1.7 mm. Head, thorax and basal two thirds of first exposed abdominal sternite dark brown. Elytra ochreous, with base dark brown to black; basal dark area expanded along suture to form triangular spot. Appendages, elytral apices and most of abdomen light ochreous. Antennae similar to those of pre- ceding species; length ratio of antennal segments 3 to 11 as follows: 5: 12: 20: 19: 23: 19: 24: 23: 28. Pronotum with lateral margins oblique in dorsal view; lateral keels exposed; punctation fine and dense near base, becoming very fine and more sparse anteriorly; punctures not well delimited, slightly smaller than puncture interval near base. Apex of scutellum exposed. Elytra each with lateral margins arcuate anteriorly and posteriorly, oblique in middle; lateral keel distinct in dorsal view; apical margin slightly concave, dentate at and near inner angle; inner apical angle prominent, lying posterior to level of outer angle; sutural margin not raised; adsutural area flat; sutural stria straight, not curved near base, gradually, very slightly converging to suture apically; punctation coarse and dense, punctures not well delimited, partly about as large as puncture intervals. Prohypomera smooth. Mesepimeral ridge slightly longer than interval to mesocoxa. Metasternum slightly convex in middle, flattened medio- apically, covered by microsculpture consisting of transverse striae; metasternal punctation very fine and sparse, but dense and coarse on medio-apical area; antecoxal puncture row distinct, lying in impressed line. Mesocoxal line arcuate, coarsely punctate, mesocoxal area 0.04 mm long. Metepisterna convex, moderately narrowed anteriorly, lying below plane of metasternum; suture slightly arcuate anteriorly and posteriorly, straight in middle. Abdominal segments with microsculpture consisting of tranverse striae but punctulate on laterobasal area of first sternite. Abdominal punc- Fics 51 to 57 Aedeagi in Scaphisoma; 51 to 55, S. paravarium sp. n., aedeagus in dorsal and lateral view, scale bar = 0.2 mm (51, 52), paramere in ventral view, scale bar = 0.1 mm (53), internal sac, dorsal view, scale bar = 0.1 mm (54), apical portion of median lobe, in lateral view, scale bar = 0.1 mm (55): 56 and 57, S. fibrosum sp. n., aedeagus in dorsal view, scale bar = 0,2 mm (56), paramere in ventral view, scale bar = 0.1 mm (57). SCAPHIDIINAE OF CHINA, II 64] fe Y EN < \ DS c x T \ AD: È A al AL Ten A 4 mG Ran et Un nm RNC / DEN \ n vor | 55 53 54 642 IVAN LOBL tation extremely fine. Metacoxal line arcuate, coarsely punctate; metacoxal area 0.05 mm long. Sexual characters of male. Segments | and 2 of protarsi strongly widened. Segment 3 of protarsi and segments | and 2 of mesotarsi slightly widened. Aedeagus (Figs 56, 57) 0.78 mm long. Median lobe symmetrical. Basal bulb weakly sclerotized, expanded over proximal part of distal process of median lobe, lacking ventral tubercles. Distal process inflexed, gradually narrowed, with tip truncate in dorsal view. Dorsal valves incompletely separated by fissure. Parameres wide, moderately curved, with outer margin almost straight up to apical portion, with wide, inner, membranous lobe; parameral basis strongly enlarged. Internal sac with membranes bearing very small squamose or denticulate structures, lacking sclerotized rods or teeth. Comments. This species is obviously closely related to S. varium, S. pinni- gerum, S. pseudovarium, S. heishuiense, and S. paravarium with which it shares the structuraly similar aedeagus. Scaphisoma fibrosum may be easily separated from these species by the coarse elytral punctation, the antecoxal puncture row lying in an impressed line, the broad parameres, and the absence from sclerotized structures in the internal sac. Scaphisoma incisum sp. n. Figs 58 to 61 Holotype d: China, Yunnan, Lijiang, 1800m, 26°53’ N, 100°18’ E, 21.vii.1992, S. Beévat (MHNG). Paratypes: same data as holotype, 4 2 (MHNG). Description. Length 1.70-1.75 mm. Head and most of pronotum very dark, almost blackish, with or without reddish lustre. Elytra as pronotum, or with large portion of disc somewhat lighter and reddish; elytral apices ochreous. Femora and tibiae ochreous, tarsi and abdominal apex yellowish. Length ratio of antennal segments 3 to 11 as follows: 6: 10: 15: 15: 17: 13: 17: 15: 19 (holotype). Pronotum with lateral margins evenly arcuate; lateral keels visible; punctation very fine, sparse, most punctures very shallow and not well delimited, but punctures near base appear slightly deeper and better delimited. Apex of scutellum exposed. Elytra each with lateral margin almost evenly arcuate; lateral keel visible in dorsal view; apical margin truncate, inner apical angle posterior to level of outer angle, not prominent; sutural stria slightly diverging from apex to mid-length of elytron, parallel from mid-length to level of pronotal lobe, slightly curved and ending near pronotal lobe; discal punctation fine and rather sparse, consisting of well delimited punctures, much larger than those on pronotum, puncture intervals mostly 3 to 5 times as large as puncture diameters. Pygidium conspicuously notched medio-apically, very finely punctate, with micro- sculpture consisting of transverse striae. Prohypomera smooth. Mesepimeral ridge about 1.5 times as long as interval to mesocoxa. Metasternum slightly convex in middle, lacking impressions, without microsculpture. Metasternal punctation very fine and sparse on lateral and on medio-anterior portions, more dense and less fine medio- apically and on areas between mesocoxae and metacoxae; antecoxal puncture row barely indicated. Mesocoxal lines arcuate, finely punctate, mesocoxal area 0.05 mm long. Metepisternum flat, lying below plane of metasternum, moderately narrowed SCAPHIDIINAE OF CHINA, II 643 anteriorly, with almost straight suture. Abdominal sternites with microsculpture consisting of transverse striae; punctation very fine and sparse. Metacoxal line arcuate, fairly coarsely punctate; metacoxal area 0.08 mm long. Tibiae straight. Male sexual characters. Segment | of protarsi strongly widened, about as wide as apex of tibia. Segment 2 of protarsi slightly widened, segment 3 of protarsi and segments | and 2 of mesotarsi barely widened. Aedeagus (Figs 58 to 61) 0.82 mm long. Median lobe almost as in S. pseudovarium, but with distal process oblique, not sinuate ventrally, bearing numerous dorsal denticles and single ventral denticle, as in the latter species. Parameres less curved apically, with inner lobe very narrow, incon- spicuous. Internal sac with two proximal, rather large, teeth-like sclerites, lacking any particular structures in middle, denticulate in apical portion. Comments. This species has the aedeagus similar to that of S. varium and the three species described above. It is obviously closely related to these four species but may be readily distinguished by the colour pattern and incised pygidium. The presence of a pair of teeth-like sclerites in the internal sac may indicate relationships to S. pseudovarium and S. heishuiense. Scaphisoma migrator sp. n. Figs 62 to 65 Holotype d: China, Shaanxi, east of Xian, Mt. Huashan, 500m, 12.v.1994, S. A. Kurbatov (MHNG). Paratypes: same data as holotype, 2 d (MHNG); Sichuan, Mt. Emei, 1700m, 22.1x.1994, S. A. Kurbatov, 3 d and with sama data but 1500, 21.ix., 1 & (MHNG); Sichuan, Wolong Nat. Res., 24.v.1994, S. A. Kurbatov, 1 3 and with same data but 1700m, 19.v., 2 G (MHNG); Shaanxi, On Ling Shan [=An Ling Shan], 110°06°E 34°27’N, Hua Shan, 118km east xian, N valley, 1200-1400m, 12-20.viii.1995, D. W. Wrase, 16 (PCMS); Shaanxi, On Lin Shan, 107°56’E 32°45’N, autoroute 93km south Zhouzhi, 108km southwest Xian, 1650m, mountaine forest, 2.ix.1995, D. W. Wrase, 1 d (PCMS); Shaanxi, Taibai Shan above Houshenzi, 130-1700m, 9.vi.-3.vii.1998, P. Jäger & J. Martens, 1 d (SMNS). Description. This species is very similar to, and possesses most diagnostic features of S. morosum Löbl, S. khao Löbl, S. karen Lobl, S. prehensor Champion, and S. tortile Löbl. The colour pattern of the elytra is fairly variable, the apical fifth to third of the disc is light. The metasternal antecoxal puncture row is irregular and usually less distinct than in similar species. This new species is characterized by the male genitalia. Aedeagus (Figs 62 to 65) 0.76-0.87 mm long. Median lobe symmetrical, fairly strongly sclerotized. Basal bulb very large, partly overlapping basal bulb. Ventral tubercles small, not prominent. Distal process of median lobe inflexed and arcuate, fairly long in lateral view, appearing short and straight in dorsal view, with small subapical denticle on dorsal side and tip acute. Dorsal valves completely separated, short, parallel. Internal sac complex, with large, sclerotized, apicolateral denticles joined to long, straight, tubular portion consisting of spinose structures. Parameres gradually narrowed from base to subapical portion, widened and slightly curved in apical portion, with small, inner, membranous lobe. Comments. This species appears closely allied to S. morosum with which it shares structural characters of the aedeagus, in particular the presence of a small, 644 IVAN LOBL subapical parameral lobe, and similar internal sac. Scaphisoma migrator may be distinguished from S. morosum by the teeth-like apical sclerites joined by complex membranous structures, the absence of dorso-apical plate of the internal sac, and wider apical portion of the parameres. In the absence of non sexual specific characters I have tentatively associated 14 females with the males listed above. The females were obviously collected together with the males. Scaphisoma mutator sp. n. Figs 66 to 69 Holotype d: China, Sichuan, Wolong Nat. Res., 1500m, 21.v.1994, S. A. Kurbatov (MHNG). Paratypes: China, Sichuan, Wolong Nat. Res., 1000m, 24.v.1994, litter, S. A. Kurbatov, 2 & (MHNG), same data but ca 500m, 15-16.v.94, 1 4 (MHNG); Mt. Emei, 1500m, 21.ix. 1994, litter, S. A. Kurbatov, 1 d (MHNG); same data but 1700m, 22.ix., 4 6 (MHNG); same data but 1800m, 1¢ (MHNG); Sichuan, Quingchen-Shan, 650-700m, 30°53°56’’N, 103°33’01”’E, 18.v.1997, M. Schülke, 2 4 (PCMS); same but Qiencheng Shan NW Chengdu, 30°54’N, 103°33’E, D. Wrase, 3 & (PCMS, MHNG); same but 65km NW Chengdu, 8km W Taiping, 800-1000m, A. Pütz, 3 d (PCAP, MHNG); Shaanxi, Qin Lin Shan 110°06’E, 34°27’N, N Hua Shan 118km east Xian, N valley, 1200-1400m, leafy wd., 18-20.viii.1995, D. Wrase, 1 & (PCMS); same but sifted, M. Schülke, 4 & (PCMS, MHNG); Shaanxi, east Xian, Huashan, 700m, 9-12.v.1994, S. A. Kurbatov, 17 & (MHNG). Description. Length 1.70-1.85 mm. Body dark reddish-brown, in some spe- cimens almost blackish, apices of elytra usually distinctly lighter; tarsi, antennae and apex of abdomen light brown to yellowish. In external charaters very similar to S. tortile Löbl but elytra with lateral keels usually concealed in dorsal view and sutural striae slightly diverging up to base; medio-apical portion of metasternum slightly impressed. This new species may be distinguished by the aedeagal characters. Aedeagus (Figs 66 to 69) 0.86 - 0.95 mm long. Median lobe fairly strongly sclerotized, symmetrical. Basal bulb very large. Ventral tubercles small, not pro- minent. Distal process of median lobe short, inflexed ventrally and oblique, very narrow in lateral view, with single, fairly large apicodorsal denticle. Dorsal valves short, completely separated, gradually narrowed apically. Internal sac complex, with long, central row of oblique or transverse teeth-like sclerites pointed ventrally, short subapical denticles, and two groups of apical denticles pointed toward apex. Para- meres sinuate, wide in middle, with very narrow inner lobe, slightly widened in apical portion (dorsal view), gradually narrowed from middle toward apex in lateral view. Comments. This species is a member of the S. haemorrhoidalis group and appears to be closely related to S. tortile Löbl and S. prehensor Champion. It shares with these two species, in addition to an overall similarity, the internal sac of the Fics 58 to 64 Aedeagi in Scaphisoma; 58 to 61, S. incisum sp. n., aedeagus in dorsal and lateral views, scale bar 0.2 mm (58, 59), apical portion of median lobe, in lateral view, scale bar = 0.1 mm (60), paramere in ventral view, scale bar = 0.1 mm (61); 62 to 64, S. migrator sp. n., aedeagus in dorsal view, scale bar = 0.2 mm (62); internal sac, scale bar = 0.1 mm (63), paramere in ventral view, scale bar = 0.1 mm (64). 645 SCAPHIDIINAE OF CHINA, II Cr 7. à Ay ren Cd GAIA za gS \ 646 IVAN LOBL aedeagus bearing a long row of teeth-like sclerites, and the apical denticles pointed toward apex. Scaphisoma mutator may be distinguished from S. tortile and S. pre- hensor by the absence of a large sclerotized plate or tooth in central part of the internal sac, by the apical denticles grouped to form two clusters, and the parameres sinuate and widened apically. An additional 32 females from Huashan are likely conspecific but are not included in the type series because this species can be recognized only by the male characters. Scaphisoma suspiciosum sp. n. Figs 70 to 73 Holotype ¢: China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont (MHNG). Description. Length 1.7 mm. Body, femora and tibiae light reddish-brown, apical third of elytra and apical abdominal segments lighter, ochreous. Antennae and tarsi yellowish. Pronotum with slightly arcuate lateral margins; lateral keels exposed; discal punctation sparse and very fine, punctures particularly shallow and not clearly delimited. Tip of scutellum exposed. Elytra each with lateral margin rounded near base, straight in posterior two thirds; apical margin truncate, inner apical angle lying posterior to level of outer apical angle, not prominent; sutural margin not raised, adsutural area flat; sutural stria slightly diverging from apex to mid-length, parallel to suture in anterior half, slightly curved near base; punctation dense and fairly fine, puncture intervals mostly as large as to 3 times larger than puncture diameters. Exposed abdominal tergites extremely finely punctate. Prohypomera smooth. Mesepi- meral ridge longer than interval to mesocoxa. Metasternum without microsculpture and with very fine and sparse punctation on lateral portions, lacking antecoxal punc- ture row. Middle portion of metasternum flattened, with shallow impression in center, fine and dense punctation, and very finely microsculptured. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.08 mm long. Metepisterna flat, moderately narrowed anteriorly, lying below plane of metasternum, with slightly sinuate suture. Abdomen with microsculpture consisting of transverse striae; sternites very finely punctate. Metacoxal line arcuate, fairly coarsely punctate; metacoxal area 0.07 mm long. Sexual characters of male. Segments | and 2 of protarsi and 1 of mesotarsi strongly widened, about as wide as tibia, segment 2 of mesotarsi moderately widened. Aedeagus (Figs 70 to 73) 0.93 mm long. Median lobe symmetrical, fairly strongly sclerotized. Basal bulb large, in dorsal view completely overlapping distal process and dorsal valves. Ventral tubercle robust, not prominent. Distal process strongly inflexed, almost perpendicular to axis of median lobe, gradually narrowed, with minute sub- apical denticle (tip broken off). Dorsal valves very short, completely separated, slightly curved. Parameres sinuate, with wide, relatively short lobe in posterior basal third, equally wide posterior lobe in dorsal view, gradually narrowed from lobe to apex in lateral view. Internal sac complex, with very dense spinose structures in basal half, followed by oblique, widened, central portion bearing large and strongly sclero- tized teeth-like sclerites and apical portion consisting of small teeth and squamose structures. SCAPHIDIINAE OF CHINA, II 647 N S IN N RISK TION UN TE ICH 70 68 Fics 65 to 70 Aedeagi in Scaphisoma; 65, S. migrator sp. n., lateral view, scale bar = 0.2 mm; 66 to 69 S. mutator sp. n., aedeagus in dorsal and lateral views, scale bar = 0.2 mm (66, 67), paramere in ventral view, scale bar = 0.1 mm (68), internal sac in dorsal view, scale bar 0.1 mm (69); 70, S. suspiciosum sp. n., parameres in ventral view, scale bar = 0.2 mm. 648 IVAN LOBL Comments. This species is a member of the S. haemorrhoidale group. It appears to beclosely related to S. insulanum Löbl, which possesses a long median lobe, with very short dorsal valves, similar shape of parameres und structurally simi- lar internal sac. Both species differ notably in the shape of the distal process of the median lobe which in S. suspiciosum is much more inflexed and overlapped by the basal bulb in dorsal view. The holotype has the segments 3 - 11 of both antennae broken off. Scaphisoma irruptum sp. n. Figs 74 to 76, 81 Holotype d : China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont (MHNG). Description. Length 1.6 mm. Body and femora dark reddish-brown, apical third of elytra and apex of abdomen light ochreous. Tibiae and tarsi ochreous. Pronotum with lateral margins evenly arcuate; lateral keels visible in dorsal view; discal punctation very fine and sparse, consisting of minute and not well delimited punctures. Tip of scutellum exposed. Elytra each with lateral margin slightly arcuate near base and in apical third, oblique in middle; lateral keel entirely exposed; apical margin slightly rounded, inner apical angle posterior to level outer apical angle, not prominent; sutural margin not raised, adsutural area flat; sutural striae diverging from middle to apex, parallel to anterior middle, curved along pronotal lobe, not extended laterally; punctation fine and dense in apical half, puncture intervals mostly as large as, to 3 times larger than, puncture diameters in apical half, 3 to 5 times larger than puncture diameters near base. Prohypomera smooth. Mesepimeral ridge longer than interval to mesocoxa. Middle and lateroposterior portions of metasternum with micro- sculpture consisting of transverse striae, most of lateral portions of metasternum without microsculpture. Metasternum flattened in middle, lacking impression, with very fine and dense punctation. Lateral portions of metasternum with punctation sparse and very fine; antecoxal puncture row distinct. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.08 mm long. Metepisterna flat, narrowed anteriorly, lying below plane of metasternum; suture oblique, rounded near angles. Exposed abdominal segments with sparse and very fine punctation, and with distinct microsculpture consisting of transverse striae. Metacoxal line arcuate, finely punctate; metacoxal area 0.07 mm long. Sexual characters of male. Segments | and 2 of protarsi strongly widened, about as wide as tibiae. Segment 3 of protarsi moderately widened. Aedeagus (Figs 74 to 76, 81) 0.72 mm long. Median lobe symmetrical, with basal bulb large, moderately sclerotized, overlapping distal process in dorsal view. Ventral tubercles distinct, not prominent. Distal process strongly inflexed ventrally, thick basally, abruptly angulate Fics 71 to 76 Aedeagi in Scaphisoma; 71 to 73, S. suspiciosum sp. n., aedeagus in dorsal and lateral views, scale bar = 0.2 mm (71, 72), internal sac in dorsal view, scale bar = 0.2 mm (73): 74 to 76, S. irruptum sp. n., aedeagus in dorsal and lateral views, scale bar = 0.2 mm (74, 76), internal sac in dorsal view, scale bar = 0.1 mm (75). 649 SCAPHIDIINAE OF CHINA, II 76 IRSA 0 NO SI x eo, ~ IN N 75 Dae 44, 357 yp yp) vy) at RE Se, 19 LOI 9 DIR: 5 RD =. ST € OS ; De (Ge = D teelecel it ic x SMA AS YADA > SARA RC > 650 IVAN LOBL posterior to ventral process (lateral view), gradually narrowed toward tip in lateral view and bearing small, subapical tooth on dorsal side; tip acute. Dorsal valves short, completely separated, slightly curved. Internal sac complex, with long, straight basal portion consisting of spinose structures, central plate covering two rows of robust teeth, followed be rows of denticles becoming larger apically, and by fine squamose structures. Parameres sinuate, large in basal portion, abruptly widened in anterior mid-length, forming large inner lobe. Comments. This species is an other member of the S. haemorrhoidale group. It is characterized by the the aedeagus with angulate ventral side of the median lobe, internal sac bearing long, narrow rod, and wide parameres. Scaphisoma volitatum sp. n. Figs 77 to 80 Holotype d: China, Yunnan, Xishuangbanna, 24.1.1993, G. de Rougemont (MHNG). Description. Length 1.6 mm. Body and femora dark reddish-brown, apical third of elytra, apex of abdomen, tibiae, tarsi and antennae light ochreous to yellowish. Basal half of elytra slightly lighter than narrow dark area delimiting apical ochreous third of elytra, and lighter than pronotum. Pronotum with lateral margins evenly arcuate; lateral keels entirely, but hardly, visible; discal punctation fairly dense, very fine, particularly shallow and not clearly delimited. Tip of scutellum exposed. Elytra each with latertal margin slightly arcuate; lateral keel entirely distinct; apical margin truncate; inner apical angle lying posterior to level of outer apical angle, not prominent; sutural margin not raised, adsutural area flat; sutural stria parallel to suture in anterior mid-length, converging to suture from middle to apex, curved near base, ending near pronotal lobe; discal punctation fairly sparse and fine, punctures mostly well delimited, puncture intervals mostly 2 to 3 times as large as puncture diametrers in middle of disc, becoming larger anteriorly. Prohypomera smooth. Mesepisterna very finely punctate. Mesepimeral ridge about twice as long as interval to mesosternum. Metasternum with microsculpture consisting of transverse striae, covering large middle portion and extending to level of lateral edges of mesocoxae; surface lateral of mesocoxae lacking microsculpture. Medio-apical area of metasternum flattened, without microsculpture, with punctation dense and rather fine. Lateral portions of metasternum very finely punctate, with antecoxal puncture row distinct, dense, coarse and irregular. Mesocoxal line arcuate, finely punctate; mesocoxal area 0.07 mm long. Metepisterna flat, strongly narrowed anteriorly, lying below plane of metasternum, with suture oblique in middle, curved near angles. Exposed abdominal segment with microsculpture consisting of transverse striae, except laterobasal portions of first sternite which lacks microsculpture. Punctation of first sternite sparse and very fine. Metacoxal line arcuate, finely punctate, metacoxal area 0.07 mm long. Sexual characters of male. Segments | to 3 of protarsi strongly widened. Seg- ments | and 2 of mesotarsi moderately widened, segment 3 of mesotarsi hardly widened. Aedeagus (Figs 77 to 80) 0.86 mm long. Median lobe large, symmetrical. Basal bulb partly overlapping distal proces, moderately sclerotized. Distal process inflexed, gradually narrowed, with acute tip in dorsal view; narrowed subapically, SCAPHIDIINAE OF CHINA, II 651 with tip dorsoventrally expanded and obliquelly truncate in lateral view. Dorsal valves slightly curved, tapering apically. Ventral tubercles small, not prominent. Internal sac complex, with lateral row of strongly sclerotized, blunt basal teeth, numerous sub- apical denticles, one large, long rod curved and pointed apically, and membranes bearing spinose and squamose structures. Parameres wide, curved posterior of mid- length, with fairly large inner lobe in middle, apical portion distinctly widened and weakly sclerotized. j Comments. This species is another membrer of the S. haemorrhoidale group. It may be distinguished from related species possessing a similar colour pattern only by the aedeagal characters, in particular by the shape of the rod of the internal sac, in combination with the presence of a row of blunt teeth and wide parameres. The aeadeagal characters indicate close relationships to S. tortile Löbl and S. prehensor Champion. The latter species differs drastically by the presence of a flat sclerite in the middle portion of the internal sac. Scaphisoma tortile may be distinguished from S. volitatum by the shape of the rod and the subapical denticulate and spinose area of the internal sac being distinctly delimited. Scaphisoma vestigator sp. n. Figs 82 to 85 Holotype dé: China, Sichuan, Mt. Emei, 1700m, 22.1x.1994, litter, S. A. Kurbatov (MHNG). Paratype d : same data as holotype (MHNG). Description. Length 1.65 - 1.80 mm. Body very dark reddish brown to blackish-brown. Apical sixth to fifth of elytra, apical abdominal segments, femora and tibiae ochreous, tarsi and antennae lighter, yellowish. Ochreous apical band of elytra extending more anteriad, and about as long as one fourth of elytral length, near lateral margins. Pronotum with lateral margins evenly arcuate; lateral keels visible only near base; discal punctation dense and very fine, consisting of shallow, not well delimited punctures. Point of scutellum exposed. Elytra each with lateral margin slightly arcuate anteriorly and in apical third, oblique in middle; lateral keel entirely visible; apical margin truncate; inner apical angle lying posterior level of outer apical angle, not prominent; sutural margin not raised; adsutural area flat; sutural striae diverging from apex to mid-length, parallel from middle to level of pronotal lobe, curved along pronotal lobe to reach base near pronotal lobe; punctation similar to that on pronotal disc near base, along sutural stria and lateral margin, becoming gradually coarser and more dense posteriorly; punctures in middle and on most of posterior half not well delimited, puncture intervals about 2 to 5 times as large as puncture diameters. Prohypomera smooth. Mesepisterna extremely finely punctate. Mesepimeral ridge about as long as interval to mesocoxa. Metasternum with shallow medio-apical impression; finely punctate and with microsculpture consisting of transverse striae in middle. Lateral portions of metasternum lacking microsculpture, with punctation very fine and sparse; antecoxal puncture row inconspicuous, consisting of few, fine punctures. Mesocoxal line arcuate, coarsely punctate; mesocoxal area 0.05 mm long. Metepisterna flat, moderately narrowed anteriorly, lying below plane of metasternum, with suture straight except near angles. Exposed abdominal segments with micro- sculpture consisting of transverse striae, laterobasal portions of first sternite lacking 652 IVAN LOBL microsculpture. Abdominal punctation very fine and sparse. Metacoxal line arcuate, coarsely punc tate; metacoxal area 0.10 mm long. Sexual characters of male. Segments 1 to 3 of protarsi strongly widened, segments 1 and 2 of mesotarsi slightly widened. Aedeagus (Figs 82 to 85) 0.88-0.92 mm long. Median lobe large, symmetrical. Basal bulb weakly sclerotized, almost completely overlapping apical process (dorsal view). Ventral tubercles small, not prominent. Distal process short, obliquely inflexed, with subapical denticle on dorsal side. Dorsal valves slightly curved, blunt. Internal sac complex, with basal portion strongly incurved, bearing dense spinose and squamose structures. Central portion of internal sac narrow, with one long, slender rod. Apical portion of internal sac straight, formed by two lobes joined apically, bearing dense, fine, squamose structures, and larger, strongly sclerotized, subapical denticles. Parameres wide and straight at base, narrowed in middle portion, arcuate in apical half (dorsal view), with long, narrow, inner lobe; slightly sinuate and widest anterior middle in lateral view. Comments. This new species is also a member of the S. haemorrhoidale group.It is unique in having the aedeagus with short distal process overlapped by basal bulb in combination with the internal sac curved basally and bearing a median rod. The aedeagal characters indicate close relationships to S. irruptum. It may be distinguished from S. irruptum by the apical half of the parameres being narrower and the median lobe lacking an angulate area posterior to ventral tubercles. Scaphisoma dumosum sp. n. Figs 85 to 89 Holotype d: China, Yunnan, above Dali, 2000-2200m, 4.-17.iv.1999, W. Schawaller (SMNS). Paratypes: same data as holotype, | d (MHNG); Yunnan, Gongga Shan, Moxi, 1300m, 11.vii.1996, 20°13’N 102°10’E, A. Smetana, J. Farkaë and P. Kabätek, 2 d (MHNG). Description. Length 1.65-1.75 mm. Body dark brown to black, apical fourth of elytra, apical abdominal segments, tarsi and antennae light ochreous, femora and tibiae reddish-brown. Pronotum with evenly arcuate lateral margins; lateral keel not exposed; discal punctation extremely fine, sparse, consisting of very shallow and not clearly delimited punctures. Point of scutellum exposed. Elytra each with lateral margin slightly arcuate; lateral keel entirely visible; apical margin trunctate; inner apical angle posterior to level of outer apical angle, not prominent; sutural margin not raised; adsutural area flat; sutural striae diverging from apex to mid-length, parallel from middle to level of pronotal lobe, curved along pronotal lobe to reach base near pronotal lobe; punctation similar to that on pronotal disc, near base, along sutural stria and lateral margin, becoming gradually coarser and denser posteriorly; punctures in middle and on most of posterior half not well delimited, puncture intervals about 2 to 5 times as large as puncture diameters. Prohypomera and mesepisterna extremely Fics 77 to 82 Aedeagi in Scaphisoma; 77 to 80, S. volitatum sp. n., aedeagus in dorsal and lateral views, scale bar = 0.2 mm (77, 78), internal sac in dorsal view, scale bar 0.1 mm (79), paramere in ventral view, scale bar = 0.1 mm (80); 81, S. irruptum sp. n., paramere in ventral view, scale bar = 0.1 mm; 82, S. vestigatum sp. n., paramere in ventral view, scale bar = 0.1 mm. 653 SCAPHIDIINAE OF CHINA, II 81 RAUM 80 TPE Son D RAT = DI iy DAS PAR 22 erg N Giorno qu > ( dt Nr = Fa i 4 Sat UES N Ÿ 2 . oy CLeccte CIA LE beer ur Mi at ee e ni 654 IVAN LOBL finely punctate. Mesepimeral ridge about twice as long as interval to mesocoxa. Metasternum without median impressions, flattened medio-apically; with micro- sculpture consisting of transverse striae present only on narrow medio-apical area, most of surface lacking microsculpture. Punctation sparse and very fine on lateral portions of metasternum, less fine on center of metasternum. Mesocoxal line subtrian- gular, coarsely punctate; mesocoxal area 0.07-0.08 mm long. Antecoxal puncture row absent. Metepisterna flat, hardly narrowed anteriorly, lying below plane of meta- sternum, with straight suture between rounded angles. Exposed abdominal segments with microsculpture consisting of transverse striae. First abdominal sternite sparsely and coarsely punctate, lacking microsculpture on basolateral area. Metacoxal line arcuate, very finely punctate; metacoxal area 0.08-0.10 mm long. Sexual characters of male. Segments | to 3 of protarsi strongly widened, seg- ments | and 2 of mesotarsi moderately widened. Aedeagus (Figs 85 to 89) 0.69 - 0.72 mm long. Median lobe symmetrical, moderately sclerotized. Basal bulb large, partly overlapping distal process. Distal process strongly inflexed, almost straight, gradually narrowed, with subapical denticle; tip acute. Dorsal valves slightly curved, acute. Ventral tubercles small, inconspicuous. Internal sac complex, with straight, wide and long basal portion consisting of rows of fairly strongly sclerotized central denticles covered by fine spicules, and lateral, weakly sclerotized, irregularly structured areas. Subapical portion of internal sac with moderately sclerotized, triangular plate rising apically and curved (lateral view); apical portion with six blunt, strongly sclerotized, robust denticles joined to finely denticulate membranous area. Parameres almost on same plane as basal bulb, arcuate in apical half (dorsal view), fairly wide, with narrow inner lobe; widest in middle and slightly narrowed toward apex in lateral view. Comments. This species exhibits on internal sac of the aedeagus similar to that of S. migrator. Both species have robust apical denticles and basal portion of the internal sac straight, bearing fine spicules. Scaphisoma dumosum may be separated from S. migrator by the parameres being more curved and lacking a subapical lobe, and by the presence of a plate in the subapical part of the internac sac. ACKNOWLEDGEMENTS My thanks are due to following colleagues and friends who provides material, either as loan from collections of which they are in charge, or as donation: S. Be¢var (Ceské Budèjovice), N. Berti (Paris), D. Burckhardt (Basel), J. Cooter (Hereford), Z. Kejval (Domazlice), D. Kral (Praha), V. Kuban (Brno), S. A. Kurbatov (Moscow), J. Jelinek (Prague), C. Johnson (Manchester), A. Piitz (Eisenhiittenstadt), G. de Rougemont (London), W. Schawaller (Stuttgart), M. Schiilke (Berlin), A. Smetana (Ottawa), and E. Sprecher (Basel). I thank also G. de Rougement (London) for comments on the manuscript which improved the study. FIGs 83 to 89 | Aedeagi in Scaphisoma; 83 to 85, S. vestigatum sp. n., aedeagus in dorsal and lateral views (83, 84), internal sac in dorsal view (85); 86 to 89, S. dumosum sp. n., aedeagus in dorsal and lateral views (86, 87), internal sac in dorsal view (88), paramere in ventral view (89). Scale bars = 0.1 mm. 655 II ’ SCAPHIDIINAE OF CHINA 89 za N A I 2% == r STAR { io aya, a } rA See ef LE = Erz — our! LAYMAN ig 07> RA, = 0m cn SS BS EEE a de y Se D te eZ III I CUS CA O. LAT rer yen az EG Oo ZE z CEN ERDE SZ Dre PVE 2 WPA II ARS 87 7 WILL 00 ) rn SLI) Gy, Ir LT 000 e re TA QE WWW IK yeh SPAN vesti ORTE | Se SSS os UN, \ > 656 IVAN LOBL REFERENCES ASHE, J. S. 1984. Description of the larva and pupa of Scaphisoma terminata Melsh. and the larva of Scaphium castanipes Kirby with notes on their natural history (Coleoptera: Scaphi- diidae). The Coleopterists Bulletin 38: 361-373. HANLEY, R. S. 1996. Immature stages of Scaphisoma castaneum Motschulsky (Coleoptera: Sta- phylinidae: Scaphidiinae), with observation of natural history, fungal hosts and develop- ment. Procedings of the Entomological Society of Washington 98: 36-43. JÄCH, M. A. & Ji, L. 1995. Introduction. /n: Water beetles of China. Vienna, 410 pp. LAWRENCE, J. F. & NEWTON, A. F. 1980. Coleoptera associated with the fruiting bodies of slime molds (Myxomycetes). The Coleopterists Bulletin 34: 129-143. LESCHEN, R. A. B. 1988. The natural history and immatures of Scaphisoma impunctatum (Cole- optera: Scaphidiidae). Entomological News 99: 225-232. LESCHEN, R. A. B. 1994. Retreat-building by larval Scaphidiinae (Staphylinidae). Mola 4: 3-5. Lost, I. 1965a. Beitrag zur Kenntnis der japanischen Arten der Gattung Scaphosoma Leach (Scaphidiidae). Entomologische Blätter 61: 44-58. Los, I. 1965b. Beitrag zur Kenntnis der Scaphosoma-Arten Chinas (Coleoptera, Scaphidiidae). Reichenbachia 6: 25-31. LÔBL, I. 1970. Revision der Gattungen Scaphisoma Leach und Caryoscapha Ganglbauer der Tribus Scaphisomini (Col. Scaphidiidae). Revue suisse de Zoologie 77: 727-799. LößL, I. 1973. Über einige orientalische Scaphidiidae (Coleoptera) aus dem Museo Civico di Storia Naturale di Genova und Museum National d’Histoire Naturelle de Paris. Nouvelle Revue d’Entomologie 3: 149-160. LOBL, I. 1979. Die Scaphidiidae (Coleoptera) Südindiens. Revue suisse de Zoologie 86: 77-129. LößL, I. 1980. Beitrag zur Kenntnis der Scaphidiidae (Coleoptera) Taiwans. Revue suisse de Zoologie 87: 91-123. Los, I. 1981a. Über die Arten-Gruppe Rouyeri der Gattung Scaphisoma Leach (Coleoptera Scaphidiidae). Archives des sciences (Geneve) 34: 153-168. LOBL, I. 1981b. Über einige Arten der Gattung Scaphisoma Leach (Coleoptera, Scaphidiidae) aus Vietnam und Laos. Annales historico-naturales musei nationalis Hungarici 73: 105-112. LOBL, I. 1982. Uber die Scaphidiidae (Coleoptera) der japanischen Ryukyu-Inseln. Mitteilungen der Schweizerischen entomologischen Gesellschaft 55: 101-105. LosL, I. 1986a. Les Scaphidiidae (Coleoptera) du nord-est de l’Inde et du Bhoutan II. Revue suisse de Zoologie 93: 133-212. LosL, I. 1986b. Contribution à la connaissance des Scaphidiidae (Coleoptera) du nord-ouest de I’ Inde et du Pakistan. Revue suisse de Zoologie 93: 341-367. LosL, I. 1990a. Contribution à la connaissance des Scaphisoma (Coleoptera, Scaphidiidae) de 1’ Himachal Pradesh, Inde. Archives des sciences (Genève) 43: 117-123. LosL, I. 1990b. Review of the Scaphidiidae (Coleoptera) of Thailand. Revue suisse de Zoologie 97: 505-621. LÒBL, I. 1992. The Scaphidiidae (Coleoptera) of the Nepal Himalaya. Revue suisse de Zoologie 99: 471-627. Los, I. 1993. Contribution to the knowledge of the Scaphidiinae (Coleoptera) of the Far East Region of Russia. Russian Entomological Journal 2: 35-40. LosL, I. 1997. Catalogue of the Scaphidiinae (Coleoptera: Staphylinidae). Instrumenta biodiver- sitatis I: XII + 190 pp. LosL, I. 1999. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People’s Republic of China, I. Revue suisse de Zoologie 106: 691-744. Nuss, I. 1975. Zur Okologie der Porlinge. Untersuchungen iiber die Sporulation einiger Porlinge und die an ihnen gefundenen Käferarten. Bibliotheca Mycologica (Vaduz) 45: 1-258. Pic, M. 1954. Coléoptéres nouveaux der Chine. Bulletin de la Société entomologique de Mulhouse 1954: 53-59. SMETANA, A. 1996. Two new species of Deinopteroloma from China (Coleoptera: Staphylinidae: Omaliinae). Koleopterologische Rundschau 66: 77-81. REVUE SUISSE DE ZOOLOGIE Tome 107 — Fascicule 3 PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphy- linidae). 261. Contribution to the knowledge of Steninae............ GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar. . . . . MUSTER, Christoph & Konrad THALER. Das Männchen von Zelotes zellensis GrimmiAraneae: Gnaphosidae) =... „u. A VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Note sur les musaraignes (Soricidae, Mammalia) du Parc National du Souss-Massa, Maroc..... Los, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the People ssRepubliciof@hima le Re Pages 419-559 561-577 579-589 591-599 601-656 REVUE SUISSE DE ZOOLOGIE Volume 107 — Number 3 PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphy- linidae). 261. Contribution to the knowledge of Steninae............ GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar. . ... MUSTER, Christoph & Konrad THALER. The male of Zelotes zellensis Grimm (AraneaeAGnaphosidae). 105. CRIER VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Shrews (Soricidae, Mammalia) of the National Park Souss-Massa, Morocco............ LÔBL, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the Beople;s/Republie/ot China: Il. Ce si as RE eee Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 419-559 561-577 579-589 591-599 601-656 PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N9s 1-17 (1908-1926)... .. série Fr. (prix des fascicules sur demande) REMUFIDEIPÄBEOBIOLOGIER. mn. ee: Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) COLLEMBOLENFAUNA EUROPAS von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Fr. THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION WITH KEYS FOR DETERMINATION ER ial par fascicule Fr. EVAANOSER S4oipacesslultioures mytext LINSE ER Fr. CLASSIFICATION OF THE DIPLOPODA Pamkachand i HOFEMANS237)PA ESA LOTS) N EPA RO Net Fr LES OISEAUX NICHEURS DU CANTON DE GENEVE par P. GEROUDET, C. GUEX et M. MAIRE 351 pages, nombreuses cartes et figures, 1983 CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ A LA CONNAISSANCE DES ECHINODERMES ACTUELS PamVichelAncouxeG/ pages. Ml planches; 198977... en ee Fr. RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) Pal NeOkINE DO WUE Ke IMIAREDANO2 pases il SOM Mii creme NE NEA Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN; SHELLS AND RADULAS bya YeO RINE Po UESiiAn Gs KGIMARED A992 ee Fr. O. SCHMIDT SPONGE CATALOGUE par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992 .......... Fr. ATLAS DE REPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr. THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST DANAVESIEINER SO: PASTE Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE palean Claude CAmirIEZE4 Oipages lOO SES ARE rl le el er hentai eal clive eters Er: PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) (Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997.................... Fr. CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis II) par Henri G. DIRICKx, 188 pages, 1998 ............ Fr. A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Er. 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Phone: +41 22 418 63 33 - Fax +41 22 418 63 01 e-mail: volker.mahnert @ mhn.ville-ge.ch Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 1IDO TOOZ JG ASSINS ANAAY ADOTOOZ 40 IVNYNOÏ SSIMS eve N N BRE 2000 ISSN 0035 - 418 X le de Gen REVUE SUISSE DE ZOOLOGIE TOME 107 — FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève MANUEL RUEDI Chargé de recherche au Muséum d’histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d'histoire naturelle de Genève Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des Instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le sujet étudié. 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Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L'ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 230 — (en francs suisses) Les demandes d'abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6. Suisse REVUE SUISSE DE ZOOLOGIE 107 (4): 657-662; décembre 2000 Redescription of Hymenaphorura alticola (Bagnall, 1935) from the Alps and description of a new related species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola: Onychiuridae) Romuald J. POMORSKI & Dariusz SKARZYNSKI Zoological Institute, Wroctaw University, Sienkiewicza 21, PL-50-335 Wroctaw, Poland. E-mail: onychus @biol.uni.wroc.pl, hypogast@biol.uni.wroc.pl Redescription of Hymenaphorura alticola (Bagnall, 1935) from the Alps and description of a new related species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola: Onychiuridae). - Hymenaphorura alticola (Bagnall, 1935) is redescribed based on material from Switzerland, France, Austria and Italy. A neotype is designated. Hymenaphorura improvisa sp. n. from Poland is described and figured. Key-words: Collembola - Onychiuridae - Hymenaphorura — taxonomy - Europe. INTRODUCTION During faunistic studies in the Sudetes (SW Poland) some specimens, which fit Bagnall’s (1935) description and Gisin’s redescription (1953, 1960) of A. alticola (Bagnall, 1935), were found. A detailed comparison with Gisin’s material which we have obtained thanks to the kindness of Dr. Charles Lienhard from the Muséum d’histoire naturelle (Genève) made it possible to ascertain that the Polish specimens represented a new species and H. alticola needed a modern redescription. Hymenaphorura alticola (Bagnall, 1935) Material. Neotype — female on slide, by present designation; (CZ 44) Schweiz (Kanton Bern), Berner Jura, Höhle Gitziloch bei Court, 12. IX. 1954, leg. P. Strinati (in collection of the Museum d’histoire naturelle in Geneva). Other material examined - 2 females, 1 male juv.; (Fn 7) Suisse (Neuchatel), Jura neucha- telois, Val de Travers, grotte de Môtiers, 21. VII. 1952. leg. P. Strinati. 3 males, 3 females; (Jb Rr 1/4 and 1/5) Osterreich, Tirol, Zillertaler Alpen, Gegend um Schönbichler Horn, Rossrucken, unterhalb Rossruckspitz, nivale Pionier-Polster, 3100 m, Sommer 1946, leg. H. Janetschek. 3 females; (Jb Rr 4/1) Österreich, Tirol, Zillertaler Alpen, Gegend um Schönbichler Horn, Rossrucken, Schneeboeden, Polytrichetum sexangularis, 2620 m, Sommer 1946, leg. H. Janetschek. 5 females; (Kg 191) France (Jura), St. Claude, grotte de la Grusse B, 40 m sous terre, II. 1963, leg. J. Colin. 4 juv.; (Ma 2a) Italie (Dolomites), Marmolada, 2000 m, prairie sur alluvions, 1956, leg. G. Marcuzzi. 2 females, 2 males, 4 juv; (Ma 5a) Italie (Dolomites), Marmolada, 2100 m, prairie alpine sur roche, 1956, leg. G. Marcuzzi. | male, 2 juv.; (Ma 16a) Italie (Dolomites), Marmolada, 2400 m, prairie alpine sur roche, 1956, leg. G. Marcuzzi (the material is preserved in the collection of the Muséum d’histoire naturelle in Geneva). Manuscript accepted 29.05.2000 658 R. J. POMORSKI & D. SKARZYNSKI Diagnosis. H. alticola is closely related to Hymenaphorura creatricis Pomorski, 1990. Both species have p, setae longer than p, on abdominal terga I-II, a similar chaetotaxy of abdominal tergum V, the same localisation of microsensillum on anten- nomere IV and a similar shape and proportions of tibiotarsi and claws. H. alticola distinctly differs from H. creatricis in its pseudocellar formula (20/111/11112 - A. alticola, 10/011/11112 - H. creatricis) and finer granulation of terga. Redescription. Colour in alcohol white. Size, without antennae, 1.6-2.0 mm (neotype female — 2.0 mm). Shape of body cylindrical, as in Fig. 1. Antennae as long as head. Trace of reduced furca in shape of 2 symmetrical small patches of fine granulation with 2 small setae posteriorly. Granulation of dorsal side of the body very fine — usually 12-13 grains around each pseudocellus. Granular areas almost invisible. Antennal III sense organ consists of 5 relatively long papillae; 2 sensory rods; 2 finely granulated, ovoid sensory clubs; 4 guard setae. Antennal segment IV with sub- apical organite and microsensillum at level of first row of setae. On antennal segment III microsensillum localised laterally, slightly below antennal HI sense organ (Fig. 3). Postantennal organ in a long cuticular groove, with 11 (9-16) simple vesicles (Fig. 2). Labium without lateral papilla E. Pseudocellar formula 20/111/11112, ventral pso absent. Parapseudocellar formula typical for the genus - dorsally O1/111/1111, ventrally 1/000/11111. Parapseudocelli on subcoxa and femora invisible. Dorsal chaetotaxy symmetrical, well differentiated into macrochaetae and microchaetae, as in Fig. 1. Thorax II-III with lateral microsensilla. On I-IV abdominal terga, setae p, shorter than p3. Granular area on abdominal tergum V with 4+4 more or less distinct macrochaetae. Subcoxae with 3, 5 (4), 5 (4) setae, between legs on meso- and metathorax 1+1 setae, tubus ventralis usually with 9+9 setae. Male ventral organ absent. Claw without denticle. Empodial appendage long with narrow basal lamella, appendage length equal to that of inner edge of claw (Fig. 5) (specimens from Italian Dolomites have a shorter appendage and a slightly broader basal lamella, as in Fig. 6). Distal whorl of setae on tibiotarsi symmetrical, with 11 setae. Anal spines strong, weakly curved, pointed, with relatively small basal papillae, as long as the claw. Remarks. Handschin (1920) summarised the knowledge about the fauna of Swiss Onychiuridae and briefly described individuals with 2+2 pseudocelli at the base of antenna as Onychiurus sibiricus Tullberg, 1876. Bagnall (1935) pointed out, that “true” O. sibiricus had 1+1 pseudocellus at the base of antenna and proposed to classify the individuals examined by Handschin as a new species Onychiurus alticola. Later, Gisin (1953, 1960), on the basis of new materials from Austria, France, Switzerland and Italy, redescribed this species. Examination of Gisin’s material allowed us to redescribe this species using a set of modern, diagnostic characters (Pomorski, 1990, 1998). Moreover, we decided to designate of neotype, since Handschin’s syntypes could not be located [Dr. Charles Lienhard - Muséum d’histoire naturelle (Genève) and Dr. Daniel H. Burckhardt - Naturhistorisches Museum (Basel); personal communication]. REDESCRIPTION OF HYMENAPHORURA ALTICOLA 659 Fics 1-6. Hymenaphorura alticola: 1 - habitus and dorsal chaetotaxy; 2 - postantennal organ and pseudocelli at base of antenna; 3 - antennal III sense organ; 4 - chaetotaxy of abdominal terga V-VI; 5 - tibiotarsus and claw of legs III (specimen from French Jura); 6 - tibiotarsus and claw of legs III (specimen from Italian Dolomites). 660 R. J. POMORSKI & D. SKARZYNSKI Hymenaphorura improvisa sp. n. Type material. Holotype, male on slide; Poland, Sudetes, Karkonosze Mts., neigh- bourhood of Szklarska Poreba, 17. X. 1999; flood debris of the river Kamienna, leg. D. Skarzynski. Paratypes 1 male and 3 females on 2 slides, same locality, 10. V. 1999, leg. D. Skarzynski; 4 females on slide, same locality, 15 V 2000, leg. D. Skarzynski; 33 specimens in alcohol, same locality, 15 V 2000, leg. D. Skarzynski (type material preserved in the collection of the Department of Systematic Zoology and Zoogeography, Wroctaw University, Poland; one paratype female housed in the collection of the Muséum d’histoire naturelle in Geneva). Etymology. The species name is derived from Latin word “improvisus” — unexpected. Diagnosis. Four guard setae in antennal III sense organ, labial lateral papilla E absent, the shape of remnant of furca and the presence of dorso-internal pseudocelli indicate that the new species belongs to the genus Hymenaphorura. The new species resembles H. alticola only in its pseudocellar formula, but two important characters: 9 setae in the distal whorl of tibiotarsi and the presence of median seta mg on abdominal tergum V, place this species in an isolated position within the genus. Description. Size, without antennae, 1.4-1.5 mm males, 1.5-1.7 mm females (holotype male — 1.5 mm). Colour in alcohol white. Shape of body cylindrical, elon- gated as in Fig. 7. Antennae as long as head. Trace of reduced furca in shape of small patches of fine granulation with 2 small setae posteriorly. Granulation of the dorsal body side distinct, with very well visible granular areas - usually 10-11 grains around each pseudocellus. Antennal III sense organ consists of 5 simple papillae; 2 sensory rods; 2 granulated, spherical sensory clubs of which internal is bent; 4 guard setae (Fig. 8). Antennal segment IV with subapical organite and microsensillum in latero- external position, at level of second row of setae. On antennal segment III micro- sensillum localised laterally, slightly below antennal III sense organ (Fig. 8). Postantennal organ in short cuticular groove, with two border setae, composed of 10-11 simple and bilobed vesicles (Fig. 9). Labium without lateral papilla E and usually with thickened and blunt-tipped sensilla on papilla C. Pseudocellar formula (pso) 20/111/11112, ventral pso absent. Parapseudocelli poorly visible (we are sure of the presence of 1+1 parapseudocelli on abdominal sternum V). Dorsal chaetotaxy symmetrical, well differentiated into macrochaetae and microchaetae, as in Fig. 7. On abdominal terga I-III, setae p2 distinctly longer than p3. Granular area on abdominal tergum V with 3+3 distinct macrochaetae and median seta m0 (Fig. 10). Subcoxae with 3, 4, 4 setae. Between legs on meso- and metathorax 1+1 setae. Tubus ventralis with 7+7 (6+6) setae. Male ventral organ absent. Claws relatively short, always without denticle. Empodial appendages with narrow basal lamella, appendage length equal to that of the inner edge of claw. Distal whorl of setae on tibiotarsi symmetrical, with 9 setae (Figs. 11-13). Anal spines strong, weakly curved, pointed, with distinct basal papillae, as long as the claw. Ecological remarks. H. improvisa was collected singly in flood debris and numerously under stones on gravel bed of the river Kamienna. REDESCRIPTION OF HYMENAPHORURA ALTICOLA 66] i= RE i acaNeie AG LI c{ DA EO) re D > 4 > ES LA COUR 73, 2 NZ AO “ NANI: ceo di exe Cote? 4 A vHIEO PARSO 2 ) A SS ALI 3 e D 220 ) ati Zare 26 RZ "E 2 : s Kw e > Ze n= = CMOS x A) SI DE 568 IONI ai EE? d Vy Ar, oa ves E O CITI MSA ATA ‘SCOLA ROL 500 DEIR sdnn, a < J IRA 3 < a in € ce ee 3 9, o a BER Lim È ° o! Fics 7-13. Hymenaphorura improvisa sp. n.: 7 - habitus and dorsal chaetotaxy (holotype): 8 - antennal III sense organ; 9 - postantennal organ and pseudocelli at base of antenna; 10 - chaetotaxy of abdominal terga IV-VI; 11 - tibiotarsus and claw of legs III (dorsal side); 12 - tibiotarsus and claw of legs III (ventral side); 13 - claw of legs III. 662 R. J. POMORSKI & D. SKARZYNSKI Discussion. Within the genus Hymenaphorura Bagnall, 1948 three species with 2+2 pseudocelli at the base of antenna are known - Hymenaphorura alticola (Bagnall, 1935) which lives in Swiss, French, Italian and Austrian Alps (Gisin, 1960), Hymenaphorura rafalskii Weiner & Szeptycki, 1997 from Korea and Hymenaphorura improvisa sp. n. from Poland. However, they do not constitute a natural group of related species. Besides the common character mentioned above, they differ in some essential, exceptional characters. H. improvisa sp. n. is easy to identify based on the presence of 9 setae in the distal whorl of tibiotarsi (in the remaining species 11 setae), the Korean species has 3+3 pseudocelli on abdominal tergum V (in the remaining species 2+2 pseudocelli) and A. alticola has pz setae longer than p, on abdominal terga I-II (in A. improvisa p3 is distinctly shorter than p,, in H. rafalskii these setae are subequal). ACKNOWLEDGEMENTS We wish to express our sincere thanks to: Dr. Charles Lienhard from Geneva for loans of Gisin’s material of A. alticola; Prof. Erhard Christian from Vienna for kind information and search for H. alticola in Austria; Dr. Daniel Burckhardt from Basel for kind information and search for H. alticola in Handschin’s collection. The study was sponsored by the Wroctaw University - grant No. 2020/W/IZ/99. REFERENCES BAGNALL, R. S. 1935. Notes on British Collembola. Entomologist’s Monthly Magazine 75: 61- 63. GISIN, H. 1953. Notes sur les Collemboles, aves description de cing espèces nouvelles, décou- vertes dans le canton de Genève. Mitteilungen der Schweizerischen entomologischen Gesellschaft 26: 56-62. GISIN, H. 1960. Collembolenfauna Europas. Muséum d'histoire naturelle, Genève, 312 pp. HANDSCHIN, E. 1920. Die Onychiurinen der Schweiz. Verhandlungen der Naturforschenden Gesellschaft in Basel 32: 1-37. POMORSKI, R. J. 1990. New data on the genus Hymenaphorura (Collembola, Onychiuridae) from Europe. Mitteilungen der Schweizerischen entomologischen Gesellschaft 63: 209- 225; POMORSKI, R. J. 1998. Onychiurinae of Poland (Collembola: Onychiuridae). Genus, supple- ment: 1-201. REVUE SUISSE DE ZOOLOGIE 107 (4): 663-674; décembre 2000 Allozyme discrimination of three species of Loricartichthys (Siluriformes: Loricariidae) from Southern Brazil! Claudio H. ZAWADZKI’, Roberto E. REIS? & Erasmo RENESTO! | Universidade Estadual de Maringä, Nücleo de Pesquisas em Limnologia, Ictiologia e Agüicultura G-90, Av. Colombo, 5790, 87020-900 — Maringä, PR, Brasil. E-mail: chzawadzki @nupelia.uem.br or e-mail: renesto@nupelia.uem.br 2 Pontificia Universidade Catölica do Rio Grande do Sul, Museu de Ciéncias e Tecno- logia, Caixa Postal 1429, Av. Ipiranga, 6681, 90619-900 — Porto Alegre, RS, Brasil. E-mail: reis @pucrs.br Allozyme discrimination of three species of Loricariichthys (Siluri- formes: Loricariidae) from Southern Brazil. - Three species of armored catfish of the genus Loricariichthys were clearly discriminated by starch gel isozyme electrophoresis on heart, liver and muscle tissues. We scored 21 enzyme loci from 14 isozyme systems (ACP, ADH, AAT, EST, GCDH, G3PDH, G6PDH, GPI, ICDH, LDH, MDH, MDHP, PGM and SOD). Expected mean heterozygosity (H,) ranged from 0.023 in Loricartichthys anus to 0.050 in L. platymetopon and 0.035 in Loricariichthys sp. The percentage of polymorphic loci (Pp 99) was 14.3% for the three analyzed species. Nei’s genetic identity (1) was found to be 0.663 between Lorica- riichthys anus and Loricariichthys sp., 0.329 between L. anus and L. platy- metopon, and 0.478 between L. platymetopon and Loricariichthys sp. Genetic variability, tissue specific differences, as well as applicability of isozyme electrophoresis as a tool for neotropical fish systematists are discussed. Key-words: Genetic diversity - isozymes - polymorphism - Siluriformes - taxonomy - systematics - fish - Loricariichthys. INTRODUCTION The freshwater fish fauna of South America is the richest of all the continents and rivals in diversity with the coral reef fishes. In a recent estimate of the diversity of neotropical ichthyofauna Schaefer (1998) suggested that the total number of species of fish in this region may reach about eight thousands. The gaps in the systematic knowledge of neotropical fishes are as great as the number of species, and can be sustained principally by the lack of overall representation in scientific collections. One of the many difficulties from dealing with neotropical fishes is related to the great ! This paper is a part of the Ph.D thesis of Claudio Henrique Zawadzki, at the State University of Maringa, granted by CAPES. Manuscript accepted 12.07.2000 664 C.H. ZAWADZKI, R.E. REIS & E. RENESTO number of forms in each taxonomic group and to the great similarity found among species of the same genus. The family Loricariidae is the largest among the Siluri- formes, with many species difficult to identify in several genera. In Loricariichthys Bleeker, 1862 as is usually the case for fish systematics and taxonomy, morphological characters forms the bulk of data used in descriptions as well as in species diagnoses (Reis & Pereira, in press). The genus Loricariichthys comprises 16 valid species (Isbriicker, 1980) and do not diverge from other neotropical taxa of fishes in regard to the gaps in the systematic knowledge. Thus, we search auxiliary tools for the morpho- logical criteria. Different cytogenetic, biochemical and molecular techniques have been developed and used to assist in the diagnosis and species identification. As suggested by Thorpe & Solé-Cava (1994), isozyme electrophoresis can become an extremely useful tool when combined with morphological methods for the discrimination of closely related species. In this work we examine the level of genetic identity and variability among three species of Loricariichthys through allozyme electrophoresis, assaying loci and tissue expression. These are first and partial results from an attempt of the first author to form a bulk of allozyme data of some Loricariidae catfishes from southern Brazil, with the objective of finding genetic markers that could help to resolve questions on the systematics of this family. We also aim to corroborate the use of allozyme electrophoresis as a tool for the discrimination of morphologically similar species of Loricariidae. MATERIALS AND METHODS After withdraw the tissues, the specimens were fixed in 10% formalin and they are alcohol preserved in the synoptic collection of Nupélia at the State University of Maringä, Brazil. Voucher specimens are deposited at the Museu de Zoologia da PUC, Porto Alegre, Brazil, L. platymetopon MCP25476 and MCP25749; Loricariichthys sp. MCP25747 and MCP25748, and also at the Muséum d'Histoire Naturelle, Geneva (Switzerland). A list of specimens was arranged in the text as follow: locality, town, collecting date, number of specimens (if there are more than one) between parenthesis and the standard length in centimeters (the smallest and biggest if there are more than one in the lot). LIST OF SPECIMENS STUDIED Brazil, Parana State: Loricariichthys platymetopon: Vila Ipiranga (25°14'12"S/54°14'02"W), rio Ocoi, rio Parana basin, Sao Miguel do Iguacu, 10.1x.1998, 27.8; 16-17.1x.1998, (4) 22.7-26.4; Passo Cöe (25°21'64"S/54°24'50"W), Itaipu lake reservoir, Foz do Iguacu, 15.1x.1998, (2) 23.7-26.3; Sao Vicente (25°01'20"S/54°23'57"W), Itaipu lake reservoir (25°01°20°S/54°23’57’’W), rio Parana basin, Santa Helena, 13.1x.1998, 30.1; Ponto do Chico Barbudo (25°22'32"S/54°26'05"W) Itaipu lake reservoir, rio Parana basin, Santa Terezinha do Itaipu, 14.1x.1998, (4) 22.9-27.8; Esquina Gaucha (25°17'42"S/54°19'20"W), Itaipu lake reservoir, rio Parana basin, Itaipuländia, 15.1x.1998, 28.9; Itaipu lake reservoir (24°28'19"S/54°18'49"W), rio Parana basin, Marechal Candido Rondom, 18.1x.1998, (3) 24.6-31.6. Loricariichthys sp.: Vila Ipiranga (25°14'12"S/54°14'02"W), rio Ocoi, rio Parana basin, Sao Miguel do Iguacu, 10.1x.1998, (5) 25.3-28.2; 17.1x.1998, (3) 223.7-25.4 Rio Sao Francisco Verdadeiro (24°40'32"S/54°13'92"W), rio Parana basin, Pato Bragado, 11.1x.1998, (2) 19.5-25.3; ALLOZYME DISCRIMINATION OF LORICARIICHTHYS 665 Ponto do Chico Barbudo (25°22'32"S/54°26'05"W), Itaipu lake reservoir, rio Parana basin, Santa Terezinha do Itaipu, 14.1x.1998, (2) 22.7-26.3; Esquina Gaucha (25°17'42"S/54°19'20"W), Itaipu lake reservoir, rio Parana basin, Itaipulandia, 15.1x.1998, 27.9 . Brazil, Rio Grande do Sul State: Loricariichthys anus: Fortaleza lagoon (30°09'02"S/50°14'30"W), Cidreira, 20.11.1999, (8) 25.7-26.4. Samples of white skeletal muscle, liver and heart tissues were collected. The tissues were stored in liquid nitrogen until analysis. Samples of tissues were homo- genized with plastic sticks, in microcentrifuge tubes with Tris/HCl 0.02 M, pH 7.5 buffer in 1:1 concentration. When homogenizing liver samples, 0.5 ml of CCl, was added to precipitate the great amount of fat present in such tissue (Pasteur er al., 1988). The homogenized samples were centrifugated at 25,000 rpm (44,720 x g) for 30 min. at temperatures between 1° C and 5° C. The supernatant fractions were applied to starch gel using straps of Whatman paper 3 MM®. Gels were prepared with corn starch (Penetrose®) 13% in a buffer solution. Two buffer systems were used for the electrophoresis: Tris- citrate, pH 7.0 (Shaw & Prasad, 1970) and Tris-borate-EDTA pH 8.3 (Boyer er al., 1963). The electrophoresis was run during six hours at 5° C with 250 V for Tris-citrate gel and 450 V for Tris-borate-EDTA gel. Gels were then cut horizontally and each slice was incubated with a solution of appropriate staining for each enzyme. The staining procedures were those described by Aebersold er al. (1987), with the exception of the AAT which followed the procedures of Morizot & Schmidt (1990). The nomenclature used was that proposed by the International Union of Biochemistry and Molecular Biology (1992). The data were analyzed using Biosys 1 software (Swofford & Selander, 1981). The genetic interpretation of the enzymatic patterns was based on the quaternary structure of enzymes described by Ward et al. (1992). RESULTS Fourteen enzymatic systems of eight Loricariichthys anus, 16 L. platymetopon and 13 Loricariichthys sp. were analyzed, allowing the detection of 21 loci (Table 1). Electrophoretic patterns of enzymes in each tissue and their genetic interpretation for the three species analyzed are shown in Figure 1. Only the most common alleles for each species are represented. The enzymes ACP, ADH, GCDH and G6PDH present a single band only in liver samples. Esterases are more active in the liver, but they are also expressed in other tissues, though less intensely. AAT, PGM and SOD zymograms show that these enzymes have the same activity in the three tissues and exhibit only one band. The enzyme G3PDH presents three bands in white muscle, probably due to presence of two loci and the formation of a heterodimer interloci. Locus G3pdh-A also presents strong activity in heart and liver, while locus G3pdh-B has a weak expression in liver and is practically absent in heart tissues of the three species. The enzyme GPI presents three regions of activity in the three species, as a result of expression of two loci Gpi-A and Gpi-B, and the intermediate region as corresponding to the formation of heterodimer interloci. In heart tissues the two loci 666 C.H. ZAWADZKI, R.E. REIS & E. RENESTO TABLE 1. Names, Number of Enzyme Commission (E.C.n°), tissues, buffers, interpreted quaternary structure (Q.S) and number of loci expressed in Loricariichthys anus, L. platymetopon and Loricariichthys sp. L = liver; M = muscle; H = Heart; TBE = Tris/borate/EDTA; TC = Tris/citrate. | | | | | | mH (© > o 5 | n | a [= oO Enzyme (Abbreviation) È Buffer Q.S. Loci Acid phosphatase (ACP) 332 L TC Dimeric I Alcohol dehydrogenase (ADH) Lee Ib TBE — Dimeric l Aspartate aminotransferase (AAT) 2.6.1.1 L,H,M TBE Dimeric 1 Esterase (EST) alal L,H,M TBE Monomeric | Glucose 1-dehydrogenase - NAD+ (GCDH) TS IST TBE Dimeric 1 Glycerol-3-phosphate dehydrogenase (G3PDH) 1.1.1.8 PANNE Dimeric 2 Glucose-6-phosphate dehydrogenase (G6PDH) 1.1.1.49 L TBE Tetrameric 1 Glucose-6-phosphate isomerase (GPI) 584109 L, MATE Dimeric 2 Isocitrate dehydrogenase - NADP+ (ICDH) LME "EMTEC Dimeric 2 L-Lactate dehydrogenase (LDH) 1.1.1.27 eM IKE Tetrameric 2 Malate dehydrogenase (MDH) S37 Sevier Dimeric 3 Malate dehydrogenase - NADP+ (MDHP) ll AO EME Tetrameric 2 Phosphoglucomutase (PGM) 5.4.2.2 SEMI Monomeric 1 Superoxide dismutase (SOD) IMSS, E,H, MONTRE PDimeric l present an equally intense activity. In liver, Gpi-A expresses itself with more intensity than that of Gpi-B, while the reverse is true for muscle tissues. The tetrameric enzyme LDH presented five bands in muscle tissues, due to the expression of two locı, Ldh-A and Ldh-B, and three interloci heterotetramers. In heart only one band occurs, apparently corresponding to the A,B, heterotetramer. A weak expression of heterotetramers and the absence of homotetramers in the three species studied were observed in liver tissues. In L. anus three weak bands (A3B,, A,B, and A,B;), were observed in liver, while in L. platymetopon and Loricariichthys sp. a two banded pattern (A,B, and AjB3) was observed. The dimeric enzyme ICDH presented two loci with different tissue-expression for the three species. Locus mIcdh-A expressed itself in heart and muscle and was absent in liver tissues. Locus sIcdh-A presents bands in liver tissues only. The dimeric enzyme MDH presents three loci for Loricariichthys. Locus sMdh- A has a greater intensity of expression in heart, and weaker intensity in liver and muscle tissues. mMdh-A locus expresses itself only in liver of L. anus and Loricariichthys sp. In L. platymetopon, despite a better resolution of the bands in liver, weak bands are also observed in heart and muscle tissues. sMdh-B locus expresses itself with greater intensity in heart and muscle tissues than in the liver. Table 2 shows allelic frequencies of the three analyzed species. Loricariichthys anus is polymorphic at loci Adh-A, G3pdh-A, and Gpi-B; L. platymetopon is polymorphic at Gpi-A, Gpi-B, and sMdh-A, and Loricariichthys sp. is polymorphic at Gcdh-A, Gpi-A, and Pgm-A. Of the enzymes studied, eleven loci were diagnostic for Fic. la. Schematic representation of enzymatic systems AAT, ACP, ADH, EST, GCDH, G3PDH, G6PDH and GPI of L. anus (La), L. platymetopon (Lp) and Loricariichthys sp. (Ls), and their genetic interpretation. =e | mM Eu mi sAat-A | mM mem | ALLOZYME DISCRIMINATION OF LORICARIICHTHYS | | | NANTERRE yw—_—_ OP >—— La Lp Ls TESA La up. Ls ADH EST mr Lo i si CJ CD) } Est-A Adh-A a || } + ONE CEE A em” La Lp Ls | GCDH = |_| \ m Gcdh-A db Se, | | La Lp___Ls La Lp ES G6PDH GPI momo) ss) } | E Gpi-A | = m | momo o Omm = = = G6pdh-A mo com = © "topa gc | + + 0 667 668 C.H. ZAWADZKI, R.E. REIS & E. RENESTO ICDH | LDH | | a = = = Ldh-A = pacata = n = = = D = = M = } nica om zum mm m | = = = mo m = = = hums | si sla | HLMHLMHLM. HLMHLMHLM. wr m La Lp TESA La Lp Ls MDH MDHP Eu | | mE } asm ES mm | u u u u u u mo ww | mMdhp-A | EE m 5 sMdh-A = os = mMdh-A zz mom mmm at Ot Fe «MALA = pra Ù + 0 CSA DA IATA MIEI HLMHLMHLM. HLMHLMHLM. La Lp Ls La Lp Ls = PGM SOD IS eS a eee sod-At mM mu mn mmm = == pres + ft HLMHLMHLM. HLMHLMHLM- Se a nt )»—> ut met et m ut rar up LS RE TONER MER EPS Fic. 1b. Schematic representation of enzymatic systems ICDH, LDH, MDH, MDHP, PGM and SOD of L. anus (La), L. platymetopon (Lp) and Loricariichthys sp. (Ls), and their genetic interpretation. one or another of the three species. Loci sAat-A, Acp-A, Est-A, sIcdh-A, sMdh-A, and sMdhp-A are diagnostic for L. platymetopon, that is, they are fixed for different alleles. Loci sAat-A, mIcdh-A, Ldh-A, Ldh-B, and sMdh-B are diagnostic for L. anus and the loci sAat-A and G6pdh-A are diagnostic for Loricariichthys sp. The genetic variability for the three species of Loricariichthys is presented in Table 3. All the polymorphic loci analyzed were in Hardy-Weinberg equilibrium. Unbiased genetic identity ( I ) and genetic distance ( D ) of Nei (1978) among the three species is represented in Table 4. ALLOZYME DISCRIMINATION OF LORICARICHTHYS 669 TABLE 2. Allelic frequencies observed for 21 loci in L. anus, L. platymetopon and Lorica- riichthys sp. EOEUS ALLELE L. anus L. platymetopon Loricariichthys sp. (n = 8) (n = 16) (= 13) sAat-A a ---- 1,000 ---- b ---- = 1,000 G 1,000 ---- ---- Acp-A a ---- 1,000 ---- b 1,000 ---- 1,000 Adh-A a 0,875 ---- 1,000 b 0425 1,000 ---- Est-A a ---- 1,000 ---- b 1,000 ---- 1,000 Gcdh-A a ---- 1,000 0,308 b 1,000 ---- 0,692 G3pdh-A a 0,063 ---- ---- b 0,938 1,000 1,000 G3pdh-B a 1,000 1,000 1,000 G6pdh-A a ---- ---- 1,000 b 1,000 1,000 ---- Gpi-B a ---- 0,906 ---- b 0,938 0,094 1,000 c 0,063 = ---- Gpi-A a ---- 0,125 0,885 b 1,000 0,500 0,115 c ---- 0,375 ---- micdh-A a ---- 1,000 1,000 b 1,000 ---- ---- sIcdh-A a ---- 1,000 ---- b 1,000 ---- 1,000 Ldh-B a 1,000 ---- ---- b ---- 1,000 1,000 Ldh-A a 1,000 ---- ---- b ---- 1,000 1,000 sMdh-A a ---- 0,156 ---- b ---- 0,844 ---- c 1,000 --- 1,000 mMdh-A a 1,000 1,000 1,000 sMdh-B a 1,000 ---- ---- b ---- 1,000 1,000 sMdhp-A a ---- 1,000 È b 1,000 ---- 1,000 mMdhp-B a 1,000 1,000 1,000 Pgm-A a 1,000 ---- 0,962 b ---- 1,000 0,038 Sod-A a 1,000 1,000 1,000 DISCUSSION The usefulness of this technique as a tool for discrimination of similar species is confirmed, especially for fish systematists, whose morphological criteria are, not always, fully efficient to discriminate taxa. Isozyme electrophoresis constitutes a practical tool, given the easiness of its application, relative little need of specialized equipment, redu- ced cost, and the ability of obtaining data in a relatively short time (Buth & Murphy, 1999). 670 C.H. ZAWADZKI, R.E. REIS & E. RENESTO TABLE 3. Genetic variability measures for 21 loci of three Loricariichthys species. Number in parentheses are standard deviation. N = number of analyzed specimens; K = mean number of alleles per locus; Pg 99 = frequency of polymorphic loci; Ho = mean observed heterozygosity; He = mean expected heterozygosity. Species n k Po 99 Ho He L. anus 8 1.1 (0.1) 14.3 0.024 (0.014) 0.023 (0.013) L. platymetopon 16 122 (Ol) 14.3 0.039 (0.028) 0.050 (0.032) Tal Loricariichthys sp. 13 0.1) 14.3 0.015 (0.009) 0.035 (0.023) TABLE 4. Nei’s unbiased genetic identity / is shown above the diagonal and genetic distance D is shown below the diagonal. Species L. anus L. platymetopon Loricariichthys sp. L. anus For 0.329 0.663 L. platymetopon 1 Fer 0.478 Loricariichthys sp. 0.411 0.738 HO Various authors have demonstrated the value of data obtained through electro- phoresis of enzymes for systematics and taxonomy of fish (Avise, 1974, 1994; Buth, 1984; Buth & Murphy, 1999; Ward & Grewe, 1995 and references therein). Besides the use of allozyme data as diagnostic characters, Buth & Murphy (1999) emphasize the importance of other electrophoretic characteristics such as tissue activity, hetero- polymer formation, differences in ontogenetic expression and post-translational modi- fications. According to Kettler et al. (1986) differences in the expression of a given enzyme in different tissues (differential tissue-specific expression, different tissular ontogenetic changes or heteropolymer fomation...) can be treated, as a set of characters for taxonomical studies, independently of the analysis of different enzymes in a given tissue (loci polymorphism, heterozygosity, presence/absence of diagnostic locı...). The patterns of tissue- specific isozyme distribution is an indirect evidence of functional divergence (Basaglia, 1991a) and interspecific differences in these patterns can reveal informative evolutionary aspects of the gene-expression divergence in phylogenetic studies (Whitt, 1983; Basaglia, 1991b). For constructing a robust data set of tissue-expression one must get information about the activity of a given enzyme through the greatest possible number of tissues, and principally, in many species within a taxon. Since the data obtained here forms a part of a broader investigation on the family Loricariidae, containing many other species (Zawadzki, in preparation), the three tissues presented were found to be the most informative in regard to the number of loci, within the limits of time and money of the whole research. Recent allozyme studies have been developed at population and species levels for neotropical fishes for example Fenerich-Verani er al. (1990a, 1990b); Almeida Val et al. (1990, 1992); D’ Avila Ferreira er al. (1991); Monteiro et al. (1991); Renesto & Zawadzki (1997): Revaldaves et al. (1997); Almeida & Sodré (1998); Chiari & Sodré (1999); Lapenta et al. (1999); Zawadzki et al. (1999a, 1999b). Nevertheless, to the ALLOZYME DISCRIMINATION OF LORICARIICHTHYS 671 present, no studies was conducted on the genus Loricariichthys preventing detailed comparison. Here, two systems, MDH and LDH, present differences among the three species of Loricariichthys in relation to the patterns of tissue activity. Locus mMdh-A is expressed as an intense band in the liver of the three species and weak bands in the heart and muscle of L. platymetopon; while in the other two species its expression is limited to the liver. The LDH isozyme in liver tissue shows differences in relation to the heterotetramer interloci formation in these species of Loricariichthys, with a three banded pattern in L. anus and a two banded pattern in the other two species. In muscle tissue, all three species show a strongly expressed five banded pattern. Zawadzki er al. (in press) found in species of Hypostomus a single banded pattern in muscle and heart tissues corresponding to the homotetramer A, for H. aff. commersonii and H. myersi and to homotetramer B, for H. derbyi. Buth & Murphy (1999) state that multimeric and multiloci enzymatic systems can vary with regard to the production of interlocus heteropolymers and that this variation when established among the species can have systematic value. Murphy (1988) states that for fishes, there is a "general" trend in the LDH heteropolymer restriction, from the primitive five banded pattern to the advanced two banded A,B, pattern condition. Thus, the LDH patterns observed for these species seem to corroborate the more basal position of Loricariinae relative to Hypostominae, or at a least to Hypostomus species, shown in recent phylogenetic analysis of the family (Schaefer, 1986, 1987; Montoya-Burgos er al., 1997, 1998). Although, these enzymatic findings are not conclusive, it shows an open branch to one questioning the applicability of LDH isozymes in assaying Loricariidae phylogeny. The presence of a single Esterase locus in these three species of Loricariichthys contrasts markedly with the six loci found by Lapenta ef al. (1999) in Hypostomus albopunctatus. Duplication or silencing loci events in this enzymatic system can have occurred in the Loricariidae family and are also to be further investigated. The heterozygosity found in the three species of Loricariichthys agree with the average 0.051, calculated by Ward er al. (1992) for freshwater fishes. The values are 0.023 for L. anus, 0.050 for L. platymetopon, and 0.035 for Loricariichthys sp. The greater values for the last two species could be explained by a more heterogeneous area sampled. However, Zawadzki et al. (1999b), studying sympatric loricariid catfish species of the genus Hypostomus in the Itaipu Hydroelectric Reservoir, found low hete- rozygosity for H. albopunctatus (0.015) and H. microstomus (0.016) and high hetero- zygosity for H. regani (0.103) and H. ternetzi (0.091), suggesting the predominance of historical factors rather than sampling artifacts in the determination of the genetic variability in some species of the family Loricariidae. The unbiased genetic identity (I) (Nei, 1978), among the three species, is low. According to Thorpe (1982) and Thorpe & Solé-Cava (1994) 85% of / values between pairs of congeneric species exceed 0.35, and 76% of the values are above 0.4, while 77% of the values of / between different genera are below 0.35. Thus, the genetic identity between L. platymetopon and L. anus is as small as that between two distinct genera. The allozyme discrimination among the three species of Loricariichthys was more efficient, in number of diagnostic characters, than the morphologic discrimination 672 C.H. ZAWADZKI, R.E. REIS & E. RENESTO of the same three species, as presented by Reis & Pereira (in press). Those authors found three morphological characters to discriminate between L. platymetopon and the new species, and two additional characters to separate the new species from the other two. From these allozymatic diagnostic characters and from that obtained by Zawadzki et al. (1999a) which found allozyme genetic markers for three species of Hypostomus from the rio Iguacu basin, we suggest the continuous use of allozyme electrophoresis as an auxiliary tool for the taxonomy of the family Loricariidae. 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Differential expression for tissue-specific isozymes in three species of Hypostomus Lacépéde, 1803 (Teleostei: Loricariidae). Biochemical Systematics and Ecology. REVUE SUISSE DE ZOOLOGIE 107 (4): 675-720; décembre 2000 Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida: Dampfiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI) Sandor MAHUNKA Zoological Department, Hungarian Natural History Museum, Baross utca 13, H-1088 Budapest, Hungary. Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida: Damp- fiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI). - Thirty-two species belonging to the superfamily Otocepheoidea Balogh, 1972 are identified and listed, twenty-one of them are described as new to science: three new species belong to the family Dampfiellidae Balogh 1961, eighteen new species to the family Otocepheidae Balogh, 1961. Key-words: Acari - Oribatida: Otocepheoidea - taxonomy - new species - Malaysia: Sabah. INTRODUCTION The Oribatid fauna of Sabah, formerly North Borneo, has been discussed in previous papers (Mahunka, 1987a, 1987b, 1988, 1990, 1991, 1995a, 1996a, 1996b). In most of my earlier papers I described various species phylogenetically far distant from each other. Later I realized that on this basis no regional zoogeographical evaluation could be made. So I decided to study larger taxonomic units. In a recent contribution I discussed the taxa belonging to the family Galumnidae (Mahunka, 1995a). Herein I propose a survey of the Sabah species belonging to the superfamily Otocepheoidea Balogh, 1972 which was earlier excellently elaborated in different ways by Grandjean (1964), Aoki (1965, 1967), Hammer (1971, 1979) and Wallwork (1962a, 1962b). The material examined yielded 32 species, of which 21 are new to science. After the present studies, I am not sure if the present classification of Otoce- pheidae Balogh, 1961 is correct or if further studies are necessary to improve it. Some genera of Otocepheinae (e.g. Acrotocepheus Aoki, 1965, Megalotocepheus Aoki, 1965 and Otocepheus Berlese, 1916) are only distinguishable by the presence or absence of prodorsal or notogastral condyles. The other main diagnostic features, such as length, size and position (in relation to the rostral setae) of the lateral lamelliform expansion and the tutorium, are very variable. Therefore all Otocepheinae species mentioned in this paper are provisionally placed in the genus Otocepheus Berlese, 1904. Manuscript accepted 08.08.2000 676 SANDOR MAHUNKA Details about the origin of the material examined have already been given and the general goal of my work has also been outlined in my earlier publications (e.g. Mahunka, 1995b). Descriptions and the terminology in this contribution follow those adopted in my previous papers. ISM OF LOCALITIES Sab-82/4: MALAISIE: Sabah (Sandakan Residency): 15 milles (24 km) W de Sandakan: Sepilok: "Kabili-Sepilok Forest Reserve", forêt pres "Orang-Utan Rehabilitation Station", prelevement de sol dans les angles formes par les contreforts ailés de grands arbres (Dipterocarpaceae), 30m: 23.1V.1982; leg. B. Hauser — (appareil Berlese à Sepilok). Sab-82/5: MALAISIE: Sabah (Sandakan Residency): 15 milles (24 km) W de Sandakan: Sepilok: "Kabili-Sepilok Forest Reserve", forét pres "Orang-Utan Rehabilitation Station", préle- vement de bois pourri, Lowland Dipterocarp Forest, 30m; 23.1V.1982; leg. B. Hauser — (appareil Berlese a Sepilok). Sab-82/15: MALAISIE: Sabah (West Coast Residency): Mt Kinabalu: “Bukit Ular Trail” (sentier reliant "Kambarangan Road" a "Power Station"), tamisage de feuilles mortes et de bois pourri, forêt de Lithocarpus-Castanopsis; 1790m; 28.1V.1982; leg. B. Hauser — (appa- reil Winkler-Moczarski a Sepilok). Sab-82/16: MALAISIE: Sabah (West Coast Residency): Mt Kinabalu: "Bukit Ular Trail" (sentier reliant "Kambarangan Road" a "Power Station"), prélèvement de feuilles mortes, forêt de Lithocarpus-Castanopsis; 1850m; 28.1V.1982; leg. B. Hauser — (appareil Berlese à Sepilok). Sab-82/23: MALAISIE: Sabah (West Coast Residency): Mt Kinabalu: "Summit Trail" (sentier reliant "Power Station" au sommet), forét brumeuse, prélèvement de sol au pied de plusieurs arbres, 2480m; 30.1V.1982; leg. B. Hauser — (appareil Berlese a Sepilok). Sab-82/27: MALAISIE: Sabah (Sandakan Residency): 15 milles (24 km) W de Sandakan: Sepilok: "Kabili-Sepilok Forest Reserve", forêt près "Orang-Utan Rehabilitation Station", tamisage de feuilles mortes et de bois pourri prélevés dans les angles formés par les contreforts ailés de grands arbres, Lowland Dipterocarp Forest, 30m; 3.V.1982; leg. B. Hauser — (appareil Winkler-Moczarski a Sepilok). Sab-82/34: MALAISIE: Sabah (Sandakan Residency): 15 milles (24 km) W de Sandakan: Sepilok: "Kabili-Sepilok Forest Reserve", forêt pres "Orang-Utan Rehabilitation Station", sentier menant vers la mangrove, prélèvement de sol dans le pré autour du "Cottage" (ancienne plantation d’hévéas): 7.V.1982; leg. B. Hauser — (appareil Berlese a Sepilok). Sab-82/41: MALAISIE: Sabah (Sandakan Residency): 15 milles (24 km) W de Sandakan: Sepilok: "Kabili-Sepilok Forest Reserve", forét pres du "Pond" (étang formant la réserve d’eau pour Sepilok), prélèvement de feuilles mortes, Secondary Lowland Forest; 10.V.1982; leg. B. Hauser — (appareil Berlese a Sepilok). Sab-82/45: MALAISIE: Sabah (Interior Residency): route de Kimanis, à 16 milles de Keningau: héliport, prélevement de feuilles mortes, forét brumeuse, 1380m; 12.V.1982; leg. B. Hauser — (appareil Berlese à Genève, Suisse). Sab-82/50: MALAISIE: Sabah (Interior Residency): route de Kimanis, a 10 milles de Keningau: prelevement de sol dans les angles formés par les contreforts ailés d’un arbre mort, 1170m; 13.V.1982: leg. B. Hauser — (appareil Berlese à Genève, Suisse). EIST OF IDENTIFIED SPECIES Dampfiellidae Balogh, 1961 Dampfiella kinabalu sp. n. Locality: Sab-82/15. ORIBATIDS FROM SABAH VIII 677 Dampfiella nebulosa sp. n. Locality: Sab-82/23. Dampfiella sepilok sp. n. Locality: Sab-82/41. Otocepheidae Balogh, 1961 Archegotocepheus singularis Mahunka, 1988 Localities: Sab-82/27: 7 specimens; Sab-82/41: 5 specimens. Distribution: Sabah (known from the type localities only); second record. Borneremaeus hauseri Mahunka, 1991 Locality: Sab-82/15: 16 specimens. Distribution: Sabah (known from the type locality only); second record. Bulbocepheus hauserorum Mahunka, 1988 Locality: Sab-82/43: 15 specimens. Distribution: Sabah (known from the type locality only); second record. Dolicheremaeus bruneiensis Aoki, 1967 Localities: Sab-82/4: 5 specimens; Sab-82/5: 2 specimens; Sab-82/41: 2 specimens. Distribution: Brunei (Aoki, 1967), Sabah (Mahunka, 1991); second record for Sabah. Dolicheremaeus cicatrichosus Mahunka, 1991 Localities: Sab-82/27: 4 specimens; Sab-82/43: 3 specimens. Distribution: Sabah (known from the type locality only); second record. Dolicheremaeus claviger sp. n. Locality: Sab-82/15. Dolicheremaeus fujikawae sp. n. Localities: Sab-82/15; Sab-82/16; Sab-82/27. Dolicheremaeus krantzi sp. n. Localities: Sab-82/4; Sab-82/15; Sab-82/16; Sab-82/43; Sab-82/45; Sab-82/50. Dolicheremaeus luxtoni sp. n. Locality: Sab-82/50. Dolicheremaeus punctatus sp. n. Locality: Sab-82/15. Dolicheremaeus sabahnus Mahunka, 1988 Localities: Sab-82/27: 8 specimens; Sab-82/41: 3 specimens; Sab-82/50: 1 specimen. Distribution: Sabah (known from the type localities only); second record. Dolicheremaeus sulcatus sp. n. Locality: Sab-82/4. Dolicheremaeus yoshii sp. n. Locality: Sab-82/4. Ikarotocepheus alatus Mahunka, 1988 Localities: Sab-82/43: 2 specimens. Distribution: Sabah (known from the type locality only); second record. Leptotocepheus orientalis Mahunka, 1988 Localities: Sab-82/4: 7 specimens; Sab-82/5: 1 specimen; Sab-82/27: 5 specimens; Sab- 82/34: 2 specimens; Sab-82/41: 17 specimens; Sab-82/43: 21 specimens; Sab-82/45: 2 specimens. Distribution: Sabah (known from the type locality only); second record. Otocepheus bajau sp. n. Locality: Sab-82/4: 6 specimens. Otocepheus berndhauseri sp. n. Locality: Sab-82/15. 678 SANDOR MAHUNKA Otocepheus burckhardti (Mahunka, 1987a) Localities: Sab-82/27: 3 specimens; Sab-82/41: 8 specimens. Distribution: Sabah (known from the type locality only); second record. Otocepheus heterosetiger (Aoki, 1965) Localities: Sab-82/27: 1 specimen; Sab-82/41: 1 specimen. Distribution: Thailand; first record for Sabah. Otocepheus kadazan sp. n. Locality: Sab-82/15. Otocepheus keningau sp. n. Locality: Sab-82/43. Otocepheus lienhardorum sp. n. Localities: Sab-82/15; Sab-82/16. Otocepheus nepenthes Sp. n. Locality: Sab-82/34. Otocepheus orangutan sp. n. Locality: Sab-82/27. Otocepheus philippinensis Aoki, 1965 Localities: Sab-82/27: 2 specimens; Sab-82/50: 4 specimens. Distribution: Philippines; first record for Sabah. Otocepheus rafflesiae sp. n. Locality: Sab-82/27. Otocepheus reniformis sp. n. Locality: Sab-82/15. Otocepheus spatulatus sp. n. Locality: Sab-82/50. Otocepheus verrucosus sp. n. Locality: Sab-82/15. DESCRIPTIONS AND REMARKS Dampfiella kinabalu sp. n. Figs 1-10 Material examined: Sabah: ol pe: Sab-82/15, 50 paratypes from the same sample. Holotype and 30 paratypes: MHNG!; 20 paratypes (1498-PO-94): HNHM*. Measurements: Length of body: 458-667 um; width of body: 180-250 um. Prodorsum: Rostrum wide, rostral setae simply arched, shorter than the geni- culate lamellar ones, all finely ciliate. Interlamellar setae shorter, exobothridial setae minute or represented only by their alveoli, arising very close to the bothridium. Sensillus (Fig. 9) comparatively long, with a well developed apex (often broken) and some small spicules on its surface. The median spots and other irregular sculptures of the prodorsum - typical for this genus - conspicous (Fig. 6). Taenidia also clearly visible. Notogaster: Narrow, humeral apophysis well developed, with cerotegument granules. Anterolateral "cuvette" very deep. Ten pairs of short notogastral setae, 5 : MHNG: deposited in the Muséum d'histoire naturelle, Geneva. ? HNHM: deposited in the Hungarian Natural History Museum, Budapest, with identification- number of the specimens in the Collection of Arachnida. ORIBATIDS FROM SABAH VIII 679 Fics 1-5 Dampfiella kinabalu sp. n. — 1: body in dorsal aspect, 2: leg I, 3: body in ventral aspect, 4: podosoma in lateral aspect, 5: femur, genu and tibia of leg IV. 680 SANDOR MAHUNKA pairs of lyrifissures and the glandular opening visible. All notogastral setae nearly equal in size, distinctly spiculate (Fig. 1), the anterior and median setae sharply pointed, 4 pairs in posteromarginal position being as long as the others but blunt at the tip and bacilliform. Lateral part of podosoma: Pedotecta I very large, pedotecta II-III triangular, discidium rounded, as shown in Fig. 4. Ventral region (Fig. 3): Coxisternal region well sculptured, and the epimeral borders clearly developed. Epimeral setal formula: 2(3) - 1 - 3 - 3. Excepting the setae lb, all setae short, or minute. Setae /c completely reduced or represented only by their alveoli. Setae 4c standing conspicuously far from the discidium, distance between setae 3a only slightly shorter than that between setae 3b. Anogenital setal formula : 3 - 1 - 2 - 2. Aggenital setae very short, anterior anal setae (an,) somewhat longer than the posterior ones (an,) and arising further from each other than the posterior ones. Setae ad, absent. Lyrifissures iad hardly discernible, located near to the anterior corner of the anal aperture. Gnathosoma: Labiogenal articulation conspicuous, mentum long (Fig. 7). Chelicera (Fig. 8) normal, both digits well developed. Legs: All femora ornamented by foveolae, one of the crests on their distal end clearly visible. All femora broadly, and all genua narrowly excavated basally, the inner pointed apices on genua III and IV sharply pointed (Fig. 4), the lateral ones rounded. All claws with 6-7 small dorsal teeth, on tarsi I-II absent. On tarsus I setae p', p" and s eupathidial, the others of normal type (Fig. 2). Leg setal formulae are: I: 1 -4- 341 - 442 - 16+2 - 1 IVa 812 71 Remarks: See the remarks after the following Dampfiella species. Derivatio nominis: Named after the Mt Kinabalu National Park. Dampfiella nebulosa sp. n. Figs 11-13 Material examined: Sabah: Holotype: Sab-82/23, 3 paratypes from the same sample. Holotype and 2 paratypes: MHNG; | paratype (1949-PO-94): HNHM. Measurements: Length of body: 556-667 um; width of body: 208-278 um. Prodorsum: Very similar to the preceding species, but the sensillus (Fig. 12) slightly thicker, without a sharply pointed apex, bearing some small spines of equal size on its distal margin. Notogaster: Broader than in the preceding species, but the position of the setae is quite similar. Ten pairs of notogastral setae well developed, six of them thicker and longer than the four other bacilliform ones which are present in the posteromarginal position (Fig. 11). Ventral region (Fig 13): Coxisternal region well sculptured, epimeral borders conspicuous. Epimeral setal formula: 2 - | - 3 - 3, the insertions of setae /c were not discernible. Setae /b longest of all, but some setae (e.g. 4a and 4b) much longer than in the preceding species. Legs: Similar to those of the preceding species. Remarks: The family Dampfiellidae Balogh, 1961, with its two genera Damp- fiella Sellnick, 1931 and Beckiella Grandjean, 1964, is rather inadequately known, as ORIBATIDS FROM SABAH VIII 681 10 Fics 6-10 Dampfiella kinabalu sp. n. — 6: lateral part of body in dorsal aspect, 7: gnathosoma, 8: chelicera, 9: sensillus, 10: genital plates. has been pointed out by Hammer (1979) and also by Pérez-Iñigo & Baggio (1986). The two new species (D. kinabalu and D. nebulosa) readily distinguishable by the shape of their notogastral setae should be grouped with D. angusta. In this respect D. angusta is closer to D. kinabalu, though its sensillus is smooth and much longer and narrower than that of the two new species. Furthermore, its rostrum is more angular in dorsal view. Derivatio nominis: An allusion to the cloud-forest of Mt. Kinabalu, where the type material was collected. Dampfiella sepilok sp. n. Figs 14-17 Material examined: Sabah: Holotype: Sab-82/41, 3 paratypes from the same sample. Holotype and 2 paratypes: MHNG; | paratype (1950-PO-94): HNHM. Measurements: Length of body: 501-610 um; width of body: 138-207 um. Prodorsum: Similar to the preceding species, but slightly longer. Rostral apex weakly foveolate. Rostral and lamellar setae simply arched, unilaterally ciliate. Interlamellar setae short, setiform, exobothridial setae reduced, represented only by their alveoli. Sensillus long, its head clavate, wide, with a small apex and some spi- cules on its distal end. 682 SANDOR MAHUNKA Fics 11-13 Dampfiella nebulosa sp. n. — 11: body in dorsal aspect, 12: sensillus, 13: body in ventral aspect. Notogaster: Extremely long and narrow. A well defined neck-region obser- vable (Fig. 14). Between the bothridial and the lateral condyles of the notogaster an intercondylar band visible (Fig. 15). Anterolateral cuvette small and not deep. Ten pairs of notogastral setae present, all blunt at tip and distinctly spiculate unilaterally. The distance between setae c, and la much smaller than that between /a and Im. The median notogastral setae gradually becoming longer posteriorly, setae h, being the longest of all. All four pairs of setae in posteromarginal position are of equal length. Lateral part of podosoma: Pedotecta I very large (Fig. 17). The extreme length of the body also reflected in the neck-region. Ventral region (Fig. 16): Ventral cuvette very large, strongly bordered, epi- meres I and II larger than normal. Behind the discidium a pair of deep hollows present. Epimeral setal formula: 3 - 1 - 2 - 2 (setae 4c near the discidium not discer- nible). Epimeral setae, excepting setae /b, very short, sometimes hardly observable. Genital and aggenital setae also very short, anal setae normal and adanal ones conspi- cuously long, setae ad, being the longest of all notogastral and ventral setae, thickened and spiculate like the posterior notogastral setae. ORIBATIDS FROM SABAH VIII 683 Fics 14-17 Dampfiella sepilok sp. n. — 14: body in dorsal aspect, 15: lateral part of body in ventral aspect, 16: body in ventral aspect, 17: podosoma in lateral aspect. Legs: All femora strongly foveolate, appearing to be polygonate. The protruding plates on genua II-IV are rounded. Remarks: On the basis of the 3 pairs of adanal setae, this species belongs to another species group than the two preceding ones. It is well characterised by the shape of the notogastral and ventral setae and primarily by the extremely elongated body with a narrow neck-part. On this basis this species may easily be distinguished from all heretofore known taxa. Derivatio nominis: The species 1s named after the region of Sepilok, where its type locality is situated. 684 SANDOR MAHUNKA Dolicheremaeus claviger sp. n. Figs 18-22 Material examined: Sabah: Holotype: Sab-82/15, 2 paratypes from the same sample. Holotype and | paratype: MHNG; | paratype (1953-PO-94): HNHM. Measurements: Length of body: 820-876 um; width of body: 319-362 um. Prodorsum: Rostrum wide, foveolate. Lamellae narrow, long, medially arching outwards (Fig. 18). Interlamellar region with strong sculpture consisting of foveolae of different sizes. Both pairs of prodorsal condyles well developed, rounded and slightly larger than the lateral ones. Tutorium sinuate and arched, not directed toward the insertion of rostral setae. Lateral lamelliform expansions straight, not touching each other. Rostral and lamellar setae setiform, lamellar setae clearly longest (Fig. 18). Interlamellar setae ensiform, straight, the much shorter and thinner exobothridial setae erect, all setae ciliate or spiculate. Sensillus (Fig. 21) short, its head clavate, with some minor spicules on its distal end. Notogaster: Its surface characteristically foveolate, ornamented by larger foveolae anteriorly and by smaller ones posteriorly (Fig. 18). Ten pairs of ensiform (needle-like), mostly straight notogastral setae present, they are spiculate or finely roughened. No great differences among them, but setae p, clearly shorter than p,, setae c, shorter than interlamellar setae. Lateral part of podosoma: Pedotecta I narrow, pedotecta H-III small, their shape typical for the genus. Sejugal region stronger, surface between the pedotecta I and II-III slightly foveolate. Discidium small. Setae /c arising at the basis of pedo- tecta I (Fig. 20). Ventral region: The shape of the apodemes and the epimeral borders typical for the genus, with a weak sculpture. The posterior border of this region (bo. 4) thin, like a tectum, but conspicuous. Great differences exist among the epimeral setae, they are longer on epimeres I and II than on epimeres III and IV. Setae /b, Ic, 3b long, setae 4c particularly short, their size and ratio are shown in Fig. 19. All setae strongly pilose. Ventral plate rarely foveolate. A pair of well sclerotized short laths observable at the anterior corner of the genital aperture. Anterior two pairs of genital setae short, posterior two pairs long. These and the aggenital setae are fine, setiform, adanal ones blunt at tip, similar to notogastral ones. Setae ad, located closer to the anal aperture than to the lateral margin of the ventral plate. Lyrifissures lad in inverse apoanal position. Legs: All tarsi with dorsal teeth, claws are spiculate dorsally. Type of ultimate setae: L - ? - ? - S. Setae v"of tibia and pv"of tarsus plumose. Remarks: The new species is characterised by the great differences among the genital setae and the form of the very short, clavate sensillus, by its characteristic notogastral sculpture, the form of its lateral condyles and the form of its notogastral setae. On this basis it may be distinguished from all the heretofore known Doli- cheremaeus species. Derivatio nominis: The species is named after the form of its sensillus. ORIBATIDS FROM SABAH VIII 685 Fics 18-20 Dolicheremaeus claviger sp. n. — 18: body in dorsal aspect, 19: body in ventral aspect, 20: podosoma in lateral aspect. 686 SANDOR MAHUNKA co.nm. Fics 21-22 Dolicheremaeus claviger sp. n. — 21: sensillus and the lateral condyles, 22: leg I. Dolicheremaeus fujikawae sp.n. Figs 23-27 Material examined: Sabah: Holotype: Sab-82/15, 1 paratype from the same sample; 1 paratype: Sab-82/16; 2 paratypes: Sab-82/27. Holotype and 2 paratypes: MHNG; 2 paratypes (1960-PO-94): HNHM. Measurements: Length of body: 331 um; width of body: 180 um. Prodorsum: Lamellae long, gradually converging anteriorly, reaching beyond the insertion of the lamellar setae but not touching them. Surface weakly ornamented, but covered by cerotegument granules. Tutorium hardly discernible, lateral lamelli- form expansion short, straight. Both pairs of median condyles very large, median one more rounded, lateral one slightly angulate (Fig. 27). Under the median condyles a transversal band observable in a deeper layer. Rostral and lamellar setae long, setiform, weakly ciliate. Interbothridial and exobothridial setae bacilliform, blunt at tip, their surface only slightly roughened. Head of sensillus asymmetrically dilated, with short spines or spicules on its distal end. Notogaster: Surface rarely foveolate. Both pairs of notogastral condyles conspicuously large. Ten pairs of short and mostly bacilliform notogastral setae pre- sent, they are nearly equal in size and length (Fig. 23), their surface slightly roughened. Five pairs of conspicous lyrifissures, ih and ips located anteriorly to seta hg (Fig. 25). ORIBATIDS FROM SABAH VIII 687 Fics 23-27 Dolicheremaeus fujikawae sp. n. — 23: body in dorsal aspect, 24: body in ventral aspect, 25: notogaster in lateral aspect, 26: podosoma in lateral aspect, 27: lateral part of the dorsosejugal region. 688 SANDOR MAHUNKA Lateral part of podosoma: Pedotecta II-II well, discidium hardly developed. Sejugal region distinctly pustulate (Fig. 26). Ventral region: Coxisternal and ventral plates somewhat sculptured, the apodema and the epimeral borders mostly observable. Bo. 4 only partly visible, interrupted. A well-developed longitudinal lath parallel with the genital aperture present. All setae in the ventral regions short and simple (Fig. 24), anal setae longer than the longest epimeral ones. Lyrifissures iad in paraanal position. Legs: Tarsus of leg I without triangular teeth. Type of ultimate setae: L - L - L - L, but these setae (u) very short and fine. No dilated setae on tibia IV and tarsus IV. Remarks: The new species is well characterised by the form of its sensillus, the position of the notogastral lyrifissures (ih and ips) and the short notogastral setae of equal length. On this basis it belongs to the species group of D. variolatus Mahunka, 1989. The new species may be distinguished from the latter by its peculiarly large condyles on the prodorsum and on the notogaster, and by the much shorter adanal setae. Derivatio nominis: I dedicate the new species to Dr. T. Fujikawa, the renow- ned Oribatidologist, for her excellent studies on the Japanese oribatids. Dolicheremaeus krantzi sp. n. Figs 28-31 Material examined: Sabah: Holotype: Sab-82/15, 10 paratypes from the same sample; 3 paratypes: Sab-82/4; 2 paratypes: Sab-82/16; 9 paratypes: Sab-82/43; 1 paratype: Sab-82/45; 3 paratypes: Sab-82/50. Holotype and 17 paratypes: MHNG; 11 paratypes (1952-PO-94): HNHM. Measurements: Length of body: 457-512 um; width of body: 226-294 um. Prodorsum: Rostrum widely rounded, lamellae long, slightly and gradually converging anteriorly. Prodorsal surface mostly smooth, with some large foveolae and wrinkles along the lamellae in the interlamellar region. Median prodorsal condyles distinct, the lateral ones weakly developed (Fig. 30). Rostral and lamellar setae setiform, both distinctly ciliated. Interlamellar setae ensiform and exobothridial setae needle-shaped, surface of the former smooth, the latter strongly spiculate. Sensillus long, with slightly lanceolate head. Notogaster: Whole surface covered by large, partly rounded, partly elongated pustules with a medial split. Notogastral condyles well developed, median pair rounded, lateral ones triangular. Great differences in length among notogastral setae. Setae c, and A, short, bacilliform, all others setiform or filiform, much longer than the preceding two pairs (Fig. 28). Lateral part of podosoma: Pedotecta I normal, pedotecta I-III well developed. Discidium with a conspicuously protruding posterior corner (Fig. 31). Ventral region (Fig. 29): Among the apodemes ap. 2, ap. sej. and ap. 3 distinct, ap. 4 weakly developed, bo. 4 only partly observable. Far laterally along the genital aperture a longitudinal crest present. Epimeral setal formula normal (3 - | - 3 - 3), setae 1b, 3b and 4b longer than the others. Ventral plate foveolate. Aggenital, anal and adanal setae fine, shorter than most of the notogastral ones. Lyrifissures iad in inverse apoanal position. ORIBATIDS FROM SABAH VIII 689 N SÌ USI 4 Exe NE Sa AN Fics 28-31 Dolicheremaeus krantzi sp. n. — 28: body in dorsal aspect, 29: body in ventral aspect, 30: lateral part of the dorsosejugal region, 31: podosoma in lateral aspect. 690 SANDOR MAHUNKA Legs: Tarsus of leg I without triangular teeth. Type of ultimate setae: L - L - L - L. Setae pv" of tibia and v" of tarsus plumose. Remarks: The new species is well characterised and well distinguished from all heretofore known Dolicheremaeus species by its characteristic sculpture on the noto- gaster and the form of its notogastral setae. Derivatio nominis: I dedicate the new species to Prof. Dr. W. Krantz (Cor- vallis, USA), an acarologist of great renown. Dolicheremaeus luxtoni sp. n. Figs 32-35 Material examined: Sabah: Holotype: Sab-82/50, 8 paratypes from the same sample. Holotype and 5 paratypes: MHNG; 3 paratypes (1966-PO-94): HNHM. Measurements: Length of body: 583-738 um; width of body: 264-390 um. Prodorsum: Rostrum smooth. Lamellae very long and narrow, running parallel to each other. Only one pair of notogastral condyles present; the median one comple- tely lacking, whereas the lateral one enlarged and very wide (Fig. 34). Tutorium hardly developed, the lateral lamelliform expansion clearly developed, directed to- ward the insertion of rostral setae but not reaching it (Fig. 35). Rostral and lamellar setae setiform, strongly ciliate. Interlamellar setae ensiform. Sensillus with asymme- trically dilated head, its distal end bearing some spicules. Exobothridial setae straight, short. Notogaster: Its surface ornamented mostly by foveolae, but posterior part clearly pustulate. Two pairs of large notogastral condyles present, co. nl. and co. mn. located conspicuously near to each other (Fig. 32), the median one is elongated. All ten pairs of notogastral setae ensiform, no great differences in length and size among them. Lyrifissures ips open between setae p; and h3. Lateral part of podosoma: All pedotecta normal in shape, as shown in Fig. 35. Sejugal region well pustulate. Ventral region (Fig. 33): Epimeral borders and the usual apodemes conspi- cuous. Epimeral setae short, no obvious differences among setae /b, Ic, 3b, 3c. Setae 4b the longest of all, setae /a, 2a, 3a and 4a very short. Ventral plate foveolate, surface of genital and anal plates smooth. Genital and aggenital setae short and simple, anal ones and especially adanal ones much longer and stronger. Setae ad, arising far laterally, near to the margin of ventral plate. All setae slightly aciculate. Lyrifissures iad in adanal position. Legs: Type of ultimate setae: L - S - S - S. No triangular teeth on tarsi I-IV. Seta v" on tibia and pv" on tarsus plumose. Remarks: Owing to the absence of the median prodorsal condyles and the posi- tion of adanal setae and lyrifissures, the new species strongly resembles D. baloghi Aoki, 1967. However, the latter is well distinguished from the new species by the sculpture of the notogaster (the posterior part also foveolate) and by the length of the interlamellar setae (short in D. baloghi). The form of the notogastral condyles also differs. Derivatio nominis: I dedicate the new species to Dr. M. Luxton, the renowned specialist of the world oribatids, as a grateful acknowledgement of his valuable help in the revision of my manuscripts. ORIBATIDS FROM SABAH VIII 69] At À Canon Fics 32-35 Dolicheremaeus luxtoni sp. n. — 32: body in dorsal aspect, 33: body in ventral aspect, 34: dorsosejugal region, 35: podosoma in lateral aspect. 692 SANDOR MAHUNKA Dolicheremaeus punctatus sp. n. Figs 36-39 Material examined: Sabah: Holotype: Sab-82/15, 23 paratypes from the same sample. Holotype and 14 paratypes: MHNG; 9 paratypes (1954-PO-94): HNHM. Measurements: Length of body: 847-1279 um; width of body: 389-667 um. Prodorsum: Rostrum slightly foveolate, prodorsal surface simple, only the exobothridial and sejugal region strongly pustulate laterally. Lamellae long and narrow, slightly bent inwards distally. Two pairs of prodorsal condyles present, median one regularly rounded and much larger than the slightly angulate lateral pair (Fig. 37). Rostral and lamellar setae arising relatively near to each other, both pairs setiform and strongly ciliate, with filiform distal end. Interlamellar setae ensiform, straight, slightly spiculate. Exobothridial setae short, sensillus with a slightly dilated lanceolate head. Notogaster: Ornamented with small but deep, well-framed foveolae (nearly punctate). Two pairs of notogastral condyles present, but the median pairs weakly developed, not reaching beyond the dorsosejugal suture (Fig. 36). The lateral condyles triangular in shape. Ten pairs of notogastral setae present, all ensiform, similar to the interlamellar ones, without essential difference between them. Lyri- fissures ih and ips located in front of setae h-. Lateral part of podosoma: Pedotecta I and II-III normal, discidium small (Fig. 39). Tutorium long, well arched, undulate dorsally, not connected with the also long and reversed S-shaped lateral lamelliform expansion. Ventral region (Fig. 38): Epimeral surface rarely, ventral plate distinctly foveolate, similar to the notogastral surface. Epimeral setae relatively long, setae Jc arising anteriorly, setae 4b and 4c located far from each other. All setae, excepting the adanal ones, in the anogenital region (e.g. genital ones) unusually long, setiform. Adanal setae ensiform like the notogastral setae, shorter than the anal setae. Setae ad, slightly longer than ad;, the former arising closer to the anal aperture than to the lateral margin. Lyrifissures iad in inverse apoanal position. Legs: Tarsi of all legs bearing sharp teeth. Type of ultimate setae: L - S - S-S. Remarks: The new species is well characterised by the strong (deep) punc- tation resembling a foveolate sculpture, by the equal length and size of the notogastral setae and primarily by the ratio of the anogenital setae (i.e. anal setae longer than the adanal ones). Derivatio nominis: The species is named after the punctate surface of the notogaster. Dolicheremaeus sulcatus sp. n. Figs 40-47 Material examined: Sabah: Holotype: Sab-82/4, 5 paratypes from the same sample. Holotype and 3 paratypes: MHNG; 2 paratypes (1951-PO-94): HNHM. Measurements: Length of body: 556-602 um; width of body: 272-300 um. Prodorsum: Surface smooth, only the surface of pedotecta weakly foveolate and the sejugal region pustulate (Fig. 40). Some very fine interlamellar spots visible, arranged in longitudinal rows. Lamellae very long, with well separated distal part, like ORIBATIDS FROM SABAH VIII 693 Fics 36-39 Dolicheremaeus punctatus sp. n. — 36: body in dorsal aspect, 37: dorsosejugal region, 38: body in ventral aspect, 39: podosoma in lateral aspect. cusps, running parallel to each other. The median prodorsal condyles connected with them, both pairs of condyles very large, the outer ones rectangular, the inner ones triangular (Fig. 42). Lamellar and rostral setae setiform, strongly ciliated. Inter- lamellar setae ensiform, exobothridial setae arising at the basis of pedotecta I. Tuto- rium absent, lateral lamelliform expansion well developed. SANDOR MAHUNKA 694 Fics 40-43 Dolicheremaeus sulcatus sp. n. — 40: body in dorsal aspect, 41: body in ventral aspect, 42: dorsoseju podosoma in lateral aspect. 2. DE gal region, 4 le ORIBATIDS FROM SABAH VII 695 Notogaster: Sejugal region wide, two pairs of well-developed condyles pre- sent, median pairs conspicuously long (Fig. 40), directed backwards. Anterior and median surface sulcate, alternating with irregular alveoli, behind them is a blister-like structure, and the posterior margin again foveolate and alveolate. Ten pairs of thin and mostly long or very long notogastral setae present, some of them with filiform and flagelliform distal ends. Lyrifissures im located far anteriorly, the glandular opening placed rather anteriorly. Lyrifissures ips present between setae p, and h, (Fig 44). Lateral part of podosoma: Pedotecta I wide, pedotecta II-III rectangular, disci- dium normal, setae 4b and 4c arising close to each other on the apex of the discidium (Fig. 43). Ventral region (Fig. 41): Transversal apodemes well developed, the two ap. 2 and the two ap. sej. connected medially. Epimeral setae (3 - | - 3 - 3) mostly clearly ciliate. Ventral plate foveolate laterally, a pair of longitudinal wrinkles observable along the genital aperture. Aggenital, anal and adanal setae similar to the notogastral ones, with filiform end. Legs: Type of ultimate setae: L - L - L - L. No triangular teeth on tarsi I-IV (Figs 45-46). Seta v"on tibia and pv"on the tarsus plumose. Remarks: The new species is distinguished from all others hitherto known from the family Otocepheidae Balogh, 1961 by the the blister-like structure of the body and by the sculpture of the notogaster. Derivatio nominis: The species is named after the sulcate surface of the noto- gaster. Dolicheremaeus yoshii sp. n. Figs 48-51 Material examined: Sabah: Holotype: Sab-82/27, 20 paratypes from the same sample. Holotype and 12 paratypes: MHNG; 8 paratypes (1955-PO-94): HNHM. Measurements: Length of body: 792-1070 um; width of body: 264-459 um. Prodorsum: Rostral apex smooth, slightly excavate medially. Lamellae long, running parallel to each other, their surface foveolate. Basal part of the interlamellar region with some wrinkles. Both pairs of prodorsal condyles large, nearly triangular. Bothridium with a small dorsal, and a very large ventral squama (Fig. 48). Tutorium weak, straight, lateral lamelliform expansion very long, arching upwards, directed rather to the lamellar than the rostral setae and ending between them. Head of sensillus dilate, with a blunt distal end, the latter with some spicules. Notogaster: Sejugal region wide, with two pairs of notogastral condyles. Lateral ones triangular, median ones asymmetrical, conspicuously long (Fig. 50). Notogastral surface ornamented with very large and irregular foveolae and with some sinuate wrinkles around the median part. Only nine pairs of notogastral setae present (setae c, absent), all ensiform, finely ciliate. Lateral part of podosoma: Pedotecta I normal, pedotecta II-III and the disci- dium weakly developed. Pedotecta and the sejugal region ornamented with smaller foveolae and a field over the acetabula III-IV with larger ones (Fig. 51). Ventral region (Fig. 49): Epimeral surface lacking sculpture, apodemes and epimeral borders well developed, typical for the genus. Epimeres well framed 696 SANDOR MAHUNKA Fics 44-47 Dolicheremaeus sulcatus sp. n. — 44: posterior part of notogaster in lateral aspect, 45: tarsus and tibia of leg IV, 46: leg I, 47: trochanter, femur and genu of leg IV. medially, between them a well-defined median field visible. Among the epimeral setae (3 - 1 - 3 - 3) great differences in length, setae /b the longest of all. Ventral plate with large foveolae, as on the notogastral surface. Genital setae strong, setiform, aggenital setae longer than adanal ones. Anal setae thin, setiform, adanal ones ensiform, slightly dilated. Setae ad, arising closer to the lateral margin of the ventral plate than to the anal aperture. Lyrifissures iad in inverse apoanal position. ORIBATIDS FROM SABAH VIII 697 nu a8 ON SY SPS Sa Fics 48-51 Dolicheremaeus yoshii sp. n. — 48: body in dorsal aspect, 49: body in ventral aspect, 50: dorsosejugal region, 51: podosoma in lateral aspect. 698 SANDOR MAHUNKA Legs: Type of ultimate setae: L - S - S - S. Tarsi I-IH with triangular teeth dorsally, they are absent on tarsus [V. Leg IV with two dilated plumose ventral setae (v" and pv"). Remarks: The new species is readily distinguished from all the heretofore described species of the genus on the basis of the missing setae c;. Derivatio nominis: I dedicate the new species to the late Prof. Dr. R. Yoshii for his help in the realization of this collecting trip. Otocepheus bajau sp. n. Figs 52-55 Material examined: Sabah: Holotype: Sab-82/4, 6 paratypes from the same sample. Holotype and 4 paratypes: MHNG; 2 paratypes (1963-PO-94): HNHM. Measurements: Length of body: 1265-1501 um; width of body: 556-657 um. Prodorsum: Rostral apex and surface of lamellae weakly foveolate. Lamellae long, narrow, slightly bent medially and convergent anteriorly. Two pairs of well developed prodorsal condyles of nearly equal size, lateral one widely rounded, median one slightly triangular (Fig. 54). Bothridium with a rounded ventral plate. Rostral and lamellar setae setiform, finely ciliate, with a filiform distal end. Interlamellar setae ensiform, exobothridial one simple and short. Sensillus small, with a lanceolate head sometimes bearing some scattered spicules. Notogaster: Surface ornamented by a weak polygonal sculpture consisting of small granules. One pair of very large notogastral condyles present, with a very long outer apex, covering the rounded prodorsal one (Fig. 52). Ten pairs of long noto- gastral setae present, setae c, and la ensiform, all the others setiform. Their length gradually increasing toward the posterior part of notogaster. Setae p, = h, the longest of all. Only one lyrifissure (ih) located in front of h3, ips visible between setae p; and p>. Lateral part of podosoma: Pedotecta I and II-III very large, posterior part of the latter larger than the anterior part, its margin foveolate (Fig. 55). Discidium with a large posterior corner, carrying only one pair of setae (4c). Ventral region (Figs 53): Completely covered by cerotegument granules, on the third and fourth epimeres they compose polygonal fields, a similar formation also on the notogaster and on the ventral plate. Epimeral borders and apodemes of usual shape, epimeral setae simple. Among them setae /b, 3b and 4b longer than their corresponding outer pairs (/c, 3c and 4c). Genital setae short, simple; the aggenital, anal and adanal setae nearly equal in length, all slightly thickened basally, their ends filiform, slightly flagellate. Lyrifissures iad in inverse apoanal position. Legs: All segments of legs very long and narrow. Tarsus I with triangular teeth, latter absent from tarsi II-IV. Type of ultimate setae: L - S - S - S. Seta v" on tibia IV and pv" on tarsus IV plumose. Remarks: The new species is related to O. holtmanni Aoki, 1965. However, it is distinguished from O. holtmanni by the ratio of the epimeral setae (3c longer than 3b in O. holtmanni) and the prodorsal sculpture (lateral part of prodorsum foveolate in O. holtmanni, smooth in the new species). 699 ORIBATIDS FROM SABAH VIII \ ù ES See 1 x D ISSN i PEAS I i » >; D E == Fics 52-55 Otocepheus bajau sp. n. — 52: body in dorsal aspect, 53: body in ventral aspect, 54: dorso- sejugal region, 55: podosoma in lateral aspect. 700 SANDOR MAHUNKA Derivatio nominis: This species is named after the Bajau people inhabiting mainly the coastal area of Sabah. Otocepheus berndhauseri sp. n. Figs 56-59 Material examined: Sabah: Holotype: Sab-82/15, 2 paratypes from the same sample; 1 paratype Sab-82/23. Holotype and two paratypes: MHNG; 1 paratype (1958-PO-94): HNHM. Measurements: Length of body: 986-1098 um; width of body: 472-556 um. Prodorsum: Rostrum strongly foveolate, some foveolae visible also in the interlamellar region. A transversal band is present in front of the lamellar cusps. Lamellae long, rather wide, running parallel to each other, ending far from the rostral apex. Both pairs of prodorsal condyles well developed and rounded, the median ones not connected with each other (Fig. 58). Rostral setae comparatively short, setiform, exobothridial setae minute, hardly discernible, lamellar and interlamellar ones very long, filiform, much resembling the other notogastral setae. All setae finely and sparsely ciliate. Sensillus comparatively small, twisted backwards, its head smooth, slightly asymmetrical in lateral aspect (Fig. 59). Notogaster: Lateral notogastral condyles very large, hardly narrower than the distance between them. Ten pairs of extremely long and filiform notogastral setae present, no essential difference between them (Fig. 56). Five pairs of lyrifissures also present, their position normal (ips located between setae I and p3). Lateral part of podosoma: Pedotecta I long, pedotecta II-III large, fishtail- shaped, ornamented by foveolae or spots (Fig. 59). Discidium very large. Ventral region: In the coxisternal region only two pairs of apodemes conspi- cuous, the epimeral borders and the other apodemes poorly developed. Epimeral setae short, the short setae /b being unusual. All setae on epimere | arising in the posterior part of this field. Both setae 3a - 3b and also setae 4b - 4c arising conspicuously near to each other (Fig. 57). All epimeral setae finely ciliate or roughened. Ventral plate foveolate. Genital, aggenital and anal setae normal, setiform, adanal setae very long, filiform and flagellate at their distal end. Setae ad, arising far from the anal aperture, near to the lateral margin of the ventral plate. Lyrifissures iad in paraanal position. Legs: Type of ultimate setae: L - S - S - S. All joints of legs conspicuously long and narrow. Femur IV with very large genual lamellae. Seta /" on genua II-IV long, spiniform, setae v on femora II-III also conspicuously long. Setae v" on tibia IV and pv' on tarsus IV dilated and plumose. Remarks: The new species has an unique body chaetotaxy within this genus. On this basis it is readily distinguished from all related species. Derivatio nominis: 1 dedicate the new species to my friend Dr. B. Hauser (Geneva), the organizer of this expedition and collector of this very rich material. Otocepheus kadazan sp. n. Figs 60-63 Material examined: Sabah: Holotype: Sab-82/15, 1 paratype from the same sample. Holotype: MHNG; paratype (1961-PO-94): HNHM. Measurements: Length of body: 1014-1029 um: width of body: 403-417 um. ORIBATIDS FROM SABAH VIII 701 FIG. 56 Otocepheus berndhauseri sp. n. — 56: body in dorsal aspect. 702 SANDOR MAHUNKA Fics 57-59 Otocepheus berndhauseri sp. n. — 57: body in ventral aspect, 58: dorsosejugal region, 59: podosoma in lateral aspect. Prodorsum: Rostrum weakly foveolate, interlamellar region with some longitudinal crests. Lamellae very long, coming close to the rostrum, bent outwards medially, inwards distally (Fig. 60). Both pairs of prodorsal condyles well developed, both rounded (Fig. 62). Rostral and lamellar setae long, simply setiform, pilose, interlamellar ones ensiform, finely spiculate. Exobothridial setae short, fine, straight. Sensillus with a small lanceolate head, without any spiculae or cilia. Notogaster: Its surface well foveolate, the foveolae are arched, small and partly connected laterally. Ten pairs of notogastral setae present, six pairs ensiform, like the interlamellar ones, 4 pairs in posteromarginal position setiform, but mostly ORIBATIDS FROM SABAH VIII 703 Fics 60-63 Otocepheus kadazan sp. n. — 60: body in dorsal aspect, 61: body in ventral aspect, 62: dorso- sejugal region, 63: podosoma in lateral aspect. 704 SANDOR MAHUNKA straight, only their distal end curved. Anterior setae arising nearly in one longitudinal row. Setae h, much shorter than setae p, - p>, h; inserted much farther from p3 than the latter one from p,. Lateral part of podosoma: All pedotecta and discidium well developed, posterior margin of pedotecta I-III ornamented by foveolae (Fig. 63). Lateral lamelli- form expansion double arched between the acetabulum and the insertion of rostral setae. Ventral region: Surface of coxisternal region without any characteristic fea- tures, apodemes and epimeral borders also typical for the genus. Bo. 4 absent, close to the anterior corner of the genital aperture a short, weakly sclerotized lath observable. Epimeral setal formula: 3 - 1 - 3 - 3. Among these setae /b, 3b and 4b longer than the others, setae /c arising far laterally and posteriorly. Ventral plate foveolate. Genital setae simple and short. No setae in postanal position and setae ad, arising far from the anal apertures, laterally (Fig. 61). All adanal setae relatively long, with a filiform distal part. Lyrifissures iad in inverse apoanal position. Legs: Type of ultimate setae: L - S - S - S. Tarsus I with triangular teeth, latter absent on the other tarsi. Remarks: The new species may be clearly characterised by the form and position of the notogastral setae and the notogastral condyles. It stands nearest to O. duplicornutus (Aoki, 1965). However, its posteromarginally placed notogastral setae are much longer than in O. duplicornutus. Derivatio nominis: This species is named after the Kadazan people which lives in the Mt. Kinabalu area. Otocepheus keningau sp. n. Figs 64-67 Material examined: Sabah: Holotype: Sab-82/43, 2 paratypes from the same sample. Holotype and | paratype: MHNG; 1 paratype (19-PO-94): HNHM. Measurements: Length of body: 792-1057 um; width of body: 326-487 um. Prodorsum: Rostrum and surface of lamellae with some foveolae. Tutorium and lateral lamelliform expansion well developed. Two pairs of prodorsal condyles present, lateral pair slightly bigger than the median ones, with some very weak light spots. Median part of the interlamellar region: Fig. 66. Rostral and lamellar setae setiform, strongly ciliate unilaterally, lamellar setae geniculate, the rostrals simply arched. Interlamellar setae ensiform, short; exobothridial setae minute. Sensillus long, its head narrowly lanceolate but thin, bearing some minute spicules on its distal end (Fig. 66). Notogaster: Its surface smooth or finely punctulate. Only one pair of lateral notogastral condyles present, not highly protruding, but with a much extending margin medially. Ten pairs of very short and fine notogastral setae present, they are setiform with flagellate ends. Lyrifissures ips located between setae p and 43. Lateral part of podosoma: Tutorium and lateral lamelliform expansion connec- ted with each other, a small field near the basal part of the tutorium weakly foveolate. Sejugal region also ornamented by foveolae. Pedotecta I and II-III large, discidium also well developed, with a large corner posteriorly (Fig. 67). ORIBATIDS FROM SABAH VIII 705 7 Lu SZ Qi Ly a Ait AWN D LTT Fics 64-67 Otocepheus keningau sp. n. — 64: body in dorsal aspect, 65: body in ventral aspect, 66: dorsosejugal region, 67: podosoma in lateral aspect. 706 SANDOR MAHUNKA Ventral region (Fig. 65): Coxisternal and ventral plates lacking sculpture. Epimeral border and apodemes conspicuous. Epimeral setal formula: 3 - 1 - 3 - 3. All setae in these two regions short and simple. Setae /c placed posteriorly and laterally, 4b and 4c arising near to each other. Setae ad; situated laterally, near to margin of the ventral plate. Legs: Type of ultimate setae: L - S - S - S. Seta v" on tibia IV and pv" on tarsus IV plumose. Remarks: The new species is characterised by the fine and short notogastral setae and the flat notogastral condyles. On this basis it 1s readily distinguished from all heretofore described otocepheoid species. Derivatio nominis: This species is named after the city of Keningau. Otocepheus lienhardorum sp. n. Figs 68-71 Material examined: Sabah: Holotype: Sab-82/15, 1 paratype from the same sample: 1 paratype Sab-82/16. Holotype and | paratype: MHNG; | paratype (1960-PO-94): HNHM. Measurements: Length of body: 847-1004 um; width of body: 336-459 um. Prodorsum: The whole body is conspicuously wide. Rostral apex foveolate, other surfaces smooth or finely punctate. Lamellae long, running clearly parallel to each other. Prodorsal condyles nearly equal in size, both pairs well rounded (Fig. 70). Rostral setae setiform, with fine distal ends, lamellar ones also setiform, but their distal end blunter (Fig.68), all distinctly ciliate. Interlamellar setae ensiform, straight, exobothridial setae minute. Sensillus simple, with a smooth lanceolate head. Notogaster: Lateral margin smooth, its inner surface distinctly pustulate (Fig.68). The shape and the density of pustules varying. A pair of very large noto- gastral condyles present, their anterolateral corner conspicuously long. Ten pairs of notogastral setae present all, excepting the 4 pairs in posteromarginal position, of nearly equal length and size, all ensiform and finely roughened or spiculate. Setae p, - p3 and A; long, simple, filiform. Lateral part of podosoma: Lateral lamelliform expansion arched, tutorium long, reaching close to the preceding one (Fig. 71). All pedotecta and the discidium well developed, the fishtail-shaped pedotecta II-II ornamented by some foveolae on their posterior margin. Ventral region: Surface of epimeres without any particular sculpture. Epimeral setae normally developed but seta /a minute or absent, setae /b and 2b longer than the rest. At the anterior corner of the genital plates a minute, well sclerotized structure observable. Ventral plate foveolate. Setae in the anogenital region short, setae ad, arising laterally, slightly farther from ad, than the latter from ad). Lyrifissure tad in inverse apoanal position (Fig. 69). Legs: Type of ultimate setae: L - S - S - S. Two small teeth visible on tarsus I, they are absent from the other legs. Remarks: The new species is well characterised by its notogastral surface, thus easily distinguishable from all related taxa. Derivatio nominis: I dedicate the new species to my friends, Dr. C. Lienhard's family (Geneva). ORIBATIDS FROM SABAH VIII 707 ~~ Tt be, EEE, el 27 Da wwe Er È DAS) à MAE! x giu ae a) - d ) (pr ia 75 la) n Fics 68-71 Otocepheus lienhardorum sp. n. — 68: body in dorsal aspect, 69; body in ventral aspect, 70: dorsosejugal region, 71: podosoma in lateral aspect. 708 SANDOR MAHUNKA Otocepheus nepenthes sp. n. Figs 72-75 Material examined: Sabah: Holotype: Sab-82/34. Holotype: MHNG. Measurements: Length of body: 959 um; width of body: 383 um. Prodorsum: Rostrum slightly convex medially. Surface foveolate and/or alveolate. Lamellar surface also foveolate, interlamellar region strongly foveolate basally, with a broad border around the foveolae. Lamellae long and wide. Two pairs of prodorsal condyles present, lateral ones well rounded and fitting into the opposite notogastral condyles. Median prodorsal condyles much smaller, slightly triangulate. Rostral and lamellar setae setiform, strongly pilose, interlamellar ones ensiform, exobothridial ones comparatively long. Sensillus lanceolate (Fig. 72). Notogaster: The whole body surface covered by a thick cerotegument layer of larger or smaller granules or tubercles, giving the surface a pustulate appearance. This ornamentation is absent from the sejugal region, including a narrow band behind the notogastral condyles. One pair of very large notogastral condyles present, with an unusually excavated anterolateral part accepting the lateral prodorsal condyles (Fig. 74). Their median part well separated and projecting forwards, giving the impression of two condyles. Ten pairs of notogastral setae present, c, and la fusiform, the others setiform with a slightly dilated basal part. Their length gradually increasing poste- riorly (Fig. 72), h, and p, being the longest setae of all. Lateral part of podosoma: Pedotecta I normal, pedotecta II-III smaller than usual, the fishtail-shaped formation hardly discernible (Fig. 75). Discidium with a very large posterior corner bearing two epimeral setae (4b, 4c). Epimeral foramen conspicuous, well bordered. Ventral region: Mentum and the epimeral surface alveolate-reticulate. The cerotegument layer thick, but not so typically granulate as the notogaster or the ventral plate. Epimeral setae comparatively long, setae 2a long and arising far from each other. Setae 4b and 4c placed very close to each other, both arising on the posterior margin of the discidium. Posteromarginal ridges (postpodosomal ornamentation opp in Aoki, 1965) continuing in longitudinal crests on the ventral plate (Fig. 73) and a small node (the genital fissure iag in Aoki 1965) conspicuous. Genital plates with a well sclerotised longitudinal ridge. Surface of anal plates also with cerotegumental tubercles. All setae in the anogenital region simply setiform. Setae ad, placed far laterally. Lyrifissures iad in adanal position, near to the anal aperture. Legs: Type of ultimate setae: L - S - S - S. Seta /" of genua I-II very strong, long, spiniform. Seta v" on tibia IV and pv" on tarsus IV plumose. Remarks: On the basis of its unique ornamentation the new species is readily distinguishable from all the heretofore described otocepheoid taxa. Derivatio nominis: Reference to the pitcher plants, which have many endemic species on Mt. Kinabalu. Otocepheus orangutan sp. n. Figs 76-79 Material examined: Sabah: Holotype: Sab-82/27, 7 paratypes from the same sample. Holotype and 4 paratypes: MHNG: 2 paratypes (1962-PO-94): HNHM. ORIBATIDS FROM SABAH VIII 709 Fics 72-75 Otocepheus nephenthes sp. n. — 72: body in dorsal aspect, 73: body in ventral aspect, 74: dorsosejugal region, 75: podosoma in lateral aspect. 710 SANDOR MAHUNKA Measurements: Length of body: 708-861 um; width of body: 250-390 um. Prodorsum: Rostral apex wide, its surface foveolate. Lamellae wide and long, running parallel to each other, their surface irregularly foveolate. Interlamellar region with some crests. Two pairs of well developed, slightly angular prodorsal condyles of nearly equal size present (Fig. 78). Rostral and lamellar setae fine, setiform, unila- terally pilose. Interlamellar ones ensiform, slightly roughened. Exobothridial setae very short. Head of sensillus wide, with a stout distal end, surface distally bearing some spicules. Notogaster: One pair of large notogastral condyles with an excavate anterior margin. Notogastral surface strongly foveolate. Among notogastral setae great differences exist in length and size. Setae c, ensiform, setae la and /m reduced, only short, arched, bacilliform setae observable (Fig. 76). Setae p3 and h, also very short, but fine and setiform. The remaining setae longer than the preceding ones, all straight and ensiform. The surface of the ensiform setae roughened or ornamented with minute spicules. Lyrifissure ips located between setae p, and r3. Lateral part of podosoma: Pedotecta I normal, pedotecta II-III typically fish- tail-shaped, the margin of the latter strongly foveolate. Lateral lamelliform expansion mostly straight, directed toward the insertion of rostral setae and touching them (1970) Ventral region: Epimeral surface lacking conspicuous sculpture, apodemes and epimeral borders conspicuous. Ventral plate foveolate, but only laterally. Epimeral setal formula: 3 - 1 - 3 - 3. Among the epimeral setae great differences exist, three pairs (/b, 3b, 4b) much longer than the others, all filiform. Setae /c, 3c and 4c conspicuously short, /a and 2a minute or very short. Genital, aggenital and adanal setae fine, short and setiform. Anal setae different in length and size: an, long but fine, clearly blunt at tip, an, resembling adanal setae (Fig 77). Lyrifissures iad in adanal position. Legs: Type of ultimate setae: L - S - S- S. Triangular projection on tarsi I and II present, on tarsi III and IV absent. Seta v" on tibia IV and pv" on tarsus IV broadened, the latter twice as large as the former. Remarks: On the basis of the fine and short notogastral and the adanal setae, the new species is related to Orocepheus heterosetiger Aoki, 1965. However, the new species is distinguished from it by lacking median prodorsal condyles and the reduced la and Im setae. Derivatio nominis: The species is named after the orang-utan, the most charac- teristic mammal of Borneo. Otocepheus rafflesiae sp. n. Figs 80-83 Material examined: Sabah: Holotype: Sab-82/27, 1 paratype from the same sample. Holotype: MHNG; paratype (1965-PO-94): HNHM. Measurements: Length of body: 1292-1515 um; width of body: 542-640 um. Prodorsum: Rostral apex and lateral part of prodorsum well foveolate. Lamellae long and wide, running mostly parallel, their surfaces also foveolate. ORIBATIDS FROM SABAH VIII TANI: Fics 76-79 Otocepheus orangutan sp. n. — 76: body in dorsal aspect, 77: body in ventral aspect, 78: dorsosejugal region, 79: podosoma in lateral aspect. 712 SANDOR MAHUNKA Tutorium conspicuous in dorsal aspect, long, reaching to the lateral lamelliform expansion. Two pairs of prodorsal condyles present, outer pairs (co.pl.) angulate, median ones very small (Fig. 82). Rostral and lamellar setae setiform, with strongly ciliate filiform distal ends. Interlamellar setae ensiform, sparsely pilose, exobothridial setae minute. Sensillus very small, its head rounded, well barbed (Fig. 80). Notogaster: Form of notogaster characteristic; the anterior third being its widest part. Surface ornamented by indistinct foveolae. Marginal ridges well deve- loped. Notogaster with only one pair of large condyles, situated laterally and removed far from each other. This distance is longer than their diameter. Ten pairs of ensiform notogastral setae present, all finely spiculate. Lyrifissures ips located between setae p3 and I. Lateral part of podosoma: Pedotecta I long and narrow, pedotecta II-III very large, with foveolae on its posterolateral margin and some pustules anteriorly (Fig. 82). Sejugal region also pustulate. Tutorium long, connected with the arched lateral lamelliform expansion. The latter reaching to the insertions of the rostral setae. Ventral region (Fig. 81): Epimeral surface smooth, apodemes normal. The posterior margin of the epimeral region with a characteristic, catenate structure (opp). Epimeral setae comparatively long, well ciliate. Ventral plate rarely foveolate, genital and anal plates smooth. Genital setae strong, not smaller than the aggenital ones, but all setae in the anal region longer than the preceding ones. They are of equal length, excepting setae ad. Legs: Type of ultimate setae: L - S - S - S. Lateral setae (/") of genua I-II long, strong, spiniform. Seta v" on tibia IV and pv" on tarsus IV plumose. Remarks: The new species is readily distinguished by the shape of the notogaster, the very small and rounded sensillus and the subequal anal and adanal setae. This combination of characters is unknown among the heretofore described species. Derivatio nominis: This species is named after the largest flower of the world, which occurs in Sabah. Otocepheus reniformis sp. n. Figs 84-87 Material examined: Sabah: Holotype: Sab-82/15, 36 paratypes from the same sample. Holotype and 19 paratypes: MHNG; 12 paratypes (1957-PO-94): HNHM. Measurements: Length of body: 657-862.um; width of body: 278-403 um. Prodorsum: Finely granulated cerotegument covering nearly the whole body surface. Prodorsum rarely foveolate. Rostrum rounded, lamellae slightly convergent anteriorly, overlapping the insertion of the lamellar setae. Interlamellar region with some longitudinal wrinkles. Tutoria long, reaching to the lateral lamelliform expansions (Fig. 87). The latter directed toward the insertion of the rostral setae and touching them. Very large prodorsal condyles present (Fig. 86), lateral ones rounded, larger than the median ones. The latter condyles connected with each other, but this connection is indistinct medially. Lamellar and rostral setae setiform, distinctly pilose, interlamellar setae blunter at tip, sparsely pilose. Sensillus simple, its head lanceolate, smooth. ORIBATIDS FROM SABAH VIII 713 Fics 80-83 Otocepheus rafflesiae sp. n. — 80: body in dorsal aspect, 81: body in ventral aspect, 82: dorsosejugal region, 83: podosoma in lateral aspect. 714 SANDOR MAHUNKA Notogaster: One pair of large notogastral condyles present, opposite to both prodorsal ones. The size of the 10 pairs of notogastral setae characteristic, setae c, ensiform and acuminate, setae da clearly dilated medially, fusiform. All other setae nearly setiform, conspicuously long, excepting setae h, being only half as long as seta p, (Fig. 84), primarily on the posterior part of the notogaster. Setae r3 also much shorter than p,. Five pairs of lyrifissures well discernible, in normal position. Lateral part of podosoma: Pedotecta I large, normal, pedotecta II-III fishtail- shaped, also large and ornamented by some spots or wrinkles. Discidium normal. Ventral region: Surface weakly sculptured. Apodemes 2 and ap. sej. distinctly developed, ap. 3, ap. 4 absent or hardly discernible. Epimeral borders also only partly visible (Fig. 85). All epimeral setae short, setae /b and 3b longer than the others. Setae 4b and 4c arising close to each other. Epimeral setal formula: 3 - 1 - 3 - 3. Anogenital setal formula: 4 - 1 - 2 - 3, typical for the genus. Setae ad; only half as long as adj. Anal and adanal setae with filiform distal ends. Lyrifissures iad in inverse apoanal position. Legs: Type of ultimate setae: L - S - S - S. Seta v on genu II conspicuously long, setiform, on genu III much shorter and spiniform. Setae v" on tibia IV and pv" on tarsus IV plumose. Legs setal formulae normal. Remarks: The new species is readily classified in the genus Otocepheus Aoki, 1965, on the basis of the position of lyrifissures iad it is related to O. duplicornutus Aoki, 1965. However, the new species is readily distinguished from all its congeners by the fusiform seta da and the ratio (very short h,) of the notogastral setae. Derivatio nominis: The species is named after the kidney-shaped condyle of the prodorsum. Otocepheus spatulatus sp. n. Figs 88-91 Material examined: Sabah: Holotype: Sab-82/50. Holotypus: MHNG. Measurements: Length of body: 1035 um; width of body: 340 um. Prodorsum: Very wide in dorsal view, rostral apex, lateral part of prodorsum and surface of the long and wide lamellae with some weak foveolae. Tutorium strong, lateral lamelliform expansion well developed. One pair of rounded prodorsal condyles present, connected by a short transversal line. Rostral and lamellar setae setiform, well ciliate, interlamellar ones very short, but ensiform, exobothridial setae minute or reduced, hardly observable. Sensillus lanceolate, but very short, ending far from the lateral margin of prodorsum (in dorsal aspect). Notogaster: One pair of large notogastral condyles laterally, the distance between them smaller than the transversal diameter of one condyle. Surface of notogaster sparsely foveolate marginally. Ten pairs of notogastral setae present, two anterior pairs simple, fusiform, six other median pairs widened distally, slightly spa- tulate. Two pairs (setae p, and h;) short, fine and simply setiform (Fig. 88). Lyri- fissures ips situated between these two setae. Lateral part of podosoma: Tutorium and the lateral lamelliform expansion arched, fused with each other and clearly extending to the insertion of the rostral setae (Fig. 91). ORIBATIDS FROM SABAH VIII 715 Fics 84-87 Otocepheus reniformis sp. n. — 84: body in dorsal aspect, 85: body in ventral aspect, 86: dorsosejugal region, 87: podosoma in lateral aspect. 716 SANDOR MAHUNKA Fics 88-91 Otocepheus spatulatus sp. n. — 88: body in dorsal aspect, 89: body in ventral aspect, 90: dorsosejugal region, 91: podosoma in lateral aspect. ORIBATIDS FROM SABAH VIII ZT Ventral region: Coxisternal region simple and unornamented, ventral plate with very fine polygonal reticulation. Apodemes, primarily ap. 2 and ap. sej., charac- teristically arched (Fig. 89). Epimeral setae simple, setae /b, 3b and 4b longer than the others. Genital setae normal, aggenital setae very short. All three pairs of adanal setae short, simple and setiform, ad, arising far anteriorly and laterally, the distance between them and setae ad, much greater than between setae ad, and ad). Anterior anal setae (an,) minute, the posterior ones much longer (Fig. 89). Lyrifissure tad in adanal position. Legs: Type of ultimate setae: L - S - S - S. Seta /" of genua I-II very strong, long spiniform. Seta v" on tibia IV and pv" on tarsus IV plumose. Remarks: The lack of median condyles of the prodorsum readily places the new species into the genus Otocepheus sensu Aoki (1965). The peculiar form of the notogastral setae distinguishes the new species not only from the hitherto known species of this genus, but from all the species of the subfamily Otocepheinae. Derivatio nominis: The species name refers to the form of the notogastral setae. Otocepheus verrucosus sp. n. Figs 92-95 Material examined: Sabah: Holotype: Sab-82/15, 3 paratypes from the same sample. Holotype and 2 paratypes: MHNG; | paratype (1959-PO-94): HNHM. Measurements: Length of body: 931-960 um; width of body: 375 um. Prodorsum: Rostrum very wide, well foveolate. Lamellae long, running pa- rallel to each other. Both pairs of prodorsal condyles present, no remarkable difference between them. Lateral condyles angulate, the median ones rounded. Some characteristic, small, longitudinal crests present in interlamellar position (Fig. 92). Tutorium short, directed towards the lateral lamelliform expansion but not touching it; the latter long and reaching beyond the insertion of the rostral setae. Rostral and lamellar setae long, simple, setiform, slightly ciliate. Interlamellar setae bacilliform, longer than the notogastral ones, but shorter than the rostral setae. Exobothridial setae completely reduced (I was unable to find their alveoli). Sensillus simple, its head weakly fusiform, with some minute spicules on its distal end. Notogaster: Lateral notogastral condyles very large, the distance between them much smaller than their transversal diameter (Fig. 94). Ten pairs of comparatively short notogastral setae present, setae c, slightly fusiform, all the others simple but erect and mostly straight. All setae slightly ciliate. Lyrifissures in normal position. Lateral part of podosoma: Pedotecta I large, distinctly foveolate, as is the surface between the acetabula of legs II and HI. Both corners of pedotecta II-III rounded, the posterior part also ornamented (Fig. 95). Posterior part of discidium distinctly protruding. Ventral region (Fig. 93): Epimeral region typical for the genus, its surface weakly foveolate anteriorly and very roughly foveolate posteriorly. Epimeral setal formula: 3 - 1 - 3 - 3, setae /c arising very far laterally, observable only in lateral aspect. Setae /b, 3b, 4b and 4c long, setae 3c conspicuously long and unusually 718 SANDOR MAHUNKA Fics 92-95 Otocepheus verrucosus sp. n. — 92: body in dorsal aspect, 93: body in ventral aspect, 94: dorsosejugal region, 95: podosoma in lateral aspect. ORIBATIDS FROM SABAH VIII 719 directed laterally. Along border IV a characteristic ornamentation of rugae obser- vable (opp). Surface of ventral plate pustulate like the notogastral one, surface of genital plates with 2-3 longitudinal crests. All setae in the anogenital region long, partly filiform, setae ad; arising in marginal position very far from the anal aperture. Lyrifissures iad located near to the anal openings, but clearly in inverse apoanal position. Legs: Surface of femora without any characteristic sculpture. Setae /" of genua II and III spiniform. Type of ultimate setae: L - S - S- S. Remarks: The attribution of the new species to the genus Otocepheus is rather problematic, because the shape of pedotecta II-III is atypical and the posterior corner of the fishtail-shaped part of the body absent. However, the habitus, primarily the form of the notogaster, and the absence of the median notogastral condyles indicate the true relationship of this species. It is readily distinguished by the conspicuously long setae 3c, which are unknown elsewhere in the family. Derivatio nominis: The species name refers to the tubercled sculpture of the notogaster. ACKNOWLEDGEMENTS I want to thank most cordially Dr. B. Hauser, former Head of the Arthropod Department of the Museum d'histoire naturelle, Geneva, for allowing me to study this valuable material and for his efforts in editing this paper. I am also very grateful to Dr. Malcolm Luxton (National Museum of Wales, Cardiff) for his critical reading of the manuscript and his many useful suggestions. This research programme was partly sponsored by the Hungarian Scientific Research Fund (OTKA 16 729). REFERENCES AOKI, J. 1965. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata) I. Subfamily Otocepheinae. Bulletin of the National Science Museum. Tokyo 8: 259-341. AOKI, J. 1967. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata) II. Subfamily Tetracondylinae. Bulletin of the National Science Museum. Tokyo 10: 297- 339. GRANDIEAN, F. 1964. Oribates mexicains (Ire serie) Dampfiella Selln. et Beckiella n. g. Acarologia 6: 694-711. HAMMER, M. 1971. On some Oribatids from Viti Levu, the Fiji Islands. Biologiske Skrifter. Kongelige Danske Videnskabernes Selskab 16: 1-60. HAMMER, M. 1979. Investigations on the Oribatid Fauna of Java. Biologiske Skrifter. Kongelige Danske Videnskabernes Selskab 22: 1-79 (1979). MAHUNKA, S. 1987a. Neue und interessante Milben aus dem Genfer Museum LV. Oribatids from Sabah (East Malaysia) I (Acari: Oribatida). Archives des Sciences 40: 293-305. MAHUNKA, S. 1987b. Neue und interessante Milben aus dem Genfer Museum LX. Oribatids from Sabah (East Malaysia) II. (Acari: Oribatida). Revue Suisse de Zoologie 94: 765- 817. MAHUNKA, S. 1988. New and interesting mites from the Geneva Museum LXI. Oribatids from Sabah (East Malaysia) III (Acari: Oribatida). Revue Suisse de Zoologie 95: 817-888. 720 SANDOR MAHUNKA MAHUNKA, S. 1990. A survey of the superfamily Euphthiracaroidea Jacot, 1930 (Acari: Oribatida). Folia Entomologica Hungarica 51: 37-80. MAHUNKA, S. 1991. New and interesting mites from the Geneva Museum LX VIII. Oribatids from Sabah (East Malaysia) IV (Acari: Oribatida). Revue Suisse de Zoologie 98: 185-206. MAHUNKA, S. 1995a. Oribatids from Sabah, East Malaysia (Acari Oribatida, Parakalummoidea n. stat. and Galumnoidea). Tropical Zoology 8: 269-308. MAHUNKA, S. 1995b. Oribatids from Brunei I (Acari: Oribatida). New and interesting mites from the Geneva Museum LXXV. Revue Suisse de Zoologie 102: 913-942. MAHUNKA, S. 1996a. Oribatids from Sabah (East Malaysia) VI (Acari: Oribatida). (Acarologica Genavensia LXXXIV). Archives des Sciences 49: 99-104. MAHUNKA, S. 1996b. Oribatids from Sabah (East Malaysia) VII (Acari: Oribatida). (Acaro- logica Genavensia LXXXV). Archives des Sciences 49: 205-212. PEREZ-INIGO, C. & BAGGIO, D. 1986. Oribates édaphiques du Brésil (II). Oribates de l'île du "Cardoso" (deuxieme partie). Acarologia 27: 163-179. WALLWORK, J. A. 1962a. Some Oribatei from Ghana VIII. The genus Tetracondyla Newell 1956 (Ist. series). Acarologia 4: 274-291. WALLWORK, J. A. 1962b. Some Oribatei from Ghana. IX. The genus Tetracondyla Newell 1956 (2nd. series). Acarologia 4: 440-456. REVUE SUISSE DE ZOOLOGIE 107 (4): 721-736; décembre 2000 More about the Buthoidea of Madagascar, with special references to the genus Tityobuthus Pocock (Scorpiones, Buthidae) Wilson R. LOURENCO Laboratoire de Zoologie (Arthropodes), Muséum National d’ Histoire Naturelle, 61 rue de Buffon, F-75005 Paris, France. E-mail. arachne@ mnhn.fr More about the Buthoidea of Madagascar, with special references to the genus Tityobuthus Pocock (Scorpiones, Buthidae). - A revised diagnosis of the genus Tityobuthus Pocock is given and three new species, i.e. 7. manonae sp. n., T. griswoldi sp. n. and T. monodi sp. n., are descri- bed. For two species previously associated with Tiryobuthus, two new genera are established, i.e. Troglotityobuthus gen. n. for Tityobuthus gra- cilis (Fage) (described under Babycurus gracilis) and Palaeogrosphus gen. n. for Tiryobuthus copalensis Lourengo. Phylogenetic relationships be- tween Madagascan scorpions and those in Africa, South America and Asia are discussed. Revised keys are given for the genera of Buthoidea present in Madagascar and for the species of the genus Tityobuthus. Key-words: Scorpiones - Buthidae - Microcharmidae - taxonomy - Mada- gascar. INTRODUCTION The genus Tiryobuthus was established by Pocock (1893) for Tityobuthus baroni (Pocock, 1890) previously included in the genus Rhoptrurus Karsch, 1886, because this name was already preoccupied by that of a snake genus, Rhoptrura Peters, 1858 (Vachon, 1979). Confusion existed about the genera Tityobuthus Pocock, Pseudobuthus Pocock and Odonturus Karsch until they were revised by Vachon (1979), who finally included only two species in the genus Tityobuthus, i.e. Tityo- buthus baroni (Pocock, 1890) and Tityobuthus gracilis (Fage, 1946). The latter was originally described in the genus Babycurus Karsch, 1886. In the last few years, and in particular since the scorpion fauna of Madagascar was summarised by Lourenco (1996a), increasing numbers of new species have been described in the genus Tityobuthus. These are: T. guillaumeti Lourenco, T. pococki Lourengo, 7. lucileae Lourengo, T. copalensis Lourengo, T. petrae Lourengo, 7. parrilloi Lourengo, T. judsoni Lourenço, T. dastychi Lourenço and T. ivohibe Lourenco & Goodman. This shows that this genus of micro-scorpions is very rich in species (Lourengo, 1995, 1996a,b,c, 1997a; Lourenço & Goodman, 1999). The same phenomenon has already been observed in other micro-scorpion genera including Manuscript accepted 02.03.2000 722 WILSON R. LOURENCO Ananteris Thorell. The number of species known in that genus increased from 3 to 23 within twenty years (Lourenço, 1993, 1994, 1997b, 1999a, b; Lourenço & Monod, 1999). The recent discovery of three additional new species of the genus Tityobuthus pointed out the necessity for a revision of this genus. A reanalysis of characters showed that two species previously associated with Tityobuthus, 1.e. T. gracilis (Fage) and T. copalensis Lourengo, deserve a separate taxonomic position. Revised keys to the genera of Buthoidea present in Madagascar, and to the known species of the genus Tiryobuthus are also provided in the following. Possible phylogenetic links and associations between Madagascan lineages and those present in Africa, South America and Asia are pointed out. TAXONOMY OF TITYOBUTHUS Tityobuthus Pocock, 1893 Type species: Rhoptrurus baroni Pocock, 1890; described from Madagascar. Revised diagnosis: Scorpions of small size, 20-25 mm in total length. Carapace with a moderate to strong concavity; median ocular tubercule markedly anterior to the center of the carapace; three pairs of lateral eyes. Chelicerae with dentition according to the buthid pattern (Vachon, 1963). Pedipalps slender and moderately long; movable fingers with 7 or 8 slightly oblique, almost straight rows of granules. Tibial spurs present in some species, reduced in others and absent in a few. Sternum sub- triangular. Pectines with 11-25 teeth; fulcra present or absent according to species; basal middle lamellae of female never dilated. Sternites with very short linear stigmata. Telson with a very slender vesicle; aculeus long and moderately curved, with a spinoid subaculear tooth of variable size. Trichobothrial pattern of type A-a - orthobothriotaxic. Remarks: The presence or absence of tibial spurs is a particularity of this genus since this character has always been considered to be invariable within a given genus. The fact that in Tiryobuthus species certain characters are present in different states had already lead Vachon (pers. comm.) to consider a division of the genus into two or more genera. I have, however, some hesitations about taking this step. Tityobuthus, like other buthid genera, may belong to archaic lineages in which some characters are not precisely differentiated (Lourenco, 1995, 1996a). Distribution: Madagascar. DESCRIPTION OF THREE NEW SPECIES Tityobuthus manonae sp. n. Figs 12-16 Type: MADAGASCAR, Mandena, Fort Dauphin (littoral forest 10 km north of Fort Dauphin), holotype male, 6-12/1/1999 (J.-B. Ramanamanjato leg.)!; deposited in the Muséum d'histoire naturelle, Geneve (MHNG). Etymology: Patronym in honor of Dr Manon Vincelette of QIT Minerals- Madagascar, Montreal, Canada. ! Collected with 11 specimens of Grosphus hirtus Kraepelin, also deposited in MHNG. BUTHOIDEA OF MADAGASCAR 723 Description (based on male holotype): Measurements in Table I. Coloration. Ground colour yellowish, symmetrically marbled with a dark reddish brown, giving an overall spotted appearence. Prosoma: carapace yellowish, moderately spotted; eyes surrounded by black pigment. Mesosoma: yellowish, with four longitudinal brown stripes, i.e. two central and two lateral ones. Metasoma: segments I to IV yellowish; V yellowish to reddish yellow. Vesicle as segment V. Venter yellowish, with a number of spots on sternites VI and VII. Chelicerae yellowish, with dark spots on the lateral edges; fingers reddish. Pedipalps: yellowish, with several dark spots on femur and tibia; chela less densely spotted; hands yellowish; fingers much darker, reddish-brown, with the extremities yellowish. Legs yellowish, with diffuse fuscous spots. TABLE I Measurements (in mm) of male holotypes of the described species T. monodi T. manonae T. griswoldi Carapace: - length 2.6 246 1:9 - anterior width 1.8 231 13 - posterior width 251 2 2.0 Metasomal segment I: - length 1.6 1:5 1.1 - width 107 5 12 Metasomal segment V: - length 33 3.4 25 - width 183 1.4 1.0 - depth 12 ies) 0.9 Vesicle: - width 0.8 1.0 0.7 - depth 0.9 11 0.7 Pedipalp: - Femur length DID. 2.6 1.8 - Femur width 0.7 0.7 0.5 - Tibia length 29 3.2 2.6 - Tibia width 12 les 0.8 - Chela length 4.1 5.0 3.4 - Chela width 0.9 1122 0.6 - Chela depth 0.8 1.0 0.6 Movable finger length N 3.3 2.4 Morphology. Carapace moderately to weakly granular; anterior margin with a weakly to moderately pronounced median concavity. Anterior median superciliary, posterior median keels and all furrows moderate to feeble. Median ocular tubercle distinctly anterior to the center of the carapace; median eyes separated by one ocular diameter. Three pairs of lateral eyes. Sternum subtriangular. Mesosoma: tergites moderately granular. Median keel moderate to strong on all tergites; confluent vesti- 724 WILSON R. LOURENCO gial keels present. Tergite VII pentacarinate. Venter: genital operculum longitudinally divided. Pectines: pectinal tooth count 19/20; basal middle lamellae not dilated; fulcra present. Sternites smooth or with very weak granulations and small, elongate stigmata; VII with two keels. Sternite V with a large smooth, bright zone on posterior edge. Metasoma: segments I to III with 10 keels, crenulate. Segment IV with 8 keels, crenulate. Intercarinal spaces weakly to moderately granular. Segment V smooth and rounded. Telson smooth, without punctuations, with a long but moderately curved aculeus; subaculear tooth very small and slightly rhomboid. Cheliceral dentition characteristic of the family Buthidae (see Vachon, 1963); basal teeth of movable fingers reduced and almost fused; ventral surfaces of finger and manus almost without setae. Pedipalps: femur pentacarinate; tibia and chelae with some keels, moderately crenulate; internal face of tibia with six spinoid granules; all faces moderately granular; movable fingers with 8/9 oblique rows of granules. Trichobothriotaxy; orthobothriotaxy A-a (cf. Vachon, 1973, 1975). Legs: tarsus with numerous fine median setae ventrally. Pedal and tibial spurs present but reduced. Relationships: From general morphology T. manonae sp. n. appears to be most closed related to 7. petrae Lourenco, 1996. The two can be readily distinguished by the following features: (1) the new species is much larger (see Table I, Lourenço, 1996a), (11) the presence of a single conspicuous white smooth and bright central zone on sternite V of the new species, whereas in T. petrae this zone is reduced and there are two small rounded white patches laterally. Tityobuthus griswoldi sp. n. Figs 17-21 Types: MADAGASCAR, Prov. Fianarantsoa, P.N. Ranomafana, Talatakely (21°15,3° S - 47°25,9° E), holotype male 9-26-IV-1998 (pitfall traps; C.E. Griswold, D.H. Kavanaugh, N.D. Penny & M.J. Raherilalao leg.); deposited in the California Academy of Sciences, San Francisco (CAS). Paratypes: Same data as for the holotype. 1 male, 19-30/IV/1998, deposited in MHNG; 1 juvenile; 3 juveniles, 19-30/1V/1998 (21°14,9 S - 47°25,6 E), deposited at the CAS. Etymology: Patronym in honor of Dr Charles E. Griswold of the California Academy of Sciences. Description (based on male holotype): Measurement in Table I. Coloration. Colour yellow, symmetrically marbled with dark brown, giving an overall spotted appearence. Prosoma: carapace yellowish, densely spotted; eyes surrounded by black pigment. Mesosoma: yellowish with three longitudinal brown stripes, the median one indistincly divided. Metasoma: segments I to V yellowish; vesicle as is segment V. Venter yellowish, with a number of dark spots on all ster- nites. Sternite V with a large bright white zone in the posterior region and with two small rounded white bright zones laterally. Chelicerae yellow, with dark spots in median zone; fingers reddish. Pedipalps yellowish, with several dark spots on femur and tibia; chela less densely spotted; hands yellowish, fingers much darker, reddish- brown. Legs yellowish, with diffuse fuscous spots. Morphology. Carapace moderately to weakly granular; anterior margin with moderate median concavity. Anterior median superciliary, posterior median keels and BUTHOIDEA OF MADAGASCAR 725 all furrows moderate to weak. Median ocular tubercle distinctly anterior to center of carapace; median eyes separated by one ocular diameter. Three pairs of lateral eyes. Sternum subtriangular to pentagonal. Mesosoma: tergites weakly granular. Median keel moderate in all tergites. Tergite VII pentacarinate. Venter: genital operculum longitudinally divided. Pectines: pectinal tooth count 14/14; basal middle lamellae not dilated; fulcra present. Sternites weakly granular with small and moderately elongate stigmata; VII without keels. Sternite V with smooth, bright white zone in posterior region and two similar small rounded zones laterally. Metasoma: segments I and II with 10 keels, crenulate. Segments HI and IV with 8 keels, crenulate. Intercarinal spaces weakly to moderately granular. Segment V smooth and rounded. Telson smooth without punctuations, with fairly short, moderately curved aculeus; subaculear tooth strong, slightly spinoid. Cheliceral dentition characteristic of the family Buthidae (cf. Vachon, 1963); basal teeth of movable fingers small; ventral surface of finger and manus with setae. Pedipalps: femur pentacarinate; tibia and chela with some keels but moderately crenulate; internal face of tibia with four spinoid granules; all faces feebly granular; movable fingers with 8/9 oblique rows of granules. Trichobothriotaxy; orthobothriotaxy A-a (cf. Vachon, 1973, 1975). Legs: tarsus with numerous fine median setae ventrally. Pedal and tibial spurs present but reduced. Variation: the pectinal tooth count in the paratypes is: 13-13, 14-13, 12-14, 14- 14 (x2). Relationships: From general morphology T. griswoldi sp. n. appears to be most closed related to 7. dastychi Lourenco. The two species can, however, be readily distinguished by the following features: (1) a different shape of the pectines and a different number of teeth, 12 to 14 in the new species and 18 in T. dastychi, (ii) the presence of a single conspicuous smooth bright white zone on sternite V of T. dastychi, while in the new species this central zone is reduced and associated with two small rounded smooth bright white zones laterally. Tityobuthus monodi sp. n. Figs 22-26 Type: MADAGASCAR, Prov. Toamasina, F.C. Andriantantely (18°41,7° S - 48°48,8° E), 530 m, holotype male, 7-10-XII-1998 (H. J. Ratsirarson leg.); deposited in MHNG. Etymology: Patronym in honor of M. Lionel Monod of the Muséum d'histoire naturelle de Geneve. Description (based on male holotype): Measurement in Table I. Coloration. Ground colour yellow, symmetrically marbled with dark brown, giving an overall spotted appearence. Prosoma: carapace yellowish, densely spotted: eyes surrounded by black pigment. Mesosoma: yellowish, with three longitudinal brown stripes more or less inter-linked by confluent stripes. Metasoma: segments I to V yellowish; vesicle yellowish but lighter than segment V. Venter yellowish, with all sternites densely spotted. Sternite V with only a vestigial bright white zone in pos- terior region. Chelicerae yellowish, densely spotted in central zone; fingers reddish. Pedipalps yellowish, with several dark spots on femur and tibia; chela less densely spotted; hands light yellow; fingers much darker, reddish, with the extremity more yellow. Legs yellowish, with diffuse fuscous spots. 726 WILSON R. LOURENCO Morphology. Carapace moderately to weakly granular; anterior margin with weak to moderate median concavity. Anterior median superciliary, posterior median keels and all furrows moderate to weak. Median ocular tubercle distinctly anterior to center of carapace; median eyes separated by more than one ocular diameter. Three pairs of lateral eyes. Sternum subtriangular to pentagonal. Mesosoma: tergites weakly granular. Median keel very weak in all tergites. Tergite VII pentacarinate. Venter: genital operculum longitudinally divided. Pectines: pectinal tooth count 16/16; basal middle lamellae not dilated; fulcra very reduced, almost vestigial. Sternites smooth, with small elongate stigmata; VII without keels. Metasoma: segments I and II with 10 keels, crenulate. Segments II and IV with 8 keels crenulate. Intercarinal spaces weakly granular. Segment V smooth and rounded. Telson smooth, without punc- tuations, with moderately long, curved aculeus; subaculear tooth strong and spinoid. Cheliceral dentition characteristic of the family Buthidae (cf. Vachon, 1963); basal teeth on movable fingers small; ventral surface of finger and manus with some setae. Pedipalps: femur pentacarinate; tibia and chela with some keels but moderately crenulate; internal face of tibia with four/five granules but not spinoid; all faces weakly granular, almost smooth. Movable fingers with 8/9 oblique rows of granules. Trichobothriotaxy; orthobothriotaxy A-a (cf. Vachon, 1973, 1975). Legs: tarsus with numerous fine median setae ventrally. Pedal and tibial spurs present but very reduced. Relationships: Tityobuthus monodi appears to be most closed related to 7. guillaumeti Lourenco. The two species can be readily distinguished by the following features: (1) a different pattern of pigmentation, 1.e. the new species is heavily spotted, whereas 7. guillaumeti is uniformily yellowish, (i1) the different shape of the pectines; in the new species these are small and have vestigial fulcra, whereas in T. guillaumeti they are large and the fulcra is completely absent. DESCRIPTION OF TWO NEW GENERA Troglotityobuthus gen. n. Figs 1-7 Type species: Babycurus gracilis Fage, 1946; described from Madagascar, Ambilobe, Ankarana, «grotte des Fanihys». Diagnosis: Scorpions of medium size, 45 mm in total length. Pigmentation almost absent. The whole body flattened dorsally, all appendices long and slender. Carapace with a moderate concavity; very flat dorsally. Median ocular tubercule anterior to center of carapace; three pairs of lateral eyes. Chelicerae with dentition according to the buthid pattern (Vachon, 1963). Pedipalps long and slender; movable fingers very long, with 9/10 slightly oblique, almost straight rows of granules. Tibial spurs present. Sternum triangular. Pectines with 20/20 teeth; fulcra strongly marked; basal middle lamellae not dilated. Sternites with short linear stigmata. Telson with a very slender vesicle; aculeus long and moderately curved, with a spinoid subaculear tooth. Trichobothrial pattern of type A-a - orthobothriotaxic. Distribution: Madagascar. BUTHOIDEA OF MADAGASCAR 727 Palaeogrosphus gen. n. Figs 8-11 Type species: Tityobuthus copalensis Lourengo, 1996; described from Mada- gascar, Diégo-Suarez, region of Antseranana (in copal). Diagnosis: Scorpions of small size, 20-25 mm in total length. Carapace with a moderate concavity; median ocular tubercule anterior to the center of the carapace; three pairs of lateral eyes. Cheliceral dentition apparently (with some uncertainty) in accordance with the buthid pattern (Vachon, 1963). Pedipalps slender but not long; movable fingers with 10 only slightly oblique, almost straight rows of granules. Tibial spurs not easily discernible, apparently reduced. Sternum sub-pentagonal. Pectines without fulcra; basal middle lamellae moderately dilated. Sternites with short linear stigmata. Telson with a moderately incrassate vesicle; aculeus moderately curved, with a short and slightly rhomboidal subaculear tooth. Trichobothrial pattern of type A-a, probably orthobothriotaxic but not all trichobothria discernible. Distribution: Madagascar (in recent palaeogeological periods; late Cenozoic- early Pleistocene). Remark: The character basal middle lamellae moderately dilated, suggests that the new genus is related to the genus Grosphus. BIOGEOGRAPHY As noted by previous authors (Fage, 1929; Millot, 1948; Vachon, 1979), there are significant affinities between the scorpion faunas of Madagascar and Africa. The following arguments can be given in support of this view. (1) Grosphus Simon, one of the most species-rich Madagascan genera, and Neogrosphus Lourengo, a genus which probably evolved more recently from Grosphus, undoubtedly have close affinities with the African genus Uroplectes Peters. These three genera share a very remarkable sexual dimorphism of the pectines (Fage, 1929). The new genus Palaeogrosphus gen. n. may represent a primitive element, older than Grosphus and Neogrosphus, but more data on this sub-fossil genus are needed before a decision can be made (Lou- renco, 1995, 1996a), (11) The recently discovered genus Pseudouroplectes Lourenço may also have some affinities with the genus Uroplectes. However, the morphology of Pseudouroplectes suggests a much more primitive lineage when compared with Uroplectes and Grosphus, which certainly evolved later (Lourenço, 1995, 1996a). Pseudouroplectes corresponds to primitive lineages, which have probably vanished in other regions of the world but are still present in Madagascar. This genus has distant affinities with the microcharmid genus Microcharmus Lourengo, but some inter- mediate elements still have to be identified in the Madagascan Buthoidea. (iii) The genus Tityobuthus was phylogenetically poorly defined until recently, but the disco- very of several new species has improved our knowledge of this group. Affinities possibly exist with the Gondwanan genus Ananteris Thorell, today present in West Africa and South America. Ananteris seems to have had its center of origin in the western part of Gondwanaland, which corresponds with present-day South America. More recently, affinities have also been shown between Ananteris, Tityobuthus and the Himalayan genus Himalayotityobuthus Lourenço (Lourenço, 1997c). These three 728 WILSON R. LOURENCO genera show several primitive characteristics, including small size and the absence of fulcra in the pectines (the absence of fulcra is observed in at least one species of Tityobuthus). The new genus 7roglotitvobuthus gen. n. represents a troglobitic element, which probably derived from epigean species close to Tityobuthus. However, our knowledge of this genus remains extremely incomplete. The discovery of the genus Microcharmus, in the north-eastern region of Madagascar, provided the first indi- cation of possible affinities with Charmus Karsch in India and Sri Lanka. Both Microcharmus and Charmus represent primitive lineages whose characteristics (see Lourenço, 1995, 1996a) show that they belong to the most early-derived extant Buthoidea. The descriptions of several other species of Microcharmus and the study of the African genus Akentrobuthus Lamoral have lead (in a preliminary decision) to the transfer of these two genera to a separate buthoid family, the Microcharmidae (Lourenco, 1998). In a more recent decision, the genus Akentrobuthus was replaced in the family Buthidae (Lourenco, 2000) and the Microcharmidae returned again to be represented by relictual elements, present only in Madagascar. At the moment there are no further indications to suggest affinities between Madagascar, Indo-Malayan region and Australia. CONCLUSIONS Although this study is only preliminary, the following findings on the Buthoidea of Madagascar can be presented: |. The majority of the buthoid taxa in Madagascar correspond with primitive lineages which no longer exist in most other regions of the world. 2. Several genera appear to be poor in species. It is, however, reasonable to expect future discoveries of several new species in certain genera of micro-scorpions such as Tityobuthus, Microcharmus and Pseudouroplectes. 3. The number of recorded species is still small, but the faunistic inventories so far carried out are extremely incomplete. 4. Neverthless the number of genera is significant, even when compared with other well-studied regions of the world. 5. The most remarkable characteristic of the buthoids of Madagascar is the impressive level of endemicity, both in species and in genera. This supports the hypothesis of a very early isolation of the island from other land masses. KEY TO THE GENERA OF BUTHOIDEA PRESENT IN MADAGASCAR I Sternum with atnancular shape: Buthidae;......- ...... 15 2 eee 2 (1) Sternum with a sub-pentagonal to pentagonal shape SERRE TE Re OA Microcharmidae - Microcharmus 2 Chelae: movable fingers with 10 to 14 oblique rows of granules ........... 3 (2) Chelae: movable fingers with 7 or 8 oblique rows of granules............ 5 3 Carapace distinctly flattened; length of movable finger 3-4 times the INCI ne Tae Meee 3 2 mes ga a oe os Troglotityobuthus gen. n. (3) Carapace not flattened; movable finger 1,5 times longer than hand ......... + BUTHOIDEA OF MADAGASCAR 729 4 Telson without subaculear tooth; pectines with fulcra ............ Grosphus (4) Telson with small subaculear tooth; pectines without fulcra AE SRI OR ORE RENTEN: PR STO SEN Palaeogrosphus gen. n. 5 Telson with moderate to strong subaculear tooth.............. Tityobuthus O eerelelsoniwithout:subaculear tooth 7 u. oo) ee ee tie NO 6 6 Pectines of female with basal middle lamellae moderately dilated Neogrosphus (6) Pectines of female with basal middle lamellae not dilated . . . Pseudouroplectes KEY TO THE SPECIES OF TITYOBUTHUS 1 Beetineswithkulerasvestieiallorabsent ARCER 2 (DEssBecimeswithswellideveloped fuleram mn. RA A I 3 2 FU CARVE SUC Re EE RITA RAR monodi sp. n. COMMENTE ADS tw: nase Are Sr T. guillaumeti 3 Aibial-spurstabsenter. ne ee N I ne T. baroni (3) MPa punsypnesemta:.. ls Gavel chy cnn pcan ete O ae 4 - Chelicerae without spots-orpiementatona TE a... T. judsoni (Des Ehelicerae with dark spots andipiementation an... fee ne 3) I Rectines: with? Oror moreiteeth m. EI 6 (5) Rectinesswath less: thame2 Oite ethics re E etree 8 6 A conspicuous smooth, white and bright central zone on sternite V RE CIN CA aE Sa RA RAS A IS ES a RIE T. manonae sp. n. (6) Noxsuchrzone ON STEAM sil ay I RIA e 7 7 Body, pedipalps and legs heavily spotted; pedipalpal chelae short and robust; internal face of tibia with four spinoid granules .......... T. lucileae (7) Body, pedipalps and legs with only vestigial spots; pedipalpal chelae long and slender; internal face of tibia with 5 or 6 spinoid granules . T. pococki 8 Pectumeswathelikteethy o wean itn, abe et cae n T. ivohibe (8) Pectnestwithdil2:{0 19 ele Ne 9 9 Rectines;wathol2 toni tects <= RO RR A ee En. 10 Ole sekecones with Stone rs ern. ee ee Il 10 A smooth white central zone on sternite V and two small rounded smoothiwhite;zones-laterallly RR me. T. griswoldi sp. n. (O)ZE=S temnitesVAWIthoutisMeothiWIIWeizZOnert neue. NR o T. parrilloi 11 Sternite V centrally with a single conspicuous smooth white zone . 7. dastychi (11) Sternite V with a reduced smooth white central zone associated with two small rounded smooth white zones laterally ................. T. petrae ACKNOWLEDGEMENTS I am very grateful to Didier Geffard and Philipe Bouchard, Laboratoire de Zoologie Arthropodes, for preparing several illustrations. Prof. John L. Cloudsley- Thompson, London and Dr. Peter Schwendinger, Geneva are acknowledged for reviewing the manuscript. 730 WILSON R. LOURENCO Fics 1-7 Male holotype of Troglotityobuthus gracilis. 1. Carapace, dorsal view. 2. The same, lateral view. 3. Sternite V. 4. Metasoma and telson, lateral view. 5. Movable finger of pedipalpal chela. 6. Pectines. 7. Pedipalp, dorsal view with trichobothrial pattern. 731 BUTHOIDEA OF MADAGASCAR ER / \ i N 7 N / \ / \ / \ it \ / ! I 1 \ 11 \ \ + We Fics 8-11 Male holotype of Palaeogrosphus copalensis. 8. Left pecten. 9. Movable finger of pedipalpal chela. 10. Metasoma and telson, lateral view. 11. Metasomal segment V and telson in detail lateral view. 732 WILSON R. LOURENCO Fics 12-16 Male holotype of Tityobuthus manonae sp. n. 12. Carapace. 13. Pectines. 14. Sternite V. 15. Metasoma and telson, lateral view. 16. Pedipalp, dorsal view with the trichobothrial pattern. BUTHOIDEA OF MADAGASCAR 733 Fics 17-21 Male holotype of Tiryobuthus griswoldi sp. n. 17. Carapace. 18. Sternite V. 19. Pecten. 20. Pedipalp, dorsal view with trichobothrial pattern. 21. Metasoma and telson, lateral view. 734 WILSON R. LOURENCO Fics 22-26 Male holotype of Tityobuthus monodi sp. n. 22. Carapace. 23. Pectines. 24. Sternite V. 25. Pedipalp, dorsal view with trichobothrial pattern. 26. Metasoma and telson, lateral view. BUTHOIDEA OF MADAGASCAR 735 ® Troglotityobuthus gracilis @ Palacogrosphus copalensis À Tityobuthus manonae © Tityobuthus griswoldi A Tityobuthus monodi Fic. 27 Map of Madagascar showing the type localities of the new genera and species described in this paper. REFERENCES FAGE, L. 1929. Les Scorpions de Madagascar. Faune des Colonies frangaises 3. Société d’Edi- tions Géographiques, Maritimes et Coloniales, Paris: 637-694. LouRENCO, W. R. 1993. A review of the geographical distribution of the genus Ananteris Thorell (Scorpiones: Buthidae), with description of a new species. Revista de Biologia Tropical 41 (3): 697-701. LOURENÇO, W. R. 1994. Scorpions (Chelicerata) de Colombie. VI. Quatre nouvelles espèces de Buthidae des regions amazonienne, sud-pacifique et de la cordillere orientale. Revista de la Academia Colombiana de Ciencias Exatas, Fisicas y Naturales 19 (73): 387-392. LOURENCO, W. R. 1995. Description de trois nouveaux genres et de quatre nouvelles especes de scorpions Buthidae de madagascar. Bulletin du Muséum National d'Histoire Naturelle, Paris, 4° ser. 17 (1-2): 95-106. 736 WILSON R. LOURENCO LOURENCO, W. R. 1996a. Scorpions. In: Faune de Madagascar. N° 87. Muséum National d'Histoire Naturelle, Paris: 102 pp. LOURENCO, W. R. 1996b. A new species of Tityobuthus from Madagascar (Scorpiones, Buthidae). Bolletino del Museo Regionale di Scienze Naturali, Torino 14 (1): 267-273. LOURENCO, W. R. 1996c. Premier cas connu d’un sub-fossile de scorpion dans le copal de Mada- gascar. Comptes Rendus de l’Académie des Sciences, Paris, Ser. Ila, t. 323: 889-891. LOURENÇO, W. R. 1997a. Another new species of Tityobuthus from Madagascar (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 12 (155) : 147-151. LOURENÇO, W. R. 1997b. A reappraisal of the geographical distribution of the genus Ananteris Thorell (Scorpiones: Buthidae). Biogeographica 73 (2): 81-85. LOURENCO, W. R. 1997c. Description of a new genus and new species of Buthidae scorpion from the Himalayas of India and Nepal, with some new biogeographic implications. Entomo- logische Mitteilungen aus dem Zoologischen Museum Hamburg 12 (156) : 183-188. LOURENCO, W. R. 1998. Une nouvelle famille est nécessaire pour des microscorpions humicoles de Madagascar et d’Afrique. Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 321: 845-848. LOURENCO, W. R. 1999a. A new species of Ananteris Thorell from the coastal region of Ecuador (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 13 (160): 95-100. LOURENCO, W. R. 1999b. New species of Ananteris from the north of Chocé, Colombia (Scor- piones, Buthidae). Anales del Instituto de Biologia, Universidad Nacional Autonoma de México, ser. Zool. 70 (2): 93-98. LOURENCO, W. R. 2000. Panbiogéographie, les familles des scorpions et leur répartition géo- graphique. Biogeographica 76 (1): 21-39. LOURENÇO, W. R. & GOODMAN, S. M. 1999. Taxonomic and ecological observations on the scorpions collected in the Forest of Ankazomivady-Ambositra and on the «RS d’Ivohibe», Madagascar. Revista de Biologia Tropical 47 (3): 475-482. LOURENCO, W. R. & Monon, L. 1999. A new species of Ananteris Thorell from French Guyana (Scorpiones, Buthidae). Revue suisse de Zoologie 106 (2): 301-306. MILLOT, J. 1948. Revue générale des Arachnides de Madagascar. Mém. Inst. Sci. Madagascar, Ser. A 1(2). In: Mémoires de la Société de Biogeographie (1953): 127-145. Pocock, R. I. 1893. Notes on the classification of scorpions followed by some observations upon synonymy, with descriptions of genera and species. Annals and Magazine of Natural History (6) 12: 303-330. VACHON, M. 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères chez les Scorpions. Bulletin du Museum National d’Histoire Naturelle, Paris, 2° ser. 35 (2): 161-166. VACHON, M. 1973. Etude des caracteres utilisés pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles tricho- bothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Museum National d’Histoire Naturelle, Paris, 3è sér., n° 140, Zool., 104: 857-958. VACHON, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des Scorpions (Arachnides) dans le classement des genres de la famille des Buthidae Simon. Comptes Rendus de l’Académie des Sciences, Paris, ser. D, 281: 1597-1599. VACHON, M. 1979. Remarques biogéographiques sur la faune des Scorpions de Madagascar a propos de l'utilisation de caractères trichobothriotaxiques permettant la distinction des genres Odonturus Karsch, 1879 et Tityobuthus Pocock, 1893. Comptes-Rendus du V° Colloque d’Arachnologie d’Expression Francaise, Barcelona: 217-224. REVUE SUISSE DE ZOOLOGIE 107 (4): 737-764; décembre 2000 Japygidés (Diplura) du Sud-Est asiatique n° 9 Dicellurata Genavensia XXIV Jean PAGES! 51, rue du Faubourg Saint-Martin, F-21121 Fontaine-les-Dijon, France. Japygidae (Diplura) from South-East Asia n° 9. - Dicellurata Gena- vensia XXIV. - This note is devoted to the descriptions or citations of 5 species belonging to 3 genera. Psalidojapyx gen. n. is proposed for Japyx murudensis Silv., 1930 described from Sarawak, here recorded from Brunei, and Psalidojapyx edentulus sp. n. from Sarawak. Burmjapyx Silv. is repre- sented by two new species: Burmjapyx lienhardi sp. n. from Sumatra and Burmjapyx murphyi sp. n. from Singapore. The generic position of Burm- Japyx huashanensis Chou, 1983 is discussed. New specimens of Paupero- Japyx (Paucijapyx) kajan Pgs are recorded from Sarawak and Brunei. Key-words: Diplura Japygidae - Indonesia - Singapore - Malaysia - Brunei - new genus and new species. INTRODUCTION Dans cette note sont étudiés des Japygidés provenant de trois des missions (1985, 1987 et 1988) organisées dans le Sud-Est asiatique par le Dr B. Hauser, ancien Conservateur du Département des Arthropodes et d'Entomologie I du Muséum d'histoire naturelle de la Ville de Genève, en collaboration avec le Dr C. Lienhard, Chargé de Recherche dans le méme Département; ils n’appartiennent pas aux omni- présents /ndjapyx, mais représentent trois genres dont un nouveau pour la Science. I Psalidojapyx gen. n. est proposé pour le Japyx murudensis Silv., 1930 du Mont Murud (2438 m ) au Sarawak (Malaisie) et dont je décris un dj du Brunei présentant des caractéristiques particulieres permettant de mieux comprendre la varia- bilité des espèces dans ce genre; Psalidojapyx edentulus sp. n. est une nouvelle espèce du Sarawak. II- Burmjapyx lienhardi sp. n. de Sumatra et Burmjapyx murphyi sp. n. de Singapour portent a 6 le nombre d'espèces de ce genre propre à l'Asie du Sud-Est asiatique, tel qu’il a été défini par Silvestri (1930). La position générique du Burm- Japyx huashanensis Chou (Chou & Chen, 1983) me paraît inappropriée et j apporte quelques éléments de discussion. III.— Pauperojapyx (Paucijapyx) kajan Pagés est signalé de 2 nouvelles sta- tions, l’une au Brunei, l’autre au Sarawak. | Professeur émérite de l'Université de Bourgogne. Manuscrit accepte le 17.07.2000 738 JEAN PAGES Tous les spécimens étudiés dans ce travail sont conservés dans les collections du Muséum d'histoire naturelle de la Ville de Genève. ETUDE TAXONOMIQUE? I.- Psalidojapyx gen. n. Appartient à la famille des Japygidae. Le premier stade à forceps est un st.III B. Espèce type: Japyx murudensis Silvestri, 1930. TETE Antennes: de 43 articles chez l’espece type, les 13 trichobothries typiques, a proximale, mais non subbasale; 6 ou 8 sensilles placoides en position typique sur l’article apical. Pieces buccales: typiques de la famille; les 5 lames du lobe interne des maxilles pectinées. ‘THORAX Chétotaxie: les 5+5 M en position typique sur les 3 notums. Pattes: peu allongées, tres pileuses; les soies spiniformes des tarses tres développées, peu nombreuses; prétarse typique. ABDOMEN Tergite 1: avec les seuls ma = M sur le scutum. Tergites 2 a 7: a chétotaxie typique. Tergite 10: sans carènes. Sternites: a chétotaxie normale. Organes subcoxaux latéraux: larges, peu saillants; soies glandulaires et soies sensorielles normales. Organe glandulaire médian: saillant, de nombreuses minuscules sensilles séti- formes (= "pseudopori") sur |’ opercule. Papilles génitales: typiques de la famille, Silvestri (1930) n’ayant rien signalé de particulier chez la 2 qu'il a décrit; je n’ai pu étudier que deux st.III B et un individu présentant des caractères de st.III B, mais avec une papille génitale d sans phanere. CERQUES Un peu moins longs que la partie normalement découverte du tergite 10. Cerque droit: marge interne rectiligne, sans dent, mais avec des denticules unisériés proximaux. Cerque gauche: régulièrement concave, avec 2 carènes sur la marge interne; 2- 3 denticules basaux nets, le reste des carenes avec des ondulations disparaissant vers l’apex des cerques. ? On trouvera la liste des abréviations et des rapports utilisés dans Pagés (1954, 1984) et Pagés & Schowing (1958). JAPYGIDES DU SUD-EST ASIATIQUE N° 9 739 AFFINITES Ce genre est bien caractérisé par ses antennes, la chétotaxie abdominale ter- gale, les organes glandulaires du premier urosternite, le tergite 10 et les cerques. Ces derniers se rapprochent de ceux de Rossjapyx Smith (Smith, 1962) et d’Hapljapyx Silv., celui-ci réuni aux Austrjapyx Silv. par Smith & Gonzalez (1964), tous les trois d’Amérique du Sud, mais l’armature des marges internes est nettement différente, ce qui joint aux autres caractéristiques détaillées ci-dessus permet de distinguer ces genres les uns des autres. DERIVATIO NOMINIS: WaALG,- doc = Ciseaux, par illusion aux cerques qui rappellent les deux lames tranchantes des anciens ciseaux connus sous le nom de "forces", dont on se servait, par exemple, pour tondre les moutons. Psalidojapyx murudensis (Silv.) comb. n. Figs 1-12 Matériel étudié: Bru-88/29: BRUNEI (Belait District): Sungai Liang, "Arboretum Forest Reserve", forét primaire ("Mixed dipterocarp forest"), prelevement de sol dans les angles formés par les contreforts de deux arbres appelés "Nyatho", 90 m; 21XI.1988; leg. B. Hauser; extraction par Berlese; 1 d | de 6, 25 mm. Je crois utile de décrire cet exemplaire trés jeune collecté a moins de 140 km a vol d’oiseau du Mont Murud (ca. 2438 m), station où a été capturée la 9 décrite par Silvestri (1930); on devrait pouvoir ainsi apprécier la variabilité des caracteres au cours du développement postembryonnaire et apporter quelques compléments a la description originale. TETE Vertex: Chétotaxie simplifiée; seuls les grands phanères A1-A4, S2, S4, S5 et S6, V2 et V4, M3 et MS, I2, [4 et I5, L1 et L4, PI et P2 sont différenciés, longs ou assez longs; seules les s6 à s9 sont reconnaissables; les autres grands phanères ou s sont soit absents, soit courts ou très courts. Antennes: la droite de 43 articles peu pileux; la gauche cassée, de 15 articles; aires pileuses sur les articles 14 à 21; 13 trichobothries typiques, a proximale, p = 0,44, du même type que celle d’un st.III B dont les autres caractéristiques se retrouvent pratiquement inchangées chez le dj (Pages, 1952, 1955); 6 sensilles placoides en position typique sur l’article apical; on note une sensille placoide sternale sur les articles 39 a 42; l’article apical étant de forme normale, conique, aussi long que large, je ne crois pas qu’il s’agisse d’une antenne en régénération, mais peut- être d’un caractère propre a murudensis (cf. Pages, 1952). Pieces buccales: typiques de la famille; les cing lames du lobe interne des maxilles pectinées, la premiere pourvue d’une dizaine de dents; palpes labiaux environ 2 fois aussi longs que large ä la base, les soies subapicales sont en moyenne 1, 50 fois (v. ex. = 1,42 - 1,56) aussi longues que le palpe qui les porte. 740 JEAN PAGES Fics 1-8 Psalidojapyx murudensis (Silv.), 3 ,.— 1. Vertex, e = 361 um. — 2. Antenne gauche, articles 3 et 4 face tergale, e = 211 um. — 3. Mésonotum, e = 361 um. — 4. Tarse et prétarse de la PII droite, face antérieure, e = 158 um. — 5. Tergites 7 à 10 et les cerques, e = 361 um. — 6. Angles postérieurs gauches des tergites 6 et 7, e = 105 um. — 7. Détail de la base des marges internes des cerques, face tergale, e = 126 um. — 8. Détail de l’apex du cerque droit, e = 126 um. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 74] Fics 9-12 Psalidojapyx murudensis (Silv.), 6 |. — 9. Ursosternite 1, les pointillés représentent les aires sur lesquelles débouchent les glandes cuticulaires, e = 211 um. — 10. Detail de l’angle interne de l’organe subcoxal gauche, e = 63 um. — 11. Urosternite 3, e = 211 um. — 12. Papille génitale de ce d,,e= 63 um. THORAX Pronotum: les 5+5 M en position typique; le M, gauche est dédoublé; les 1+1 M, typiques sont tombés, cependant compte tenu de la taille des embases on peut estimer que le rapport M,/e est compris entre 1,5 et 2. Mesonotum. Préscutum: 1+1 M. Scutum: 6+6 M, les 5+5 M typiques et 1+1 M supplémentaires latéaux, insérés au-dessus des M,; les 5+5 soies assez longues habituelles. Métanotum: ne diffère du précédent que par la présence des seuls 5+5 M typiques. Pattes: longues, les PIII atteignent la limite des urosternites 4 et 5; peu pileuses, sauf les tarses; ces derniers présentent 4 paires de soies spiniformes tres fortes et très longues; aux PIII la longueur du tarse égale 0,65 fois celle du tibia; la griffe postérieure égale les 4/10 du tarse, 1,4 fois la griffe antérieure et plus de 6,5 fois celle de l’unguiculus. 742 JEAN PAGES ABDOMEN Tergite 1: Préscutum: 1+1 M assez longs. Scutum: 1+1 M (ma = M) longs, msa, M3, mp seuls decelables. Tergite 2:4+4 M (ma = M, M, M,_5) longs, msa et m, courts, m, assez courts, m et mp, ainsi qu'un sm entre M, et Ms, longs. Tergites 3 à 7: 6+6 M (ma = M, M; 5) longs; msa et m, courts; mz, M; et mp assez longs. Tergite 8: 2 fois aussi large que long (VL = 2, 04); 4+4 M longs, 3+3 homo- logables aux M, , et 1+1 latéraux postérieurs substernaux; 1+1 soies homologables aux M3, assez longues; m,, m} très courts, mp assez courts. Tergite 9: près de 3,5 fois aussi large que long (V/L = 3,41); pas de M, mais une rangée postérieure de 4+4 soies tres courtes entre lesquelles s’intercalent de minuscules sensilles. Tergite 10: a bords subparalléles, 1,5 fois aussi long que large; sans carene; 5+5 M, tous tombés, a grandes embases, dont 2+2 discaux et 3+3 latéraux antérieurs, intermédiaires et subpostérieurs; 2+2 longues soies, sublatérales subpostérieures et latérales postérieures; en outre 2+2 soies latérales assez longues entre les M latéraux antérieurs et intermédiaires. Angles latéraux postérieurs des tergites: nuls aux tergites 1 à 6 et 9; en pointe aiguë, petite, peu saillante au tergite 7; en pointe plus forte, obtuse au tergite 8. Acropyge: saillant, en demi-cercle. Longueurs relatives des segments 7 a 10: 43-36-19-100. Sternite 1: Préscutum: 4+4 M longs et 3+3 soies assez longues. Scutum: 11+11 M typiques, A, et C3 indifférenciés, A,, A, et A; très longs, les autres longs ou assez longs; B, et B, les plus courts de tous; 1+1 M supplémentaires entre C, et C,; de nombreuses soies assez longues ou assez courtes, localisées surtout sur les bords latéraux du sternite; les soies a embase circulaire caractéristique localisées en arriere des B, relativement nombreuses, sont de 2 tailles; les plus postérieures sont assez longues, dessinant une rangée régulière en avant des organes subcoxaux latéraux; elles sont séparées des autres, nettement plus courtes, par un espace assez important sur lequel s’ouvrent les orifices de très nombreuses glandes cuticulaires dont le diamètre du réservoir est d'environ 5 um; chez toutes les espèces de Japygidés, on en rencontre toujours quelques unes, très éparses, mais je n’avais encore jamais observé une telle concentration de ces glandes sur une aire aussi bien délimitée. Organes subcoxaux latéraux: occupant un peu plus de 1/3 de la largeur inter- stylaire, saillants, les soies glandulaires nettement éloignées de la rangée postérieure de soies à embase circulaire caractéristique; 32 soies glandulaires à droite, 37 à gauche, de taille inégales, irrégulièrement bisériées, SG / st] = 0,96 en moyenne (v. ex. 0,75-1,15), canal glandulaire un peu plus long que la soie glandulaire correspon- dante, a réservoir ovoide terminé par un petit goulot saillant; 26-27 soies sensorielles, SS/ st, = 0,40; SG / SS = 2,42 en moyenne (v. ex. = 1,9-2,9). Organe glandulaire médian: saillant, occupant environ le 1/6 de la largeur interstylaire; 11 minuscules sensilles setiformes (="pseudopori") sur l’opercule. JAPYGIDÉS DU SUD-EST ASIATIQUE N° 9 743 Sternites 2 à 6: 16+16 M typiques, Ag, B, et les C de rang pair (sauf C,) indifférenciés; les A, ainsi que les B,_3 et C, très longs ou assez longs; en général 5+5+5 soles assez courtes. Sternites 8: avec 7+7 M longs, répartis en 3 rangées longitudinales de 2, 2 et 3 M en partant de l’extérieur vers la ligne médiane: on note que les phanères situés entre les M postérieurs des rangées intermédiaire et médiale sont différenciés en M assez longs dont la taille de l’embase est à peine égale a la moitié de celle d’une des 7 M normaux; il en est de méme pour 2 phaneres insérés entre les M intermédiaires de la rangée mediale; ces 6 phanères sont donc des sm (cf. Pages, 1951). Paratergites 8: avec 1 M subpostérieur long, | soie postérieure assez longue. Sternopleurites 9: largement réunis par une courbe régulière; 1 M latéral pos- térieur assez long, | assez longue soie sublatérale postérieure et une rangée posté- rieure de 3-4 soies très courtes. Vésicules exsertiles: typiques aux urites 1 à 7. Styles: typiques; cône secondaire obsolète, simple renflement à la base du style, pore énigmatique bien net aux st, et devenant de moins en moins différencié, pour être à peine discernable aux st,; sj/st] = 0,34; st,/stz = 0.86; 5/57 = 0,94; s/st7 = 0,30. Papille génitale 3: par rapport à ce que j'ai défini comme caractéristique de la papille génitale du d | (Pages, 1955, 1967) celle de cet exemplaire s’en distingue par l’absence totale de phanères; compte tenu de la difficulté d’ observer ce qui existe sur la membrane reliant les urites 8 et 9 lorsque l’individu n’est pas en extension parfaite et sans élevages, il est impossilble d’affirmer qu’il peut y avoir un stade particulier entre un st.III et un à | tels que je les ai définis. CERQUES Un peu plus courts que la partie normalement découverte du tergite 10, LealLiodec =0,82; assez élancés, L/l = 2,4; recourbés à l’extrémité distale qui est aigué; sans dent différenciée. Cerque droit: marge interne rectiligne avec sur le quart proximal 5 denticules asymétriques, à sommet arrondi dirigé vers l’apex du cerque; le reste de la marge est pratiquement lisse, sauf tout a fait a son extrémité où s’observent 2 ondulations peu élevées. Cerque gauche: marge interne légèrement concave à la base, puis rectiligne; 2 carenes, l’inferieure présentant 2 denticules basaux peu saillants, puis à une certaine distance un troisième denticule très faible; le reste de la marge est intègre: la carène supérieure montre 6 denticules dont seuls les 2 premiers sont assez saillants, les autres s’estompant graduellement; le reste de la carène est intègre, sauf à son extrêmité distale ou s’observent 2 ondulations peu élevées, comme au cerque droit. Chétotaxie: une assez longue soie subantérieure, au niveau des premiers denticules; 8 longues soies le long du bord externe des cerques; je n’ai pas observé de M lateral antérieur. 744 JEAN PAGES REMARQUES Dans l’ensemble, les caractéristiques de ce d | sont parfaitement comparables à celles de la © décrite par Silvestri; la seule discordance importante concerne le nombre de M présents sur les notums thoraciques. D’après Silvestri (l.c.) on aurait la chétotaxie suivante en admettant que ses "setis longis" soient des M: pronotum: 5+5; mésonotum: 5+5; métanotum: 3+3; d’après nos observations le d | présente 5+5 M au pronotum, 6+6 M au mésonotum et 5+5 M au métanotum. Sans un réexamen du type et l’étude d’un plus abondant matériel il n’est pas possible de savoir, soit qu’il s’agit d’une divergence réelle, d’une évolution chétotaxique au cours du développement postembryonnaire, les individus les plus "âgés" ayant moins de M que les plus jeunes soit, plus vraisemblablement, que Silvestri qui, à l’époque, ne s’occupait pas de la forme des embases pour différencier les divers types de phanères, mais uniquement de leur taille, n’a pas pris en compte aux méso- et métanotum des M trop "courts" pour être comptés avec les "setis longis" et les a inclus dans ses "setis sat longis". Psalidojapyx edentulus sp. n. Figs 13-20 Matériel étudié: Sar-87/82: MALAISIE: SARAWAK: Bau, environs de "Fairy-Cave I", prélevement de sol dans les angles formés par les contreforts de grands arbres, 20 m; 13.X11.1987; leg. B. Hauser; extraction par Berlese; 2 st.Ill B de 6,64 (holotype) et 6,44 mm (paratype); holotype s’est cassé, lors du démontage de la préparation, en plusieurs fragments, conservés dans un tube d’alcool. TETE Vertex: les 4 grands phanères Al-4 assez longs; S2, S4, S6, M2-M3, MS, 13-15, L4 et PI-P2 longs; les autres assez courts ou absents; seules s6 et s9 présentes. Antennes: de 38 articles peu pileux; aires pileuses sur les articles 15 a 19; les 13 trichobothries typiques, a est une sensille sétiforme trés courte, submédiane, p = 0,90; 8 sensilles placoides en position typique sur l’article apical, il n’en existe aucun sur les articles précédents. Piéces buccales: typiques de la famille; les 5 lames du lobe interne des maxilles pectinées; palpes labiaux 3 fois aussi longs que larges a la base, les soies subapicales sont en moyenne 1,42 fois aussi longues que le palpe qui les porte. THORAX Pronotum: les 5+5 M typiques, les M, les plus longs, M; et M, les plus courts; la longueur des M, égale un peu plus de 1 fois un tiers l’écartement de leurs embases. Méso- et métanotum: identiques à ceux du d , de P. murudensis. Pattes: assez longues, les PIII atteignant la limite des sternites 3 et 4; 4-5 paires de soies spiniformes tres développées; le tarse des PIII égale les 2/3 de la longueur du tibia correspondant; la griffe postérieure mesure environ la moitié de la longueur du tarse, 1,4 fois celle de la griffe antérieure et 5 fois celle de l’unguiculus qui est peu aigu. ABDOMEN Tergite 1. Préscutum: 1+1 M. Scutum: 1+1 M (ma = M); seuls les mp, assez longs, sont présents. JAPYGIDÉS DU SUD-EST ASIATIQUE N° 9 745 Fics 13-17 Psalidojapyx edentulus sp. n., holotype. — 13. Vertex, e = 316 um. — 14. Mésonotum, e = 316 um. — 15. Tergites 5 à 10 et les cerques, e = 316 um. — 16. Angles postérieurs des tergites 6 et 7,e = 105 um. — 17. Detail de la base des marges internes des cerques, face tergale, au centre ceux de l’holotype, de part et d’autre ceux du paratype, e = 126 um. 746 JEAN PAGES Fics 18-20 Psalidojapyx edentulus sp. n., paratype. — 18. Urosternite 1, e = 253 um. — 19. Organe glandulaire médian, e = 63 um. — 20. Urosternite 3, e = 253 um. Tergite 2: 3+3 M (ma = M, M, ;); msa très courts; m, et mp assez longs. Tergites 3 à 5: 6+6 M (ma= M, M,_;), M, et M; les plus longs; M, assez longs; m, et mp très courts; m, assez courts. Tergites 6 et 7: 4+4 M (M, nuls); msa, m,, mp tres courts; m, et m, assez courts. Tergite 8: un peu plus de 2 fois (2,15) aussi large que long; 3+3 M homo- logables aux M3, M, et m3 = M; m, assez longs, mp courts. Tergite 9: environ 4 fois 1/5 aussi large que long; pas de M, mais 5+5 soies très courtes le long du bord postérieur. Tergite 10: à bords parallèles: près de 1 fois 1/2 aussi long que large; pas de carenes; 4+4 M longs dont 1+1 discaux antérieurs et 3+3 latéraux antérieurs, intermédiaires et subpostérieurs, 1+1 assez longues soies latérales postérieures: 10- 12+3-4+10-12 très courtes soies. Angles latéraux postérieurs des tergites: nuls aux tergites 1 à 5; en pointe aiguë, large, peu saillante aux tergites 6 et 7; saillants, larges, aigus au tergite 8 et à un moindre degré au 9ème. Longueur relative des segments 7 à 10: 36-38-17-100. Acropyge: saillant, trapézoïdal à angles arrondis. Sternite 1: Préscutum: 6+6 M longs et 4+4 soies assez longues. Scutum: 12+12 M typiques (B; nuls), très longs à longs; de nombreuses soies assez longues, prati- quement une entre chaque paire de M, plus nombreuses entre les A, et B,; soies à embase circulaire caractéristique localisées en arrière d’une ligne joignant C;-B;,-B;- C;; elles forment une rangée régulière en avant des organes glandulaires postérieurs. Organes subcoxaux latéraux: saillants, occupant environ le tiers de la largeur interstylaire; 9 à 11 soies glandulaires, les médianes plus longues que les latérales, JAPYGIDES DU SUD-EST ASIATIQUE N° 9 747 SG/st, =0,94 en moyenne (v. ex. = 0,7-1,20); 19-21 soies sensorielles assez courtes, SS/st] = 0,44 en moyenne (v. ex. = 0,39-0,49); SG/SS = 2,14 en moyenne (v. ex. = 1,72-2,41). Organe glandulaire median: peu saillant; occupant environ le 1/5 de la largeur interstylaire; 10-12 minuscules sensilles setiformes sur l’opercule. Sternites 2 à 7: 16+16 M (Ag, By, C;, Cs, Cg et Cy, indifférenciés) longs, A3, As, C4 et Cy les plus longs de tous; de nombreuses soies courtes ou très courtes. Sternite 8: les 7+7 M typiques longs, en position normale. Paratergites 8 et sternopleurites 9: identiques a ceux de murudensis. Vesicules exsertiles: typiques aux sternites | à 7. Styles: typiques, identiques a ceux de murudensis; s,/st, = 0,50; 5/57 = 0,96; st,/stz = 0,85; sj/st7 =0,43. Papille génitale: aucune trace n’a pu en étre décelée. CERQUES Un peu moins longs que la partie normalement découverte du tergite 10, Leg/L 10dec = 0,90; assez élancés, L/] = 2,69 en moyenne (v. ex. = 2,50-2,92); apex peu arqué, aigu. Cerque droit: marge interne sans dent, rectiligne; près de la base 5 denticules asymétriques peu saillants, arrondis, le reste de la marge avec de très faibles ondu- lations à peine indiquées. Cerque gauche: marge interne avec 2 carènes s’étendant pratiquement de la base à l’apex du cerque; la carène inférieure présente sur son tiers proximal un léger sinus ayant 2 denticules arrondis à sa base et 1 autre à sommet aigu à son extrémité distale; la largeur du cerque au niveau de ce denticule égale les 2/3 de la largeur à la base, le reste de la carène est intègre; la carène supérieure est absolument lisse à l'exception de 2-3 denticules arrondis à son début. Chétotaxie: identique à celle de P. murudensis. AFFINITÉS Cette espèce est proche du dj de P. murudensis. Elle s’en écarte par ses antennes, la chétotaxie des tergites abdominaux et ses cerques. DERIVATIO NOMINIS D’après edentulus = édenté, allusion à l’absence de dent différenciée aux 2 cerques. IL. Burmjapyx Silv., 1930 Parona (1892) signale deux espèces de Japygidés: 9 spécimens sont rapportés au Japyx indicus Oudemans, 1890 et deux autres à une espèce nouvelle, Japyx oudemansi Parona. Ces deux derniers spécimens ont été revus par Silvestri (1930) qui constate qu’il s’agit d’un d et d’une 9; il les répartit entre deux espèces pour lesquelles il crée le genre Burmjapyx caractérisé surtout par la structure très parti- culière des organes subcoxaux latéraux: B. oudemansi (Parona) pour le & à 28 articles 748 JEAN PAGES antennaires, nombre retenu par Parona dans sa description, et B. paronae Silvestri> pour la 2 qui possede 32 articles antennaires. En 1948 Silvestri décrit B. megurus d’Indochine (le Vietnam actuel). Enfin, Pages (1977) redécrit B. inferus (Carpenter, 1932) d’après un d et une 9. Avec B. lienhardi Sp. n. et B. murphyi sp. n., décrits plus loin, le genre Burmjapyx compte donc 6 espèces: pour les différencier, outre l’armature des cerques, qui est très utile, surtout celle du droit, la chétotaxie des tergites abdominaux 1 à 7 a aussi une grande importance: sur le tableau 1 sont indiqués les M et les sm diffé- renciés en M chez chacune de ces espèces: les chétotaxies de megurus, paronae et oudemansi sont données d’après les descriptions et figures de Silvestri (1930) et peuvent être sujettes à caution, surtout pour les deux dernières. En 1957, Paclt met 8 autres genres en synonymie avec Burmjapyx; j'ai dit à plusieurs reprises (Pagés, 1961, 1977, 1994) ce que je pensais de ce système qui, pour moi, est la négation de la systématique évolutive et qui ne fait que compliquer la recherche des affinités réelles entre les diverses lignées de Japygidés; je crois que nous ne connaissons même pas le dixième des taxons existants: je pense que la bio- géographie, entre autre, doit être considérée comme ayant une importance primordiale sur l’évolution des différentes lignées et la diversification des genres, surtout chez des Insectes dont l’hybridation ou l’interfécondité entre espèces paraît invérifiable pour le moment, les élevages d’assez longue durée, étant, pour l'instant, impossibles. De plus l’utilisation de la clef dichotomique des différents genres qu'il retient, aussi simplifiée qu’elle soit, ne semble pas permettre d'aboutir à une détermination correcte d’un de ses genres. Pour ne prendre qu’un exemple voici le cas du Burmjapyx huashanensis Chou, décrit du Shaanxi par Chou & Chen (1983). En tenant compte du très court résumé en espéranto et des figures 1-23 de la planche I, très peu précises et difficiles à exploiter, on arrive en utilisant la clef de Paclt, non pas à un Burmjapyx, sensu Paclt dont le cerque droit a des tubercules uniseriés, mais, puisque les 2 cerques de B. huashanensis sont à tubercules bisériés (fig. 23), à un Teljapyx, sensu Paclt; ce serait plus correct en faisant abstraction de l’absence d’une dent basale au cerque gauche, caractere bien spécifié par Silvestri (1948) dans la définition du genre. En fait, B. huashanensis, compte tenu des données de Chou & Chen, devrait 4 mon avis se rapprocher des Sinjapyx Silvestri (1948) (considéré comme un synonyme de Teljapyx par Paclt), et plus particulièrement de Sinjapyx davidoffi Silv. de I’ Annam. Seule une révision des types de ces espèces permettrait de pouvoir tenter de les classer génériquement. Burmjapyx lienhardi sp. n. Figs 21-34, 44 Matériel étudié: Sum-85/49: INDONESIE: SUMATRA (Sumatra Utara: Langkat): Réserve naturelle de Bukit Lawang, prés de Bohorok, forét primaire le long de la riviere, 180 m, prélèvement de sol dans les angles formés par les contreforts d’un grand arbre; 20.X1.1985, leg. B. Hauser; extraction par Berlese; 1d de 9,9 mm (holotype), 1 st.II B de 4,9 mm (paratype). 3 Il décrit aussi un Indjapyx indicus var. birmana nov., récoltés par L. Fea à Bahmo, sans indiquer s’il s’agit des spécimens étudiés par Parona et provenant de la même localité. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 749 Fics 21-25 Burmjapyx lienhardi sp. n., 4 holotype. — 21. Vertex, e = 549 um. — 22. PII droite, face tergale, e = 316 um. — 23. Tergites 7 à 10 et les cerques, e = 592 um. — 24. Détail de la marge droite du tergite 7, a noter la taille du m2, e = 316 um. — 25. Angles latéraux postérieurs des tergites 3 a 7,e=316 um. 750 JEAN PAGES Fics 26-28 Burmjapyx lienhardi sp. n., 3 holotype. — 26. Urosternite 1, sur la surface délimitée par des tirets s’ouvrent les très nombreuses glandes cuticulaires, e = 421 um. — 27. Organe subcoxal droit, les point représentent l’ouverture des glandes cuticulaires, e = 140 um. — 28. Urosternite Se — 4211 im: DESCRIPTION DU d TETE Vertex: tous les grands phanères sont présents: en outre de nombreuses soies de toutes tailles, longues à courtes parsement les quarts latéraux du vertex; enfin une multitude de minuscules sensilles criblent toute la surface, plus densément en avant des branches latérales de la suture en Y, sur le territoire en-dessous duquel se trouve le cerveau; chez de très nombreuses espèces de Japygoidea cette zone est plus fortement colorée que le reste de la capsule céphalique. Antennes: de 32 articles, la gauche du d est un régénérat de 31 articles; la chétotaxie est typique, chaque article portant deux verticilles de soies longues, voire assez longues et des sensilles recourbées assez longues; aires pileuses sur les articles 14 a 19; les 13 trichobothries typiques, a assez proximale, p = 0,46; 6 sensilles placoides en position typique sur l’article apical. Piéces buccales: typiques de la famille; les 5 lames du lobe interne des maxilles pectinées; palpes labiaux environ 4 fois aussi longs que large a la base, les soies subapicales sont en moyenne 1,5 fois aussi longues que le palpe qui les porte. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 75] Fics 29-34 Burmjapyx lienhardi sp. n., paratype (st.III B). — 29. Vertex, e = 303 um. — 30. Tergite 10 et les cerques, e = 253 um. — 31. Angles latéraux postérieurs droits des tergites 6 et 7, e = 105 um. — 32. Urosternite 1, e = 211 um. — 33. Detail de l’organe subcoxal latéral gauche et de l’organe glandulaire médian, les tirets représentent les glandes cuticulaires, e = 84 um. — 34. Urosternite Ole 211 (om: 752 JEAN PAGES THORAX Pronotum: les 5+5 M typiques, M, et M, les plus longs; les M, sont tombés, mais en tenant compte de la taille de leurs embases et de leur écartement on peut admettre que e est au moins égal à 2. Mésonotum: Préscutum: 1+1 M assez longs et 2+2 soies assez courtes. Scutum: 6+6 M dont les 5+5 M typiques et 1+1 M supplémentaires situés au même niveau que les M,, mais latéraux, insérés au-dessus des M3; 6-8+6-8 soies assez longues ou courtes. Métanotum: comparable au mésonotum, mais sans la paire de M supplémen- taires. Pattes: assez longues, les PIII atteignant le milieu de l’urosternite 4; très pileuses; 6-8 paires de soies spiniformes, on notera que la soie spiniforme postérieure de chacune de 3 ou 4 paires distales est nettement plus forte et colorée que les autres; calcars bien développés, le postérieur le plus fort; la longueur du tarse égale un peu plus de la moitié de celle du tibia (53/100); la longueur de la griffe postérieure égale environ de 1/3 de celle du tarse, | fois 1/5 celle de la griffe antérieure et près de 5 fois celle de l’unguiculus. Tergite 1: Préscutum: 1+1 M assez longs. Scutum: 2+2 M, (ma = M et Ms), les M, sont indifférenciés et de même taille que les autres sm, assez longs à assez courts. Tergite 2: 3+3 M (ma = M, M, 3) longs; les M; sont des sm identiques aux m, et m3, les seuls décelables, assez longs. Tergite 3: 6+6 M (ma = M, M,_;) longs; msa assez courts. Tergites 4 et 5: 6+6 M (ma = M, M, s, m, = M); les m, sont aux moins aussi longs que M, ou M3, mais indifférenciés; les msa sont présents et assez courts. Tergite 6: 5+5 M (M,_;, m, = M); les msa indiscernables. Tergite 7: 4+4 M (M,_5); ma, msa, M; et mp totalement absents; m minus- cules, m, et m3 assez courts. Tergite 8: environ 2,23 fois aussi large que long; 4+4 M longs, dont 3+3 homologables aux M3, M, et M4 et 1+1 latéraux postérieurs substernaux; 2+2 soies assimilables aux m, et m3, ces dernières longues: 2+2 soies courtes ou très courtes représentent les m, et mp. Tergite 9: a peu pres 3,40 fois aussi large que long; pas de M; 4-5 + 4-5 soies tres courtes le long du bord postérieur; ce tergite comme ceux des urites 6 a 8, est criblé de minuscules soies ou sensilles. Tergite 10: à bords subparallèles, 1 fois 1/4 aussi long que large; carènes bien développées, fortement colorées; 4+4 M longs dont 1+1 latéraux antérieurs, 1+1 latéraux intermédiaires et 2+2 discaux subantérieurs et subpostérieurs; 2+2 longues soles carénales, subantérieures et subposterieures; 3+3 soies latérales longues ou assez longues, subantérieures, intermédiaires et postérieures; 1+1 autres aux extrémités postérieures des carènes et 1+1 plus courtes correspondant au mp; de nombreuses soies très courtes. Angles latéraux postérieurs des tergites: nuls aux tergites 1 et 2; arrondis et saillants au tergite 3; en pointe progressivement plus saillante du tergite 4 au 8ème, nuls au 9ème. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 753 Acropyge: typique, peu saillant, a angles arrondis. Longueurs relatives des segments 7 a 10: 44-45-16-100. Sternite 1: Préscutum: 6+5 M longs; 3-4+3-4 soies assez longues. Scutum: 12+12 M typiques, C, apparemment absents; 2-3+4+2-3 M supplémentaires, dont 1+1 latéraux à hauteur des Bs, 1-2+1-2 latéraux au-dessus des C3; environ 10+10 soies assez longues, presque toutes le long des bords latéraux et 15+15 autres courtes ou très courtes: les soies à embase circulaire caractéristique occupent un territoire situé en arrière d’une ligne joignant C3, B,, B,, C,; elles sont assez longues et forment une rangée assez régulière en avant des organes glandulaires postérieurs du sternite. Sur tout le territoire occupé par ces soies à embase circulaire caractéristique on observe, comme chez le dj de Psalidojapyx murudensis (Silv.) au même emplacement, une multitude de minuscules glandes cuticulaires, mais plus petites, de 3 um en moyenne. Organes subcoxaux latéraux: caractéristiques du genre; tres saillants, ils occupent environ le 1/4 de la largeur interstylaire. Les soies a tres grande embase circulaire caractéristique du genre sont au nombre de 2, l’une latérale externe, l’autre insérée au-dessus de la vésicule exsertile, elles sont égales et à peine plus longues que le st, (1,05-1,08); chaque organe compte environ 60 soies glandulaires réparties en 2 groupes, interne et externe, où elles sont quadrisériées, réunis par un pont composé de soies bisériées, elles sont courtes, SG/st, = 0,22, à sommet obtus, semblables a celles que J'ai décrites chez Kinabalujapyx Pgs et Pauperojapyx Pgs, dont elles présentent aussi le même canal glandulaire et réservoir; 2 soies sensorielles courtes, SS/ st, = 0S3:SG/SS = 0:67: Organe glandulaire médian: très saillant, occupant environ le quart de la largeur interstylaire, à opercule en demi-cercle bien net sur lequel s’observent 22 minuscules sensilles sétiformes (= "pseudopori" de Silvestri), dont 2+2 près de la ligne d’articulation de l’opercule, les 18 autres sur 2-3 rangées transversales près du bord de l’opercule. Sternites 2 à 7: 18+18 M typiques longs; C4, €, différenciés en M, les autres C de rang pair et les B, indifférenciés; un M supplémentaire peut s’observer entre B, et B;; d’assez nombreuses soies courtes disposées plus ou moins par paire et dont une vingtaine forment une rangée continue le long du bord postérieur du sternite. Sternite 8: 7+7 M répartis en 3 rangées longitudinales de 2, 2 et 3 M en partant de l’extérieur vers la ligne médiane; le M intermédiaire de la rangée interne est nettement moins développé que les autres qui sont longs; on note aussi que les 3 M de cette rangée sont parfaitement alignés alors que typiquement le M intermédiaire est plus ou moins décalé vers l’exterieur. Paratergites 8: | M long près de l’angle postérieur et 5-6 soies courtes ou très courtes. Sternopleurites 9: se raccordant par une large courbe régulière: | M postérieur assez long et quelques soies insérées le long du bord libre, assez longues a courtes. Vesicules exsertiles: typiques aux urites 1 à 7. Styles: assez élancés, peu aigus, à cône secondaire petit, obtus, pore énigma- tique parfaitement différencié; sj/st] = 0,31; 5/87 = 0,72; stj/st7 = 0,78; s,/st; = 0,26. 754 JEAN PAGES Papille génitale 3: typique: restée invaginée sous le sternite 8 je n’ai pu l’etudier correctement; elle est beaucoup plus pileuse que celle que j’ai représentée chez Burmjapyx inferus (Carpenter) (Pagés, 1977), en particulier les aires antérieure et postérieure au gonopore; les soies sans embase bordant la lèvre antérieure du gonopore sont biseriées, tout au moins vers les angles latéraux de l’orifice génital; appendices génitaux bien développés, coniques, pres de 2 fois aussi longs que larges a la base, tres pileux. Papille genitale 9: inconnue. Fossettes glandulaires sternales des & : inexistantes. CERQUES Aussi longs que la partie normalement découverte du tergite 10, assez élancés, L/I = 2,44; par rapport à la largeur à la base, celle au niveau de la dent est égale à 1 au cerque droit et aux 3/4 au gauche; nettement arqués et à apex aigu. Cerque droit: à dent en triangle isocele, obtuse, nettement prémédiane, ry = 0,58; marge prédentale rectiligne avec 2 tubercules, le proximal trés peu saillant, le second arrondi; marge postdentale montrant a la suite de la dent 2-3 ondulations peu marquées, le reste de la marge intègre. Cerque gauche: a dent en triangle isocèle obtus, nettement distale, r, = 1,38; marge prédentale concave avec 7/8 tubercules arrondis; les 5 premiers supérieurs ont leurs bases pratiquement contigués, les 2 autres sont largement séparés; les 4ème et 8ème inférieurs sont très petits et aigus. Chétotaxie: typique; le phanère latéral antérieur de chacun des cerques étant tombé il ne m'a pas été possible d'examiner correctement l’embase pour savoir s’il s'agissait d’un M ou non. DESCRIPTION DU ST.III B Le st.III B présente un certain nombre de caractéristiques qui me paraissent très importantes à décrire pour mieux appréhender la variabilité des espèces de Bumjapyx. TETE Vertex: chétotaxie très simplifiée, de nombreux grands phanères sont absents, mais la série A est complete. Antennes: de 32 articles; a proximale, p = 0,50, valeur tres voisine de celle des adultes; aires pileuses sur les articles 15-19; 6 sensilles placoides. Piéces buccales: identiques a celles des adultes; palpes labiaux 2,6 fois aussi longs que large a la base, les soies subapicales sont en moyenne 1,25 fois aussi longues que le palpe qui les porte. THORAX Pronotum: les 5+5 M typiques: les M, sont aussi longs que l’écartement de leurs embases. Méso- et métanotum: préscutum et scutum typiques. JAPYGIDÉS DU SUD-EST ASIATIQUE N° 9 755 Pattes: les PIIT atteignent à peine le sternite 3; elles sont moins pileuses que chez les adultes, mais tout les phanères principaux sont présents y compris les soies spiniformes des tarses; la longueur du tarse égale les trois quarts de celle du tibia. ABDOMEN Tergites 1 et 2: identiques à ceux des adultes. Tergites 3 a 7: identiques a ceux des adultes; on note cependant que seul le ma = M droit existe au tergite 5 et que le tergite 6, normalement sans ma = M, présente aussi ce M a droite. Tergites § et 9: identiques a ceux des adultes. Tergite 10: a bords subparalleles, 1,28 fois aussi long que large; sans carene; 4+4 M identiques à ceux des adultes; peu de soies très courtes. Angles latéraux postérieurs des tergites: nuls aux tergites 1 à 5; en lobes arrondis dirigés vers l’arrière aux tergites 6 à 9. Acropyge: typique, très peu saillant. Longueurs relatives des segments 7 à 10: 41-44-19-100. Sternite I: Préscutum: 4+4 M longs et 3+3 soies assez longues. Scutum: les 13+13 M typiques, pas de M supplémentaires; toutes les soies à embase circulaire caractéristique sont localisées en arrières des C, elles forment une rangée assez régulière en avant des organes glandulaires postérieurs; un petit nombre de minus- cules glandes cuticulaires en arrière des C. Organes subcoxaux latéraux: caractéristiques du genre; très saillants ils occupent à peine le 1/5 de la largeur interstylaire; 2 soies à très grande embase circulaire d’égale longueur, mesurant | fois 1/3 la longueur du sf; 3 soies glandu- laires, typiques du genre; SG/st, = 0,5-0,6; 2 soies sensorielles très courtes, SS/st, = 0157SG/SS =3.0. Organe glandulaire median: typique du genre, occupant un peu plus du cin- quieme de la largeur interstylaire; 8 minuscules sensilles setiformes disposées sur 2 rangées longitudinales. Sternites 2 à 7: 14+14 M typiques, Ag, By et les C de rangs pairs (sauf C,) indifférenciés. Sternite 8: 7+7 M répartis en 3 rangées comme chez les adultes. Paratergites 8: identiques à ceux des adultes. Sternopleurites 9: se raccordant par une large courbe régulière; pas de M: 3-4 soies courtes ou très courtes. Vésicules exsertiles: typiques. Styles: peu allongées, à apex arrondi; cône secondaire et pore énigmatique peu différenciés; s/st = 0,42; st, /st7 = 0,90; 5 /st7 = 0,38; 5/57 = 0,83. CERQUES Un peu plus longs que la partie normalement découverte du tergite 10, Leg’ 10 gec = 1,14 (v. ex. = 1,12-1,6); assez élancés, L/I = 2,50; la largeur au niveau de la dent par rapport à celle à la base égale 0,89 au cerque droit et 0,68 au gauche; peu arqués, a apex peu aigu. 756 JEAN PAGES Cerque droit: a dent triangulaire très saillante à sommet obtus, prémédiane, ry = 0,72; marge prédentale rectiligne avec 2 forts tubercules arrondis; marge post- dentale concave avec | seul denticule triangulaire sur son tiers proximal. Cerque gauche: a forte dent triangulaire isocèle, à sommet assez aigu, post- médiane, r, = 1,43; marge prédentale avec 1/3 forts tubercules triangulaires, les inférieurs largement espacés; marge postdentale ne montrant que 2 ondulations sur sa moitié proximale, le reste de la marge intègre. Chétotaxie: typique: le M latéral antérieur présent. AFFINITÉS Cette espèce appartient au même groupe que paronae Silv.; elle est bien carac- térisée par la chétotaxie des urotergites et les organes subcoxaux. DERIVATIO NOMINIS Le Dr. C. Lienhard collabore depuis 1983 aux missions de recherche et de récolte d’Arthropodes en Asie du Sud-Est. Spécialiste mondialement reconnu des Psocoptères il a mis au point une technique de montage en préparation dont il a eu la gentillesse de me faire profiter, ce qui m'a été d’une aide fondamentale dans |’ étude de certains Diploures délicats. C’est pour l’en remercier que Je lui dédie cette espèce. Burmjapyx murphyi sp. n. Figs 35-43, 45 Matériel étudié: Bru-88/67: SINGAPOUR: Botanic Gardens, prélèvement de sol dans les angles formés par les contreforts d’un très grand arbre, au fond de la partie "Jungle", env. 20 m; 5.X11.1988; leg. B. Hauser; extraction par Berlese; 1 4 de 10,30 mm (holotype), 1 d de 9, 50 mm (paratype). TETE Vertex: on notera l’absence de plusieurs grands phanères et le faible déve- loppement de certains autres, en particulier ceux de la série L; de nombreuses soies assez courtes Ou courtes et une multitude de minuscules sensilles criblent, mais en plus faible densité que chez B. lienhardi sp. n., la surface du vertex. Antennes: de 32 articles a chétotaxie typique; aires pileuses sur les articles 12 a 18; les 13 trichobothries typiques, a proximale, p = 0,52; 6 sensilles placoides sur l’article apical. Piéce buccales: typiques de la famille; les 5 lames du lobe interne des maxilles pectinées; palpes labiaux 2,6 fois aussi longs que larges a la base, les soies subapicales sont en moyenne égales a 1,3 fois la longueur du palpe qui les porte. THORAX Pronotum: les 5+5 M typiques, M, les plus longs; la longueur des M, égale 1,8 fois l’écartement de leurs embases: 4+4 soies assez longues et 5-6+5-6 autres assez courtes ou courtes. Mesonotum: Préscutum: 1+1 M assez longs; 2-3+2-3 soies très courtes et une douzaine de minuscules sensilles sétiformes. Scutum: les 5+5 M typiques, M, les plus longs, M; les plus courts; 4-5+4-5 soies assez longues ou assez courtes. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 757 Fics 35-40 Burmjapyx murphyi sp. n., 3 holotype. — 35. Vertex, e = 515 um. — 36. Tergite 1, e = 632 um. — 37. Tergite 2, e = 632 um. — 38. Tergite 3, e = 632 um. — 39. Tergites 6 à10 et les cerques, e = 632 um. — 40. Angles latéraux postérieurs droits des tergites 4 à 8, e = 316 um. 758 JEAN PAGES Fics 41-43 Burmjapyx murphyi sp. n., 3 paratype. — 41. Urosternite 1, e = 253 um. — 42. 6 holotype, organe glandulaire médian, e = 158 um. — 43. d paratype, urosternite 3, e = 355 um. Métanotum: identique au mésonotum, sauf qu’il n’y a que 2+2 soles courtes et quelques soies très courtes. Pattes: assez longues, PIII atteignant le milieu de l’urosternite 4; chétotaxie très fournie, semblable à celle de /ienhardi en ce qui concerne les calcars et les soies spiniformes; la longueur du tarse est égale a 1,75 fois celle du tibia; la longueur de la griffe postérieure égale environ les 2/5 de celle du tarse, 1,29 fois celle de la griffe antérieure et plus de 6 fois celle de l’unguiculus. ABDOMEN Tergite 1: Préscutum: 1+1 M longs; 1+1 soies très courtes de part et d’autre des M. Scutum: 1+1 M (M) un peu plus longs que ceux du préscutum; M, = sm courts; environ 6+6 soies courtes ou très courtes. Tergite 2: 3+3 M longs (ma = M, M, s), Ms; les plus longs; msa assez courts, M,, My, m, et mp assez courts ou courts; une dizaine de soies très courtes sur chaque moitié du tergite. Tergite 3: 6+6 M (ma = M, M, ;), M, assez courts; msa, my, m, et mp courts, m, indiscernables de la vingtaine de soies très courtes recouvrant chaque demi-tergite. Tergites 4 à 6: 4+4 M (M, nuls). M; assez longs, les autres longs; msa, m; et my courts, à peine plus développés que les très nombreuses soies très courtes. Tergite 7: 3+3 M (M,, M3, M,), l'absence de M, est peu courante chez les Japygides; m, et m, seuls sm reconnaissables; de très nombreuses soies très courtes. Tergite 8: environ 2,40 fois aussi large que long: 3+3 M longs homologables à M,, My et m; = M: my, les seuls sm reconnaissables, assez courts; de nombreuses soies très courtes et une multitude de minuscules sensilles. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 759 Fics 44-45 Burmjapyx lienhardi sp. n., 3 holotype. — 44. Details des marges internes des cerques, e = 253 um. — Burmjapyx murphyi sp. n., 6 holotype. — 45. Details des marges internes des cerques, e = 253 um. Tergite 9: a peu pres 5 fois 3/4 aussi large que long; pas de M; une rangée d’une vingtaine de soies tres courtes le long du bord posterieur; de nombreuses sensilles minuscules. Tergite 10: à côtés subparalleles, 1 fois 1/3 aussi long que large; carènes très développées, fortement colorées; 4+4 M dont 1+1 latéraux antérieurs, 1+1 latéraux intermédiaires et 2+2 discaux, subantérieurs et subpostérieurs; 1+1 soies latérales postérieures assez longues; 1+1 soies sublatérales subantérieures et 1+1 latérales postérieures courtes; d’assez nombreuses soies très courtes dont 2 (subantérieure et intermédiaire) sur chacune des carenes; les très nombreuses minuscules sensilles habituelles. Angles latéraux postérieurs des tergites: nuls aux tergites 1 à 3; peu saillants et arrondis au tergite 4; obtus, dirigés vers l’arrière au tergite 5; très saillants, aigus au tergite 6, en pointe arrondie assez longue au 7ème, en pointe très étroite au 8ème, droits au 9ème. Acropyge: typique, saillant, a angles arrondis. Longueurs relatives des segments 7 à 10: 45-36-15-100. Sternite 1: Préscutum: 4-5+4-5 M longs; 4+1+4 soies courtes. Scutum: 12+12 M, C, apparemment nuls chez le d de 9, 50 mm; un M supplémentaire peut être présent entre A, et B;; les soies à embase circulaire caractéristique occupent un territoire situé en arrière d’une ligne joignant B;, B, et C,; elles sont plus ou moins nettement bisériées en avant des organes subcoxaux latéraux; sur l’aire située en 760 JEAN PAGES arriere de la série des B, occupée par les soies a embase circulaire caractéristique, on observe de tres nombreuses glandes cuticulaires comme chez lienhardi. Organes subcoxaux latéraux: tres saillants; ils occupent environ le quart de la largeur interstylaire. Les soies a tres large embase sont au nombre de 3 par organe, sauf chez le d de 9,5 mm chez lequel il n’y en a que 2 à droite, correspondant aux soies externe et médiane normales; leur taille par rapport au st, est tres variable et ne semble pas être un caractère spécifique très important; chez le d de 10, 30 mm la soie externe égale 1,70 fois la longueur du sf, à gauche et 1,46 a droite, la soie médiane égale à gauche 1,45 fois et à droite 1,32 fois celle du st,, quant à l’interne elle en égale 0,83 à gauche et à droite; chez le d de 9,5 mm, à l’organe droit, la soie externe égale 0,83, la médiane 1,29 et l’interne 0,56 fois celle du sf,, à l’organe gauche l’externe égale 1,66 et l’interne 1,39 fois la longueur du st,; environ 70 soies glandulaires par organe réparties en 2 groupes, ou elles sont tri- ou quadrisériées, reliés par un pont de 4-5 soies uni- ou bisériées; en moyenne SG/ st, = 0,22 (v. ex. = 0,18-0,25); 6 soies sensorielles par organe, SS / st, = 0,28 (v. ex. = 0,26-0,30); SG/ SS = 0,82 en moyenne. Organe glandulaire médian: saillant, occupant environ le tiers de la largeur interstylaire, a opercule net sur lequel on observe 22 a 23 minuscules sensilles sétiformes dont les 2+2 plus antérieures sont légerement plus longues que les autres. Sternites 2 à 7: 19+19 M typiques longs, By, Cs, Cg, Cio généralement indifférenciés assez courts ou courts: 2 M supplémentaires médians, le plus constant étant inséré entre les B,, 1 autre M supplémentaire entre B, et B3; une vingtaine de soies courtes ou très courtes sur chaque moitié du sternite. Sternite 8: 7+7 M répartis en 3 rangées longitudinales de 2, 2 et 3 M en partant de l’extérieur vers la ligne médiane; dans la rangée de 3 M, le médian est moins long que les autres et décalé vers l’extérieur par rapport aux 2 autres. Paratergites 8: 1 M long subpostérieur et 5-6 soies courtes ou très courtes dont la plus postérieure est la plus développée. Sternopleurites 9: se raccordant par une large courbe régulière; 1 M long postérieur sublatéral et quelques soies courtes ou très courtes le long du bord libre. Vésicules exsertiles: typiques aux urites | à 7. Styles: assez élancés, peu aigus, à cône secondaire peu développé à apex aigu, pore énigmatique peu différencié, petit; sj/stj = 0,40; s/s7 = 0,89; st/st7 = 0,84 (v. ex. = 0,81-0,87); s,/st = 0,33. Papille génitale d : restée invaginée sous le sternite 8 elle paraît identique au point de vue de la chétotaxie à celle de lienhardi; les soies sans embase bordant le bord antérieur du gonopore n’ont pu être observées: appendices génitaux coniques, près de 2,5 fois aussi longs que larges à la base, très pileux. Papille génitale © : inconnue. Fossettes glandulaires sternales des 3: inexistantes. CERQUES Pratiquement aussi longs que la partie normalement découverte du tergite 10; assez élancés, environ 2,5 fois aussi longs que larges à la base; par rapport à la largeur JAPYGIDES DU SUD-EST ASIATIQUE N° 9 761 TABLEAU | Chétotaxie des tergites 3 a 7 des Burmjapyx connus (sous toutes réserves pour paronae et oudemansi) inferus lienhardi murphyi megurus paronae oudemansi 762 JEAN PAGES a la base, celle au niveau de la dent est égale a 0,95 au cerque droit et aux 2/3 au gauche; relativement peu arqués, a apex aigu. Cerque droit: à dent en triangle isocèle, à sommet mousse, prémédiane, ry = 0,63; marge prédentale rectiligne avec 3 tubercules saillants arrondis sans intervalles entre-eux; marge postdentale pourvue de denticules arrondis, les 2-3 premiers les plus développés, les suivants décroissant graduellement de taille. Cerque gauche: a dent peu saillante, a sommet dirigé vers la base du cerque, distale, r, = 1,16; marge prédentale concave avec 9/8 tubercules régulièrement espacés chez l’holotype, 8/11 chez le paratype; chez les 2 d les 3-5 tubercules les plus proximaux de chaque rangée ont tendance à être aigus, les autres arrondis; marge postdentale subintègre présentant de faibles ondulations peu élevées. Chétotaxie: typique; 1 M latéral antérieur long. AFFINITÉS Cette espèce appartient au groupe paronae, mais se rapproche de B. inferus dont elle se distingue par la présence de 3 tubercules au lieu de 2, la chétotaxie des tergites abdominaux et les organes subcoxaux latéraux. DERIVATIO NOMINIS Je dédie avec plaisir cette espèce au Prof. Dr. D.H. Murphy de la National University de Singapour, pionnier de l’étude des Microarthropodes du sol à Singapour dont il a offert d’intéressants échantillons au Muséum de Genève. Une raison supplémentaire est qu'il cite le genre Burmjapyx dans son ouvrage (Murphy, 1973) sur la Vie animale des forêts à Singapour. III.- Pauperojapyx (Paucijapyx) kajan Pagés Fig. 46 Matériel étudié: Sar-87/86: MALAISIE: SARAWAK: route Kuching-Serian, pres du Kampong Kuap (a 18 km de Kuching), forét secondaire, prelevement de sol dans les angles formés par les contreforts d’un grand arbre, 30 m; 13.XII.1987; leg. B. Hauser; extraction par Berlese (1 st.III B de 2,6 mm, 1 © de 3,68 mm, 3 d de 3,32 mm, 3,92 mm et 4,60 mm). Bru- 88/41: BRUNEI (Belait District): Sungai Liang, "Arboretum Forest Reserve", forét primaire ("Mixed dipterocarp forest"), prélevement de sol dans les angles formés par les contreforts d’arbres appelés "Kempas" (= Kompassia malaccensis Maing. & Benth. (Fabaceae)), 20 m; 25:X1.1988; leg. B. Hauser; extraction par Berlese (1 st.III B de 1,84 mm, 1 9 de 3,04 mm, 3 d de 3,28 mm, 3,56 mm et 3,60 mm). Ces exemplaires correspondent parfaitement a la description originale de l’espece connue d’une seule station du Sarawak. Les st.Ill B possèdent déjà la chétotaxie abdominale typique des adultes, seul le nombre de soies courtes ou très courtes est réduit. Les organes subcoxaux montrent 1-2 soies glandulaires et 2-3 soies senso- rielles. Les cerques, aussi longs ou à peine plus longs que la partie normalement découverte du tergite 10 sont élancés, L/I = 2,98-3,00, leurs marges internes montrent une faible dent triangulaire proximale, ry = r, = 0,80 en moyenne, précédée de 1-2 minuscules tubercules et suivie de 3 faibles ondulations. JAPYGIDES DU SUD-EST ASIATIQUE N° 9 763 Fics 46 Pauperojapyx (Paucijapyx) kajan Pgs, st B de Sar-87/86. — 46. Tergite 10 et les cerques, e = 126 um. Chez les 2, en particulier chez celle de 3, 68 mm, on note que les sm des tergites sont sensiblement plus longs que chez les individus déjà connus; |’ armature des marges prédentales des cerques est plus variable, en effet, si celle de la 9 de 3, 68 um est identique à celle de la 2 paratypique, celle de la 9 de 3,04 mm présente au cerque droit 2 tubercules arrondis, peu saillants, très rapprochés l’un de l’autre et 2 tubercules largement espacés, le proximal beaucoup plus développé, au cerque gauche. Les d conformes à la description originale; on notera cependant que le 4 de 4,6 mm est pourvu de 9 soies glandulaires, et celui de 3,56 mm de 8 de ces soies à chaque organe subcoxal, alors que l’holotype (4,5 mm) n’en possède que 6 ou 7; 3 soies sensorielles comme chez le type; aux cerques les marges prédentales montrent 1-2 tubercules, le d de 3,56 mm en possède 3 rapprochés. REMERCIEMENTS Je tiens à remercier Mlle F. Marteau qui a reporté avec précision mes dessins sur calque et à Mme I. Juriens-Cottet qui a dû établir les frappes de mon manuscrit. BIBLIOGRAPHIE CHou, I. & CHEN, T. 1983. Studo de Japigedoj (V) (Dipluroj: Japigedoj). Entomotaxonomia 5: 327-341. MuRPHy, D. H. 1973. Animals in the Forest Ecosystem, pp. 53-73. In: S. H. CHUANG (ed.). Animal Life and Nature in Singapore. Singapore University Press, 302 pp. 764 JEAN PAGES PACLT, J. 1957. Diplura. Genera Insectorum 212: 123 pp. Paces, J. 1952. Japygidae (Insecta, Diplura) du Congo Belge. (Première note). Revue de Zoologie et de Botanique africaines 46: 345-372. PAGES, J. 1954. Japyginae (Japygidae, Insecta Diplura) de la Yougoslavie et des régions limitrophes. Glasnik Prirodnja Ckog Muzeja Srpske Zemlje (= Bulletin du Museum d’Histoire naturelle du Pays Serbe) 5-6: 235-264. PAGES, J. 1955. Biospeologica LXXV. Mission Henri Coiffait au Liban (1951). 5. Diploures Japygides. Archives de Zoologie experimentale et generale 91: 413-421. PAGES, J. 1961. Japygidae (Insecta, Diplura) de 1’ Afghanistan II. (Contribution a l’etude de la faune de l’Afghanistan. 58.). Comptes Rendus du 86° Congrès des Sociétés savantes, Montpellier, 1961: 735-750. PAGES, J. 1967. Données sur la biologie de Dipljaxpyx humberti (Grassi). Revue d’Ecologie et de Biologie du Sol 4: 187-281. PAGES, J. 1977. Dicellurata Genavensia III. Japygidés du Sud-Est asiatique. N°1. Revue suisse de Zoologie 84: 687-698. PAGES, J. 1984. Dicellurata Genavensia XIII. Japygidés du Sud-Est asiatique. N°4. Revue suisse de Zoologie 91: 425-436. PAGES, J. 1994. Japygidés du Sud-Est asiatique n°6: Malaysia (Sabah). Dicellurata Genavensia XIX. Revue suisse de Zoologie 100: 413-434. PAGES, J. & SCHOWING, J. 1958. Diploures Japygidés du Kivu et de l’Urundi (Congo belge). Revue de Zoologie et de Botanique africaines 57: 193-240. PARONA, C. 1892. Di alcuni Tisanuri e Collembola delle Birmania, raccolti da Leonardo Fea. Atti della Societa Italiana di Scienza naturali 34: 123-135. SILVESTRI, F. 1930. Contributions to a knowledge of the Indo-Malayan Japygidae (Thysanura). Records of the Indian Museum 32: 439-489. SILVESTRI, F. 1948. Contributo alla conoscenza degli Japygidae (Insecta Diplura). Rendiconti. Societa Italiana delle Scienze della Accademia dei Quaranta (ser. III) 27: 3-115. SMITH, L. M. 1962. Japygidae of South America. 3: Japygidae of Chile. Proceedings of the Biological Society of Washington 75: 273-292. SMITH, L. M. & GONZALEZ, R. H. 1964. Japygidae of South America 4: The Genus Austrjapyx and descriptions of new species. Annals of the Entomological Society of America 57: 159-164. REVUE SUISSE DE ZOOLOGIE 107 (4): 765-776; décembre 2000 Notes sur les Psélaphines néotropicaux (Coleoptera, Staphylinidae, Pselaphinae) — 11. Un nouveau genre et sept especes nouvelles de la tribu des Metopiasini André COMELLINI Museum d’histoire naturelle, case postale 6434, CH-1211 Geneve 6, Suisse. Notes on Neotropical Pselaphines (Coleoptera, Staphylinidae, Psela- phinae) 11. A new genus and seven new species of the tribe Meto- piasini. - The genus Metopiasoides is described to accomodate nine species, all described and/or redescribed: Metopiasoides brasiliensis sp. n., M. elongatus (Schaufuss) from Brasil, M. carinipes (Raffray) from Bolivia, M. costaricensis Sp. n., M. werneri sp. n. from Costa-Rica, M. decoratus sp. n., M. grandior sp. n., M. minusculus sp. n. from Panama and M. pecki sp. n. from Ecuador. A lectotype is designated for M. carinipes (Raffray). Key-words: Coleoptera - Staphylinidae - Pselaphinae - Metopiasini - taxonomy - Neotropics. INTRODUCTION La tribu des Metopiasini est constituée actuellement de huit genres, y compris ce nouveau, répartis dans la région néotropicale. Elle est caractérisée par la forme de la tête et par les antennes coudées, avec un scape particulièrement long (Sharp, 1887; Raffray, 1908; Park, 1942). Les espèces décrites ou redécrites ci-après forment un groupe homogene; elles sont réunies dans un genre nouveau. Les dessins des mémes structures sont a la méme échelle. Les abréviations suivantes ont été utilisées: CNCI = Canadian National Collection of Insects, Ottawa; DENH = University of New Hampshire Insect Collection, Durham; FMNH = Field Museum of Natural History, Chicago; MNHN = Muséum National d'Histoire Natu- relle, Paris; MHNG = Muséum d’histoire naturelle, Geneve. SYSTEMATIQUE Metopiasoides gen. n. Espèce type: Metopiasoides werneri sp. n. Ce genre est reconnaissable en premier lieu à ses antennes (fig. 1). L’article 3 de celles-ci est tres grand et les articles 2 et 4, bien plus courts, sont a peu pres de méme longueur; de plus, tous ces articles sont de méme largeur. Les trois premiers segments abdominaux apparents ont de larges rebords latéraux soulignés, à l’ex- Manuscrit accepté le 12.07.2000 766 ANDRE COMELLINI terieur, d’une petite carene assombrie. La pilosité est longue, fine, claire et légèrement courbe. Le tégument est lisse et brillant. Les tibias des pattes antérieures possedent, sur leur face avant, une longue carene longitudinale sombre tres en relief; celle-ci est située dans la partie supérieure concave de la patte. Les sternites 5, 6 et 7 forment chez les 5 une structure caractéristique très différente pour chaque espèce. Ce nouveau genre est proche de Metopias Gory,1832 (cf. Comellini, 1993: 921-927). Il en diffère surtout par la base des fémurs grêle, très nettement pédonculée (chez Metopias, fémurs élargis dès la base, non pédonculés), par les fossettes du vertex plus ou moins rapprochées, réunies dans une dépression commune ou reliées par un sillon transversal (largement séparées, reliées par un sillon transversal en arrière duquel se trouve une bosse arrondie assez grande) et par la moitié antérieure de l'abdomen subparallèle, les trois premiers tergites apparents subégaux, leur base nettement déclive (abdomen relativement court chez les Metopias, atténué des la base du 2°M© segment apparent; 1% tergite un peu plus grand, sa base nettement déclive). On peut distinguer deux groupes d'espèces dans ce genre Metopiasoides. Le premier, formé des grandes especes (M. brasiliensis sp. n., carinipes (Raffray), elon- gatus (Schaufuss), grandior sp. n. et M. pecki sp. n.), se caractérise par les fémurs postérieurs moins longuement pédonculés (moins de la moitié de la longueur du fémur), par l'apex des tibias postérieurs peu élargi, droit, par les tarses avec le 3ème article presque aussi long que le 2°™®, par la pubescence des élytres très longue, par les fossettes du vertex bien séparées, reliées par un sillon transversal, par les yeux formés de petites ommatidies, par l'article 9 des antennes soit un peu plus long que large (brasiliensis sp. n., carinipes (Raffray) et pecki sp. n.), soit presque deux fois plus long que large (elongatus (Schaufuss), grandior sp. n.). Le deuxième groupe, formé des petites especes (costaricensis sp. n., decoratus sp. n., minusculus sp. n. et werneri sp. n.), est bien caractérisé par les fémurs postérieurs très longuement pédonculés, par la partie apicale des tibias postérieurs élargie et un peu courbée, par les tarses avec le 3°M® article nettement plus court que le 2°M®, par la pubescence des élytres moins longue, par les fossettes du vertex très rapprochées, réunies dans une dépression commune, par les yeux formés de grosses ommatidies, enfin par l'article 9 des antennes au moins deux fois plus long que large. Metopiasoides brasiliensis sp. n. Figs 4, 12 Holotype ¢: Brésil, province de Sao Paulo, Station biologique Boracea (leg. J. M. et B. A. Campbell) (CNCI). Paratypes: 4 4, 1 2, même provenance (CNCI et MHNG). Téte avec deux fossettes profondes sur le vertex impressionant celui-ci trans- versalement. Un sillon longitudinal net partage le dessus de la téte de l’avant de la saillie antennaire au vertex. Yeux du d gros et saillants. Tégument ponctué, un peu granuleux sur l’avant et sur les côtés de la saillie antennaire. Pilosité plus dense et plus épaisse sur les côtés de la tête. Pronotum aussi long que large, cordiforme, avec un large sillon transversal au tiers postérieur, s’etendant sur toute la largeur, avec une fossette plus profonde au centre; une faible dépression longitudinale médiane part de l’avant du pronotum et se PSELAPHINES NEOTROPICAUX, 11. 767 Fics 1-11 1, Metopiasoides elongatus (Schaufuss), antenne. Dessus de la tête: 2, Metopiasoides carinipes (Raffray); 3, M. elongatus (Schaufuss). Derniers sternites des d: 4, M. brasiliensis Sp. n.; 5, M. costaricensis sp. n.; 6, M. decoratus sp. n.; 7, M. elongatus (Schaufuss); 8, M. grandior sp. n.9, M. minusculus sp. n.; 10, M. pecki sp. n.; 11, M. werneri sp. n. 768 ANDRE COMELLINI termine dans cette fossette. Tégument assez finement ponctué; une zone granuleuse transversale a l’avant et a |’ apex. Pilosité comme celle de la tête, plus dense et un peu plus épaisse sur les côtés. Elytres réunis nettement plus larges que longs. La suture des élytres est trés saillante, plus en avant qu’en arrière. Deux petites fossettes basales sur chaque élytre; l’une pres de la suture, dans une dépression de la base de l’élytre, l’autre plus petite a l’intérieur de la bosse humérale. Angles huméraux soulignés par une bosse allongée arrondie. Tégument finement et éparsément ponctué. Pilosité assez dense dirigée un peu obliquement vers l’arrière; cette pilosité est plus dense sur les côtés et à l’apex. Abdomen. Chaque tergite est fortement déprimé à la base sur toute la largeur de la partie centrale. Tégument finement et régulièrement ponctué. Pilosité plus dense et plus épaisse sur les côtés et a l’apex de chaque tergite. Derniers sternites des d (fig. 4). Longueur: d : 3, 25 à 3, 5 mm; l’unique 9: 3, 30 mm. Edéage: longueur 0, 73 à 0, 89 mm. (fig. 12). Cette espèce ressemble au M. carinipes par ses caractères généraux, en diffère nettement par la structure de l’édéage. Metopiasoides carinipes (Raffray) comb. n. Riss 22418 Metopias carinipes Raffray, 1904 Lectotype ¢: Bolivie, Yuracaris. L’étiquette porte comme indications: Yucaris / Ger- main / ex. Fauvel. Coll. Raffray (MNHN). Tête (fig. 2) avec un léger bourrelet transversal a l’arriere, fortement gra- nuleux; ce bourrelet est précédé de deux profondes fossettes réunies par un sillon. Yeux moyens, en cônes arrondis avec l’extrémité dirigée vers l’arriere. Tégument finement, irrégulièrement et peu densément granuleux-ponctué. Pilosité dense, recour- bée vers l’avant, sur les côtés de la tête. Tégument des antennes finement et assez éparsément granuleux. Pilosité de l’antenne assez longue, dirigée vers l’avant. Pronotum un peu plus large que long, cordiforme, partagé au tiers postérieur, sur toute sa largeur, par un profond sillon transversal. Tégument comme celui de la tête. Pilosité assez dense sur les côtés du pronotum, mais moins longue que celle de la tête. Elytres réunis un plus larges que longs avec deux fossettes basales profondes sur chacun. Un sillon longitudinal, plus profond en avant, partant d’une fossette basale, longe la suture sur toute sa longueur: un large dépression longitudinale partant de la seconde fossette basale, située plus près de la bosse humérale que de la suture, s’efface vers le milieu de l’élytre. Angles huméraux arrondis mais très marqués. Tégument très finement, irrégulièrement et éparsément ponctué. Pilosité assez dense sur les côtés, longue comme celle de la tête. Abdomen. Tergites avec les rebords latéraux relevés, garnis d’une longue pilosité recourbée vers l’arrière. Derniers tergites à forte pilosité dirigée vers l’apex. Tégument finement et éparsément ponctué. Longueur de l’unique exemplaire: 2, 8 mm. Edéage: longueur 0, 49 mm (fig. 13). PSELAPHINES NEOTROPICAUX, 11. 769 Metopiasoides costaricensis sp. n. Figs 5, 14, 15 Holotype d: Costa-Rica, Turrialba, Catie (alt. 600 m) (Leg. J.M. et B.A. Campbell) (CNCI). Paratype: 1 à, réserve de Montevarde (alt. 1500 m.) Costa-Rica (Leg. J.M. et B.A. Campbell) (MHNG). Téte avec une grande dépression sur le vertex a fond lisse, atténuée en avant et ornée en arrière de deux fossettes. Saillie antennaire avec, à la base, deux mamelons séparés par un sillon lisse, longitudinal, prolongé en avant presque jusqu’à l’extré- mité. Yeux des d gros et saillants. Tégument finement ponctué. Pilosité peu dense, dirigée vers l’extérieur. Antennes du d avec l'article 9 trois fois plus long que large. Pronotum plus long que large avec une dépression transversale bien marquée au tiers postérieur, s’etendant sur toute sa largeur et une faible dépression médiane longitudinale. Tégument avec une faible ponctuation irrégulière; une bande trans- versale plus ponctuée et granuleuse à l’avant et une autre à l’arrière du pronotum. Pilosité comme celle de la tête, un peu plus dense sur les côtés. Elytres réunis nettement plus longs que larges. Sur chaque élytre, à la base, une fossette pres de la suture et une autre vers le milieu de l’avant; ces fossettes marquent le départ de larges sillons longitudinaux, arrondis, s’effagant avant le milieu. Angles huméraux assez effacés. Tégument finement et irregulierement ponctué-granuleux. Pilosité comme celles de la tête et du thorax mais un peu épaisse, surtout sur les côtés et a l’apex. Abdomen. Rebords latéraux de chaque tergite saillants fortement a l’arriere sur le suivant. Pilosité dirigée ves l’ apex. Derniers sternites des d (fig. 5). Tibias postérieurs du d nettement élargis et un peu coudés un peu avant l'apex. Longueur des d: 2, 60 et 2, 75 mm. Edéage: longueur 0, 40 mm. (figs 14 et 15). Cette espèce peut être distinguée de M. brasiliensis et de M. carinipes par sa taille plus petite et ses antennes bien plus allongées. Metapiasoides decoratus sp. n. Figs 6, 16 Holotype d : Panama, Bocas del Toro (leg. H. Wolda) (FMNH). Paratypes: 5 6, même provenance (DENH et MHNG). Tête avec une dépression sur le vertex. Un faible sillon longitudinal au milieu de la saillie antennaire. Yeux des 4 gros. Tégument finement et irrégulièrement ponc- tue. Pilosité plus dense sur les côtés. Antennes du d avec l'article 9 deux fois plus long que large. Pronotum aussi long que large, cordiforme, avec un profond sillon transversal après le tiers postérieur, s'étendant sur toute sa largeur. Tégument et pilosité comme la tête; la pilosité est dirigée vers l’extérieur et est plus dense sur les côtés. Elytres réunis à peine plus longs que larges. Deux petites fossettes basales sur chaque élytre; l’une près de la suture au départ d’une faible dépression longeant celle- ci et s'étendant, en s’affaiblissant, jusqu’à l’apex; l’autre près de l’épaule au départ d’une faible dépression longitudinale s’effaçant déjà au tiers de la longueur de l’élytre. Angles huméraux assez effacés. Tégument finement ponctué. Pilosité un peu 770 ANDRE COMELLINI plus épaisse que sur la tête et le pronotum, assez dense, régulière, dirigée vers l'arrière. Abdomen. Rebords latéraux des tergites un peu bombés, plus larges en arrière qu’en avant. Tégument comme les élytres. Pilosité comme les élytres, dirigée aussi vers l’arrière, mais plus dense et plus épaisse sur les rebords latéraux et à l’apex de chaque tergite où elle forme une frange très nette. Derniers sternites des d (fig. 6). Longueur des d: 2, 12 à 2, 25 mm. Edéage: longueur 0, 36 mm (fig.16). Cette espèce ressemble à M. costaricensis, en diffère par la forme de l’édéage et par les tibias postérieurs à peine élargis dans la partie apicale. Metopiasoides elongatus (Schaufuss) comb. n. Fes sy Aalen Metopias elongatus Schaufuss, 1872 Holotype 9: Brésil, Amazone, Coll. Raffray (MNHN); 1 d, Bolivie, Yucaris, Coll. Raffray (MNHN). Tête (fig. 3) avec une dépression transversale à l’arrière. Yeux des d gros, en cônes arrondis, leur sommet dirigé vers l’arrière. Tégument très finement et épar- sément ponctué. Pilosité plus épaisse sur les côtés. Antenne (fig. 1).Tégument des antennes finement granuleux. Pilosité des antennes dense, orientée vers l’extrémité. Pronotum un peu plus large que long, cordiforme. Au tiers postérieur, un pro- fond sillon transversal s’étendant sur toute sa largeur; une dépression longitudinale, plus ou moins marquée, part du milieu de l’avant et se termine au sillon transversal. Tégument comme celui de la tête. Pilosité dense. Elytres réunis plus larges que longs. Deux fossettes basales peu profondes sur chaque élytre. Un sillon longitudinal partant de la fossette centrale longe la suture sur toute sa longueur; une légère dépression longitudinale part de la deuxième fossette et s’efface déjà vers le tiers antérieur de l’élytre. Angles huméraux arrondis, assez marqués. Tégument finement, irrégulièrement et éparsément ponctué. Pilosité plus dense sur les côtés et à l’apex, dirigée vers l’arrière. Abdomen. Rebords latéraux des trois premiers tergites apparents terminés à l’angle postérieur extérieur en une dent obtuse plus ou moins forte. Bord postérieur de chaque tergite formé d’une large bande de très petites alvéoles rectangulaires alignées et étagées sur plusieurs rangs; cette bande alvéolaire est plus large au centre. Tégument finement et irrégulièrement ponctué. Pilosité latérale et apicale dense, plus épaisse et plus courte que celle des élytres; pilosité du dessus fine et éparse. Derniers sternites des d (fig. 7). Longueur des deux exemplaires à : 4 mm. Edéage: longueur 0, 75 mm (figs 17 et 18). Les édéages des deux seuls spécimens connus ne sont pas indentiques; je ne suis pas sûr qu'ils soient de la même espèce. Metopiasoides grandior sp. n. Figs 8, 19 Holotype d: Panama, province de Chiriqui, Cerro Colorado alt. 1475 m (leg. W. Suter) (FMNH). PSELAPHINES NEOTROPICAUX, 11. 771 Fics 12-18 Edeages: Metopiasoides brasiliensis sp. n., 12, face; M. carinipes (Raffray), 13 face; M. costaricensis sp. n., 14, face, 15, profil; M. decoratus sp. n., 16, profil; M. elongatus (Schau- fuss), 17, face, 18, profil. 7/72 ANDRÉ COMELLINI Tête triangulaire avec un faible rebord en arrière du vertex. Angles postérieurs en angles aigus. Yeux moyens, saillants. Deux petites fossettes arrondies sur le vertex. Tégument finement et assez densément ponctué, finement granuleux sur la saillie antennaire et sur la Zone médiane du dessus de la tête. Pilosité dirigée vers l’avant, plus dense sur les côtés. Pronotum un peu plus long que large, en quadrilatere plus large en avant; les côtés sont arrondis et les angles sont saillants, les antérieurs plus fortement que les postérieurs; un profond sillon transversal sur toute la largeur au tiers postérieur; une faible dépression longitudinale médiane dans la partie antérieure. Tégument finement ponctué avec, par place, de fines granulations. Pilosité comme celle de la tête, mais dirigée vers le centre du pronotum. Elytres réunis un peu plus larges que longs. Sur chaque élytre, deux petites fossettes basales peu apparentes, l’une près de la suture, l’autre près de la bosse humérale. Angles huméraux soulignés par une forte bosse allongée arrondie. Tégu- ment finement ponctué, irrégulièrement granuleux à la base. Pilosité comme celles de la tête et du pronotum, mais un peu plus longue et dirigée vers l'arrière; plus dense sur les côtés et à l’apex. Abdomen. Tergites avec les rebords latéraux épais et obliques. A l’apex de chaque tergite, une forte bande transversale alvéolée. Tégument finement ponctué. Pilosité comme celle des élytres, dirigée en arrière, plus dense sur les rebords latéraux et à l’apex de chaque tergite. Derniers sternites du d (fig. 8). Longueur de l’unique d : 4, 20 mm. Edéage: longueur 0, 69 mm, mais l’extrémité est cassée (fig. 19). L’espece peut être facilement distinguée de ses congénères par la tête aux tempes anguleuses et par le pronotum aux angles antérieurs saillants. Metopiasoides minusculus sp. n. Figs 9, 20 Holotype d : Panama, Bocas del Toro (leg. H. Wolda) (DENH). Paratypes: 16 d , même provenance (DENH et MHNG). Tête avec une dépression triangulaire sur le vertex, la pointe dirigée en avant, avec deux petites fossettes à l’arrière. Un sillon longitudinal au milieu de la saillie anternnaire qui est soulignée à la base par un sillon oblique transversal. Yeux des d gros. Tégument finement et éparsément ponctué. Arrière de la tête plus foncé que le reste. Pronotum à peine plus long que large, cordiforme, avec un sillon transversal s'étendant sur toute sa largeur, après le tiers postérieur. Tégument et pilosité comme la tête; cette dernière plus dense sur les côtés. Avant et arrière du pronotum plus foncés que le reste. Elytres réunis plus longs que larges avec la base, l’apex, les côtés et la suture plus foncés que le dessus. Deux petites fossettes basales sur chaque élytre; l’une près de la suture, au départ d’une dépression longitudinale s'étendant jusqu’à l'apex de l’élytre, l’autre plus petite, au milieu de la base. Angles huméraux en bosses longi- tudinales arrondies. Tégument et pilosité comme la tête et le pronotum; la pilosité est plus dense sur les côtés et à l’apex et est dirigée vers l’arrière. PSELAPHINES NEOTROPICAUX, 11. Aa Fics 19-24 Edéages: Metopiasoides grandior sp. n., 19, face; M. minusculus sp. n., 20, profil; M. pecki sp. n., 21, face, 22, profil; M. werneri sp. n., 23, face, 24, profil. 774 ANDRÉ COMELLINI Abdomen. Les rebords latéraux sont un peu bombés. Tégument et pilosité comme le reste du corps, mais cette dernière est beaucoup plus dense et un peu plus épaisse à l’apex de chaque tergite. Derniers sternites des d (fig. 9). Tibias postérieurs du 4 particulièrement robustes, élargis peu après la base, ornés un peu avant l'apex de deux dépressions allongées profondes, l'une sur la face externe du tibia, l'autre sur la face interne; ces deux dépressions déterminent en leur centre une petite fenêtre transparente. Longueur des à : 2 à 2,2 mm. Edéage: longueur 0, 36 et 0, 38 mm (fig. 20). Espèce proche de M. decoratus, bien caractérisée par les tibias postérieurs du d et par l'édéage. Metopiasoides pecki sp. n. Figs 10, 21, 22 Holotype d: Equateur, Pastaza (alt. 1000 m.) à 25 km. au N. de Puyo (leg. S. et J. Peck) (CNCI). Paratypes: 1 4,3 2, même provenance (CNCI et MHNG). Tête avec une dépression transversale sur le vertex terminée de chaque côté par une fossette. Un sillon longitudinal bien marqué au milieu de la saillie antennaire. Yeux des d gros, en cônes arrondis, leur sommet dirigé vers l’arrière. Tégument assez densément granuleux-ponctué. Pilosité assez dense, dirigée vers le centre de la tête. Pronotum bien plus large que long, en parallépipède plus large en avant qu’en arrière. Un profond et large sillon transversal au tiers postérieur. Tégument assez finement et éparsément ponctué; à la base, une mince bande transversale densément guillochée. Pilosité comme celle de la tête, mais dense, dirigée vers l’extérieur et vers l'arrière. Elytes réunis un peu plus longs que larges. Sur chaque élytre, une fossette bien marquée près de la suture, au départ d’un profond sillon s'étendant jusqu’à l’apex, une autre fossette, au milieu de la base, prolongée par une courte dépression longi- tudinale très rapidement effacée. Angles huméraux soulignés par une forte bosse allongée. Tégument très finement et très éparsément ponctué. Pilosité comme celles de la tête et du pronotum, mais plus longue et dirigée vers l’extérieur et vers l’arrière. Abdomen. Tergites relevés de chaque côtés, dépassant latéralement les sui- vants. Apex de chaque tergite souligné d’une forte bande plus claire très finement alvéolée Le tégument est très finement et éparsément ponctué. Pilosité comme celle des élytres, encore un peu plus longue, dirigée vers l’arrière, plus dense à l’apex de chaque tergite. Derniers sternites des d (fig. 10). Longueur des d : 3, 38 et 3, 75 mm. Edéage: longueur 0, 76 et 0, 77 mm (figs 21 et 22) L'espèce réssemble au M. elongatus, en diffère par la pubescence plus longue et par la forme de l’édéage. Etymologie: Espèce dédiée au Dr Stuart Peck de l’Université Carlton d’Otta- wa, Canada. PSELAPHINES NEOTROPICAUX, 11. 775 Metopiasoides werneri sp. n. Figs 11, 23, 24 Holotype d: Costa-Rica, volcan Poas (alt. 1500 m) (leg. P. Werner) (MHNG). Paratypes: 15,2 2, méme provenance (MHNG). Téte avec une dépression triangulaire sur le vertex, la pointe dirigé en avant. Un sillon longitudinal au milieu de la saillie antennaire, effacé vers l’arrıere Yeux gros chez le d, très petits chez la 9. Tégument très finement et éparsément ponctué. Pilosité assez serrée, surtout sur les côtés de la tête. Antennes du d et de la 2 avec l'article 9 deux fois et demie plus long que large. Pronotum plus long que large, en ovale court, avec un profond sillon trans- versal au tiers postérieur s’étendant sur toute sa largeur et une faible ponctuation irreguliere; une bande transversale plus ponctuée et granuleuse a l’avant et une autre bande transversale identique à l’arrière. Pilosité assez dense, dirigée du centre vers les côtés, moins dense dans la zone longitudinale médiane. Elytres réunis aussi longs que larges. Deux fossettes basales sur chaque élytre, au départ d’un large sillon arrondi disparaissant déjà après le premier tiers de la longueur de celui-ci. Angles huméraux bien marqués chez le d, complètement effacés chez la 2. Tégument finement et densément ponctué-granulé. Pilosité comme celle du pronotum, mais un peu plus longue; plus dense sur les côtés et a |’ apex. Abdomen. Chaque tergite est bordé a l’apex d’une large bande transversale légèrement bombée plus ponctuée et granulée que le reste du segment, soulignée en avant et en arriere d’une mince ligne sombre. Pilosité comme celles du pronotum et des élytres, mais dirigée en arrière, plus dense à l’apex de chaque tergite et sur les côtés. Derniers sternites du d (fig. 11). Tibias postérieurs du d élargis dans la moitié apicale, ornés sur la face externe, un peu avant l'apex, d'une dépression allongée assez profonde. Longueur: d : 2, 50 à 2, 75 mm; 2: 2, 40 à 2, 55 mm. Edéage: longueur 0, 38 et 0, 40 mm (figs 23 et 24). Cette espèce ressemble au M. costaricensis, elle en diffère par la forme de l’édéage. Etymologie: espèce dédiée au Dr Philippe Werner, Ollon s/ Sierre, Valais, Suisse. REMERCIEMENTS Madame N. Berti, Paris et Messieurs J.M. Campbell, Ottawa, D.S. Chandler, Durham et J.A. Wagner, Chicago m'ont communiqués le matériel étudié. Monsieur G. Roth, Genève a mis au net les dessins et M. C. Besuchet, Genève, a pris la peine d'apporter quelques corrections et compléments à mon manuscrit. BIBLIOGRAPHIE COMELLINI, A. 1993. Notes sur les Psélaphides néotropicaux (Coleoptera). 8. Le genre Metopias Gory de la tribu des Metopiasini. Revue suisse de Zoologie 100: 921-927. PARK, O. 1942. A study in Neotropical Pselaphidae. Northwestern University Studies in Biological Sciences and Medecine, Number 1: 403 p., 21 pls. 716 ANDRE COMELLINI RAFFRAY, A. 1904. Genera et catalogue des Psélaphides. Annales de la Société entomologique de France: 1-400. RAFFRAY, A. 1908. Coleoptera. Fam. Pselaphidae. Genera Insectorum 64: 487 p., 9 pls. SCHAUFUSS, L. W. 1872. Beschreibung einiger Pselaphiden. Numguam Otiosus 2: 259-279. SHARP, D. 1887. Fam. Pselaphidae. Biologia Centrali-Americana. Insecta. Coleoptera, vol. 2 (1): 1-46, pl. 1. REVUE SUISSE DE ZOOLOGIE 107 (4): 777-791; décembre 2000 A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygosominae): resurrection of C. simulans (Vaillant, 1884) and description of a new species Wolfgang BOHME, Andreas SCHMITZ & Thomas ZIEGLER! Zoologisches Forschungsinstitut und Museum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygosominae): resurrection of C. simulans (Vaillant, 1884) and description of a new species. - The taxonomy of the West African scincid genus Cophoscincopus is reviewed. Based on external and genital morphology the so far monotypic genus can be partitioned into three very similar species. Besides Cophoscincopus durus (Cope, 1862) the name Cophoscincopus simulans (Vaillant, 1884) 1s available for one of these forms. The third, up to now, unnamed form is herein described as Cophoscincopus greeri sp. n. A key to the species is provided and informations about ecology and distribution are given. We revealed the presence of large, multicellular integumentary glands (located post-cloacally), which are described for the first time in the genus. Key-words: Reptilia - Scincidae - Lygosominae - Cophoscincopus - C. durus - C. simulans - C. greeri sp. n. - taxonomy - postcloacal glands - West Africa. INTRODUCTION The genus Cophoscincopus consists of small, semi-aquatic skinks which are known to inhabit the closed forests of Africa north of the Gulf of Guinea, from Sierra Leone and southern Guinea east to Togo (Müller, 1910; Mertens, 1933). Until recently the genus Cophoscincopus was considered to be monotypic. The single species C. durus was originally described as Tiliqua dura by Cope (1862) and was based upon a single West African specimen. Cope (I. c.) characterised this species, among other traits, by the presence of supranasals, a very tiny outer ear opening and strongly tricarinate scales on the back. A second West African lizard of similar habitus was described by Vaillant (1884) in his new genus Cophoscincus as C. simulans. According to the original description, this species is in part characterised by the absence of either supranasals or an outer ear opening and by tricarinate scales on the back, in which the middle keel is dominating in size. | Present address: Staatliche Naturhistorische Sammlungen Dresden, Museum für Tier- kunde, A.B.-Meyer-Bau, Königsbrücker Straße 159, D-01109 Dresden, Germany. Manuscript accepted 21.06.2000 778 W. BÖHME, A. SCHMITZ & TH. ZIEGLER Despite these seemingly clear differences Boulenger (1887) indicated that he thought Cophoscincus simulans to be a synonym of Tiliqua dura, but without further discussing this issue. Boulenger (l.c.) also transferred both forms to the genus Lygosoma, as the genus Tiliqua had been restricted to Australasian forms. In contrast to this point of view, Tornier (1901) considered Cophoscincus simulans and C. durus to be two distinct species within the genus Lygosoma. He argued that Lygosoma simulans differed in part from L. durum in the following characters: no supranasals, rostral in contact with the frontonasal, the latter not touching the frontal, no enlarged nuchals, no ear opening, three keels on the back scales, the middle one the largest. Müller (1910), after having received a larger series of new specimens from Liberia, discovered that a tiny ear opening was always present in both L. durum and L. simulans, though it can be often hidden by overlapping scales. Furthermore he found one specimen in which the internasal was divided by an azygous scale, which otherwise exactly fitted the description of Tiliqua dura Cope, 1862. He concluded that Cope had misidentified the divided internasal as two supranasals, and the azygous scale as an internasal. Also as Cope did not specifically describe the exact status of the keels on the dorsal scales, he considered the different descriptions of the middle keel as being within the limits of variability. Having discarded the principal differences between the two species, he concluded that Lygosoma simulans is a junior synonym of Lygosoma durum. Mertens (1933) reviewed Lygosoma durum and principially agreed with the arguments of Müller (1. c.). He nonetheless found the species to be distinct enough from the other known species of Lygosoma to put it in a genus of its own. As Tiliqua was still restricted to Australasian species, the next available name was Cophoscincus Vaillant, 1884. But Mertens also noted that this name was preoccupied by Cophoscincus Peters, 1867 (type species Lygosoma quadrivittatum Peters, 1867). Therefore he erected the new name Cophoscincopus, with the single species Cophoscincopus durus. This arrangement has been followed by all subsequent authors (e.g. Mittleman, 1952; Hoogmoed, 1973; Greer, 1974; Joger, 1981). The history of nomenclatural changes is summarized in the following list: 1862 Tiliqua dura Cope, Proc. Acad. Philadelphia 1862: 190. Type specimen: USNM 5996; western Africa. 1884 Cophoscincus simulans Vaillant, Bull. Soc. Philomath. 8 (7): 170. Type specimen: MNHN 6457; Crouacrou, Ghana. 1884 Cophoscincus simulans — Vaillant, Bull. Soc. Zool. France 1884: 349. 1887 Lygosoma durum — Boulenger, Cat. Lizards III: 304. 1893 Cophoscincus simulans — Matschie, Mitteil. deutsch. Schutzgeb. 6 (3): 4. 1901 Lygosoma simulans — Tornier, Arch. F. Naturgesch. 1901, Beiheft: 86. 1910 Lygosoma durum — Müller, Zool. Anz. 35 (9/10): 266-269. 1921 Lygosoma simulans — Chabanaud, Bull. Comité d'Etudes Hist. Sci. l'Afrique Occi- dentale Française: 445-472. 1933 Cophoscincopus durus — Mertens, Zool. Anz. 102: 188-190. 1938 Cophoscinopus durus — Loveridge, Proc. New England Zool. Club 17: 49-74. 1952 Cophoscincopus durus — Mittleman, Smithson. misc. Coll. 117: 1-35. THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 779 1973 Cophoscincopus durus -Hoogmoed, Die Aquarien- u. Terr.-Zeitschr. 5: 174-178; 7: 217-222, 8: 282-283. 1974 Cophoscincopus durus — Greer, Aust. J. Zool., Suppl. Ser. No 31; 67 pp. 1979 Cophoscincopus (durus) — Greer, Rec. Aust. Mus. 32: 339-371. 1981 Cophoscincopus durus — Joger, Bonn. zool. Beitr. 32: 297-340. Based on unpublished data kindly provided to us by Allen Greer, further investigations based on morphological and hemipenial data confirmed that the genus Cophoscincopus actually consists of three, though very similar, species. Whereas names are available for two of these forms, the third yet unnamed species 1s described below. MATERIAL AND METHODS Specimens included in the present study are kept in the following museums: BMNH - Natural History Museum, London; FMNH - Field Museum of Natural History, Chicago; HLMD - Hessisches Landesmuseum Darmstadt; MCZ - Museum of Comparative Zoology, Cambridge/Mass.; MHNG - Muséum d’histoire naturelle, Geneve; MNHN - Muséum National d'Histoire Naturelle, Paris; RMNH - Natuur- historisch Nationaalmuseum, Leiden; ZFMK - Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn; ZMB - Zoologisches Museum Berlin. Measurements were taken as follows: External morphology: Snout-vent length (SVL) from tip of snout to vent and tail length (TL) from vent to tail tip were measured with a ruler to the nearest 1.0 mm; other measurements were done with dial callipers and recorded to the nearest 0.1 mm. Genital morphology: Whereas the hemipenes of the holotype of Cophoscincopus greeri sp. n. were available to us in a completely everted condition, the available hemipenes of C. durus (ZFMK 36127-8, 36137-8, 36142-3, 36146, 36153) were only partially everted. Due to a new technique (Pesantes 1994, Ziegler & Böhme 1997) which allows to evert and study the hemipenes not only of fresh material, but also of specimens previously preserved in alcohol, it was possible to reconstruct the shape of completely everted hemipenes in Cophoscincopus durus of ZFMK 36128 (SVL: 4.8; TL: 4.5; HPL: 0.5 cm). Hemipenes of C. simulans were not available in the framework of the present study. Terminology of genital morphology follows Böhme (1988) and Ziegler & Böhme (1.c.). HPL - Hemipenis length, from cloacal base-point to apex. RESULTS RESURRECTION OF COPHOSCINCOPUS SIMULANS As explained above, the description of Cophoscincopus simulans as a distinct species was the direct consequence of Cope's misinterpretation of the head scales in his original description of C. durus and the fact that some of the used diagnostic characters are highly variable in the genus. Therefore it seemed justifiable to keep C. simulans as a synonym to C. durus until today. Our analysis of the new material, which was collected since the last revision of the genus by Mertens (1933), clearly 780 W. BOHME, A. SCHMITZ & TH. ZIEGLER demonstrates C. simulans not to be conspecific with C. durus. However, we are basing its diagnosis on additional key characters and the two species can consequently be differentiated as follows: Cophoscincopus durus (Cope, 1862) Tiliqua dura Cope, 1862: 190; type specimen: USNM 5996; western Africa. Diagnosis. Differs from C. simulans in the following combination of charac- ters: frontonasal longitudinally divided (comp. Fig. 1); prefrontals either usually separated by a medial azygous scale or, less frequently, in direct contact vs. usually separated or, less frequently, in direct contact in C. simulans (see Figs. 1-3 in Miiller 1910; Table 1, this paper); anterior loreal double; posterior loreal and ventral preocular distinct; size slightly smaller (max. SVL = 55 mm vs 61 mm for C. simulans). Cophoscincopus simulans (Vaillant, 1884) Cophoscincus simulans Vaillant, 1884: 349; type specimen: MNHN 6457; Crouacrou, Ghana. Diagnosis. Differs from C. durus in the following two characters: frontonasal entire; posterior loreal fused to lower preocular; Furthermore it is characterised by: paravertebral scales 39-48; external ear opening small but generally evident in lateral view (comp. Fig. 2). Remarks. Our new careful examination of the specimens also revealed for the first time the presence of post-cloacal glands in the genus Cophoscincopus. The openings of these glands are located along the central part of the first row of scales posterior to the soft scaleless lining of the cloaca. They lie either between the cloacal lining and a scale of the first row or between two scales. This area is normally covered by the pre-anal flap and this flap must be reflected and the area cleaned in order to see the gland openings to full advantage. They are identified most easily by their semi-solid, yellowish-brown exudate. The glands occur only in males and their openings are first identifiable grossly only in males approaching sexual maturity (on gross gonadal criteria). The post-cloacal glands can furthermore be used to differenciate between Cophoscincopus durus and Cophoscincopus simulans: In C. durus the gland openings occur between the scales of the first row and hence are rather obliquely elongate whereas in C. simulans they open anterior to the scales and are circular. In all three species the openings are largest medially and diminish in size laterally. DESCRIPTION OF A THIRD SPECIES Furthermore we found a third form which seems to be closely related to C. simulans, but being at the same time clearly distinct from it. For this species, there is no name available and it is described here as: THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 78] Fic. 1 Head portrait of Cophoscincopus durus (ZFMK 36141) 782 W. BOHME, A. SCHMITZ & TH. ZIEGLER Cophoscincopus greeri sp. n. Holotype: ZFMK 57599, 3, Mt. Nimba, 1800 m a.s.l., SE-Guinea, coll. Wolfgang Bohmettis 5 4s Vilas TE V7 EP: 0:67em: Paratypes. BMNH 1903.4.24.1-3: Abuasi, Ghana; BMNH 1930.6.9.14: Okurom River, Begoro, Akim, Ghana; BMNH 1921.11.12.6-7: N'Zebela, French Guinea; BMNH 1960.1.3.60: Tingi Hills, Sierra Leone; BMNH 1962.912: W.A.C.R.I., Tafua, Ghana; FMNH 190998: 48 km N of Koforidua, Tafo, Ghana. MNHN 20.128-130, MNHN 20.132 (two specimens), MNHN 20.135 (four specimens), MNHN 20.134 (one of two specimens): N'Zebela, SE Guinea; MNHN 1943.53-54: Yalenzou, SE Guinea; MNHN 1943.56-57: Kéoulenta, SE Guinea; MNHN 1951.118: Crete de Nion, 1500 m, Massif du Nimba, SE Guinea; MNHN 1951.124- 126, MNHN 1951.142: Ziéla, SE Guinea; MNHN 1951.134-137: Zougue, 1050 m, SE Guinea; MNHN 1963.404-405: Macenta, SE Guinea; MNHN 1963.412-416: Mt. Nimba, 1580 m, SE Guinea; RMNH 17242, RMNH 18806, RMNH 18809: Amedzofe, Togo; MCZ 51512: Bintumane Mts., Sierra Leone; ZFMK 20208-9: Mt. Tonkoui, Ivory Coast; ZMB 11251: Togo; ZMB 16061: Misahöhe, Togo. Other material: The following series was recently discovered far from the known area of the species (and genus!) and has therefore not been included in the type series: HLMD 2515- 19, S Niokolo Koba NP, SE Senegal. Diagnosis: A large species of Cophoscincopus (for definition of the genus see Mertens, 1933 and below) with a total maximum length of 165 mm. It differs from all other species of Cophoscincopus in the following combination of characters: frontonasal entire; posterior loreal fused to lower preocular (vgl. Fig 3); size slightly larger (max. SVL = 66 mm vs. 61 mm for next largest species); paravertebral scales 45-57, but only one specimen below 48 vs. 39-48 for other species; external ear opening very small and almost always hidden by overlapping scales. Body elongated, habitus + mabuiform with well developed extremities. Tail round, slightly flattened laterally. Head relatively pointed. Prefrontal in contact or widely divided by the frontal. A tiny outer ear opening is present but generally hidden by overlapping scales. Anterior loreal undivided. Dorsal head scales with weak longitudinal ridges or smooth. Dorsal neckscales weakly keeled. As in C. durus the gland openings occur between the scales of the first row and hence are rather obliquely elongate. Colour of dorsum dark brown, ventral colour whitish. The hemipenes of C. greeri sp. n. (Fig. 4a) differ from those of C. durus (Fig. 4b) mainly in lacking (1) well developed terminal lobes (the uniformly unpaired apex which Böhme [1988] stated for the hemipenes of C. durus was based on terminally not completely everted hemipenes), (2) a terminally distinctly divided sperm groove, and (3) a well defined collar-like ornamentation ("kragenartiger Ringsaum" sensu Böhme, 1988) at the upper trunk of the organ. Description of holotype: Snout-vent length: 59 mm, tail length: 77 mm. Body slender, elongated, tail round, slightly flattened laterally; degree of lateral flattening increases towards tail tip. Limbs well developed, long, pentadactyl; if adpressed to the body, slightly overlapping. Head relatively pointed, its width smaller than that of the back, only marginally set off from the latter. Distance of tip of snout from anterior edge of eye two-thirds of the distance from the posterior edge of the eye to ear opening. Latter tiny, generally hidden, about twice as wide as high. Frontonasal undivided, about one-third as wide as deep. Two undivided loreal scales. Anterior loreal small, triangular. Posterior loreal more or less quadrangular, about three times THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 783 Head portrait of Cophoscincopus simulans (MNHN 1967.187) (The subdivided frontoparietals of C. simulans are a unique variant). 784 reeri sp. n. (ZFMK 57599) THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 785 as long as high, four times as large as the anterior loreal, slightly narrowing towards the latter. Prefrontals widely separated from each other by the frontal, subtriangular, about as large as the nasal. Frontal longer than its distance from the tip of snout, narrowing to a tip posteriously, about as large as the parietals. Frontoparietals half the size of the frontal, equally long but twice as large as the interparietal. Latter narrowing to a tip posteriously. Parietals about twice as long as large, touching each other only in a small suture. A tiny round occipital scale present. 5-6 supralabials, the fourth bordering the eye. 51 paravertebral scales. Nearly no ridges on the head scales. Dorsal neck scales slightly keeled. Ventrals slightly enlarged. 67 subcaudals, number of subcaudal scale rows decreasing from 5 at the anal opening to | at the tailtip. Dorsal and lateral scales on tail strongly keeled. Dorsal colour (in preservative) dark brown, brightening laterally. On the lateral sides four rows of round white spots, which run from the neck to about the middle of the tail. Throat grey, slowly merging into the light colour of the ventral scales. Ventral side whitish, unspotted (in life, it was bright salmon-red: see Fig. 26 in Böhme, 1994). Dorsally, above the vertebral column two very weak rows of white spots, slowly disappearing from the neck to the anal region. Hemipenes slender, longish, partially transparent and apically slightly widened. At the upper truncus an inconspicuous, hardly recognizable collar-like ring (purposely slightly accentuated in Fig. 4a), which is closed around the truncus except for the sulcus spermaticus. The sulcus is surrounded by inconspicuous sulcal lips and widens laterally at the lower apex. Terminally the apex is medially slightly divided but is lacking well-defined lobes. At the sulcal side the terminal division of the apex turns into a longish flap nearly reaching the forking point of the sperm groove. No remar- kable structures were discernible at the asulcal surface of the apex. Besides the whole apical area is densely covered with delicate pustules, not discernible from Fig. 4a. Variation of paratypes: The paratypes agree generally well with the above description. Only in about 5% of the specimens examined the prefrontals are in contact with each other (see Tab. 1). In the living female paratypes, the colouration of the belly is distinctly sexually dimorphic: light yellowish instead of salmon-red in the male. Etymology: We dedicate the new skink to Dr. Allen Greer to acknowledge his valuable help, which was indispensible for the completion of this paper. - REDEFINITION OF THE GENUS A new, generalized definition of the genus Cophoscincopus can be given as follows: It belongs to the beta subgroup of the Eugongylus group sensu Greer (1974). It differs from all other members of this subgroup in the following combination of derived characters: the supranasals are absent; the upper preocular and prefrontal are in contact (they thereby separate the first supraciliary and posterior loreal); the external ear opening is small; there are usually six supralabials, the fourth lies usually below centre of eye; the dorsal scales are densely shagreened. - Type species: C. durus (Cope, 1862); Species included: C. simulans (Vaillant, 1884); C. greeri sp. n. W. BOHME, A. SCHMITZ & TH. ZIEGLER 786 Imm THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 787 KEY TO THE SPECIES OF COPHOSCINCOPUS I Frontonasal longitudinally divided; prefrontals either separated by an azygous median scale or in contact; anterior loreal double; posterior loreal and lower preocular distinct; 38-48 paravertebral scales........ durus - Frontonasal entire; prefrontals either separated by frontonasal-frontal or in contact; anterior loreal single; posterior loreal and lower preocular LESER EIN RE a e PL 2 2 Size slightly larger (max. SVL = 66 mm); paravertebral scales 45-57 (but only one specimen with fewer than 48); external ear opening minute and generally hidden by overlapping scales; on neck, dark colour of dorsum grades gradually into light ventral colour; dorsal head scales with only weak longitudinal ridges or none at all; dorsal neck sealessweaklyekeeledi..n. es san a N Sa greeri - Size slightly smaller (max. SVL = 61 mm); paravertebral scales 39-48; external ear opening small and generally evident; on neck, usually a black mid-lateral stripe slightly distinct from brown of dorsum and sharply distinct from brownish yellow of venter; head scales with moderately strong longitudinal ridges; dorsal neck scales moderately to stronelyakeeled es: una ee en en simulans DISTRIBUTION OF COPHOSCINCOPUS The genus Cophoscincopus is restricted to West Africa and is distributed from Senegal, Sierra Leone and southern Guinea east to Togo (Chabanaud, 1921; Barbour & Loveridge, 1930; Loveridge, 1938; Hoogmoed, 1973; Joger, 1981) (see Fig. 6). C. durus is known only from Sierra Leone, Liberia and southern Guinea. It is recorded herein for the first time from the Ivory Coast (Voucher specimen: ZFMK 68759, Tai National Park). C. simulans occurs from Sierra Leone, Liberia and southern Guinea east through the Ivory Coast and Ghana to Togo. C. greeri sp. n. 1s the most widespread of the three species: It is currently known from Senegal and Sierra Leone as well as from the extreme SE of Guinea eastwards through Ivory Coast and Ghana to Togo. It has not yet been recorded from Liberia. Fic. 4 a) Sulcal view of the right hemipenis of the holotype of Cophoscincopus greeri (ZFMK 57599), b) Left sulcal, right asulcal view of the left hemipenis of C. durus (ZFMK 36127) from Liberia (SVL: 4.9; HPL: 0.4 cm). The mode of completely everted lobes in the hemipenis of C. durus is indicated by the broken lines (in the case of completely everted lobes the lateral branches of the sulcus spermaticus are ending laterally of each lobe); not discernible from the hemipenis drawings is a delicate and dense pustular apical surface. 788 W. BOHME, A. SCHMITZ & TH. ZIEGLER TABLE | Comparison between the three species of Cophoscincopus in certain interspecifically variable characters. durus simulans greeri Prefrontals (% of NB) Meet 13 25 5 Separated by azygous scale 87 - - Separated - 15 95 N 179 79 20 Paravertebral Scales Range 38-48 39-48 (45) 48-57 Mean 43.7 43.0 51.4 SD: 1.62 1.89 2.24 N 100 45 48 Snout-vent Lenght (mm) Range 23-55 24-61 27-66 N 196 118 47 Tail Length (% SVL) Range 1.12-1.45 1.29-1.61 1.31-1.50 N 83 20 8 Size of gravid 9 9 Range 43-54 47-58 48-63 N 28 19 Ü Clutch Size | | - - 2 25 16 5 3) | 3 2 4 - - 2 There are several localities where the species occur in sympatry: All three species have been recorded from Kéoulenta and Ziela, Guinea; durus and simulans from Macenta, Mt. Nimba and Yalenzou, Guinea, and simulans and greeri from Amedzofe, Ghana. No doubt more thorough collecting would reveal additional cases. HABITATS, ECOLOGY AND REPRODUCTION OF COPHOSCINCOPUS Up to now, there is only little information available regarding the habitats and the autecology of Cophoscincopus. All species are mainly found in the vicinity of small creeks and pools in forested areas. The animals are active on the surface by day. If disturbed, they flee either into earth holes, under various rotten logs etc. or mostly into water. They swim under the water surface by lateral undulations with limbs adpressed to body and tail (Scherer in Miiller, 1910; Chabanaud, 1921; Bequaert in Barbour & Loveridge, 1930; Harley in Loveridge, 1938; Hoogmoed, 1973 and Joger, 1981). Their prey comprises, in part, termites and spiders (Loveridge, 1938). All species are oviparous with clutch sizes of 1-4 eggs (Loveridge, 1938; Hoogmoed, 1973 and Table 1, this paper). THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 789 IGES Cophoscincopus greeri sp. n., holotype in life. The holotype of the new species was collected at the Guinean side of Mt. Nimba at an altitude of ca. 1200 m a.s.l., in a fast-running, cold mountain creek within the submontane forest zone (among else characterised by the tree fern Cyathea manniana). Next to the specimen itself (Fig. 5), also its habitat has been figured by Bohme (1994), however, still using the name C. simulans. The same name was used for the two female paratypes from a partly cleared lowland forest site at Malweta river (Forét de Ziama, SE Guinea), with interspersed cocoa (Theobroma cacao) trees. Joger (1981: 329), who collected two ZFMK paratypes of C. greeri sp. n., reports that they "were collected in a cool, shallow creek, where they dived like newts, and hid themselves under stones". MNHN 1951.126 was collected in or on the edge of a "marigot"; MNHN 1951.142 has the word "swamp" associated with it in the registers, and MNHN 1951.134-137 the words "gallery forest". In view of the habitual, external overall similarity, it is surprising how many cases of broad sympatry have already been found (cf. Fig. 6). Clearly it would be interesting to learn more about the niche segregation of these highly specialized lizards. 790 W. BOHME, A. SCHMITZ & TH. ZIEGLER Liberia \., ) @ C. simulans A C.greeri FIG. 6 West Africa north of the Gulf of Guinea showing the distribution of the three species of Cophoscincopus. ACKNOWLEDGEMENTS First of all we wish to thank Dr. Allen Greer, Sydney for his indispensible help with an earlier draft of this paper. Furthermore, we thank Phyllis Koshland for providing us with drawings of the heads of the 3 species. Drs. E. N. Arnold, London, E. R. Brygoo, Paris, R. Giinther, Berlin, M. S. Hoogmoed, Leiden, R. F. Inger, Chicago, U. Joger, Darmstadt, and J. P. Rosado, Cambridge/Mass. made specimens of their respective collections available to us. M.-O. Rödel kindly donated some voucher specimens to the ZFMK collection. To all of them, we extend our deep gratitude. LITERATURE CITED BARBOUR, T. & LOVERIDGE, A. 1930. Reptiles and amphibians from Liberia (pp. 769-786). In: Strong's report of the Harvard-African Expedition upon the African Republic of Liberia and the Belgian Congo. Cambridge, Mass., Vol. 2. [Not seen by us.] BOHME, W. 1994. Frösche und Skinke aus dem Regenwaldgebiet Südost-Guineas, Westafrika. II. Ranidae, Hyperoliidae, Scincidae; faunistisch-6kologische Bewertung. Herpetofauna Weinstadt 16(93): 6-16. BÖHME, W. 1988. Zur Genitalmorphologie der Sauria: funktionelle und stammesgeschichtliche Aspekte. Bonner zoologische Monographien 27: 1-176. BOULENGER, G.A. (1887): Catalogue of the Lizards in the British Museum London 3: 575 pp. CHABANAUD, P. 1921. Contribution à l'étude de la faune herpétologique de l'Afrique occiden- tale. Deuxième note. Bulletin du Comité d'Etudes Historiques et Scientifiques de l'Afrique Occidentale Francaise: 445-472. Cope, E. D. 1862. On Lacerta echinata and Tiliqua dura. Proceedings of the Academy of Natural Science Philadelphia 1862: 189-191. THE WEST AFRICAN SKINK GENUS COPHOSCINCOPUS 79] Greer, A.E. 1974. The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, Suppl. Ser. 31, 67 pp. GREER, A. E. 1979. A phylogenetic subdivision of Australian skinks. Records of the Australian Museum 32: 339-371. HOOGMOED, M. S. 1973. Herpetologische Beobachtungen in Ghana. Die Aquarien- u. Terra- rien-Zeitschrift 5: 174-178; 7: 217-222, 8: 282-283. JOGER, U. 1981. Zur Herpetofaunistik Westafrikas. Bonner zoologische Beiträge 32: 297-340. LOVERIDGE, A. 1938. On a collection of reptiles and amphibians from Liberia. Proceedings of the New England Zoological Club 17: 49-74. MERTENS, R. 1933. Die Scincidengattung Cophoscincus Vaillant. Zoologischer Anzeiger 102: 188-190. MITTLEMAN, M. B. 1952. A generic synopsis of the Lizards of the subfamily Lygosominae. Smithsonian miscellanious Collections 117: 1-35. MULLER (or MUELLER), L. 1910. Uber Lygosoma durum (Cope). Zoologischer Anzeiger 35 (9/10): 266-269. PESANTES, O. S. 1994. A method for preparing the hemipenis of preserved snakes. Journal of Herpetology 28(1): 93-95. PETERS, W. 1867. Herpetologische Notizen. Monatsberichte der Königlichen Akademie der Wissenschaften Berlin: 13-37. VAILLANT, L. 1884. Catalogue raisonné des reptiles et batraciens d'Assinie donnés par M. Chaper au Muséum d'Histoire Naturelle. Bulletin de la Société Zoologique de France 1884: 343-354. ZIEGLER, T. & BOHME, W. 1997. Genitalstrukturen und Paarungsbiologie bei squamaten Reptilien, speziell den Platynota, mit Bemerkungen zur Systematik. Mertensiella, Rheinbach 8: 1-207. REVUE SUISSE DE ZOOLOGIE 107 (4): 793-834; décembre 2000 Review of the Lithoscirtus genus group (Orthoptera, Acrididae, Proctolabinae) with description of new species CAE ROWELL Zoologisches Institut der Universitat Basel, Rheinsprung 9, CH-4051 Basel, Switzerland. Review of the Lithoscirtus genus group (Orthoptera, Acrididae, Proc- tolabinae) with description of new species. - The genera Lithoscirtus Bruner, 1908, Drymophilacris Descamps, 1976, and Drymacris Descamps & Rowell, 1978, occurring in Costa Rica and Panama, are reviewed and in part redescribed. Five species are described as new: Lithoscirtus tesselatus sp. n., Drymophilacris glyphocerca sp. n., Drymophilacris nigrescens sp. n., Drymophilacris veraguensis sp. n., and Drymophilacris melanopsis sp. n. The previously unknown female of Drymacris panamae (Descamps) is described, and the species restored to Drymophilacris on the basis of female characters, giving the reinstated combination Drymophilacris panamae Descamps; Drymacris is thereby left monospecific, and is redescribed. New data on the distribution and coloration in life of pre- viously described species are presented. Keys to the three genera and to the species of Lithoscirtus and Drymophilacris are given. Key-words: taxonomy - Neotropics - Central America - Orthoptera - Acrididae - Proctolabinae. INTRODUCTION The term "Lithoscirtus genus group" is used here for three closely related genera, Lithoscirtus Bruner, 1908, Drymophilacris Descamps, 1976 and Drymacris Descamps & Rowell, 1978, belonging to the subtribe Lithoscirtae of the tribe Proctolabini. Phylogenetic analysis of mitochondrial ribosomal DNA genes (Flook and Rowell, in preparation) confirm that the three genera are a clade and each others' nearest relatives. The first species to be described (Rehn, 1905) were erroneously placed by the author in the genus Dellia, which is not a proctolabine but possibly a primitive ommatolampine, and is restricted to the Greater Antilles. Later Rehn (1929) combined these and other newly described species with the genus name Lithoscirtus and characterised that genus for the first time, the name having been created by Bruner (1908) without documentation. The genus Drymophilacris was split from Lithoscirtus by Descamps (1976), based on the shape of the male cerci and of the anterior margin of the endophallic sclerites. Subsequently, Descamps & Rowell Manuscript accepted 08.08.2000 794 C. H. F. ROWELL (1978) further split off from Drymophilacris the genus Drymacris, on the basis of differences in the male and female cerci, the female subgenital plate, the ovipositor and the elytra. To date there are 3 described species of Lithoscirtus, two of Drymacris and three of Drymophilacris, and the combined range of all three genera extends from northern Costa Rica to western Panama. Six of these are montane, but D. bimaculata and L. viceitas extend into the Caribbean lowlands. Early larvae of all three genera have a characteristic black coloration with an orange head and dorsal yellow stripe (Descamps & Rowell, 1978). The adult males are notable for the species-specific coloured patches, usually yellow or pale blue, on the subgenital and supraanal plates, which appear to serve in intraspecific communication (Rowell, 1983). All these species are small, flightless, brilliantly coloured grasshoppers of neotropical wet forest, especially of tree-fall clearings and forest edges, and all are foodplant spe- cialists, eating a few species of Solanaceae and in a few cases (Lithoscirtus minia- tulus, Drymophilacris nigrescens sp. n.) some Asteraceae as well. The spatulate dorsal ovipositor valves are associated with the habit of laying eggs in foam sandwiched between leaves of the foodplant, and not in the ground. In this paper I describe one new species of Lithoscirtus from Costa Rica and four new species of Drymophilacris, one from Costa Rica and three from Panama. All the new species are from montane forest of the Caribbean slope. Keys to these genera are given; their range is extended to Central Panama. The previously unknown female of Drymacris panamae Descamps is also described, and the species reassigned to Drymophilacris. Additional distributional and natural history records for the pre- viously described species of the three genera are given. Abbreviations of depositories: ANSP, Academy of Natural Sciences, Philadel- phia, USA; INBio, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; MHNP, Museum National d'Histoire Naturelle, Paris, France; RC, the author's collection; UMMZ, University of Michigan Museum of Zoology, Ann Arbor, USA; UP, Museo de Invertebrados G.B. Fairchild, Universidad de Panama, Panama. CHARACTERS OF THE GENUS GROUP The Lithoscirtae were recognized as a subtribe of the tribe Proctolabini by Amedegnato (1974, 1977), and this usage was continued by Descamps in his 1976 revision. Within the subtribe the Lithoscirtus genus group is separated from other genera by the incurved posterior edge to the pronotum and by its microptery (except for Drymacris nebulicola (Descamps, 1976)); also by the fact that the upper ovipositor valves are to a greater or lesser extent dorsoventrally flattened into spatula-like organs (e.g. Figs. 3A, 7A), which diverge distally (unlike the situation in Tela and Paratela, where the valves are also spatulate, but parallel). The fastigium is more or less produced, subtriangular in dorsal view, ending in a low ridge separating it from the flattened frontal ridge. The prosternal process is wide and low. The male supra-anal plate and the posterior rim of the 10th. abdominal tergite anterior to it are usually covered with long vertical white hairs with bent-over horizontal tips (e.g. Figs 1C, AB). NEW PROCTOLABINE GRASSHOPPERS 795 The male internal genitalia were first described and figured by Descamps (1976). All the Lithoscirtae have a flat, shield-shaped epiphallus (rather than disc- shaped, the term used by Descamps), devoid of ancorae and with small lophi (e.g. Fig. 2A). In the entire subfamily the ventrolateral sclerites are elongate and fused together at both ends, and posterior to the cingulum are united dorsally by a more or less sclerotised membrane. The resultant structure resembles the soleless upper of a shoe (e.g., Figs 2B, 5B): the cingular apodemes and endophallic processes project from the opening for the "ankle", the "heel" encloses the endophallus anteriorly, and the "toe" overhangs the aedeagal valves. Amédégnato (1977) pointed out that in the Litho- scirtae and most other proctolabines the anterior apodemes of the endophallus are lost, and replaced by very similar-looking structures derived from flattened projections of the central zone of the endophallus. For this reason the term "processes" rather than "apodemes" is used here). The cingulum is rather small and weak; paired lobes of thin, densely ornamented membrane run posteriorly from the rami, sheathing the lower aedeagal valves ventrolaterally midway to their tip (e.g. Figs 2B; 5B & D). The anterior processes of the endophallus are large, compressed laterally and provided with wide semimembranous margins (e.g. Figs 2B, 5B). The dorsal aedeagal sclerites are thin and bound by the sheathing membrane into a single medial dorsal aedeagal valve (e.g. Figs 2D, 5B & D), lying between the lower valves. In the Lithoscirtus genus group (as in the genera Tela and Paratela) the ventral sclerites of the aedeagus have a laterally directed hook (or hooks) at the tip (e.g. Figs 2B, D & E; 5B & D). KEY TO GENERA (external characters) MALES I Cerci long, only slightly incurved, with an abrupt embayment in the ventral edge at 2/3 of the length, tip laterally flattened and symme- trically rounded. Elytra cycloid, extending into 2nd abdominal segment and partially covering tympana............ Drymacris Descamps & Rowell - Cerci of various forms, but not as above. Elytra squamiform, extending maximally intommMetano turn’: Sr Ba Er oa LAN ay RTE ee 2 2 Cerci abruptly flexed inwards by some 70° at about half length, comple- tely or almost completely melanized; furcula absent (Fig. 1B-D) ER ED ALICI N TE IR AREA Lithoscirtus Bruner = Cerci curved smoothly inwards, forceps-like, usually only the distal region melanized; furcula absent or present (Figs. 4, 8,11,14 & 16, B-D IMAI CASES Ore ARS BEER AR NR N Drymophilacris Descamps FEMALES 1 Elytra cycloid, extending into 2nd abdominal segment and partially covering tympana. Cerci short, with blunt rounded tips (Fig. 19A, D). Dorsal ovipositor valves slightly excavated on dorsal surface, with raised internal edge (Fig. 19A, D). Subgenital plate flat (Fig. 19C) Ea bee eam) ER EN > CARDEN VICARI, 5 ORA DICI EEE Drymacris Descamps & Rowell 796 C. H. F. ROWELL = Elytra squamiform, extending maximally into metanotum. Dorsal ovi- positor valves flattened, spatulate. Cerci long and straight, with fine sharp points (Figs 3A, 7A, 10A, 13A, 18A). Subgenital plate domed or withipaired:swellings at anterior margin. i OR 2 2 Subgenital plate smoothly domed in anterior region (Fig. 3C). de DS A nr Nee Lithoscirtus Bruner - Subgenital plate with paired ventral or ventrolateral swellings near anterior margin (Figs 7B & C, 10B & C, 13B & C, 18B & C) Bere ee Pid ne ORS cid ts gets Been ee Drymophilacris Descamps Lithoscirtus Bruner, 1908 Bruner, 1908: 273. Type species: Dellia miniatula Rehn, 1905: 423. Rehn, 1929: 14. Amé- dégnato, 1974: 200. Descamps, 1976: 68, 77-80 (revision of genus). Amédégnato, 1977: 256. Amédégnato & Poulain, 1987: 400. Medium-small grasshoppers (>9.9, <20 mm in length); body size 14-17 mm (males), 17-19 mm (females). Hind femur 3.8-3.9X as long as pronotum, 0.20-0.23X as deep as long. Hind foot 0.39-0.46X as long as femur, second tarsal joint makes up 0.22-0.24 of foot. Interocular space narrow, 0.36-0.48X (males), 0.58-0.64X (females) as wide as antennal pedicel. Antennae long and filiform, 4.5-4.9 (males), 3.3-3.7X (females) as long as pronotum. Micropterous: elytra extend maximally to the anterior part of the metanotum. The thickened transverse ridge typically separating the fastigium from the frontal ridge in the Proctolabinae is obsolete in this genus (Descamps, 1976). Male characters. Male cerci short, thick, inwardly flexed at their tip, typically heavily melanized throughout (that of L. tesselatus sp. n. has however a green outer face proximally). Epiphallus (Fig. 2A) subrectangular, as long or longer than its greatest width, widening posteriorly towards the lophi, which are more erect than in Drymophilacris. The anterior processes of the endophallic sclerites are remarkable and diagnostic of the genus, being large thin irregular crumpled sheets of chitin, resembling miniature cabbage leaves (see Figs 2B-D). Ventral sclerites of the aedeagus rather short, dorsoventrally flattened, wide proximally in ventral view, tapering to the tip, ending in recurved hooks (Fig. 2B-E). Female characters (Fig. 3). Cerci straight, tapering to a fine point (as in Drymophilacris). Subgenital plate simple, triangular, but with concave posterior edges; the anterior region smooth and domed, without paired swellings. Posterior extremity of subgenital plate ("egg-guide") short, straight, laterally compressed and bluntly pointed in lateral view, about half as long as the visible portion of the ventral valves of ovipositor. Bursa copulatrix rather short and thin-walled relative to the other two genera. Lateral diverticulum of spermatheca with an initial wide chamber follo- wed by a thinner, rather simply convoluted appendage without secondary diverticula. NEW PROCTOLABINE GRASSHOPPERS 797 KEY TO SPECIES OF LITHOSCIRTUS (males) Males of this genus are traditionally separated on the complex shapes of the cerci (see Figs in Rehn, 1929 & Descamps, 1976, and Fig. | of this paper), which is undoubtedly the best method when dealing with museum specimens, but requires a microscope. The key given below is based on the colour differences of living animals, which are unambiguous for all except the two very similar species miniatulus and viceitas. These latter however are nowhere sympatric and when alive differ markedly in ground colour (though not when dried). Females are best identified by association with their males, to which they are similarly coloured; all females and no males have white-tipped antennae. A rarer bronze-coloured female morph exists in all species. 1 Apex of subgenital plate entirely black without other markings. Body generally black, but with dark green iridescence in life; paired dorsoventral spots at posterior margin of pronotum whitish yellow (not green, as in other spp.). All legs green. Cercus in side view vertically CRUINC ALE ARP RR tee N EEE ame L. daedalus Rehn, 1929, southern (nominate) race. Noßassabover. u. 22 Ba ee ER ale a MORI st eee 2 Apex of subgenital plate with paired coloured spots. Elytron extends INCOME CATT OM i RANCE NR RR DEI 3 - Apex of subgenital plate with an undivided pale blue medial patch (or a pair of medially confluent ones); femora of mesothoracic legs red, other legs green. Elytron does not reach posterior border of mesonotum......... 4 3 Apex of subgenital plate with paired pale blue patches. Fastigium and vertex markings, metallic blue green; femora of mesothoracic legs, red. Cercus in side view vertically truncate at tip. . ...... L. daedalus Rehn, 1929, northern race (cf. Descamps & Rowell, 1984). - Apex of subgenital plate with paired dull green patches (Fig. 1B). Legs all green. Cercus in side view (Fig. IC) tapering to a blunt, upwardly inflected tip, with a grey-green area near the base. Antennae black proximally and in distal half, but pale green in segments 2-7 (Fig. 1A). BENENNEN RENE RIED Heck OSEAN NTS PERG th TS orn ace Nc RIO L. tesselatus sp. n. 4 Colour in life principally metallic green. Cercus in side view obliquely truncateratitips (NE CostalRica) nen at re. L. miniatulus (Rehn, 1905) - Colour in life principally metallic blue. Cercus in side view tapering to POINT (S: BC ostaskacay Ne banana). ru ea L. viceitas Rehn, 1929 N | 1. Lithoscirtus miniatulus (Rehn, 1905) Dellia miniatula Rehn, 1905: 423. Bruner, 1908: 273. Kirby, 1910: 423. Lithoscirtus miniatulus Rehn, 1929: 19. Descamps, 1976: 77. Amédégnato, 1977: Figs 364-366. Holotype male: COSTA RICA: Prov. S. Jose: Carrillo, June 1903 (J.A.G. Rehn) (ANSP). 798 C. H. F. ROWELL Dimensions: see Table 1. Distribution: NE Caribbean slope of Costa Rica from at least 1700 m down to about 300 m altitude. The species is typical of montane forest and does not occur in true lowland rain forest, where it is replaced ecologically by Drymophilacris bimaculata. As noted by Rehn (1929), Bruner's locality of Pozo Azul (on the Pacific slope) 1s almost certainly false. Rehn listed specimens from Carrillo, Juan Vinas, and Guäpiles. Other specimens have now been taken from the Sarapiqui valley at Cinchona, La Virgen del Socorro and Chilamate (Provinces of Heredia and Alajuela); from the ridge between the Rio Peje and the Rio Sardinalito (Prov. Heredia); from Cerro Zurqui through the entire P.N. Braulio-Carrillo to Carrillo itself, and from the region around S. Jer6nimo, Alto la Palma and Bajo la Hondura (Prov. S. José); along the upper reaches of the Rio Toro Amarillo (Prov. S. José) down to Guäpiles, and from near Tucurrique (Prov. Limön); and from Turrialba and Chitaria (Prov. Cartago). This species is usually found on selected species of Solanum and Witheringia, but also occurs on Datura and (more rarely) on Verbesina and Vernonia spp. (Asteraceae). 2. Lithoscirtus viceitas Rehn, 1929 Rehn, 1929: 20. Descamps, 1976: 78. Holotype male: COSTA RICA: Prov. Limon: Suretka trail, between the Sixaola and Estrella Valleys, 19. May 1924 (Bradley JC) (ANSP). Dimensions: see Table |. Apart from the ground colour when alive (see Key) and the form of the male cerci, almost indistinguishable from L. miniatulus. The pregenicular part of the hind femur is thicker in viceitas, as noted by Rehn (1929), but the difference is only 0.05 mm and requires simultaneous comparison of the two species to be seen. Distribution: mid-elevation and lowland forest of the Caribbean slope, from the city of Limon (Costa Rica) south and east at least to Colon in Panama. The most widely distributed species of the genus, in many areas sympatric with one of several species of Drymophilacris. Rehn's (1929) localities were from the Suretka trail (Prov. Limon) and the Ujarras de Terraba (Prov. Cartago), both in montane forest in SE Costa Rica. It has since been found along the valley of the Rio Pacuare down to near Turrialba (Prov. Cartago), and throughout the valleys of the Rio Bananito, of the rivers of the Valle de la Estrella and of the Rio Sixaola almost to sea level (Prov. Limön). In Panama it has been recorded from Prov. Bocas del Toro (Changuinola, Guabita, and above Chiriqui Grande), Prov. Coclé (Cerro Copé; El Valle de Anton, Cerro Gaital), Prov. Panama (Cerro Campana) and Prov. Colon (Colön; Gatün; Porto Bello). The Panamanian localities range from 1100 to 20 m altitude. A strict specialist on a few species of Solanaceae. NEW PROCTOLABINE GRASSHOPPERS TABLE |. Dimensions of Lithoscirtus spp. and Drymacris Lithoscirtus daedalus L. miniatulus L tesselatus 799 Drymacris nebulicola L. viceltas MALES MEAN MEAN MEAN MEAN MEAN Dimensions in millimetres: N=5 N=3 N=3 N=3 N=3 F: Hind femur, length 8.90- 9.15 ( 9.06) 10.30- 9.33 ( 9.74) 8.07 - 10.20 ( 9.49) 10.03 - 10.58 (10.31) 10.40 - 11.44 (10.84) FD: Hind femur, depth 1.93- 2.08 ( 2.00) 2.13- 1.98 ( 2.04) 1.72- 2.15 ( 1.98) 1.96- 2.16 ( 2.07) 1.97- 2.07 ( 2.02) L: Rostrum-subgen. plate 14.64-14.72 (14.68) 15.79 - 14.94 (15.26) 12.32 - 16.98 (14.74) 16.14 - 18.04 (16.91) 15.06 - 15.78 (15.47) P: Pronotum (midline) 2.36. DAS 0239), 2:61 2149, (2:55))12.05 12772! (DAT) 12:59... 21721 (62165), 03:01 13.23. 03:13) Interocular space 0.15- 0.17 (0.16) 0.16- 0.14 ( 0.15) 0.16- 0.16 ( 0.16) 0.18- 0.21 ( 0.20) 0.15 - 0.19 ( 0.17) EE: (eye-eye) 3.19.3240 (3.18) 3232 3.13, ( 319) 2584. -" 336.037), #331922 3136. (6328), 3.402 3.432342) Fastigium 0.38- 0.52 (0.44) 0.49- 0.34 ( 0.42) 0.38- 0.40 ( 0.39) 0.40- 0.51 ( 0.44) 0.51- 0.56 ( 0.54) Antennal pedicel (width) 0.39- 0.44 (0.41) 041- 040 ( 0.41) 0.37- 0.41 ( 0.39) 0.39- 0.42 ( 0.41) 0.46- 0.50 ( 0.48) Antenna 10.14-11.43 (10.84) 13.37- 11.61 (12.47) 10.30 - 13.40 (12.12) 11.16- 12.65 (12.11) 11.60 - 13.07 (12.38) TI: hind tarsus 1 LOAM CIS) Ne OO IM as (000) NESS Mey CS) 5721748 01,64) T2: hind tarsus 2 0.86- 0.92 ( 0.89) 0.93- 0.87 ( 0.91) 0.76- 0.84 ( 0.80) 1.09- 12.15 ( 1.12) 1.35- 1.47 ( 1.43) T3: hind tarsus 3 MSC ES ED M PE 186 (1.96) 1.49 1,824. € 1.69) 21261221328 229) 2180 247. ( 2133) Ratios FD/F 0.21- 0.23 (0.22) 0.22- 0.21 ( 0.21) 0.20- 0.21 ( 0.21) 0.20- 0.21 ( 0.20) 0.18- 0.19 ( 0.19) F/P 3.73- 3.88 ( 3.79) 3.95- 3.75 ( 3.82) 3.75- 3.94 ( 3.85) 3.69- 4.01 ( 3.89) 3.31- 3.54 ( 3.47) L/P 5.98- 6.24 (6.14) 6.19- 5.76 ( 5.98) 5.66- 6.24 ( 5.97) 6.11- 6.63 ( 6.38) 4.80- 5.24 ( 4.95) Interoc./P 0.06- 0.07 (0.07) 0.06- 0.05 ( 0.06) 0.06- 0.08 ( 0.07) 0.07- 0.08 ( 0.07) 0.05- 0.06 ( 0.05) Interocular/pedicel 0.38- 0.43 (0.40) 0.39- 0.34 ( 0.36) 0.39- 0.43 ( 0.41) 0.43- 0.54 ( 0.48) 0.30- 0.41 ( 0.36) Fast/EE 0.12- 0.17 (0.14) 0.15- 0.11 ( 0.13) 0.12- 0.14 ( 0.13) 0.12- 0.15 ( 0.13) 0.15- 0.16 ( 0.16) (T1 + T2 + T3)/F 0.40- 0.42 ( 0.40) 0.43- 0.42 ( 0.42) 0.37- 0.40 ( 0.39) 0.45- 0.47 ( 0.46) 0.49- 0.50 ( 0.50) T2/ (T1+T2+ T3) 0.24- 0.25 (0.24) 0.43- 0.42 ( 0.22) 0.21- 0.23 ( 0.22) 0.23- 0.24 ( 0.24) 0.26- 0.27 ( 0.27) Ant/Pronotum 4.28- 4.67 (4.53) 0.23- 0.21 ( 4.89) 4.80- 5.02 ( 4.92) 4.10- 4.83 ( 4.57) 3.59- 4.34 ( 3.97) EE/F 0.34- 0.36 (0.35) 5.12- 4.55 ( 0.33) 0.32- 0.35 ( 0.33) 0.32- 0.32 ( 0.32) 0.30- 0.33 ( 0.32) FEMALES MEAN MEAN MEAN MEAN MEAN Dimensions in millimetres: N=3 N=3 N=3 N=2 N=3 F: Hind femur, length 10.73 - 10.93 (10.83) 11.03 - 10.20 (10.75) 8.56 - 10.82 ( 9.88) 11.10-12.11 (11.61) 11.59 - 12.36 (12.00) FD: Hind femur, depth DOTE 2.47, (239), 2352 2:21 (02:27) LOT = 2.42.0235), 2.26. 2.40.( 2:33), 2141) 2:46 (12.43) L: Rostrumsubgen. plate 16.74-18.58 (17.38) 17.38 - 18.28 (18.69) 14.30 - 18.63 (16.81) 17.86- 19.39 (18.63) 19.46 - 19.27 (18.86) P: Pronotum (midline) 2.81- 2.90 ( 2.85) 2.88- 2.84 ( 2.86) 2.44- 3.07 ( 2.74) 2.74- 3.14 ( 294) 3.61- 3.78 ( 3.68) Interocular space 0.21- 0.23 (0.22) 0.24- 0.21 ( 0.23) 0.21- 0.25 ( 0.24) 0.29- 0.32 ( 0.31) 0.26- 0.28 ( 0.27) EE: (eye-eye) Sis 1135451352) 3550, 3.34 N( 3140) 2.99.2347, (328) 355°. 35710856) 3412 3.61.3354) Fastigium 0.43- 0.56 ( 0.50) 0.46- 0.40 ( 0.42) 0.41- 0.52 ( 0.45) 0.46- 0.48 ( 0.47) 0.54- 0.65 ( 0.60) Antennal pedicel (width) 0.37- 0.39 ( 0.38) 0.40- 0.36 ( 0.38) 0.32- 0.36 ( 0.34) 0.35- 0.40 ( 0.38) 0.39- 0.46 ( 0.42) Antenna 9.43- 947 ( 9.45) 10.00- 9.67 ( 9.83) 7.89 - 12.40 (10.01) 10.15- 10.27 (10.21) 11 - 11.87 (11.40) TI: hind tarsus 1 1422 MSI AT) Ps 133 (135) 0446101230) 1.4953 1.5470152), 1.69 1.79, (1.75) T2: hind tarsus 2 0.98- 1.01 (1.00) 1.10- 0.97 ( 1.04) 0.71- 1.00 ( 0.87) 1.20- 1.30 ( 1.25) 1.42- 1.56 ( 1.50) T3: hind tarsus 3 2.07- 2.13 (2.10) 2.26- 1.85 ( 2.10) 1.56- 1.94 ( 1.80) 2.41- 2.60 ( 2.51) 231- 2.63 ( 2.49) Ratios FD/F 0.21- 0.23 (0.22) 0.22- 0.20 ( 0.21) 0.22- 0.23 ( 0.23) 0.20- 0.20 ( 0.20) 0.19- 0.21 ( 0.20) F/P 3.77- 3.85 (3.80) 3.86- 3.59 ( 3.76) 3.51- 3.80 ( 3.61) 3.86- 4.05 ( 3.95) 3.19- 3.38 ( 3.26) L/P 5.80- 6.54 ( 6.10) 6.03- 5.69 ( 5.83) 5.86- 6.48 ( 6.14) 6.18- 6.52 ( 6.35) 4.88- 5.27 ( 5.12) Interoc./P 0.07- 0.08 ( 0.08) 0.08- 0.07 ( 0.08) 0.08- 0.09 ( 0.09) 0.09- 0.12 ( 0.10) 0.07- 0.08 ( 0.07) Interocular/pedicel 0.54- 0.61 (0.58) 0.67- 0.53 ( 0.59) 0.66- 0.76 ( 0.70) 0.73- 0.01 ( 0.82) 0.61 - 0.72 ( 0.65) Fast/EE 0.12- 0:16 ( 0.14) 0.14- 0.12 ( 0.12) 0.12- 0.15 ( 0.14) 0.13- 0.13 ( 0.13) 0.15- 0.18 ( 0.17) (T1+T2+T3)/F 0.42- 043 ( 0.42) 043- 041 ( 0.42) 0.38- 041 ( 0.40) 0.76- 0.83 ( 0.80) 0.47- 0.50 ( 0.48) T2/ (T1+T2+T3) 0.21- 0.22 ( 0.22) 0.23- 0.23 ( 0.23) 0.21- 0.23 ( 0.22) 0.24- 0.24 ( 0.24) 0.26- 0.26 ( 0.26) Ant/Pronotum 3.26- 3.37 (332) 347- 3.38 ( 3.44) 3.23- 4.04 ( 3.63) 3.23- 3.75 ( 3.49) 3.04- 3.14 ( 3.09) EE/F 0.32- 0.33 (0.33) 0.33- 0.30 ( 0.32) 0.32- 0.35 ( 0.33) 0.29- 0.32 ( 031) 0.28- 0.31 ( 0.30) 3. Lithoscirtus daedalus Rehn, 1929 Rehn, 1929: 22. Descamps & Rowell, 1984: 155. Holotype male: COSTA RICA: Prov. Cartago: Navarro, 3800-3950 m, 24. July 1927 (Lankester CH, Rehn JAG) (ANSP). Dimensions: see Table 1. This Costa Rican species occurs in two distinct colour forms which are geographically separated (Descamps & Rowell, 1984). It is apparently confined to the upper reaches (above 1000 m altitude) of the Rio Reventazön and its trıbutaries. A strict specialist on a few species of Solanaceae. The ranges of the three preceding species adjoin each other in the neigh- bourhood of Turrialba. Nowhere are they known to be sympatric. 800 C. H. F. ROWELL Fic. 1 Lithoscirtus tesselatus sp. n., male. A, Habitus. B-C, external genital region, in axial, lateral and dorsal views. NEW PROCTOLABINE GRASSHOPPERS 801 Fic. 2 Lithoscirtus tesselatus sp. n., male. Internal genitalia. A. epiphallus. B, C, phallic complex in lateral and dorsal views. D, E, endophallus in dorsal and ventral views. Abbreviations: ape, anterior process of endophallus; dav, dorsal aedeagal valve; rc, rami of cingulum and associated membranes; vas, ventral aedeagal sclerite; vls, ventrolateral sclerite. 802 C. H. F. ROWELL FIG. 3. Lithoscirtus tesselatus sp. n., female. A-C, external genital region in dorsal, ventral and lateral views. D, ventral ovipositor valves and bursa copulatrix. E, spermatheca. NEW PROCTOLABINE GRASSHOPPERS 803 4. Lithoscirtus tesselatus sp. n. Holotype male: COSTA RICA: Guanacaste Prov.: Volcan Tenorio: nr. summit of rd. from Tierras Morenas to Bajo Los Cartagos. 1040 m, LN287250_426500, 21 July 1991 (Rowell CHF & Elsner N), specimen no. 91108. Allotype female: same data as holotype, specimen no. 91109 (both ANSP). Paratypes: As holotype, male, specimen no. 91107 (RC). Guanacaste Prov.: Tilarin, Zona Protectora Tenorio A.C.A., Rio San Lorenzo, 1050 m, LN287800_427600, May 1994 (G. Rodriguez), specimen nos. CRIOO1 873901 (male), CRIOOI 873902 (female); (INBio). Etymology: Latin tesselatus, inlaid with square mosaic stones, referring to the pattern of green and black markings. MALE (Fig. 1). Elytron squamiform, large for the genus, reaching to anterior margin of metanotum. Wing absent. Cerci (Fig. 1B-D) short, bent abruptly inwards halfway to tip at 45°, tip rounded, concavely excavated on ventral edge. Subgenital plate (Fig. 1B-D) with a short round tubercle medially and apically, at the junction of the plate and the pallium. Furcula absent. Supra-anal plate (Fig. 1D) longer than wide, ending in a blunt-tipped medial process. Internal genitalia (Fig. 2) generally typical of the genus. The hooks at the tips of the ventral valves are more robust and less curved than in the other species. Coloration: Antennae: scape, pedicel and first flagellar segment, glossy black; flagellar segments 2-7, pale green, 8 to 21 (tip), dull black. Eyes black (brown when dried). Palps green; mandibles, labrum and clypeus, glossy black. Rest of head basically green; inverted V on vertex, post-ocular stripe, ventral rim of frons, ventral and posterior rim of genae, antennal sockets, rostrum, and an hourglass-shaped mark on frontal ridge, with narrowest point ventral to the medial ocellus: glossy black. Disc of pronotum, glossy black. A pair of dorsolateral green spots in front of each sulcus, and a pair of yellow spots behind the third sulcus, reaching the hind margin. Lateral lobes glossy black, blotched with green ventrally, proepimeron black. Mesonotum black. Elytron, black, dorsal margin green-brown. Mesonotum black with central green blotch and paired yellow spots dorsolaterally at hind margin. Meso- and metepimera and episterna black dorsally, green ventrally. Front and middle legs yellow proximally, shading to green by the middle of the femur, each with 4 brown tibial spines. Claws and pulvilli black. Hind femur green, hind tarsus green proximally, blue-green distally, tibial spines (7 externally, 8 internally) and spurs brown, tipped black. Tarsi green, claws and pulvillus brown, tipped black. Abdominal segments with same pattern as metanotum, but with paired medial green marks, increasing in relative size towards the distal segments. Underside black, with a pale posterior rim to each segment. Genital region black. Supraanal plate (Fig. 1D) lemon yellow, bordered black, covered thinly with long white hairs. Cerci (Fig. 1B-D) mostly black but greenish proximally on outer surface. Subgenital plate (Fig. 1B, C) black, with paired green spots. FEMALE. Elytron squamiform, overlapping anterior margin of metanotum. Supra-anal plate (Fig. 3A) triangular, longer than wide, tip rounded, almost as long as 804 C. H. F. ROWELL FIG. 4 Drymophilacris panamae Descamps, male. A, habitus. B-D, external genitalia in lateral, dorsal and ventral views. dorsal ovipositor valves, transversely divided, with a shallow medial depression proximally. Subgenital plate (Fig. 3B, C) melanized in medial region, smoothly domed anteriorly, with paired patches of long hairs (in the position of the swellings of Drymophilacris), flat posteriorly, with a central semi-membranous area, posterior edges concave, apex ("egg-guide") short, not more than half the length of the ventral valves in ventral view; lateral lobes densely hairy. Internal genitalia (Fig. 3D, E). Typical of genus (see above). Basal external shoulder of ventral ovipositor valves sclerotized and provided with long lateral hairs. NEW PROCTOLABINE GRASSHOPPERS 805 Coloration other than in antennal and genital regions as in male, but markings somewhat less clearly defined and colours less brilliant. Antennae black, terminal 2 segments white, the preceding one reddish brown. Supraanal plate olive green, shading to olive brown, blackish at tip. Ovipositor valves, brown-black; dorsal face of upper valves, green. Subgenital plate blackish-brown, lateral lobes green. Dimensions: see Table 1. L. tesselatus is the largest species of the genus to date; its proportions, however, are normal. This species appears to be more closely related to L. daedalus than to the miniatulus/viceitas complex. The former has a trace of the protuberance at the apex of the dorsal edge of the male subgenital plate, seen in tesselatus, as well as the longer elytra. Natural History. On Solanaceae in clearings and at edges of mid-altitude montane forest in the Tilaran area. At first sight looks like a small example of Drymo- philacris monteverdensis, which has a very similar coloration and is geographically close. Males are however readily distinguished by the subgenital plate with its twin small green spots, that of D. monteverdensis is entirely pale turquoise blue. The pale proximal third of the male antenna is also distinctive; those of D. monteverdensis are entirely black. Drymophilacris Descamps, 1976 Descamps, 1976: 80. Type species: Dellia bimaculata Rehn, 1905: 426. Amédégnato, 1977: 256. Descamps & Rowell, 1978: 358. Amédégnato & Poulain, 1987: 400. Medium-small (>10, <20 mm) to medium (>20, <30 mm) sized grasshoppers: body length 14-19 mm (males), 17-22 mm (females). Hind femur 3.5-4.0X as long as pronotum, 0.20-0.24X as deep as long. Hind foot 0.42-0.48X as long as femur, second tarsal joint makes up 0.21-0.25 of foot. Interocular space narrow but somewhat wider than that of Lithoscirtus, 0.48-0.52X (males), 0.69-0.82X (females) as wide as antennal pedicel. Antennae 3.7-4.8X (males), 2.5-3.5X (females) as long as pronotum, thus tending to be slightly shorter than those of Lithoscirtus. Micropterous, elytra do not exceed posterior border of metanotum (D. panamae), and are still smaller in the remaining spp. Male characters. Cerci long, gradually curving inwards, forceps-like, nearly Straight or somewhat sigmoid in lateral view, with tips of variable shape. Rim of subgenital plate pinched together at apex and prolonged medially, like the lip of a jug. Endophallus has large regularly oval anterior processes, with doubly flanged margins, sharply differentiating the genus from Lithoscirtus. Ventral aedeagal sclerites thin and tapering in both lateral and ventral view, ending in one or more sets of recurved hooks. Epiphallus usually relatively broader than that of Lithoscirtus, nearer square than rectangular in dorsal view, less narrowed anteriorly, lophi less erect. Female characters. Cerci long and slender at tip, acutely pointed, similar to those of Lithoscirtus. Upper ovipositor valve flattened dorsoventrally, upper face flat (as in Lithoscirtus). Subgenital plate simple, triangular, posterior edges almost 806 C. H. F. ROWELL straight, with 2 more or less prominent swellings near anterior margin; in some species these form large laterally projecting bulges visible from above. Apex of subgenital plate ("egg-guide") of variable length, from short in D. bimaculata to equalling the ventral ovipositor valves in D. rubripes and D. nigrescens; tip upturned, straight or downturned in lateral view. Spermatheca similar to Lithoscirtus, but with a more complex appendage of the lateral diverticulum, usually provided with pouches and small diverticula (but simple in D. bimaculata). KEY TO SPECIES OF DRYMOPHILACRIS (adult males) 1 Subgenital plate black with a pair of yellow spots. Hind knees wholly orRpartlyzreddish brown. «0... 02g. 4 Vide 2.20. ILA Pe 2 - Subsemitaliplate not as above... ........ ..... 2.2.2.2... RENE 4 2 Clypeus, labrum, frontal ridge, frons, genae: yellow, lightly mottled with small black markings. Antenna in dorsal view black, minutely ringed with white at joints, terminal segment white. Furcula absent (Ganibbeanw@ostarRica): 2. 27.0 N eee D. bimaculata (Rehn, 1905) — Face not as above: at least clypeus, labrum and antennal sockets black. Eureulaspresent: 0728 LUEUR ARR PE 00000090 3 3 Frontal ridge, frons and ventral part of genae entirely black; terminal two segments of antennae white: furcula well-developed (Fig. 16) (CocleiProve;Panamia)? eo ee D. melanopsis sp. n. - Frontal ridge black only in dorsal region; remainder of frons and genae, yellow; spots of subgenital plate relatively small, covering only about half of total area; furcula small (Fig. 14) (Veraguas Prov., Panama). TARRA Se te ARE ete Sie re nl AES D. veraguensis sp. n. 4 Cercus with ventral distal extremity angled obliquely inwards and shorter thanidorsal'extremity (Ficsi4 & 8)... IO 5 Cereus.nobasiabove:. En er Se O TE 6 5 Subgenital plate black with broadly confluent green spots; supra-anal plate yellow-green; hind knees red; antennae black with indistinct pale annulations proximally; furcula absent (Fig. 8) (highlands of Central GOs tayRicay ae RR D. glyphocerca sp. n. _ Subgenital plate entirely pale blue, supra-anal plate yellow brown; all femora light reddish brown: furcula present but minute (Fig. 4) (highlands of SW Costa Rica and SW Panama). . D. panamae Descamps, 1976 6 Cercus long and nearly parallel-sided in lateral view, almost completely black: subgenital plate entirely turquoise, supra-anal plate emerald green in life, yellow green when dried; furcula absent (Cordillera del Norte CostayRica)g jas) aise D. monteverdensis Desc. & Rowell, 1978 — Cercus not as above; subgenital plate with confluent or narrowly separated blue-grey spots; furcula small but present. .................... 7 7 Supra-anal plate blue-grey; femora of middle legs red basally, blue distally; pronotum mostly green (lowlands, Bocas del Toro, Panama). IE rt alk LÉ SE ANT che eee RS D. rubripes Desc. & Rowell, 1984 NEW PROCTOLABINE GRASSHOPPERS 807 - Supra-anal plate yellow; femora of middle legs yellow basally, green distally; pronotum mostly black (Fig. 11) (highlands, Bocas del Toro, AAA) Re go Tee D. nigrescens sp. n. 5. Drymophilacris bimaculata (Rehn, 1905) Dellia bimaculata Rehn, 1905: 426. Bruner, 1908: 272. Lithoscirtus bimaculatus Rehn, 1929: 26. Drymophilacris bimaculata Descamps, 1976: 81. Amédégnato, 1977: 256, Fig. 346. Descamps & Rowell, 1978: 359. Rowell, 1983: 714. Holotype male: COSTA RICA: Prov. S. José: Carrillo, June 1903 (ANSP). The species differs from all others in the noticeably upturned tip to the dorsal aedeagal valve, as illustrated in Descamps (1976). The recurved tips of the ventral valves are also proportionately longer than in other species. Dimensions, see Table 2. A specialist on species of Solanum and Witheringia. Distribution. As indicated by Descamps & Rowell (1978), the species is widely distributed on the Caribbean slope of Costa Rica, from sea level to over 1000 m. It does not however seem to extend south of the Valle de la Estrella; though recorded from Puerto Viejo on the coast, it is absent from the valley of the R. Sixaola and in the adjoining area of Panama is replaced by D. rubripes. Its possible occurrence in the lowlands of Nicaragua is undocumented. 6. Drymophilacris monteverdensis Descamps & Rowell, 1978 Descamps & Rowell, 1978: 358. Holotype male: COSTA RICA: Prov. Puntarenas: Monteverde, 1400 m, LN 253500_448400, 10. Sept. 1975 (Rowell CHF) (MHNP). New material examined: COSTA RICA: Prov. Guanacaste: W side of Volcan Cacao, Estac. Mengo, 1400 m, LN 323700_376700, 11 July 1988 (Janzen D, Hallwachs W), specimen no. CRIOO1 013274, and other later specimens from same locality (INBio). Dimensions, see Table 2. A specialist on species of Solanum and Lycium. Previously recorded only from the region around the type locality, this species is now also known in a particularly large and robust form from Volcan Cacao, some 95 km to the NW. 7. Drymophilacris rubripes Descamps & Rowell, 1984 Descamps & Rowell, 1984: 154. Holotype male: PANAMA: Prov. Bocas del Toro; Changuinola District, United Fruit Co. Plantation, 3. Oct. 1925 (Walker FW) (UMMZ). New material examined: PANAMA: Prov. Bocas del Toro: Parque Internacional La Amistad, WEKSO-Region Teribé, 20.10.99 (DeGrazia L & Santos A), specimen nos. 99502 & 99503 (UP). Dimensions, see Table 2. 808 C. H. F. ROWELL TABLE 2. Dimensions of Dr. bimaculata Dr. monteverdensis Dr. rubripes Dr. panamae MALES MEAN MEAN MEAN MEAN Dimensions in millimetres: N=3 N=3 N=2 N=5 F: Hind femur, length 9.29 - 10.23 ( 9.88) 12.18- 11.95 (12.05) 9.11- 9.66 ( 9.39) 9.07- 9.93 ( 9.50) FD: Hind femur, depth 2.15- 2.27 ( 2.21) 2.51- 2.45 ( 2.48) 1.99- 2.04 ( 2.02) 1.90- 2.06 ( 1.97) L: Rostrum-subgen. plate 15.05 - 16.91 (15.88) 19.70 - 18.61 (19.26) 15.00 - 15.81 (15.41) 14.06 - 15.82 (14.82) P: Pronotum (midline) 2.46- 2.67 ( 2.59) 3:45-.32130 (03:32) 2.55 - 2.70 ( 2.63) 2.70- 2.93 ( 2.77) Interocular space 0.18- 0.19 ( 0.19) 0.27- 0.21 ( 0.24) 0.18- 0.20 ( 0.19) 0.19- 0.22 ( 0.20) EE: (eye-eye) 3.16- 3.48 ( 3.30) 3:82. 531697. 03511). 3.14- 3.35 ( 3.25) 3.16- 3.28 ( 3.23) Fastigium 0.38- 0.41 ( 0.40) 0.50- 0.47 ( 0.48) 0.19- 0.31 ( 0.25) 0.33- 0.59 ( 0.46) Antennal pedicel (width) 0.38- 0.40 ( 0.39) 0.55- 0.46 ( 0.49) 0.38- 0.41 ( 0.40) 0.41- 0.43 ( 0.42) Antenna 11.19 - 12.26 (11.66) 13.39 - 12.76 (13.07) 113720570159) 9.37 - 10.22 ( 9.61) TI: hind tarsus | 12715537 (1739) 1777-1700 173) 1.13- 1.27 ( 1.20) 1.24- 1.50 ( 1.33) T2: hind tarsus 2 0.80- 1.16 ( 0.98) 1.45- 1.40 ( 1.43) 0.92- 0.92 ( 0.92) 1.04- 1.20 ( 1.12) T3: hind tarsus 3 1.37 - 2.18 ( 2.05) 2:53 -.2:52°( 2.53) 1.76- 1.94 ( 1.85) 1.80- 2.00 ( 1.88) Ratios FD/F 0.21- 0.24 ( 0.22) 0.21- 0.20 ( 0.21) 0.21- 0.22 ( 0.21) 0.20- 0.22 ( 0.21) F/P 3.78- 3.88 ( 3.82) 3.82- 3.53 ( 3.64) 3.57- 3.58 ( 3.58) 3.25 - 3.66 ( 3.44) L/P 5.94- 6.33 ( 6.13) 6.22- 5.52 ( 5.82) 5.56- 6.20 ( 5.88) 5.20- 5.57 ( 5.36) Interoc./P 0.07 - 0.08 ( 0.07) 0.08 - 0.06 ( 0.07) 0.07 - 0.07 ( 0.07) 0.06 - 0.08 ( 0.07) Interocular(pedicel 0.45 - 0.50 ( 0.48) 0.51 - 0.46 ( 0.49) 0.47- 0.49 ( 0.48) 0.44 - 0.54 ( 0.48) Fast/EE 0.11- 0.13 ( 0.12) 0.14- 0.12 ( 0.13) 0.06 - 0.09 ( 0.08) 0.10- 0.18 ( 0.14) (T1+T2+T3)/F 0.42- 0.46 ( 0.46) 0.48- 0.46 ( 0.47) 0.42- 0.43 ( 0.42) 0.43- 0.48 ( 0.46) T2/ (T1+T2+T3) 0.20- 0.25 ( 0.22) 0.25- 0.25 ( 0.25) 0.22- 0.24 ( 0.23) 0.25- 0.27 ( 0.26) Ant/Pronotum 4.24 - 4.69 ( 4.51) 4.28- 3.78 ( 3.95) 4.54- 4.54 ( 4.54) 3.30- 3.77 ( 3.48) EE/F 0.32- 0.34 ( 0.33) 0.31 - 0.30 ( 0.31) 0.34- 0.35 ( 0.35) 0.33- 0.35 ( 0.34) FEMALES Dimensions in millimetres: Ni=3 N=3 N=2 N=4 F: Hind femur, length 10.57 - 11.82 (11.12) 13.18 - 12.71 (13.01) 10.26 - 10.76 (10.51) 10.13 - 10.67 (10.40) FD: Hind femur, depth 2.54 - 2.60 ( 2.56) 2.70- 2.69 ( 2.73) 2.26- 2.47 ( 2.37) 2.09- 2.24 ( 2.14) L: Rostrum-ovip. 17.11 - 19.36 (18.41) 22.18-21.76 (21.91) 17.28 - 17.43 (17.36) 18.06 - 16.89 (16.37) P: Pronotum (midline 3.00- 3.07 ( 3.04) 3.84- 3.72 ( 3.77) 2.86- 2.86 ( 2.86) 2.84- 3.05 ( 2.92) Interocular space 0.28- 0.31 ( 0.29) 0.33- 0.32 ( 0.32) 0.24- 0.25 ( 0.25) 0.26 - 0.33 ( 0.31) EE: (eye-eye) 3.44- 3.54 ( 3.48) 4.09- 3.94 ( 4.01) 3.36- 3.52 (3.44) 3.33- 3.54 ( 3.43) Fastigium 0.46- 0.56 ( 0.51) 0.64- 0.55 ( 0.59) 0.35- 0.38 ( 0.37) 0.51 - 0.59 ( 0.54) Antennal pedicel (width) 0.37- 0.41 ( 0.38) 0.45- 0.44 ( 0.44) 0.36- 0.37 ( 0.37) 0.34 - 0.38 ( 0.36) Antenna 9.37 - 10.79 (10.17) 11.40 - 10.00 (10.50) no data 7.45- 8.25 ( 7.98) Tl: hind tarsus | 1.41- 1.54 ( 1.49) 1.90 - 1.74 ( 1.83) 1.20- 1.47 ( 1.34) 1.44- 1.55 ( 1.50) T2: hind tarsus 2 0.99- 1.11 ( 1.03) 1.69- 1.49 ( 1.57) 0.95 - 1.03 ( 0.99) 0.28- 1.21 ( 0.96) T3: hind tarsus 3 2.14- 2.41 ( 2.24) 2.85- 2.68 ( 2.79) 2.10- 2.13 ( 2.12) 1.97- 2.16 ( 2.10) Ratios FD/F 0.22- 0.24 ( 0.23) 0.21- 0.20 ( 0.21) 0.22- 0.23 ( 0.22) 0.20- 0.21 ( 0.21) F/P 3.52- 3.85 ( 3.65) 3.53- 3.38 ( 3.45) 3.59- 3.76 ( 3.67) 3,323 722035577) L/P 5.70- 6.31 ( 6.05) 5.85- 5.78 ( 5.81) 6.04 - 6.09 ( 6.07) 0.00- 5.89 ( 4.27) Interoc./P 0.09- 0.10 ( 0.10) 0.09- 0.08 ( 0.09) 0.08- 0.09 ( 0.09) 0.09- 0.11 ( 0.11) Interocular/pedicel 0.71- 0.84 ( 0.77) 0.75- 0.71 ( 0.73) 0.65 - 0.69 ( 0.67) 0.76- 0.94 ( 0.87) Fast/EE 0.13 - 0.16 ( 0.15) 0.16- 0.14 ( 0.15) 0.10- 0.11 ( 0.11) 0.11- 0.15 ( 0.14) (T1+T2+T3)/F 0.43- 0.43 ( 0.43) 0.49- 0.46 ( 0.48) 0.21- 0.24 ( 0.22) 0.38- 0.47 ( 0.44) T2/ (T1+T2+T3) 0.21- 0.22 ( 0.22) 0.26- 0.25 ( 0.25) 0.42- 0.42 ( 0.42) 0.07- 0.25 ( 0.21) Ant/Pronotum 3.12- 3.51 ( 3.34) 3.06- 2.63 ( 2.78) no data 2:60 - 2.84 ( 2.73) EE/F 0.29- 0.33 ( 0.31) 0.03 - 0.02 ( 0.02) 0.33- 0.33 ( 0.33) 0.31- 0.35 ( 0.33) The original description was based on a 60-year old museum series. Fresh new specimens from a nearby region (some 20 km higher up the Rio Teribé) show that the types are badly discoloured. The description of the coloration must be amended as below. MALE. General colour, dark green. Antennae, black; 21 flagellar segments. Eyes, black (brown when dried). Frons and genae blueish-white, with a black band running from eyes through antennal sockets and including the rostrum; lower edge of frons and genae edged black. Mouthparts black, palps white. Postocular stripe and vertex, black. Fastigium and inverted V marking on vertex, blue grey. Pronotum green, with black postocular stripe produced over lateral lobes to posterior border, Dr. glyphocerca n. sp. 9.45 - 9.97 2.10- 2.24 14.01 - 15.31 2.55- 2.89 0.20 - 0.26 52328339 0.42- 0.53 0.39 - 0.47 9.09 - 10.80 1.31- 1.39 0.86 - 0.98 1.89- 2.05 0.22 - 0.23 3.34 - 3.82 5.17- 5.74 0.07 - 0.10 0.45 - 0.57 0.13- 0.16 0.43- 0.46 21 - 0.23 3.35- 4.24 0.34- 0.35 9.93 - 10.65 2.38- 2.56 17.22 - 18.27 3.16- 3.30 0.31- 0.34 3.42- 3.61 0.50- 0.60 0.39- 0.41 7.79 - 8.72 1.32- 1.50 0.97- 1.06 1.84- 2.07 0.23 - 0.25 3.01 - 3.29 5.22- 5.66 0.09- 0.10 0.76- 0.87 0.14- 0.18 0.41- 0.44 0.23- 0.24 2.36- 2.67 0.33- 0.34 Drymophilacris spp. ISSO IO BIN QU © Il un NSH PO OWIOWUINSZ © © à à B à à à D IEW CEICOLONDOS® NEW PROCTOLABINE GRASSHOPPERS Dr. melanopsis n. sp. 10.42 - 11.00 2.30- 2.32 16.44 - 18.30 2.83- 3.02 0.20- 0.22 3.39- 3.58 0.39 - 0.45 0.41- 0.43 12.91 - 13.42 1.41- 1.56 IS 9122 2.21- 2.42 0.21- 0.22 3.45- 3.82 5.55- 6.47 0.07 - 0.08 0.48 - 0.51 0.11- 0.13 0.44- 0.47 0.23- 0.25 4.27- 4.70 0.32- 0.33 12.23 - 12.66 2.68- 2.77 20.66 - 21.07 3.47 - 3.63 0.29- 0.31 3.74- 3.79 0.532, 0:57. 0.43 - 0.44 9.92 - 11.60 1:67:= 91.73 1.27- 1.44 22 TA 0.21- 0.23 3.49 - 3.52 5.69- 6.07 0.08 - 0.09 0.66- 0.72 0.14- 0.15 0.46- 0.47 0.22- 0.25 2.86- 3.20 0.30- 0.31 MEAN N=3 (10.74) ( 2.31) (17.27) ( 2.94) ( 0.21) ( 3.48) ( 0.42) Dr. nigrescens n. sp. 9.92 - 10.41 1.98- 2.16 15.75 - 16.30 2.532.753 0.17- 0.23 32320337 0.33- 0.49 0.38- 0.47 12.04 - 13.39 1.22- 1.41 1.07- 1.25 2.06- 2.18 0.20- 0.21 3.66- 3.93 5.75- 6.40 0.06 - 0.09 0.40 - 0.56 0.10- 0.15 0.44 - 0.46 0.24- 0.26 4.39- 5.29 0.32- 0.34 10.96 - 11.63 2.29- 2.41 17.57 - 20.38 2.83- 3.30 0.29- 0.36 3.55 - 3.68 0.45- 0.52 0.37 - 0.40 9.59 - 10.46 1.36- 1.54 1.18- 1.31 2.20- 2.39 0.20 - 0.21 3.48 - 3.87 5.61- 6.70 0.10- 0.11 0.69 - 0.84 0.12- 0.14 0.42- 0.45 0.24- 0.26 3.15- 3.47 0.31- 0.33 MEAN N=5 (10.09) ( 2.10) (16.01) ( 2.67) ( 0.19) ( 3.29) ( 0.39) ( 0.42) (12.47) Dr. veraguensis n. sp. 9.11- 9.69 1.90- 2.06 13.80 - 14.84 2.50- 2.64 0.14- 0.23 3.09 - 3.21 0.27- 0.29 0.37 - 0.37 8.28- 8.28 1 1320123 0.96 - 1.07 1.89- 1.92 0.21- 0.21 3.64- 3.67 5.52- 5.62 0.05 - 0.09 0.38 - 0.62 0.09 - 0.09 0.43 - 0.44 0.24 - 0.26 3.14 - 3.14 0.33 - 0.34 MEAN N=2 ( 9.40) ( 1.98) (14.32) (257) ( 0.19) ( 3.15) ( 0.28) ( 0.37) ( 8.28) ( 1.18) ( 1.02) (91) ( 0.21) ( 3.66) (SE) ( 0.07) ( 0.50) ( 0.09) ( 0.44) ( 0.25) ( 3.14) ( 0.34) 809 where the black band is punctuated by a single green spot; sulci black. Pterothorax and abdomen, green. Preterminal abdominal segments pale yellow ventrolaterally, genital region black. Spots on subgenital plate, supraanal plate and outer face of cerci, blue-grey. Fore and middle legs distal to midfemur, blue; proximal femur red, coxa yellow. Hind femora dark green, hind knee and hind tibia darker, suffused with blueish; hind feet brownish. FEMALE. Generally similar to male. Metathoracic and abdominal segments 1-7 with an interrupted lateral yellow stripe, flanked dorsally and ventrally with diffuse black. Black stripes on pronotum bordered dorsally by a thin yellow stripe. Supranal plate yellow green. Hind femora paler green proximally. 810 C. H. F. ROWELL 8. Drymophilacris panamae Descamps, 1976: reinstated combination Descamps, 1976: 82. Drymacris panamae Descamps & Rowell, 1978: 357, 361. Holotype male: PANAMA: Prov. Chiriqui: El Volcan: Cerro Punta, 5600 feet, Feb. 1937 (D.W. Bishop) (ANSP). Type examined. Note that Descamps omitted the true locality, Cerro Punta, from his original description; there is both a village and a mountain of that name, and they lie some 10 km NE of the town of El Volcan. New records: PANAMA: Prov. Chiriqui: Volcan Bart, eastern slope, 1800 m, 26 Sept. 1999 (Rowell CHF, Bentos-Pereira A), 1 male, 1 allotype female, specimen nos. 99248 & 99246 (ANSP); 1 male, 1 female, specimen nos. 99247 & 99251 (UP); 4 males, 3 females, specimen nos. 99245, 99249-50, 99252-55 (RC). COSTA RICA: Prov. Puntarenas: Cerro Cruces, 6 km. S. of San Vito de Jaba, 5000', 16 Mar 1969 (Rentz DC), specimen no. 69001 (RC). (This locality appears to be Cruces in the Cerro Zapote, 1476 m, LS 300600_575100, now completely deforested). The holotype male is discoloured (possibly pinned from alcohol?) and the female has not been described previously. REDESCRIPTION Small for genus, and with relatively short antennae. Measurements: see Table 2. MALE (Fig. 4). Elytra squamiform, reaching posterior edge of metathoracic notum. Furcula present, in form of two minute melanized projections. Supra-anal plate (Fig. 4C) 1.48X as long as broad, with a medial proximal depression; having a somewhat up-rolled, thickened margin, and ending in a bluntly pointed cylindrical process. Apical lip of subgenital rim rather long. Cercus (Fig. 4B-D) longer than supraanal plate, curving gently inwards, tapering smoothly to a rounded tip; lower edge bearing a ventrally and inwardly directed lobe 3/4 of the way to the tip. Internal genitalia (Fig. 5). Phallic complex of same form as that of other members of the genus, but elongated along anterior-posterior axis until relatively nearly twice as long as that of e.g. D. bimaculata; epiphallus correspondingly unusually long for genus, only 0.62X as broad as long. Dorsal aedeagal valve straight, slightly shorter than ventral valves, symmetrically rounded at tip. Ventral valves project beyond posterior edge of capsule, ending in single recurved hooks, barbed on the inner surface (Fig. 5D). Coloration. Antennae, black. Eyes, black (brown when dried). Mandibles blackish distally. Postocular stripe, marks on genae and V-mark on vertex dark green. Remainder of head metallic pale green with golden sheen (drying to dull light green). Nota of pterothorax and disc of pronotum olive green, pronotal lobes shading to olive brown. Metallic green marks dorsolaterally at anterior rim of pronotum and anterior to each pronotal sulcus (vanish in dried specimens). Elytra olive green. Front and middle legs pale reddish brown proximally shading to olive green at knee. Tibia and tarsi olive green. Tarsal spines and tarsal claws tipped black, pulvillus blackish. Basal 3/4 of hind femur, reddish brown, distally green, upper part of knee tinged brown. Hind tibia blue-green, shading to reddish brown distally, spurs brown; tarsal spines (6/7) blue-green, tipped brown. Hind tarsi pink, claws brown. NEW PROCTOLABINE GRASSHOPPERS 811 vis D Fic. 5 Drymophilacris panamae Descamps, male. Internal genitalia. A, epiphallus. B-D, phallic complex in lateral, dorsal and ventral view. The ventral aedeagal valves are here coloured black for clarity, they are not heavily melanized in life. Arrow in D shows the "barb" on the appendage of the ventral valve. Abbreviations: ape, anterior process of endophallus; dav, dorsal aedeagal valve; rc, rami of cingulum and associated membranes; vas, ventral aedeagal sclerite: vis, ventrolateral sclerite. 812 C. H. F. ROWELL First 8 abdominal segments olive green, paired darker green blotches dorso- laterally on each tergum. Segment 8 with metallic iridescence in life, lost at death. Segments 9-10 reddish brown, furcula and rim of 10th. tergum blackish. Cerci pale blue-green basally, black distally. Supraanal plate pale yellow-brown proximally, dark brown distally; it lacks the striking colours seen in other species of the genus. Subgenital plate a conspicuous light blue in life, medium dark grey-blue when dried, undivided by black medially; rim and pallium blackish. Underside purplish grey, darker medially. Sternum of 9th abdominal segment blackish. FEMALE (Figs 6, 7). Elytron squamoid, extending to posterior margin of meta- notum. Cerci (Figs. 7A-C) short, but abruptly narrowed to thin points. Subgenital plate (Fig. 7B) flat posteriorly, somewhat concave medially; proximally two paired swellings, easily visible in profile in side view of the abdomen (Fig. 7C); distal margin subtriangular, ending in an thick straight "egg guide" nearly equal in length to the ventral ovipositor valves; outer lobes densely haired. Internal genitalia. Bursa copulatrix (Fig. 7D) long, thin, thick-walled, with two prominent dorsal thickenings at the base; the lumen is mostly as narrow as the duct and is only expanded over a short length near the entrance. Spermatheca (Fig. 7E): terminal ampulla rather long and thin, preapical diverticulum with a long narrow initial chamber, and a convoluted appendage bearing three small angular accessory diverticula. Coloration. All adult and late larval females seen were of the bronze morph frequently found in this genus group; if there is a form coloured like the males, it is not common. Ste setences MAIER N 5 mm if x FIG. 6 Drymophilacris panamae Descamps, female. Habitus. NEW PROCTOLABINE GRASSHOPPERS 813 FIG. 7 Drymophilacris panamae Descamps, female. Genitalia. A-C, external genitalia in dorsal, ventral and lateral views. D, ventral ovipositor valves and bursa copulatrix. E, spermatheca. 814 C. H. F. ROWELL Antennae: scape and pedical yellow, flagellar segments 1-10 red, 11-20 black, 21 white. Eyes light purple-grey, with black pseudopupil (brown when dried). Labrum and clypeus, yellow brown. Palps greenish, blackish at base. Frons and anterior portion of genae, pinkish bronze. Posterior genae iridescent green (dries pale yellow brown) with a few black spots. Postocular stripe black. Vertex and fastigium brown, V-shaped mark on vertex green. Dorsal surface of thorax, pinkish bronze. Postocular stripe continued onto upper lobe of pronotum as an irregular black stripe, bordered ventrally with iridescent green. Metathoracic epimeron and episternum metallic blue-green dorsally, shading to olive green ventrally. Elytron blackish brown. Front and middle legs green; tibiae blackish distally, tarsi and pulvilli blackish brown. Hind femur and tibia green, upper part of knee tinged brown. Tarsi pink. Spines and spurs minutely tipped black. Dorsal surface of abdomen, including supranal plate, pinkish bronze; all segments with grayish posterior border. Conspicuous black spot on the side of each notum of abdominal segments 2-4, each containing an ill-defined yellow blotch posteriorly. Abdomen widest at level of segment 7, thereafter narrowing abruptly to genital region. Tips of cerci, dorsal surface of superior ovipositor valves and tips of ventral valves blackish brown. Underside of abdomen and thorax dark green laterally, in each segment shading to dark blue medially and posteriorly. Dried specimens lose almost all the colour described above and become a uniform pale rufous brown with green legs and some black markings. The antennae retain their colours. Natural History. The type locality, Cerro Punta, is now a centre of intensive vegetable horticulture with very heavy insecticide application, little natural vegetation and few living insects. It is reassuring that the species occurs in suitable localities on Volcan Bart, which is a National Park. It lives there on Solanum bushes at the forest edge, sharing them with a very large population of Ampelophilus olivaceus (Giglio- Tos), another lithoscirtan. In September, when we made our collection, the majority of individuals of D. panamae were still larvae; adults, especially females, were scarce, but the species is not uncommon in this locality. Comment. At the time of our transfer of this species to Drymacris only the holotype male was available. The rather large squamiform elytra suggest this attri- bution, and the male cercus has a weak resemblence to that of Drymacris nebulicola. The diagnostic structures of the female subgenital plate and ovipositor, however, leave no doubt that the real genus is Drymophilacris. The specimen from Puntarenas Province, Costa Rica, is a fifth instar female larva, and precise determination is not easy. Its antennal coloration indicates either Drymacris nebulicola or Drymophilacris panamae, which are also the geographically closest members of the genus group. The shorter antennae, small wing rudiment, long thin pointed cerci and flattened upper ovipositor valves confirm the latter identi- fication. This is the first record of the species from Costa Rica; the locality lies some 50 km W of Volcan Bart. NEW PROCTOLABINE GRASSHOPPERS 815 9. Drymophilacris glyphocerca sp. n. Holotype male: COSTA RICA: Prov. Cartago: P.N. Tapanti: Rancho Negro, 1735 m, LN 186300_560000, 7. Oct 1999 (Rowell CHF), specimen no. 99382. Allotype female: same data, specimen no. 99380 (both ANSP). Paratypes: As holotype, but specimen nos. 99364, 99372, 99378 ,99379, 99383, 99384, 99387, 99388, 99389, 99395 and 99396 (all ANSP). Same, but specimen no. 99367 (f); COSTA RICA: Prov. Cartago: P.N. Tapanti: 0-1 km past entrance to reserve, LN 194000_559800, 8.Oct 1999 (Rowell CHF), specimen nos. 99403 & 99404 (2 m) (all MHNP). As holotype, but specimen nos. 99365, 99366, 99368, 99370, 99371, 99375, 99381 & 99416 (RC). COSTA RICA: Cartago: P.N. Tapanti: Sector La Represa: Rancho Negro, LN 186300_560000, 15. Aug 1997 (Delgado R), specimen nos. CRIO02 592428 & 592429 (INBio). COSTA RICA: Cartago: P.N. Tapanti: R. Dos Amigos, 1480 m, LN 189500_560200, 15. Mar 1994 (Mora G), specimen no. CRIOO1 964083 (larva) (INBio). Etymology: (Gr. glyphis, penknife, and kerkos, tail), referring to the form of the male cercus. MALE (Fig. 8). Elytra rounded, squamiform, reaching to first 1/3 of meta- thoracic notum. Furcula barely perceptible. Supra-anal plate 1.33X as long as broad, in form like that of D. panamae. Lip of rim of subgenital plate shorter than in D. panamae. Sheathed aedeagus very large, projects above and beyond cerci. Cerci (Figs 8B-D) very similar to those of D. panamae, ventral lobe somewhat smaller and more angular at extremity. Antennal flagellum with 20 segments. Internal genitalia (Fig. 9). Rather elongate phallic complex, similar to but less extreme than that of D. panamae. Epiphallus (Fig. 9A) approximately square in dorsal view, about equally long as broad. Superior valve of aedeagus (Fig. 9B, D) short, straight and very slender in both lateral and ventral view. Ventral valves of same form as in D. panamae, but project further and are less obviously barbed (Fig. 9C, D). Coloration. Antennae black, proximally annulated in grey. Eyes black (brown when dried). Frons, clypeus, entire frontal ridge, genae, blue green. Labrum and mandibles, black distally, brownish proximally. Palps dark at base, tipped green. Fastigium green-gold; vertex greenish blue. Postocular stripe and some pitting on adjacent parts of gena, black. Pronotum blue-green, its margins, sulci and continuations of post-ocular stripe, black. Meso- and metathoracic epimera and episterna, blue-green, sutures black. Elytron brown, central area green. Thoracic and abdominal nota blue-green with paired black chevrons, each bearing a gold mark. The black area diminishes in the later abdominal segments. Underside blueish green, medial area black. Legs pale olive green, pulvilli and claws black. All feet red-brown. Hind knees, red brown. Hind tibiae green proximally, shading to red-brown distally; spines red-brown. Genital area, black. Supra-anal plate metallic green-gold. Cerci black at tip, yellow-grey on outer face proximally. Subgenital plate gold-green, undivided in adult, medially divided into two spots in late larva. In dried specimens the blue, gold and some black elements fade, leaving predominantly shades of green. The black of the antennae, mouthparts and genital area is however unaffected. 816 C. H. F. ROWELL Fic. 8 Drymophilacris glyphocerca sp. n., male. A, habitus. B-D, external genitalia in lateral, dorsal and ventral views. NEW PROCTOLABINE GRASSHOPPERS 817 Fic. 9 Drymophilacris glyphocerca sp. n. male. Internal genitalia. A, epiphallus, dorsal view. B, phallic complex, lateral view. C, D, as B, but dorsal and ventral views respectively. Arrow in D shows the "barb" on the appendage of the ventral valve - compare with Fig. 5D. 818 C. H. F. ROWELL FEMALE. Elytra rounded, squamiform, reaching to middle of metathoracic notum. Cerci (Fig. 10A-C) abruptly narrowed to long thin points. Subgenital plate (Fig. 10B) flat or slightly concave posteriorly; proximally two paired swellings, which are produced anteriorly onto the next abdominal sternum; distal margin subtriangular, ending in an "egg guide" equal to 2/3 the length of the ventral ovipositor valves and markedly curved upwards at the tip; outer lobes densely haired. Antennal flagellum with 19-20 segments, median 20. Internal genitalia. Bursa copulatrix (Fig. 10D) as in D. panamae. Spermatheca (Fig. 10E): preapical diverticulum with a short initial chamber and an appendage with 2 rounded accessory diverticula. Coloration. Highly polymorphic in colour, with at least 4 variants - green like male, though with less black on pronotum; bronze; dull brown; and almost black. The first two forms can additionally have either red or green legs. Antennae always red proximally, blue-black distally, with white tip separated from the blue area by a thin red band. Relationships. The species seems close both ecologically and morphologically to D. panamae; both are alpine, small, have shorter antennae than usual in the genus, similar flanged male cerci, minute furculae, and barbed hooks on the tips of the ventral aedeagal valves. Natural history. Occurs commonly above 1600 m on Solanum spp. in wet montane forest in the upper drainage of the Rio Grande de Orosi and its tributary, the Rio Macho, both draining the north-eastern Talamanca mountains. At high altitude (>2000 m) it is sympatric with Drymacris nebulicola, below 1600 m it is replaced by Lithoscirtus daedalus. Of the 28 specimens of the type series, three were parasitised by mermithid nematodes. 10. Drymophilacris nigrescens sp. n. Holotype male: PANAMA: Prov. Bocas del Toro: 1-3 km past watershed on road Fortuna-Chiriqui Grande, 1035-850 m, 24. Sept. 1997 (Rowell CHF Bentos A), specimen no. 97488. Allotype female: same data but 23. Sept 1997, specimen no. 97477 (both ANSP). Para- types: same data as holotype, but dates between 18. and 27. Sept. 1997: specimen nos. 97452, 97453, 97462, 97480, 97486, 97487 & 97569 (all ANSP). As above, but specimen nos. 97472 & 97473 (2 males); as above, but 18.09.99, specimen no. 99126 (female) (all MHNP). As above, but specimen nos. 97451 (female) & 97461 (male) (UP). As above, but specimen no. 97509 (female) (RC). PANAMA: Prov. Bocas del Toro: Quebrada Felix, 2 km NW of summit of rd. to Chiriqui Grande, 900 m, 26. Sept. 1997 (Rowell CHF Bentos A), specimen nos. 97535 & 97536 (2 males) (ANSP). Etymology: Latin nigrescens, dusky, blackish. MALE (Fig. 11). Elytron extending over 50% to 90% of mesonotum, not reaching the posterior margin of the tergum. Furcula small but distinct. Tip of cercus with rounded obtuse upper and angular acute ventral corners. Supra-anal plate 1.18X as long as broad, tapering to a rounded tip, largely without the appendage seen in panamae and glyphocerca. Subgenital plate with a short medial lip. NEW PROCTOLABINE GRASSHOPPERS 819 KE. \ \\ E MURGIA Fic. 10 Drymophilacris glyphocerca sp. n., female. Genitalia. A-C, external genitalia in dorsal, ventral and lateral views. D, ventral ovipositor valves and bursa copulatrix. E, spermatheca. 820 C. H. F. ROWELL 5mm Fic. 11 Drymophilacris nigrescens sp. n. Male. A, habitus. B-D, external genitalia in lateral, dorsal and ventral views. NEW PROCTOLABINE GRASSHOPPERS 82] Internal genitalia (Fig. 12). Upper aedeagal valve slightly shorter than lower valves, laterally compressed, rather pointed, straight. Terminal appendages of lower valves a simple straight hook, unbarbed. Coloration: Antennae black. Eyes black (red-brown when dried). Lateral ocelli, yellow. Palps black basally, green at tip. Mandibles, labrum, clypeus, ventral rim of genae, black. Antennal socket, black. Postocular stripe and most of genae, black. Frons and anterior central part of genae, grey-green or grey-blue, variably suffused with black. Frontal ridge in upper part always black, below medial ocellus variably so. Fastigium and vertex, dull yellow-green or blue-grey; V-marking on vertex black. Pronotum mainly black. Anterior ventral angle green, a pair of green marks dorsolaterally at posterior margin, just anterior to elytra. In some individuals a variable amount of dark green marking medially on pronotal disc. Meso- and metathorax, black laterally and ventrally, dark green dorsally and medially, variously suffused black; a pair of poorly defined yellow dots dorsolaterally and posteriorly, in the black area of each segment. Elytra brown, traces of wings visible in some individuals. Front and middle legs yellow proximally, dark green in distal femur and from thence distally. Pulvilli and claws blackish. Hind femur pale green proximally, dark green or blue-green distally, ventral surface light blue. Ventral lobes of knee light blue. Tibiae mainly blue, but green proximally and black distally; spines 6/7, blue, tipped red brown, spurs brown. Tarsi pinkish brown, claws and pulvilli blackish. Fic. 12 Drymophilacris nigrescens sp. n., male. Internal genitalia. A, phallic complex, dorsal view. B, the same, lateral view. C, epiphallus, dorsal view. 822 C. H. F. ROWELL Terga of abdominal segments 1-6 repeat pattern of meso- and metathorax; segments 7-9 are lighter. Sides of segments yellow, underside blackish, with blue posterior margins and yellow-brown anterior margins. Genital area (Figs 11B-D) mostly black. Furcula black. Tip of cercus with rounded dorsal and angular ventral corners, black, proximal part of outer face grey-blue or grey-green. Paired confluent spots of subgenital plate grey-blue. Supraanal plate with black rim and distal half, proximal central part yellow; distal half with long dense hairs. FEMALE (Fig. 13). Elytra extend to posterior margin of mesonotum. Cerci typical of genus, but slightly downturned at tip in lateral view (Fig. 13C). Subgenital plate flat or concave distally, ending in an egg guide (Fig. 13D) that equals or exceeds the length of the ventral valves, downwardly hooked at the tip; anteriorly provided with two large laterally directed melanized swellings, clearly visible as lateral projections from above or below (Figs 13A, B). Internal genitalia (Figs 13E, F) typical of the genus. Coloration. Females are dimorphic in coloration. Green form: coloration as male, except for antennae (distal 2 segments white), hind tarsi (grey) and genital area. Elytron greenish yellow, brown basally, reaching posterior edge of mesothoracic tergum. Sides of last three abdominal tergites black, forming a pair of conspicuous dark spots. Cerci yellow-green basally, black distally. Proximal half of supraanal plate green, terminal half blackish. Ovipositor valves light brown, blackish on dorsal face of superior valve. Subgenital plate brown laterally, blue grey medially; basal swellings, blackish. Bronze form: entire body except legs reddish brown, with metallic sheen in life, lost after death; distal half of antenna black with white tip, postocular stripe darker brown. Yellow spots of the green form represented by light brown flecks. Front and middle legs green, tarsi and tibiae tinged brown. Hind femur green, knee area reddish brown, underside blue. Hind tibia red-brown, ventral surface blue, blackish at distal extremity. Tarsi light brown. Claws, spurs and pulvillus dark brown. Relationships. Morphologically similar to the geographically adjacent D. rubripes, especially in the form of the female subgenital plate, but differing in the ventrally hooked egg guide. It also differs in its larger size and slightly longer fastigium, and markedly in coloration. Natural history. On Solanaceae (mostly Solanum spp., esp. S. ochraceo- ferrugineum, but also Witheringia solanacea) and sometimes on Vernonia (Aster- aceae) in clearings in montane forest, up to 1200 m (the highest available point). Not recorded below 550 m. Found to date solely on the Caribbean side of the watershed. Adults and all stages of larvae present in September. Mature females contain 2-6 eggs. Comment. The species is remarkably similar in general coloration to a large undescribed Hylopedetes sp. (Rowell in prep.) with which it is sympatric. Especially green females of D. nigrescens, lacking the characteristic markings of the male genital region, can readily be confused with this rhytidochrotine; the best field NEW PROCTOLABINE GRASSHOPPERS 22 (GS) Fic. 13 Drymophilacris nigrescens sp. n., female. Genitalia. A-C, external genitalia in dorsal, ventral and lateral views. D, ventral ovipositor valves and bursa copulatrix. E, spermatheca. 824 C. H. F. ROWELL character under these circumstances is the length of the antennae, which in Hylopedetes are much shorter. The black and green males with twin yellow patches on the posterior edge of the pronotum are strongly reminiscent of Lithoscirtus daedalus, but differ in the bright yellow supraanal plate and the blue spots of the subgenital plate. 11. Drymophilacris veraguensis sp. n. Holotype male: PANAMA: Veraguas Prov.: Santa Fé: Alto la Piedra, 900 m, 16. Sept 1999 (Rowell CHF, Bentos A), specimen no. 99096 (ANSP). Paratype male: same data, but 16. Nov 1999 (De Gracia L, Santos A, Gonzalez P), specimen no. 99520 (UP). Etymology: coming from Veraguas, the province in which the type locality is located. MALE (Fig. 14). Elytron minute, falling well short of posterior margin of the mesonotum; wing absent. Furcula quite well developed. Cerci incurving, rounded at the dorsal tip, angular at the ventral tip, similar to those of D. nigrescens. Supraanal plate 1.23X as long as wide, and tip blunter and more rounded, most similar to that of D. nigrescens. Internal genitalia (Fig. 15). Very similar to those D. nigrescens, but dorsal aedeagal valve equal in length to ventral valves, truncate at tip. Coloration. Antennae black (but flagellar segments distal to no. 16 missing in both specimens). Eyes black (brown when dried). Clypeus, labrum and mandibles black, palps yellow. Remainder of head yellow, except for black antennal sockets, black mark on upper part of frontal ridge between the two antennal sockets, black postocular stripe, small black fleck in centre of fastigium, and inverted black V on vertex. Pronotum yellow, with one medial and two lateral black stripes continuing the markings of the vertex and the post-ocular stripe posteriorly; these stripes continue as far as the first abdominal segment. Pronotal epimeron black. Elytron brown. Meso- and metathoracic epimera and episterna yellow with longitudinal black marks. Fore and mid legs, yellow. Hind femora green, upper lobe of knee tinged red-brown. Hind tibia green; 6 external and 7 internal tibial spines, tipped red-brown, tarsal spurs tipped black. Tarsi yellow, claws tipped black. Abdominal segments greenish-yellow. Dorsa of segments 6-8 bright yellow medially. Genital region (see Fig. 14B-D) mostly black; central area of suprananal plate, spots on subgenital plate, bright yellow. Cerci black, with a grey-green region on the central part of the outer surface of the shaft. FEMALE unknown. Relationships. As far as can be judged from the male alone, this species is most closely related to nigrescens and rubripes, which are also the closest geogra- phically. Natural history. Found on Solanum plants at forest edges. NEW PROCTOLABINE GRASSHOPPERS 825 ec LNG 1mm Fic. 14 Drymophilacris veraguensis sp. n., male. A, habitus. B-D, external genitalia in lateral, dorsal and ventral views. 826 C. H. F. ROWELL Fic. 15 Drymophilacris veraguensis sp. n., male. Internal genitalia. A, phallic complex, dorsal view. B, the same, lateral view. C, epiphallus, dorsal view. NEW PROCTOLABINE GRASSHOPPERS 827 12. Drymophilacris melanopsis sp. n. Holotype male: PANAMA: Prov. Coclé: Cerro Copé: 830 m, 20 Sept 1997 (Rowell CHF, Bentos A), specimen no. 97400 (ANSP). Paratypes: as above, but 13-15 Sept 1999: one male, specimen no. 99084, one female, specimen no. 99076 (ANSP); one male, specimen no. 99094, one female, specimen no. 99085 (RC). Etymology: melanopsis Gr., black face, noun in apposition. MALE (Fig. 16). Elytron rather large for the genus, reaching clearly into the metathoracic notum. Furcula well developed, with widely separated points directed obliquely upwards. Supraanal plate almost glabrous anteriorly, entirely yellow with a thin black/red-brown rim, 1.37X as long as broad, with a prominent appendage terminally; cerci somewhat sigmoid and narrowly waisted behind tip in lateral view, tip broad and spatulate, dorsal corner rounded, ventral corner more acute. Internal genitalia (Fig. 17). Epiphallus (Fig. 17C) longer than broad, deeply notched on anterior edge, and wider anteriorly than posteriorly, unlike other species of the genus; oval sclerites small. Phallic complex rather short and wide, relative to other species, and unique in having double recurved hooks at the tip of the ventral aedeagal valves (Fig. 17 A, B, D). Coloration. Antennae black, terminal 2 segments white (uniquely for males of the genus); 21 flagellar segments. Eyes black (red-brown when dried). Palps green. Mandibles, labrum, clypeus, frons, antennal sockets, postocular stripe, lower edge of genae, glossy black. Upper part of genae, lemon yellow. Fastigium and vertex, lemon yellow; inverted V-marking on vertex, black. Pronotum green dorsally, with darker dorsolateral stripes continuing line of post-ocular stripes. Ventral half of lateral lobes lemon-yellow, continuing line of genae. Two short yellow stripes dorsolaterally posterior to hind sulcus, continuing the line of the elytra. Pronotal epimeron, green. Meso- and metasternal interspace, blackish. Elytron, yellow. Fore and middle legs green, pulvilli black. Hind femur green, knee reddish brown. Hind tarsus green, blackish-brown distally, spurs and tibial spines (6/7) blackish brown. Tarsi blackish brown, pale olive dorsally, claws and pulvilli tipped black. Abdominal segments green dorsally, blackish brown ventrally. A pair of longitudinal yellow lines run dorsolaterally from the metathoracic notum to the genital region. Genital region (see Fig. 16B-D). Tergum 9 black, tergum 10 (with furcula) brown anteriorly, black posteriorly. Supraanal plate entirely yellow with a thin black/red-brown rim. Cerci green proximally, black distally. Paired spots of subgenital plate lemon yellow. FEMALE (Fig. 18). Elytron extends on to metathoracic notum. Cerci typical of genus. Subgenital plate (Fig. 18B) flat, with low, laterally spreading, paired swellings with rugose cuticle anteriorly, triangular posteriorly, terminating in a straight, pointed, upward slanting egg guide (Fig. 18F), 0.7X as long as the ventral ovipositor valves. 828 C. H. F. ROWELL Fic. 16 Drymophilacris melanopsis sp. n., male. A, habitus. B-D, external genitalia in lateral, dorsal and ventral views. NEW PROCTOLABINE GRASSHOPPERS 829 Fic. 17 Drymophilacris melanopsis sp. n., male. Internal genitalia. A, phallic complex, lateral view. B, the same, dorsal view. C, epiphallus, dorsal view. D, tip of aedeagus, ventral view. 830 C. H. F. ROWELL Internal genitalia. Bursa copulatrix long and wide (Fig. 18D). Spermatheca (Fig. 18E) with a long convoluted appendage to the lateral diverticulum, bearing 3 or 4 accessory diverticula. Coloration: as in male, except white tip of antenna includes the 3 terminal segments, and except for genital region. Supraanal plate, mottled olive green, tip tinged brown. Cerci olive brown, darker at tips. Ovipositor valves brown, dorsal valves black distally. Subgenital plate predominantly blackish in colour. Relationships. A morphologically distinctive species, which does not seem to be closely related to any other member of the genus, and the second largest described to date. Natural history. The insect is found on species of Solanum and Witheringia along paths and in treefall clearings in montane forest, on both sides of the Pacific/ Caribbean watershed. One male (99084) contained a mermithid nematode and unde- veloped testes. The remaining specimens were sexually mature, the females each containing 6 eggs. Drymacris Descamps & Rowell, 1978 Descamps & Rowell, 1978: 358, 361. Type species: Lithoscirtus nebulicola Rehn, 1929: 26. Amédégnato & Poulain, 1987: 400. The genus originally contained three species: nebulicola (Rehn, 1929), pana- mae Descamps, 1976, then known only from the male holotype and here transferred to Drymophilacris, and ovatipennis (Rehn, 1905), then known only from the female holotype and transferred to Paratela by Descamps & Rowell (1984) following dis- covery of the male. The genus is now monospecific. REDESCRIPTION Medium-small grasshoppers (>9.9, <20 mm in length); body length 15 (males) - 19 (females) mm. Pronotum relatively longer than in Lithoscirtus and Drymo- philacris. Hind femur 3.2 (females) - 3.5X (males) as long as pronotum, rather slender, 0.19 - 0.20X as deep as long. Hind foot longer than in the other genera, 0.48X (females), 0.50X (males) as long as femur, second tarsal joint makes up 0.27 of foot. Interocular space narrow, 0.36X (males), 0.65X (females) as wide as antennal pedicel. Antennae 4.0X (males), 3.1X (females) as long as pronotum. Fastigium slightly longer than in the other genera. Elytra much larger than in the other two genera, cycloid and extending beyond the posterior margin of the metanotum. Male cerci abruptly notched on the ventral margin towards the tip. Internal genitalia very similar to those of Drymophilacris, but rather shorter and wider than usual in that genus; epiphallus 1.6X as wide as long. Female cerci (Fig. 19A, C) short, bluntly rounded at the tip, unlike the other two genera. Upper ovipositor valves (Fig. 19A, C) relatively unmodified, less flattened dorsoventrally than in the other genera, slightly excavated dorsally, with a raised, minutely toothed, internal edge. Subgenital plate (Fig. 19B, C) flat, neither NEW PROCTOLABINE GRASSHOPPERS 831 SS È Ly u”, SH ‘Me Fic. 18 Drymophilacris melanopsis sp. n., female. Genitalia. A-C, external genitalia in dorsal, ventral and lateral views. D, ventral ovipositor valves and bursa copulatrix. E, spermatheca: EF, extremity of subgenital plate ("egg-guide") in lateral view. 832 C. H. F. ROWELL Fic. 19 Drymacris nebulicola (Rehn, 1929), female. A-C, external genitalia in dorsal, ventral and lateral views. D, bursa copulatrix. E, spermatheca. NEW PROCTOLABINE GRASSHOPPERS 833 domed (Lithoscirtus) nor provided with paired anterior swellings (Drymophilacris); terminal process ("egg-guide") very short. Terminal ampulla of spermatheca (Fig. 19E) rather long and narrow relative to the other two genera; appendage of lateral diverticulum with several small diverticula. 13. Drymacris nebulicola (Rehn, 1929) Lithoscirtus nebulicola Rehn, 1929: 26. Transferred to Drymophilacris, Descamps, 1976: 81. Transferred to Drymacris, Descamps & Rowell, 1978: 361. Holotype male and allotype female: COSTA RICA: La Palma, between Volcan Barba & Volcan Irazu, 4960-5100 feet, 1 Sept. 1923 (J.F. Tristan & J.A.G. Rehn) (ANSP). New records: COSTA RICA: Prov. Cartago: Reserva Forestal Rio Macho, Alto Roble, 2200 m, LN 190000_552300, 12 Oct. 1999 (Marshall S), specimen no. 99471 (RC). Tres Rios: Cerro la Carpintera, 1700-1800 m, LN 207600_538600 21 Sept 1993 (Rowell CHF), specimen nos. 93310 & 93311 (RC). Below pass between Volcan Irazü and Volcan Turrialba, S. side, 1900-2000m, LN 217000_562500, 6 July 1980 (Rowell CHF, Rowell-Rahier M, Hyde C). Prov. Heredia: 5 km. S of Los Cartagos (crossing of Rta 9 & R. Tabor), 1750 m, LN 232730_518480, 8 Sept 1993 (Rowell CHF), specimen no. 93203 (RC). San Rafael de Vara Blanca, 1800-2000 m, LN 239800_524200, 13 April 1986 (Rowell CHF, Braker HE), specimen nos. 86019-86022 (RC). Prov. Puntarenas: Potrero Grande: Tres Colinas, 1945-2150 m, LS 342100_ 565600, 12 Sept 1995 (Rowell CHF, Meier V), specimen nos. 95540-95542 (RC). Monteverde: Sendero Nuboso, 1480-1520 m, LN 253500_449400, 14 July 1980 (Rowell CHF, Rowell-Rahier M, Hyde C, Braker HE). Prov. S. José: Between Cascajal and Tierras Morenas, 1600-1700 m, LN 224000_ 543400, 18 Sept 1993 (Rowell CHF). S. Gerardo de Dota, 2000-2500 m, LS 387400_482700, 24 Feb 1992 (Curso Tachinidae y Syrphidae), specimen no. CRIOOO 407614 (INBio). Fig. 19 shows the previously unillustrated female subgenital plate (19B, C) and internal genitalia (Figs 19D, E). Dimensions, Table 1. Distribution. The species is now known to be widely distributed in the higher montane forest zone of Costa Rica on both Atlantic and Pacific watersheds. Descamps & Rowell (1978) noted occurences in Chompipe (near the type locality in the Cordillera Central) and in Monteverde (Cordillera del Norte). Further new records see above. ACKNOWLEDGEMENTS I thank A. Solis (INBio, Costa Rica), Prof. D. Quintero (Universidad de Panama), D. Azuma (Academy of Natural Sciences, Philadelphia, USA) and Dr. M. O'Brien (University of Michigan, USA) for access to their collections and the loan of specimens, Drs. D. Rentz (Canberra, Australia) and S. Marshall (Guelph, Canada) for the gift of specimens, the authorities of Costa Rica and Panama for permission to collect in their National Parks and elsewhere, and Profs. A. Bentos (Montevideo) and N. Elsner (Gottingen) for help and companionship in the field. 834 C. H. F. ROWELL REFERENCES AMEDEGNATO, C. 1974. Les genres d'Acridiens neotropicaux, leur classification par familles, sous-familes et tribus. Acrida 3: 193-204. AMEDEGNATO, C. 1977. Etude des Acridoidea Centre et Sud Americains (Catantopinae sensu lato): Anatomie des genitalia, classification, repartition, phylogenie. Thése, Université Pierre et Marie Curie, Paris, 385 pp. (mimeo.). AMEDEGNATO, C. & POULAIN, S. 1987. Les acridiens néotropicaux I: Proctolabinae Amazoniens (Orthoptera, Acrididae). Annales de la Société Entomologique de France (NS) 23(4): 399-434. BRUNER, L. 1908. Acrididae. Biologia Centrali Americana. Insecta, Orthoptera 2: 1-342, plates 1-4 (1900-1909). DESCAMPS, M. & ROWELL, C. H. F. 1978. Acridiens des clairieres de Costa Rica. Diagnoses, signalisations, notes biologiques, polymorphisme (Acridomorpha, Acrididae). Annales de la Societe Entomologique de France (N.S.) 14(3): 351-367. Descamps, M. 1976. La faune dendrophile néotropicale. I. Revue des Proctolabinae (Orth. Acrididae). Acrida 5: 63-167. KIRBY, W. F. 1910 A synonymic catalogue of Orthoptera. Vol. 3. Orthoptera Saltatoria. Part. 2. (Locustidae vel Acrididae). British Museum, London, 674 pp. REHN, J. A. G. 1905. A contribution to the knowledge of the Acrididae (Orthoptera) of Costa Rica. Proceedings of the Academy of Natural Sciences of Philadelphia 57: 400-454. REHN, J. A. G. 1929. Studies in Costa Rican Dermaptera and Orthoptera. II. New genera and species of Acrididae. Transactions of the American Entomological Society 55: 9-77. ROWELL, C. H. F. 1983 Drymophilacris bimaculata (pp. 714-716). In: JANZEN, D. H. (ed.). Costa Rican natural history. University of Chicago Press, Chicago and London, 816 pp. REVUE SUISSE DE ZOOLOGIE 107 (4): 835-841; décembre 2000 A new species of the genus Stumpffia (Amphibia: Anura: Microhylidae) from a small forest remnant of the central high plateau of Madagascar Denis VALLAN Universitat Bern, Zoologisches Institut, Baltzerstrasse 3, CH-3012 Bern, Switzerland. E-mail: denis.vallan@zos.unibe.ch A new species of the genus Stumpffia (Amphibia: Anura: Micro- hylidae) from a small forest remnant of the central high plateau of Madagascar. - A new frog species of the terrestrial and leaf litter dwelling genus Stumpffia is described. Stumpffia helenae sp. n. has enlarged finger tips and partially reduced fingers and toes. It is the first record of the genus from the central high plateau of Madagascar. Since this frog was discovered in small forest fragments, this record shows how important small forest remnants could be for nature conservation. Key-words: Amphibia - Anura - Microhylidae - Stumpffia - new species - Madagascar - high plateau - forest fragment - nature conservation. INTRODUCTION In Madagascar the family of Microhylidae is represented by ten endemic genera distributed in three subfamilies: Scaphiophryninae, Dyscophinae and Cophylinae (Glaw & Vences, 1994). Stumpffia, a genus containing small to very small species that often have reduced fingers and toes, belongs to the latter subfamily. At present, the genus Stumpffia contains seven species: Stumpffia psologlossa Boettger, 1881, S. tridactyla Guibé, 1975, S. pygmaea Vences & Glaw, 1991, S. tetradactyla Vences & Glaw, 1991, S. grandis Guibé, 1974, S. roseifemoralis Guibé, 1974, and S. gimmeli Glaw & Vences, 1992. In this paper a new species of Stumpffia is described. It was found in a forest remnant of the nature reserve of Ambohitantely and is the first record of this genus in the central high plateau of Madagascar. MATERIALS AND METHODS Animals were collected during a survey for ecological research. After anaes- thesia with ether specimens were fixed in 90% ethanol and stored in 70% ethanol. All measurements were taken with dial callipers to a precision of 0.1 mm. Unless otherwise stated the following parameters were measured according to Duellman (1970): snout-vent length; head width; forearm length; hand length (from the tubercle Manuscript accepted 19.06.2000 836 DENIS VALLAN at the base of finger I to the tip of the longest digit); tibia length; foot length including tarsus (from the joint between the tibiofibula and ankle to the tip of toe IV); foot length (from the base of the metatarsal tubercle to the tip of toe IV); horizontal tympanum diameter; horizontal eye diameter; distance from eye to nostril; distance from nostril to snout tip. All measurements are in millimetres. Photos were taken to document the colour of the living animals. The following abbreviations are used: NMBE = Naturhistorisches Museum Bern; SMF = Senckenberg Museum Frankfurt; ZFMK = Zoologisches Forschungs- institut und Museum Alexander Koenig. The following specimens were used for comparison: Stumpffia psologlossa (SMF 7337; holotype), S. grandis (NMBE 1034195), S. gimmeli (ZFMK 52536; paratype). For the other species data were taken from literature (Vences & Glaw, 1991: Glaw & Vences, 1994). RESULTS Stumpffia helenae sp. n. Holotype: NMBE 1044802 (Figs la, lc, 2a and 2c) Adult d from the “Reserve Spéciale” (RS) d’Ambohitantely, central high plateau of Madagascar, Ankazobe Fivondronana, Antananarivo Province, 18° 09° S, 47° 15° E, 1500 m a. s. 1., 25 March 1997, D. Vallan leg. Paratype: NMBE 1044801 (Figs Ib, Id, 2b and 2d) Adult 2, same locality and collector as the holotype, 23 March 1997. Diagnosis: Due to its general features (small size, partially reduced fingers, connected lateral metatarsals, inner metatarsal tubercle present, outer metatarsal tubercle absent, webbing between fingers and toes absent, no signs of external vocal sac are visible, terrestrial habits and slow motion moving) I include this species to the genus Stumpffia. Differentiation from other Stumpffia species: The most similar species are S. gimmeli and S. psologlossa. Both species have a similar size and can be similarly coloured. In all three species (S. gimmeli, S. psologlossa and S. helenae Sp. n.) fingers and toes are clearly recognisable but S. gimmeli, S. psologlossa differ in having less reduced fingers and toes. They also lack the enlarged fingertip. As shown in Tab. 1 S. helenae sp. n. differs from all other Stumpffia species by the combination of size, digital reduction and the enlarged fingertips. Enlarged fingertips are usual in species of several genera (Anodonthyla, Plethodontohyla, Cophyla and Platypelis) of the microhylid family but not known from species of the genus Stumpffia. | presume that Stumpffia helenae sp. n. belongs to the genus Stumpffia due its enlarged finger tips mainly because of its reduced finger and toes and the slow motion moving. In Malagasy anurans these two characteristics are only known in species of the genus Stumpffia. Description of the holotype: A 3 of 13.8 mm length. Also if the animal was not found calling, the presence of nuptial pads indicate that the holotype is an adult d. Four fingers and five toes present. Finger II is slightly and finger I is more reduced. Toe I is also reduced (Figs 2a and 2c). Tip of finger III is slightly enlarged and triangular (due to the small size of the only known holotype and paratype, further 837 A NEW SPECIES OF THE GENUS STUMPFFIA PO6T SSOUSA 9 METH PUP [661 MIO 2 SIIUIA WOI BIE ¢ “GESTS MINAZ + S6IVEOI AGINN ¢ ‘LEEL ANS 7 ‘I08++OI AGN | ou ou ou ou ou ou ou SoA dy 193u1J "€ posiejua 0/0/S 0/0/7 0/1/t 0/1/€ O/1/+ 0/1/€ [/0/ÿ 0/1/€ 0/1/t O/T/E [/T/T 1/€/0 0/0/S 0/0/t 0/1/f O/T/T $90} s.193u1J jo Jaquinu jo Jaquinu E/I 249 : C/E 249 © 1/E 249 : Ce El 249 : SIT - I unueduA : PIE - S/T wnuedwA] ' 1/1 aka" ul 249 : OUI 949 ** OY} SIYPLII UOTE] jumuedwä] -nI1.AIe [es BOI qu], 6 CI CC O81 Sl GE=OI SO orl 8 €I (wu) Yu] ; pwund 'S ç S1jpaowafıosoA 'S £ SIPUDAS 'S ç PIUSDPP.A101 'S c PADUS "S ç Popp 'S - DSSO]80J0Sd 'S ‘u ‘ds avuajay pıf/dums TOMO) TO) SOM Oc: Or. I x9S sInads ‘S207 pur s193UIy paonpai AJ21>]dU109 Jo JOQUINU ay} 01 saojaı IOQUUNU PITY) Sy) pur sao) pur sıoduny juosoid ınq pasnpai JO JaquINU ay} 0) PUO99S Sy) ‘S20] Pur sıodur) poonpol JOU JO JaquINU dy} 0) S19JO1 JOQUINU JSIIJ Sy) „S90) Jo Joqunu,, pur .SIISULJ JO Jaquinu,, UUINJOS ay} UT 'snusd ou} Jo sarsads Joyo pue ‘ds "u apuajay pıfldums usomaq SONSUAeILYD oY) Jo UOSLIVdWOD ‘| “AVI, 838 DENIS VALLAN Fic. 1 Dorsal (a) and ventral (c) view of & Stumpffia helenae sp. n. (holotype: NMBE 1044802). Dorsal (b) and ventral (d) view of 2 Stumpffia helenae sp. n. (paratype: NMBE 1044801). osteological analyses - e.g. enlarged end of the phalanx - were impossible without damage them). Dorsal skin slightly granular. Tibiotarsal articulation reaches the posterior edge of the eye. Measurements of the holotype: head width: 4.5; forearm length: 3.0; hand length: 2.1; tibia length: 6.6; foot length including tarsus: 9.3; foot length: 5.4; tympanum diameter: 0.7; eye diameter: 1.3; distance from eye to nostril: 0.9; distance from nostril to snout tip: 0.6. The left forearm is broken. Description of the paratype: A 2 of 15.0 mm length (since the size of the paratype is similar to that of the holotype (adult 4 ) and nuptial pads are absent, it can be assumed that the paratype is an adult 2). Similar to the holotype except fingertip less enlarged (Figs 2a and 2b), and the skin on the back smoother than in the holotype. Measurements of the paratype: head width: 5.1; forearm length: 3.5; hand length: 3.1; tibia length: 7.2; foot length including tarsus: 10.7; foot length: 6.2; tympanum diameter: 1.0; eye diameter: 1.6; distance from eye to nostril: 0.9; distance from nostril to snout tip: 0.7. Colour in life: The 3 had a dark grey, marbled colour on the back (Fig. la). In the inguinal region two black spots were present (probably an imitation of eyes). An A NEW SPECIES OF THE GENUS STUMPFFIA 839 FIG. 2 Hand (a) and foot (c) of & Stumpffia helenae sp. n. (holotype: NMBE 1044802). Hand (b) and foot (d) of 2 Stumpffia helenae sp. n. (paratype: NMBE 1044801). orange line was present on the middle of the back, extending from the middle of the head to the posterior end of the body. On both sides of the dorsal line two lines of small weakly elevated light spots were visible. A thin creamy and golden line extended from the humeral joint of the arm passing above the eye to its posterior edge. The sides of the head were dark grey but not marbled. The dark grey colour of the flanks (same colour as the sides of the head) was covered by lighter spots. The colour of the dorsal side of arms and legs was similar to the colour of the back but a 840 DENIS VALLAN bit lighter. The throat, the belly and the ventral sides of the extremities were light grey with a few indistinct lighter spots (Fig. Ic). The chest was also light grey but covered by irregular white patches, giving to the chest a netlike impression. No distinct border present between the coloration of the dorsal and ventral side. The oval (horizontally flattened) pupil is surrounded by a golden iris. Compared to the coloration of the d, the coloration of the 2 shows the following differences (Fig. 1b and Id): the colour of the back was rather brown than grey, the orange line present in the d was nearly not visible in the © and the primary colour of the chest was not light grey but pink. Colour in preservative: After 22 month in preservative the colour has changed little. The brown colour of the back of the 2 and the orange line on the back of the d changed into grey and light grey respectively. The pink colour of the © chest faded out. On the ventral side of the legs the indistinct white spots became visible. Etymology: I wish to dedicate this new species to my wife and friend Helena Bigler for her support of my research and her enthusiasm for Madagascar. Distribution: Stumpffia helenae sp. n. is just known from the type locality. Habitat and habits: Both known animals were found in the high plateau rain forest of the RS d'Ambohitantely, which was described by Koechlin (1972) as “high- altitude lichen forest”. This forest contains wetter (riparian) and dryer (on slopes, plateau and ridges) forest types as described by Razakanirina (1993). The 2 was active during the day in the leaf litter of the riparian forest 1 m from a brook. The d also was active during the day in the leaf litter but far from brooks in the dryer slope forest. The riparian forest is characterised by trees to a canopy height of 20 m and a relatively sparse undergrowth vegetation, whereas the slope forest is characterised by a lower canopy and a denser undergrowth vegetation. DISCUSSION It was surprising to find a Stumpffia in Ambohitantely since this genus was mainly known from middle or lower altitudes. S. gimmeli is known from the Montagne d'Ambre, S. roseifemoralis from the Marojezy massive and S. grandis from both mountains, but they do not reach altitudes much higher than 1000 m a.s.l.. Raxworthy & Nussbaum (1996) mentioned a not specified Stumpffia species from the Tsaratanana mountains in 2050 m a.s.l.. Stumpffia helenae sp. n. is one of the few records of this genus from higher altitudes (1500 m a. s. 1.) and the first record in the central high plateau of Madagascar. The forests of the high plateau of Madagascar are among the most threatened forests of Madagascar. The "Réserve Spécial" of Ambohitantely is a nature reserve containing one of the last forests. It consists of hundreds of islands of forest, their sizes ranging from 0.16 to 1250 ha making up a total of 2737 ha about half the total reserve area (Langrand and Wilmé, 1997). The paratype was found in the 1250 ha fragment, whereas the holotype lived in a 30 ha fragment. The fragmentation of the forest started centuries ago. Small forest fragments may maintain stable amphibian populations since outside the forest, in the almost completely cleared landscape, the A NEW SPECIES OF THE GENUS STUMPFFIA 84] conditions for the survival of this species are not favourable (high temperature and low air and soil humidity). The discovery of this new species shows the importance of even small forest remnants for conservation. ACKNOWLEDGMENTS I am very thankful to Monica Biondo for drawings and to John E. Cadle, Frank Glaw, and an anonymous reviewer for the review of the manuscript. I thank Wolfgang Nentwig and Walter Hödl for their support, Solofo and Armand for the help in the field, Wolfgang Böhme (Museum Koenig, Bonn) and Gunther Kohler (Sencken- bergmuseum, Frankfurt) for the loan of comparative specimens. Claudio Vallan kindly reviewed an early draft of this paper. The research in Madagascar was made possible by scientific co-operation with the University of Antananarivo, the financial support of the Swiss Academy of Natural Sciences (SANS) and the Natural History Museum of Bern. The research was also supported by Air Madagascar, Photo-Wyss (Ziirich) and Tramp Store (Trimbach) and authorized by the Direction des Eaux et Foréts (DEF), Association Nationale pour la Gestion des Aires Protégées (ANGAP), and the Commission Tripartite. REFERENCES DUELLMAN, W. E. 1970. The Hylid frogs of Middle America. Monographs of the University of Kansas Natural History Museum 1: 1-753. GLAW, F. & VENCES, M. 1994. A fieldguide to the amphibians and reptiles of Madagascar. 2nd edition, including mammals and freshwater fish. Vences and Glaw, Koln. KOECHLIN, J. 1972. Flore and vegetation of Madagascar (pp. 144-190). In: BATTISTINI, R. & RICHARD-VINDARD, G. (eds). Biogeography and ecology of Madagascar. Dr. W. Junk B. V., Publishers, The Hague. LANGRAND, O. & WILME, L. 1997. Effects of forest fragmentation on extinction patterns of the endemic avifauna on the Central High Plateau of Madagascar (pp. 280-305). In: GOODMAN, S. M. & PATTERSON, B. D. (eds). Natural change and human impact in Madagascar. Smithsonian Institution Press, Washington D.C. RAZAKANIRINA, D. 1993. Le jardin botanique de la réserve speciale d’Ambohitantely. Akon'ny Ala 11: 50-52. RAXWORTHY, C. J. & NUSSBAUM, R. A. 1996. Montane Amphibian and Reptile Communities in Madagascar. Conservation Biology 10: 750-756. VENCES, M. & GLAw, F. 1991. Revision der Gattung Stumpffia Boettger 1881 aus Madagaskar mit Beschreibung von zwei neuen Arten (Amphibia, Anura, Microhylidae). Acta bio- logica Benrodis 3: 203-219. REVUE SUISSE DE ZOOLOGIE 107 (4): 843-869; décembre 2000 Nuclear and mitochondrial markers reveal the existence of two parapatric scorpion species in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha Caporiacco, 1950, stat. nov. (Euscorpiidae) Benjamin GANTENBEIN!, Victor FET?, Mark BARKER? & Adolf SCHOLL! ! Institute of Zoology, Division of Population Biology, University of Berne, Baltzerstrasse 3, CH-3012 Berne, Switzerland. E-mail: gantenbein @zoo.unibe.ch 2 Department of Biological Sciences, Marshall University, Huntington, West Virginia 25755-2510, USA. E-mail: fet@marshall.edu Nuclear and mitochondrial markers reveal the existence of two para- patric scorpion species in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha Caporiacco, 1950, stat. nov. (Euscorpiidae). - A molecular (mtDNA and allozyme) data set reveals a clear divergent phylogeny within the Alpine scorpion species Euscorpius (Alpiscorpius) germanus (C. L. Koch, 1837). Two distinct (ca. 7 % DNA sequence diver- gence), monophyletic clades exist which are geographically separated by the Adige (Etsch) River in northern Italy. At the allozyme level, these population groups are fixed for alternative alleles at eight out of 18 gene loci and correspond roughly to the morphological subspecies E. g. germa- nus and E. g. alpha. No evidence of introgressive hybridization between the two groups is shown by the allozyme data. The branching points of the two population groups are found at unusually high distances compared with the outgroup taxa E. gamma Caporiacco, 1950 and E. flavicaudis (De Geer, 1778). The subspecies E. g. alpha is therefore elevated to species level: Euscorpius alpha Caporiacco, 1950, stat. nov. A neotype for E. germanus (C. L. Koch, 1837) and lectotypes for E. alpha stat. nov., E. germanus beta syn. nov. of E. alpha, and E. germanus croaticus are designated. Phylo- genetic and biogeographic implications are discussed. Key-words: Scorpions - trichobothria - allozymes - 16S mtDNA - biogeo- graphy - parapatric species - Alps - phylogeny. INTRODUCTION Several species of the scorpion genus Euscorpius Thorell, 1876 (Euscorpiidae) are common in the circum-Mediterranean region and in southern Europe. Taxonomy of this genus is not well resolved; there are currently seven “good” species, and 40 (!) formally valid subspecies in Euscorpius (Fet & Sissom, 2000; Scherabon er al., Manuscript accepted 25.04.2000 844 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL 2000). Recently, we began to apply molecular data to solve complicated taxonomic problems in this genus (Gantenbein & Scholl, 1998; Gantenbein er al., 1998, 1999a; Fet et al., 1999; Scherabon er al., in press). This paper reports new phylogenetic data, based on the analyses of mtDNA sequences and allozymes, from the Alpine populations, which have been traditionally placed in Euscorpius germanus (C. L. Koch, 1837). E. germanus is a mountainous species, recorded from the western Balkans, Austria, northern Italy and southern Switzerland (Capra, 1939; Caporiacco, 1950; Valle et al., 1971; Bonacina, 1980; Scherabon, 1987; Crucitti, 1993: Fet & Braunwalder, 1997; Gantenbein er al., 1998). Scorpion taxa are traditionally classified using morphological characters such as variation in the trichobothrial numbers and patterns (‘trichobothriotaxy’) (Birula, 1900, 1917; Hadzi 1931; Vachon 1962, 1981). However, molecular markers have recently become a powerful tool for evaluating the taxonomic status of populations and subspecies/species. The combination of nuclear and mitochondrial markers has been efficiently applied to detect introgression between taxa (Barton & Hewitt, 1989: Harrison, 1990; Bernatchez et al., 1995). The subspecies Euscorpius germanus gamma Caporiacco 1950 was recently elevated to species rank (Scherabon er al., in press) after using allozyme and mtDNA data. Gantenbein et al. (1998) and Gantenbein & Scholl (1998) demonstrated that the Swiss E. germanus populations probably have originated from two different refuges during the glaciations, forming two genetically highly divergent population groups. In order to further clarify the taxonomic status of the two subspecies E. g. germanus and E. g. alpha and to confirm the hybrid zone between these taxa assumed by Bonacina (1980), we initiated a molecular survey applying previously established nuclear (allo- zymes) and mitochondrial (16S mtDNA sequences) gene markers. METHODS AND MATERIALS SPECIMENS ANALYSED A map of Switzerland, northern Italy, Austria and Slovenia from where E. germanus samples were collected is given in Fig. 1. Two outgroup species were collected, Euscorpius gamma Caporiacco, 1950 (see Scherabon er al., 2000) from Koschuta (Carinthia, Austria) and Euscorpius flavicaudis (DeGeer, 1778) from Lauris (Vaucluse, France). The animals were caught in a sampling area of about 100-300 m? and were brousht alive to the laboratory where they were killed by deep-freezing and stored in -80°C prior to biochemical analyses. In general, only few animals were taken from one site because small sample sizes (N < 10) are already expected to result in relatively good estimates of gene frequences at allozyme loci. Previous studies reported a low genetic variability within E. germanus populations (Gantenbein er al. 1998, 1999a). Sample sizes are given in appendix I. After biochemical analyses, the animals were transferred to 70 - 80% ethanol for morphological analysis. 845 PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) BINYISONY ES !punund 7 dno1TINO “(VILIOD PAON IVAU) SAU) TE UNEIOPIT [TS “NNYIS OF “Yosie.qod Gp ‘SINQNEIPIOGO Bp ‘YOR|[Ed Lt ‘Yyorzuesy Op ‘ZuauE], Sp ‘YOEQUINIPIS pp ‘o1opeD IP ozuoiny Ep “MOPV) IP CIO Th ‘APUEIOZZIAIN [t ‘OSELIOL URS Op ‘OTEIOA HE “OUNT[IG YE ‘OTOLMOA LE “OWOIG 9€ ‘SOA SE ‘UOXIIg PE ‘SUIPIOA EE “SUIOPULTYOS TE “SULOPN|YOS TE “S1oquayyor'] YE ‘ELET LIS 67 ‘75-67 Sa[dues (snunundd ‘7) APE] use ‘zes peg 87 ‘Burpy LT ‘0IPOT Ip BUIIOIN IT ‘9997794 CT 'OJOSIA FT ‘EIOUIDALI €T ‘OIQUION 77 ‘OUTA]9S [Z ‘PABA 07 ‘LUOILD GI ‘VAPUO, IP LOS] 8] ‘OPJOZZOW LI ‘QUONEZZEIA 9] “OqUiolg [E OO ST “O[[SUIOD vIvIOWIR FI “OURIG TUURAOIDH UES £] ‘OULISa]Joq URS 7] BIOygad II ‘PSOI99 OI TIquaig 6 ‘ORD URS 8 ‘AIUOdONOS / ‘apnuod 9 ‘aovUIO.] ¢ ‘UO f ‘ROURY £ ‘AIOUQUTEJUO 7 ‘Opuor) | ‘87-] sojdues :(pydyv 7) pero UI9JS9 MA “SOUS Surdues [ 'DIH ONV9OH38 crs 2 equaig SE 6 SR . o BSO1a5 So wy 001 ° OL Je ooueig obL 7e 3 Elo)\uuenog Ss 224684 x oljawog oquiaig |e SL ounoL eızaus\ -BZZEIJo opiozza eueqwaug 3lleA yonıqsuu| \eugsny 846 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL DNA ANALYSIS DNA extraction and sequencing techniques: A comparative analysis of the mitochondrial 16S ribosomal RNA and allozymes has been recently used for resolving species-level phylogeny of Euscorpius (Gantenbein et al., 1999a); this contribution should be consulted for the technical details and protocols. Total DNA was extracted from fresh or preserved (95% ethanol) muscle tissue using a standard extraction method. An approximately 400 bp long fragment of the mitochondrial (mt) 16S rRNA gene was amplified by the polymerase chain reaction (PCR) using the primers 16Sbr (= LR-J-12887), (Simon et al. 1994; CGATTTGAACTCAGATCA; forward, 18-mer) and a scorpion-specific reverse primer (GTGCAAAGGTAGCA- TAATCA, 20-mer). A total of 25 mtDNA sequences was used for the analysis (Table 1). For further analysis, all ambiguities and indels were excluded, as suggested by Swofford et al. (1996), with 357 characters remaining. Haplotype diversity: We calculated the haplotype (gene) diversity (Nei, 1987), the nucleotide diversity m (Nei & Li, 1979) and the number of segregating (poly- morphic) sites (S) among sequences of E. germanus. Neutrality of mutations within each species was examined by using Tajima’s D test (1989). The genetic variability estimates and the neutrality tests were calculated using the computer program DnaSP (Rozas & Rozas, 1999). Phylogenetic analyses: We applied character-matrix-based methods (maxi- mum parsimony (MP) and maximum likelihood (ML)) methods (Felsenstein, 1981a) as well as distance-based methods (neighbour-joining (NJ) cluster algorithm) (Saitou & Nei, 1987). The beta-version of the computer program PAUP* 4.0 (Swofford, 1998) was used for all phylogenetic DNA analyses. We calculated hierarchical likelihood ratio tests, in order to find the most appropriate model of DNA substitution, using the program MODELTEST 2.0 (Posada & Crandall, 1998). This program calculates the likelihood ratio statistic à = —2 log A where A is defined as __ max [Ly(NullModel |Data)| ‘max [Li(AlternativeModel |Data)] with Ly being the likelihood under the null hypothesis (simple model) and L, being the likelihood under the alternative hypothesis (more complex, parameter rich, model). When the models compared are nested (the null hypothesis is a special case of the alternative hypothesis) and the null hypothesis is correct, the 6 statistic is asymptotically distributed as x2 with q degrees of freedom (q is the difference in number of free parameters between the two models). In the next step, a test for the molecular clock hypothesis (i. e. rate constancy among lineages) was calculated (x2 distributed, df= N - 2 OTUs). Details on model testing using maximum likelihood ratios are given in Huelsenbeck & Rannala (1997) and in Huelsenbeck & Crandall (1997). The likelihood ratio tests suggested the Tamura & Nei (1993) model with rate heterogeneity (TrN93 + I), which is a submodel of the general-time-reversible (GTR) substitution model (Rodriguez er al., 1990, Yang et al., 1994). All parameters (base frequencies, rate matrix) were estimated via maximum likelihood. The rate PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 847 heterogeneity among sites was assumed to follow a gamma distribution (shape para- meter a was ML-estimated) with four categories, each represented by its mean (Yang, 1994). Phylogenetic analysis is facilitated when rates are equal among lineages. However, the test for the molecular clock was rejected at the 0.01 level, therefore, the tree search was carried out without enforcing the clock. For ML analyses the tree space was explored using the heuristic search option implemented in PAUP* with random addition of sequences (100 replicates, tree bisection-reconnection (TBR) branch-swapping algorithm). For the MP analysis the transitions (ti) were weighted twice over transversions (tv) according to the ML estimated ti / tv ratio using the HK Y85 (Hasegawa er al., 1985) model, and the tree search was done using the branch-and-bound search option. To save computing time, identical haplotypes were eliminated. The consistency index (CI) and the retention index (RI) (Kitching er al., 1998) were calculated as measures for tree stability with PAUP*. Alternatively, pairwise ML-distances were estimated using the TrN93 + [ model. Estimating distances via ML has the advantage of constant parameters over all pairwise comparisons and consequently the variance of distances is reduced. These distances were used as a matrix for neighbour-joining (NJ) clustering (Saitou & Nei, 1987). NJ is assumed to be a good heuristic approach for estimating the minimum evolution tree (Page & Holmes, 1998). The trees were rooted using two outgroup species: E. gamma and E. flavicaudis. The trees were bootstrapped resampling 1,000 data sets with 357 characters. DNA SEQUENCE AVAILABILITY All sequences were deposited in the EMBL Nucleotide Sequence Database with the following accession numbers: E. alpha stat. nov.: EalGO (= EalFO) = AJ389379; EalRA = AJ271886; EalSP = AJ286751; EalSG = AJ286752; EalCA = AJ286753; EalOL = AJ286754; EalSO = AJ286755; EalTA = AJ286756; EalMA (= EgeML) = AJ286757; E. germanus: EgeVO (= EgeSH = EgeSM = EgeTz = EgeKR = EgeVE) = AJ389380; EgeOB (= EgeDE = EgeST) = AJ249553; EgeBO = AJ286758; EgeME = AJ286759; EgeCR = AJ249552; E. gamma: EgaKO = AJ249554; E. flavicaudis: EfILA = AJ389381. Abbreviations for haplotypes are given in appendix I. ALLOZYME ANALYSIS Horizontal starch gel electrophoresis of allozymes was carried out according to the protocols described in Harris & Hopkinson (1976) and Murphy er al. (1996). We scored the same 18 loci as described in Gantenbein er al. (1998): N-(3-Aminopropyl)- morpholine-citrate (AC, pH 6.2, modified from Clayton & Tretiak 1972), Tris-citrate (TC, pH 7.3, Ayala et al. 1972) and Tris-borate-EDTA (TBE, pH 9.3, modified from Ayala et al., 1972). The loci scored were: AAT-1 and AAT-2 (aspartate amino- transferase; EC 2.6.1.1), ALPDH (alanopine dehydrogenase; EC 1.5.1.17), ARK (arginine kinase; EC 2.7.3.3), DDH (dihydrolipoamide oxidase; EC 1.8.1.4), GAPDH (glyceraldehyde-3-phosphate dehydrogenase; EC 1.2.1.12), GTDH (glutamate dehy- 848 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL drogenase; EC 1.4.1.2), GPI (PGI) (glucose-6-phosphate isomerase; EC 5.3.1.9), HK (hexokinase; EC 2.7.1.1), IDH-1 and IDH-2 (isocitrate dehydrogenase; EC 1.1.1.42), MDH-1 and MDH-2 (malate dehydrogenase; EC 1.1.1.37), MPI (mannose-6- phophate isomerase; EC 5.3.1.8), PEP (pepdidase; EC 3.4.-.-), PGM (phospho- glucomutase; EC 5.4.2.2), 6-PGD (6-phosphogluconate dehydrogenase; EC 1.1.1.44), and PK (pyruvate kinase; EC 2.7.1.40). We refer to the observed electromorphs as alleles which are identified by their electrophoretic mobility relative to the most common mobility in the E. flavicaudis population from Lauris, France (assigned mobility=100) as described in Gantenbein er al. (1998). To assess the genetic variability within each population, the mean number of alleles per locus, the percentage of polymorphic loci and the mean heterozygosity were calculated by the direct count method and by Nei’s (1978) unbiased estimate. Calculations were done using BIOSYS-1 (Swofford & Selander, 1989). Cavalli-Sforza & Edwards’ (1967) chord distance was calculated from pairwise comparisons of populations using the program GENDIST from the PHYLIP 3.5 package (Felsenstein, 1995). Using Nei’s pairwise distances as an input matrix, an additive tree was created by the neighbour- joining algorithm (NJ). Alternatively, an unrooted maximum likelihood tree was calculated using the computer program CONTML. This estimates phylogenies by the restricted maximum likelihood (REML) method, based on the Brownian motion model (Cavalli-Sforza & Edwards, 1967). The REML algorithm was described in Felsenstein (1973, 1981b). It uses less parameters than the full ML analysis and is therefore considered to be more consistent. Additionally, the program calculates branch lengths and rough confidence intervals for the branches. Bootstrap values were obtained from 1,000 pseudo-replicates of allele frequencies using the SEQBOOT routine in PHYLIP. MORPHOLOGICAL ANALYSIS We scored the number of pectinal teeth (Dp) and the numbers of trichobothria on the ventral (Pv) and external (Pe) aspects of the palpal patella (called tibia by some authors; see Hjelle, 1990). & d have higher numbers of pectinal teeth. We tested the differences between two discovered clades using a one-sided r-test. RESULTS MOLECULAR ANALYSES mtDNA data: We analysed 25 mtDNA sequences representing 17 different haplotypes. The heuristic tree search (100 replicates) using maximum likelihood (ML) revealed a single tree with a In likelihood of —904.57 (Fig. 2A). The nucleotide frequencies within the 16S mtDNA were estimated via ML to A = 0.332, C = 0.13, G = 0.12, and T = 0.41, respectively. The substitution rate matrix (R) was estimated via Mi to A <--> C=1, A <>+G=19.0, A<>T=1,C<>G=l € < 2 207 and G <--> T = 1. The shape parameter a of the gamma distribution was estimated via ML to 0.085. The relatively low estimate of a indicates a high rate of hetero- geneity among nucleotide sites. The tree topology of the ML tree was identical to that PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 849 EalFO EalFO EalSP EalSP 79 87 EalSG (71) a EalSG EalCA Western clade EaCA = E, alphastat. nov. 13 EalOL. # EalOL 95 EalSO EalSO EalTA EalTA EaMA 70 EgeOB EgeOB EgeBO EgeBO 94 EgeVO Eastern clade EgeVO = E. germanus 93 EgeME 64 EgeCR EgaKO = EfILA 0.01 substitutions/site A B Fic. 2 (A) Maximum likelihood (ML) phylogeny based on the 16S mtDNA gene sequences in the “western” clade (Euscorpius alpha stat. nov., Eal) and in the “eastern” clade (E. germanus, Ege). The -In Likelihood was 904.57 using the model by Tamura & Nei (1993) with rate heterogeneity (TrN93 + I). (B) Strict consensus tree of six equally parsimonious trees (91 steps, CI = 0.80, RI = 0.85) calculated by weighted maximum parsimony (MP). Numbers at nodes refer to bootstrap values calculated from 1,000 pseudoreplicates. Bootstrap values in parentheses (in A) are from neigbour-joining (NJ) analysis, which resulted in the same tree topology showing a deep split between both clades (species). Abbreviations for haplotypes are explained in appendix I. of a NJ tree (tree not shown) which was built using TrN93 + I distances. The bootstrap values for the ML tree and the NJ analysis (in parentheses) are given in Fig. 2A. This phylogeny splits all analysed mtDNA sequences of E. germanus into two clearly distinct clades (“western” and “eastern”), both supported by relatively high bootstrap values. The analysis using weighted maximum parsimony (MP) revealed six equally parsimonious trees with a tree score of 91 steps. 297 characters were constant, 27 characters were parsimony-informative. The consistency index (CI) for all eight trees was 0.80 and the retention index (RI) was 0.85, respectively. Both indices indicate a high tree stability. The strict consensus tree is shown in Fig. 2B. The deep splitting of 850 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL two population groups in the MP analysis is consistent with the ML tree and the NJ tree. However, the sequence from Crnice (EgeCR) showed an ambiguous grouping in ML and NJ analysis. Therefore, the bootstrap values for the two clades were moderate (about 70%) in ML and NJ analyses. This was not the case in MP analysis where both clades were supported by high values (about 90%). The DNA polymorphism of the 16S data is listed in Table I and in Appendix II. The analysis of DNA variation revealed that 16 sites out of 357 characters were polymorphic (segregating) among the eleven “western clade” sequences whereas 14 sites out of 364 were polymorphic among the twelve “eastern clade” sequences. The probability that two randomly chosen haplotypes are different (= gene diversity) was 0.94 and 0.72, respectively (Table 1). Within the “western clade”, the average nucleo- tide diversity m was 0.03 + 0.00, whereas in the “eastern clade” it was close to zero. Tajimais D test statistics were not significant for both species. TABLE 1: mtDNA diversity measures within the “western” and “eastern” clades (E. alpha stat. nov. and E. germanus, respectively). haplotype Average Sample size Number of (gene) Polymorphic Total Number numberof Nucleotide 6 (=2 Ne p) per Tajima’s n haplotypes diversity sites S of sites* nucleotide diversity 7 site DEE differences Western clade = E. alpha stat. Il 8 0.95 16 357 5.78 0.0322 0.016 0.26 nov. Eastern clade = E. germanus 12 5 0.72 8 364 3.12 0.009 0.008 -1.39 * excluding indels and ambiguities. ** P>0.10 Allozyme data: The NJ tree based on allozyme gene frequency data at 18 loci (appendix I) independently revealed a tree topology comparable to those obtained by 16S mtDNA sequence analysis. It splits all populations of E. germanus examined into two highly divergent groups, a “western” and an “eastern” one (Fig. 3). However, bootstrap values of these two groups were not as high as in DNA analysis. This is caused by the Crnice population, which in some cases exhibited ambiguous clade groupings. The In Likelihood of the best tree (tree not shown) using the Restricted ML (REM) criterion by Felsenstein (1981b) was -3647.51970 (14,688 trees explored) and the topology was identical to that revealed by NJ analysis. The genetic variation estimates within both clades are given for samples N 2 4 in Table 2. The populations were fixed at many gene loci, therefore, the mean number of alleles was approximatively one. Low genetic variability estimates were found for the mean number of loci polymorphic (16%) and for the average heterozygosity (0.03 + 0.02) (Table 2). MORPHOLOGICAL ANALYSIS We measured the number of pectinal teeths (Dp) in d d and © 2 from each of the two major clades recognized by molecular analyses (Figs. 2-3). 22 of the “eastern clade” had significantly higher number (t = 2.29, P = 0.011) of pectinal teeth ((Dpjeg + DPiiopt) / 2) (mean = 6.05, sz = 1.10, N = 83) than the © © of the “western 851 PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) "soyeaıpdaaopnasd 000° | JOAO POPEINITEI sanyea dENSI0OQ 0) Jaya sapou sy} je SIOQUUNN ‘190] QUIAZO][e g] UO poseq ale sooURIsSIG 'xıyew Jndur ue se (/96] “SpreMpy W BZIOJS-I[[LALI) SOUPISIP PIOU9 VZIOJS-1][LALD Sursn (sa1sads) sope[o Om) au} JO (+ = N) sejdures Jo sısAjeur (N) Surutol-mMoqgysioN ‘€ ‘OI SOUE]SIP P1OU9 (2961) 2Z10/S-IRACI 0 0010 0070 00€°0 00r'0 006'0 Lr EE SIPNDIIADY "7 DWWDD 'J =— SUN ounyag 69 9J0PD) Ipozuoiny bINAQNDIPpijogO L6 ur: sos ened SNUDW.AOD 'J = UDDQUENIDIS SPD U19}SD9 911PD) IP DIIOg ODD}}OA 91DA UIDZUDJY UDXII 0101157 v8 J9W91q 6/ SUIP19A SIQA SUJOPN|UIS b13qua}u217 JIDISNW D239zz9g DIOUJSADL bunNJDW 01p91 Ip DUNOW opauod ouI163}]94 UDS OZ QUIA}9S 4 pniquusi “AOU *}D}S DYG]D ‘7 = D4APUOY Ip Bios È) et hel be DUOIDI 87 SPD uU194S3M ODUDIg IUUDAOID UDS oqwaJg 10 OWIO 2110}0ZZDId OPIOZZaW O}1DD UDS 92DU104 9}DIUDY ajuodo}}os AJUOW opuog 3JOWIUIDJUOY 81 852 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL TABLE 2: Genetic variability estimates between the “western” and “eastern” clades (E. alpha stat. nov. and E. germanus, respectively) based on 18 allozyme loci (see appendix I) (N = 4). Population See Un À D © 00 SA À © D 16 Western clade = E. alpha stat. nov. Gondo Fontainemore Rancate Monte Fornace Pontide Sottoponte San Carlo Brembilla San Pellegrino San Giovanni Bianco Olmo al Brembo Piazzatorre Mezzoldo Isola di Fondra Carona Selvino Tavernola Bezzecca Molina di Ledro Marling Mean # ** Unbiased estimate (Nei, 1978) Eastern clade = E. germanus Sta Maria Lichtenberg Schluderns Verdins Brixen Völs Bremer Vetriolo Belluno Voltago San Tomaso Borca di Cadore Auronzo di Cadore Starkenbach Tarrenz Kranzach Dellach Oberdrauburg Dobratsch Schiitt Federaun Crnice Mean aos Overall Mean exceed 0.99 (n ui | | _ { nd dt pt nd pt pt pd BWI RERBRUWNDAWRWADRNNANWNOW tt H+ HH H+ HH HH HH HH H+ HH tt lt o pos 0 DDD DD 000000 D IINNDNNTNTNNNNNNNTTNCcINOT I+ HHHHHHHHHHHHHHHH I Ht IE + sS|o|SSSSSSSSSS92992990999099 mim DI DD SS SR ut I+ 3 SENIO pp pe EE ER i) — D = D De ee © 19 09 ee Dom DD el I+ Mean no. of alleles Percentage of polymorphic loci — jean ©. n D | DDD- D = 09 = DD WWND DH UD ni ni IONI 9909 SISO OS ON EE DN 00 WIN © 1 10 © Lo Us 00 D YW Yee ON olSoo009800000000000000000 oo go Mean heterozygosity Direct count expected Hardy-Weinberg o csosssssssssssssssssss here) eee) Un DUT IRLCAUNULROROERRLO OO I+ It 1+ 1 HH HH 1+ 4 HH HH HH 14 EH SO ossssssssssssssssssss Sree '@' Sle |S SiS'S SSN) SI OO DODODODPOLOODPRUNUDDRDDWOO I+ RO RO OO BR 100% NN ND D D = ONO EEE A AA AR REA RER AIRE SOSEO00222092.99090999009090 HH 1 IH 1 18 1 1 18 HE 1 1 1 HI IH SS _ SO SO _ SS OSO SO SO _SO_ SO OSO SOS SO _S _ SO SOS a [USSR RUE ES GERS RON ls SSIS | Oo} OCC eee) EF OOF OSSO | OO mr N 0010 BONO CORP OR SDA [DO OO ee nm OOWRhWABRDORE NRK ORF O- SSSSSSSSSSSSssssoSessssss SSscococoscococsoccecosce | I+ | | sean) SU IA OS © © DA © Li oO © D BR © © © BR À D HHHHHHHHHHHHHHH 1 1 HI IE Sooooo00o0000000000000 À © GS Un © © © PD Ca © Un © = © BW © © © SSSSSSSSSSsSssssssssesss =) ne O OOO DD SS DOO 22220 o o0o00005000000 000000000 Oo I+ (N°) 10.04 +0.03 Go I+ © i “ A locus is considered polymorphic if the frequency of the most common allele does not 0.06 + 0.04 853 3) PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE = 970 OLO i ant 90 0 IO 90°0 [TO 90 0 ITO 90°0 110 90°0 OLO 500 ILO 500 ITO 900 ant 900 [TO 90 0 ant 90 0 (and 90°0 010 90 0 VIUA OMA 30934 S9 0 dat ÿ0 0 €00 €00 €00 90°0 90 0 L0°0 L0°0 L0°0 L0°0 L0°0 LO 0 at vs 0 80 0 000 100 00°0 90°0 900 LO'0 L0°0 LO'0 LO'0 LO'0 90 0 AN°$4 OA°$4 90 er 0 L0°0 00°0 00°0 00°0 90 0 900 90 0 L0°0 LO'O LO'O 90 0 90 0 6S 0 9¢ 0 LO°0 10°0 00°0 00 0 soo 900 900 L0°0 90°0 90°0 90 0 90 0 O4°54 S9 0 ev 0 90°0 00°0 00°0 00°0 90 0 900 90 0 L0°0 LO'O L0°0 90°0 90°0 doesq VINIPA. VIII Osea TOI SV 0 60°0 810 6€ 0 IE 0EO [ce 0 10°0 c0°0 c0°0 c0°0 c0°0 c0°0 co 0 870 O10 610 6€°0 [LEO 0EO Ic 0 100 c0°0 c0°0 c0°0 c0°0 c0°0 €0°0 590 0c 0 sro 19°0 670 LVO 670 €0°0 €00 00 0 000 c0°0 10°0 c0°0 c9°0 610 0Oÿ 0 OLO 96 0 750 950 €0°0 £00 00 0 000 co 0 co 0 c0°0 590 STO 9C°0 [80 790 c9°0 790 ÿ0 0 S0°0 00°0 00°0 c0°0 c0°0 co 0 VOIE 980 (ay) OS 0 690 ps0 cs 0 750 700 S00 €0°0 €0°0 ÿ0 0 00 0 c0°0 OST EH 6L'0 0c 0 970 c9'0 6r 0 Ly0 670 70°0 00 co 0 co 0 £00 000 c0°0 dSI'd sro STO 8€0 [SO Iv 0 6£ 0 [7 0 po 0 C0 0 €O 0 €00 c0°0 00 co 0 Oaled VIVA ONCSA 49954 Aw>34 OA»3AI og>34 40954 vneg vLeA OSICA TOA Voled Osea dsıed OAICA ‘saouanbes YNGI Sg] Jo suosLeduos osımamed Woy payent (Joy JOMO]) JT + E6NAL JO pue (ysıı soddn) (d pagoariooun) sdudSJ0AIp oouonbos oy) JO xLmew F9UBISIC :€ AIAVL 854 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL clade” (mean = 5.68, s = 1.41, N = 111). This corresponded to a unimodal distri- bution of this character in the “eastern clade” (common Dp = 6) versus bimodal in the “western clade” (Dp = 5 or 6). 6 d of the “eastern clade” also had significantly higher number (t = 1.73, P = 0.045) of pectinal teeth (mean = 7.58, s* = 2.1, N = 24) than 36 of the “western clade” (mean = 6.98, s? = 1.97, N= 60). This corresponded to a bimodal distribution of this character in the “eastern clade” (Dp = 7 or 8) versus unimodal in the “western clade” (common Dp = 7). The average number of trichobothria on the ventral aspect of the pedipalp patella ((Pvjeg + PVright) / 2) showed more variation. Among populations belonging to the “eastern clade”, the LI of ventral trichobothria ((Pvjg + Den) / 2) was more constant (mean = 4.96, s? = 0.24, N = 126) and no geographic pattern was detectable. Within the “western clade” the mean Pv was higher and more variable (mean = 5.60, s? = 0.34, N = 146) and also exhibited a considerable geographic variation. In the center of its geographical range (Bergamascan Alps) the Pv character was fixed around 6. However, in the marginal populations of the “western clade”, this character was fixed at ca. 5. This holds true for both, the westernmost (Fontainemore and Gondo), as well as the easternmost populations (Tavernola, Molina di Ledro, Bezzecca, Marling and Bad Salz). DISCUSSION TAXONOMIC SUBDIVISION OF EUSCORPIUS GERMANUS (C. L. KOCH) Before the advent of chaetotaxy (trichobothrial pattern) analysis, taxonomy of Euscorpius species was extremely confusing and was based mainly on morpho- sculpture and coloration characters. Large and conspicuous trichobothria of Euscorpius as taxonomic characters were first studied in detail by Hadzi (1929, 1931) and Caporiacco (1950), who used overall trichobothrial counts of pedipalp chela and patella for identification of species and subspecies. Euscorpius germanus (C. L. Koch, 1837) has been originally described from “southern Tirol [i.e. today’s Trentino - Alto Adige in Italy] and northern Italy”; see Fet & Braunwalder (1997) for the detailed taxonomic history and authorship discussion. This species traditionally included several subspecies with rather unclear diagnostics (Caporiacco, 1950). Although Birula (1900) already clearly demonstrated species-level differences between E. germanus and the Caucasian E. mingrelicus (Kessler, 1876), several authors later often confused these two species. Until 1980, E. germanus was treated as a widely distributing species (from Italy to Caucasus) with a number of subspecies (Hadzi, 1929; Caporiacco 1950; Curcic, 1971; Kinzelbach, 1975). Bonacina (1980) limited E. germanus (C. L. Koch) to the Alpine regions of Italy (from Piemonte in the West to Goriziano in the East), Switzerland, and Austria, plus some Balkan populations; and reestablished as a “good” species E. mingrelicus (western Balkans and Anatolia to Caucasus, with notable absence in Greece) (Bonacina, 1980; Fet, 1993; Fet & Sissom, 2000). Most recently, we (Scherabon er al., 2000) demonstrated PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 855 presence of more than one species within the “E. mingrelicus complex”, including E. gamma Caporiacco, 1950. Hadzi (1929) was the first to establish subspecies of E. germanus; however, his names (polytrichus, mesotrichus and oligotrichus) are invalid since they are homonyms, and therefore replacement names are necessary. Moreover, they were not sufficiently defined to allow identification of these taxa (Fet, 1997). Besides, most populations of “E. germanus” from ex-Yugoslavia treated by Hadzi (1929) and by Curcic (1971) do not belong to this species as it is currently defined (Bonacina, 1980; Fet & Sissom, 2000; Scherabon et al., in press). Capra (1939) separated E. germanus from Italy into four forms (A, B and C, as well as a “typical form”) based on the number of pectinal teeth (Dp) and the number of trichobothria on the ventral aspect of the pedipalp patella (Pv). Within Italy, the “typical form” of E. germanus (Pv = 5) was limited by Capra to the region of Trentino, Alto-Adige and Cadore; the ‘A-form’ (Pv = 6) to the Lombardian Alps; the ‘B-form’ (Pv = 5) was assigned to the Piemontese Alps and the ‘C-form’ (Pv = 5 or 6) to the Goriziano and the Karawanken Alps. Caporiacco (1950) confirmed all of Capra’s forms and formally described three new subspecies (E. g. alpha, E. g. beta, E. g. gamma) in addition to the nominotypical E. g. germanus. Caporiacco (1950) also described E. g. croaticus from Croatia. Finally, Valle et al. (1971) described E. g. marcuzzii from the Dolomites and Slovenia. Bonacina (1980) in his revision restricted E. germanus to four subspecies: E. g. germanus (= E. g. beta), E. g. alpha, E. g. marcuzzii and E. g. croaticus. E. g. gamma was treated as a subspecies of E. mingrelicus (see Scherabon et al. (in press) for a detailed taxonomic history). Bonacina (1980) synonymized the disjunct western (Piemonte) E. g. beta with the eastern E. g. germanus since both taxa shared character of Pv = 5. Furthermore, he suggested hybridization between the taxa E. g. alpha and E. g. germanus. Bonacina (1980) also carried out a thorough statistical study of trichobothrial numbers (on ventral and external surfaces of the pedipalp patella) for numerous populations in Piemonte and Lombardy, Italy. He postulated that the number of ventral patellar trichobothria is Pv = 5 for E. g. germanus and Pv = 6 in E. g. alpha. He also suggested a hybridogenic origin for populations in the Bergamascan Alps (Valle Brembana, north of Bergamo), because of intermediate forms (5-5 or 6-6) and a high number of asymmetric (5-6 or 6-5) individuals. However, our allozyme and DNA data do not confirm any assumptions of hybridogenic origin by Bonacina (1980), or by Kinzelbach (1975); see also Gantenbein er al. (1999a). Applying methods based on models of evolutionary change (pairwise distance methods / maximum likelihood) and the maximum parsimony criterion, in our study, two independent systems of molecular markers (allozymes and 16S mtDNA sequences) revealed an almost identical phylogenetic pattern. The phylogeny based on both mtDNA and allozyme data suggests a clear, distinct topology of two major clades (Figs 2-3). These clades are well supported statistically. They include parapatric population groups which are geographically separated by the river Adige (Etsch). The genetic distance which is found between these two groups is comparable to the genetic distance found between each of these clades and the outgroup species 856 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL E. gamma. Therefore, we propose to treat these two population groups as two species: Euscorpius germanus (C. L. Koch, 1837) sensu stricto (“eastern” clade) and Euscorpius alpha Caporiacco, 1950 stat. nov. (“western” clade). The deep divergence and parapatry of the “eastern” and “western” clades inferred by using independent genetic markers is not entirely consistent with geogra- phical ranges of the subspecies E. germanus germanus and E. g. alpha as shown by Caporiacco (1950) and Bonacina (1980). Nevertheless, type localities of these two taxa (as designated below) would fall well inside the ranges of two papapatric clades, thus making the existing taxonomic names applicable to the monophyletic clades. THE STATUS OF EUSCORPIUS GERMANUS BETA CAPORIACCO, E. G. CROATICUS CAPORIACCO, AND E. G. MARCUZZII VALLE ET AL. (a) Euscorpius germanus beta Caporiacco, 1950. Originally delineated by Capra (1939) as “Form B” from Val d’ Aosta, including the marginal, westernmost populations of “E. germanus” (sensu lato). It was formally described as a subspecies by Caporiacco (1950) and also limited to Piemonte populations. Bonacina (1980) synonymized E. g. beta with eastern E. g. germanus since both taxa shared the character Pv = 5. Other diagnostic characters listed by Caporiacco (1950) (i.e. meta- somal granulation and carination of chela) were considered too variable to be diagnostic. This synonymy, however, created a disjunct distribution for E. g. germa- nus sensu Bonacina (1980). Our molecular analysis shows that “E. g. beta” populations from Italy (Fon- tainemore) and bordering Switzerland (Gondo, Zwischbergental) occupy the most derived position in the “western clade”, or E. alpha. Both populations from this area for which allozyme and DNA data were available (i.e. Fontainemore and Gondo) grouped together and were supported by boostrap values in all analyses (Figs 2-3). Thus, we cannot confirm Bonacinais synonymy E. g. beta = E. g. germanus. On the other hand, assigning a separate taxonomic status to these two populations would create a paraphyletic subspecies “E. alpha beta”. Subsequently we would be required to treat other E. alpha subclades as monophyletic assemblages as well and assign at least three other new “subspecies” names. Thus, it seems reasonable not to retain E. g. beta as a valid taxon, but to place it into synonymy: Euscorpius alpha stat. nov., elevated from Euscorpius g. alpha Caporiacco, 1950 = Euscorpius g. beta Capo- riacco, 1950 syn. nov. (b) Euscorpius germanus croaticus Caporiacco, 1950. This taxon remained enigmatic since its description. It was mentioned but not revised by Bonacina (1980). We analysed the morphology of the only existing type specimen of this taxon (MZUF 5580, a male from Mali Halam, Velebit Mountains, Croatia; here designated as lectotype, see below). Its trichobothrial pattern on the pedipalps, i.e. number of ventral trichobothria on patella (Pv = 6) and position of trichobothria on the fixed finger, is identical with that of many populations of E. alpha (but not of E. germanus s. str.). The external face of the pedipalp patella in E. g. croaticus bears 22 tricho- bothria (Pe = 22) in the following serial arrangement: et = 5, est = 4, em = 3, esb = 2, PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 857 eb, = 4, eb = 4). However, the number ef = 5 is not found in other populations of E. germanus or E. alpha, which have et = 4 or even et = 3. A number of other mor- phological characters, first of all the very clear presence of carinae on the metasomal segments, shape of pedipalps and spination of legs, indicate that this form is not close to E. germanus and falls into the “species complex” of E. carpathicus (L., 1767). Similar forms have been observed by one of us (V.F.) from the Rhodope Mountains in Bulgaria. Reduction of trichobothrial numbers is not uncommon in E. carpathicus. In fact, reduction of the trichobothria in the series em from 4 to 3 in E. c. banaticus from Romania has been the reason for confusion (Vachon & Jaques, 1977) since this single character was considered to be diagnostic for E. germanus (sensu lato, including E. alpha, E. gamma and E. mingrelicus). A detailed study of E. carpathicus and related taxa is now being carried out by us (V. Fet, M. Soleglad, B. Gantenbein, in preparation). Pending the completion of this study, we treat E. germanus croaticus Caporiacco, 1950 as a form belonging to “E. carpathicus complex”, but not to E. germanus C. L. Koch. Its exact taxonomic status has to be determined. (c) Euscorpius germanus marcuzzü Valle, Berizzi, Bonino, Gorio, Gimmi- laro-Negri & Percassi, 1971. Marcuzzi & Fabris (1957) first recorded a form of E. germanus from the Dolomites (Italy) with 20 trichobothria (in contrast to the common 21) on the external face of the pedipalp patella (Pe = 20). Valle er al. (1971, p. 95-96) very briefly (one line!) described this subspecies from the “refugial massifs of the Venetian Pre-Alps (Italy) and from northern Slovenia’, without designating any type specimens (Valle’s syntypes of this subspecies are in the Museo Civico di Scienze Naturali "Enrico Caffi", Bergamo). The sole morphological character distinguishing this taxon from other subspecies is the presence of 3 trichobothria instead of 4, in the accessory basal series (eb, = 3) on the external face of the pedipalp. This character is unique and indeed accords with the general trend of trichobothrial number reduction in the subgenus Alpiscorpius (see Gantenbein et al. 1999a). Bonacina (1980) mentions a number of populations from northeastern Italy and Slovenia, some “pure” E. g. marcuzzii and others mixed (and “hybrid”, i.e. asymmetric eb =3-4 or 4-3) with E. g. germanus. Our molecular analysis of E. alpha from Italy presently includes assymetrical specimens (eba = 3-4 or 4-3) which were found to be rare at Bezzecca (only one find) but are much more common at Auronzo di Cadore, Belluno, Schluderns, Voltago, Vetriolo, Völs, and Crnice. None of the populations mentioned corresponds with a monophyletic, clearly divergent clade. The only Slovenian locality studied (Crnice) includes both symmetric “E. g. marcuzzii” (eb, = 3-3) and regular E. germanus speci- mens. However, further molecular analysis of all populations with eb, = 3 and a thorough analysis of the syntype series and of additional material are necessary to clarify the status of E. g. marcuzzii, which currently appears to be a taxon of dubious validity. 858 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL DESIGNATION OF TYPE SPECIMENS None of the taxa treated in here have a holotype or lectotype specimen. Types designated by C. L. Koch (1837) are lost. For the taxon described by Caporiacco (1950) only syntypes were designated; they are deposited in the collection of the Museo Zoologico "La Specola" dell'Università de Firenze, Florence, Italy (MZUF) (Bartolozzi et al., 1988). Here we designate the necessary type specimens for the following taxa: Euscorpius germanus (C. L. Koch, 1837) Originally described as Scorpius germanus C. L. Koch, 1837 (pp. 110-112, plate 108, figs 250-252) from “southern Tyrol (now Trentino-Alto Adige, Italy) and upper (= northern) Italy”. Neotype: d, Brixen (Bressanone), Trentino - Alto Adige, Italy, 9. 10. 98, coll. B. Gantenbein & I. Gantenbein, sample No. BG-109-07, deposited in Natur- historisches Museum Bern, Switzerland, accession number Scl. Trichobothrial for- mula: Pv = 5, Pe = 21 (et =4, est= 4, em = 3, esb= 2, eb,=4, eb =4). Euscorpius alpha Caporiacco, 1950 stat. nov. Originally described as Euscorpius germanus alpha Caporiacco, 1950: 211. Lectotype: 2 (MZUF 5569), Lago di Como near Varenna, Lombardy, Italy (August 1879, coll. Cantoni). Trichobothrial formula: Pv = 6, Pe = 21 (et = 4, est = 4, emi—34esb:—2.eba —4.eb — 4): Paralectotypes: 1 d& (MZUF 5571), 5 2 2(MZUF 5568, 5570-5574), from the type locality; 6 d 4,5 2 ?(MZUF 5575-5579), Varese, Lombardy, Italy (1879, coll. Cantoni); 1 2 (MZUF 5567), Monte Stelvio, Trentino - Alto Adige, Italy (August 1877, coll. P. Magretti). Euscorpius germanus beta Caporiacco, 1950: 211 Here considered as syn. nov. of Euscorpius alpha Caporiacco, 1950. Lectotype: d (MZUF 5588), Monte Massone (Cesara, Novara), Piemonte, Italy (12 August 1879, coll. C. Parona). Trichobothrial formula: Pv = 5, Pe = 21 (er = 4, est=4,em=3,esb=2,eb,=4,eb=4). Paralectotypes: 5 dd and 5 22 (MZUF 5589, 5590-5593), type locality; 1 d, 1 2 (MZUF 5584, 5585), Colle della Piccola Mologna, 2000 m, (Biella, Vercelli), Piemonte, Italy; 2 2 2 (MZUF), Lamorano, 1879, Piemonte, Italy. Euscorpius germanus croaticus Caporiacco, 1950: 215 Lectotype: d (MZUF 5580), Mali Halam, Velebit Mountains, Croatia (other specimens were listed in the original description but are absent in the MZUF collection). Probably belongs to Euscorpius carpathicus (L., 1767) “complex” (see above). Trichobothrial formula: Pv = 6, Pe = 22 (et = 5, est = 4, em = 3, esb = 2, eb, = 4,eb=4). PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 859 MORPHOLOGICAL VERSUS MOLECULAR DATA The classical morphological characters in the species of Euscorpius, such as the number of pectinal teeth and the number of trichobothria, are quantitative threshold characters similar the number of bristles in Drosophila (Futuyma, 1986). Such traits are expressed discontinuously at the phenotypic level, but are affected by a continuous distribution of some underlying trait. Capra (1939), Caporiacco (1950) and Bonacina (1980) have found geographic variation in these characters and also asymmetries which are very common. We confirm their observations that within the E. alpha clade the number of the ventral trichobothria of the patella (Pv) is about 6 in the region of the Bergamscan Alps (Valle Brembana) and changes to Pv = 5 in the western populations (Fontainemore and Gondo). The same pattern is found in the eastern populations at Molina di Ledro, Bezzecca (Lago di Garda) and Marling, Bad Salz (Trentino-Alto Adige). Here the subclades detected by using molecular markers (Figs. 2-3) correspond to the observed pattern of the Pv character. However, no hybridization or gene flow was observed between E. alpha and E. germanus clades. Bonacina (1980) gave a detailed morphological account of the distribution of Pv = 5-5, 6-6 or 5-6 individuals within the Bergamascan Alps. Ana- lysing the same populations (Fig. 1, box) we found no confirmation of his assumption of hybridisation between two distinct taxa; it appears that the described morphological variation occurs at the phenotypic level within a single genetically coherent species, E. alpha. Apparently character states Pv = 5 or 6 alone are not sufficient to define both parapatric species since E. germanus usually has Pv = 5 but in E. alpha this character varies from 5 to 6 with various degree of fixation. The same 1s true for the pectinal teeth number, Dp (see Results). Neither trichobothrial nor pectinal teeth scores give us a clear-cut delineation which would reflect the deep divergence of two clades revealed by molecular data and treated here as species-level taxa. In order to characterize these two clades morpho- logically, more complex characters should be searched for and analysed. Among these, the morphology of the hemispermatophore could possibly be applied as a species-level character set for Euscorpius as suggested by some previous authors (Kinzelbach, 1975; Bonacina, 1980; Scherabon 1987). PHYLOGENETIC IMPLICATIONS ON BIOGEOGRAPHY In their analysis of molecular phylogeny and historical biogeography of the genus Euscorpius, Gantenbein et al. (1999a) noticed the deep split between the endemic Alpine clade (subgenus Alpiscorpius Gantenbein, Fet, Largiadér & Scholl, 1999 which included traditional E. germanus) and the major Asia Minor-Trans- mediterranean lineage (subgenera Euscorpius Thorell, 1876 and Polytrichobothrius Birula, 1917). This led to the assumption that in the ancestors of modern E. germanus (sensu lato) ecological differentiation and adaptation to orophylic and mesophylic habitats (in contrast to xerophylic habitats occupied by E. carpathicus and especially E. italicus) could have been an ancient event. In other words, modern forms in- habiting the Alpine zone of Europe are not necessarily a result of speciations due to recent (Pleistocene) glacial events (Klicka & Zink, 1997) but these taxa may have 860 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL evolved in this area since the beginning of the Alpine orogenesis. This long time scale can explain the high level of genetic divergence observed within the subgenus Alpiscorpius, which we separate here into two species, Euscorpius (Alpiscorpius) germanus (C. L. Koch, 1837) and E. (A.) alpha Caporiacco, 1950. The period of divergence time between these two parapatric taxa can be estimated by using the genetic distance and by assuming a constant evolutionary rate through time. The calibration of a molecular clock for Mesobuthus gibbosus (Brullé, 1832) (Scorpiones, Buthidae) in the mainland Greece and Turkey and on several Aegean islands reveals an average rate of about 3% sequence divergence per Myr for the 16S rRNA gene (our data, unpublished). A comparable rate was reported for another scorpion species, Buthus occitanus (Amoreux, 1789) (Buthidae) across the Strait of Gibraltar (Gantenbein er al., 1999b). Applying this “scorpion clock” to the mtDNA sequence divergence between E. germanus and E. alpha (ca. 7%) reveals a separation time of about 2-3 million years. Such a time scale contradicts the Late Pleistocene Origin model (LPO), which is widely accepted today (see also Klicka & Zink, 1997). The deep split between the evolutionary lineages in E. germanus and E. alpha remains high even when genetic distances for superimposed substitutions are corrected by using the most appropiate model (TrN93 in this case), as proposed by Arbogast & Slowinski (1998). A similar deep split (> 1.6 Myrs) was uncovered by a recent allozyme survey of Alpine species of Glomeris (Diplopoda: Glomeridae) (Hoess & Scholl, 1999) which presumably have similar dispersal rates as Euscorpius species. Taberlet et al. (1998) identified several so-called “suture zones” in Europe, including the Alps, where different taxa meet after postglacial isolation. The situation for Euscorpius is similar but lacks hybridisation. No gene flow was detected between the two clades, which appear to be true parapatric species separated by the geographic divide of the Adige River valley. Hedin (1997) recently demonstrated for Nesticus spiders (Aranae, Nesticidae) in the Appalachian Mountains that mtDNA analysis allows to discover considerable genetic divergence, both between and within recognized morphological species; most of these probably predate the Pleistocene. The divergence between E. alpha and E. germanus clearly predates the Pleistocene glaciations, as it is the case for a number of other Alpine taxa (Taberlet er al., 1998). This confirms the hypothesis that speciation events mainly occurred during the Pliocene (Zink & Slowinski, 1995). Possible Pleistocenic refugia for these two species were the Bergamascan Alps for E. alpha and the Venetian Prealps for E. germanus. Evidence that these regions could have served as refugia for small terrestrial arthropods comes from a palyno- logical analysis (Kral, 1989) which indicates that a relatively mild climate prevailed during the Pleistocene. Other arguments for this interpretation are provided by the genetic data presented in this study. The genetic variability, expressed in the average heterozygosity and the mean number of alleles per locus, in E. alpha populations is relatively higher in the region around Bergamo than in regions near the edges of the refuge. Low levels of heterozygosity in E. germanus indicate possible genetic bottlenecks in the history of this species. Further investigations are required to test this hypothesis. PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 861 ACKNOWLEDGEMENTS B. G. thanks his wife Iris for her efforts and enthusiasm when collecting Euscorpius germanus and E. alpha. Dietmar Huber and Bernhard Scherabon kindly donated additional material of E. germanus and E. gamma from Austria, and Matjaz Kuntner provided specimens from Slovenia. We are grateful to Carlo Largiadèr for critical discussions and assistance in DNA data analyses. We thank Matt Braunwalder for his enthusiastic help and guidance in the issues related to the natural history of Euscorpius. Sarah Whitman Mascherini allowed the loan of Caporiacco’s specimens from MZUF. David Sissom and Michael Soleglad advised on taxonomic issues. The Karl-Bretscher Foundation Berne; award to A. S. supported this study. The comments of two anonymous reviewers greatly improved the final version of the manuscript. REFERENCES ARBOGAST, B. S. & SLOWINSKI, J. B. 1998. Pleistocene speciation and the mitochondrial DNA clock. Science 282 (5396): 1955. AYALA, F. J., POWELL, J. R., TRACEY, M. L., Mourao, C. A. & PEREZ-SALAS, S. 1972. Enzyme variability in the Drosophila willistoni group. IV. 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Also given are the identified 16S rDNA haplotypes. alpdh ark ddh gapdh aatl Nr. Sample Country N Sequence 100 95 104 100 98 93 101 100 100 78 88 96 100 104 Western clade = E. alpha stat. nov. Gondo CHO) “EalkGO 1.00 1.00 1.00 1.00 1.00 I 2 Fontainemore CH (10) Eal FO 1.00 1.00 1.00 1.00 1.00 3 Rancate CH (15) EalRA 1.00 1.00 1.00 1.00 1.00 4 Monte CH (11) 1.00 1.00 1.00 1.00 1.00 5 Fornace CH (10) 1.00 0.05 0.95 1.00 1.00 1.00 6 Pontide I (5) 1.00 1.00 1.00 1.00 0.10 0.90 7 Sottoponte CH (13) Eal SO 1.00 1.00 1.00 1.00 1.00 8 San Carlo CH (12) 1.00 1.00 1.00 1.00 1.00 9 Brembilla I (5) 1.00 1.00 1.00 1.00 1.00 10 Gerosa I (2) 1.00 1.00 1.00 1.00 1.00 11 Peghera I (2) 1.00 1.00 1.00 1.00 1.00 12 San Pellegrino I (4) Eal SP 1.00 1.00 1.00 1.00 025M07S 13 San Giovanni Bianco I (11) Eat SG 1.00 1.00 1.00 1.00 0.09 0.05 0.82 0.05 14 Camerata Comello I (1) 1.00 1.00 1.00 1.00 1.00 15 Olmo al Brembo I (10) Eal OL 1.00 1.00 1.00 1.00 1.00 16 Piazzatorre er) 1.00 1.00 1.00 1.00 1.00 17 Mezzoldo I (7) 1.00 1.00 1.00 1.00 1.00 18 Isola di Fondra I (13) 1.00 1.00 1.00 1.00 1.00 19 Carona I (13) EalCA 1.00 1.00 1.00 1.00 1.00 20 Valleve I (1) 1.00 1.00 1.00 1.00 1.00 21 Selvino I (5) 1.00 1.00 1.00 1.00 1.00 22 Membro I (3) 1.00 1.00 1.00 1.00 0.17 0.83 23 Tavemola I (4) EalTa 1.00 1.00 1.00 1.00 1.00 24 Vigolo I (3) 1.00 1.00 1.00 1.00 1.00 25 Bezzeca I (5) 1.00 1.00 1.00 1.00 1.00 26 Molina di Ledro I (10) Eal ML 1.00 1.00 1.00 1.00 0.75 0.25 27 Marling I (10) EalMA 1.00 1.00 1.00 1.00 0.30 0.70 28 Bad Salz I (1) 1.00 1.00 1.00 1.00 1.00 Eastern clade = E. germanus 29 Sta Maria CH (22) Ege SM 1.00 1.00 1.00 1.00 1.00 30 Lichtenberg I (5) 1.00 1.00 1.00 1.00 1.00 31 Schluderns I (10) Ege SH 1.00 1.00 1.00 1.00 0.55 0.35 0.10 32 Schlanderns I (1) 1.00 1.00 1.00 1.00 1.00 33 Verdins 1240) 0.05 0.95 1.00 1.00 1.00 1.00 34 Brixen I (4) 1.00 1.00 1.00 1.00 1.00 35 Vols I (10) Ege VO 0.40 0.60 1.00 1.00 1.00 0.95 0.05 36 Bremer I (7) 0.21 0.79 1.00 1.00 1.00 1.00 37 Veltiolo I (9) Ege VE 1.00 0.94 0.06 1.00 1.00 1.00 38 Belluno I (4) 1.00 1.00 1.00 1.00 1.00 39 Voltago I (5) 1.00 1.00 1.00 1.00 0.70 0.30 40 San Tomaso I (3) 1.00 1.00 1.00 1.00 0.67 0.33 41 Mezzocanale I (2) Ege ME 1.00 1.00 1.00 1.00 0.50 0.50 42 Borca di Cadore I (10) Ege BO 1.00 1.00 1.00 1.00 0.65 0.35 43 Auronzo di Cadore I (6) 1.00 1.00 1.00 1.00 1.00 44 Starkenbach A (4) Ege ST 1.00 1.00 1.00 1.00 1.00 45 Tarrenz A (2) EgeTZ 1.00 1.00 1.00 1.00 1 46 Kranzach A (4) Ege KR 1.00 1.00 1.00 1.00 1.00 47 Dellach A (8) Ege DE 1.00 1.00 1.00 1.00 1.00 48 Oberdrauburg A (5) Ege OB 1.00 1.00 1.00 1.00 1.00 49 Dobratsch A (4 1.00 1.00 1.00 1.00 1.00 50 Schütt A (6) 1.00 1.00 1.00 1.00 1.00 51 Federaun A (4) 1.00 1.00 1.00 1.00 1.00 52 Cmice SLO (8) 1.00 1.00 1.00 1.00 1.00 E. gamma 53 Koschuta A (6) EgaKO 1.00 1.00 1.00 1.00 1.00 E. flavicaudis Lauris ™ a = = & > © (©) | 8 866 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL APPENDIX I (2) aat2 gtdh hk idh1 idh2 mdh1 Nr. Sample 88 100 107 113 90 95 10071007 10777947 “95100 87S 937710076) 87 89 100 Western clade = E. alpha stat. nov. 1 Gondo 1.00 1.00 1.00 1.00 1.00 1.00 2 Fontainemore 1.00 1.00 1.00 1.00 1.00 1.00 3 Rancate 1.00 1.00 1.00 1.00 1.00 1.00 4 Monte 1.00 1.00 1.00 1.00 1.00 1.00 5 Fornace 1.00 1.00 1.00 1.00 1.00 1.00 6 Pontide 1.00 1.00 1.00 1.00 1.00 0.80 0.20 7 Sottoponte 1.00 1.00 1.00 1.00 1.00 1.00 8 San Carlo 1.00 1.00 1.00 1.00 1.00 1.00 9 Brembilla 1.00 1.00 1.00 1.00 1.00 0.90 0.10 10 Gerosa 1.00 1.00 1.00 1.00 1.00 1.00 11 Peghera 1.00 1.00 1.00 1.00 1.00 1.00 12 San Pellegrino 0.25 0.75 1.00 1.00 1.00 1.00 1.00 13 San Giovanni Bianco 1.00 1.00 1.00 1.00 1.00 1.00 14 Camerata Cornello 1.00 1.00 1.00 1.00 1.00 1.00 15 Olmo al Brembo 1.00 1.00 1.00 1.00 1.00 1.00 16 Piazzatorre 1.00 1.00 1.00 1.00 1.00 1.00 17 Mezzoldo 1.00 1.00 1.00 1.00 1.00 1.00 18 Isola di Fondra 0.08 0.92 1.00 0.92 0.08 1.00 1.00 0.04 0.92 0.04 19 Carona 1.00 1.00 1.00 1.00 1.00 1.00 20 Valleve 1.00 1.00 1.00 1.00 1.00 1.00 21 Selvino 1.00 1.00 1.00 1.00 1.00 0.80 0.20 22 Nembro 1.00 1.00 1.00 1.00 1.00 0.83 0.17 23 Tavernola 1.00 1.00 1.00 1.00 1.00 0.12 0.88 24 Vigolo 0.33 0.67 1.00 1.00 1.00 1.00 1.00 25 Bezzecca 1.00 1.00 1.00 1.00 1.00 1.00 26 Molina di Ledro 1.00 1.00 1.00 1.00 1.00 1.00 27 Marling 1.00 1.00 0.95 0.05 1.00 1.00 1.00 28 Bad Salz 1.00 1.00 1.00 1.00 1.00 1.00 Eastern clade = E. germanus 29 Sta Maria 1.00 1.00 1.00 1.00 1.00 1.00 30 Lichtenberg 1.00 1.00 1.00 1.00 1.00 1.00 31 Schluderns 1.00 1.00 1.00 1.00 1.00 1.00 32 Schlanderns 1.00 1.00 1.00 1.00 1.00 1.00 33 Verdins 1.00 1.00 1.00 1.00 1.00 1.00 34 Brixen 1.00 1.00 1.00 1.00 1.00 1.00 35 Vols 1.00 1.00 1.00 1.00 1.00 1.00 36 Bremer 1.00 1.00 1.00 1.00 1.00 1.00 37 Vetriolo 1.00 1.00 1.00 1.00 1.00 1.00 38 Belluno 1.00 1.00 1.00 1.00 1.00 1.00 39 Voltago 1.00 1.00 1.00 1.00 1.00 1.00 40 San Romaso 1.00 1.00 1.00 1.00 1.00 1.00 41 Mezzocanale 1.00 1.00 1.00 1.00 1.00 1.00 42 Borca di Cadore 1.00 1.00 1.00 1.00 1.00 0.20 0.10 0.70 43 Auronzo di Cadore 1.00 1.00 1.00 1.00 1.00 0.33 0.67 44 Starkenbach 1.00 1.00 1.00 1.00 1.00 1.00 45 Tarrenz 1.00 1.00 1.00 1.00 1.00 1.00 46 Kranzach 1.00 1.00 1.00 1.00 1.00 1.00 47 Dellach 1.00 1.00 1.00 1.00 1.00 1.00 48 Oberdrauburg 1.00 1.00 1.00 1.00 1.00 1.00 49 Dobratsch 1.00 1.00 1.00 1.00 1.00 1.00 50 Schütt 1.00 1.00 1.00 1.00 1.00 1.00 51 Federaun 1.00 1.00 1.00 1.00 1.00 1.00 52 Crnice 1.00 1.00 1.00 1.00 1.00 1.00 Outgroup species E. gamma 53 Koschuta 1.00 1.00 1.00 1.00 1.00 1.00 E. flavicaudis 1.00 1.00 Lauris 0.03 0.97 1.00 1.00 0.09 0.91 PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) 867 APPENDIX I (3) mdh2 mpi pep Nr. Sample LOOMMOSi£ TOON LOM MO7 S110) 121187 125 0130) 3578 87 94 98 100 104 107 Western clade = E. alpha stat. nov. 1 Gondo 1.00 0.17 0.83 1.00 2 Fontainemore 1.00 1.00 1.00 3 Rancate 1.00 0.54 0.27 0.12 0.08 1.00 4 Monte 1.00 0.14 0.77 0.09 1.00 5 Fornace 1.00 0.30 0.10 0.60 1.00 6 Pontide 1.00 0.70 0.10 0.20 0.40 0.60 7 Sottoponte 1.00 0.92 0.08 1.00 8 San Carlo 1.00 0.14 0.86 0.95 0.05 9 Brembilla 1.00 0.25 0.50 0.13 0.13 0.90 0.10 10 Gerosa 1.00 0.75 0.25 1.00 11 Peghera 1.00 0.25 0.25 0.25 0.25 0.75 0.25 12 San Pellegrino 1.00 0.13 0.13 0.13 0.64 0.50 0.25 0.25 13 San Giovanni Bianco 1.00 0.05 0.18 0.59 0.18 0.59 0.41 14 Camerata Cornello 1.00 1.00 1.00 15 Olimo al Breembo 1.00 0.05 0.45 0.35 0.15 0.95 0.05 16 Piazzatorre 1.00 0.12 0.39 0.31 0.12 0.08 0.89 0.12 17 Mezzoldo 1.00 0.64 0.14 0.14 0.07 0.83 0.17 18 Isola di Fondra 1.00 0.08 0.58 0.12 0.23 0.92 0.08 19 Carona 1.00 0.65 0.19 0.15 1.00 20 Valleve 1.00 1.00 1.00 2] Selvino 1.00 0.10 0.70 0.20 1.00 22 Membro 1.00 0.50 0.50 1.00 23 Tavernola 1.00 0.50 0.50 1.00 24 Vigolo 1.00 0.50 0.50 1.00 25 Bezzecca 0.90 0.10 0.10 0.40 0.50 0.20 0.80 26 Molina di Ledro 1.00 0.10 0.35 0.45 0.10 0.10 0.90 27 Marling 1.00 0.10 0.30 0.10 0.50 1.00 28 Bad Salz 1.00 1.00 1.00 Eastern clade = E. germanus 29 Sta Maria 1.00 1.00 1.00 30 Lichtenberg 1.00 1.00 1.00 31 Schluderns 1.00 1.00 1.00 32 Schlanderns 1.00 1.00 1.00 33 Verdins 1.00 1.00 1.00 34 Brixen 1.00 1.00 1.00 35 Vols 1.00 1.00 1.00 36 Bremer 1.00 1.00 1.00 37 Vetriolo 1.00 1.00 1.00 38 Belluno 1.00 1.00 0.50 0.50 39 Voltago 1.00 1.00 0.60 0.40 40 San Tomaso 1.00 1.00 0.17 0.83 41 Mezzocanale 1.00 1.00 0.25 0.75 42 Borca di Cadore 1.00 1.00 0.05 0.45 0.50 43 Auronzo di Cadore 1.00 1.00 0.83 0.17 44 Starkenbach 1.00 1.00 0.38 0.62 45 Tarrenz 1.00 1.00 1.00 46 Kranzach 1.00 1.00 1.00 47 Dellach 1.00 1.00 0.31 0.69 48 Oberdrauburg 1.00 1.00 0.80 0.20 49 Dobratsch 1.00 1.00 0.88 0.12 50 Schütt 1.00 1.00 0.70 0.30 51 Fereraun 1.00 1.00 0.75 0.25 52 Crnice 1.00 0.06 0.94 0.42 0.58 Outgroup species E. gamma 53 Koschuta 1.00 1.00 0.17 0.83 E. flavicaudis Lauris 1.00 0.10 0.87 868 B. GANTENBEIN, V. FET, M. BARKER & A. SCHOLL APPENDIX I (4) 6-pgd pgi pgm pk Nr. Sample 88 93 98 100 104 107 80 88 93 94 100 80 91 94 98 100 98 100 101 Western clade = E. alpha stat. nov. 1 Gondo 1.00 1.00 0.83 0.06 0.11 1.00 2 Fontainemore 1.00 1.00 1.00 1.00 3 Rancate 1.00 1.00 0.57 0.43 1.00 4 Monte 1.00 1.00 0.73 0.27 1.00 5 Fornace 1.00 1.00 0.70 0.30 1.00 6 Pontide 1.00 0.10 0.90 0.40 0.30 0.30 1.00 7 Sottoponte 1.00 0.23 0.77 0.85 0.15 1.00 8 San Carlo 1.00 0.62 0.38 0.75 0.25 1.00 9 Brembilla 1.00 0.10 0.90 0.38 0.13 0.50 1.00 10 Gerosa 1.00 0.25 0.75 0.75 0.25 1.00 11 Peghera 1.00 0.25 0.75 0.50 0.50 1.00 12 San Pellegrino 1.00 0.50 0.50 0.50 0.25 0.25 1.00 13 San Giovanni Bianco 0.91 0.09 0.09 0.91 0.05 0.96 1.00 14 Camerata Cornello 1.00 1.00 1.00 1.00 15 Olmo al Brembo 1.00 0.15 0.85 0.75 0.25 1.00 16 Piazzatorre 1.00 0.08 0.92 0.65 0.35 1.00 17 Mezzoldo 1.00 1.00 0.71 0.29 1.00 18 Isola di Fondra 1.00 0.73 0.27 0.08 0.92 1.00 19 Carona 1.00 1.00 0.13 0.88 1.00 20 Valleve 1.00 1.00 1.00 21 Selvino 1.00 0.10 0.90 0.13 0.13 0:75 1.00 22 Nembro 1.00 0.17 0.83 0.17 0.33 0.50 1.00 23 Tavernola 1.00 1.00 0.50 0.50 1.00 24 Vigolo 1.00 1.00 0.50 0.50 | 1.00 25 Bezzecca 0.90 0.10 1.00 0.10 0.50 0.20 0.20 | 1.00 26 Molina di Ledro 1.00 1.00 0.25 0.30 0.45 | 1.00 27 Marling 1.00 0.05 0.95 0.61 0.22 0.17 1.00 28 Bad Salz 1.00 1.00 1.00 1.00 Eastern clade = E. germanus 29 Sta Maria 1.00 1.00 0.80 0.21 1.00 30 Lichtenberg 1.00 1.00 0.40 0.60 1.00 31 Schluderns 1.00 1.00 0.93 0.07 1.00 32 Schlanderns 1.00 1.00 1.00 1.00 33 verdins 1.00 1.00 1.00 1.00 34 Brixen 1.00 1.00 0.75 0.13 0.13 1.00 35 Vols 1.00 1.00 0.70 0.20 0.10 1.00 36 Bremer 1.00 1.00 0.86 0.14 1.00 37 Vetriolo 1.00 1.00 0.89 0.11 1.00 38 Belluno 1.00 1.00 1.00 1.00 39 Voltago 1.00 1.00 0.40 0.60 1.00 40 San Tomaso 1.00 1.00 1.00 1.00 41 Mezzocanale 1.00 1.00 1.00 1.00 42 Borca di Cadore 1.00 1.00 0.85 0.15 1.00 43 Auronzo di Cadore 1.00 1.00 0.08 0.92 1.00 44 Starkenbach 1.00 1.00 0.75 0.25 1.00 45 Tarrenz 1.00 1.00 1.00 1.00 46 Kranzach 1.00 1.00 0.50 0.50 1.00 47 Dellach 1.00 1.00 0.06 0.94 1.00 48 Oberdrauburg 1.00 1.00 0.10 0.30 0.40 0.20 1.00 49 Dobratsch 1.00 0.13 0.88 0.25 0.75 1.00 50 Schütt 1.00 1.00 0.33 0.25 0.42 1.00 51 Federaun 1.00 1.00 0.75 0.25 1.00 52 Crnice 1.00 1.00 0.56 0.44 1.00 Outgroup species E. gamma 53 Koschuta 1.00 1.00 1.00 1.00 E. flavicaudis 1.00 1.00 1.00 Lauris 0.35 0.65 PARAPATRIC ALPINE SCORPIONS (EUSCORPIIDAE) APPENDIX II Polymorphic sites (indels, parsimony informative sites, transitions (ti), and transversions (tv)) in the 16S rRNA gene sequences analysed. Abbreviations for the haplotypes are explained in 869 appendix I. position lel eas ee 2 2 222222222722222222227222222333333333 position 33378012 2223344455 5555666890 1111122222 2333346677 778999022233555 position 312352760 2381812923 4569056598 0123723467 9023471901 232246145702029 Indel ===p=====,;==========/=-=======\=====p==== 1==2===D22=]==22=222=222222=== Informative====FF=F= ==FFFF===F F=FFFFFFFF FF==F==F== ==FFFF==F= FFFF=F=F======= Ti ===NNNNN =NNNNN=NNN NNNNN=NNNN NN=NN==N=N NNNNNN==N= NNNNNNNNN=NN=NN Tv =VV==V=== V==V=VV==V V====VV=VV V=V===VVV= V=VV===V=V =======V=V==V=V EalFO TTATATTGT TTACGTTGAT CCTCATATAG GGTTA*TGTA ATAGAAGAAT AACAAGAGGTTTCGA EalGO --------- ---------- ---------- ---------- ---------- --------------- mals 9 9 Se ee em Hee ae Se een Gen == D — G EFalSG = --------- ---------- TT ---~------- -- Greco leccese = AT EalRA Sal Seema TR eS ees ae == CM PRIGN - INR RE eo en CN enne Cole Geena Bao nn m AR EEE CEE En A-- wenige) | Ne re ee PESTE te EE GE Ce CE meg ee Be) men TEST An aa sestese CC ee EUM NT RDS me IN ee a Ca (© het liblllotto BEINEN E er ee A at Nees ee Ben @S en FGeOB RE en GA "AG -CIT-C-G-C-CT A=--G--A-—. =-GAG===—= GGT--A-T------- EgeDE CA AC CUOCERE GGT--A-T------- FES TR GAME EE AG CAT CC G-CF A==-G==A-= CAC GGT--A-T------- EgerOMM === PREME RC CRTC CCE CINA GR NE CAC nen GGT--A-T------- EgeSM ----- Ne See Nero, Meter een er eher GGT--A-T------- EgeVE -----G-A- ---TAG---- T-C-G-G-GT A---G--A-- --GAG----- GGT--A-T------- EgeSH GA = TAG = === TC =G-GaGT Aj==G=—A==) —-GAG====— CGT A == == EG EVO === — Gah TAG une Ge e Tan EA CAGE GGT--A-T------- EGEME ee G=A=e---TAG==—— “T=C=G=G-CTA---G--A-—= =ZGAGE-- GET AT seen FockR ee GERA = TAG. UT C-G-C-CT A---G--A=—) =-GAG=*==— CET An === nea iL esa GERME TAG====—T=C-C-G=CT A-=-G=-A-===CGAG-= === GGT--A-T------- EgeCR ----GG-A- ----AGG--A T-C-G--C-T AA-----A-- -C-AG---G- GGT--A-T------- EC aK OM IVe DENMOSCICSSACANAR---A-C-TWan=-ccoh=- CC Kern CAT === Ef1LA CGC-GAC-C -CGAA---G- A----AT--T TAACGTGTAG T-TT-G-T-A G-T--A---GCC--- felpa to REVUE SUISSE DE ZOOLOGIE 107 (4): 871-882; décembre 2000 A new genus of Prionoglarididae from a Namibian cave (Insecta: Psocoptera) Charles LIENHARD Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. E-mail: charles.lienhard@ mhn.ville-ge.ch A new genus of Prionoglarididae from a Namibian cave (Insecta: Psocoptera). - Sensitibilla strinatii gen. n., sp. n. is described and illus- trated based on three females and several nymphs collected in Arnhem Cave (124 km SE of Windhoek). The new genus is closely related to the North American genus Speleketor Gurney. The description of its type species, Speleketor flocki Gurney, is complemented and the systematic position of both genera within the family Prionoglarididae is discussed. The presence of trichobothria on legs, unique in Psocoptera, is considered as a striking synapomorphy of Speleketor and Sensitibilla. Key-words: Psocoptera - Prionoglarididae - new genus - new species - trichobothria - cave fauna - Namibia. INTRODUCTION The family Prionoglarididae (suborder: Trogiomorpha, family-group: Psoc- athropetae) groups some of the most astonishing psocids currently known. It contains two genera, Prionoglaris Enderlein, 1909 and Speleketor Gurney, 1943, each comprising 3 species which have been revised by Lienhard (1988) and Mockford (1984) respectively. The Palaearctic genus Prionoglaris is characterized by the absence of a bulging postclypeus and the corresponding reduction of hypopharyngeal structures (which are present in all other psocids and implicated in the active uptake of water vapour from the atmosphere, cf. Lienhard, 1988: 104) and by the strong metamorphosis of mouthparts during the last moult, also unique in the order Pso- coptera: adults nearly lack laciniae (only a microscopical remnant is visible at high magnification), while nymphs have normal laciniae; adults have strongly modified sickle-shaped mandibles, while these organs are of the normal chewing-type in nymphs (cf. Lienhard, 1988, 1998). In the Nearctic genus Speleketor only a slight tendency to such a metamorphosis of mouthparts can be observed (Mockford, 1984), postclypeus and hypopharynx are not modified and exhibit the characters which are typical for the suborder Trogiomorpha. Both genera have very similar wing venation characterized by a strongly developed, arched basal segment of sc, joining rl close to the base of the pterostigma. The very particular morphology of male genitalia, unique in Psocoptera, can also be interpreted as a synapomorphy of these genera. Manuscript accepted 23.06.2000 872 CHARLES LIENHARD In 1999 the Swiss biospeleologist Pierre Strinati discovered the first African prionoglaridid, a very distinct Namibian species which will be described in the following. It is much more closely related to Speleketor than to Prionoglaris. One of the most striking diagnostic characters is the presence of trichobothria on all tibiae and on hindtarsus, while in Speleketor such sensilla are only present on femora and on mid- and hindtrochanter. This feature and the very characteristic development of female genitalia warrant the erection of a new genus for this species, in spite of its close general resemblance to Speleketor. It would have been of particular interest to compare also the male genitalia, but unfortunately the male of the new species remains unknown. Due to the kindness of my colleague Edward Mockford I have been able to examine a paratype female of Speleketor flocki Gurney, 1943, the type species of Speleketor. In the following comparative discussions some complements to the description of this species are made, enabling a better comparison of these closely related genera. The following abbreviations are used in the descriptions: P1-P4 = segments of maxillary palp; fl, f2, f3, ...= antennal flagellomeres; tl, 12, 13 = tarsomeres of hindtarsus; c = costal vein; sc = subcostal vein; rl = first branch of radial vein; rs = radial sector; m = medial vein; pcu = postcubital vein. Sensitibilla gen. n. Diagnosis. General habitus very similar to Speleketor (cf. Gurney, 1943: fig. 3). Differing form Speleketor by the following characters. Mandibles with slender incisor region lacking subapical tooth in adults (fig. 9), this region broader and with well developed subapical tooth in nymphs (fig. 13); chewing sculpture on molar region very weakly developed or absent in adults and nymphs. Distal segment of maxillary palp in apical half with a small subglobular sensillum on outer margin (figs 7, 8). Forefemur (fig. 17) lacking the row of articulated spines which is present in Speleketor (fig. 22). Legs with the following pattern of trichobothria on dorsal (outer) margin: 2 on foretibia (fig. 17) and midtibia (fig. 18) in adults and nymphs (one of them in proximal half, the other in distal half); 2 also on hind tibia in nymphs (fig. 21), but only the distal one present in adults (fig. 19); 1 on second article of hindtarsus in adults and nymphs (figs 19-21), this article is distinctly thickened at the place where the trichobothrium is located (fig. 20). Female paraproct concave in ventral half, this region densely setose, of about circular shape (fig. 29, the setose ventral part is transversally folded in slide mounted terminalia). Ovipositor valvulae (= third valvulae, fig. 28) with a long distal process, bearing several spinelike setae. Ovi- positor valvulae basally articulated to their abdominal tergite on outer side and fused by a sclerified zone to the subgenital plate medially. Apical lobe of subgenital plate free but ventrally almost completely covered by the valvulae. Subgenital plate and main lobe of the ovipositor valvulae sparsely pilose (fig. 28). Spermapore (fig. 25) with a caplike structure and a longitudinal sclerite near the opening of the very long, narrow spermathecal duct. Distal part of the spermathecal duct spirally curled. A NEW GENUS OF PRIONOGLARIDIDAE 873 Spermatheca (fig. 25) thin-walled, its wall with irregular rows of small circular pores (fig. 26), near origin of duct a slightly sclerified oval region bearing 2 strong denticles directed to the inner side of the spermathecal sac. Spermatophore simple, more or less pear-shaped (fig. 25). Type species. S. strinatii sp. n. Etymology. The name is of feminine gender and alludes to the presence of trichobothria on tibiae [sensilis (lat.) = sensible]. Discussion. A detailed revised diagnosis of Prionoglaris has been published by Lienhard (1988, 1998), the diagnosis of Speleketor is given by Mockford (1984, 1993). One of the most striking diagnostic characters mentioned above is the presence of a characteristic pattern of trichobothria on legs. In other Psocoptera, trichobothria only occur on sense cushions of paraprocts. The genera Speleketor and Sensitibilla are the only psocids, so far as I know, where such sensilla are present on legs. In Priono- glaris the general pilosity of legs is short and uniform, similar to that in the two other genera of the family, but trichobothria are lacking. In Speleketor, the presence of trichobothria on ventral margin of femora has already been mentioned by Gurney (1943: 198), and Mockford (1984) considers the presence of two trichobothria on forefemur as a generic character. The reexamination of a female paratype of S. flocki showed the following pattern of trichobothria, which probably can be considered as diagnostic for the genus Speleketor (figs 22-24): 2 on forefemur; 1 on midfemur and 1 on midtrochanter; 1 on hindfemur and 1 on hindtrochanter; all trichobothria are situated on the ventral (inner) margin of the correponding segment. In Sensitibilla the trichobothria are located dorsally on tibiae and hindtarsus. Female terminalia have been illustrated for Speleketor by Gurney (1943: figs 24-26) and Mockford (1984: figs 29-32). The only pair of well developed ovipositor valvulae (= third valvulae) are large rounded flaps bearing numerous slender setae on most of their margin and 2-3 stout setae close together in mediodistal region. They are not fused to the subgenital plate medially and no distal process is present. In Speleketor the paraproct is of normal shape, its ventral half is not concave as in the new genus and the paraproctal setal field is subdivided in a transversal median area and a roughly circular ventral area, both areas are separated by a large zone without hairs (cf. Gurney, 1943: fig. 26). Nothing is known about the spermatheca of Speleketor flocki, which is not present in the slide mounted genitalia of the paratype female examined by me. The spermatheca of the allotype female of Speleketor irwini Mockford can be characterized as follows (Mockford, in litt. 2000): the relatively short, straight duct is wide throughout, with a well-sclerotized cap but with no longitudinal sclerite at the spermapore end; the sac is thin-walled with no ornamen- tation; within the sac are three vague bodies which are probably spermatophores. Female genitalia of Prionoglaris have been illustrated by Lienhard (1988, 1998). The subgenital plate is simple, with a broadly rounded somewhat sclerified distal margin. The ovipositor valvulae are not fused to the subgenital plate medially and no distal process is present; they are similar in shape to those of Speleketor but densely pilose throughout, with some stout setae near apical margin. The paraproct of Prionoglaris is simple, pilose and bears some stout setae resembling those on ovi- 874 CHARLES LIENHARD positor valvulae. The spermatheca of Prionoglaris has a narrow, straight duct of medium length, and there are no ornamentations on the thin-walled sac; spermapore without sclerotized cap but with a longitudinal sclerite; spermatophore not yet observed. The characters of trichobothrial pattern and female genitalia warrant the separation of Speleketor and Sensitibilla. Some other useful diagnostic characters are the presence of a row of small spines on anterior face of forefemur in Speleketor (fig. 22) (only a row of normal short hairs at the same place in Sensitibilla), the pre- sence of a small flattened double sensillum in apical half of P4 in Speleketor (fig. 15) (unique subglobular sensillum in Sensitibilla) and the presence, on mandibles, of a small but distinct subapical tooth and a weakly but distinctly sculptured chewing plate in Speleketor (fig. 16) (no subapical tooth and no chewing sculpture in Sensitibilla). Sensitibilla strinatii sp. n. MATERIAL Holotype 2. NAMIBIA: Arnhem Cave (124 km SE of Windhoek), 1550m, 21 October 1999, leg. P. Strinati. Paratypes: 29 and 7 nymphs (different stages), same collecting data. The specimens were collected at the end of a large gallery going down from the entrance, at about 120 m from the entrance, in a completely dark zone (the gallery being straight, the opening of the entrance is visible from the collecting place), under stones on dusty and sandy soil (the psocids were sitting on the underside of the stones, not on the soil itself). Air temperature at this place: 21°C. At the same place, some pseudoscorpions and beetles were also collected. The cave is inhabited by many bats and guano deposits are important. The type series is deposited in the Psocoptera collection of the Muséum d'histoire naturelle, Geneva, Switzerland: No. 7435 (© holotype, mounted on two slides), 7436 (9 paratype, in alcohol), 7437 (© paratype, head mounted on slide, other parts in alcohol), 7438- 7441 (4 nymphs paratypes, each mounted on a slide) and, without collection numbers, 3 nymphs paratypes, in alcohol. ETYMOLOGY The species is dedicated to its collector, Pierre Strinati, in acknowledgement of his tireless efforts in collecting psocids in caves all over the world. DESCRIPTION (2, with some indications on nymphal morphology) Coloration. Head (fig. 5) marked with some light brown zones of cuticular pigmentation, a small and sometimes very indistinct crescent-like brown spot on each side near antennal socket. Compound eyes black, ocelli each with a dark brown pig- ment crescent on the inner side. Antennae and legs grey-brown, thorax brown, abdomen whitish with some red-brown hypodermal pigment on several tergites, arranged in irregular transversal bands. Wings clear, unmarked, pterostigma only very slightly opaque. Morphology. Head shape in frontal view as in Speleketor flocki (cf. Gurney, 1943: fig. 1), compound eyes relatively small and very much in lateral position, close to antennal sockets. Vertical suture well visible, frontal suture absent, ocelli well developed (fig. 5). Antennae longer than body, one antenna of holotype damaged, the other apparently intact (i.e. with last flagellomere regularly rounded apically), with 13 A NEW GENUS OF PRIONOGLARIDIDAE 875 articles. All flagellomeres very thin and densely annulated (sculpture), fl somewhat curved, f2-f11 straight. Lacinia with reduced apical denticles in adults (fig. 11), with three apical tines in nymphs (fig. 12). Maxillary palp (fig. 7) very long and slender, P2-P4 with annulated sculpture, general pilosity relatively dense, consisting of ordinary hairs of about the same length as the diameter of the palpal segments (not figured in fig. 7), 2 short and slightly thickend setae on P1, | small relatively weakly differentiated spur sensillum subbasally on P2, a very characteristic subglobular marginal sensillum in apical half of P4, slightly trilobate at apex (fig. 8). Mandibles as described in generic diagnosis (adult: fig. 9, nymph: fig. 13); molar lobe of right mandible subbasally with a well developed and slightly caudally directed fringed lobe. Cibarial sclerite of hypopharynx, epipharyngeal sclerite and oval lingual scle- rites well developed, the latter with a distinct hexagonal sculpture, hypopharyngeal filaments separate (fig. 6), hypopharyngeal brush also well developed (not figured in fig. 6). Labial palp 2-segmented (fig. 27), first segment with a short, thickened seta on outer margin, second segment with a slightly sclerified tubercle near distal margin and 3 small thin-walled subapical sensilla. Labrum (fig. 30) with 4 spiniform marginal setae on each side, a row of 6 slender setae in the semicircular hyaline area in the middle of the anterior margin and a row of 5 small setiform marginal sensilla situated in a membraneous fold. Legs with uniform short general pilosity (e.g. fig. 20, these hairs are not figured in figs 17-19). Forefemur with a longitudinal row of normal short hairs on anterior face (at the same place where in Speleketor there is a row of articulated spines, cf. fig. 22). Trichobothria pattern of legs as described in generic diagnosis (figs 17-21). Pretarsal claws symmetrical in nymphs and adults, lacking basal process or basal seta, with one preapical tooth in adults (fig. 10), in nymphs with a small additional tooth or short slender process just distally to it (fig. 14). Hindtibia with two ventral marginal spines in distal half and, on apex, 4 large ventral spines and 2 smaller dorsal ones. Hindtarsus with 2 apical spines and 5 plantar spines on fl and | apical spine on #2. Same number of apical and plantar spines on tibia and tarsus of foreleg and midleg, the basal 2 or 3 plantar spines of foretarsus weakly differentiated, no ventral marginal spines in apical half of foretibia and midtibia. All leg spines with broadly rounded apex (fig. 20) (in Speleketor flocki the leg spines are pointed apically, cf. fig. 24). Midcoxa with a hyaline tubercle on ventral side. Pearman's organ of hindcoxa only represented by a mirror, coxal rasp not differentiated. Forewing venation as in fig.1, distal segment of sc nearly perpendicular to c (much more inclined towards wing apex in Speleketor flocki), in one of the forewings of the holotype only anterior half of the crossvein between base of pterostigma and rs developed, wing margin and veins with very loose microscopical pilosity (fig. 2), pcu bald. Hindwing relatively small in comparison to forewing. Hindwing venation as in figs 3 and 4, sc with basal thickening on lower surface of wing, rs and m unbranched, rl sometimes lacking. Pilosity very sparse or completely lacking on margin and veins of hindwing. Distal segment of sc, on lower surface of forewing, with a row of small denticles, diminishing in length towards wing margin (fig. 2) (no such denticles in Speleketor flocki), c in forewing with scale-like sculpture (fig. 2). 876 CHARLES LIENHARD Terminalia of female as described in generic diagnosis (figs 25, 26, 28, 29), with the following complements. Epiproct simple, sense cushion of paraproct with 5-6 trichobothria in normal sockets (no basal rosettes) and 1 shorter ordinary hair. Proximal lobe of ovipositor valvulae with 5-6 long ventral hairs, 3 short hairs on inner margin and a transversal row of 4 short dorsal hairs basally to distal process. Distal process with 7-8 spiniform setae distributed throughout its length: 2 of them on dorsal face, 2-3 on inner margin, 2 on ventral face, | on apex. This terminal spine with flattened apex, slightly spoon-shaped (cf. detail of fig. 28). Subgenital plate on each side with a pair of long setae just anterior to the sclerified link with ovipositor valvulae. Basal part of subgenital plate with a characteristic pattern of cuticular pigmentation, apical lobe hyaline (fig. 28). Spermapore, spermatheca and sperma- tophore (= "sperm packet" of Mockford, 1993) as described in generic diagnosis (figs 2526); Measurements (2 holotype, mm). Body length = 2.5. Forewing length = 2.9. Hindwing length = 1.46. Length of hindfemur = 0.86. Length of hindtibia = 1.45. Length of hindtarsomeres (measured from condyle to condyle): rl = 0.55; #2 = 0.15; W308: Remark. The male is not yet known, but the presence of spermatophores in the spermatheca of the dissected female gives evidence of the bisexuality of the species. GENERAL DISCUSSION PHYLOGENETICS Within the family Prionoglarididae the genus Prionoglaris is characterized by the autapomorphic characters of head morphology already mentioned in the intro- duction; another autapomorphy of this genus is the asymmetrical structure of pretarsal claws and their metamorphosis during the last moult (presence of a membraneous vesicle on inner side of anterior claw in adults, no vesicle in nymphs, cf. Lienhard, 1988, 1998). The clade comprising the remaining two genera, Speleketor and Sensitibilla, is defined by the synapomorphic presence of trichobothria on legs. Within this group, Speleketor is characterized by the presence of a row of articulated spines on forefemur and Sensitibilla by the very specialized genitalia of females. The presence of tricho- bothria on ventral side of femur in all known species of Speleketor, mentioned by Mockford (1984), indicates that the trichobothria pattern is of diagnostic value on generic level. The more complex pattern seen in Sensitibilla may be interpreted as the apomorphic character state. GEOGRAPHICAL DISTRIBUTION The distribution of the currently known species of Prionoglarididae is the following (cf. Badonnel & Lienhard, 1994; Lienhard, 1988, 1996, 1998; Mockford, 1984, 1993): Genus Prionoglaris (type species: P. stygia): Prionoglaris stygia Enderlein, 1909: Western Palaearctic, usually in caves, sometimes also under stones and within A NEW GENUS OF PRIONOGLARIDIDAE 877 rocky débris, recorded from the following countries: Belgium, France (type locality), Germany, Greece, Morocco, Portugal, Switzerland, Turkey, former Yugoslavia. — Prionoglaris dactyloides Lienhard, 1988: Greece: Peloponnese (type locality, within rocky débris) and Crete (in cave). — Prionoglaris lindbergi Badonnel, 1962: Afghanistan, only one nymph known (holotype), under stone. Genus Speleketor (type species: S. flocki): Speleketor flocki Gurney, 1943: USA: Southern Arizona (Tucson Mountains, in cave, type locality) and Southeastern Nevada (near Las Vegas, in cave). — Speleketor irwini Mockford, 1984: USA: Southern California, on the skirts of dead leaves of the palm Washingtonia filifera in native stands. — Speleketor pictus Mockford, 1984: USA: Southern California, collected at a light (exact habitat unknown). Genus Sensitibilla gen. n.: Sensitibilla strinatii sp. n.: Namibia, Arnhem Cave. The few populations known are strongly localized, therefore all these species are considered as extremely rare, even the widely distributed P. stygia is very rarely collected. Two species, Sp. flocki and S. strinatii are only known from caves. P. stygia and P. dactyloides are also regularly found in caves. But probably none of the species is exclusively cave inhabiting; they are all fully winged, well pigmented and have well developed compound eyes and ocellı. It is interesting to see that all species of Speleketor and Sensitibilla are known from desert regions only. The strikingly disjunct distribution pattern (SW North America and SW Africa) of these closely related genera may be interpreted as an indication of a relatively old age of this clade. FUNCTIONAL MORPHOLOGY Some of the morphological characters observed in the new genus reveal several interesting problems of functional morphology which have not been studied in detail but which I would like to point out here very briefly. The presence of trichobothria on legs in Sensitibilla and Speleketor, unique in Psocoptera, has been described in detail in the discussion following the description of the new genus. These sensilla are very long and extremely fine smooth filiform hairs, inserted in very large and deep sockets (relative to the diameter of the hairs, cf. fig. 20). Their arrangement is apparently constant within a species and essentially the same in nymphs and adults. The absence, in adults of Sensitibilla, of the basal trichobothrium on hindtibia (fig. 19) can be explained by the position of the forewing, which touches and covers the basal half of hindtibia in resting position (cf. figs a and b of pl. 9 in Lienhard, 1998, showing the analogous situation in Prionoglaris stygia which has a very similar habitus). The resting position of wings is the same in Speleketor flocki, where the forewing covers also most of hindfemur and basal part of hindtibia; this situation is incorrectly figured by Gurney (1943: fig. 3). These trichobothria are thin but relatively rigid hairs. When preserved spe- cimens of Sensitibilla strinatii are moved in alcohol, near to the surface of the liquid, it can be observed that the trichobothria easily inflect at their base, without being bent. No histological investigation has been made but the observations mentioned above confirm the hypothesis that these sensilla are real trichobothria, as they are 878 CHARLES LIENHARD known in arachnids, for example (cf. Foelix, 1996: 71-73). In spiders, the tricho- bothria are stimulated by air currents and low frequency air vibrations (sound). Probably their function could be the same in Sensitibilla and Speleketor and they may be interpreted as adaptations to cave life, even if only one of the three species of Speleketor is known to be cavernicolous. But they could also play an important role in the more general context of a life under the conditions of desert climate (cf. above: "Geographical distribution"). The different position of trichobothria in these genera (ventral side of basal segments of legs in Speleketor, dorsal side of apical segments of legs in Sensitibilla) may indicate a certain difference in function and therefore in biology of these species. The row of articulated spines on forefemur in Speleketor could perhaps also be related to this biological specialization. The row of denticles on distal segment of sc in forewing, present in Sensitibilla (absent in Speleketor and Prionoglaris) could even- tually be involved in a specialized ethology; it could be interpreted as a sound pro- ducing rasp, but there is no evidence of a corresponding frictional structure. The presence of morphologically different mouthparts (laciniae, mandibles) in adults and nymphs of Sensitibilla (and to a somewhat lesser extent in Speleketor) indicates a difference in nutritional biology of these life stages. As in Prionoglaris, an interpretation is not possible due to the lack of direct observations on animals in captivity (cf. Lienhard, 1988: 103). The reduction of chewing sculpture and the presence of a brush-like fringed basal lobe on right mandible of nymphs and adults could also be related to some nutritional specialization. In this context, it seems interesting to mention the presence of densely fringed mandibles in Arixenia esau Jordan (Dermaptera), which lives on bat guano in tropical caves (cf. Strenger, 1977). The fusion of the ovipositor valvulae with the subgenital plate in Sensitibilla results in a unique functional structure which in this configuration is unknown in other psocids. The strong distal process of ovipositor valvulae, bearing several spiniform setae, is reminiscent of the situation in the Namibian desert psocid Spina- tropos philippi Lienhard, 2000 (Trogiomorpha: Atropetae: Trogiidae), where similar spines have been tentatively interpreted as digging organs (Lienhard, 2000). Sensi- tibilla strinatii lives under stones on dusty and sandy soils; that is why this inter- pretation could also be true for this species, which is only very distantly related to Spinatropos (same suborder, different family-group). The particular shape and pilosity of the ventral half of the paraproct in Sensitibilla could be in functional relation with the unit "ovipositor valvulae-subgenital plate" during oviposition. ACKNOWLEDGEMENTS I am very grateful to the renown biospeleologist Dr P. Strinati (Geneva: Cologny) for having made this interesting material available to me. I also thank Dr E. L. Mockford (Normal, Illinois) for the loan of a female paratype of Speleketor flocki and for reading the manuscript and making some valuable suggestions. Dr P. Schwendinger (Geneva) kindly indicated to me some literature concerning problems of functional morphology. A NEW GENUS OF PRIONOGLARIDIDAE 879 Fics 1-16 Figs 1-11. Sensitibilla strinatii, female: 1, forewing; 2, detail of distal segment of sc in forewing; 3, hindwing (holotype); 4, variant of venation (anteroapical part of hindwing, paratype); 5, head (dorsal view); 6, hypopharyngeal sclerites (dorsal view); 7, maxillary palp (ordinary pilosity not figured); 8, marginal sensillum of P4; 9, mandibles (frontal view): 10, pretarsal claw; 11, lacinia. — Figs 12-14. Sensitibilla strinatii, nymph: 12, lacinia; 13, mandibles (frontal view); 14, pretarsal claw. — Speleketor flocki, female (paratype): 15, double marginal sensillum on P4; 16, mandibles (frontal view). — N.B. All corresponding figures of laciniae and mandibles at same magnification. 880 CHARLES LIENHARD Fics 17-24 Figs 17-20. Sensitibilla strinatii, female: 17, foreleg; 18, midleg; 19, hindleg; 20, hindtarsus. — Fig. 21. Sensitibilla strinatii, nymph: hindleg. — Figs 22-24. Speleketor flocki, female (paratype): 22, foreleg; 23, midleg; 24, hindleg. — N.B. Figs 17-19 and 21 at same magni- fication, figs 22-24 at lower magnification. Ordinary pilosity not represented on these figures (only spinelike setae and trichobothria figured). A NEW GENUS OF PRIONOGLARIDIDAE 88] Fics 25-30 Sensitibilla strinatii, female: 25, spermatheca containing three spermatophores; 26, rows of pores on spermathecal wall (detail to fig. 25); 27, labial palp (ventral view); 28, subgenital plate and ovipositor valvulae (ventral view); 29, epiproct and left paraproct; 30, sensilla on distal margin of labrum (right side and middle). 882 CHARLES LIENHARD REFERENCES BADONNEL, A. & LIENHARD, C. 1994. Psocoptera (pp. 301-305). Jn: JUBERTHIE, C. & DECU, V. (eds). Encyclopaedia Biospeologica, tome I, XII + 834 pp. Société de Biospéologie, Moulis. FOELIX, R. F. 1996. Biology of spiders (second edition). Oxford University Press, New York, 330 pp. GURNEY, A. B. 1943. A synopsis of the psocids of the tribe Psyllipsocini, including the descrip- tion of an unusual new genus from Arızona (Corrodentia: Empheriidae: Empheriinae). Annals of the entomological Society of America 36: 195-220. LIENHARD, C. 1988. Vorarbeiten zu einer Psocopteren-Fauna der Westpaläarktis. IV. Die Gattung Prionoglaris Enderlein (Psocoptera: Prionoglarididae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 61: 89-108. LIENHARD, C. 1996. Psocopteres nouveaux ou peu connus de quelques îles atlantiques (Canaries, Madère, Açores, Ascension) et de l'Afrique du Nord (Insecta: Psocoptera). Boletim do Museu municipal do Funchal (Historia Natural) 48(267): 87-151. LIENHARD, C. 1998. Psocoptères euro-méditerranéens. Faune de France 83: XX + 517 pp. LIENHARD, C. 2000. A new desert psocid from Namibia (Insecta: Psocoptera: Trogiidae). Revue suisse de Zoologie 107(2): 277-281. MOCKFORD, E. L. 1984. Two new species of Speleketor from southern California with comments on the taxonomic position of the genus (Psocoptera: Prionoglaridae). South- western Naturalist 29(2): 169-179. MOCKFORD, E. L. 1993. North American Psocoptera (Insecta). Flora and Fauna Handbook 10: XVIII+455 pp. Sandhill Crane Press, Gainesville, Florida. STRENGER, A. 1977. Arixenia esau J., kein Parasit. Eine funktionsmorphologische Studie. Zoologischer Anzeiger 199: 95-106. REVUE SUISSE DE ZOOLOGIE 107 (4): 883-892; décembre 2000 Two new Tephritidae (Diptera) from the Western Palaearctic region Bernhard MERZ Muséum d histoire naturelle, c.p. 6434, CH-1211 Geneve 6, Switzerland. E-mail: bernhard.merz @ mhn.ville-ge.ch Two new Tephritidae (Diptera) from the Western Palaearctic region. - Chaetostomella baezi sp. n. from the Canary Islands (El Hierro, La Palma, Teneriffa), which was reared from flowerheads of Cheirolophus duranii, Ch. junonianus and Ch. sventeni gracilis, is an unusual species. Although having an anterior dorsocentral seta, which is a character of Chaetorellia, it is placed in Chaetostomella because of the shape of the male terminalia and the strong setae along the vibrissal angle. Its generic position is discussed. Campiglossa freidbergi sp. n. is described from high altitude in the Sierra Nevada mountains of Southern Spain. It belongs to the C. difficilis-group and differs from C. difficilis (Hendel) mainly in the wing pattern and details of the terminalia. Its hostplants are not known. Key-words: Diptera - Tephritidae - new species - Spain - Canary Islands. INTRODUCTION Knowledge of the western Palaearctic Tephritidae is rather good (Norrbom et al., 1998), with about 250 species recorded from this area. Nevertheless, new species may still be found, mostly from areas which harbour a rich endemic flora and fauna. In particular, the Mediterranean region is the least studied. It is therefore not Surprising, that two quite large, characteristic species have been found recently in Southern Europe and the Canary Islands which are apparently new to science. The fauna of the Canary Islands has been studied by several authors during the past 100 years. These studies have been summarized by Merz (1992) who listed 34 species from these islands. Shortly after the publication of that paper the author received independently from B. van Aartsen and M. Baéz specimens of an as yet undescribed and problematic species. It shows an unusual set of characters which make its correct generic position difficult to determine, and it is described here as Chaetostomella baezi sp. n. The Tephritidae of mainland Spain have never been treated in a comprehensive manner. The most important papers are listed in Merz & Blasco-Zumeta (1995) who estimated that about 80 species are known from Spain. However, knowledge of the fauna of the adjacent countries suggested that further species should be found on the Iberian Peninsula. The species described here was found at high altitude in the Sierra Nevada and it belongs to Campiglossa. This genus is almost worldwide in distri- Manuscript accepted 04.09.2000 884 BERNHARD MERZ bution, with about 200 described species. It was revised for Europe by Merz (1994) and for the Eastern Palaearctic region by Korneyev (1989). Only C. producta (Loew) is widespread throughout the entire Mediterranean region, whereas some other species are found sporadically at higher altitudes. None of these species is known to be endemic in this area. MATERIAL The specimens studied belong to the following institutions and personal collections: CBVA: private collection Bob van Aartsen, t Harde, Netherlands MHNG : Muséum d’histoire naturelle, Genève, Switzerland TAU: Tel Aviv University, Israel ZCUL: Zoological collection, University of La Laguna, Tenerife The terminology follows Merz & Haenni (2000) for external morphology and Korneyev (1985) for structures of male terminalia. SYSTEMATIC PART Chaetostomella baezi sp. n. Figs 1-11 MATERIAL Holotype 9. [Canary Islands:] La Palma, Fuencaliente, IIT.1996, leg. R. Mesa / ex larva Cheirolophus junonianus (ZCUL). The specimen is glued laterally on to a card and is in good condition. Paratypes, Canary Islands: La Palma, same data as holotype, 66 4, 19; Bco [= Barranco] Fagundo, 400m, 9.XII.1994, leg. M. Baéz, 1 9. El Hierro, Sabinosa, 14.VIII.1985, leg. M. Baéz, 1¢, 29 2; Sabinosa, III.1995, leg. R. Meso, ex larva Cheirolophus duranii, 26 6; Frontera, 14.VIII.1985, leg. M. Baéz, 14. Tenerife, El Tablado, IV.1995, leg. R. Meso, ex larva Cheirolophus sventeni, 13; Buenavista, 2.1V.1968, leg. B. van Aartsen, 1 ©. The para- types are deposited in the following institutions: MHNG, ZCUL and CBVA. Additional material: La Palma, Barranco Fagundo, 400m, 9.XII.1994, leg. M. Baéz, 1 specimen without abdomen (also | wing missing); Barranco del Jorao, Tijarafe, 4.1X.1994, leg. R. Mesa Coello, cultivad en Cheirolophus sventeni gracilis, Chaetorellia n.sp. ?, det. M. Baéz, 13 (without wings). El Hierro, Sabinosa, III.1995, leg. R. Meso, ex larva Cheirolophus duranii, 13 (without wings). Tenerife, Guimar, 17.V.1993, leg. M. Baéz, 1d (teneral); Barranco del Rio, 21.VII.1996, leg. M. Baéz, 1 without abdomen (also head missing). Remark: The specimens sent by M. Baéz were badly damaged in the post. Whereas the paratypes show at most some minor damage, all other specimens with heavy damage are excluded from the type series. Nine specimens dropped of their pins and are not listed here because it is impossible to assign them to their locality labels. ETYMOLOGY Named in honour of Marcos Baéz, La Laguna, in recognition of his long-term investigations of the Diptera of the Canary Islands. DIAGNOSIS Medium sized species of Chaetostomella. Differs from all other species of the genus by the presence of a presutural dorsocentral seta and the arrangement of the TWO NEW TEPHRITIDAE 885 black spots on the mesonotum, with one large black spot in the triangle of the two supraalar setae and the wing base. DESCRIPTION Head (Fig. 1). Entirely yellow, only around ocelli with darker spots; in profile about 1.3 times higher than long; gena and vibrissal angle densely covered with long black and shorter whitish setulae, the anteriormost 2-3 black setulae stronger; frontal plate densely covered with dark setulae; frons bare; scape and pedicel with black setulae; chaetotaxy as usual in the genus: 3 dark frontal setae, 2 dark orbital setae, the posterior pair inclinate; | dark ocellar seta, 1 dark medial vertical and 1 dark lateral vertical seta, | whitish, upright postocellar seta, between them usually with 1-3 small medial postocellar setulae, all postocular setae dark, but just behind them with several rows of whitish occipital setulae. Thorax (Fig. 2). Brownish-yellow in ground colour, but mesonotum with lyre- like pattern, which may be somewhat more or less developed as in Fig. 2; katepis- ternum usually pale reddish, but central spot in a few specimens with blackish upper margin; 6 pairs of dark spots visible on the mesonotum: 3 pairs of small dark spots present at the bases of the prescutellar seta and the anterior and posterior dorsocentral setae; 3 pairs of large dark spots present at the base of the presutural seta, in a triangle formed by the anterior and posterior supraalar seta and the base of the wing, and on the lower postalar callus; scutellum with a large apical and two smaller laterobasal dark spots; anepisternum, anepimeron and katepisternum densely covered with predo- minantly pale, rough setulae; posterior margin of anepisternum with 1-3 black setae; posterodorsal corner of katepisternum with | black seta, anepimeron with | dark, central seta; legs yellowish. Wing (Fig. 3). Pattern as usual in the genus; discal and preapical crossbands always separated; preapical and apical crossbands usually connected, although some- times only for a short distance; lower extension of posterior cubital cell reaching posteriorly beyond cell bm; halteres yellowish; fringes of upper calypter darkened. Abdomen (Fig. 4). Ground colour brownish-yellow; tergites 2-5 (in d ) and 2-6 (in 2) each with 2 pairs of small dark spots; setulae usually entirely dark, only on first tergite mainly pale; last tergite of male slightly shining, other tergites mat. Male terminalia (Figs 5-8). Epandrium, hypandrium and associated structures as usual in the genus; distiphallus without spines or setulae; general structure of glans as in other Chaetostomella; ejaculatory ductus evenly curved; at base of glans with an ill-defined serrate sclerite; ligula at least half as long as maximum length of glans, slightly brownish towards tip. Female terminalia (Figs 4, 9-11). Oviscape orange, but at base usually with a pair of small, dark lateral spots; all setulae dark; length about equal to that of the preceeding 3 tergites combined; spermathecae rather narrow, cylindrical, densely covered with elongate papillae; aculeus evenly acute, only at tip of sternite 8 with slight shoulder, tip evenly rounded; ratio of length of sternite 8 / total length of aculeus about 0.7. Wing length: 4.9-5.4 mm (male), (4.0-) 4.8-5.5 mm (female). 886 BERNHARD MERZ Fics 1-4 Chaetostomella baezi sp. n.: 1, head, lateral view; 2, mesontoum and scutellum, dorsal view; 3, wing; 4, abdomen of 2, dorsal view. TWO NEW TEPHRITIDAE 887 Fics 5-8 Chaetostomella baezi sp. n, male: 5, external terminalia, caudal view; 6, external terminalia, lateral view; 7, glans, lateral view; 8, glans, ventral view. 888 BERNHARD MERZ BIOLOGY Some specimens were reared from Cheirlolophus duranii, Ch. junonianus and Ch. sventeni gracilis. This genus belongs to the tribe Cardueae within the Asteraceae. Although not stated on the labels, it may be assumed that the larvae live in the flowerheads. DISCUSSION With the well developed presutural dorsocentral seta and the presence of 12 dark spots on the mesonotum this species exhibits the diagnostic characters of Chaetorellia. However, the body is more robust than in Chaetorellia, the male terminalia are almost identical with other species of Chaetostomella (Korneyev, 1985; Freidberg & Kugler, 1989, pers. obs.), the vibrissal angle bears some strong, dark setulae and the lyre-pattern on the mesonotum is very similar to the known species of Chaetostomella, so that its placement in the latter genus seems justified. The discovery of this new species with some characters intermediate between Chaetorellia and Chaetostomella raises the question about the status of the two genera. According to V.A. Korneyev (pers. comm.) also few or most specimens of other species of Chaetostomella (Ch. rossica Hendel) and Terellia (T. blanda Richter) may have a presutural dorsocentral seta. Therefore, the number of dorsocentral setae does not seem of generic value and the synonymy of Chaetostomella and Chaetorellia should probably be proposed. However, this action is premature without the study of the terminalia of all species, which is outside the scope of this paper. The main external differences between the two genera may be summarized as follows: | Vibrissal angle with some well developed dark setae; if presutural dorsocentral seta present, then mesonotum with dark spot laterad of supraalar setae; in general larger, more robust species . Chaetostomella Hendel - Vibrissal angle setulose, but without outstanding dark setae; presutural dorsocentral seta always present and inserted on a black spot; never with dark spot laterad of supraalar setae; more slender species So Soa aera ER ER Li. MO VII Chaetorellia Hendel Campiglossa freidbergi sp. n. Figs 12-17 MATERIAL Holotype 3. Spain: Sierra Nevada Veleta, 2500m, 4.X.1985, A. Freidberg (TAU). Paratypes: same data as holotype, 6d 4, 39 9. The paratypes are deposited in MHNG and TAU. The holotype is double mounted with a minutien pin on a polyporus strip and is in excellent condition. ETYMOLOGY Named in honour of Amnon Freidberg, the leading specialist on Tephritidae, the finder of the new species, and an excellent colleague. TWO NEW TEPHRITIDAE 889 Fics 9-11 Chaetostomella baezi sp. n., female: 9, aculeus, ventral view; 10, tip of aculeus, ventral view: 11, spermatheca. DIAGNOSIS Medium sized species of Campiglossa, wing length 3.5-4 mm. It is charac- terized by the mixed black and whitish postocular setae, the black posterior noto- pleural seta and in particular by its wing pattern, which shows some quite isolated, much reduced black areas and very large, often fused spots over the entire surface. DESCRIPTION Head (Fig. 12). General colour yellow, with occiput dark and sometimes with a dark band between the two vertical setae; in profile slightly higher than long; frons with indistinct longitudinal vitta; antennae at base separated from each other by half the width of diameter of scape; labellae 1.7-2 times as long as length of Ist flagello- mere; scape and pedicel with dark setulae; chaetotaxy: 2 dark frontal setae, 1 dark and 890 BERNHARD MERZ 1 white orbital setae; 1 dark ocellar seta; 1 whitish lateral vertical and | dark medial vertical seta; 1 pale postocellar seta, 1 pale paravertical seta; postocular setae mixed dark and pale. Thorax. General colour black with thick greyish microtrichosity; mesonotum with 5 indistinct darker longitudinal stripes; postpronotal lobe slightly yellowish; setulae on pleura whitish, rough; chaetotaxy as usual in the genus, posterior noto- pleural seta, anepisternal seta and katepisternal seta black; anepimeral seta whitish; apical scutellar setae inserted on slightly paler ground, about half as long as basal scutellar setae; coxa, trochanter and femur of all legs mostly black, parts of distal half of femur, tibia and tarsus yellow, sometimes hind tibia with dark ring at middle. Wing (Fig. 13). Dark areas rather narrow, hyaline parts of wing large, most spots below vein R4+5 are fused; r2+3 always with a large hyaline spot anterior to R-M; cell cual and anal lobe with only a very indistinct pattern of isolated bars. Abdomen. Of the same greyish microtrichosity as thorax; the paired medial dark spots on tergites 2-5 well developed; all setulae pale. Male terminalia (Fig. 14). Epandrium with a small caudal plate (as in Merz, 1994, Fig. 13y); distal end of distiphallus with two clusters each of 10-15 small setulae; glans elongate, rostrum rather long, distinctly, but weakly sclerotized along entire length; vesica narrow. Female terminalia (Figs 15-17). Oviscape black, about as long as preceding 2- 3 tergites of preabdomen; aculeus evenly narrowed towards tip, but at apex with a distinct step; length of aculeus 1.0 mm (one specimen dissected); spermathecae ovoid, entirely covered by small papillae. BIOLOGY No hostplants are known for this species. DISCUSSION Together with €. difficilis (Hendel), this species forms the C. difficilis-group of Merz (1994). Both species share the same chaetotaxy and the same type of male terminalia with a cylindrical acrophallus, a narrow vesica, and an elongated rostrum. However, both species differ clearly in the wing pattern and in details of the male terminalia: The preapical lappet of the vesica of €. difficilis is absent in C. freidbergi. Further, the internal structure of the acrophallus shows some slight differences (see Fig. 15a in Merz, 1994). Another species which may be confused with C. freidbergi is C. cain (Hering) from Ethiopia and Kenya, which has a very similar wing pattern and head shape. However, the two species differ clearly in the structure of the terminalia in both sexes: The epandrium bears long projections in C. cain, as in C. argentata (Munro) (see Fig. 79 in Munro, 1957), which are entirely absent in C. freidbergi. The rostrum in the glans of C. cain is much narrower and bears an apical sclerotized semicircle, which is somewhat similar to C. anomalina (Bezzi), as illustrated in Fig. 114 in Munro (1957). The females differ in the tip of the aculeus, which is truncated at the tip and without any step in C. cain. TWO NEW TEPHRITIDAE 891 Fics 12-17 Campiglossa freidbergi sp. n: 12, head, lateral view; 13, wing; 14, 5 glans, ventral view; 15, © aculeus, ventral view; 16, © tip of aculeus, ventral view; 17, 2 spermatheca. ACKNOWLEDGEMENTS My sincerest thanks are offered to B. van Aartsen (t?Harde) M. Baéz (La Laguna) and A. Freidberg (Tel-Aviv) for making these flies available for study. Further, I thank V. Korneyev (Kiev) and A. C. Pont (Goring-on-Thames) for checking a first draft of this manuscript. REFERENCES FREIDBERG, A. & KUGLER, J. 1989. Diptera: Tephritidae. Fauna Palaestina - Insecta IV: 212 pp. & 8 plates. KORNEYEV, V. A. 1985. Fruit flies of the Tribe Terelliini Hendel, 1927 (Diptera, Tephritidae) of the fauna of the USSR. Entomologiseskoe Obozrenie 49 (4): 626-644. (in Russian; English translation in: Entomological Review, Washington 65 (1): 35-55). 892 BERNHARD MERZ KORNEYEV, V. A. 1989. A review of Sphenella and Paroxyna series of genera (Diptera, Tephritidae, Tephritinae). Insects of Mongolia 11: 359-470. (in Russian). MERZ, B. 1992. The fruit flies of the Canary Islands (Diptera: Tephritidae). Entomologica Scandinavica 23 (2): 215-231. MERZ, B. 1994. Diptera, Tephritidae. Insecta Helvetica, Fauna 10: 1-198. MERZ, B. & BLASCO-ZUMETA, J. 1995. The fruit flies (Diptera, Tephritidae) of the Monegros region (Zaragoza, Spain), with the record of the host plant of Rhagoletis zernyi Hendel, 1927. Zapateri, Revista aragonensis de entomologia 5: 127-134. MERZ, B. & HAENNI, J. P. 2000. 1.1. Morphology an terminology of adult Diptera (other than terminalia) (pp. 21-51). In: PAPP, L. & DARVAS, B. (eds). Contributions to a manual of Palaearctic Diptera, Volume 1. Science Herald, Budapest, 978 pp. Munro, H. K. 1957. Trypetidae. Ruwenzori Expedition 2 (9): 853-1054. NORRBOM, A. L., CARROLL, L. E. & FREIDBERG, A. 1998. Status of knowledge (pp. 9-47). In: THOMPSON, F. C. (ed). Fruit Fly Expert Identification System and Systematic Infor- mation Database. Myia 9: 1-524. WHITE, I. M. & MARCQUART, K. 1989. A revision of the genus Chaetorellia Hendel (Diptera: Tephritidae) including a new species associated with spotted knapweed, Centaurea maculosa Lam. (Asteraceae). Bulletin of Entomological Research 89: 453-487. REVUE SUISSE DE ZOOLOGIE Tome 107 — Fascicule 4 POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymena- phorura alticola (Bagnall, 1935) from the Alps and description of a new related species from the Sudetes, Hymenaphorura improvisa sp. n. (eollembeola: Onychiundae) 7. 2 EN Dal ee ZAWADZKI, Claudio H., Roberto E. REIS & Erasmo RENESTO. Allozyme discrimination of three species of Loricariichthys (Siluriformes: Lori- Canidae) ItomsSouthern Brazil IRR MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Ori- batida: Dampfiellidae and Otocepheidae). (Acarologica Genavensia RODELN a ME PAR AA, LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special references to the genus Tityobuthus Pocock (Scorpiones, RUI RO fe hr cs nen, mcd Oem OA ae ee PAGES, Jean. Japygidés (Diplura) du Sud-Est asiatique n° 9. - Dicellurata GEMGVEMSIGIONIING Seek anne hs ee ee RE Nec kr COMELLINI, André. Notes sur les Psélaphines néotropicaux (Coleoptera, Staphylinidae, Pselaphinae) — 11. Un nouveau genre et sept especes nouvelleside la tabuides Metopiasini 2 as. - a. fu CE eae BOHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scin- cidae: Lygosominae): resurrection of C. simulans (Vaillant, 1884) and descnpton oh a NEW ASPECIES. hie EU EI a ee ry Ile ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae, Proctolabinae) with description of new species. .......... VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura: Microhylidae) from a small forest remnant of the central high plateau OMNIA SASCARI vl ten N Re GANTENBEIN, Benjamin, Victor FET, Mark BARKER & Adolf SCHOLL. Nuclear and mitochondrial markers reveal the existence of two para- patric scorpion species in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha Caporiacco, 1950, stat. nov. (Euscorpiidae)...... LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave ([MSecta PSOCOpiErA) o I MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palae- ACEC ME TION CATIA RRR RATA ee a Pages 657-662 663-674 675-720 721-736 737-764 765-776 777-791 793-834 835-841 843-869 871-882 883-892 REVUE SUISSE DE ZOOLOGIE Volume 107 — Number 4 POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymena- phorura alticola (Bagnall, 1935) from the Alps and description of a new related species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola\Onychiuridae) ER ZAWADZKI, Claudio H., Roberto E. REIS & Erasmo RENESTO. Allozyme discrimination of three species of Loricariichthys (Siluriformes: Lori- Canidae) fromsS outhemvrBrazils . LEE MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Ori- batida: Dampfiellidae and Otocepheidae). (Acarologica Genavensia REED) NRA SEE de TT ed EE LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special references to the genus Tityobuthus Pocock (Scorpiones, BUTS) PNR ine een RISI TE Sent an ee ee NE PAGES, Jean. Japygidae (Diplura) from South-East Asia n° 9. - Dicellurata Genaven a PON Vin nu. le ae ee SIRIO E COMELLINI, André. Notes on Neotropical Pselaphines (Coleoptera, Staphy- linidae, Pselaphinae) 11. A new genus and seven new species of the tnibesMetopiasınn. Ae er EO RE ak REN a BÖHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scin- cidae: Lygosominae): resurrection of C. simulans (Vaillant, 1884) and description. OP aE W SPECIES. yo) ve ae A ee U ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae, Proctolabinae) with description of new species........... VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura: Microhylidae) from a small forest remnant of the central high plateau OEMAdADASCAR LE. we hoist Rashid ee Rods gale tip een RE GANTENBEIN, Benjamin, Victor FET, Mark BARKER & Adolf SCHOLL. Nuclear and mitochondrial markers reveal the existence of two para- patric scorpion species in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha Caporiacco, 1950, stat. nov. (Euscorpiidae)...... LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave (Insecta Psocoptera).i 2... 2. ei ee MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palae- ACUCHEITON: VIRALI EEE Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 657-662 663-674 675-720 721-736 737-764 765-776 777-791 793-834 835-841 843-869 871-882 883-892 PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N° 1-17 (1908-1926)... . . serie Fr. (prix des fascicules sur demande) REVUEDEPALEOBIOLOGIE . .. Ara... Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) ....... par fascicule Fr. COLLEMBOLENFAUNA EUROPAS von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Fr. THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION WITH KEYS FOR DETERMINATION by J. NOSEK, 346 pages, 111 figures in text, 1973 .............................. Fr. CLASSIFICATION OF THE DIPLOPODA pawRachardilE} FIOFEMAN) 230 pages: 1979) En. en cis sale se eee ees er Fr. LES OISEAUX NICHEURS DU CANTON DE GENEVE par P. GEROUDET, C. GUEX et M. MAIRE Solipasessnombreusesicartes et fieures, 1983 0.6 22. ae ne cna is ese eee Fr. CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ A LA CONNAISSANCE DES ECHINODERMES ACTUELS panMiuchel JANGOUX. 67 pages, Il planches; 1989... a an. nn. een. Er: RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) par Y. FINET, J. WÜEST et K. MAREDA, 62 pages, 1991 .......................... Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN; SHELLS AND RADULAS BYAYSEINESEISWIUESTSand KOIMAREDAS LOD FEN ee NO Er: O. SCHMIDT SPONGE CATALOGUE par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992.......... Fr. ATLAS DE REPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr. THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST PALSNVESIEINET ZL: 0 only 19932 ROOT oboe EE ON Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE pawJean-@laude/@AILLIEZ, 49 pages; 1995 eel sie ee oe oer etn a yee Sees Fr. PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr. CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) (Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997 CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis II) par Henri G. DIRICKX, 188 pages, 1998 ............ Br A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Fr. THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE SOKOTRA ARCHIPELAGO (Instrumenta Biodiversitatis IV) par Beat SCHÄTTI & Alain DESVOIGNES, 178 pages, 1999 285.— 30.— 40.— . 70.— Boia as 7 mt PMR Ho À VI) le ly j ala iat = Tor Den, Y Teer Dur in Volume 107 - Number 4 - 2000 Revue suisse de Zoologie: Instructions to Authors The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies that it has been approved by all named authors, that it reports their unpublished work and that it is not being considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of colour plates and large manuscripts. All papers are refereed by experts. 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