\ LAS NH ANNALES de la SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Geneve tome 112 fascicule 4 2005 g EI GENÈVE DÉCEMBRE 2005 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY — © ®, N nm En lead) „N 2 a Sf) ca è > hertz! = REVUE SUISSE DE ZOOLOGIE TOME 112—FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d’histoire naturelle de Genéve CHARLES LIENHARD Chargé de recherche au Muséum d’ histoire naturelle de 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 a des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 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. 250.— (en francs suisses) Les demandes d’abonnement doivent étre adressées a la rédaction de la Revue suisse de Zoologie, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse ANNALES de la SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D’HISTOIRE NATURELLE de la Ville de Genéve tome 112 fascicule 4 2005 JAN 3 12006 | LIBHAKIED E 2 HI GENEVE DECEMBRE 2005 ISSN 0035 - 418 X REVUE SUISSE DE ZOOLOGIE SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE TOME 112—FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE VOLKER MAHNERT Directeur du Muséum d’histoire naturelle de Genéve CHARLES LIENHARD Chargé de recherche au Muséum d’histoire naturelle de Genéve Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Geneve et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis 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. 250.— (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 Genéve 6, Suisse REVUE SUISSE DE ZOOLOGIE 112 (4): 771-805; décembre 2005 Additional notes on the fauna of Agromyzidae (Diptera) in Switzerland Milos’ CERNY CZ-763 63 Halenkovice 1, Czech Republic. E-mail: cerny.milos@centrum.cz Additional notes on the fauna of Agromyzidae (Diptera) in Switzerland. - New faunistic data on the Swiss Agromyzidae are given. 144 species are recorded including 92 species found in Switzerland for the first time. Each species included is accompanied by information on its host plants and geographical distribution. The current list of the Swiss Agromyzidae thus embraces 232 species. Keywords: Diptera - Agromyzidae - faunistics - biology - Switzerland. INTRODUCTION The Agromyzidae, with its 1165 species described from the Palaearctic Region (Scheirs et al., 1999) and 843 species from Europe (Martinez, 1998) is one of the larger families of Diptera. The Swiss fauna of Agromyzidae is little studied with 140 recorded species in 19 genera (Martinez, 1998). On the other hand, the study of Agromyzidae in Europe and especially in the other countries of Central Europe is relatively advanced (cf. e.g. Griffiths, 1980; Nowakowski, 1973; Spencer, 1964, 1966, 1972b, 1976; Zlobin, 1992, 1993a, 1993b, 1993c, 1993d, 1994, 1995, 2003a, 2003b). The following species were described directly from the territory of Switzerland (type locality in parentheses, abbreviation of cantons in following chapter): Agromyza alnibetulae Hendel, 1931 (type locality: Switzerland); Chromatomyia gentianella (Hendel, 1932) (Switzerland); Ch. hoppiella Spencer, 1990 (GR: Lenzerheide); Liriomyza polygalae Hering, 1927 (TI: Lugano); Napomyza cichorii Spencer, 1966 (VS: Orsiéres); Phytobia mallochi Hendel, 1924 (GR: Peist); Phytomyza abdita Hering, 1927 (ZH: Ziirichberg); P. alpestris Hendel, 1920 (GR: St. Moritz); P. hoppi Hering, 1925 (SG: Pfafers); P. narcissiflorae Hering, 1928 (GR: Parpan); P. hoppiana Hering, 1931 (GR: Zernez) as a synonym of P. pulsatiallae Hering, 1924 (Spencer, 1976); P. thalictri Escher-Kiindig in Rougemont, 1912 (Switzerland); P. virgaureae Hering, 1926 (SZ: Rigi); P. thymi Hering, 1928 (GR: Lenzerheide) and Pseudonapomyza europaea Spencer, 1973 (VS: Stalden). P. atragenis Hendel, 1931 (GR: Zernez) and P. philactaeae Hering, 1932 (GR: Vals) were synonymized with P. kaltenbachi Hendel, 1922 (Hendel, 1931-6; Papp, 1984). P. prenanthidis Hering, 1932 (SG: Pfäfers) was synonymized with P. sonchi Robineau-Desvoidy, 1851 by Hendel (1931-6). Later Spencer (1965) synonymized P. sonchi with P. marginella Fallén, 1823. Manuscript accepted 07.02.2005 T2 M. CERNŸ The new Material examined by the author provided records on 144 species including 92 species which represent faumistic novelties for Switzerland. In total 232 species are now known from Switzerland representing 28% of the European fauna. Agromyzids are mostly small to very small flies with a dark body sometimes showing a light metallic shine (Melanagromyza spp.), or a distinct yellow pattern (e.g. Liriomyza spp., Phytoliriomyza spp., Cerodontha spp.). The length of wings varies betwen 0.9 and 4.5 mm. The adult thorax is massive and the relatively broad abdomen consists of 6 visible segments. The eyes are broadly separated in both sexes, 1-2 rows of short orbital setulae are developed in addition to the regular orbital setae (except for Selachops). Trophic preferences are phytophagous, larvae typically feed upon tissues of living plants and form characteristic mines. The larvae are cylindrical, slightly tapered on both ends, their body consists of the cephalic segment, 3 thoracic and 8 abdominal segments. The length of body varies between 1.5-3.0 mm but the body of Phytobia is unusually long, sometimes reaching 25 mm. Larvae of most species live in the parenchym of leaves, some mine the surface of stems. Larvae of Phytobia feed upon bast of some wood plants. Hexomyza larvae form galls on twigs of trees and bushes. Larvae of Melanagromyza, Napomyza and Ophiomyia (partially) live within stems and on roots of host plants. Some Phytomyza larvae are specialized for life in flowers. Most Agromyzidae are oligophagous and their larvae prefer closely related species, genera or families of host plants. Some species are even monophagous and mine only one host plant. Only a relatively small group of agromyzids are widely polyphages and their spectrum of host plants includes numerous plants from many different families. Chromatomyia horticola (Goureau, 1851) may be mentioned as the best known and widely distributed polyphagous species mining a large series of plants. A review of Central European agromyzids with a certain economic importance was published by Spencer (1973). Some species used to be included among serious pests of crops, e.g. Liriomyza huidobrensis (Blanchard, 1926) recorded also from Switzerland (Martinez, 1998). However, many species damage only the general or aesthetical appearance of plants, and economical losses are not too serious. More important economic losses are known only during the mass occurrence of some species living on cereals, (Agromyza megalopsis Hering, 1933 or A. nigrella [Rondani, 1875]), on Fabaceae and Brassicaceae (Phytomyza rufipes Meigen, 1830) and some ornamental plants (Ch. horticola, L. huidobrensis). Larvae of Phytobia carbonaria (Zetterstedt, 1848) bore the cambium of apple twigs. The density of agromyzid populations is very often seriously influenced by hymenopteran parasites from families Braconidae, Chalcididae, Eulophidae and Ichneumonidae. The strictly monophagous species with their close relations to host plants may be used as suitable models for ecological research. MATERIAL AND METHODS Species originating from Switzerland were found in the following collections: CBM private collection Bohuslav Mocek, Hradec Kralové, Czech Republic, 8 spe- cimens. AGROMYZIDAE IN SWITZERLAND 773 CMB private collection Miroslav Bartäk, Praha, Czech Republic, 75 specimens. These specimens were collected with a Malaise Trap of the type M. Bartak (Rohäëek et al., 1998) above the tree line on 2000 m a.s.l. in the Alps of the Canton Graubiinden. MHNG Muséum d’histoire naturelle, Genéve, Switzerland (Bernhard Merz), 284 specimens. ZMUC Zoological Museum, University Copenhagen, Denmark (Rudolf Meier), 33 specimens. Genera and species are arranged alphabetically in the two subfamilies Agromyzinae and Phytomyzinae. All relevant data from locality labels are included. Only selected important synonymy and new References (with regard to Switzerland) are quoted here, for full data see Martinez (1998), Papp (1984), Spencer & Martinez (1987) and Spencer (1990, 1992). Nomenclature of Agromyzidae and plant names follow Spencer (1990). The localities are grouped according to cantons (districts) of Switzerland and the following abbreviations are used: AG = Aargau; BE = Bern; FR = Fribourg; GE = Genéve; GL = Glarus; GR = Graubiinden (= Grisons); JU = Jura; NE = Neuchatel; SG = St. Gallen; SH = Schaffhausen; SZ = Schwyz; TI = Ticino; VD = Vaud; VS = Valais; ZH = Zürich. MT on locality lables means Malaise Trap. The species which are new to Switzerland are marked with an asterisk (*) before the name. LIST OF SPECIES Subfamily AGROMYZINAE Agromyza abiens Zetterstedt, 1848 Reference: Martinez, 1998: 269. Material examined: GE: Cartigny, Moulin de Vert, 350 m.a.s.l., 1 2 ,2.vi.2002. GR: Lenzerheide near Sanaspans, 1500 m a.s.l.,1 2, 7.viii.1992. VS: Leuk Bahnhof, 625 ma.s.l., 1 d, 15.v.2000. ZH: Zürich, 500 ma.s.l.,1 2, 10.viii.1991. All B. Merz leg. (MHNG). Notes: This relatively large species with the orange frons and a broad epistoma was described from Sweden. A. abiens is generally distributed and locally common in Europe. Its occurrence in Japan was also confirmed. Larvae form broad leaf mines on Boraginaceae. Records include the genera Anchusa, Asperugo, Borago, Cynoglossum, Echium, Lycopsis, Pentaglottis, Pulmonaria and Symphytum. Agromyza albipennis Meigen, 1830 Reference: Martinez, 1998: 269. Material examined: GE: Chancy Bord du Rhône, 350 m a.s.l., 1 ¢ , 28.vii.2002. B. Merz leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 à , 1.vii.2002, B. Merz & Herrmann leg. (MHNG). Notes: A. albipennis is a Holarctic species which is common in Europe, espe- cially in its temperate and northern parts. It is less known from the Mediterranean area 774 M. CERNY (only Italy and Spain). Its occurrence in Japan, Kamchatka and North America (United States and Canada) is also documented. Larvae feed singly, forming linear-blotch mine on Gramineae, particularly on Phalaris arundinacea, less commonly on Poa and Hordeum, rarely on other grasses. *A gromyza bicaudata (Hendel, 1920) Material examined: VS: Leuk Pfynwald, 630 ma.s.l., 1 4, 21.iv.1998, B. Merz & Botta leg. (MHNG). Notes: The type series includes localities in Germany and Austria. It is distri- buted in temperate Europe (Austria, British Isles, the Czech Republic, Germany, Hungary, Lithuania, Poland and Slovakia). Hendel (1931-6) mentioned this species from Finland a South Russia. This species is unusual in having a strongly developed presutural de and specific male terminalia. Very probably a grass-feeder though its biology is not precisely known. New for Switzerland. Agromyza bromi Spencer, 1966 Reference: Martinez, 1998: 269. Material examined: VS: Leuk Platten, 630 m a.s.l., 1 4 ,22.iv.1998, B. Merz & B. Botta leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.l., 1 d, 16.viii.1998, B. Merz leg. (MHNG). Notes: The species is known from temperate Europe. The only recorded host is Ceratochloa unioloides (= Bromus catharticus) but other grasses are doubtless also attacked. *A gromyza cinerascens Macquart, 1835 Material examined: GE: Bernex, Signal, 510 ma.s.l., 1 &,21.111.1999; Chancy, La Laire, 350 ma.s.l., 1 d, 1.iv.2002; Russin, Les Baillets (Allondon), 390 m a.s.l., 1 6, 24.111.2003. ZH: Zürich Albisgütli, 490 m a.s.l., 1 4, 9.iv.1997; Zürich Allmend, 440 m a.s.l., 1 2, 17.iv.1996. All B. Merz leg. (MHNG). Notes: This Palaearctic species was described from France; it is known from many countries of Europe (Austria, Belarus, Belgium, Croatia, the Czech Republic, Denmark, England, Estonia, Finland, Germany, Hungary, Italy incl. Sicily, Lithuania, the Netherlands, Norway, Poland, Romania, Slovakia, Spain, Russia, and Sweden), North Africa (Egypt and Tunisia) and Japan. Host plants are Gramineae, most frequently Dactylis glomerata, sometimes Secale cereale. New for Switzerland. *A gromyza frontella (Rondani, 1875) Material examined: GE: Bernex, Signal, 510 m a.s.1., 1 4, 8.vii.2001, B. Merz leg. (MHNG). VS: Leuk Brentjong, 920 m a.s.l., 1 d, 15.v.2000, B. Merz leg. (MANG). Notes: This Holarctic species was described from Italy (Parma). It is common in Austria, Belarus, the Czech Republic, Denmark, England, France, Germany, Hungary, Italy, Lithuania, Poland, Russia, Slovakia, Spain, Sweden, Canada, United States, Afghanistan, Israel, and Turkey. A. frontella belongs to the Agromyza species feeding on Leguminosae. The larva forms a characteristic mine on Medicago sativa and other Medicago spp., but also on Melilotus. New for Switzerland. AGROMYZIDAE IN SWITZERLAND 775 *Agromyza hendeli Griffiths, 1963 Material examined: ZH: Zürich Irchel, 500 m a.s.1., 1 & , 16.viii.1998, B. Merz leg. (MHNG). Notes: A. hendeli is distributed in temperate Europe and known from Austria, Belgium, the Czech Republic, Denmark, England, Estonia, Germany, North Italy, Latvia, Lithuania, the Netherlands, Poland, Scotland and Slovakia. The larvae form a broad blotch mine on Phragmites communis. New for Switzerland. *Agromyza idaeiana Hardy, 1853 Material examined: GR: San Vittore, Rebberg, 290 m a.s.l., 1 dé, 8.iv.1997, B. Merz leg. (MHNG). VS: Leuk Platten, 625 m a.s.l., 1 d, 30.v.2002, B. Merz leg. (MHNG). Notes: A Holarctic species known in the literature under the junior synonym Agromyza potentillae (Kaltenbach, 1864) (Bland, 2000). The species seems to be common in Europe (Austria, Belgium, the Czech Republic, England, Estonia, Finland, Germany, Hungary, Italy (Sardinia), Lithuania, the Netherlands, Norway, Poland, Russia, Slovakia, Spain, Sweden) as well as in Central and East Palaearctic (Uzbekistan, Kuril Islands and Japan). Largely distributed also in North America (Canada and United States). The larva forms a linear mine which later widens to a conspicuous blotch on Rosaceae (Agrimonia, Alchemilla, Comarum, Filipendula, Fragaria, Geum, Potentilla, Rosa, Rubus and Sanguisorba). New for Switzerland. *Agromyza lucida Hendel, 1920 Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.1., 1 4,9.vii.2002, B. Merz leg. (MHNG). Notes: A. lucida is known from the Holarctic Region and distributed especially in Europe (Austria, the Czech Republic, Denmark, England, Finland, France, Germany, Hungary, Ireland, Italy, Latvia, Lithuania, the Netherlands, Poland, Russia, Scotland, Slovakia, Spain, Sweden) and North America (Canada). Its occurrence is also confirmed in India. The larvae form a broad blotch mine on Gramineae, known foodplants are Glycera maxima and Deschampsia caespitosa. New for Switzerland. *A gromyza marionnae Griffiths, 1963 Material examined: ZH: Zürich Albisgiitli, 500 m a.s.1., 1 6, 2.1v.1997, B. Merz leg. (MHNG). Notes: The species was described from England and later recorded from Germany, Ireland, Lithuania and Spain. According to PakalniSkis (1996) the larva mines stems of Vicia cracca and V. tetrasperma at borders of mixed forest and in urban cenoses. New for Switzerland. Agromyza mobilis Meigen, 1830 Reference: Martinez, 1998: 269. Material examined: GE: Bernex Chante Merle, 420 m a.s.l., 1 6, 7.v111.2000, B. Merz & G. Bächli leg. (MHNG). GR: Zernez, Gondas, 1480 m a.s.l., 1 dg, 8.viii.1996, B. Merz & G. Bächli leg. (MHNG). SH: Bargen-Mülital, 650 ma.s.l.,1¢, 776 M. CERNY 28.viii.1992, B. Merz leg. (MHNG). TI: Gordola, 210 m a.s.l., 1 ¢, 14.ix.1989, B. Merz leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.l., 1 à, 24.v.1996, B. Merz leg. (MHNG). Notes: A. mobilis is a species with a Palaearctic distribution, known to occur especially in Europe but also in China and Japan. It occurs namely in countries of temperate and northern Europe. In the Mediterranean area it was recorded only from Italy, Spain and the former Yugoslavia. Triticum represents the only confirmed known host plant but the larvae mine very probably also other species of Graminae. Agromyza nana Meigen, 1830 Reference: Martinez, 1998: 269. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 14.-21.vii.2000, 2 36, 23.-31.viii.2000, pasture, MT, B. Merz leg. (CMB). SH: Merishausen, Ladel, 700 m a.s.1., 1 d, 13.v.1991, B. Merz leg. (MHNG). VS: Eggerberg, 850 ma.s.l.,1 g, 24.vi.2000, B. Merz leg. (MHNG); Visperterminen Rothorn, 2250 m a.s.l., 1 26.vili.2001, B. Merz & Landry leg. (MHNG); Saas-Fee, 2400 m a.s.l., 1 22.vii.1965, 2 366, 23.vii.1965, O. Lomholdt leg. (ZMUC). Notes: The Palaearctic A. nana belong to the species of the orobi-group (Zlobin, 2000), which live in Leguminosae (Medicago sativa, Trifolium, Melilotus). With its morphology and the structure of male genitalia it is very similar to other species of this group, especially to A. frontella (Rondani, 1875), but stridulation border is species- specific (Tschirnhaus, 1971). Generally common in the Palaearctic Region including Egypt, Marocco, Tunisia, Turkey and eastern Siberia. Also recorded from India. DI 3 9 Agromyza nigripes Meigen, 1830 References: Griffiths, 1963: 132; Martinez, 1998: 269. Material examined: GE: Jussy, Prés-de-Villette, 475 m a.s.l., 1 4, 2.viii.2002, B. Merz leg. (MHNG). ZH: Zürich Albisgiitli, 470 m a.s.l., 1 4, 19.viii.1997, B. Merz & Botta leg. (MHNG); Zürich Hönggerberg, 600 m a.s.1., 1 4 ,18.vi.1992, B. Merz leg. (MHNG); Zürich Zürichberg, 650 m a.s.l., 1 4, 26.vii.1995, S. Ungricht leg. (MHNG). Notes: The species is known mainly from the Holarctic Region, (Europe and Canada), but it is also recorded from the Oriental Region (India). The larva forms a long, widening mine on the upper leaf surface of Glyceria maxima and the species also commonly occurs on Holcus. Agromyza polygoni Hering, 1941 Reference: Martinez, 1998: 269. Material examined: GR: Zuoz, Nüd, 1700 m a.s.1., 1 4, 7.viii.1996, B. Merz & G. Bächli leg. (MHNG). Notes: The species was described from Germany and recently it was also recorded from the Czech Republic, France, Lithuania, Poland and Switzerland. The larva forms on Polygonum bistorta an irregular linear-blotch mine, which is not associated to the midrib. *A gromyza prespana Spencer, 1957 Material examined: TI: Gordola, Bolle di Magadino, Zeltfalle, 220 m a.s.l., 1 d, 19.vi.1995, B. Merz & G. Bächli leg. (MHNG). AGROMYZIDAE IN SWITZERLAND TOG, Notes: Spencer (1957) described this species from Macedonia. It is also known from Austria, British Isles, Croatia (Dalmatia), the Czech Republic, France, Hungary, Poland, Slovakia and Sweden. The larva forms a broad mine on Triticum aestivum and other grasses. New for Switzerland. *A gromyza pseudoreptans Nowakowski, 1967 Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 d, 1.vii.2001; Dardagny, Roulave, 420 m a.s.l., 1 4, 30.vi.2001. GR: Valbella, Casoja, 1550 m a.s.l., 1 d,13.vil.1996. SH: Merishausen-Ladel, 600 m a.s.l., 1 2, 28.vii.1992. ZH: Flaach, Thurauen, 350 m a.s.l., 1 2, 1.ix.1993; Zürich Waldgarten, 450 m a.s.l., 2 dd, 24.v.1996; 460 m a.s.l., 1 d, 1.v.1997. All B. Merz leg. (MHNG). Notes: This Holarctic species is common in Austria, Belgium, Canary Islands, the Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Hungary, Ireland, Italy, Lithuania, the Netherlands, Norway, Poland, Russia, Scotland, Slovakia, Sweden and North America (Canada and United States, incl. Alaska). The larva forms elongate linear-blotch mine, normaly adjoining margin of leaves of Urtica dioica. New for Switzerland. *Agromyza pseudorufipes Nowakowski, 1964 Material examined: GR: Valbella, Casoja, 1550 m a.s.l., 1 d, 13.vii.1996, B. Merz leg. (MHNG). Notes: The species was described from Poland and relaible records are known from the Czech Republic, England, Germany, Lithuania, Poland, Romania, Slovakia and Japan. New for Switzerland. *Agromyza quadriseta Zlobin, 2001 Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 6, 24.v.2002. ZH: Embrach, Haumiili, 430 m a.s.l., 1 6, 29.v.1998, B. Merz leg. (MHNG). Notes: This species was recently described from Estonia, Lithuania, Finland and Russia (holotype). A. quadriseta belongs to the large group of Palaearctic species having long presutural dorsocentrals and the costal vein strongly extending to Mj 45. Biology of this species is unknown. New for Switzerland. *A gromyza rondensis Strobl, 1900 Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.l., 1 4, 4.v.2003, B. Merz & M. Eggenberger leg. (MHNG). Notes: The species was described from Spain and later on recorded in other countries of West Palaearctic (Austria, Belarus, Belgium, Canary Islands, Crete, Croatia, the Czech Republic, England, Estonia, France, Germany, Hungary, Italy, Lithuania, Norway, Poland, Portugal, Romania, Scotland, Slovakia, Sweden and Turkey). The larva forms a leaf mine on Gramineae (Arrhenaterum, Bromus, Calamagrostis, Dactylis, Hordeum, Poa, Secale and Triticum). New for Switzerland. Agromyza sulfuriceps Strobl, 1898 Reference: Martinez, 1998: 269. Material examined: VS: Saas-Fee, 2400 m a.s.1., 1 6, 22.vii.1965, O. Lomholdt leg. (ZMUC). 778 M. CERNY Notes: This small species with yellow frons and bright yellow antennae is known especially from the Holarctic Region but also recorded from tropical Africa (Nigeria). Host plants of this species are Potentilla and Sanguisorba. The larva forms a narrow upper surface gallery to start, then broadening and zigzagging to create a false blotch. *Melanagromyza astragali Spencer, 1976 Material examined: SZ: Brunni, 900 m a.s.l., 1 4, 26.vi.1990, B. Petersen leg. (ZMUC). VS: Leuk Brentjong, 930 m a.s.l., 1 4 ,3.v.1990,900 m a.s.l.,1 6, 1.vi.1991, B. Merz leg. (MHNG). Notes: This species was described by Spencer (1976) from Sweden from stems of Astragalus glyciphyllus and later on it was recorded from other countries of Europe (Czech Republic, France, Germany, Lithuania and Slovakia). New for Switzerland. Melanagromyza cunctans (Meigen, 1830) Reference: Martinez, 1998: 269. Material examined: VS: Visperterminen Dorf, 1400 m a.s.l., 1 d, 26.viii.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.l., 1 3, 16.viii.1998, B. Merz leg. (MHNG). Notes: The species was described from Spain and its known distrubution area includes other countries of temperate Europe and Scandinavia (Sweden) but it seems to be more common in the Mediterranean area (Canary Islands, Corsica, Crete, Croatia, Egypt, Italy, Maltese Islands and Turkey) and the Afrotropical Region (including Madagascar). Its occurrence was also confirmed in Central Palaearctic (Uzbekistan) and the Oriental Region (India). The larva forms a slender stem-gall on Lotus corni- culatus (Leguminosae). *Ophiomyia aeneonitens (Strobl, 1893) Material examined: VS: Leuk Pfynwald, 600-650 m a.s.l., 1 &, 15.v.1996, B. Merz & G. Bächli leg. (MHNG). Notes: The sole member of the genus with typically white squamae and fringes, and lacking the posterior crossvein on the wing. This is the second record of this species in Europe. It was described from a warm area of Austria. Biology of this species is not known. New for Switzerland. *Ophiomyia campanularum Stary, 1930 Material examined: NE: St. Blaise, Les Riedes, 470 ma.s.l., 1 4,19.v.2001, B. Merz leg. (MHNG). Notes: This species was described from the the Czech Republic and now it is known only from Germany, Slovakia and Sweden. The larva forms an external stem- mine with frass in large, widely-spaced grains on Campanula rotundifolia. New for Switzerland. *Ophiomyia cunctata (Hendel, 1920) Material examined: ZH: Zürich Oerlikon, 420 ma.s.l.,1 2, vi.-x.1987, B. Merz leg. (MHNG). AGROMYZIDAE IN SWITZERLAND 779 Notes: Widespread throughout much of Europe (Albania, Austria, Canary Islands, the Czech Republic, Denmark, England, Finland, France, Germany, Hungary, Italy, Lithuania, Norway, Poland, Russia, Scotland, Spain, Sweden, Turkey and the former Yugoslavia). The larva forms a whitish blotch mine along the midrib, with irregular offshoots into the leaf-blade on Crepis, Hypochoeris, Lapsana, Mycelis, Picris, Sonchus and Taraxacum. New for Switzerland. Ophiomyia curvipalpis (Zetterstedt, 1848) Reference: Martinez, 1998: 269. Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 3d, 24.v.2002, B. Merz leg. (MHNG); Corsier-Port, 19, 16.-31.viii.2003, C. Besuchet leg. (MHNG). TI: Biasca Loderio, 300 m a.s.l., 1 3d, 5.vii.1991, B. Merz leg. (MHNG). VS: Leuk Pfynwald, 680 m a.s.1., 1 2, 7.vi.2001, B. Merz & Landry leg. (MHNG); Leuk Platten, 625 m a.s.l., 1 5, 2.v.1999, B. Merz leg. (MHNG); St. German, Brücke, 624 m a.s.l., 1 d,3.viii.1998, B. Merz & G. Bächli leg. (MHNG). Notes: This species is distributed chiefly in the West Palaearctic but it is also known from Japan. The larva forms a narrow, inconspicuous stem-mine on Achillea millefolium, A. ptarmica, Anthemis tinctoria, Artemisia vulgaris, Matricaria inodora, and possibly also on Medicago sativa. *Ophiomyia galii Hering, 1937 Material examined: VS: Visperterminen, Kreuz, 1400 m a.s.l., 1 4, 3.vi.2003, B. Merz leg. (MHNG). Notes: The species was recorded from Corsica, the Czech Republic, England, France, Germany, Lithuania, Poland, Slovakia, and Spain. O. galii was described from Germany, and the type series was reared from Galium mollugo. The larva forms an external stem mine, with frass in large, widely spaced grains. According to Pakalniskis (1998b) it may also attack G. verum. New for Switzerland. *Ophiomyia inaequabilis (Hendel, 1931) Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 ¢, 2.vii.2001, B. Merz leg. (MHNG). Notes: This Mediterranean species was described from a single male originating from Turkey; later it was also recorded from the Czech Republic. Our record represents the westernmost boundary of the distribution area in Europe. Its biology is unknown. New for Switzerland. Ophiomyia nasuta (Melander, 1913) Reference: Martinez, 1998: 269. Material examined: SH, Merishausen, Ladel, 600 m a.s.l., 1 d, 28.11.1992, B. Merz leg. (MHNG). Notes: This species occurs in the Holarctic Region. It is known especially from temperate and northern Europe, North America (Canada, United States) and Japan. Its only known host plant genus is Taraxacum sp. Ophiomyia orbiculata (Hendel, 1931) References: Spencer, 1964: 800; Martinez, 1998: 269. 780 M. CERNY Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 6, 24.v.2002, B. Merz leg. (MHNG). NE: St. Blaise Les Riedes, 470 ma.s.l.,2 ¢ 4, 19.v.2001, B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600-650 m a.s.l., 1 4, 15.v.1996, B. Merz & G. Bächli leg. (MHNG); Leuk Platten, 630 m a.s.l., 1 d, 22.iv.1998, B. Merz & Botta leg. (MHNG). Notes: The species was described from Austria based on two males and its distribution area includes mainly temperate and northern Europe. In the Mediterranean area it is known only from the former Yugoslavia and Turkey. The larva feeds as a stem-miner on Pisum sativum, and probably also on Vicia. Ophiomyia pinguis (Fallén, 1820) References: Spencer, 1973: 140; Martinez, 1998: 269. Material examined: GR: Valbella Casoja, 1500 m a.s.l., 1 d, 19.vii.1997, B. Merz leg. (MHNG). VS: Saas-Fee, 2300 m a.s.1.,1 6,1 2, 19.vii.1965, 1800 m a.s.1.,1 9, 21.vii.1965, O. Lomholdt leg. (ZMUC). Notes: O. pinguis is known to occur in the Palaearctic Region. The species is recorded from European countries including the Mediterranean area (Italy, Spain and the former Yugoslavia). It is also confirmed from Egypt and Turkey and penetrates through Tajikistan and Uzbekistan to China. The natural hosts of this species are Cichorium intybus, C. endivia, Lactuca sativa and Leontodon. *Ophiomyia submaura Hering, 1926 Material examined: GR: Ardez, Bahnhof, 1350 m a.s.l., 1 gd, 6.viii.1996, B. Merz & G. Bächli leg. (MHNG). Notes: O. submaura occurs in the West Palaearctic. It is confirmed from the Czech Republic, Germany, Hungary, Lithuania, Poland, Spain and Turkey. Pakalniskis (1996) reared it from Medicago falcata at borders of mixed forests and in brighter pine woods. New for Switzerland. *Ophiomyia vimmeri Cerny, 1994 Material examined: GR: Valbella, Casoja, 1500 m a.s.l., 1 d, 14.vii.1998, B. Merz leg. (MHNG). Notes: O. vimmeri was described from the Czech Republic, it is also known to occur in Slovakia. Its biology is unknown. New for Switzerland. Subfamily PHYTOMYZINAE *Amauromyza (Cephalomyza) flavifrons (Meigen, 1830) Material examined: SH: Rüdlingen, 370 m a.s.l., 1 3d, 7.1x.1996, B. Merz leg. (MHNG). TI: Gordola, Bolle, 220 m a.s.., 1 gd, 1.vii.1993, B. Merz & M. Eggenberger leg. (MHNG). ZH: Zürich Oerlikon, 430 m a.s.l., 1 d, 23.v.1992, B. Merz leg. (MHNG). Notes: This Holarctic species is distributed in Alabania, Austria, Belgium, Corsica, the Czech Republic, Denmark, England, Finland, France, Germany, Hungary, Kyrgyzstan, Lithuania, the Netherlands, Norway, Poland, Romania, Sardinia, Scotland, Spain, Sweden, Turkey and North America (Canada and United States). The larva AGROMYZIDAE IN SWITZERLAND 781 forms a white linear-blotch mine on many genera of Caryophyllaceae, particularly on Dianthus, Lachnis, Melandrium, Saponaria, Silene, Stellaria but also on Beta vulgaris and Spinacia oleracea (Chenopodiaceae). New for Switzerland. *Amauromyza (Cephalomyza) mihalyii Spencer, 1971 Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.1.,2 4 4 ,19.v.2001, B. Merz leg. (MHNG). Notes: This species was described from Hungary and later recorded from the Czech Republic. This record is a further evidence of A. (C.) mihalyii in Central Europe and represents the westernmost boundary of its occurrence. Its biology is unknown. New for Switzerland. Amauromyza (Cephalomyza) monfalconensis (Strobl, 1909) References: Spencer, 1992: 142; Martinez, 1998: 269. Material examined: GR: Lenzerheide Sundroina, 1600 m a.s.l., 1 à, 16.vi1.2000, B. Merz leg. (MHNG). Notes: This species is distributed especially in temperate Europe. It is also recorded from Scandinavia (Sweden) and some countries of the Mediterranean area (Italy and Spain). Host plants and immature stages unknown; larva almost certainly feeds as internal stem-borer. *Aulagromyza luteoscutellata (de Meijere, 1924) Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 6, 25.viii.2001, B. Merz & Landry leg. (MHNG). Notes: The species is distributed in temperate and northern Europe (Belgium, the Czech Republic, Denmark, Finland, France, Germany, Lithuania, the Netherlands, Norway, Poland, Sweden) and Canada. It has not been recorded from the Mediterranean area, but it is known from India. The larva forms a short, funnel-shaped leaf mine, filled centrally with dark-green frass on Lonicera and Symphoricarpos. New for Switzerland. *Aulagromyza orphana (Hendel, 1920) Material examined: ZH: Zürich Katzensee, 440 m a.s.l., 1 d, 25.v.1999, B. Merz leg. (MHNG). Notes: Locally distributed particularly in Central Europe. Its occurrence is confirmed for Austria, Belgium, the Czech Republic, Denmark, England, France, Germany, Hungary, the Netherlands, Poland, Scotland, Slovakia, Spain and Turkey. The larva forms an external stem mine on Galium aparine, probably also on G. palustre. New for Switzerland. *Aulagromyza similis (Brischke, 1880) Material examined: SH: Merishausen, 550 m a.s.l., 1 2, 26.iv.1990, B. Merz leg. (MHNG). Notes: A European species occurring in Austria, the Czech Republic, England, Finland, France, Germany, Hungary, Ireland, Italy, Lithuania, Norway, Poland, Romania, Sweden and Wales. The larva forms a large whitish linear blotch leaf mine, 782 M. CERNY frequently associated with the midrib on Knautia arvensis and Succisa pratensis. New for Switzerland. Aulagromyza trivittata (Loew, 1873) Reference: Martinez, 1998: 270. Material examined: ZH: Zürich Albisgiitli, 490 m a.s.l., 1 & ,9.iv.1997, B. Merz leg. (MHNG). Notes: This species is distributed mainly in temperate Europe. In Scandinavia it is recorded from Norway and Sweden. It is not known from the Medirranean area. The larva feeds as internal stem borer on Galium mollugo. Calycomyza humeralis (von Roser, 1840) Reference: Martinez, 1998: 270. Material examined: GR: Lenzerheide, 2000 m a.s.l.,3 à à , viii.2000, pasture, MT, B. Merz leg. (CMB). VS: Leuk Platten, 630 m a.s.l., 1 4, 22.iv.1998, B. Merz & Botta leg. (MHNG); Visperterminen Giw-Gebidemsee, 1900-2200 m a.s.l., 1 à, 28.v111.2001, B. Merz & Landry leg. (MHNG). Notes: A semicosmopolitan species which is locally distributed in Europe but also known from East Palaearctic, North and South America, the Afrotropical and the Oriental Regions, Australia and the Hawaiian Islands. The larva forms a leaf mine on Aster, Bellis and Erigeron. *Cerodontha (Butomomyza) angulata (Loew, 1869) Material examined: ZH: Zürich Zürichberg, 500-600 m a.s.., 3 dd, 16.viii.1992, B. Merz leg. (MHNG). Notes: The species is distributed in the Holarctic Region including Japan and Kuril Islands. The larva forms a long greenish leaf mine on Carex, rarely on Scirpus silvaticus. New for Switzerland. *Cerodontha (Butomomyza) rohdendorfi Nowakowski, 1967 Material examined: ZH: Embrach, Haumiili, 400 m a.s.l., 1 d, 28.v.1997, B. Merz leg. (MHNG). Notes: Described from Poland and later recorded from the Czech Republic, Finland and Italy. The larva forms an upper or lower surface mine, filling the entire width of the leaf of Poa chaixii, Festuca pratensis and probably other related grasses. New for Switzerland. Cerodontha (Cerodontha) affinis (Fallén, 1823) References: Spencer, 1976: 177; Martinez, 1998: 270. Material examined: ZH: Horgen, 550 m a.s.l., 1 dé, 24.v.1989, B. Merz leg. (MHNG). Notes: The species is common in Europe, particularly in the temperate zone. Its biology is unknown. Cerodontha (Cerodontha) denticornis (Panzer, 1806) Reference: Martinez, 1998: 270. Material examined: GR: Lenzerheide, 1600 ma.s.l., 1 £,8.x.1991, B. Merz & M. Eggenberger leg. (MHNG); Lenzerheide, 2000 m a.s.l., 1 6, 23.viii.-9.ix.2000, AGROMYZIDAE IN SWITZERLAND 783 pasture, MT, B. Merz leg. (CMB). SH: Merishausen, Grate, 520 m a.s.l., 1 d, 1.vi.1996, B. Merz & M. Eggenberger leg. (MHNG). VS: Saas-Fee, 1800 ma.s.l.,1 d, 3 22, 21.vii.1965, O. Lomholdt leg. (ZMUC). TI: Lavertezzo, 560 m a.s.l., 1 9, 12.ix.1989, B. Merz leg. (MHNG). ZH: Zürich Allmend, 460 m a.s.l., 2 dû, 17.v.1995; Zürich Irchel, 500 m a.s.l., 1 4 , 24.v.1996; Zürich Katzensee, 440 m a.s.l., 1 8, 25.v.1996, B. Merz leg. (MHNG). Notes: A common Palaearctic species described from Germany. Known to occur also in the Afrotropical and Oriental Regions including Taiwan. The larva feeds mainly in the leaf sheath of Gramineae. Cerodontha (Cerodontha) fulvipes (Meigen, 1830) References: Spencer, 1976: 177, 180; Martinez, 1998: 270. Material examined: GL: Pragelpass, 1500 m a.s.l., 1 d, 5.viii.1991, G. Bachli, Beuk & B. Merz leg. (MHNG). GR: Lenzerheide, 2000 m a.s.l., 1 3, viii.2000, pasture, MT, B. Merz leg. (CMB). TI: Gordola, Bolle d. M., 205 m asl.,2 dd, 6.v11.2001, B. Merz leg. (MHNG). VS: Baltschieder, Rotten-Ufer, 650 m a.s.l.,1 4, 19.v.1996, B. Merz leg. (MHNG); Saas-Fee, 2000 m a.s.1.,1 6,1 2, 18.vii.1965, 2300 ma.s.1.,2 2 9, 19.vii.1965, 1800 ma.s.1., 1 £,21.vii.1965, O. Lomholdt leg. (ZMUC). ZH: Embrach Haumiili, 400 m a.s.l.,3 646, 24.vi.1995, 1 3, 19.vi.1998, Zürcher leg. (MHNG); Zürich Irchel, 500 m a.s1., 1 3, 16.viii.1998, B. Merz leg. (MHNG). Notes: A Palaearctic species which is generally distributed in Europe and recorded also from China. Poa trivialis is known as a host plant but the larva feeds probably on other grasses as well. *Cerodontha (Cerodontha) unguicornis Hendel, 1932 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 23.-31.viii.2000, pasture, MT, B. Merz leg. (CMB); Valbella, Casoja, 1600 m a.s.1., 1 9, 18.vii.2000, B. Merz leg. (MHNG). VS: Saas-Fee, 1800 m a.s.l., 1 ¢, 21.vii.1965, O. Lomholdt leg. (ZMUC). Notes: The species was described from Central Europe. It is known from Austria, the Czech Republic and Germany. Its biology is still unknown. New for Switzerland. *Cerodontha (Dizygomyza) bimaculata (Meigen, 1830) Material examined: BE: Tramelan, La Tourbière, 995 m a.s.l., 1 4, 4.vi.2003, Merz, Haenni & Rapp leg. (MHNG). GR: S. Vittore, Rebberg, 300 m a.s.l., 1 g, 4.v111.1997, B. Merz leg. (MHNG). TI: Biasca, 350 m a.s.l., 1 4, 18.v.1991, B. Merz leg. (MHNG); Mte. Caslano, 400 m a.s.l., 1 4, 20.v.1991, B. Merz leg. (MHNG). VS: Morgins, Portes du Soleil (Monthey), 1700-1950 m a.s.l., 1 4, 22.vi.2003, B. Merz leg. (MHNG). Notes: This species is common in Europe. It is also known from the Kuril Islands and Japan. The larva forms a narrow mine on Luzula, particularly on L. pilosa. New for Switzerland. *Cerodontha (Dizygomyza) crassiseta (Strobl, 1900) Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 3, viii.2000, pasture, MT, B. Merz leg. (CMB). 784 M. CERNY Notes: A European species which is distributed especially in temperate Europe. In Fennoscandia known only from Sweden, and in the Mediterranean area recorded only from Italy and Spain. Dactylis glomerata and Poa compressa are known as host plants of this species. New for Switzerland. *Cerodontha (Dizygomyza) fasciata (Strobl, 1880) Material examined: VS: Leuk Pfynwald, 630 m a.s.l., 1 6, 21.iv.1998, B. Merz & Botta leg. (MHNG); Visperterminen Dorf, 1400 ma.s.l., 1 3, 26.viii.2001, B. Merz & Landry leg. (MHNG). Notes: A species with a Holarctic type of distribution, known from temperate and northern Europe. It has not been recorded from the Mediterranean area. In North America occurring in Canada and the United States. Its biology is unknown. New for Switzerland. *Cerodontha (Dizygomyza) griffithsi Nowakowski, 1967 Material examined: VS: Saas-Fee, 2000 m a.s.l., 1 d, 22.vii.1965, 1 6, 27.vii.1965, O. Lomholdt leg. (ZMUC). Notes: This species was described from one male originating from the Austrian Alps. Later it was found in France. Its discovery in the Swiss Alps is thus not surprising. Recently it was captured in the High Tatra Mts in Slovakia (Cerny & Vala, 2005). Its biology is unknown. New for Switzerland. *Cerodontha (Dizygomyza) luctuosa (Meigen, 1830) Material examined: GR: Zernez, Gondas, 1480 m a.s.l., 1 d, 4.viii.1996, B. Merz & G. Bächli leg. (MHNG). Notes: The species occurs in the Holarctic Region. It is known particularly from Europe, but also from Tunisia, China, Japan and North America (Canada, United States, incl. Alaska). The larva feeds on Juncus effusus. New for Switzerland. *Cerodontha (Dizygomyza) luzulae (Groschke, 1957) Material examined: ZH: Zürich Albisgiitli, 450 m a.s.l., 1 4, 1.v.1995, B. Merz leg. (MHNG). Notes: This European species was recorded from some Central European countries (the Czech Republic, Germany, Hungary and Poland) and the British Isles. Luzula sylvatica is the only known host plant of C. (D.) luzulae. New for Switzerland. Cerodontha (Dizygomyza) morosa (Meigen, 1830) References: Hendel, 1931-6: 92; Martinez, 1998: 270. Material examined: GR: S. Vittore Rebberg, 300 m a.s.l., 1 d, 4.viii.1997, B. Merz leg. (MHNG). Notes: This Holarctic species is common in Europe and known to occur also in the East Palaearctic (China and Japan) and the Oriental Regions (Philippines). The larva forms a relatively short and broad mine, which not extends to the leaf base of Carex. *Cerodontha (Dizygomyza) spinata (Groschke, 1954) Material examined: VS: Visperterminen Rothorn, 2250 m a.s.., 2 dd, 26.viii.2001, B. Merz & Landry leg. (MHNG). AGROMYZIDAE IN SWITZERLAND 785 Notes: The species is known from Austria, the British Isles, the Czech Republic, Germany and Poland. The larva feeds on Carex sylvatica and C. pilosa. New for Switzerland. *Cerodontha (Poemyza) alpina Nowakowski, 1967 Material examined: GR: Lenzerheide, St. Cassian, 1450 m a.s.l., 1 à, 12.vii.1998, B. Merz leg. (MHNG). Notes: This mountain species was described from the upper forest zone of the East Carpathians (Poland) and later it was found in Austria. Our record of C. (P.) alpina in the Swiss Alps could be expected. The larva forms a leaf mine on Poa alpina and Trisetum alpestre. New for Switzerland. Cerodontha (Poemyza) atra (Meigen, 1830) References: Hendel, 1931-6: 38; Nowakowski, 1973: 91; Martinez, 1998: 270. Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 6, 24.v.2002, B. Merz leg. (MHNG). Notes: This species is common in Europe and known to occur in Japan. Some species of Gramineae are host plants of this species, in particular Calamagrostis epigeios and Agrostis alba. *Cerodontha (Poemyza) beigerae Nowakowski, 1973 Material examined: GE: Russin, Les Baillets, 405 m a.s.l., 1 d, 30.vi.2001, B. Merz leg. (MHNG). Notes: Originally known only from Central Europe (the Czech Republic, Germany, Hungary and Poland). Recently Zlobin (1986, 1992) recorded C. (P.) beigerae also from East Siberia and the Far East. The larva feeds on Calamagrostis canescens, C. arundinacea, C. villosa and Agrostis canina. New for Switzerland. *Cerodontha (Poemyza) calamagrostidis Nowakowski, 1967 Material examined: GE: Bernex, Chante-Merle, 415 m a.s.1.,1 4, 20.v.2002, B. Merz leg. (MHNG). Notes: This Holarctic species is distributed in Europe but it has not been found in the Mediterranean area. In North America known from Canada. Calamagrostis sp. and Alopecurus pratensis are the known host plants. New for Switzerland. *Cerodontha (Poemyza) incisa (Meigen, 1830) Material examined: ZH: Zürich Zürichberg, 450-650 m a.s.l., 1 4, 20.viii.1992, B. Merz leg. (MHNG). Notes: This Holarctic species is common in Europe but also known from China, Pakistan, Japan and North America. The larva forms a leaf mine on Gramineae, most commonly on Agropyron, Calamagrostis, Festuca and Phalaris but Nowakowski (1973) recorded 17 additional European genera. New for Switzerland. *Cerodontha (Poemyza) lateralis (Macquart, 1835) Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 23.-31.viii.2000, pasture, MT, B. Merz leg. (CMB). VS: Branson, Follatères, 450-950 m a.s.l., 1 à, 29.111.2002; Leuk Pfynwald, 630 m a.s.l., 1 6, 25.v.1997, 600 m a.s.l., 1 &,2.vii.2001. All B. Merz leg. (MHNG). 786 M. CERNY Notes: This Palaearctic species is common in the whole of West Palaearctic and penetrates to North Africa (Tunisia), Turkey, China and Japan. The larva forms a linear mine running towards the apex of the leaf and this can widen and become almost blotch-like on Gramineae. New for Switzerland. *Cerodontha (Poemyza) lyneborgi Spencer, 1972 Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 gd, 24.iv.2003, B. Merz & F. Amiet leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 ¢, 30.vi.2001, B. Merz leg. (MHNG). Notes: This species was described by Spencer (1972a) from Spain and later on it was found in other countries of Europe (the Czech Republic, Hungary, Italy, Lithuania and Russia). Its biology is unknown. New for Switzerland. Cerodontha (Poemyza) muscina (Meigen, 1830) Reference: Martinez, 1998: 270. Material examined: GR: Lenzerheide, 2000 m a.s1., 2 à d, viii.2000, pasture, MT, B. Merz leg. (CMB); Lenzerheide, gegen Sanaspans, 1500 m as.l., 1 à, 7.vili.1992, B. Merz leg. (MHNG). VS: Visperterminen Dorf, 1400 m a.s.l., 1 à, 26.vi1i.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich Katzensee, 450 m a.s.l., 1 g, 1.vii.1995; Zürich Ziegelhütte, 460 m a.s.l., 1 d, 29.vi.1997, B. Merz leg. (MHNG). Notes: This Holarctic species is common in Europe including the Mediterranean area (Spain, Italy and Slovenia), but also in North America. The larva forms a leaf mine on Gramineae. Mines were recorded on Dactylis, Festuca, Holcus, Milium and Poa. *Cerodontha (Poemyza) pygmaea (Meigen, 1830) Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 23.-31.viii.2000, pasture, MT, B. Merz leg. (CMB). TI: Biasca, Loderio, 350 m a.s.l., 1 dg, 26.11.1998, B. Merz leg. (MHNG). Notes: This species from the incisa-group shows a Holarctic type of distribution and it is well known from a greater part of Europe. The larva forms a leaf mine on Gramineae. New for Switzerland. *Cerodontha (Poemyza) spencerae Zlobin, 1993 Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.1.,4 dd, 16.v1.1997, B. Mocek leg. (CBM). Notes: This Holarctic species is based on the type series originating from Russia, Estonia and Kazakhstan. In the Palaearctic Region C. (P.) spencerae was recorded from the British Isles, the Czech Republic, Finland, Italy, Latvia, Lithuania, Slovakia, East Siberia and the Russian Far East incl. Sakhalin. In Europe it was once bred from Agropyron repens. New for Switzerland. *Cerodontha (Xenophytomyza) atronitens (Hendel, 1920) Material examined: VS: Leuk Pfynwald, 600-650 m a.s.l., 1 4, 19.v.1996, B. Merz & G. Bachli leg. (MHNG). AGROMYZIDAE IN SWITZERLAND 787 Notes: This species is distributed in temperate and northern Europe, but not yet found in the Mediterranean area. Host plants are unknown but certainly it is an internal feeder in the leaf-sheath or stem of Gramineae. New for Switzerland. Cerodontha (Xenophytomyza) biseta (Hendel, 1920) Reference: Martinez, 1998: 270. Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.l., 1 ¢,9.vii.2001, B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 d, 6.vi.2001, 1 à, 7.vi.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich Hönggerberg, 530 m a.s.l., 1 3, 9.vi.1998; Zürich Waldgarten, 460 m a.s.l., 1 4, 7.vi.1997; Zürich Ziegelhütte, 460 m a.s.l., 2 d d, 29.vi.1997, B. Merz leg. (MHNG). Notes: C. (X.) biseta shows a Holarctic distribution and it is known to occur particularly in Europe including the Mediterranean area but also in Japan and North America (Canada and United States). The single specimen of C. (X.) biseta collected from the Jamaican Blue Mountains represents the first Neotropical record (Boucher, 2003). Host plants unknown, certainly an internal feeder in Gramineae. *Chromatomyia fuscula (Zetterstedt, 1838) Material examined: GR: Lenzerheide, 2000 m a.s.l., pasture, 1 d, 23. 31.viii.2000, MT, B. Merz leg. (CMB). SZ: Unteriberg, O. Weid, 1600 m a.s.l., 1 à, 24.vii.2001. TI: Biasca, Loderio, 350 m a.s.l., 1 6, 15.i11.1994, 1 6, 23.iii.1996, 1 à, 26.111.1998. VS: Branson, Follatères, 450-950 m a.s.1., 2 & d, 29.111.2002, B. Merz leg. (MHNG); Leuk Pfynwald, 600 m a.s.l., 1 d, 6.vi.2001, B. Merz & Landry leg. (MHNG); Visperterminen Dorf, 1550 m a.s.l., 1 d, 27.viii.2001, B. Merz & Landry leg. (MHNG). Notes: Ch. fuscula has a Holarctic distribution. In Europe this species is known to occur chiefly in temperate and northern parts. Also recorded in the East Palaearctic: Russia (Asian part), Kamchatka and Japan. The larva forms leaf-mines on Gramineae. New for Switzerland. Chromatomyia gentianella (Hendel, 1932) References: Hendel, 1931-6: 311; Spencer, 1990: 396; Martinez, 1998: 270. Material examined: GR: Lenzerheide, 2000 m a.s.l., 2 6 d, 21-31.viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This mountain species is only known from Central Europe (Austria, North Italy, Poland and Switzerland). Larva forms a linear mine on Gentiana. Chromatomyia hoppiella Spencer, 1990 References: Spencer, 1990: 396; Martinez, 1998: 270. Material examined: GR: Ausserferrera, 1300 m a.s.l., 1 Sd, 15.viii.1991, B. Merz & A. Freidberg leg. (MHNG); Lenzerheide, 2000 m a.s.l., 1 d, viii.2000, pasture, MT, B. Merz leg. (CMB); Valbella Casoja, 1500 m a.s.l., 1 4, 19.vii.2000, B. Merz leg. (MHNG). Notes: Ch. hoppiella was described recently from Switzerland (Lenzerheide), based on the male holotype which was bred from Gentiana excisa. Our records represents the first additional faunistic data after the description. 788 M. CERNY Chromatomyia horticola (Goureau, 1851) Reference: Martinez, 1998: 270. Material examined: VS: Branson Follatéres, 450-950 m a.s.1., 1 6, 29.11.2002, B. Merz leg. (MHNG). ZH: Zürich Oerlikon, 430 m a.s.l., 1 9, 23.v.1992, leaf mine ex Centaurea sp., B. Merz leg. (MHNG). Notes: A polyphagous species, common in the Palaearctic, Afrotropical and Oriental Regions. Ch. horticola is a well known and common species mining a large spectrum of host plants, Griffiths (1967) and Spencer (1973) recorded host plants belonging to 34 families. The larva forms a long whitish upper surface corridor, which may go to the lower surface. *Chromatomyia isicae (Hering, 1962) Material examined: TI: Biasca, Loderio, 300 m a.s.l., 1 4, 9.iv.1995, B. Merz leg. (MHNG). Notes: This species was described from Austria. It is also recorded from the Czech Republic, Finland, Germany, Ireland, Lithuania, Norway and Sweden. Its biology is unknown. New for Switzerland. Chromatomyia milii (Kaltenbach, 1864) Reference: Martinez, 1998: 270. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 3, viii.2000, pasture, MT, B. Merz leg. (CMB); Valbella Casoja, 1550 m a.s.l., 1 dg, 13.vii.1996, 1 gd, 15.vii.1996, B. Merz leg. (MHNG). TI: A. d’ Arena, 1700 m a.s.1., 1 3, 20.vii.1983, C. Besuchet leg. (MHNG); Biasca Loderio, 350 m a.s.l., 1 4, 23.i11.1996, 300 m ass. 1 3, 9.iv.1995, B. Merz leg. (MHNG); Gordola, Bolle d. M., 205 m a.s.l., 1 6, 6.vii.2001, B. Merz leg. (MHNG). ZH: Embrach Haumiili, 430 m a.s.l., 1 à, 3.vii.1996, B. Merz leg. (MHNG); Zürich Albisgiitli, 450 m a.s.l., 1 4, 27.11.1994, 500 m a.s.l., 1 dg, 2.iv.1997, B. Merz leg. (MHNG); Zürich Allmend, 450 m a.s.1.,2 66, 30.11.1994, B. Merz leg. (MHNG); Zürich Katzensee, 440 m a.s.l., 1 4, 19.iv.1997, B. Merz leg. (MHNG); Zürich, 1 ¢, 26.vii.1987, B. Merz leg. (MHNG). Notes: A Holarctic species which is common in the West Palaearctic and in North America (Canada) but it is also recorded from India. This species feeds in the larval stage on a variety of soft-leaved wild grasses in mesic and woodland habitats. It is not a pest of cultivated cereals, and large marsh grasses, such as Phragmites; Phalaris and Glyceria are also apparently avoided. Chromatomyia nigra (Meigen, 1830) Reference: Martinez, 1998: 270. Material examined: GR: Lenzerheide Sundroina, 1550 m a.s.l., 1 dg, 18.vii.1997, B. Merz leg. (MHNG). SG: Chäserrugg, 1850 m a.s.l., 1 G , 15.viii.1997, B. Merz leg. (MHNG). Notes: A Holarctic species. Its distribution area in the Palaearctic and Nearctic Regions seems to be larger than in the related Ch. milii. It was also recorded in the Oriental Region. The larva forms a narrow, whitish linear mine on many genera of Gramineae. AGROMYZIDAE IN SWITZERLAND 789 *Chromatomyia opacella (Hendel, 1935) Material examined: GR: Lenzerheide, Sanaspans, 2150 m a.s.l., 1 à, 14.vii.1996, B. Merz leg. (MHNG). Notes: This mountain species is distributed up to the subalpine and alpine zones of the Polish Tatra Mts. It penetrates far to the North (Iceland and Faroes Islands). Recently recorded from Central Europe (Cerny & Vala, 2005), known also from the British Isles and North Italy. One record from Greenland is mentioned by Griffiths (1980). Immature stages and the mine are unknown. Sesleria tatrae and Trisetum fuscum are recorded as host plants. Our record extends its range of alpine localities from Austria and Italy to Switzerland. New for Switzerland. *Chromatomyia pseudomilii Griffiths, 1980 Material examined: VS: Visperterminen Dorf, 1550 m a.s.l., 1 4, 27.viii.2001, B. Merz & Landry leg. (MHNG). Notes: This mountain species was described from Germany; it is also known from mountains of the Czech Republic (Cerny & Vala, 1996), from the High Tatra Mts in Slovakia (Cerny & Vala, 2005) as well as from North America (United States and Alaska). Its occurrence in further European mountains is very likely. The larva forms leaf mines on Gramineae and Carex. New for Switzerland. Chromatomyia ramosa (Hendel, 1923) References: Spencer, 1992: 142; Martinez, 1998: 270. Material examined: JU: Alle, 450 m a.s.l., 1 3d, 6.1x.1989, B. Merz leg. (MHNG). ZH: Volketswil, Homberg, 500 m a.s.l., 1 d, 7.vi.1995, B. Merz leg. (MHNG). Notes: A common species in Europe which was also recorded from China. The larva feeds primarily along midrib, forming short offshoots into the leaf-blade in Dipsacus, Knautia and Succisa. *Galiomyza galiivora (Spencer, 1969) Material examined: GE: Jussy, Prés-de-Villette, 475 m a.s.l., 1 9, 2.viii.2002, B. Merz leg. (MHNG). Notes: G. galiivora was described by Spencer (1969) from specimens bred from Galium in the United States. It is also recorded in Canada and Europe (Belgium, Corsica, the Czech Republic, Germany, Lithuania, Poland and Slovakia). New for Switzerland. Galiomyza morio (Brischke, 1880) Reference: Martinez, 1998: 270. Material examined: NE: St. Blaise Les Riedes, 470 m a.s.l., 1 à, 19.v.2001, B. Merz leg. (MHNG). Notes: Also this species is mining Galium, but it was also bred from Asperula odorata (= Galium odoratum). Its distribution area includes Europe and Japan. The larva forms a narrow linear mine which may largely fill small leaves and produce a secondary blotch. 790 M. CERNY *Liriomyza approximata (Hendel, 1920) Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 &, 21-31.viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This species of mixed forests is known from Central and Northern Europe where it was found in Austria, Finland, Germany, Hungary, Northern Italy and Poland. The larva forms an upper surface blotch mine on Daphne mezereum. New for Switzerland. Liriomyza congesta (Becker, 1903) Reference: Martinez, 1998: 270. Material examined: SH: Merischausen, Ladel, 700 m a.s.l., 1 ¢, 13.v.1991, B. Merz leg. (MHNG). ZH: Zürich Katzensee, 440 m a.s.l., 1 à ,25.v.1996, B. Merz leg. (MHNG). Notes: Generally distributed in the West Palaearctic but apparently ranging to Japan. Common in the Mediterranean area (Albania, Corsica, Egypt, Italy incl. Sardinia, Maltese Islands, Spain and Turkey). The larva forms an upper surface linear mine exclusively on Leguminosae, with many genera being attacked, in particular Medicago, Lathyrus, Pisum and Vicia. *Liriomyza erucifolii de Meijere, 1944 Material examined: TI: Biasca, Loderio, 300 m a.s.l., 1 &, 9.iv.1995, B. Merz leg. (MHNG). Notes: This species is based on the male holotype from France and later recorded from the Czech Republic, Denmark, England, the Netherlands and Poland. Recently found on the Maltese Islands (Cerny, 2005b). The larva forms a relatively long linear mine, normally beginning near the apex of the leaf on Senecio erucufolius and S. jacobaea. New for Switzerland. Liriomyza flaveola (Fallén, 1823) Reference: Martinez, 1998: 271. Material examined: GE: Chancy, La Laire, 350 m a.s.l., 2 dd, 24.iv.2003, B. Merz & F. Amiet leg. (MHNG). ZH: Zürich, 600 m a.s.1.,1 4, 10.vi.1991, B. Merz leg. (MHNG). Notes: A common European species but known to occur also in Turkey, eastern Russia (East Siberia, Far East, Kamchatka, Kuril Islands), Uzbekistan, Japan and India. The larva forms a narrow, whitish leaf mine on Gramineae, occurring on many genera, particularly Bromus, Dactylis, Holcus and Poa, also on cultivared Avena sativa and Hordeum vulgare. *Liriomyza lutea (Meigen, 1830) Material examined: VS: Visperterminen, 1460 m a.s.l., 1 6, 23.vii.1992, B. Merz leg. (MHNG). Notes: Widespread in Europe but local, frequently occurring in large numbers in association with the food-plants. The larva feeds in individual seeds on Angelica sylvestris, Heracleum sphondylium and Pastinaca sativa, probably also on Laserpitium. New for Switzerland. AGROMYZIDAE IN SWITZERLAND 791 *Liriomyza obliqua Hendel, 1931 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 ¢, 21-31.viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: L. obliqua is distributed in temperate Europe and in the Mediterranean area (Austria, Bulgaria, Germany, Lithuania, Romania, Slovakia, former Yugoslavia and Ukraine). Its biology is unknown. New for Switzerland. Liriomyza polygalae Hering, 1927 Reference: Martinez, 1998: 271. Material examined: GR: Lenzerheide, 2000 m a.s1., 2 d à , viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: The species was described by Hering (1927) from Lugano (TI) in Switzerland. It is known to occur mainly in Central Europe but also in England. This is the second record from Switzerland. The larva forms an irregular linear-blotch mine on Polygala vulgaris. *Liriomyza taraxaci Hering, 1927 Material examined: VS: Leuk Pfynwald, 630 m a.s.l., 2 dd, 23.iv.1998, B. Merz & Botta leg. (MHNG). Notes: A Holarctic species which is common in Europe, but known only from Slovenia and Spain in the Mediterranean area. In North America recorded from Canada and United States. The larva forms a somewhat irregular, elongate blotch leaf mine on Taraxacum. New for Switzerland. *Metopomyza flavonotata (Haliday, 1833) Material examined: VS: Leuk Platten, 625 m a.s.., 1 d, 30.v.2002; Visperterminen Kreuz, 1400 m a.s.l., 1 ¢, 3.vi.2003, B. Merz leg. (MHNG). Notes: This is a type species of the genus Metopomyza. It is common in Europe as well as in Russia (incl. Siberia and Far East) and Japan. Its biology is unknown, Pakalniskis (1998a) recorded Deschampsia caespitosa as a host plant for the first time. New for Switzerland. *Metopomyza nigriorbita (Hendel, 1931) Material examined: GE: Bernex, Chante-Merle, 415 m a.s.1., 1 4, 16.viii.2002, B. Merz leg. (MHNG). Notes: A Palaearctic species which is common in Europe but not known in southern parts. Recorded also from Japan. The puparium is dark brown, with posterior spiracles each having a bunch of about 6 irregular bulbs (Spencer, 1976) but the host plant is unknown. New for Switzerland. *Metopomyza scutellata (Fallén, 1823) Material examined: GE: Jussy, Prés-de-Villette, 475 m a.s.l., 1 d, 2.vii.2002, B. Merz leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 &, 1.vii.2002, B. Merz & Herrmann leg. (MHNG). VS: Leuk Pfynwald, 630 m a.s.l., 1 4, 6.vi.1997, B. Merz leg. (MHNG), 625 m a.s.l., 1 6, 16.v.2000, B. Merz & Ulrich leg. (MHNG). ZH: 792 M. CERNY Zürich, 500 m a.s.l.,2 od, 10.viii.1991, B. Merz leg. (MHNG); Zürich Waldgarten, 460 ma.s.l., 1 8, 30.vii.1997, B. Merz leg. (MHNG). Notes: A Palaearctic species which is, in addition to Europe, also recorded from Kazakhstan, Kyrgyzstan, eastern Russia (East Siberia, Far East) and Japan. Species of Carex are host plants. New for Switzerland. *Metopomyza xanthaspis (Loew, 1858) Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.1.,2 & d,19.v.2001, B. Merz leg. (MHNG). VS: Grächen, 1450 m a.s.l., 1 ¢, 5.vi.1987, B. Merz leg. (MHNG); Leuk Pfynwald, 630 m a.s.l., 1 d, 6.vi.2001, B. Merz & B. Landry leg. (MHNG). Notes: A common species in Europe, ranging to the Mediterranean area (Croatia, Spain, Tunisia), Kazakhstan, Asian Russia and Japan. The larvae feed as leaf miner on Carex humilis. New for Switzerland. *Napomyza bellidis Griffiths, 1967 Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 d, 18.1v.2003, B. Merz leg. (MHNG). Notes: This species is based on a type series reared from Bellis perennis in England and later recorded from further European countries (Austria, Belgium, England, Estonia, France, Germany, Hungary, Italy, Latvia, Lithuania, Maltese Islands, Portugal, Russia, Sweden and Ukraine) and Central and East Palaearctic (Uzbekistan, Japan). New for Switzerland. Napomyza cichorii Spencer, 1966 Reference: Martinez, 1998: 271. Material examined: VS: Saas-Fee, 2400 m a.s.l., 1 4, 22.vii.1965, O. Lomholdt leg. (ZMUC). Notes: This species was described from a pair reared from Cichorium intybus originating from the Netherlands and one male from an alpine meadow in Switzerland (VS: Orsiéres). Our male confirms the occurrence in the canton Valais. It is distributed mainly in temperate Europe, the Mediterranean region and penetrates to Armenia, East Siberia, Far East and Japan. The larvae feed in stems and roots of Cichorium intybus and C. endivia (Asteraceae). *Napomyza elegans (Meigen, 1830) Material examined: GR: Valbella, 1500 m a.s.l., 1 2, 28.vii.1999, leg. B. Merz & Müller (MHNG). Notes: A European species which is more common in temperate and northern Europe. Host plants and early stages are unknown but flies are not infrequently caught on flowers of Valeriana officinalis and it is believed the larva may feed in the root of this plant. New for Switzerland. *Napomyza maritima von Tschirnhaus, 1981 Material examined: TI: Biasca, 350 m a.s.l., 1 dg, 18.v.1991, B. Merz leg. (MHNG). AGROMYZIDAE IN SWITZERLAND 793 Notes: This species was described from Germany and France and it is also recorded from the Czech Republic, Finland, Hungary, Kazakhstan, Russia, Slovenia and Uzbekistan. The larva feeds in stems of Artemisia maritima. This plant does not occur in Switzerland. Other species of Artemisia, like A. campestris and A. vulgaris, both common in Biasca, may be infested by this species. New for Switzerland. *Napomyza nigriceps van der Wulp, 1871 Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 4, 1.1v.2002, 1 6, 18 .iv.2003, B. Merz leg. (MHNG). Notes: The species is known from Western, Central and Northern Europe, but it is not recorded in the Mediterranean area. Its biology is unknown. New for Switzerland. *Napomyza tripolii Spencer, 1966 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 &, 21-31.viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: The type series of this species was caught on Aster tripolium in England. Later records includes localities in Denmark, Germany, Ireland and Spain. The larva feeds in the stem of Aster tripolium. This plant does not occur in Switzerland. On the other hand, A. alpinus is a common plant in the collecting and may be host of N. tripolii. New for Switzerland. Phytoliriomyza arctica (Lundbeck, 1901) Reference: Martinez, 1998: 271. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, viii.2000, pasture, MT, B. Merz leg. (CMB); Rothenbrunnen, 600 m a.s.l., 1 d, 25.x1.1992. VS: Baltschieder, 670 m a.s.l., 12, 12.v.1991; Branson, Follatéres, 450-950 m a.s.l., 1 à, 29.111.2002; Leuk Pfynwald, 650 m a.s.l., 1 6, 2.v.1997, B. Merz leg. (MHNG). Notes: Almost cosmopolitan species but not known from the Afrotropical Region; common in Europe. Early stages unknown, but once reported as stem miner on Sonchus, probably also on other Asteraceae. Phytoliriomyza melampyga (Loew, 1869) Reference: Martinez, 1998: 271. Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 d, 25.viii.2001, B. Merz & Landry leg. (MHNG). Notes: This Holarctic species is common in temperate and northern Europe and also recorded from North America and Oriental Region (India). The larva forms a leaf mine on /mpatiens particularly on J. noli-tangere and I. parviflora. *Phytoliriomyza perpusilla (Meigen, 1830) Material examined: VS: Branson, Follatères, 450-950 m a.s.l., 1 6, 29.111.2002, B. Merz leg. (MHNG). Notes: P. perpusilla is distributed everywhere in Europe including the Mediterranean area and penetrates to the Afrotropical Region (Cape Verde Is., Lesotho, 794 M. CERNY South Africa). Host plant unknown but probably several genera of Asteraceae (Spencer, 1976). New for Switzerland. Phytomyza affinis Fallén, 1823 References: Hendel, 1931-6: 334; Martinez, 1998: 271. Material examined: GR: Valbella Casoja, 1500 m a.s.l., 1 6, 19.vii.2000, B. Merz leg. (MHNG). Notes: A species known to occur in the West Palaearctic, in particular in temperate and northern Europe but also recorded in the Mediterranean area and the Far East (Kuril Islands). The larva feeds in seed-heads on Euphrasia. *Phytomyza albipennis Fallen, 1823 Material examined: GR: S. Vittore, Monticello, 280 m a.s.l.,2 gd d, 8.iv.1997, B. Merz leg. (MHNG). Notes: This species is recorded from Europe including the Mediterranean area (Croatia, Canary Islands, Italy, Spain and the former Yugoslavia). Host plant unknown but some specimens have been caught on Ranunculus and the larvae possibly feed as an internal stem-borer (Spencer, 1972b, 1990). New for Switzerland. Phytomyza angelicae Kaltenbach, 1872 Reference: Martinez, 1998: 271. Material examined: JU: Bonfol, 450 m a.s.l., 1 &, 30.iv.1990, B. Merz leg. (MHNG). Notes: A Holarctic species which is known from temperate and northern Europe (not recorded from southern Europe), Kazakhstan, Turkey and North America. The larva forms an upper surface blotch leaf mine on Angelica silvestris, A. archangelica, Laserpitium latifolium, infrequently also on Aegopodium podagraria. Phytomyza artemisivora Spencer, 1971 Reference: Martinez, 1998: 271. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This species was described from England, Denmark and Germany. P. artemisivora is distributed in the Palaearctic Region from temperate and northern Europe to Kyrgysztan and Japan. Not confirmed in the Mediterranean area. The larva forms a white linear leaf mine on Artemisia vulgaris. Phytomyza brischkei Hendel, 1922 References: Hendel, 1931-6: 364; Martinez, 1998: 271. Material examined: GR: Lenzerheide, 2000 m a.s.l., 25 d gd, viii.2000,2 dd, 21.-31.v111.2000, pasture, MT, B. Merz leg. (CMB); Lenzerheide, Sundroina, 1520 m a.s.l., 1 0, 17.v.1997, B. Merz leg. (MHNG). VS: Saas-Fee, 1800-2100 m a.s.1.,2 6 6, 26.11.1965, O. Lomholdt leg. (ZMUC). Notes: The species is only known from temperate Europe, no records are available from northern and southern Europe. Host plants are species of Trifolium, particularly 7. repens, T. pratense, T. fragiferum and T. alpinum. AGROMYZIDAE IN SWITZERLAND 795 *Phytomyza calthivora Hendel, 1934 Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.l., 1 d,, 6.iv.2002, B. Merz leg. (MHNG). Notes: A rare species of temperate and northern Europe which can be found at localities where Caltha palustris, the host plant, occurs. The larva forms a short and broad leaf mine. New for Switzerland. *Phytomyza calthophila Hering, 1931 Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.l., 1 d, 6.iv.2002, B. Merz leg. (MHNG). Notes: P. calthophila is more common in temperate and northern Europe than the preceding species, though both species may occur together at same sites on Caltha palustris. Compared with P. calthivora the leaf mine formed by the larva is long and narrow. New for Switzerland. Phytomyza chaerophylii Kaltenbach, 1856 Reference: Martinez, 1998: 271. Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 Sd, 25.viii.2001, B. Merz & Landry leg. (MHNG). Notes: This species is known from Europe including the Mediterranean area and also recorded from Turkey and Japan. The larva forms a linear leaf mine on Chaerophyllum, Anthriscus sylvestris, less frequnetly on Conium maculatum, Conopodium majus, probably also on Carum, Daucus, Sison and Torilis. Phytomyza continua Hendel, 1920 Reference: Martinez, 1998: 271. Material examined: ZH: Zürich, 420 m a.s.l., 1 ©, 30.vi.1989, B. Merz leg. (MHNG). Notes: Very common in Europe and also recorded from China, Kamchatka and Japan. The larva feeds along the midrib of leaves of Cirsium and Carduus. *Phytomyza crassiseta Zetterstedt, 1860 Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 d, 8.vii.2001, B. Merz leg. (MHNG). Notes: P. crassiseta has a typically dilated arista. This Holarctic species is common in Europe and also recorded from Russia (including the Asian part), Kuril Islands, Japan and the United States. The larva forms a linear mine, initially adjoining leaf margin on Veronica. New for Switzerland. *Phytomyza eumorpha Frey, 1946 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 21.-31.vii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This species was described from Finland and Spencer (1976) recorded it later from Sweden. Our locality is the first one in Central Europe. The biology of P. eumorpha is unknown. New for Switzerland. 796 M. CERNY *Phytomyza evanescens Hendel, 1920 Material examined: ZH: Zürich Oerlikon, 430 ma.s.l., 1 & ,23.v.1992, B. Merz leg. (MHNG). Notes: Zlobin (1994) reclassified this species and placed it in Phytomyza where it belongs to the albipennis-group. P. evanescens is characterized by a flap-like appendage on the epandrium. This Holarctic species is distributed in Europe from Island to the Mediterranean area (Dalmatia, Sicily and Spain), and also recorded from Tajikistan, Kuril Islands and North America. The larva feeds internally in the stem of Ranunculus acris and R. lanuginosus. New for Switzerland. Phytomyza fallaciosa Brischke, 1880 Reference: Martinez, 1998: 271. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: P. fallaciosa was described from Poland. Its distribution area includes Europe and the easternmost Palaearctic (Kuril Islands). The larva forms a leaf mine on Ranunculus, particularly on R. repens, but also on R. lingua. *Phytomyza flavicornis Fallén, 1823 Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 3d, 18.iv.2003. JU: Lucelle, 560 m a.s.l., 1 2, 30.iv.1990. All B. Merz leg. (MHNG). Notes: This Holarctic species is common in Europe and North America. The larva feeds as an internal stem borer on Urtica dioica. New for Switzerland. Phytomyza glechomae Kaltenbach, 1862 Reference: Martinez, 1998: 271. Material examined: GE: Chancy, La Laire, 350 m a.s.l., 2 66, 18.iv.2003; Chancy Vers Vaux, 335 m a.s.l., 1 36, 1.vi.2002, B. Merz leg. (MHNG). Notes: P. glechomae is common in Europe, Japan and North America. The larva forms a distinctive leaf mine on Glechoma hederacea. *Phytomyza gymnostoma Loew, 1858 Material examined: SZ: Brunni, 900 m a.s.l., 1 2, 26.vi.1990, B. Petersen leg. (ZMUC). VS: Leuk Pfynwald, 630 m a.s.l., 1 d, 21.1v.1998, B. Merz & Botta leg. (MHNG); Saas-Fee, 2000-2200 m a.s.l., 1 2, 21.vii.1965, O. Lomholdt leg. (ZMUC). Notes: The species was described from Poland and later recorded from further European countries, Turkey and Turkmenistan. Recently, this large species is considered to be a pest on Allium sp. with a certain economic importance (Spencer, 1973). New for Switzerland. *Phytomyza heracleana Hering, 1937 Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.l., 1 à, 1 2, 16.vi.1997, B. Mocek leg. (CBM). Notes: P. heracleana was described from Germany and it is distributed mainly in countries of temperate Europe. Rarely it is also recorded from the Mediterranean AGROMYZIDAE IN SWITZERLAND 797 area (Bulgaria and Italy). The larva forms a regular, interparenchymal blotch leaf mine on Heracleum sphondylium, H. sibiricum, and H. mantegazzianum. New for Switzerland. *Phytomyza hirsuta Spencer, 1976 Material examined: GR: Lenzerheide, Piz Danis, 2250-2490 m a.s.l., 1 d, 12.vii.1996, B. Merz leg. (MHNG). Notes: This species was described from Finland and Norway. Later it was also recorded from Sweden. Recently this species was found in the High Tatra Mts in Slovakia (Cerny & Vala, 2005). Our record represents the southernmost locality in Europe. New for Switzerland. *Phytomyza homogyneae Hering, 1927 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 21-31.viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This mountain species was described from Germany, where it was reared from Homogyne alpina, where the larva forms a linear mine. The species is known from Central European mountains (recently found in the Sumava Mts, Czech Republic, Cerny & Vala, 2005) but also from China, Japan and Taiwan. New for Switzerland. Phytomyza nigripennis Fallén, 1823 References: Hendel, 1931-6: 439; Martinez, 1998: 272. Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 d, 24.iv.2003, B. Merz & F. Amiet leg. (MHNG). ZH: Zürich Allmend, 450 m a.s.l., 1 2, 1.v.1995, B. Merz leg. (MHNG); Zürich Waldgarten, 1 à , 31.i11.1997, B. Merz leg. (MHNG). Notes: This species is characterized by conspicuously dark wings. It is distributed in temperate and northern Europe, with a few localities in Northern Italy and former Yugoslavia. Its host plant is not known, but it may attack Anemone nemorosa. The larva feeds probably in the stem or the root, early stages unknown (Spencer, 1976). Phytomyza notata Meigen, 1830 Reference: Martinez, 1998: 272. Material examined: GR: Lenzerheide, 2000 m a.s.1., 8 d d, viii.2000, pasture, MT, B. Merz leg. (CMB). Notes: This species is known from temperate and northern Europe including northern Italy. The larva forms a short, broad, linear mine recorded on Ranunculus acer, R. auricomus, R. bulbosa and particularly on R. repens. *Phytomyza origani Hering, 1931 Material examined: GE: Cartigny, Moulin de Vert, 470 m a.s.l.,1 6, 4.vi.2001, M. Eggenberger & B. Merz leg. (MHNG). Notes: P. origani was described from Germany and later recorded from Bulgaria, the Czech Republic, Denmark, England, France, Hungary, Lithuania, Poland 798 M. CERNY and Spain. The larva forms a typical leaf mine on Origanum vulgare (Spencer, 1976). New for Switzerland. *Phytomyza pauliloewii Hendel, 1920 Material examined: VS: Branson, Follatéres, 450-950 m a.s.1., 1 4, 29 .iii.2002, B. Merz leg. (MHNG). Notes: This species is only locally distributed in temperate Europe but also known from northern Europe (Finland and Sweden) and the Mediterranean area (Italy). The larva forms a small regular blotch leaf mine on Pimpinella spp. and Peucedanum oreoselinum. New for Switzerland. Phytomyza plantaginis Robineau-Desvoidy, 1851 Reference: Martinez, 1998: 272. Material examined: VS: Leuk Pfynwald, 650 m a.s.l., 1 5, 23.ix.1992, B. Merz & Otto leg. (MHNG), 630 m a.s.l., 1 gd, 6.viii.1997, B. Merz leg. (MHNG). Notes: P. plantaginis is a common species of the West Palaerctic and North America (Canada, United States). Rarely also known from the East Palaearctic, Afrotropical, Oriental and Australasian Regions. The larva forms a narrow, white linear leaf mine on Plantago lanceolata and P. major. *Phytomyza platystoma (Hendel, 1920) Material examined: VS: Branson, Follatéres, 450-950 m a.s.l., 3 dd, 29 111.2002, B. Merz leg. (MHNG). Notes: This alpine species was described as Napomyza platystoma but Spencer & Martinez (1987) transferred it to Phytomyza. It was described from Austria (Piestingtale, Nördliche Voralpen). This is the second record from Central Europe. Its biology is unknown. New for Switzerland. *Phytomyza pubicornis Hendel, 1920 Material examined: GE: Avusy, Moulin de la Grave, 360 m a.s.l., 1 dg, 18 .iv.2003, B. Merz leg. (MHNG). ZH: Zürich Allmend, 440 m a.s.l., 1 ¢, 17.iv.1996, B. Merz leg. (MHNG). Notes: This species is distributed in temperate and northern Europe. No records are known from the Mediterranean area. The larva forms a short linear leaf mine on Aegopodium podagraria. New for Switzerland. *Phytomyza pullula Zetterstedt, 1848 Material examined: ZH: Affoltern a. A., 600 ma.s.l., 1 d, 8.viii.1987, B. Merz leg. (MHNG). Notes: P. pullula belongs to the species with a Holarctic distribution. It is common in Europe and North America. The larva forms a narrow linear leaf mine on Achillea, Anthemis, Chrysanthemum vulgare, Matricaria, and most commonly on M. inodora. New for Switzerland. Phytomyza ranunculi (Schrank, 1803) References: Hendel, 1931-6: 467; Martinez, 1998: 272. AGROMYZIDAE IN SWITZERLAND 799 Material examined: GR: Ausserferrera, 1300-1550 m a.s.l., 1 4, 18.vi.1994, B. Merz leg. (MHNG); Lenzerheide, 2000 m a.s.l., 1 6, 14.-21.vii.2000, pasture, MT, B. Merz leg. (CMB); Lenzerheide See, 1500 m a.s.l., 1 d, 15.vii.2000, B. Merz leg. (MHNG); Lenzerheide Sundroina, 1520 m a.s.l., 1 d, 17.v.1997, B. Merz leg. (MHNG). TI: Monte San Giorgio, 600-1100 m a.s.l., 1 4, 5.vii.2001, B. Merz leg. (MHNG). ZH: Zürich Zürichberg, 600 m a.s.l., 1 8, 5.v.1993, B. Merz leg. (MHNG), 1 3d, 8.v.1996, P. Walser leg. (MHNG). VD: Bonvillars en fauchant [= sweeping], 630 m a.s.1., 1 2, 25.iv.1988, D. Burckhardt leg. (MHNG). Notes: P. ranunculi is a common species mining Ranunculaceae in the Holarctic Region. *Phytomyza ranunculicola Hering, 1949 Material examined: GE: Bernex, Chante-Merle, 415 m a.s.1.,1 6 ,20.v.2002, B. Merz leg. (MHNG). Notes: The species was described from Germany. It is distributed in temperate Europe but it is probably absent from Scandinavia and the Mediterranean area. The larva forms a secondary blotch leaf mine on Ranunculus acer. New for Switzerland. *Phytomyza rapunculi Hendel, 1927 Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.l., 1 à, 1 ®, 16.vi.1997, B. Mocek leg. (CBM). GR: Lenzerheide, 2000 m a.s.l., 1 à, v111.2000, pasture, MT, B. Merz leg. (CMB). Notes: This European species was described from Austria and later recorded particularly in temperate and southern Europe. In Scandinavia it is known only from Sweden (Spencer, 1990). The larva forms a linear mine on Campanula rapunculoides, C. persicifolia and Phyteuma. New for Switzerland. *Phytomyza rhabdophora Griffiths, 1964 Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, viii.2000, pasture, MT, B. Merz leg. (CMB). Lenzerheide, Sundroina, 1550 m a.s.l., 1 3, 18.vii.1997, B. Merz leg. (MHNG). VS: Saas-Fee, 1800 m a.s.1.,1 d ,21.vii.1965, O. Lomholdt leg. (ZMUC). Notes: The type locality is in Germany and the species is distributed in temperate Europe, but it is apparently missing in the Mediterranean area. Its biology is unknown, larvae feed probably on Leontodon (Tschirnhaus, 1969). New for Switzerland. *Phytomyza rostrata Hering, 1933 Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 6, 2.vii.2001, B. Merz leg. MHNG). Notes: The species was described from Germany and it is distributed in temperate Europe, but apparently absent from the Mediterranean area. The larva forms a typical mine on Euphrasia, Melampyrum, Odontites and Rhinanthus (Spencer, 1976). New for Switzerland. Phytomyza rufipes Meigen 1830 Reference: Martinez, 1998: 272. 800 M. CERNY Material examined: GE: Cartigny, Moulin de Vert, 360 m a.s.1.,1 8, 1.v.1999, B. Merz leg. (MHNG). Notes: This pale species is common in the West Palaearctic and North America. The larva feeds inside the stem or midrib of larger leaves on Brassicaceae, mainly on Brassica. *Phytomyza sedi Kaltenbach, 1869 Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.1.,2 6d, 19.v.2001, B. Merz leg. (MHNG). Notes: This species was described from Boppard (Germany) and later recorded from Spain under the synonym Ph. catalaunica Spencer, 1960. Our record confirms the occurrence in Central Europe. Sedum is the host plant of this species. New for Switzerland. *Phytomyza soenderupi Hering, 1941 Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.l., 1 d, 6.1v.2002, B. Merz leg. (MHNG). Notes: This species was described from Denmark and later recorded from Belgium, the British Isles, the Czech Republic, Estonia, Germany and Norway. The larva feeds within the petiole of Caltha palustris. New for Switzerland. Phytomyza tenella Meigen, 1830 Reference: Martinez, 1998: 272. Material examined: GR: Lenzerheide, 2000 m a.s.l., pasture, 2 dd, 23.viii.- 9.1x.2000, MT, B. Merz leg. (CMB); Juf-Stallerberg, 2400 m a.s.l., 1 à , 18.viii.1991, B. Merz & A. Freidberg leg. (MHNG). Notes: Phytomyza tenella is distributed in Europe, China, Turkey, the United States and the Afrotropical Region (Ethiopia). The larva feeds in seed-heads of Pedicularis palustris and almost certainly other Pedicularis. *Phytomyza varipes Macquart, 1835 Material examined: GR: Lenzerheide, Sundroina, 1550 m a.s.., 1 à, 18.vi1.1997, B. Merz leg. (MHNG). Notes: This rare species was described from Northern France. The larva feeds in seed-heads of Rhinanthus. New for Switzerland. Phytomyza vitalbae Kaltenbach, 1872 Reference: Martinez, 1998: 272. Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 d, 14.-21.vii.2000, pasture, MT, B. Merz leg. (CMB). VS: Leuk Pfynwald, 600 m asl., 2 dd, 25.v111.2001, B. Merz & Landry leg. (MHNG). Notes: A species known from various European countries and also from Canada, South Africa, China, Nepal, Taiwan and Australia. The larva forms a long, narrow, upper surface leaf mine on Clematis vitalba, C. alpina and some other Clematis. Phytomyza wahlgreni Rydén, 1944 References: Griffiths, 1964: 411; Spencer, 1976: 527; Martinez, 1998: 272. AGROMYZIDAE IN SWITZERLAND 801 Material examined: AG: Thalheim, 550 m a.s.l., 1 5, 20.v.1992, B. Merz leg. (MHNG). GR: Lenzerheide, 1600 m a.s.l.,1 5, 8.x.1991, B. Merz & M. Eggenberger leg. (MHNG); Lenzerheide, 2000 m a.s.l., 3 Sd, 14.-21.vii.2000, 2 dd, 23.- 31.viii.2000, pasture, MT, B. Merz leg. (CMB). NE: St. Blaise, Les Riedes, 470 m a.s.l., 1 d, 19.v.2001, B. Merz leg. (MHNG). VS: Saas-Fee, 2000 m a.s.l., 3 dd, 18.vii.1965, 2400 m a.s.1.,1 2, 19.vii.1965, 2200 m a.s.l.,2 2 2, 21.vii.1965, 1800 m a.s.1.,1 2, 24.vii.1965, O. Lomholdt leg. (ZMUC). Notes: A Holarctic species distributed in temperate and northern Europe. In the Mediterranean area known to occur in Italy. It is also recorded from the United States, China and the Kuril Islands. From Switzerland recorded by Griffiths (1964) for the first time (VS: Bérisal, under the synonym P. taraxacocecis Hering, 1949). The larva feeds in the midrib of leaves on Taraxacum spp. *Pseudonapomyza atra (Meigen, 1830) Material examined: FR: Mt. Vully, Bas Vully, 460-650 m a.s.l., 1 6, 6.vi.2003, B. Merz & Amiet leg. (MHNG). GE: Bernex, Chante-Merle, 415 m a.s.l., 1 à, 16.viii.2002, B. Merz leg. (MHNG); Cartigny, Moulin de Vert, 360 m a.s.l., 1 à, 2.vi.2002, B. Merz leg. (MHNG); Chancy, La Laire, 350 m a.s.l., 1 4, 9.vii.2002, B. Merz leg. (MHNG). VS: Visperterminen, Giw-Gebidemsee, 1900-2200 m a.s.l., 1 à, 28.viii.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich-Hönggerberg, 600 m a.s.l., 1 3, 26.iv.1993, B. Merz leg. (MHNG). Notes: Ps. atra is generally distributed in the Holarctic Region. It is common in northern and temperate Europe (Cerny 1992, 1998, 2005a). Spencer (1973) considered it to be rare in the Mediterranean area. The larva forms a short, narrow mine on Gramineae, feeding on many genera, e.g. Apera, Avena, Hordeum, Lolium, Phalaris, Poa, Secale and Triticum. New for Switzerland. *Pseudonapomyza errata Zlobin, 1993 Material examined: GE: Russin, Les Baillets, 405 m a.s.l., 1 d, 30.vi.2001, B. Merz leg. (MHNG). NE: St. Blaise, Les Riedes, 470 m a.s.l., 2 66, 19.v.2001, B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 d, 2.vii.2001; Visperterminen, 1400 m a.s.l., 1 4, 27.vii.1991, B. Merz leg. (MHNG). Notes: This species was described from Russia, Uzbekistan and Mongolia and it is also recorded from the Czech Republic and France. Its biology is unknown. New for Switzerland. *Pseudonapomyza eurasiatica Zlobin, 2003 Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 à, 6.vi.2001, 1 à, 7.v1.2001, B. Merz & Landry leg. (MHNG). Notes: Recently described from the male holotype originating from Tajikistan and paratypes from Greece, Kazakhstan, Turkmenistan and Ukraine by Zlobin (2003b). This is the first record from Central Europe and represents the westernmost boundary of its distribution area. Biology is unknown. New for Switzerland. Pseudonapomyza europaea Spencer, 1973 References: Papp, 1984: 313; Spencer, 1973: 270; Martinez, 1998: 272. 802 M. CERNY Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 ¢, 25.v.1997, B. Merz leg. (MHNG), 2 6 4, 6.vi.2001; 1 d, 2.vii.2001; 1 d ,25.viii.2001, B. Merz & Landry leg. (MHNG). Notes: Widely distributed in Europe and also recorded from Turkey and Japan (Cerny, 2005a). It is less common than Ps. atra. Spencer (1973) described this species from Italy (holotype), Austria, Macedonia, Serbia, Slovenia and Switzerland (VS: Stalden). Its biology is unknown. *Pseudonapomyza strobliana Spencer, 1973 Material examined: GE: Russin, Les Baillets, 405 m a.s.1.,1 4, 8.viii.2002, B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 d, 6.vi.2001, 1 6, 25.viii.2001, B. Merz & Landry leg. (MHNG). Notes: The species was described from Dalmatia and Austria. The center of its distribution area is probably in the Mediterranean area (Spain, Bulgaria, Turkey) but recently it was also recorded from the Czech Republic, France, Germany, Hungary, Sweden and Ukraine (Cerny, 1998, 2005a). Its biology is unknown. New for Switzerland. *Pseudonapomyza vota Spencer, 1973 Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 dg, 7.vi.2001; Leuk Platten, 625 m a.s.l., 1 6, 8.vi.2001, all B. Merz & Landry leg. (MHNG). Notes: This species was described from a single male originating from Spain. The distribution area of this species ranges from Spain to Israel, i.e. throughout the Mediterranean area. Biology of the species is unknown. New for Switzerland. CONCLUSIONS This review embraces new data on the fauna of mining flies or Agromyzidae occurring in Switzerland and adds 92 species to the 140 species listed by Martinez (1998) bringing the list of Swiss agromyzids to 232 species. The potential number of Swiss Agromyzidae species is probably much higher, given the different origins of the native fauna and the unusally diverse pattern of ecosystems. Such a conclusion seems to be confirmed by a comparison of species richness in neighbouring countries. Franz (1989) listed 270 species from Austria and Tschirnhaus (1999) 552 species from Germany. According to Spencer (1992) many species known from the Italian alpine valleys will probably also be found in the Swiss Alps. Siiss (1999, 2001, 2002) recorded 224 species in Italy (173 from North Italy); of this number 112 species are common for both countries and 112 species are known exclusively from Italy. So far, no species of Hexomyza Enderlein, 1936, has been recorded from Switzerland, although 6 species are known in Europe and 5 of them live in Italy. Likewise Ptochomyza Hering, 1942, with three European species and Nemorimyza posticata (Meigen, 1830) known from adjoining countries, have not been found so far in Switzerland. Of 10 European species of Phytobia Lioy, 1864, only P. mallochi is known from Switzerland, while Melanagromyza Hendel, 1920, is represented by four species only (11 % of the European species) and Liriomyza Mik, 1894, by 18 species (13 %). There is thus good evidence that the list of Swiss agromyzids is far from final. AGROMYZIDAE IN SWITZERLAND 803 ACKNOWLEDGEMENTS My sincerest thanks go to Miroslav Bartäk (Praha), Rudolf Meier (Copenhagen), Bernhard Merz (Genève) and Bohuslav Mocek (Hradec Kralové) for allowing me to use the collections in their care and Rudolf Rozkoënÿ (Brno, Czech Republic) for invaluable support during preparation of the manuscript. REFERENCES BLAND, K. P. 2000. James Hardy and some new synonyms for British Flies (Diptera: Cecidomyiidae, Agromyzidae and Anthomyiidae). Dipterists Digest 7: 9-14. BOUCHER, S. 2003. The New World species of Cerodontha (Xenophytomyza) Frey (Diptera: Agromyzidae). Zootaxa 178: 1-8. Cerny, M. 1992. A revision of Czechoslovak species of Pseudonapomyza Hendel, with description of four new species (Diptera, Agromyzidae). Acta Entomologica Bohemoslovaca 89: 451-465. CERNY, M. 1998. Two new species of Pseudonapomyza, with notes on distribution of the European species (Diptera: Agromyzidae). Folia Heyrovskyana 6 (1): 7-14. CERNY, M. 2005a. 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Memoirs of an amateur entomologist. SPB Academic Publishing. The Hague, 414 pp. SPENCER, K. A. & MARTINEZ, M. 1987. Additions and corrections to the Agromyzidae section of the catalogue of Palaearctic Diptera (PAPP, 1984). Annales de la Societe Entomologique de France (N. S.) 23 (3): 253-271. Süss, L. 1999. Su alcuni Agromizidi recentemente raccolti in Italia. Bollettino di Zoologia agraria e di Bachicoltura, Series II, 31 (2): 127-137. Suss, L. 2001. Cerodontha (Poemyza) unisetiorbita Zlobin (Diptera Agromyzidae) nuova per l’Europa. Bollettino di Zoologia agraria e di Bachicoltura Series I, 33 (1): 73-77. Suss, L. 2002. Morphological and biological observations on Ptochomyza czerny (Strobl) new for Italy. Bollettino di Zoologia agraria e di Bachicoltura, Series II, 34 (1): 17-23. TSCHIRNHAUS, M. von 1969. Zur Kenntnis der Variabilität, Eidonomie und Verwandtschaft bemerkenswerter Agromyzidae. Senckenbergiana biologica 50 (3/4): 143-157. TSCHIRNHAUS, M. von 1971. Unbekannte Stridulationsogame bei Dipteren und ihre Bedeutung für Taxonomie und Phylogenetik der Agromyziden. Beiträge zur Entomologie 21 (7/8): 551-579. TSCHIRNHAUS, M. von 1999. Agromyzidae (pp. 118-130). In: SCHUMANN, H., BAHRMANN, R. & STARK, A. (eds). Checkliste der Dipteren Deutschlands. Entomofauna Germanica 2. Studia Dipterologica, Supplement 2: 1-354. ZLOBIN, V.V. 1986. Review of mining flies of the genus Cerodontha Rondani (Diptera: Agro- myzidae) of the fauna of the USSR. III. Subgenus Poemyza atra-group. Proceedings of the Zoological Institute, Leningrad 140: 85-91. (In Russian, English summary) AGROMYZIDAE IN SWITZERLAND 805 ZLOBIN, V. V. 1992. Review of mining flies of the genus Cerodontha. IV. Subgenus Poemyza (Diptera: Agromyzidae). Zoosystematica Rossica 1: 117-141. ZLOBIN, V. V. 1993a. Review of mining flies of the genus Napomyza Westwood (Diptera: Agromyzidae). III. New European species of lateralis-group. International Journal of Dipterological Research 4 (4): 225-235. ZLOBIN, V. V. 1993b. Review of mining flies of the genus Cerodontha. V. Subgenus Poemyza (Diptera: Agromyzidae). Zoosystematica Rossica 2 (1): 179-184. ZLOBIN, V. V. 1993c. Review of mining flies of the genus Cerodontha Rond. (Diptera: Agromyzidae). VI. Subgenus Xenophytomyza Frey. Proceedings of the Zoological Institute, St.-Petersburg 251: 135-156. (In Russian, English summary) ZLOBIN, V. V. 1993d. Review of mining flies of the genus Napomyza Westwood (Diptera: Agromyzidae). I. elegans - group. Proceedings of the Zoological Institute, St.- Petersburg 251: 157-174. (In Russian, English summary) ZLOBIN, V. V. 1994. Review of mining flies of the genus Napomyza Westwood (Diptera: Agromyzidae). V. Phytomyza species currently placed in the genus Napomyza West- wood. International Journal of Dipterological Research 5 (4): 289-311. ZLOBIN, V. V. 1995. Review of mining flies of the genus Metopomyza Enderlein (Diptera: Agromyzidae). International Journal of Dipterological Research 6 (2): 143-168. ZLOBIN, V. V. 2000. Contribution to the knowledge of Agromyza species (Diptera: Agromyzidae) feeding on Leguminosae. I. International Journal of Dipterological Research 11 (1): 37-45. ZLOBIN, V. V. 2003a. Review of mining flies of the genus Liriomyza Mik (Diptera: Agro- myzidae). I. The Palaearctic flaveola-group species. International Journal of Diptero- logical Research 13 (3): 145-178. ZLOBIN, V.V. 2003b. Contribution to the knowledge of the genus Pseudonapomyza Hendel (Diptera: Agromyzidae), with descriptions of twenty four Old World species. Inter- national Journal of Dipterological Research 13 (4): 205-245. SORA TST Dea TRS Ae Een ia = Li iM a ba N VD Auen Ais A) a“ E a ple que i fe. de Ba ict ee me ee fi REVUE SUISSE DE ZOOLOGIE 112 (4): 807-830; décembre 2005 The millipede family Paradoxosomatidae in Paraguay, with descriptions of five new species (Diplopoda, Polydesmida) Sergei I. GOLOVATCH Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071 (V-71), Russia. E-mail: sgol@orc.ru The millipede family Paradoxosomatidae in Paraguay, with descrip- tions of five new species (Diplopoda, Polydesmida). - A review is pro- vided of the millipede family Paradoxosomatidae in Paraguay, with keys compiled to 14 genera and 63 species occurring in this country and/or adjacent areas. Five species are described as new: Catharosoma bilineatum sp. n., C. mahnerti sp. n., Broelemannopus minutus sp. n., Mestosoma sim- plex sp. n. and M. crassipes sp. n. Keywords: Diplopoda - Polydesmida - Paradoxosomatidae - taxonomy - Paraguay. INTRODUCTION The family Paradoxosomatidae is among the largest among the Diplopoda, with nearly 200 genera currently accepted as valid. The American fauna is highly peculiar due to the absence of indigenous paradoxosomatids north of Costa Rica. In South America two major centres of diversification are distinguishable, one in the area of southern Brazil, Paraguay and northern Argentina, the other in Peru, northern Bolivia and possibly Ecuador. The vast regions of northern Brazil, Guiana, Venezuela and Colombia appear to harbour very few species (Jeekel, 1968, 2002). The Neotropical fauna itself is strongly dominated by relatively few (about a dozen) genera of the endemic tribe Catharosomatini. Of these genera, Mestosoma Silvestri, 1897 is certainly the largest and particularly widespread, with about 75 species occurring throughout South America and reaching both Costa Rica and the island of Dominica (introduced?) in the north. Catharosoma Silvestri, 1913 is the second largest genus, its 13 species being known from Brazil, Paraguay and Argentina (Hoffman, 1980, 1999). The remaining genera are mono- to oligotypic. The present paper provides a review of, and keys to, the bulk of the paradoxo- somatids occurring in the southern Neotropical diversification centre, based both on all available literature sources and on the important collection of Paradoxosomatidae from Paraguay housed in the Muséum d’histoire naturelle, Geneva (MHNG). Five new species from three genera of Catharosomatini have been revealed there, all described below and thus considerably enriching our knowledge of the fauna of Paraguay. A few paratypes from the material have been retained for the collection of the Zoological Museum, State University of Moscow (ZMUM), Russia. Manuscript accepted 07.02.2005 808 S. I. GOLOVATCH FAUNISTIC REVIEW Since detailed reviews of the history of research on Neotropical Paradoxo- somatidae are available and still fully relevant (Jeekel, 1963, 1968), only a short account appears necessary here concerning the species list of the particular region involved. Virtually all later contributions (e.g. Hoffman, 1977, 1999; Golovatch, 1992; Jeekel, 2002; Golovatch et al., 2003) are irrelevant in the present context, as they only or chiefly treat more northern faunas. The following Paradoxosomatidae from Paraguay and/or the adjacent parts of Bolivia, northern Argentina and southern Brazil have hitherto been recognized as valid: Broelemannopus Verhoeff, 1938 Broelemannopus escaramucensis (Schubart, 1944) - Brazil (Säo Paulo) (Schubart, 1944, 1952) B. glabratus (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b) B. ibitiensis (Schubart, 1945) - Brazil (Sao Paulo) (Schubart, 1945a) B. pirassunungensis (Schubart, 1944) - Brazil (Säo Paulo) (Schubart, 1944, 1945a, 1952) Catharosoma Silvestri, 1897 Catharosoma curitibense Schubart, 1953 - Brazil (Parana) (Schubart, 1953) C. digitale Schubart, 1953 - Brazil (Parana) (Schubart, 1953) C. hoffmani Kraus, 1956 - Paraguay (Kraus, 1956) C. intermedium (Carl, 1902) - Brazil (Rio Grande do Sul) (Carl, 1902; Attems, 1914, 1937) C. mesorphinum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 1937; Schubart, 1953) C. mesoxanthum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 1937; Schubart, 1953) C. mixtum Kraus, 1956 - Brazil (Santa Catarina) (Kraus, 1956) C. myrmekurum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 1937; Schubart, 1953) C. palmatum Schubart, 1953 - Brazil (Parana) (Schubart, 1953) C. palustre Schubart, 1943 - Brazil (Mato Grosso) (Schubart, 1943) C. paraguayense (Silvestri, 1895) - Paraguay, Argentina and Brazil (Santa Catarina and Mato Grosso) (Silvestri, 1895b, 1902; Attems, 1898, 1914, 1937; Schubart, 1953; Kraus, 1956; Jeekel, 1965) («South America», according to Mauriès, 1998) C. peraccae Silvestri, 1902 - Paraguay (Silvestri, 1902; Attems, 1914, 1937) C. taeniatum (Brolemann, 1929) - Brazil (Santa Catarina) (Brolemann, 1929; Attems, 1937; Schubart, 1953) Chondromorpha Silvestri, 1897 Chondromorpha xanthotricha (Attems, 1898) - pantropical, introduced (e.g. Jeekel, 1963) PARADOXOSOMATIDAE IN PARAGUAY 809 Gonodrepanoides Schubart, 1945 Gonodrepanoides travassosi Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) Gonodrepanum Attems, 1914 Gonodrepanum drepanephoron (Attems, 1898) - Brazil (Sao Paulo) and Argentina (Attems, 1898, 1901, 1914, 1937; Brölemann, 1902b; Mauries, 1998) G. falciferum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) G. flavolineatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) G. furcatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) G. grajahuense Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) G. levisetum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 1937; Schubart, 1953; Mauriés, 1998) G. levisetum var. coniferum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 1937; Schubart, 1953; Mauriès, 1998) G. torresae Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) Habrodesmoides Attems, 1943 Habrodesmoides costalimai (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b) H. perturbans Attems, 1943 - Brazil (Rio de Janeiro) (Attems, 1943) Mestosoma Silvestri, 1897 Mestosoma alticola (Attems, 1931) - Bolivia (Cochabamba) and Peru (Attems, 1931, 1937; Kraus, 1956) M. balzanii (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems, 1914, 1937) M. bicolor Silvestri, 1898 - Paraguay and Brazil (Mato Grosso) (Silvestri, 1898, 1902; Attems, 1899, 1914, 1937; Jeekel, 1965) M. boliviae (Chamberlin, 1957) - Bolivia (near Potosf) (Chamberlin, 1957) M. borellii (Silvestri, 1895) - Argentina (Tucuman) and Paraguay (Silvestri, 1895b; Attems, 1914, 1937; Mauriés, 1998) M. camerani (Silvestri, 1895) - Argentina (Chaco) (Silvestri, 1895b; Attems, 1914, 1937; Jeekel, 1965; Mauriés, 1998) M. carioca (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b) M. derelictum (Silvestri, 1895) - Bolivia (Misiones Mosetenes) (Silvestri, 1895a; Attems, 1914, 1937) M. differens Kraus, 1956 - Bolivia (Cochabamba) (Kraus, 1956) M. femorale (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943) M. kalliston (Attems, 1898) - Brazil (Rio Grande do Sul) (Attems, 1898, 1914, 1937) M. luctuosum Silvestri, 1897 - Bolivia (Caiza) (Silvestri, 1897; Attems, 1914, 1937) 810 S. I. GOLOVATCH M. lugubre Silvestri, 1897 - Argentina (Buenos Aires) and Uruguay (Silvestri, 1897, 1902; Attems, 1914, 1937; Mauriés, 1998) M. montanum (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems, 1914, 1937) M. perfidum (Schubart, 1943) - Brazil (Sao Paulo) (Schubart, 1943) M. pseudomorphum (Silvestri, 1895) - Paraguay (Silvestri, 1895b; Carl, 1902; Attems, 1914, 1937; Jeekel, 1965) M. pulvillatum (Attems, 1898) - Paraguay (Attems, 1898, 1901, 1914, 1937) M. salvadorii (Silvestri, 1895) - Argentina (Salta), Bolivia (Chaco) and Paraguay (Silvestri, 1895b, 1902; Attems, 1914, 1937; Hoffman, 1977; also Costa Rica, according to Mauries, 1998) M. schindleri Kraus, 1956 - Bolivia (Silhuencas) (Kraus, 1956) M. tricuspis (Verhoeff, 1938) - Paraguay (Verhoeff, 1938) M. truncatum (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943) M. vittatum (Attems, 1898) - Paraguay (Attems, 1898, 1914, 1937) Mogyella Schubart, 1944 Mogyella nana Schubart, 1944 - Brazil (Sao Paulo) (Schubart, 1944, 1952) Mogyosoma Schubart, 1944 Mogyosoma hamatum Schubart, 1944 - Brazil (Sao Paulo) (Schubart, 1944) Ologonosoma Silvestri, 1898 Ologonosoma iguassuense (Schubart, 1953) - Brazil (Parana) (Schubart, 1953; Mauries, 1998) O. sanctum (Silvestri, 1895) - Paraguay (Silvestri, 1895a; Attems, 1914, 1937; Mauriés, 1998) Orthomorpha Bollman, 1893 Orthomorpha coarctata (De Saussure, 1860) - pantropical, introduced (e.g. Jeekel, 1963) Oxidus Cook, 1911 Oxidus gracilis (C. L. Koch, 1847) - subcosmopolitan, introduced (e.g. Jeekel, 1963) Promestosoma Silvestri, 1898 Promestosoma boggianii Silvestri, 1898 - Paraguay (Silvestri, 1898; Attems, 1899, 1914, 1937; Jeekel, 1965; Mauriés, 1998) and Brazil (Mato Grosso and Mato Grosso do Sul) (original data) Pseudogonodrepanum Schubart, 1945 Pseudogonodrepanum scitum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) PARADOXOSOMATIDAE IN PARAGUAY 811 A few more species, i.e. Mestosoma laetum Silvestri, 1897 (Bolivia, San Francisco — Silvestri, 1897), «Strongylosoma» nitidum Brölemann, 1902 (Brazil, Cubatäo, Sao Paulo — Brölemann, 1902a), «Strongylosoma» pustulatum Brölemann, 1902 (Brazil, Sao Paulo — Brölemann, 1902b) and Catharosoma bromelicola Schubart, 1945 (Brazil, Rio de Janeiro — Schubart, 1945b), the male characters of which are not or are insufficiently known, still remain incertae sedis; none can be included in a key (Jeekel, 1963). The same concerns /ulidesmus Silvestri, 1895, with its type-species /. typicus Silvestri, 1895 known only from a female holotype from Bolivia (Yungas) (Silvestri, 1895a). Jeekel (1963) correctly excluded /ulidesmus from Paradoxosomatidae on the basis of a later redescription of /. typicus from Chilean material provided by Silvestri (1905). But Hoffman (1980), rightly doubting the conspecificity of the Bolivian and Chilean samples, believes that the holotype of /. typicus represents a Mestosoma. If so, then Mestosoma becomes a junior synonym of /ulidesmus. Only topotypic material in connection with revisionary work can shed additional light on the identity of all of the enigmatic taxa mentioned above. DESCRIPTIONS OF NEW SPECIES Catharosoma bilineatum sp. n. Figs 1-9 Material: Holotype 6 (MHNG), Paraguay, Alto Parana Prov., Forestry Centre (C.F.A.P.), Puerto Presidente Stroessner, plot 7, 18.03.1983, leg. P. Berner & C. Dlouhy. — Paratypes: 1 2 (MHNG), same locality, together with holotype; 1 4,3 2 2 (MHNG), 16,1 9 (ZMUM), same locality, around Pindo trunk, plot 3, 24.03.1983, leg. C. Dlouhy; 1 2 (MHNG), same locality, soil sample, plot 4 (Monte natural), Winkler extraction, 10.03.1983, leg. C. Dlouhy; 1 2 (MHNG), same plot 4 (Monte natural), 10.03.1983, leg. P. Berner & C. Dlouhy; 1 2 (MHNG), same locality, soil sample, plot 2 (Monte natural), 29.02.1983, leg. C. Dlouhy; 1 d nn: same locality, soil sample, plot 8 (Monte natural), 18.03.1983, leg. P. Berner & C. Name: To emphasize the presence of two dark paramedian stripes divided by a similar- ly wide but light axial stripe. Diagnosis: Differs from congeners by the characteristic coloration, the strong- ly reduced paraterga visible only on segments 2-4, the presence of relatively short an- tennae, the deeply divided sternal lobe present between coxae 6 of the à , the subunci- form tip of the hypoproct, the bifid tip of the epiproct, coupled with certain details of solenophore structure. Description: Length 21-27 mm (d, 2), width of midbody metazona 1.9-2.1 (3) or 2.1-2.8 mm (@); © usually larger and somewhat broader than &. Holotype ca 23 mm long and 2.0 mm wide. Coloration in alcohol pale yellowish brown to brown, with a characteristic pair of brown to dark brown paramedian stripes divided by an equally wide axial stripe of background coloration; legs and sterna yellowish gray, slightly paler than background coloration; tip of antennae pallid, penultimate anten- nomere dark brown; posterior half of body always somewhat paler than anterior one. Body subcylindrical, not moniliform. Postcollar constriction faint, width of head = 5-16 > collum = 4 > 2 = 3; on segments 17-20 trunk gradually and gently ta- pering toward telson both in width and in height. Antennae relatively short, slightly clavate, in situ reaching beyond segment 2 dorsally (Fig. 1), a little shorter in 9. 812 S. I. GOLOVATCH Fics 1-9 Catharosoma bilineatum sp.n., d holotype: 1) anterior body portion, lateral view; 2) telson, lat- eral view; 3) sternal structures between coxae 4-7 (coxae 4 on top); 4) sternal structures between midbody coxae; 5) leg 12; 6-9) left gonopod, medial, ventral, lateral and ventromedial views, re- spectively. Scale bars 1.0 (1-5) and 0.25 mm (6-9). Paraterga strongly reduced, only on segment 2 fully developed as low keels with a caudal tooth, discernible on segments 3 and 4 as arcuated sulci (Fig. 1), on following segments totally missing. Tegument smooth, shining, only rear parts of metazona often faintly rugulose. Limbus thin, caudal margin entire. Metaterga fully devoid of a trans- verse sulcus; setae medium-sized, often abraded, arranged 2+2 in a transverse line behind a shallow stricture between pro- and metazona. Ozopores lateral, opening level to metatergal surface ca 1/3 metazonital length away from caudal edge. Pleurosternal carinae shaped like narrow keels with a caudal spinule (Fig. 1) traceable until segment 16 (2) or 17 (6), on following segments wanting. Epiproct (Fig. 2) rather long, coni- form, bifid, in ¢ a little longer than in 2 . Hypoproct (Fig. 2) subtriangular, tip pointed, unciform, directed ventrad; 1+1 paramedian setae at caudal edge considerably separated from each other. Paraterga, pleurosternal carinae and hypoproct uncus in d a little more strongly developed than in ©. Sterna densely setose. Sternum between coxae 3 of d with a paramedian, nearly contiguous pair of bunches of setae. Sternum between coxae 4 of d with a prominent, linguiform, laterally setose outgrowth directed anteroventrad (Fig. 3); a similar but PARADOXOSOMATIDAE IN PARAGUAY 813 much lower outgrowth between coxae 5 of d (Fig. 3); a more or less prominent and deeply divided outgrowth between coxae 6 of d poorly delimited caudally against a similar but less prominent bulge with peculiar paramedian bunches of setae between coxae 7 of d (Fig. 3). Postgonopodial sterna with shorter (between anterior coxae) or longer (between posterior coxae), coniform, paramedian spines developed a little bet- ter in 3 (Fig. 4) than in 9. Legs without tarsal brushes but densely setose ventrally; setation gradually thinning out toward telson; tibiae swollen ventrally and supporting characteristic brushes (Fig. 5) between leg-pair 9 until two last pairs. Legs in d a little longer than in 2, as usual becoming a little longer and slenderer toward telson. Each coxa 2 of d with a prominent, somewhat sinuate, distoventral spine carrying a gonopore at base. Gonopods (Figs 6-9) highly complex. Coxite elongate, subcylindrical, setose distoventrally; cannula normal. Telopodite strongly unciform, with a hypertrophied, as usual densely setose prefemoral part somewhat longer than femorite. A few undu- lations on ventrocaudal face and a few longitudinal ridges anterodorsally at base of solenophore. Solenophore coiled, ventrally with both a slightly folded base of lamina medialis (m) and a subtriangular parabasal lobe (p) similarly well-developed; p followed by a spine (k) supporting the tip of a flagelliform solenomere at base of a hyaline lobule (h), this lobule supporting the penultimate !/4 extent of both lamina lateralis and lamina medialis; distalmost 1/4 extent of both laminae slender, unciform, free of support. Remarks: Judging from the colour pattern and gonopod structure, this new species seems to be especially close to C. mesoxanthum, but both differ in size, in structure of the process between coxae 6 of the d, and in direction of the apical papillae on the epiproct (see also key below). Catharosoma mahnerti sp. n. Figs 10-17 Material: Holotype ¢ (MHNG), Paraguay, Prov. Concepcion, Estancia Laguna Negra, 15 km E of Paso Barreto, dead wood, 13.10.1985, leg. Expédition zoologique du Muséum de Genève. — Paratypes: 1 d,2 22 (MHNG), 1 d,1 2 (ZMUM), same locality, together with holotype; 1 d (MHNG), Prov. Concepciön, near mouth of Ao River, Trementina (Estancia Laguna Negra), forest, 13.10.1985, leg. Expédition zoologique du Muséum de Genève. Name: Honours Dr Volker Mahnert, who entrusted me this valuable material for study. Diagnosis: Differs from congeners by the submoniliform body, the longer antennae, the absence of a distinct colour pattern and of an unciform hypoproct, the presence of two paramedian pilose ridges between coxae 3 of the d, and in certain details of gonopod structure. Description: Length 20-23 mm (d, £), width of midbody metazona 1.7 (d) to 2.3-2.5 mm (2); © usually larger and somewhat broader than d. Holotype ca 20 mm long and 1.7 mm wide. Coloration in alcohol uniformly light brown to red-brown; only penultimate antennomere rather dark brown, tip of antenna pallid; legs paler, yellow- brown to brown. Body subcylindrical, submoniliform. Postcollar constriction faint, width of head = 5-16 > collum = 4 > 2 = 3; on segments 17—20 trunk gradually and gently tapering toward telson both in width and in height. Antennae medium-sized, slender, in situ reaching beyond segment 3 dorsally (Fig. 10), a little shorter in 2. Paraterga 814 S. I. GOLOVATCH Fics 10-17 Catharosoma mahnerti sp. n., 3 paratype: 10) anterior body portion, lateral view; 11) metater- gum 10, dorsal view; 12) telson, lateral view; 13) hypoproct, ventral view; 14) sternal structures between coxae 2-7 (legs 2 at left); 15) sternal structures between midbody coxae; 16) leg 15; 17) right gonopod, medial view. Scale bars 1.0 (10-16) and 0.2 mm (17). strongly reduced, only on segment 2 fully developed as low keels devoid of a caudal tooth, discernible on segments 3 and 4 as arcuated lines, on segment 5 as modest swellings (Fig. 10), on following segments totally missing. Tegument smooth, shining, only metazona at places faintly rugulose. Limbus thin, caudal margin entire. Metaterga fully devoid of a transverse sulcus; setae short, often abraded, arranged 3+3 on seg- ments 2 and 3, on following segmens 2+2 in a transverse line behind a rather deep stricture between pro- and metazona (Fig. 11). Ozopores lateral, opening level to metatergal surface ca 1/3 metazonital length away from caudal edge (Fig. 10). Pleurosternal carinae like narrow keels with a caudal spinule (Fig. 10) traceable until segment 16 (2) or 17-18 (6), onward wanting. Epiproct (Fig. 12) rather long, coni- form, faintly emarginate at tip in dorsal view, in 6 a little longer than in 2 . Hypoproct PARADOXOSOMATIDAE IN PARAGUAY 815 (Figs 12, 13) subtriangular, tip narrowly rounded, nearly pointed, straight; 1+1 para- median setae at caudal edge poorly separated. Paraterga, pleurosternal carinae and hypoproct a little more strongly developed in d than in 9. Sternal structure (Figs 14, 15) much like in C. bilineatum sp. n., but with two paramedian pilose ridges between coxae 3 of d and paramedian bunches of setae on poorly developed knobs between coxae 6 of à. Legs without tarsal brushes but densely setose ventrally; setation gradually thinning out toward telson; tibiae swollen ventrally and supporting characteristic brushes (Fig. 16) between leg-pair 7 and two last pairs. Legs in d a little longer than in 2, as usual becoming a little longer and slenderer toward telson. Each coxa 2 of 4 with a strong, distoventral, sigmoid, apically pointed process carrying a gonopore at base (Fig. 14). Gonopods (Fig. 17) highly complex, much like in C. bilineatum n. sp., but cox- ite more elongate, telopodite circular and elongate, while solenophore ventrally with a much larger, hyaline lobe/base of lamina medialis (m), an apically strongly unciform parabasal lobe (p) followed by a spiniform process supporting a flagelliform solenomere at base of a hyaline lobule (h), this lobule supporting the penultimate 1/4 extent of both lamina lateralis and lamina medialis; distalmost 1/4 extent of both laminae likewise slender, unciform, free of support. Remarks: Judging from the particularly elongate gonopod telopodite, which includes the femorite, this new species seems to be especially close to C. curitibense and C. intermedium, but the circular telopodite and the shape of its individual elements in C. mahnerti sp. n. are quite characteristic. At first Jeekel (1963) merged Catharosoma and Mestosoma together, but later (1968) he separated them on the account of the apparently more apomorphic character states observed in Mestosoma species. This viewpoint has also been shared by Hoffman (1980). Broelemannopus minutus sp. n. Figs 18-25 Material: Holotype ¢ (MHNG), Paraguay, Alto Parana Prov., Forestry School, 12 km S of Puerto Presidente Stroessner, soil sample 1, 04.1983, leg. C. Dlouhy. — Paratypes: 20 6 G , 22 22,2 juv. (MHNG), 2 dd,2 2 2 (ZMUM), same locality, together with holotype; 1 d,1 © (MHNG), same locality, soil sample 8, 04.1984; 2 4 4,1 2 (MHNG), same locality, soil sample 24,06.1984; 3 4 d (MHNG), same locality, soil sample 10, 05.1984; 1 © ,3 juv. (MHNG), same locality, soil sample 22, 25.06.1984; 1 dg, 1 2 (MHNG), same locality, soil sample 25, 25.06.1984; 1 d (MHNG), same locality, soil sample 7, 04.1984; 1 d ,3 juv. (MHNG), same lo- cality, soil sample 23, 25.06.1984; 2 6 4 (MHNG), same locality, soil sample 3, 21.05.1984; 1 3,1 2,1 juv. (MHNG), same locality, soil sample 13,04.1984; 1 d ,1 2, 1 juv. (ZMUM), same locality, soil sample 30, 04.1984; 14 dd, 17 22 (MHNG), same locality, soil sample 12, 04.06.1984, all leg. C. Dlouhy. Name: To emphasize the small body size. Diagnosis: Differs from congeners by the particularly small size, the especially strongly reduced paraterga visible only on segment 2, the lack of transverse metatergal sulci, the absence of a distinct colour pattern and of an unciform hypoproct, and in certain details of solenophore structure. Description: Length 7-9 mm (d, 2), width of midbody metazona 0.6-0.7 (4) to 0.8-0.9 mm (2); 2 usually larger and somewhat broader than d . Holotype ca 8 mm 816 S. I. GOLOVATCH Fics 18-25 Broelemannopus minutus sp. n., d paratype: 18) anterior body portion, lateral view; 19) caudal body portion, lateral view; 20) sternal structures between coxae 3-7 (coxae 3 on top); 21) ster- nal structures between midbody coxae; 22) leg 10; 23-25) right gonopod, medial, lateral and an- terodorsal views, respectively. Scale bars 0.5 (18-19), 0.25 (20-22) and 0.1 mm (23-25). long and 0.7 mm wide. Coloration in alcohol uniformly brown to red-brown, only juveniles, antennae and legs paler, yellow-brown to brown; tip of antenna pallid. Body subcylindrical, not moniliform. Postcollar constriction faint, width of head > collum > 2 = 3 < 4 < 5-16; on segments 17-20 trunk gradually and gently tapering toward telson both in width and in height. Antennae medium-sized, slender, in situ reaching the end of segment 2 dorsally (Fig. 18), a little shorter in 2. Paraterga strongly reduced, only present as low lines devoid of a caudal tooth on segment 2 (Fig. 18), on following segments totally missing. Tegument smooth, shining. Limbus thin, caudal margin entire. Metaterga fully devoid of a transverse sulcus; setae relatively long, sometimes abraded, arranged 2+2 on segment 2, on following seg- ments 1+1 in a transverse line behind a shallow stricture between pro- and metazona (Fig. 18). Ozopores lateral, opening level to metatergal surface ca 1/3 metazonital length away from caudal edge. Pleurosternal carinae like narrow arcuated lines (Fig. 18) traceable until segment 16 (2) or 17 (3), on following segments wanting. Epiproct (Fig. 19) rather long, coniform, in ¢ a little longer than in £, tip with a stong claw- shaped uncus directed dorsocaudally. Hypoproct (Fig. 19) subtriangular, tip narrowly rounded, nearly pointed, straight; 1+1 very long paramedian setae at caudal edge poorly separated from each other. Paraterga, pleurosternal carinae, epi- and hypoproct in d a little more strongly developed than in 9. PARADOXOSOMATIDAE IN PARAGUAY 817 Sterna mostly sparsely setose. Sternum between coxae 3 of d furnished with several transverse rows of relatively small, subcontiguous, scale-like structures on a boss (Fig. 20). Sterna between coxae 4 and 5 of d with a very prominent, linguiform, laterally setose outgrowth directed anteroventrad (Fig. 20); a paramadian pair of bunches of setae between coxae 5 and 7 of d (Fig. 20); a central, compact and thicker bunch of setae on a swelling between coxae 6 of d (Fig. 20). Postgonopodial sterna without modifications (Fig. 21). Legs without tarsal brushes but densely setose ventrally, without modifications (Fig. 22); setation gradually thinning out toward telson; legs in 4 a little longer than in 2, as usual becoming a little longer and slenderer toward telson. Each coxa 2 of d nearly unmodified, with a small distoventral cone surmounted by a gonopore. Gonopods (Figs 23-25) not particularly complex. Solenophore ventrally with a distinct but not hypertrophied, rounded, hyaline lobe/base of lamina medialis (m); another simple, parabasal lobe (p) supporting the tip of a flagelliform solenomere at base of a smaller hyaline lobule (h), the latter supporting the subterminal part of both lamina lateralis and lamina medialis; distalmost extent of both laminae very short and rather broad, subunciform and subtruncate. Remarks: Judging from the particularly small size, the similarly uncigerous epiproct, the paraterga present on segment 2 only, as well as from several other traits, including those of the gonopod, the new species seems to be especially close to B. ibitiensis, from Monte Alegre, Sao Paulo State, Brazil (Schubart, 1945a). The distinctions of B. minutus lie in the total absence of sternal cones on postgonopodial segments of the d, as well as the presence of a smaller and differently armed lobe between coxae 3 of the d, and of a smaller lobe p on the solenophore (see also key below). B. ibitiensis has only tentatively been placed in Broelemannopus (see Jeekel, 1963) but, since this species shows especially close affinities with B. minutus sp. n., there can no longer be any doubt that both are congeners best to be assigned to Broelemannopus. In general the genus Broelemannopus is so poorly distinguished from Mestosoma that at first Jeekel (1963) merged these two genera together and treated the former as only a species group of the latter. However, later (1968) he separated them on the basis of certain apomorphies observed in both (see key below), a view which is also shared by Hoffman (1980). Mestosoma simplex sp. n. Figs 26-32 Material: Holotype d (MHNG), Paraguay, Prov. Cordillera, 5 km N of Emboscada, Rio Piribebuy, forest with bromeliads, dead wood and leaves, 05.04.1985, leg. Expédition zoologique du Muséum de Genève. — Paratype: 1 juv. (MHNG), same locality, together with holotype. Name: To emphasize the only slightly modified legs of the 4 and the relatively simple gonopod structure. Diagnosis: Differs from congeners by the relatively small size, the only slight- ly modified legs of the d, and the relatively simple solenophore structure. Description: Length of holotype ca 12 mm, width of midbody metazona 1.5 mm. Coloration in alcohol uniformly light red-brown, anterior body portion 818 S. 1. GOLOVATCH Fics 26-32 Mestosoma simplex sp. n., è holotype: 26) anterior body portion, lateral view; 27) left halves of segments 10 and 11, dorsal view; 28) sternal structures between coxae 4 and 5 (coxae 4 on top); 29) leg 15; 30-32) right gonopod, medial, submedial and lateral views, respectively. Scale bar 0.6 (26-29) and 0.3 mm (30-32). slightly more intensely reddish; penultimate antennomere darker brown, tip of antenna pallid; legs paler, yellow-brown. Body strongylosomoid, not moniliform, with poorly developed but evident paraterga. Postcollar constriction apparent, width of head = collum > 2 = 4 << 5-16; on segments 17-20 trunk gradually and gently tapering toward telson both in width and in height. Antennae medium-sized, slightly clavate, in situ reaching beyond segment 2 dorsally (Fig. 26), a little shorter in juvenile. Paraterga 2 and 5-19 delimited by an evident sulcus not only dorsally but also ventrocaudally (Fig. 26), all set low, considerably larger on pore-bearing segments than on poreless ones (Fig. 27), only on segments 18 and 19 slightly projecting beyond rear tergal contour. Tegument smooth and shining. Limbus thin, caudal margin entire. Metaterga 5-17 with a faint but PARADOXOSOMATIDAE IN PARAGUAY 819 apparent transverse sulcus far from reaching base of paratergum, the sulcus being very slightly sinuate anteromedially; axial impression on metaterga interrupted in the mid- dle, barely visible; tergal setae untraceable (Figs 26, 27). Stricture between pro- and metazona thin and shallow (Figs 26, 27). Ozopores lateral, lying on paraterga at ca 1/4 metazonital length away from caudal edge (Figs 26, 27). Pleurosternal carinae like narrow keels delimited by a sulcus dorsally, devoid of a caudal spinule (Fig. 26), trace- able until segment 17, on further segments wanting. Epiproct rather long, coniform, narrowly truncate at tip in dorsal view, in d a little longer than in juvenile. Hypoproct roundly triangular, tip evidently rounded, straight; 1+1 paramedian setae at caudal edge rather strongly separated from each other. Sterna modestly setose, mainly unmodified but both with an evident, deeply emarginate, setose, linguiform outgrowth directed anteroventrally between coxae 4 of 3 and with a paramedian pair of bunches of setae between coxae 5 of d (Fig. 28). Legs only slightly modified, in d with dense tarsal brushes starting from leg- pair 1 and with dense tibial brushes starting from leg-pair 2 (Fig. 29), both these brushes thinning out toward telson but absent only from last two leg-pairs. Each coxa 2 of & with a low distoventral cone bearing a gonopore. Gonopods (Figs 30-32) relatively simple. Coxite subcylindrical, not particularly elongate, sparsely setose distoventrally. Telopodite subfalcate, a densely setose prefemoral part normal in shape, much shorter than femorite. Solenophore unciform, somewhat constricted at base, relatively stout and simple; lobe m at base of both solenomere and lamina medialis highly inconspicuous, ledge-shaped, lamina lateralis slightly better developed than lamina medialis, tips of both subtruncate. Remarks: Due to the basally strongly constricted solenophore, M. simplex sp. n. joins group III in the sense of Jeekel (1963), which has hitherto been known to comprise only three species, one each from the Brazilian states of Pernambuco, Alagoas and Sao Paulo. However, the new species differs in its smaller size, in the absence of femoral and some sternal modifications in the d, and in the particularly simple gonopod structure. Mestosoma crassipes sp. n. Figs 33-42 Material: Holotype d (MHNG), Paraguay, Prov. Concepciön, Ao. Tagatya-mi, small grove, near “gué”, sifted litter, 23.10.1985, leg. Expédition zoologique du Muséum de Genève. Name: To emphasize the modified and inflated legs of the 4. Diagnosis: Differs from-congeners by the relatively small size, the conspicu- ously modified legs of the d, and by the relatively complex gonopod structure. Description: Length of holotype ca 12 mm, width of midbody metazona 1.3 mm. Coloration in alcohol uniformly light brown; antennae brown, penultimate an- tennomere darker brown; tip of antenna pallid; legs slightly paler. Body strongylosomoid, not moniliform, much like in M. simplex sp. n. Post- collar constriction apparent, width of head ? collum = 4 > 2 = 3 <5 < 6(7)-16; on seg- ments 17-20 trunk gradually and gently tapering toward telson both in width and in height. Antennae medium-sized, slightly clavate, in situ reaching beyond segment 2 dorsally (Fig. 33). All paraterga set low, delimited by an evident sulcus not only dorsally but also ventrally to ventrocaudally (Fig. 33), on segment 19 particularly flat 820 S. I. GOLOVATCH Fics 33-42 Mestosoma crassipes sp. n., ¢ holotype: 33) anterior body portion, lateral view; 34) left half of metatergum 10, dorsal view; 35) caudal body portion, lateral view; 36) sternal structures be- tween coxae 3 and 4 (coxae 3 on top); 37) leg 7; 38) leg 9; 39-42) left gonopod, submedial, sub- lateral, ventrolateral and anterodorsal views, respectively. Scale bars 1.0 (33-35),0.5 (36-38) and 0.3 mm (39-42). in lateral view, considerably larger on pore-bearing segments than on poreless ones (Figs 33, 34), never projecting beyond rear tergal contour. Tegument smooth and shin- ing. Limbus thin, caudal margin entire. Metaterga 5-18 with a faint but apparent trans- verse sulcus far from reaching base of paratergum, the sulcus being slightly sinuate an- teromedially; axial impression on metaterga wanting; tergal setae untraceable (Figs 33, 34). Stricture between pro- and metazona thin and shallow (Figs 33, 34). Ozopores lat- eral, lying on paraterga at ca 1/4 metazonital length away from caudal edge (Figs 33, 34). Pleurotergal carinae like narrow keels delimited by a sulcus dorsally, devoid of a caudal spinule (Fig. 33), traceable until segment 17, on following segments wanting. Epiproct rather long (Fig. 35), coniform, narrowly truncate at tip in dorsal view. Hypoproct triangular, tip pointed, straight; 1+1 paramedian setae at caudal edge widely separated from each other (Fig. 35). Sterna mainly modestly setose, postgonopodial ones unmodified, pregonopodi- al ones with a paramedian pair of bunches of setae on small cones between coxae 3 and PARADOXOSOMATIDAE IN PARAGUAY 821 4 of & (Fig. 36); sterna between coxae 5-7 of d concave and with some particularly long setae (like in Fig. 36). Legs modified, with dense tarsal brushes starting from leg-pair 1 and with dense tibial brushes starting from leg-pair 2 (Figs 37, 38), both these brushes thinning out to- ward telson and virtually absent only from last two leg-pairs. Femora 4-6 conspicu- ously inflated and each with an evident distoventral tooth. Legs 7 (Fig. 37) with a dis- toventral cone on each coxa and a little less strongly incrassate femora, the tooth situ- ated more ventro-apically. Femora 9-11 (Fig. 38) nearly normal, ventral knobs in dis- tal one-third gradually disappearing toward leg-pair 12. Each coxa 2 with a low dis- toventral cone bearing a gonopore. Gonopods (Figs 40-42) somewhat more complex than in the previous congener, especially so due to a longitudinal sulcus on dorsal face of femorite, a digitiform process (m), not a ledge-shaped structure lying at base of lamina lateralis, as well as a short, unciform and pointed solenophore tip. Remarks: This new species seems to be particularly close to M. alticola and to a few other congeners possessing a small but evident process (m) at base of both the solenophore and solenomere (see Attems, 1937). The colour pattern in M. crassipes sp. n. resembles that of M. femorale, yet the gonopod structure is quite different (see Schubart, 1943). Due to the presence of two small tubercles between coxae 3 of the d and an attenuated tip of the solenophore, the new species is similar to M. perfidum, but the latter species is larger (17-18 mm long and 1.9-2.0 mm wide versus 12 and 1.3 mm), and their gonopods are very different as well. IDENTIFICATION KEYS KEY TO TRIBES AND GENERA OF PARADOXOSOMATIDAE OCCURRING IN PARAGUAY AND ADJACENT AREAS 1 Paraterga strongly developed, wing-like. (Introduced, normally synan- TOPIC, SPECIES) FR El Nee ENTE SL OR ER: 2 - Paraterga poorly developed to missing (Figs 1, 10, 18, 26, 33) (Tribe Catharosomatinisautochthonous;species) IE an we. 4 2 Metaterga densely papillate and setose; a bituberculate process between coxae 4 of d ; femora 4-7 of d each with a ventro-parabasal knob; gono- pod femorite stout, solenophore with a large basal prong ventrally 2. bed MR: TATE Qi oi N LL Tribe Sulciferini: Chondromorpha (C. xanthotricha) = Metaterga not papillate, at most poorly setose; sternum between coxae 4 of d without process; legs of 4 without such femoral tubercles; gono- femorite not so stout, almost as long as to longer than solenophore . ....... 3 3 Caudal corners of paraterga surpassing rear tergal contour only on cau- dalmost segments (16-19); gonofemorite relatively short, distinctly broadened distally; solenophore bifid, at base with a long apical process as well as a prong and a lobe ventrally . . Tribe Sulciferini: Oxidus (O. gracilis) 2 Caudal corners of paraterga surpassing rear tergal contour on all body segments; gonopod telopodite very slender and simple, rod-shaped; go- nofemorite slightly longer than solenophore, latter without outgrowths dubase Dut witha minute lobule-attpr RR le Sate oe SSN DAME ER. 30 SOE Ie Tribe Orthomorphini: Orthomorpha (O. coarctata) 822 S. I. GOLOVATCH Gonopod devoid of a solenophore, solenomere free ................... D Gonopod with a solenophore sheathing and supporting a flagelliform solenomerer....22..2: 2 n er IEEE 9 Gonopod telopodite extremely simple, subfalcate to subcircular, devoid of any outgrowths at base of a subflagelliform solenomere ... Gonodrepanum Gonopod telopodite more complex, with 1-2 processes or dilatations in femorallorpostfemoraliregion®. 2/3 CE. wa N 6 Gonofemorite with a large, medial, lobuliform dilatation; solenomere Wwithrarsmalletoothineartip iss aes Be BEE Mogyella (M. nana) Gonofemorite without a large dilatation but sometimes with a tooth dis- tally or parabasally; 1-2 dilatations or processes at base of solenomere, latter sometimes stout and'thickK®® SCE NME | ee EEE Re 7 Body small, 0.7 mm wide. Gonopod telopodite subfalcate, much like in Gonodrepanum; femorite with a small ventro-parabasal tooth; a small but evident, elongated, subtriangular dilatation/lobe fused ventrally at base with a flagelliform solenomere ....... Pseudogonodrepanum (P. scitum) Body medium-sized, >1.0 mm wide. Gonopod telopodite mostly sub- erect, usually with two independent teeth/outgrowths at base of soleno- Entire gonopod telopodite suberect, with an evident process at base of a rather thick, sometimes bifid solenomere; a setose, linguiform, sternal Processibetweenteoxae ATO Habrodesmoides Only gonopod femorite elongated and suberect; solenomere subflagelli- and unciform, acuminate, at base with two relatively small teeth/out- growths; sternal process between coxae 4 of 4 either missing or repre- sentedibytwoiparamedian'knobst e Ologonosoma Solenophore simple, subflagelliform, acuminate, supporting a similarly Haselliform:solenomere...... ur. lecita era E 10 Solenophore complex, sometimes with a process or lobe at base, usually consisting of several more or less folded lobes/laminae, almost entirely sheathing a flagelliform solenomere 20 RNC 11 Body width 0.7 mm. Prefemoral (setose) part of gonopod much shorter than acropodite; femorite with a conspicuous lamella on medial side; both solenophore and solenomere somewhat reduced, about half as long AS Pe MONIC ge ct g tc tata Mogyosoma (M. hamatum) Body width 1.1-1.5 mm. Prefemoral portion of gonopod elongated, nearly as long as acropodite; femorite without a lobe on medial side; both solenophore and solenomere longer than femorite ............. ann este are eg Gonodrepanoides (G. travassosi) Solenophore relatively simple but with a large lateral branch/process at aS gables nerina iii ernennt Promestosoma (P. boggianii) Solenophore more complex, usually consisting of several lobes but devoidiof.alarselateral\branchtatibaser 162 CP MERE See RREEREE 12 Sternal cones usually present (Figs 4 & 15); each coxa 2 of d with a strong distoventral process carrying a gonopore (Figs 1 & 14); tibiae of 13 PARADOXOSOMATIDAE IN PARAGUAY 823 d usually conspicuously inflated (Figs 5 & 16). Prefemoral (densely setose) portion of gonopod hypertrophied, about as long as femorite RO AIM MTS OEM TREE SODIO, EROS, de) SES rL OI Catharosoma Sternal cones usually absent; normally each coxa 2 of d with a small cone carrying a gonopore; tibiae of d not inflated. Prefemoral portion of gonopod normal, shorter than femorite 2 529 un... oes aa an en 13 Tarsal brushes often absent even in à ; sternite between coxa 3 of 4 with a conspicuous comb- or tongue-shaped structure (Fig. 20); sternite between coxae 5 of & with a particularly large process directed antero- ventrally and covering a boss or protuberance if any between coxae 4 5-010 90 GEES RE O RS RE LN Te Reh GS Me Pree aE AT eae Broelemannopus Tarsal brushes in d mostly present; sternite between coxa 3 of d usually without protuberances; a particularly large process directed anteroven- trally and placed between coxae 4, not 5, of d ................ Mestosoma KEY TO BROELEMANNOPUS SPECIES OF PARAGUAY AND ADJACENT AREAS 1 Body of adult < 10 mm long and < 1.0 mm wide; epiproct topped with a prominent claw-shaped uncus directed slightly dorsad (Fig. 19); cones absent at least between front coxae of postgonopodial sterna ............. 2 Body > 14 mm long and > 1.6 mm wide; epiproct without central uncus at tip, latter usually bifid; sternal cones in d present between both pairs oigleostolecachisegment Brazil... RT 3 Lobe between coxae 4 and 5 of d linguiform, entire (Fig. 20); sternal cones on midbody segments of d totally absent (Fig. 21); solenophore lobeyp small (Figsi23=25): Paraguay aptly. dor ee E B. minutus Lobe between coxae 4 and 5 of d divided distally; sternal cones in à present between posterior coxae of segments 8—18; solenophore lobe p larce#Sa0}PauloiState Brazil eee tenes B. ibitiensis Body 1.6-2.2 mm (d) or 1.6-2.4 mm wide (9). Paraterga traceable as low arcuated sulci on segments 2-4, on following segments only as slight impressions visible near caudal margin; pleurosternal carinae (raccablesuntiliseomentil 4 So ee Peete B. pirassunungensis Body 2.3-3.3 mm wide (d, ®). Each paratergite traceable as a low arcuated crest at least on segment 2; pleurosternal carinae visible until SELMEN WON. are taggare SEHR Sr Anti re Ton 4 Sternal lobe between coxae 3 of d comb-shaped, entire distally; coxae 2-6 of d each with a distoventral coniform process; central protuberance between coxae 7 of d absent; distal end of solenophore not divided VIII rotante B. escaramucensis Sternal lobe between coxae 3 of d linguiform, deeply incised distally; only coxae 2 and 3 of d each with a coniform process; central protu- berance between coxae 7 of d present; distal end of solenophore deeply divided into two large lobes, p being one of these ............. B. glabratus 824 S. I. GOLOVATCH KEY TO CATHAROSOMA SPECIES 1 10 Sternum between coxae 6 of d with a single, undivided process or OUT ro 2a e N ee N 00000 0 2 Sternum between coxae 6 of d either with a deeply divided process or with two distinct paramedian processes, or with two paramedian knobs with) bunches of long setae ..... 2.0.00. 00. 2 hs ee | I Hypoproct tip unciform; gonopod femorite with a prominent, subtrian- eularadıstaklobe 7 roi Geen DIRE C. digitale Hypoproct tip straight; gonofemorite without a prominent, subtrian- sularsdistalllebe =... cirrosi STE 3 Body width about 2.2 mm; no sternal modifications except pilosity be- tween coxae 5 of d; tip of solenophore simple, like a broad and sub- ICUMIMMCANODE nn C. mixtum Body width 3.0-3.2 mm; a swelling and/or a paramedian pair of knobs between coxae 5 of 6; tip of solenophore more complex —erpreree er 4 Paraterga 2 like swellings, paraterga 3 and 4 sulciform; light subtrian- gular spots on posterior parts of proterga and on anterior parts of metaterga against a dark background; apical piece of solenophore short, acuminate RR, Ne MN Ret 10 eft00 (ho eet IN C. peraccae Paraterga 2-4 like low crests; background coloration pale yellowish with a wide castaneous axial stripe; apex of solenophore very broad and of tatherirte sularıshape:... .. 2 oh) axa e's nen aoe IE ORE C. mesorphinum Even paraterga 2 expressed as sulci; no sternal modifications except pilosityabetweenicoxae SION ERRE Co PER RME ee C. myrmekurum Crest-like paraterga present at least on segment 2 (Figs 1 & 10), often ridge/crest-like even on segments 2-4; sternal modifications present between coxae stot Gis: AMOR SL WRG, SU RER 6 Paraterga 2-4 distinctly crest-like, onward sulciform on segments 5, 7,9 and 10; a distinct bilobed process present between coxae 3 of dC. palmatum Paraterga even more poorly developed, totally untraceable on segments behind AR ANT, JR N ED AO eo E 7 Body about 12 mm long and 1.3 mm wide; coxae 2 of d produced distally into a small mammiform process; gonofemorite with a medio- parabasaldisitiform process er RARE RP RE C. curitibense Body width > 1.7 mm; coxae 2 of 4 usually produced into a more or less prominent process (Fig. 14); gonofemorite devoid of such a process ....... 8 Hypoproct uneiform; its tip directed ventrad? Sto 2808 NE 9 Hypoproct:tip’straight 2... 0) IR OS, DO ee ee ee 12 Dorsum rather dark, red-brown, with two yellowish paramedian stripes; pleurosternal carinae visible only until segment 15............ C. taeniatum Dorsum pale but with two dark, brownish paramedian stripes; pleu- rosternal’carinae visible at least until’sesment16 Re 10 Body of d 1.9-2.1 mm wide; a large and deeply incised process present belweenicoxae 6.086,68 (Fie. 3). | ot ee eee C. bilineatum 11 12 PARADOXOSOMATIDAE IN PARAGUAY 825 Body width = 2.6 mm; only a pair of small paramedian knobs between COX OO CK Gaudin sa as Soe sheds, tele: 11 Apical papillae on epiproct acuminate, elongate and directed distodor- sad; gonofemorite strongly broadened distad .............. C. mesoxanthum Apical papillae on epiproct small, simple, inconspicuous; gonofemorite Stendere IE Vate EEE TI rain cea EEE C. intermedium A paramedian pair of low setigerous crests present between coxae 3 of d; gonopod telopodite elongate and circular (Fig. 17) ......... C. mahnerti At most a paramedian pair of bunches of setae present between coxae 3 Of 7 gonopod telopodite falcate and stouti Elric ein... 13 A single protuberance between coxae 5 of d and a paramedian pair of Conicaliprocesses!betweenicoxae CON di line e C. palustre Two setigerous knobs on a swelling between coxae 5 of d and either a swelling or a paramedian pair of spinules between coxae 6 of d ......... 14 Body width 2.0-2.3 (¢) to 2.5 mm (2); lobe p of solenophore ancori- COMANO I UE a SE C. paraguayense Body width 2.6 mm (d); tip of lobe p of solenophore blunt, devoid of LECCESE RE SURE ee OBER RAR (i hme. ME eee en C. hoffmani KEY TO GONODREPANUM SPECIES 1 Coloration of adults normally castaneous brown with a wide, uninter- rupted, axial, contrastingly creamy stripe; Rio de Janeiro, Brazil .......... 2 Coloration uniformly castaneous to black, usually devoid of a contrast- ingly pallid stripe, or axial stripe/line thin and interrupted; Argentina Ain OTA LAZU en ee Bun en RO I I EN: 3 Smaller, only known from 2: 16-18 mm long and 1.8-2.0 mm wide; clear sulciansplace of paratersay 2 4a na ug G. flavolineatum 2 18-21 mm long and 2.2-2.4 mm wide, d 15-16 mm long and 1.5-1.6 mm wide; paraterga on segment 2 like crests, on segments 3 and 4 like clear sulci, on following segments like striae; paramedian pairs of setigerous protuberances present between coxae 3-5 of d ....... G. torresae Body particularly slender: ¢ 20 mm long and 1.2 mm wide. Dorsum and sides uniformly blackish; paraterga 2-4 crest-like, on following segments sulciform; sternal cones absent; Brazil and Argentina....... IE ee eye! ER AE E G. drepanephoron Body neither so long (< 14 mm even in ©) nor so slender. Coloration of adults usually castaneous, sometimes yellowish; paraterga represented Dyzsulcgaumost3sternalleonesipresent?Brazilere u 0 ae 4 Epiproct topped with two long, claw-shaped, diverging unci directed dossocaudadzahpaleraialistape/linezabsentz un nsf oa aces 5 Epiproct without such unci; a pale, axial, interrupted stripe/line usually [DIALS ale ES es PO OG SEMI LARIO RR 6 Body 12-14 mm long and 1.3-1.7 mm wide; a small ventral tubercle present only on femora 3 and 4 of à ; sternal protuberance between coxae 826 S. I. GOLOVATCH 3 of d bifid, sterna between coxae 4 and 5 devoid of protuberances; solenomeressubeiteular Sy ICE G. grajahuense Body 9-12 mm long and 1.0-1.3 mm wide; a distoventral tubercle present/traceable on femora 3-9 of d; median sternal protuberances present between coxae 3-5 of d ; solenomere regularly and gently sub- CRIARI an hehe stake NC OR CE EEE G. furcatum Body about 14 mm long and 1.3-1.5 mm wide. Metatergal sulci absent; femora of d not modified; a large median process with two knobs behind and between coxae 4 of d; solenomere relatively short and subereetSanta Catarina; Brazile Jae eae RIO ERA G. levisetum (G. levisetum var. coniferum seems to only represent a colour form dis- tinguished by a yellowish body devoid of a pattern) Body 11-12.5 mm long and 1.1 mm wide. Metatergal sulci light but present; a pair of diverging processes between coxae 5 of à ; solenomere relatively long, subcircular; Rio de Janeiro, Brazil ........... G. falciferum KEY TO HABRODESMOIDES SPECIES OF PARAGUAY AND ADJACENT AREAS | Paraterga replaced by a low arcuated ridge on segment 2, by sulci on segments 3 and 4; pleurosternal carinae traceable until segment 18; a small protuberance present between coxae 5 of d ; solenomere retrorse, with a slender, subfalcate, simple process at base ............ H. costalimai Segments 2-18 with only slight sulci/striae in place of paraterga; pleuro- sternal carinae present on segments 2-7; sternite between coxae 5 of d devoid of a protuberance; solenomere directed distoventrad, process at itsibase massiveystoutand branched! 214.000 73 IRE AU H. perturbans KEY TO MESTOSOMA SPECIES OF PARAGUAY AND ADJACENT AREAS 1 Sterna between coxae (3)4-6(7) of d each with a more or less distinct median process or protuberance; sternal cones in d absent.............. 2 At least some of these sterna devoid of a protuberance; sternal cones in Sf oltenipresent n. LUE ET RT IP PRE RE 5) Body width > 2.3 mm; metatergal sulcus present on segments subse- quent to 5!h; no single sternal process between coxae 6 and 7 of d........ 3 Body width < 1.5 mm (d); metatergal sulci absent; a single, prominent, sternal process between coxae 6 and 7 of d; Bolivia ................... 4 Coloration uniformly brownish; sternal processes or protuberances present between coxae 3-6 of ¢; Rio de Janeiro, Brazil ......... M. carioca With a wide pale axial stripe against a dark olive-brown background; sternal process present between coxae 4 of d, only protuberances or swellings between coxae 5 and 6 in gd; Bolivia.............. M. schindleri Body width 1.0 mm (dé); colour pattern in adults indistinct, body uni- formly yellowish; femora 3-5 of 4 each with a ventral tubercle . . . M. alticola Body width about 1.5 mm (d); a wide, pale, axial stripe against a cho- colate-brown background; femora in d apparently without ventral tubercles’ PI ee Ss a ee a PNR ee eee M. boliviae ZI I OO 12 13 PARADOXOSOMATIDAE IN PARAGUAY 827 Sterna of segment 6 in d with a pair of processes at anterior border, sterna of segment 5 of d devoid of processes; neither tibial nor tarsal bzushes;presenter. eriwellch bue sen. 2.2 ii oly. tr M. differens Sterna of segment 6 in d without processes, those of segment 5 of d with or without processes; tibial and tarsal brushes in d normally present . . . 6 Colour pattern relatively distinct, usually entire dorsum or axial stripe pale to yellowish, sometimes this stripe divided into two paramedian stripes or broken into series of spots, more or less strongly contrasting with a dark, normally brown background coloration of the sides .......... 7 Colour pattern indistinct, normally dorsum and sides uniformly brown . ... 14 Sternal process between coxae 4 of d present, conspicuous .............. 8 Sternal process between coxae 4 of ¢ absent to very small ............. 10 Body width < 2.0 mm. Dorsum with axial stripe(s), background coloration brown; gonofemorite not very broad, solenophore evidently longerithansi/2zfcmonitena a sts Ya. rn ae ne 9 Body width = 2.0 mm. Dorsum with an axial series of yellowish spots, background coloration dark (red-)brown to black; gonofemorite conspicuously broadened, solenophore scarcely half as long as femorite; Rara CUA or ne e SIRIO RIE RATE DATO Ot 10 Dorsum with two yellowish paramedian stripes; tibial and tarsal brushes in d present; a bilobate process present between coxae 4 of d ; Paraguay anta LIRE LIL: M. vittatum Dorsum with two series of paramedian spots; tibial and tarsal brushes in d absent; a lobe-shaped process present between coxae 4 of 4 ; Bolivia à 0.020: 0 io 6e MDI Rene IA RN EEE M. montanum Solenophore with 2-3 prongs/spines subapically ...................... 11 End of solenophore subtriangular, often rounded ..................... 12 Legs uniformly blackish; a pair of paramedian knobs between coxae 3 of 3; femora 4-7 in d each with a ventral tubercle, each coxa 7 of d Withiadistoventral process ste m ee M. tricuspis Legs brown; a process present between coxae 4 of d ; apparently neither femora nor coxae of d with ventral tubercles/processes . . M. pseudomorphum Body width about 3.0 mm. Dorsum with a pale, sometimes yellowish axial stripe; solenophore strongly curved proximad, its end almost in gontactswithybasesoftemoriter een ee M. salvadorii Body width about 2.0 mm. Dorsum with an axial series of conspicuous yellowish spots; solenophore not so strongly curved ................... 13 Median spots situated on proterga and in anterior parts of metaterga; solenophore in medial view almost twice as slender as distofemoral part BERN REED. AIRE RAIN, SHARE TERN DADI RE AE DIMMI AE ARE EE M. camerani Median spots lying only in middle of metaterga; solenophore in medial view nearly as broad as distofemoral part ................... M. kalliston /NIsinolesprocessipresentibetWeenicoxae lot OS CR ee 15 Either two cones/tubercles or nothing between coxae 4 in gd ............ 17 Body width about 2.0 mm. Sternal process between coxae 4 of d short and subquadrate; solenophore bifid apically .................. M. balzanii 828 Wy 18 23 24 S. I. GOLOVATCH Body width 1.3-1.8 mm. Shape of sternal process between coxae 4 in d andtof solenophore different. ro. Ri 2 EE 16 Metatergal sulci distinct on segment 5 and following segments; sternal process between coxae 4 of d short and conical; gonofemorite much broaderthanvsolenophore re yest a EEE M. lugubre Metatergal sulci absent; sternal process between coxae 4 of d long and rectangular; gonofemorite considerably slenderer than solenophore M. borellii Solenophore base conspicuously constricted (Figs 30-32) ........ M. simplex Solenophore base broader- | i RAI ER 18 Body length about 12 mm, width 1.3 mm. Metatergal sulcus visible also on segment 18; pleurosternal carinae present until segment 17; a disto- ventral tubercle present on femora 4-12 of à , and a process on coxae 9 of d (Fig. 38); a characteristic process (m) at base of solenophore, tip OMsolenophore pointed (Figs'39=42)\ ree ne ee M. crassipes Body length at least 17 mm, width 1.8 mm. Metatergal sulci either absent or at most traceable only until segment 17; distofemoral tubercles present until leg 10 of d at most; coxae 9 of d normal; no process at base of solenophore, tip of solenophore more or less rounded ........... 19 Body length 30-34 mm, width 3.8-4.2 mm; tip of solenophore very broadlysubtruncate ME Rn com, SRE M. truncatum Body length < 29 mm, width < 3.5 mm; tip of solenomere not subtruncate . 20 Faint sternal cones traceable between coxae 3 and behind segment 7 of d ; distoventral tubercles present only on femora 9 and 10 of d . . M. perfidum Noteworthy sternal modifications absent; distoventral tubercles usually present only untilfemora FONC SE 21 Tip of solenophore bilobate, rather deeply emarginate/notched in the middle, forming more or less equal, rounded lobes . ter aso 22 ipiofisolenophore- differente" 3. 3465. OCR LE CETTE CEE 23 Coloration blackish, legs red-brown; both lobes of solenophore tip sub- equal: Boliviar. 1056 Sagen Se are oe Soleo Se RE M. derelictum Coloration dark brown, legs light brown; proximal lobe of solenophore tip somewhat smaller than distal one; Paraguay ............. M. pulvillatum Tip of solenophore subacuminate, narrowly rounded; Bolivia . . . M. luctuosum Tip of solenophore broadly rounded; Brazil and/or Paraguay ............. 4 Body coloration dark, legs and venter contrastingly pale; pleurosternal carinae traceable until segment 15; a distoventral tubercle present only on femora 7 of à ; ventral brushes on tibiae of d absent......... M. bicolor Body coloration brown, legs and venter only slightly paler than remain- ing body; pleurosternal carinae traceable until segment 16; distoventral tubercles present on femora 4-7 of d; ventral brushes present both on tibiae and) tarsi indi: sock eR ee ee ae RER M. femorale KEY TO OLOGONOSOMA SPECIES 1 Sternal process between coxae 4 of d missing; solenomere retrorse; Brazili(Parand).. s.c.« <<..." eee RC TER EEE O. iguassuense PARADOXOSOMATIDAE IN PARAGUAY 829 = Sternal process between coxae 4 of d represented by two paramedian knobs; solenomere directed distoventrad; Paraguay ............ O. sanctum ACKNOWLEDGEMENTS I am most grateful to Dr Volker Mahnert (MHNG), who entrusted me this valu- able material for study. Peter Schwendinger (MHNG) kindly edited an advanced draft. REFERENCES ATTEMS, C. 1898. System der Polydesmiden. I. Theil. Denkschriften der mathematisch-natur- wissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften 67: 221-482. ATTEMS, C. 1899. System der Polydesmiden. II. Theil. Denkschriften der mathematisch-natur- wissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften 68: 251-436. ATTEMS, C. 1901. Neue Polydesmiden des Hamburger Museums. Mitteilungen aus dem Natur- historischen Museum Hamburg 18: 85-107. ATTEMS, C. 1914. Die indo-autralischen Myriopoden. Archiv fiir Naturgeschichte 80: 1-398. ATTEMS, C. 1931. Die Familie Leptodesmidae und andere Polydesmiden. Zoologica (Stuttgart) 79: 1-149. ATTEMS, C. 1937. Myriopoda 3. Polydesmoidea I. Fam. Strongylosomidae. Das Tierreich 68: i-xxii, 1-300. ATTEMS, C. 1943. Myriopoden von Brasilien, gesammelt von E. Bresslau in den Jahren 1913/14. Senckenbergiana 26: 434-458. BROLEMANN, H. W. 1902a. Myriapodes du Musée de Sao Paulo. Revista do Museu Paulista (1901) 5: 35-236. BROLEMANN, H. W. 1902b. Myriapodes recuillis par M. E. Gounelle au Brésil. Annales de la Société entomologique de France 71: 649-694. BROLEMANN, H. W. 1929. Myriapodes recueillis au Brésil par M. le Professeur Gaullery, membre de l’Institut. Mémoires de la Société zoologique de France 29: 1-37. CARL, J. 1902. Exotische Polydesmiden. Revue suisse de Zoologie 10: 563-679. CHAMBERLIN, R. V. 1957. The Diplopoda of the Lund University and California Academy of Sciences expeditions. Reports of the Lund University Chile Expedition 1948-1949, No. 30. Lunds Universitets Arsskrift (N. F. 2) 53 (8): 1-44. GOLOVATCH, S. I. 1992. Review of the Neotropical millipede genus Onciurosoma Silvestri, 1932, with the description of three new species from near Manaus, Central Amazonia, Brazil (Diplopoda, Polydesmida, Paradoxosomatidae). Amazoniana 12 (2): 227-237. GOLOVATCH, S. I., HOFFMAN, R.L., MARMOL, A. & Apis, J. 2003. A new, apparently arboricolous species of the millipede genus Mestosoma Silvestri, 1897 from near Iquitos, Peruvian Amazonia (Diplopoda: Polydesmida: Paradoxosomatidae). Amazoniana 17 (3/4): 343-348. HOFFMAN, R. L. 1977. The milliped genus Mestosoma in Costa Rica (Polydesmida: Paradoxosomatidae). Studies on the Neotropical Fauna and Environment 12: 207-215. HOFFMAN, R. L. 1980. Classification of the Diplopoda. Muséum d’histoire naturelle de Geneve (1979), 237 pp. HOFFMAN, R. L. 1999. Checklist of the millipeds of North and Middle America. Virginia Museum of Natural History Special Publication, Number 8, Martinsville, 584 pp. JEEKEL, C. A. W. 1963. Diplopoda of Guiana (1-5). In: GEISKES, D. C. & WAGENAAR HUMMELINCK, P. (eds). Studies on the fauna of Suriname and other Guyanas 4: 1-157. JEEKEL, C. A. W. 1965. A revision of the South American Paradoxosomatidae in the Museo Civico di Storia Naturale di Genova (Diplopoda, Polydesmida). Annali del Museo Civico di Storia Naturale di Genova 75: 99-125. 830 S. I. GOLOVATCH JEEKEL, C. A. W. 1968. On the classification and geographical distribution of the family Paradoxosomatidae (Diplopoda, Polydesmida). Academisch Proefschrift, Rotterdam, 162 pp. JEEKEL, C. A. W. 2002. Paradoxosomatidae from Venezuela, with the description of a new species (Diplopoda, Polydesmida). Myriapod Memoranda 5: 40-51. Kraus, O. 1956. Uber neotropische Strongylosomatidae (Diplopoda). Senckenbergiana bio- logica 37 (5/6): 403-419. Mauris, J.-P. 1998. Diplopoda (pp. 475-484). In: MORRONE, J. J. & COSCARÖN, S. (eds). Biodiversidad de artrépodos argentinos: Una perspectiva biotaxonémica. Sur, La Plata, 599 pp. SCHUBART, O. 1943. Espécies novas das familias Strongylosomidae e Leptodesmidae da ordem Proterospermophora do interior dos estados de Säo Paulo e de Mato-Grosso. Papéis Avulsos do Departamento de Zoologia Sao Paulo 3 (8): 129-163. SCHUBART, O. 1944. Os diplopodos de Pirassununga. Acta Zoologica Lilloana 2: 321-440. SCHUBART, O. 1945a. Diplopodos de Monte Alegre (Municipio de Amparo, Est. de Sao Paulo). Papeis Avulsos do Departamento de Zoologia Sao Paulo 6 (23): 283-320. SCHUBART, O. 1945b. Os Proterospermophora do Distrito Federal (Myriapoda, Diplopoda). Arquivos do Museu Nacional Rio de Janeiro 38: 1-156. SCHUBART, O. 1952. Diplopoda de Pirassununga IV. Adenda à fauna regional. Dusenia 3 (6): 403-420. SCHUBART, O. 1953. Sobre os Diplopoda dos estados do Parana e Santa Catarina. Arquivos do Museu Paranaense Curitiba 10 (3): 77-140. SILVESTRI, F. 1895a. Chilopodi e diplopodi raccolti dal Capitano G. Bove e dal Prof. L. Balzan nell’ America meridionale. Annali del Museo Civico di Storia Naturale di Genova, Serie 2, 14 (34): 764-783. SILVESTRI, F. 1895b. Chilopodi e diplopodi. Viaggio del dottor Alfredo Borelli nella Repubblica Argentina e nel Paraguay. XIV. Bollettino dei Musei di Zoologia ed Anatomia compara- ta della Reale Universita di Torino 10 (203): 1-12. SILVESTRI, F. 1897. Chilopodi e diplopodi. Viaggio del Dott. Alfredo Borelli nel Chaco boliviano e nella Repubblica Argentina. IV. Bollettino dei Musei di Zoologia ed Anatomia com- parata della Reale Universita di Torino 12 (283): 1-11. SILVESTRI, F. 1898. Descrizione di alcuni nuovi diplopodi raccolti nell’ Alto Paraguay dal Cav. Guido Boggiani. Annali del Museo Civico di Storia Naturale di Genova, Serie 2, 18 (38): 671-675. SILVESTRI, F. 1902. Diplopodi. Viaggio del Dr. A. Borelli nel Matto Grosso. VII. Bollettino dei Musei di Zoologia ed Anatomia comparata della Reale Università di Torino 17 (432): 1-25. SILVESTRI, F. 1905. Myriapoda. Zoologische Jahrbiicher, Supplementband 6. Fauna Chilensis 3: 715-702. VERHOEFF, K. W. 1938. Über Diplopoden des Zoologischen Museums in München. Zoologische Jahrbücher, Abteilung für Systematik, Okologie und Geographie der Tiere 71: 1-54. REVUE SUISSE DE ZOOLOGIE 112 (4): 831-848; décembre 2005 On the systematic position of Podopterocus Banks and Dinopsocus Banks, with a revised diagnosis of the genus Sigmatoneura Enderlein (Psocodea: ‘Psocoptera’: Psocidae) K. YOSHIZAWAI, C. LIENHARD? & A. B. IDRIS 1 Systematic Entomology, Graduate School of Agriculture, Hokkaido University, Sapporo, 060-8589 Japan. E-mail: psocid @res.agr.hokudai.ac .jp 2Muséum d’histoire naturelle, c.p. 6434, CH-1211 Genéve 6, Switzerland. 3Center for Insect Systematics, Faculty of Sciences and Technology, Universiti Kebangsaan Malaysia, 43600 Bangi, Malaysia. On the systematic position of Podopterocus Banks and Dinopsocus Banks, with a revised diagnosis of the genus Sigmatoneura Enderlein (Psocodea: ‘Psocoptera’: Psocidae). - The genus Podopterocus Banks has been known only from male specimens and has been characterized by the thickened first antennal flagellomere and the expanded hind tibia. The genus Dinopsocus Banks has been characterized by the thickened first antennal flagellomere. Detailed morphological examinations and brief molecular- based identification (only for one species) suggest that both taxa are congeneric. Based on characters of female terminalia and fore wing venation, both taxa are furthermore considered to be closely related to the genus Sigmatoneura Enderlein. The discovery of two new species and observations on Sigmatoneura spp. reveal the limited significance of the diagnostic characters of Podopterocus and Dinopsocus. Therefore we con- sider both Podopterocus and Dinopsocus as junior synonyms of Sigma- toneura. Monophyly of the genus Sigmatoneura, including Podopterocus, Dinopsocus, and the subgenus Longifolia Li, is well supported by the unique fore wing venation of females and by sexually dimorphic fore wing venation and coloration. Two new species, Sigmatoneura kakisayap sp. n. and Sigmatoneura lemahsayap sp. n., which would be classified under Podopterocus or Dinopsocus by the previous generic definitions, are des- cribed. Sigmatoneura longicornis comb. n., the type species of Podo- pterocus, is redescribed and transferred to Sigmatoneura and the female of this species is described for the first time. Dinopsocus atratus, the type species of Dinopsocus, is synonymized with S. longicornis. Sigmatoneura semicolorata comb. n. is redescribed and transferred from Dinopsocus to Sigmatoneura. Keywords: Psocodea - Psocidae - Sigmatoneura - Podopterocus - Dinopsocus - systematics - Malaysia - Indonesia - Brunei - Singapore. Manuscript accepted 01.09.2005 832 K. YOSHIZAWA ET AL. INTRODUCTION Podopterocus Banks, 1920 and Dinopsocus Banks, 1920 are enigmatic taxa of the family Psocidae. The former genus has been known only from one species, Podopterocus longicornis Banks, 1920, and only from male specimens (see New, 1975) characterized by highly specialized, broadened hind tibiae (Figs 1D, 3C) and a thickened first antennal flagellomere (Fig. 1D). Podopterocus was once treated as a subgenus of Eremopsocus McLachlan, 1866 by Roesler (1944) but is now regarded as a separate genus (Mockford, 1975) and is assigned to the tribe Cerastipsocini of the subfamily Psocinae (Lienhard & Smithers, 2002). The genus Dinopsocus on the other hand contains two species, which are also characterized by a thickened first antennal flagellomere. As Podopterocus, Dinopsocus was once treated as a subgenus of Eremopsocus but is now regarded as a separate genus and assigned to the tribe Cerastipsocini (Lienhard & Smithers, 2002). However, a detailed examination of phylogenetically relevant characters of these genera has not yet been carried out. Therefore their exact taxonomic status and systematic position remained unclear. In addition, New (1978) tentatively suggested that Podopterocus and Dinopsocus may represent the same taxon, closely related to the genus Sigmatoneura Enderlein, 1908, a member of the tribe Metylophorini (Lienhard & Smithers, 2002). Recently we obtained some specimens that can be identified as Podopterocus and/or Dinopsocus collected in Brunei, Indonesia (Sumatra), Singapore and Malaysia (peninsula and Sabah). By comparing these specimens and after having examined the type material of D. atratus, the type species of the genus Dinopsocus, we concluded that these two genera are actually congeneric and show extreme sexual dimorphism. Furthermore, extensive examination of specimens of Sigmatoneura revealed that the diagnostic characters for Podopterocus and Dinopsocus are not clear-cut but rather continuous or homoplastic. In the following, we synonymize Dinopsocus and Podopterocus with Sigmatoneura and discuss their systematic position based on a morphological examination of this material. We also describe two new species that would be assigned to Podopterocus and/or Dinopsocus in the previous sense, based on specimens from Malaysia and Sumatra. Methods of morphological examination follow Yoshizawa (2002). Specimens stored in 70% or 99.5% ethanol were used (except for dry preserved Dinopsocus types). Depositories of specimens are abbreviated as follow: MCZ - Museum of Comparative Zoology, Harvard University, USA (type specimens of Dinopsocus); MHNG - Muséum d’histoire naturelle, Genève, Switzerland; SEHU - Hokkaido University Insect Collection, Sapporo, Japan; UKM - Center for Insect Systematics, Universiti Kebangsaan Malaysia, Bangi, Malaysia. SYSTEMATICS Sigmatoneura Enderlein Sigmatoneura Enderlein, 1908: 761. Type species: Cerastipsocus subcostalis Enderlein, 1903. Podopterocus Banks, 1920: 308, syn. n.; Mockford, 1975: 251. Type species: Podopterocus longicornis Banks, 1920. Eremopsocus (Podopterocus): Roesler, 1944: 147. Dinopsocus Banks, 1920: 307, syn. n.; Mockford, 1975: 251. Type species: Dinopsocus atratus Banks, 1920. Eremopsocus (Dinopsocus): Roesler, 1944: 147. PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 833 Revised diagnosis. Large psocids, fore wing length about 6 mm in male and about 8 mm in female. Antenna very long, first flagellomere normal or sometimes thi- ckened in both sexes (Fig. 1AB vs Fig. ICDE). Male fore wing hyaline or sometimes with pale pigmentation or blackish brown markings, female fore wing blackish brown (Fig. 2); male fore wing venation normal, female fore wing R4+5 strongly sinuate; Rs and M fused for a short distance, meeting at a point, or connected by a cross vein of variable length in both sexes. Male hind tibia usually weakly flattened and somewhat enlarged, rarely paddle-shaped, on each side with a flat wing-like expansion (Fig. 3); female hind tibia of normal cylindrical shape (Fig. 1BE). Sigmatoneura kakisayap Yoshizawa & Lienhard, sp. n. Figs 1AB, 2AB, 3A, 4,5 Podopterocus sp. Johnson, Yoshizawa & Smith, 2004: 1774. Podopterocus sp. KY240. GenBank (online database for gene sequences): accession number for 18S rDNA of holotype male is AY630557. Podopterocus sp. KY329. GenBank (online database for gene sequences): accession number for 18S rDNA of paratype female is DQ116946. Holotype. Male. MALAYSIA (peninsula), Gunung Berembun, Cameron Highlands, Pahang, 14. vii. 2003, leg. H. Kojima et al. (canopy fogging) (UKM). Paratypes. MALAYSIA (peninsula): 2 males, same data as for holotype (SEHU and UKM); 1 male, from the type locality, 15. vii. 2003, leg. H. Kojima et al. (canopy fogging) (UKM); 1 female, Gunung Jasar, Cameron Highlands, Pahang, 14. iii. 2003, leg. K. Yoshizawa (beating dead branches) (SEHU); 1 female, from the type locality, 15. vii. 2003, leg. N. Takahashi (sweeping branches) (UKM). MALAYSIA (Sabah - West Coast Residency): 1 male (heavily damaged, lacking antennae and hind legs), Mt. Kinabalu, 1500 m, Liwagu Trail Section 2, 30. iv. 1987, leg D. Burckhardt & I. Löbl (MHNG); 1 female, Mt. Kinabalu, 1750- 1850 m, Liwagu Trail, 20. ii. 1983, leg. C. Lienhard (MHNG). INDONESIA (Sumatra): 1 male, Pematang Siantar, 1. vii. 1984 - 13. iv. 1985, leg. E. W. Diehl (MHNG). Description. MALE (Fig. 1A). Head black, with narrow pale markings on frons along internal margins of eyes; gena white. Appendices blackish brown; first flagello- mere not thickened. Eye small, IO/D=2.7. Thorax black, except for white membranous regions, propleuron, posterior margin of mesoscutum and metascutellum. Fore wing (Fig. 2A) hyaline, with blackish brown tinge along apical margin between veins R1 and M1, veins and pterostigma black; venation normal, Sc reaching C, Rs-M cross vein long, first section of CuA1 almost perpendicular to the anterior wing margin, CuA1+M connection long. Hind wing hyaline, with brownish tinge posteroproximally, veins black. Legs (Fig. 1A) blackish brown, distal ends of femora and basal ends of tibiae of front and middle legs brown, trochanter and basal end of tibia of hind leg white; hind leg long, hind tibia (Fig. 3A) with symmetrical wing-like expansions along distal 2/3. Abdomen whitish in ground color, dorsally black except for 7th and 8th segments, gray in distal ventral region. Terminalia. Dorsal shelf of clunium absent (Fig. 4AB), dorsal region of clunium with round shallow concavity in the middle. Epiproct (Fig. 4B) without membranous region anteromedially. Paraproct as in Fig. 4AB. Hypandrium (Fig. 4C) covered with wrinkles, anterolaterally fused with clunium, anteromedially with semicircular mem- branous region, posteriorly with roughly trapezoidal projection and with longitudinal crest medially. Phallosome (Fig. 4D): parameres very weakly sclerotized; aedeagus gradually narrowing to truncated posterior end, posterior margin slightly rounded and 834 K. YOSHIZAWA ET AL. D FIG. 1 Habitus of Sigmatoneura spp. (C dried specimen, others in alcohol). A: S. kakisayap sp. n., male paratype; B: S. kakisayap sp. n., female paratype; C: S. semicolorata, male holotype; D: S. lon- gicornis, male from Brunei; E: S. longicornis, female from Singapore. with tiny denticules laterally; phallobase gradually tapering anteriorly, with very short projection (anteriorly). Length (in mm): body 4.3-4.5; fore wing 5.8-6.1; hind wing 3.6-3.8. FEMALE. Almost as in male, except as follows. Head (Fig. 1B): white mar- kings along eyes broad, expanded from antennal sockets to vertex. First and second PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 835 FIG. 2 Fore wings of Sigmatoneura spp. A: S. kakisayap sp. n., male holotype; B: S. kakisayap sp. n., female paratype; C: S. semicolorata, male holotype; D: S. lemahsayap sp. n., male holotype; E: S. lemahsayap sp. n., female paratype; F: S. longicornis, male from Brunei; G: S. longicornis, female from Singapore. segments of maxillary palpus white. IO/D=2.9. Thorax: laterally with broad longitu- dinal white band. Mesoscutellum yellowish white. Fore wing (Fig. 2B) blackish brown, with white region along first section of CuAl and along posterior wing margin between veins CuA2 and Anl; veins and pterostigma black, except the following 836 K. YOSHIZAWA ET AL. yellowish white veins: basal section of Rs, basal 2/3 of R2+3, basal 3/4 of R4+5, Rs- M cross vein, M, basal parts of M branches, and CuA; R4+5 strongly sinuate in the middle, Rs-M cross vein long, first section of CuAl directed anterodistally, CuA1+M connection short. Hind wing pale brown, its veins black. Legs (Fig. 1B): hind leg nor- mal, hind tibia white in distal 2/3. Genitalia. Subgenital plate (Fig. 5A): egg guide pointed apically, with transver- sal crest basally; body of subgenital plate wide, connected with egg guide by narrow sclerite, anteromedian part broadly membranous. Gonapophyses (Fig. 5B): ventral valve long; dorsal valve widely membranous, without distal process; external valve short and wide, densely covered with long setae along posterior margin. Length (in mm): body 4.6-5.1; fore wing 7.9-8.4; hind wing 5.2-5.3. Distribution. Malaysia (peninsula and Sabah), Indonesia (Sumatra). Etymology. The specific epithet, a noun in apposition, is a combination of two Malayan words, kaki (= leg) and sayap (= wing). The synonymized generic name Podopterocus is a combination of the Greek words for «leg» and «wing». Remarks. This new species can be distinguished from the other species of the genus Sigmatoneura by the long hind tibia with dorso-ventrally symmetrical wing- like expansions in the male (Fig. 3A, cf. 3BC) and by the morphology of the subgenital plate in the female. Males and females of this species (Fig. 1AB) are significantly different in fore wing coloration and venation, and in hind leg morphology. Therefore male-female conspecifity was also checked by analysing a 957 bp fragment of the 18S rDNA gene sequence (GenBank accession numbers: AY630557 for holotype male and DQ116946 for one female paratype from Gunung Jasar). No substitution has been identified between the gene sequences of the male holotype and the female paratype, which supports conspecifity. Sigmatoneura semicolorata (Banks) comb. n. RSI 2, © Dinopsocus semicoloratus Banks, 1920: 307. Eremopsocus (Dinopsocus) semicoloratus: Smithers, 1967: 98. Specimen examined. Holotype male. PHILIPPINES, Luzon, Makiling, leg. C. F. Baker (MCZ: 10801). Redescription (coloration in dried condition). MALE (Fig. 1C). Head blackish brown. Appendices blackish brown; first flagellomere thickened. Eye small, IO/D=2.4. Thorax blackish brown. Fore wing (Fig. 2C) hyaline, basal 1/3 and apical part blackish brown, anterior part of apical marking extending proximally to apical part of Rs and CuA1+M connection, veins and pterostigma black; venation normal, Sc reaching C, Rs and M fused for a short distance, first section of CuAl directed anterodistally, CuAl+M connection short. Hind wing hyaline, basal 1/3 brown, veins black. Legs (Fig. 1C) blackish brown; hind tibia almost normal, very slightly flattened apically. Abdomen blackish brown, with whitish longitudinal band laterally. Terminalia. Dorsal shelf of clunium absent (Fig. 6AB). Epiproct (Fig. 6B) with narrow membranous region anteromedially, and with slight transversal swelling medially. Paraproct as in Fig. 6AB. Hypandrium (Fig. 6C) with smooth surface pos- teriorly, lateral region with some weak wrinkles, anterolaterally fused with clunium, PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 837 Fic. 3 Male hind tibiae of Sigmatoneura spp. A: S. kakisayap sp. n., male holotype; B: S. lemahsayap sp. n., male holotype; C: S. longicornis, male from Brunei. anteromedially with triangular membranous region, posteriorly with roughly trapezoi- dal projection. Phallosome (Fig. 6D) relatively long, opened basally; parameres rather well sclerotized; aedeagus gradually narrowing to truncated posterior end, posterior margin slightly concave and with tiny denticules posterolaterally; phaliobase tapering anteriorly, without projection. Length (in mm): body 3.8; fore wing 5.9; hind wing 4.1. Distribution. Philippines (Luzon). Remarks. S. semicolorata can be distinguished from the other species of the genus Sigmatoneura by the unique male fore wing markings. S. kakisayap sp. n., des- cribed above, has also hyaline fore wings with some dark brown markings, but it differs from S. semicolorata by the absence of a thickened first flagellomere and by the greatly expanded hind tibia. The presence of a thickened first flagellomere in S. semi- colorata indicates some relationships with the two species described below, but colo- ration of fore wing in male is remarkably different. The female of S. semicolorata is so far unknown. Sigmatoneura lemahsayap Yoshizawa & Lienhard, sp. n. Figs 2DE, 3B, 7, 8A (?) Dinopsocus atratus Banks sensu New, 1978: 45 (see remarks below). Not Dinopsocus atratus Banks, 1920: 307 (see S. longicornis, below). Holotype. Male. INDONESIA (N-Sumatra), «Holzweg 2», 10 km NE Prapat, 1050 m, 98°57 E, 2°44’ N, 9. i. 1988, leg. E. W. Diehl (MHNG). 838 K. YOSHIZAWA ET AL. Fic. 4 Male terminalia (holotype) of Sigmatoneura kakisayap sp.n. A: terminalia, lateral view; B: epi- proct and left paraproct (trichobothrial field not figured), dorsal view; C: hypandrium, ventral view; D: phallosome, ventral view. Paratypes. INDONESIA (N-Sumatra): 1 female, same locality as for holotype, 27. x1. 1989, leg. E. W. Diehl (MHNG); 1 female, Huta Padang, 99°14’ E, 2°45’ N, 400 m, 2. ii. 1991, leg. H. Malicky (MHNG). Description. MALE. (Male holotype in bad condition, coloration of body, except for wings, almost indistinguishable: i.e., just uniformly pale brown without any distinct marking). First flagellomere thickened. Eye small, IO/D=2.6. Fore wing (Fig. 2D) pale brown, veins black; venation normal, Sc ending in cell c, Rs-M cross vein short, first section of CuAl directed anterodistally, CuA1+M connection short. Hind wing pale brown, basal 1/3 darker, veins brown. Hind tibia (Fig. 3B) with very weakly developed asymmetrical expansions along distal half, ventral expansion stronger (confirmed for both hind legs). PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 839 FIG. 5 Female genitalia (paratype) of Sigmatoneura kakisayap sp.n. A: subgenital plate, showing struc- ture (left) and color pattern (right); B: gonapophyses. Terminalia. Dorsal shelf of clunium absent (Fig. 7A), dorsal region of clunium with round shallow concavity in the middle. Epiproct (Fig. 7A) with membranous region anteromedially. Paraproct as in Fig. 7A. Hypandrium (Fig. 7B) with smooth sur- face, only laterally with few wrinkles, anterolaterally fused with clunium, antero- medially with broad membranous region, posteriorly with semicircular projection. Phallosome (Fig. 7C): parameres very weakly sclerotized; aedeagus gradually narrowing to truncated posterior end, posterior margin slightly rounded and with tiny denticules; phallobase gradually tapering anteriorly, with short projection (anteriorly). Length (in mm): body 4.6; fore wing 6.2; hind wing 4.3. FEMALE. Head brown; gena white. Appendices brown; first and second seg- ments of maxillary palpus paler. First flagellomere thickened. Eye black, IO/D=3.0. Thorax brown, laterally with broad longitudinal white band; meso- and metascutellum paler. Fore wing (Fig. 2E) brown, with white region along first section of CuAl and along posterior wing margin between veins CuA2 and Anl; veins and pterostigma black, except the following yellowish white veins: basal section of Rs, basal 2/3 of R2+3, basal 3/4 of R4+5, Rs-M cross vein, M, basal parts of M branches, and CuA; R4+5 strongly sinuate in the middle, Rs-M cross vein long, first section of CuAl directed anterodistally, CuA1+M connection short. Hind wing pale brown, with darker portion anteroproximally; veins black. Legs brown; trochanters and base of femora paler; hind leg of normal cylindrical shape. Genitalia. Subgenital plate (Fig. 8A): egg guide with a pair of very shallow trenches anterolaterally, truncated apically, apical margin folded upwards and some apical setae arising from dorsal surface of egg guide; body of subgenital plate wide, connected with egg guide by narrow sclerite, anteromedian part broadly membranous. Gonapophyses as in S. kakisayap sp. n. (see description above). 840 K. YOSHIZAWA ET AL. Fic. 6 Male terminalia (holotype) of Sigmatoneura semicolorata. A: terminalia, lateral view; B: epi- proct and left paraproct (trichobothrial field not figured), dorsal view; C: hypandrium, ventral view; D: phallosome, ventral view. Length (in mm): body 4.2-5.0; fore wing 7.2-7.6; hind wing 5.2-5.3. Distribution. Indonesia (Sumatra). Etymology. The specific epithet, a noun in apposition, is a combination of two Indonesian words, lemah (= weak) and sayap (= wing). The synonymized genus Podopterocus was characterized by the wing-like hind tibia of males, but in this species the hind tibia is only weakly expanded in males. Remarks. S. lemahsayap sp. n. is most similar to S. longicornis, redescribed below, because of the thickened first flagellomere in both sexes and the widely pig- mented fore wing in males. However, these species can be clearly distinguished by the shape of the male hind tibia. In having a less expanded hind tibia, S. Jemahsayap is similar to most Sigmatoneura species, but it differs from them by its thickened first flagellomere. The female from Kuala Lumpur described by New (1978) as Dinopsocus atra- tus shows an egg guide which is strikingly similar to that of S. lemahsayap (cf. Fig. 8A PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 841 Fic. 7 Male terminalia (holotype) of Sigmatoneura lemahsayap sp. n. A: epiproct and left paraproct; B: hypandrium, ventral view; C: phallosome, ventral view. and New, 1978: fig. 6). Therefore we consider it as very probable that New’s specimen does not belong to S. longicornis (= D. atratus, see below) but to S. lemahsayap. The specimen from Kuala Lumpur should be re-examined to check this hypothesis. Sigmatoneura longicornis (Banks) comb. n. Figs 1DE, 2FG, 3C, 8B, 9 Podopterocus longicornis Banks, 1920: 308. Eremopsocus (Podopterocus) longicornis: Roesler, 1944: 147; New, 1975: 245 (redescription of male, see remarks below). 842 K. YOSHIZAWA ET AL. Dinopsocus atratus Banks, 1920: 307, syn. n. (?) Not Dinopsocus atratus Banks sensu New, 1978: 45 (see remarks on S. lemahsayap, above). Eremopsocus (Dinopsocus) atratus: Roesler, 1944: 147. Specimens examined. Lectotype female of Dinopsocus atratus (present designation), [MALAYSIA (Sabah):] Borneo, Sandakan, leg. C. F. Baker (MCZ: 10800); paralectotype female of D. atratus (present designation), PHILIPPINES [Luzon], Mt. Banahao, leg. C. F. Baker (MCZ: 10800); non-type female of D. atratus mentioned in the original description, SINGA- PORE, leg. C. F. Baker (MCZ). BRUNEI: 1 male (reared from nymph) and 4 nymphs, Bru-88/30 (Belait District), Sungai Liang, Arboretum of Forest Service, primary forest (mixed dipterocarp forest), alt. about 50m, 21. xi. 1988, leg. C. Lienhard (MHNG); 1 male, Bru-88/42, same locality and collector, alt. 20-50 m, 25. xi. 1988 (SEHU). MALAYSIA (Sabah): 1 female, Sepilok, 24 km W of Sandakan, v-vili, 1983, leg. S. Nagai (MHNG). SINGAPORE: 1 female, Bukit Timah Nature Reserve, 16. xi. 1981, leg. D. H. Murphy (MHNG); 1 female, same locality, no date, leg. D. H. Murphy (MHNG). Diagnosis. The male of this species is easily distinguishable from any other psocid by the unique structure of its hind tibia, which is distinctly paddle-shaped due to a conspicuous, flattened, wing-like expansion on each side; these expansions are dorso-ventrally asymmetrical, the ventral expansion is broadest at basal 1/3 and the dorsal expansion broadest at distal 1/3 (Fig. 3C). The female of this species is very similar to that of S. lemahsayap sp. n., described above, but can be distinguished by the shape of egg guide (Fig. 8B). Length of male (Bru-88/42, in mm): body 4.5; fore wing 5.8; hind wing 4.1. Redescription of male terminalia. Clunial dorsal shelf absent (Fig. 9AB). Epiproct (Fig. 9B) with membranous region anteromedially. Paraproct as in Fig. 9AB. Hypandrium (Fig. 9C) with smooth surface, only mediolaterally with few wrinkles; anterolaterally fused with clunium; anteromedially with broad membranous region; posteriorly with semicircular projection. Phallosome (Fig. 9D): parameres very weakly sclerotized; aedeagus narrowing to truncated posterior end, posterior margin slightly concave; phallobase gradually tapering anteriorly, with very short projection (an- teriorly). Description of female. Very similar to the previous species, S. lemahsayap sp. n., but smaller. Apical margin of egg guide of subgenital plate truncated in S. le- mahsayap but arched in S. longicornis (Fig. 8AB). Length (in mm): body 4.5-4.8; fore wing 6.6-6.7; hind wing 4.6-4.8. Distribution. Singapore (Banks, 1920; New, 1975; present study), Malaysia (Sabah [Banks, 1920 and present study]; peninsula [Banks, 1938]), Brunei (present study), Philippines (Banks, 1920). NOTE: Soehardjan (1958) lists Dinopsocus atratus for Sarawak, probably because of an erroneous interpretation of Banks’ original state- ment («Borneo: Sandakan»), and Endang er al. (2002) erroneously list it for Indonesia. Remarks. S. longicornis, the type species of Podopterocus, has been described from a male collected in Singapore (sex not mentioned in the original description, but the holotype of Podopterocus longicornis must be a male because of having a broadly expanded hind tibia). Banks’ description and figures (Banks, 1920: pl. 1, fig. 4, fore wing and hind tibia) closely correspond to our specimens from Brunei, and we are convinced that they belong to the same species. S. longicornis has briefly been redes- cribed by New (1975) on the basis of three males from Singapore. The only significant difference between New’s description and our observations on the Brunei males PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 843 FIG. 8 Female subgenital plates. A: Sigmatoneura lemahsayap sp. n. (paratype); B: Sigmatoneura lon- gicornis (specimen from Singapore). concerns the «broad rugose apex» of the phallosome mentioned by New, which contrasts with the slightly concave, smooth apical margin of the phallosome in the Brunei material. The differences concerning the shape of the expanded hind tibia and of the phallobase between our Figs 3C and 9D and New’s figures 8 and 11, are pro- bably due to deformations by slide-mounting (phallosome) and to some teratological phenomena or optical distortion (hind leg). In our opinion, it is not very likely that 844 K. YOSHIZAWA ET AL. Fic. 9 Male terminalia of Sigmatoneura longicornis (specimen from Brunei). A: terminalia, lateral view; B: epiproct and left paraproct (trichobothrial field not figured), dorsal view; C: hypan- drium, ventral view; D: phallosome, ventral view. New’s specimens belong to an additional species closely related to S. longicornis. However, this possibility can only be excluded with certainty after a re-examination of New’s material. Concerning shape of hind tibia, our Brunei males correspond more closely to Banks’ than to New’s figure. In our opinion this justifies the assumption that these specimens are conspecific with the male holotype of S. longicornis. Male and female specimens of the present material assigned to S. longicornis are superficially highly divergent and have not been collected simultaneously. There- PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 845 fore the proposed male-female combination remains somewhat tentative. However, three females examined above were collected in Singapore, type locality of S. longi- cornis. Their morphological features do not contradict the present observations concer- ning sexual dimorphism in the genus Sigmatoneura (see Discussion, below). The only other Sigmatoneura species with a thickened first flagellomere, from which female ge- nital morphology has been described, is S. lemahsayap sp. n., which is distinguished by the shape of its egg guide (Fig. 8AB). DISCUSSION In the present study we examined some psocid species that have been or that can be assigned to the genera Podopterocus and/or Dinopsocus which are here placed in the synonymy of Sigmatoneura. Although a greatly expanded hind tibia has been recognized as the most prominent diagnostic character of Podopterocus, our present examination revealed that in some species the hind tibia is sexually dimorphic in shape. Since Podopterocus has been diagnosed by such a dimorphic character, the genus has been known only from males. The genus Dinopsocus on the other hand has been characterized by the thickened first flagellomere, but this character is also present in males and females of Podopterocus longicornis, the type species of Podopterocus (Fig. 1DE). Furthermore, the lectotype of Dinopsocus atratus (type species of Dinopsocus) is considered to be a female of P. longicornis. As already suggested by New (1978), it is now evident that females of Podopterocus have been assigned to the genus Dino- psocus because of this extreme sexual dimorphism. Male-female combination of these highly sexually dimorphic psocids was confirmed genetically only for Sigmatoneura kakisayap sp. n., described and discussed above, the female of which is not of «Dinopsocus type» (i.e., lacking a thickened first flagellomere). However, the result of the molecular male-female match clearly indicates that the expanded hind tibia is a secondary sexual character only developed in males, while corresponding females have a normal cylindrical hind tibia. Similar sexual dimorphism is also known in some psocids of the family Caeciliusidae. For example, in the flavidus group of the genus Valenzuela Navas (see Mockford, 1993; Lienhard, 1998) and in the genus Phymo- caecilius Li (see Li Fasheng, 2002) a more or less conspicuous swelling of the fore ti- bia (and sometimes middle tibia) has been observed in males, never in females. Based on these observations, we concluded that the species described under Podopterocus and Dinopsocus are actually congeners. The above mentioned molecular-based match of a «Podopterocus type» male with a «Sigmatoneura type» female definitively confirmed close relationships between these taxa, as already suggested by New (1978). In the following we discuss the arguments in favor of synonymizing Podopterocus, Dinopsocus and Sigmatoneura instead of assigning subgeneric status to Podo- pterocus/Dinopsocus Within the genus Sigmatoneura s. 1. Mosaic distribution of the diagnostic characters of Podopterocus and Dino- psocus is evident. S. longicornis, S. lemahsayap sp. n. and S. semicolorata belong to the «Dinopsocus type» in having a thickened first flagellomere in both sexes. In contrast, S. longicornis and S. kakisayap are clearly of the «Podopterocus type» in having a greatly expanded paddle-shaped hind tibia in males. S. lemahsayap has a slightly but distinctly expanded hind tibia and therefore is close to «Podopterocus 846 K. YOSHIZAWA ET AL. type». Checking our material of Sigmatoneura s. str. (i. e., species without thickened first flagellomere and without significant expansion of male hind tibia: e. g., S. kolbei), we observed a general tendency for very slight expansion and flattening of the hind tibia in males. The apomorphic presence of a long Rs-M cross vein in the fore wing was observed in all possible Podopterocus and/or Dinopsocus species (except for S. semi- colorata). However, an Rs-M cross vein was also observed in some Sigmatoneura s. str. species, showing a certain variability from almost fused at one point to as long as in S. kakisayap. All known species of the subgenus Sigmatoneura (Longifolia) Li, 2002 also have a long Rs-M cross vein (Li Fasheng, 2002). Therefore, this character state also cannot be used to diagnose Podopterocus and/or Dinopsocus. The thickened first flagellomere is shared by three of four species examined above (S. longicornis, S. lemahsayap and S. semicolorata), but a secondary reversal of the character state in S. kakisayap is also possible (see also discussion below). New (1978) mentioned that the subcosta of Dinopsocus is long and approaching R, whereas that of Sigmatoneura is short and approaching the costa, and he tentatively considered this character as diagnostic for these genera. However, in all specimens examined here, the subcosta is short and approaching the costa (Fig. 2). No other apomorphic charac- ters clearly characterizing Podopterocus and/or Dinopsocus could be found. Judging from these observations, we concluded that the differences between Sigmatoneura s. str. and Podopterocus/Dinopsocus are continuous or highly homoplastic and that it is impossible to clearly define the latter genera. Thus, synonymizing these genera with Sigmatoneura is the most reasonable and practical solution. Monophyly of Sigmatoneura, including Podopterocus, Dinopsocus and the subgenus Longifolia, is well supported by the following unique autapomorphies: female fore wing blackish brown in ground color, with veins R2+3, R4+5 and first section of CuAl white, and with vein R4+5 strongly sinuate; sexual dimorphism in fore wing coloration and venation. Molecular data (18S rDNA: Johnson et al., 2004) also show that Sigmatoneura, including Podopterocus/Dinopsocus (at least S. kakisayap), form a monophyletic group. Among the species examined above, S. longicornis, S. lemahsayap and S. semi- colorata share an apomorphic character state, the thickened first flagellomere, whereas S. kakisayap has normal antennae in both sexes. In contrast, S. longicornis and S. kaki- sayap share an apomorphic condition, the well expanded hind tibia, whereas in S. lemahsayap and S. semicolorata the male hind tibia is only slightly expanded. As already mentioned above, there are conflicts in the distribution of apomorphic character states among these species. Well expanded paddle-shaped hind tibia in males might have evolved independently in S. /ongicornis and S. kakisayap, because the ex- pansion in S. kakisayap is symmetrical, whereas that in S. longicornis and S. lemah- sayap is clearly asymmetrical, although weakly developed in the latter species. The subgenus Longifolia was recognized in the genus Sigmatoneura by Li Fasheng (2002). This subgenus shares the characteristic sexual dimorphism in the fore wing coloration and venation with the subgenus Sigmatoneura. Longifolia is characte- rized by the elongate egg guide of the female subgenital plate (autapomorphic for the subgenus) and thus monophyly of the subgenus can be confirmed. However, no PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 847 apomorphy supporting the monophyly of the subgenus Sigmatoneura is known. Although we do not propose here synonymy of Sigmatoneura and Longifolia, we would like to point out the fact that Li’s treatment probably established a paraphyletic taxon (the subgenus Sigmatoneura). Podopterocus and Dinopsocus have previously been assigned to the tribe Cerastipsocini, whereas Sigmatoneura has been classified under the tribe Metylophorini sensu Mockford (1993) (Smithers, 1980; Lienhard & Smithers, 2002). Placement of Sigmatoneura into Metylophorini is based on the shape of the gonapo- physes, i. e., absence of the distal process of the dorsai valve, which is considered to be an autapomorphy for the tribe. Females of possible Podopterocus and Dinopsocus examined here also have this character state (Fig. 5B); therefore placement of Sigmatoneura, including Podopterocus and Dinopsocus, into Metylophorini is jus- tified. Li Fasheng (2002) established the subfamily Sigmatoneurinae for the genus Sigmatoneura, and assigned the tribes Cerastipsocini and Metylophorini (sensu Mockford, 1993) to a different subfamily (Cerastipsocinae). As mentioned above, the genus Sigmatoneura shares an apomorphic character with the other genera of Metylophorini, and Li’s subdivision of the family Psocidae has to be rejected. ACKNOWLEDGEMENTS We thank P. Perkins and S. Cover for the loan of the type specimens of Dinopsocus spp. KY thanks H. Kojima, S. Nomura, N. Takahashi, M.Y. Ruslan, Y.F. Ng, M. Fauzi and J. Saiful Firdaus for assistance in the field. KY’s collecting trip to Malaysia was supported by the Japan Society for the Promotion of Science (JSPS) grant 14255016 to O. Yata. This study was partly supported by the JSPS grant 15770052 to KY. CL thanks B. Hauser for organizing joint collecting trips to Sabah (1983) and Brunei (1988), H. Malicky, S. Nagai, D. H. Murphy and the late E. W. Diehl for the generous gift of psocids collected by them, and P. Schwendinger for technical assistance and for critical reading of the manuscript. REFERENCES BANKS, N. 1920. New Neuropteroid insects. Bulletin of the Museum of Comparative Zoology, Harvard College 64(3): 299-362, 7 pls. BANKS, N. 1938. Further Neuropteroid insects from Malaya. Journal of the Federated Malay States Museum 18: 220-235, 2 pls. ENDANG SRI KENTJONOWATI, THORNTON, I. W. B. & NEw, T. R. 2002. The Psocidae (Insecta: Psocoptera) of Java and the eastern islands of Indonesia. Invertebrate Systematics 16(1): 107-176. 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Genus Eremopsocus McLachlan: distinction from Cerastipsocus Kolbe and review of species (Psocoptera: Psocidae). Psyche 82(2): 244-258. MOCKFORD, E. L. 1993. North American Psocoptera (Insecta). Flora and Fauna Handbook 10: XVIII+455 pp. Sandhill Crane Press, Gainesville, Florida. New, T. R. 1975. Psocidae (Psocoptera) from Malaysia and Singapore. Oriental Insects 9: 243-259. New, T. R. 1978. Notes on some Oriental genera of Cerastipsocinae (Psocoptera: Psocidae). Oriental Insects 12: 43-48. ROESLER, R. 1944. Die Gattungen der Copeognathen. Stettiner Entomologische Zeitung 105: 117-166. SMITHERS, C. N. 1967. A catalogue of the Psocoptera of the world. Australian Zoologist 14(1): 1-145. SMITHERS, C. N. 1980. The position of Sigmatoneura Enderlein (Psocoptera: Psocidae). Australian Entomological Magazine 6(5): 93. SOEHARDJAN, M. 1958. First contribution to a study of Copeognatha (Corrodentia) of the Indonesian Archipelago. /dea 11(1): 25-33. YOSHIZAWA, K. 2002. Phylogeny and higher classification of suborder Psocomorpha (Insecta: Psocodea: ‘Psocoptera’). Zoological Journal of the Linnean Society 136: 371-400. REVUE SUISSE DE ZOOLOGIE 112 (4): 849-857; décembre 2005 Review of the species of Platypalpus Macquart from Guangxi, China (Diptera, Hybotidae, Tachydromiinae) Ding YANG! 2 & Bernhard MERZ? ! Department of Entomology, China Agricultural University, Beijing 100094, China. E-mail: dingyang @cau.edu.cn 2Key Lab of Insect Evolution & Environmental Changes, Capital Normal University, Beijing 100037, China. 3 Departement d’entomologie, Muséum d’histoire naturelle, C. P. 6434, CH-1211 Genève 6, Switzerland. E-mail: bernhard.merz@mhn.ville-ge.ch Review of the species of Platypalpus Macquart from Guangxi, China (Diptera, Hybotidae, Tachydromiinae). - The species of the genus Platy - palpus Macquart from Guangxi (Southern China) are reviewed. The following 3 species are described, illustrated and discussed: P. alamaculatus sp. n., P. maoershanensis sp. n., P. xanthodes sp. n. A key to the species of Platypalpus from Guangxi is presented. Keywords: Hybotidae - Tachydromiinae - Platypalpus - Guangxi - new species. INTRODUCTION Platypalpus Macquart is one of the largest genera of the subfamily Tachy- dromiinae (Hybotidae). It can be easily recognized by the following characters: eyes narrowly separated on face, mid leg raptorial, and anal cell present (Chväla, 1975; Grootaert & Chväla, 1992). The genus has a worldwide distribution and its center of diversity seems to be the Western Palaearctic region with some 250 described species (Chvala & Kovalev, 1989; Grootaert & Chväla, 1992). In contrast, only some 31 species were described from the Oriental region up to 1975 (Smith, 1975). Since then, 25 species from China have been added by Yang, D. & Yang, C. (1989, 1990, 1992, 1997), Yang, C. & Yang, D. (1992), Saigusa & Yang (2002), Yang et al. (2002). Therefore, 33 species of Platypalpus have been reported from China. The major references dealing with Platypalpus are as follows: Brunetti, 1920; Frey, 1943; Smith, 1965; Chväla, 1975, 1989; Grootaert, 1984; Grootaert & Chväla, 1992. The genus Platypalpus from Guangxi (Southern China) is poorly studied with only Platypalpus guangxiensis Yang & Yang, 1992 known from this province. In the present paper, three species are described as new to science, based on the specimens collected by Ms Shuwen An and Mr Xingyue Liu by sweep net and light trap in 2003. A key to all four species from Guangxi is presented. Manuscript accepted 01.02.2005 850 D. YANG & B. MERZ MATERIAL AND METHODS The specimens on which this study is based on are deposited in the following collections: CAU = Insect collection of China Agricultural University, Beijing, China MHNG = Muséum d’histoire naturelle, Genéve, Switzerland In order to allow an easy comparison of descriptions of various authors, the ter- minology of morphological terms follows generally Grootaert & Chväla (1992), except for crossveins which are written in capitals. The following abbreviations for bristles are used: acr-acrostichial, ad-antero- dorsal, av-anteroventral, dc-dorsocentral, h-humeral, oc-ocellar, npl-notopleural, pd- posterodorsal, psa-postalar, pv-posteroventral, sa-supraalar, sc-scutellar, vt-vertical. KEY TO THE SPECIES OF PLATYPALPUS FROM GUANGXI 1 Thorax mainly yellow; wing without dark spots; mid femur entirely yellow . . 2 - Thorax black; wing with brown spot between Costa and R-M (Fig. 1); midifemuriblackwith yellowitip 2... sere. as eee alamaculatus sp. n. 2 Mesonotum entirely yellow. 2.2... ONCE EME 1 RE 3 - Mesonotum yellow with rather large black mid-longitudinal spot CRI A RI IRONIA EIA maoershanensis sp. n. 5 Scutellum yellow: apical projection of right genital lamella with wide ODIUSETADEX (FLO) RS RE oe eer eae xanthodes sp. n. - Scutellum black; apical projection of right genital lamella with acute APERTA ITA NIN guangxiensis Yang, C. & Yang, D., 1992 SPECIES TREATMENT 1. Platypalpus alamaculatus sp. n. Figs 1-5 MATERIAL Holotype ¢, CHINA: Guangxi: Maoershan National Nature Reserve, 1100-1600 m, 29.VI.2003, Xingyue Liu (CAU). Paratypes: 56 dg, 79 9, same data as holotype (CAU); 56 d, 1099, Guangxi: Maoershan National Nature Reserve, Hongjunting, 1600 m, light trap, 28.VI.2003, Xingyue Liu (CAU); 35 8, 39 2, Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, light trap, 30.VI.2003, Xingyue Liu (MHNG); 46 d, 19, Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, light trap, 30.VI 2003, Shuwen An (CAU). ETYMOLOGY The specific name refers to the maculate wing. DIAGNOSIS Mainly black species: head with 2 vt; wing with a brown spot between C and R-M (Fig. 1); legs yellow, but mid and hind coxae black; mid femur black with yellow tip; mid femur with row of pv bristles; mid tibia with long acute apical spur bearing 1 curved apical hair (Fig. 2). DESCRIPTION Male. Body length 2.6-2.8 mm, wing length 2.9-3.1 mm. PLATYPALPUS FROM GUANGXI 851 Fics 1-5 Platypalpus alamaculatus sp. n, male. 1, wing; 2, apex of mid tibia; 3, right genital lamella; 4, genitalia, dorsal view; 5, left genital lamella. Head, including occiput and clypeus black, gray microtrichose. Eyes narrowly separated on face; frons wider than face. Jowls in lateral view very narrow, almost invisible. Hairs on head pale, bristles black. Ocellar tubercle with 1 oc and 2 posterior hairs; 2 vt (curved inward vt short). Antenna black; pedicel with a circle of apical hairs; first flagellomere short conical, about 1.3 times longer than wide, with some short pale apical hairs; arista 4.5-4.7 times as long as first flagellomere, black, short pubescent. Proboscis blackish with blackish bristles; palpus brown to blackish with 3 long pale bristles (apical bristle long). Thorax black, grey microtrichose. Hairs on thorax pale, bristles blackish; 1 short weak h curved medially, 2 npl (anterior npl short), acr in about 4 irregular rows, 1 short psa, long de and presc absent, scutellum with 2 pairs of sc, basal sc one third as long as apical sc. Legs yellow; fore coxa yellow, mid coxa and posterior half of hind coxa black; mid trochanter black; fore and hind femora yellow, mid femur black with yellow tip; tarsomeres 3-5 of all legs gradually becoming darker towards tip. Hairs on legs blackish, bristles black. Coxae with pale bristles and hairs. Fore femur 1.3 times and mid femur 2.0 times as wide as hind femur; fore femur with one row of short av and one row of long pv (4 apical av the longest); mid femur with two rows of very short black v spine-like bristles (but ventral bristles on basal quarter longer), and with one row of rather long brownish yellow pv; fore tibia with 3 ad, apically with 1 ad; mid tibia (Fig. 2) with a row of short weak black ventral bristles and long acute apical spur 852 D. YANG & B. MERZ (distinctly longer than thickness of mid tibia) bearing 1 curved apical hair. Wing (Fig. 1) hyaline with a brown spot from tip of cell c running to R-M; veins dark brown, R4,5 and M slightly convergent apically; crossveins almost contiguous. Squama yellow with brownish margin, with pale hairs. Halter yellow with brownish yellow base. Abdomen subshining black, very thinly grey microtrichose. Hairs on abdomen pale. Male genitalia (Figs 3-5): Left genital lamella rather narrow in dorsal view, its apical projection finger-like; right genital lamella rather wide in dorsal view, its apical projection rather small and subtriangular; both cerci equally long, right cercus slightly narrower. Female. Body length 2.3-2.5 mm, wing length 2.8-3.0 mm. Similar to male, but av and pv on fore femur shorter except 3 bristle-like apical av. DISTRIBUTION China (Guangxi). REMARKS The new species belongs to the P. pallidiventris-cursitans species-group (Chvala, 1989). It can be easily separated from other known species from China by the colouration of legs and wing pattern. 2. Platypalpus guangxiensis Yang, C. & Yang, D. Platypalpus guangxiensis Yang, C. & Yang, D., 1992: 46. Type locality: Guangxi (Tianlin). DIAGNOSIS Thorax yellow but scutellum black. Abdomen yellow, but tergites medially and apically brownish. Apical projection of right genital lamella spine-like, left cercus spine-like and curved inwards apically. DISTRIBUTION China (Guangxi, Sichuan, Hubei). 3. Platypalpus maoershanensis sp. n. Figs 6-9 MATERIAL Holotype ¢ , CHINA: Guangxi: Maoershan National Nature Reserve, 2100 m, light trap, 5.V11.2003, Shuwen An (CAU). Paratype: 19, Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, light trap, 30.V1.2003, Shuwen An (CAU). ETYMOLOGY The species is named after the type locality. DIAGNOSIS Mainly yellow species with 2 vt. Mesonotum subshiny with one rather large, black mid-longitudinal spot; h, de and row of pv on mid femur absent. Legs yellow with dark brown tarsomere 5. Antenna yellow; first flagellomere short oval. PLATYPALPUS FROM GUANGXI 853 Fics 6-9 Platypalpus maoershanensis sp. n, male. 6, apex of mid tibia; 7, right genital lamella; 8, geni- talia, dorsal view (long apical bristles on left genital lamella omitted, cf. Fig. 9); 9, left genital lamella. DESCRIPTION Male. Body length 2.7 mm, wing length 3.3 mm. Head black, grey microtrichose. Eyes narrowly separated on face; face slightly narrower than frons. Hairs and bristles on head pale. Ocellar tubercle with 2 oc and 2 posterior hairs; 2 vt. Antenna pale yellow; scape without dorsal hairs; pedicel with a circle of pale apical hairs; first flagellomere short oval, 1.2 times longer than wide, with short pale apical hairs; arista 4.2 times as long as first flagellomere, black, short pu- bescent. Proboscis yellow with blackish tip, with sparse black hairs; palpus yellow with 4 pale bristles (of which apical bristle is rather long). 854 D. YANG & B. MERZ Thorax yellow, thinly pale grey microtrichose; mesonotum subshiny with one rather large mid-longitudinal black spot; scutellum, metanotum and lateropleuron (= anatergite & katatergite) (except lateral portion) black. Bristles and hairs on thorax pale; h absent, 2 npl (anterior npl short, hair-like), 1 long psa, short, sparse acr nearly 6-seriate, dc absent, 1 presc (shorter and weaker than psa) in front of scutellum, scutel- lum with 2 pairs of sc (basal pair 1/3 as long as apical pair). Legs yellow with dark brown tarsomere 5. Bristles and hairs on legs pale; fore coxa with 4 dorsal apical bris- tles, mid coxa with 4 dorsal apical bristles, hind coxa with 3 dorsal apical bristles. Fore femur weakly and mid femur distinctly thickened, fore femur 1.1 times and mid femur 1.7 times as wide as hind femur; fore femur apically with 1 dorsal bristle, 1 av and 1 pv; mid femur with two rows of spine-like ventral bristles (posterior row slightly longer than anterior row, but basal ventral bristle rather long and mostly brownish yel- low), without row of pv; mid tibia with a row of short blackish ventral bristles and fin- ger-like apical spur which is longer than thickness of tibia. Wing hyaline; veins brown- ish yellow, R4,5 and M weakly convergent apically. Squama yellow with pale hairs. Halter yellow. Abdomen yellow, thinly pale grey microtrichose; tergites 1-5 and hypopygium dark brown. Hairs on abdomen pale. Male genitalia (Figs 7-9): Left genital lamella rather elongate in dorsal view, its apical projection wide and obtuse, apically with long bristles; right genital lamella rather wide in dorsal view, its apical projection very short and with wide apex; left cercus with acute apex, right cercus with wide rounded apex. Female. Body length 2.8 mm, wing length 3.1 mm. Similar to male. DISTRIBUTION China (Guangxi). REMARKS This new species cannot be placed in any of the species group defined by Chväla (1989). It is similar to Platypalpus hamulatus Yang & Yang, 1989, from Tibet, but the mid-longitudinal spot on mesonotum is uniformly wide. In P. hamulatus, the mid-longitudinal spot on mesonotum is widened towards the scutellum (Yang, D. & Mans 1989): 4. Platypalpus xanthodes sp. n. Figs 10-13 MATERIAL Holotype d, CHINA: Guangxi: Maoershan National Nature Reserve, 1100-1600 m, 29.VI.2003, Xingyue Liu (CAU). Paratypes: 1d 19, same data as holotype (CAU); 39 2, Guangxi: Maoershan National Nature Reserve, Hongjunting, 1600 m, light trap, 28.VI.2003, Xingyue Liu (CAU); 14,399, Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, light trap, 30.VI.2003, Xingyue Liu (CAU); 18,3 2 2, Guangxi: Maoershan National Nature Reserve, 350 m, light trap, 3.VII.2003, Shuwen An (MNHG). ETYMOLOGY The specific name refers to the yellow thorax and abdomen. It is an invariable epithet. PLATYPALPUS FROM GUANGXI 855 Fiss 10-13 Platypalpus xanthodes sp. n, male. 10, apex of mid tibia; 11, apical projection of right genital lamella; 12, genitalia, dorsal view (postgonite [?] drawn as contour behind the cerci); 13, left genital lamella. DIAGNOSIS A mainly subshining yellow species with only the head capsule black; 2 vt; first flagellomere about as long as wide, arista much longer; 4 irregular rows of acr; mid femur without pv; mid tibia with an obtuse apical spur about as long as diameter of tibia; male cerci very asymmetrical; male left genital lamella with long bristles. DESCRIPTION Male. Body length 2.7-3.0 mm, wing length 2.9-3.1 mm. Head black, grey microtrichose, but frons and face orange-brown micro- trichose. Eyes narrowly separated on face; face narrower than frons. Bristles and hairs on head pale. Ocellar tubercle with 1 oc and 2 very short posterior hairs. Jowls almost invisible in lateral view; 2 vt. Antenna yellow; scape without dorsal hairs; pedicel with a circle of mostly pale apical hairs; first flagellomere short oval, 1.1 times as long as 856 D. YANG & B. MERZ wide, with some hairs; arista very long, 3.5-3.7 times as long as first flagellomere, brown, short pubescent. Proboscis yellow with brown hairs; clypeus yellow, mictro- trichose; palpus yellow with 3 long pale bristles, the apical one much longer than the others. Thorax yellow, thinly grey microtrichose; mesonotum subshiny, without darker stripe; katepisternum with indistinct shining yellow spot medially; meron with a small longitudinal, black spot. Hairs on thorax pale, bristles brownish yellow; 1 short hair- like h curved inward, 2 npl (anterior npl short and hair-like), acr in 4-6 irregular rows, dc not longer than acr, 1 long psa, 1 weak presc in front of scutellum, scutellum with 2 pairs of sc (basal pair very short, 1/3 of apical pair). Legs yellow, but tarsomere 5 slightly brownish apically. Hairs on legs pale. Fore femur weakly and mid femur dis- tinctly thickened, fore femur 1.2 times and mid femur 1.7 times as wide as hind femur; mid femur with two rows of short black ventral bristles (posterior row longer than anterior row, but ventral bristles on basal quarter long and brownish yellow), without row of pv; mid tibia (Fig. 10) with a row of blackish ventral bristles, finger-like apical spur about as long as diameter of tibia. Wing hyaline; veins brownish yellow; crossveins contiguous; Ry, 5 and M parallel apically. Squama yellow with pale hairs. Halter yellow. Abdomen yellow, pale grey microtrichose; hypopygium mostly yellow. Hairs on abdomen pale. Male genitalia (Figs. 11-13): Left genital lamella rather narrow, its apical projection wide and obtuse apically, and with long apical bristles; right genital lamella slightly wide in dorsal view, its apical projection short with wide and obtuse apex; left cercus long with spine-like apex, right cercus shorter with obtuse apex. Female. Body length 2.7-3.1 mm, wing length 2.8-3.2 mm. Similar to male. DISTRIBUTION China (Guangxi). REMARKS The new species belongs to the P. longicornis species-group of Chväla (1989). Itis very similar to Platypalpus guangxiensis Yang & Yang from Guangxi, Sichuan and Hubei, but can be separated from the latter by the yellow scutellum and right surstylus with wide obtuse apex. In guangxiensis, the scutellum is black, and the right surstylus has an acute apex (Yang, C. & Yang, D., 1992). CONCLUSIONS Guangxi belongs to South China which is considered to be part of the Oriental region with a subtropical and tropical climate. Although Platypalpus is basically a genus of temperate regions, it is noteworthy that 4 species are known to occur in this province. However, they were all found at higher altitude (above 1000 m, only one record of P. xanthodes from 350 m). Among them the three new species are endemic to Guangxi so far. The only already known species has a much wider distribution with records extending to Sichuan and Hubei in Central China region. Taking into account the bad collecting effort in Guangxi it is well possible that more species will eventually be found in this province. PLATYPALPUS FROM GUANGXI 857 ACKNOWLEDGEMENTS Our sincere thanks are due to Ms Shuwen An and Mr Xingyue Liu (China Agricultural University, Beijing) for collecting specimens and for their help during the study. The research was funded by the National Natural Science Foundation of China (No. 30070100_No. 30225009). REFERENCES BRUNETTI, E. 1920. Diptera Brachycera Vol. 1. The Fauna of British India, including Ceylon & Burma. London, 401 pp. CHVALA, M. 1975. The Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. I. Fauna Entomologica Scandinavica 3: 1-336. CHVALA, M. 1989. Monograph of northern and central European species of Platypalpus (Diptera, Hybotidae), with data on the occurrence in Czechoslovakia. Acta Universitatis Carolinae - Biologica 32: 209-376. CHVALA, M. & KOVALEV, V. G. 1989. Family Hybotidae (pp. 174-227). In: Soös, A. & Papp, L. (eds). Catalogue of Palaearctic Diptera. Vol. 6. Akademiai Kiad6, Budapest, 435 pp. Frey, R. 1943. Übersicht der paläarktischen Arten der Gattung Platypalpus Macq. (=Coryneta Meig.). (Dipt., Empididae). Notulae Entomologicae 23: 1-19. GROOTAERT, P. 1984. Two new species of Platypalpus Macquart (Diptera, Empidoidea, Hybo- tidae) from Papua New Guinea. Bulletin et Annales de la Société Royale d’Entomologie de Belgique 120: 283-290. GROOTAERT, P. & CHVALA, M. 1992. Monograph of the genus Platypalpus (Diptera: Empidoidea, Hybotidae) of the Mediterranean region and the Canary Islands. Acta Universitatis Carolinae - Biologica 36: 3-226. SAIGUSA, T. & YANG, D. 2002. Empididae (Diptera) from Funiu Mountains, Henan, China (1). Studia Dipterologica 9 (2): 519-543. SMITH, K. G. V. 1965. Diptera from Nepal: Empididae. Bulletin of the British Museum (Natural History), Entomology 17 (2): 61-112. SMITH, K. G. V. 1975. Family Empididae (pp. 185-211). Jn: DELFINADO, M. D. & HARDY, D. E. (eds). A Catalog of the Diptera of the Oriental Region. Vol. 2. The University Press of Hawaii, Honolulu, 459 pp. YANG, C. & YANG, D. 1992. Three new species of Empididae from Guangxi — Diptera: Brachy- cera. Journal of the Guangxi Academy of Sciences 8 (1): 44-48. YANG, D. & YANG, C. 1989. The dance flies of Xizang (II) (Diptera: Empididae). Acta Agri - culturae Universitatis Pekinensis 15 (4): 415-424. YANG, D. & YANG, C. 1990. Eleven new species of the subfamily Tachydrominae from Yunnan (Diptera: Empididae). Zoological Research 11 (1): 63-72. YANG, D. & YANG, C. 1992. Diptera: Empididae (pp. 1089-1097). In: CHEN, S. (ed.). Insects of the Hengduan Mountains Region. Vol. 2. Science Press, Beijing, 1547 pp. YANG, D. & YANG, C. 1997. Diptera: Empididae (pp. 1469-1476). In: YANG, X. (ed.). Insects of the Three Gorge Reservoir Area of Yangtze River. Chongqing Publishing House, Chongqing, 1847 pp. YANG, D., AN, S. & GAo, C. 2002. New species of Empididae from Henan (Diptera) (pp. 30-38). In: SHEN, X. & SHI, Z. (eds). The Fauna and Taxonomy of Insects in Henan. Vol. 5. China Agricultural Scientech Press, Beijing, 453 pp. EIERN EL AU N TANT — i si sta m “Rs ture N amine Le [ven 0 è so Ia lié A AL Key ta rape its hms te wae $ 3 un u eg mos È. er | ah ell = ug utp 0 ¥ 42} piedi i . 2 u REVUE SUISSE DE ZOOLOGIE 112 (4): 859-867; décembre 2005 A contribution to the knowledge of the Chyromyidae (Diptera) of Italy with description of a new species of Aphaniosoma Becker Martin J. EBEJER Hendre, Colwinston, Cowbridge, South Glamorgan CF71 7NL, United Kingdom. E-mail: martinebejer@evemail.net A contribution to the knowledge of the Chyromyidae (Diptera) of Italy with description of a new species of Aphaniosoma Becker. - The family Chryomyidae is reported from Sicily (eleven species) and from Sardinia (nine species). Additional records of species for mainland Italy are also given. A new species of Aphaniosoma is described from France, Sicily and Sardinia. A total of eleven species are new to Italy: Chyromya miladae, Gymnochiromyia fallax, G. flavella, G. inermis, G. mihalyii, Aphaniosoma collini, A. claridgei, A. grisescens, A. micromacro, A. proximum, and A. bi- falcatum sp.n. A list is given of all the seventeen species so far recorded for Italy and its larger islands, together with their data and their distribution in the West Palaearctic. Keywords: Diptera - Chyromyidae - Aphaniosoma bifalcatum sp. n. - Italy - Sardinia - Sicily - France. INTRODUCTION The family Chyromyidae of Italy has not been well studied and there are no published records from Sardinia or from Sicily. In the Checklist of the Diptera of Italy (Canzoneri et al., 1995) only three species are listed, namely: Chyromya flava (Linnaeus, 1758), C. oppidana (Scopoli, 1763) and Aphaniosoma nigrohirtum (Hendel, 1933). All occur in the north of Italy. The two species of Chyromya, according to the Checklist, also occur in the south. The Catalogue of Palaearctic Diptera (Soés, 1984) gives only A. nigrohirtum as specifically occurring in Italy. Thus, both works overlooked or did not accept the record, based on a single specimen deposited in the Museum of Natural History in Vienna, of A. approximatum Becker, which was given in Die Fliegen der Paläarktischen Region (Czerny, 1927). However, this specimen was re-examined and it belonged not to approximatum, but to a new species, egregium Ebejer (1998b); in the same paper, A. melitensis Ebejer and A. propinquans Collin were also recorded from Italy. Recent studies of this family of Diptera (Ebejer, 1998a, b; Carles-Tolra, 2001) have revealed that there are many more species in the Mediterranean. Thus, the list given in the Catalogue of Palaearctic Diptera (Soös, 1984) is now very out of date. During a brief visit to Sicily in the spring of 1999, eleven species of Chyromyidae were Manuscript accepted 26.11.2004 860 M. J. EBEJER encountered. Dr Merz found nine species in Sardinia in the spring of 2002 and Dr Gatt collected three species in Tuscany in the summer of 2003. They are listed in this article with their data and summarised in Table 1. Eleven of these species are new records for Italy. TABLE 1: The Italian species of Chyromyidae. (The north and south Italian regions are according to the Checklist of the Diptera of Italy.) North Italy South Italy Sardinia Sicily 1. Chyromya flava (Linnaeus) x x 2. Chyromya miladae Andersson x 3. Chyromya oppidana (Scopoli) x x 4. Gymnochiromyia fallax Ebejer x 5. Gymnochiromyia flavella (Zetterstedt) xi 6. Gymnochiromyia inermis (Collin) x x 7. Gymnochiromyia mihalyii So6s x X 8. Aphaniosoma bifalcatum sp. nov. x x 9. Aphaniosoma claridgei Ebejer x x x 10. Aphaniosoma collini Lyneborg x 11. Aphaniosoma egregium Ebejer x x 12. Aphaniosoma grisescens Ebejer Xe 13. Aphaniosoma melitensis Ebejer x x X 14. Aphaniosoma micromacro Carles-Tolrä x 15. Aphaniosoma nigrohirtum Hendel x 16. Aphaniosoma propinquans Collin X x 17. Aphaniosoma proximum Ebejer X x MATERIAL AND METHODS Specimens were collected by hand-net sweeping only. All the material, which is cited in this article, is dry mounted and deposited in the personal collections of the collectors or their Institutions [Dr Deeming — National Museum of Wales, Cardiff, UK (NMWC); Dr P. Gatt — Rabat, Malta; Dr B. Merz — Museum of Natural History, Geneva, Switzerland (MHNG); and in the author’s collection (MJE)]. In the course of this study a number of specimens similar to A. claridgei Ebejer could not be identified with certainty. Detailed examination showed these to belong to a new species and this is described below. For the indentification to species, the reader is referred to the following litera- ture: for the genus Chyromya - Andersson, 1971, 1976; for Gymnochiromyia - Soés, 1979, Ebejer, 1998a; and for Aphaniosoma - Collin, 1949; Ebejer, 1993, 1998b, Carles- Tolra, 2001. The nomenclature of the hypopygium that is used in this article takes into account the terminology given in the Contributions to a Manual of Palaearctic Diptera (Sinclair, 2000). The term gonostylus is retained for the time being, since there are un- certainties of homologies with regard to this structure. It is so identified because it articulates with the posterior end of the hypandrium and in many Aphaniosoma it bears sensory setulae (absent in the species). The term corresponds to terminology used in recent literature on the Chyromyidae (Ebeier, 1998a, b). CHYROMYIDAE OF ITALY 861 RESULTS Chyromya flava (Linnaeus, 1758) Material examined. ITALY: Sictty: 38 6,22 9, Catania, Randazzo, Monte Spagnolo, 1200 m, grassland, edge of woodland of Acer, Castanea, Pinus, 6.vi.1999, J.C. Deeming; 19, same data, but M.J. Ebejer; 18,82 2, same data, but B. Merz; 16 and 19 , Messina, Nebrodi, Cesarò, 1150 m, on Tilia, 8.vi.1999, J.C. Deeming; 19, Messina, Nebrodi, Troina, Elia River, 1000 m, 8.vi.1999, MJ. Ebejer; 88 d, 122 2, Catania, Randazzo, Lago di Gurrida, Quercus, Populus, 870 m, 11.vi.1999, M.J. Ebejer; 36 3,52 9 , same data, but B. Merz. This is one of the commonest and most widespread species in the family. It is also known from North America and North Africa. It has been observed in good num- bers on and beneath leaves of isolated broad-leaved trees, especially Tilia and Populus, growing at the edge of open woodland. New to Sicily. Chyromya miladae Andersson, 1976 Material examined. ITALY: SıcıLy: 19, Catania, Randazzo, Lago di Gurrida, Quercus, Populus, 870 m, 11.vi.1999, M.J. Ebejer. Described from the Czech Republic and later recorded from Slovakia, this species was recently listed as occurring in Britain (Chandler, 1998). This uncommon species is probably associated with broad-leaved woodland. New to Sicily. Chyromya oppidana (Scopoli, 1763) Material examined. ITALY: SıcıLy: 19, Catania, Randazzo, Lago di Gurrida, Quercus, Populus, 870 m, 11.vi.1999, M.J. Ebejer. This species is as frequent as flava and it has the same wide distribution and habits. New to Sicily. Gymnochiromyia fallax Ebejer, 1998 Material examined: ITALY: SARDINIA: 59 2, Nuoro reg., 120 m, 8 km E of Oliena, Hotel su Gologone, 16.vi.2002, B. Merz & M. Eggenberger. This species is typically found in saltmarshes and coastal regions. It was described from Malta, but it is also found in Mallorca. New to Sardinia. Gymnochiromyia flavella (Zetterstedt, 1848) Material examined: ITALY: SARDINIA: 19, Baunei reg., Santa Maria Navarrese, 18.vi.2002, B. Merz & M. Eggenberger. Widespread in Europe and probably all the Palaearctic including North Africa. New to Sardinia. Gymnochiromyia inermis (Collin, 1933) Material examined: ITALY: SARDINIA: 19, Villacidro reg., 600 m, Nuxeddu Mts., Montimannu forest, 12.vi.2002, B. Merz & M. Eggenberger; 12, Nuoro reg., Monte Albo 1120 m, Punta Catirina, 16.vi.2002, B. Merz & M. Eggenberger. Sicity: 26 6,39 9, Messina, Nebrodi, Troina, Lago D’Ancipa, mixed woodland, 8.vi.1999, M.J. Ebejer; 12 same data, B. Merz; 19, Catania, Randazzo, Monte Spagnolo, 1200 m, grassland, Acer, Castanea, Pinus, 6.vi.1999, M.J. Ebejer; 12, same data, but B. Merz; 12, Messina, Bronte, Monte Minardo, Quercus forest, 345 m, 11.vi.1999, M.J. Ebejer; 19, same data, but J.C. Deeming. 862 M. J. EBEJER When Collin (1933) described this species, he gave a number of examples of birds’ nests from where it was reared. Deeming (1998) described the puparium and cephalopharyngeal skeleton. It is a common species and it is the most likely member of the genus to be found at some altitude and in broad-leaved woodland. New to Sardinia and Sicily. Gymnochiromyia mihalyii Soös, 1979 Material examined: ITALY: SARDINIA: 16, 59 ?, Nuoro reg., Monte Albo 1120 m, Punta Catirina, 16.vi.2002, B. Merz & M. Eggenberger; 14 , Dorgali, Rio Flumineddu, Gola di Gorropu, 15.vi.2002, B. Merz & M. Eggenberger. SICILY: 19, Catania, Etna, 3 km NW of Milo, Castanea, Corylus, 1000 m, 5.vi.1999, M.J. Ebejer. Carles-Tolra (1992) recorded this species from Spain and Merz (1997) recorded it from Switzerland. The present author has specimens from Britain, Turkey and Greece. It is probably widespread in several Mediterranean countries. Unlike inermis, it seems to prefer open countryside including dunes and cultivated fields. New to Sardinia and Sicily. Aphaniosoma bifalcatum sp. n. Figs 1-3 Material examined: Holotype: d, ITALY: SARDINIA: Dorgali reg., 350 m, Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger (MHNG). Paratypes: 18,22 2, same data and depository; 1d, same data, but in MJE; 14, Bosa reg., 0 m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger (MHNG). Sicity: 16, 19, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. Ebejer (MJE). FRANCE: 23 4,4 2, Montpellier, Lotte salt- marsh, 23.v111.2000, J.C. Deeming (NMWC). A dark brownish black species of the approximatum Becker group characterized by a pair of long frontal setae anterior to the ocellar triangle and long narrow dark surstyli. Holotype, male. Head: yellow, ocellar triangle black, occiput deep black except for narrow, yellow postocular margin; frons at vertex about half width of head; frons, antenna, gena and mouth parts all yellow; third antennal segment with distinct setulae anteriorly, arista black except for yellow basal segments; gena about half as high as eye; chaetotaxy: a pair of very long frontal setae present on frons in front of ocellar triangle and 8 pairs of pale setulae scattered on frons, 2 long orbitals and 2 shorter ones in front of these, 1 internal and 1 external vertical. Thorax: black with dark grey dusting becoming brownish in prescutellar de- pression and on scutellum; scutellum with pale apical margin; humerus and notopleur- al depression clear yellow; pleura all dark brownish black except for narrow yellow margins to sutures; metanotum black; chaetotaxy: 1 humeral, 1 posthumeral, 2 noto- pleural, the anterior, strongly developed — about twice as long as posterior; 2+5 dorso- centrals, but only the prescutellar strong, 2+4 acrostichals, 3 intra-alar, 1 post-alar and 0 supra-alar, 2 pairs of marginal scutellars; 1 mesopleural with 4 short setulae in vertical row below this, 1 sternopleural at posterior upper corner, and 4 setulae in vertical row at anterior part; stigmatical setula present. Wing: broad, hyaline with anterior cross vein pale yellow and other veins all brownish, becoming darker towards apex; distance between cross veins about 1.2 times length of posterior cross vein; distance between R,,; and R,,5 on costa is half that between R,,; and M,,5. Haltere: CHYROMYIDAE OF ITALY 863 gon bac scl hyp Fics 1-3 Aphaniosoma bifalcatum sp. n., male terminalia. 1, hypopygium lateral aspect; 2, pregenital sternite; 3, hypopygium posterior aspect; abbreviations: bac scl = bacilliform sclerite, ep = epandrium, gon = gonostylus, hyp = hypandrium, surs = surstylus, t6 = sixth tergite; scale bar = 0.1mm. 864 M. J. EBEJER yellow. Legs: all yellow except for 5! tarsal segment of all legs — this being broadly infuscated around margin; hind coxa simple and brownish at base. Abdomen: tergites black with narrow pale posterior margins; epandrium with shiny pale brown spot at base near cerci, which are relatively long, yellow and pale- haired. Hypopygium (Figs 1-3): typical of the approximatum Becker - creperum Collin group, having an internalised and reduced epandrium with prominent dark surstyli visible externally; distiphallus poorly sclerotized and relatively amorphous; bacilliform sclerite and gonostylus prominent, darkly pigmented and heavily sclerotized; gono- stylus articulates with posterior arm of hypandrium very close to where this articulates with inferior margin of epandrium. Female. As in male, but without secondary sexual characters; apex of abdomen with tergite 7 strongly curved on sides to form a broad and short conical cavity with the cerci deep within; apical 2 sternites, in middle third, heavily sclerotized and dark shiny brown pigmented. Variation. The males from France and the one from Bosa in Sardinia have a greyish yellow scutellum and a dusky yellow prescutellar area; the grey mesonotal pattern fades posteriorly except in the middle where it reaches the scutellar margin and, laterally, the yellow area extends forwards along the intra-alar line; the pleura have the sutures more broadly yellow and the hind marginal bands of the tergites are broader, especially laterally. The colour variations are present also in the females from France. A male paratype from France was dissected and is identical to the paratype from Sardinia, which is illustrated in the figure. Similar species. The nominate form resembles the dark species of the claridgei Ebejer group, whereas the paler form resembles grisescens Ebejer and proximum Ebejer. If the surstyli are properly extruded, then bifalcatum can be recognised because these structures are sinuous and the apex is somewhat spatulate. Etymology. The curved surstylus together with the sinuate gonostylus gives an appearance of two sickles (Latin singular: falx) on each side of the postabdomen. Aphaniosoma claridgei Ebejer, 1995 Material examined: ITALY: Toscana: 38 8,72 2, Parco Nazionale della Maremma, Collelungo, Bonifica Canal, 2.ix.2003, P. Gatt; 26 4, 1522, plus 89 © in alcohol, Parco Nazionale della Maremma, Bocca D’Ombrone estuary, saltmarsh, 2.ix.2003, P. Gatt; 38 4,49 9, Parco Nazionale della Maremma, Collelungo, beach and dunes, 2.1x.2003, P. Gatt. SARDINIA: 33 d, 19, Dorgali reg., 350 m, Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger; 38 8,32 2, Bosa reg., 0 m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger; 133 8,32 9, Cuglieri Reg., Sinis Peninsula, 0 m, Capo Mannu, 21.vi.2002, B. Merz & M. Eggenberger, SICILY: 66 3, 39 9, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. Ebejer; 118 4,169 2, same data, but B. Merz; 26 4,59 9, same data, but J.C. Deeming. A typical salt marsh species with a preference for coastal areas, it is widespread in the Mediterranean. It is often the dominant species of the genus in such habitats. There are many similar species in this group. New to Italy: Sardinia and Sicily. Aphaniosoma collini Lyneborg, 1973 Material examined: ITALY: SARDINIA: 26 à , Bosa reg., Bosa Marina, 0 m, 20.v1.2002, B. Merz & M. Eggenberger. CHYROMYIDAE OF ITALY 865 This species was described from Spain. It is common in Tunisia, Mallorca, France and Malta and may be one of the more widespread species in the group with projections on the hind trochanter in the male. New to Sardinia. Aphaniosoma egregium Ebejer, 1998 Material examined: ITALY: SıcıLy: 16, Siracusa, Noto, Vendicari, coastal marshes and dunes, J.C. Deeming (MJE). This is a distinctive, but one of the most uncommon species of Aphaniosoma. Other than the types, very few specimens of this species have been seen. New to Sicily. Aphaniosoma grisescens Ebejer, 1998 Material examined: ITALY: SARDINIA: 26 à , Dorgali reg., 350 m, Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger; 14 , Baunei reg., Santa Maria Navarrese, 18.vi.2002, B. Merz & M. Eggenberger; 23 d , 19, Cuglieri Reg., Sinis Peninsula, 0 m, Capo Mannu, 21.vi.2002, B. Merz & M. Eggenberger; 36 d, Bosa reg., 0 m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger. Described from Tunisia, it is also known from Malta. Not a common species, but it can easily be confused with claridgei and proximum. New to Sardinia. Aphaniosoma melitensis Ebejer, 1993 Material examined: ITALY: ToscaNa: 18, 22 2, Parco Nazionale della Maremma, Collelungo, Bonifica Canal, 2.ix.2003, P. Gatt; 12, Parco Nazionale della Maremma, Bocca D’Ombrone estuary, saltmarsh, 2.ix.2003, P. Gatt. SARDINIA: 26 6, 8£ © , Dorgali reg., 350 m, Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger; 29 9, Baunei reg., Santa Maria Navarrese, 18.vi.2002, B. Merz & M. Eggenberger; 88 4, 19, Bosa reg., 0 m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger; 16, 19, Cuglieri Reg., Sinis Peninsula, 0 m, Capo Mannu, 21.vi.2002, B. Merz & M. Eggenberger. Sicity: 38 d, 39 2, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. Ebejer; 26 8,16% © , same data, but B. Merz; 2d 6, 89 2, same data, but J.C. Deeming. This is another common species known from several European countries where it inhabits flushed meadows, marshes and dunes. It is a polymorphic species with small very dark forms appearing quite different from the large pale specimens. There is also variation as to how rounded are the apical projections on the ventral aspect of the fourth tergite in the male. All intermediates exist within the same population if a large enough sample is examined. New to Sicily and Sardinia. Aphaniosoma micromacro Carles-Tolrä, 2001 Material examined: ITALY: Toscana: 26 6,62 2, Parco Nazionale della Maremma, Collelungo, beach and dunes, 2.ix.2003, P. Gatt. Described from Spain, this species is one of the very few with modified tarsi. It is also known from Malta. New to Italy. Aphaniosoma propinquans Collin, 1949 Material examined: ITALY: Sictty: 244,299, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. Ebejer. Originally described from Britain, this species is now known to have a wide dis- tribution in Europe. It is relatively easy to identify in both sexes and so it should not be overlooked. New to Sicily. 866 M. J. EBEJER Aphaniosoma proximum Ebejer, 1998 Material examined: ITALY: SARDINIA: 19, Bosa reg., 0 m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger; 24 4, Cuglieri Reg., Sinis Peninsula, 0 m, Capo Mannu, 21.vi.2002, B. Merz & M. Eggenberger. SiciLy: 68 4,79 ©, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. Ebejer; 15,42 2, same data, but B. Merz. This species belongs to the approximatum Becker group. It is difficult to iden- tify without dissection. It is probably widespread in the Mediterranean, but overlooked. New to Sardinia and Sicily. ACKNOWLEDGEMENTS I would like to thank my colleagues Dr John Deeming (Cardiff, UK), Dr Paul Gatt (Rabat, Malta) and Dr Bernhard Merz (Geneva, Switzerland) who kindly put at my disposal all their material of Chyromyidae. I thank the authorities of the Parco dei Nebrodi (Sicily) for allowing access and collecting in the National Park (permit number 3294). Dr Gatt collected in the Parco Nazionale della Maremma as a parti- cipant in the MECO Project. Finally, I am grateful to Dr Giorgio Sabella of the University of Catania for sparing his time to discuss with us the best sites to visit and for arranging permission to collect on Etna and at Vendicari. REFERENCES ANDERSSON, H. 1971. The Swedish species of Chyromyidae (Diptera), with lectotype designations. Entomologisk Tidskrift 92: 95-99. ANDERSSON, H. 1976. Chyromya miladae n.sp. from Czechoslovakia (Diptera: Chyromyidae). Entomologica scandinavica 7: 187-189. CANZONERI, S., GORODKOV, K., KRIVOSHEINA, N.P., MUNARI, L., NARTSHUK, E., PAPP, L. & SUss, A. 1995. Diptera Opomyzoidea, Carnoidea, Sphaeroceroidea. /n: MINELLI, A., RUFFO, S. & LA POSTA, S. (eds). Checklist delle specie della fauna Italiana 75, Bologna, Calderini, 27 pp. CARLES-TOLRA, M. 1992. New and interesting records of Diptera Acalyptrata from Spain. Part II: Heleomyzidae, Trixoscelididae, Chyromyidae, Curtonotidae, Camillidae, Diastatidae and Campichoetidae. Graellsia 48: 19-24. CARLES-TOLRA, M. 2001. Eight new chyromyid species from Spain (Diptera, Chyromyidae). Boletin de la Asociacion espanola de Entomologia 25(3-4): 45-62. CHANDLER, P. 1998. Chyromyidae (p. 151). Jn: CHANDLER, P. J. (ed.). Checklists of Insects of the British Isles (New Series), Part 1: Diptera. Handbooks for the Identification of British Insects 12: ili-xx + 1-234. Royal Entomological Society, London. COLLIN, J.E. 1933. Five new species of Diptera. Entomologists monthly Magazine 69: 272-275. COLLIN, J.E. 1949. On Palaearctic Species of the Genus Aphaniosoma. Annals and Magazine of Natural History (12) 2: 127-147. CZERNY, L. 1927. 53c. Chiromyidae (pp. 51-54). In: LINDNER, E. (ed.). Die Fliegen der Paläarktischen Region, Vol. 5. E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, 56 pp. DEEMING, J.C. 1998. A description of the puparium of Gymnochiromyia inermis (Collin, 1933) from England (Diptera: Chyromyidae). Studia Dipterologica 5 (1): 30-32. EBEJER, M.J. 1993. The Chyromyidae of Malta with descriptions of new species of Aphaniosoma (Dipt., Schizophora). Entomologists Monthly Magazine 129: 125-133. EBEJER, M.J. 1998a. A new species of Gymnochiromyia Hendel (Diptera: Chyromyidae) from the Mediterranean, with notes, lectotype designations and a key to the species from the West Palaearctic. Studia Dipterologica 5 (1): 19-29. CHYROMYIDAE OF ITALY 867 EBEIER, M.J. 1998b. A Review of the Palaearctic Species of Aphaniosoma Becker (Diptera, Chyromyidae), with Descriptions of New Species and a Key for the Identification of Adults. Mitteilungen Museum fiir Naturkunde Berlin, Deutsche Entomologische Zeitschrift 45 (2): 191-230. MERZ, B. 1997. Die Megamerinidae, Strongylophthalmyiidae, Pseudopomyzidae, Chyromyidae und Camillidae der Schweiz (Diptera, Acalyptrata). Mitteilungen der entomologischen Gesellschaft Basel 47(4): 130-138. SINCLAIR, B. 2000. Morphology and terminology of Diptera male terminalia, 1.2. (pp. 53-74). In: Papp, L. & Darvas, B. (eds). Contributions to a Manual of Palaearctic Diptera. Vol. 1. Science Herald, Budapest, 978 pp. Soos, A. 1979. Einige neue Acalyptraten Musciden aus der Paläarktischen Region (Diptera). Acta Zoologica Academiae Scientiarum Hungaricae 35 (3-4): 409-414. Soös, A. 1984. Family Chyromyidae (pp. 56-60). In: Soos, A. & Papp, L. (eds). Catalogue of Palaearctic Diptera. Vol. 10. Akademiai Kiado, Budapest, 402 pp. = > mi tuo vel a AO love: ane 64 neck at Coeliac ine (homies EE in (KE ve cat A a Az pa, PESA Re — is ai vale Md u La À 2 : = a REVUE SUISSE DE ZOOLOGIE 112 (4): 869-941; décembre 2005 Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new species and notes on biogeography and phylogenetic relationships Lionel MONOD! & Wilson R. LOURENCO? 1 Muséum d’histoire naturelle, route de Malagnou 1, case postale 6434, CH-1211 Genève 6, Switzerland. E-mail: liocheles@ gmail.com 2 Département de Systématique et Evolution, USM 0602 — CP 53, Section Arthropodes (Arachnologie), Muséum National d’Histoire Naturelle, 61 rue de Buffon, F-75005 Paris, France. E-mail: arachne@mnhn fr Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new species and notes on biogeography and phylogenetic relationships. - The family Hemiscorpiidae is closely related to the Liochelidae. Within the Hemiscorpiidae, the Oriental species are particularly interesting. Most of them exhibit highly derived characters in comparison to their African rela- tives. Males possess a strongly elongated metasoma and a similarly elon- gated telson bearing a pair of tuberculiform processes at the base of the aculeus. Furthermore, Hemiscorpius lepturus Peters, 1861, which occurs in Iraq and Iran, is known to have an extremely virulent venom with cytotoxic and haemolytic components. It is responsible for severe dermonecrotic scorpionism in southern Iran. This is the only non-buthid scorpion that is potentially lethal. In this paper an overview of the species of Hemiscorpius in Iran is presented with revised diagnoses and descriptions. Two new species from western Iran, H. enischnochela sp. n. and H. acanthocercus sp. n., are described. The genus Habibiella Vachon, 1974 is synonymised with Hemiscorpius Peters, 1861. A thorough analysis of hemisperma- tophores shows close phylogenetic relationships with several genera of the family Liochelidae. A hypothesis on the geological events that probably triggered the present distribution of Hemiscorpius is finally proposed. Keywords: Scorpiones - Hemiscorpiidae - Hemiscorpius - new species - new synonymy - Iran - hemispermatophore - Liochelidae. INTRODUCTION The family Hemiscorpiidae Pocock, 1893 was formerly considered as a sub- family (Hemiscorpiinae) of the Scorpionidae Latreille, 1802 (Fet, 2000). Sissom (1990) criticised the placement of the Hemiscorpiinae in the Scorpionidae, but did not suggest an alternative. Stockwell (1989) first suggested the transfer of Hemiscorpiinae directly from the Scorpionidae to the Ischnuridae, where they retained their subfamily rank. Recently Lourenco (2000) and Prendini (2000) simultaneously elevated the Manuscript accepted 11.08.2005 870 L. MONOD & W. R. LOURENCO Hemiscorpiinae to family level. While Lourengo did not give any justification, Prendini provided a detailed phylogenetic analysis where he demonstrated that Hemiscorpiidae is the sister group of Ischnuridae Simon, 1879, now Liochelidae Fet & Bechly, 2001. Solegad & Fet (2003) confirmed the phylogeny of Scorpionoidea estab- lished by Prendini (2000), but downgraded the Hemiscorpiidae from family to sub- family rank under Liochelidae. Until the discussion is settled, we decided to follow Lourengo’s and Prendini’s view and consider the hemiscorpiids as a family. Two genera were traditionally listed in the family Hemiscorpiidae (see Fet, 2000), Habibiella Vachon, 1974 and Hemiscorpius Peters, 186la. Habibiella is a monotypic genus, the type species, H. gaillardi, was described by Vachon (1974) from a single female collected in eastern Iran. This genus is here placed in the synonymy of Hemiscorpius. The genus Hemiscorpius in the traditional sense includes six species which occur from Somalia to Pakistan, most of them in the Middle East (Fet, 2000). These are: A. arabicus (Pocock, 1899a), H. lepturus Peters, 186la, H. maindroni (Kraepelin, 1900), H. persicus Birula, 1903, H. socotranus Pocock, 1899b and A. tellinii Borelli, 1904. Species of Hemiscorpius and Habibiella gaillardi (now under Hemiscorpius) are morphologically very similar and were distinguished only by their trichobothriotaxy. Habibiella is neobothriotaxic major, with 10-12 trichobothria on the ventral side of the pedipalp patella instead of 3 as in the species previously comprising Hemiscorpius, and 15 trichobothria on the external side of the pedipalp patella instead Of 13: Three species belonging to the family Hemiscorpiidae are known to occur in Iran: A. gaillardi, H. lepturus and H. persicus. H. lepturus is of particular medical interest. This species is the only dangerous and potentially lethal scorpion that does not belong to the family Buthidae. It is responsible for significant scorpionism problems in the southern provinces of Iran. The venom of H. lepturus is highly cytotoxic and haemolytic and can cause serious wounds and skin inflammations whereas other scor- pions have a neurotoxic venom. The deep dermonecrotic ulcers and blisters caused by such a sting are slow and difficult to heal and therefore usually result in very un- pleasant scars. Severe complications such as serious haemolysis, internal haemor- rhages, secondary renal failure and death were also reported (Radmanesh, 1990, 1998). Venoms with such cytotoxic and hemolytic effects are also quite uncommon in spiders and only known in three genera, i. e. Cheiracanthium C.L. Koch, 1839 (Miturgidae), Loxosceles Heineken & Lowe, 1832 (Sicariidae) and Sicarius Walckenaer, 1847 (Sicariidae), (Filmer, 1999). While severe envenomations in humans by Cheira- canthium and Sicarius are not known, the virulent poison of the Brown Recluse Spider, Loxosceles reclusa Gertsch & Mulaik, 1940, and its enzymatic activities have been extensively studied (Anderson, 1998; Butz er al., 1971; Dillaha er al., 1964; Foil et al., 1979; Patek er al., 1994). Among the 13 species of Loxosceles present in the United States (at least 5 of them are associated with necrotic arachnidism), L. reclusa is most commonly responsible for dermonecrotic arachnidism in North America. Brown spi- ders are also of significant clinical concern in South America, with L. laeta (Nicolet, 1849) being responsible for several deaths each year. In Iran cases of scorpion enven- omation with cutaneous and viscerocutaneous reactions are usually attributed to the species H. lepturus. Species of Hemiscorpius are morphologically very close to each HEMISCORPIIDAE FROM IRAN 871 other and difficult to distingish for a non-specialist, therefore we assume that H. lep- turus is probably not the only species responsible for all envenomations in that coun- try. Other Hemiscorpius species probably possess venoms with similar necrotic effects. As stated by Fet (2000), there is no recent revision and no dichotomic key for Hemiscorpius. In the past 40 years several checklists of scorpions from Iran, which also include Hemiscorpius species, have been published (Vachon, 1966; Habibi, 1971; Farzanpay & Pretzmann, 1974; Pérez-Minnocci, 1974; Farzanpay, 1988; Kinzelbach, 1985; Kovarik, 1997). However, neither precise descriptions, nor specific diagnoses were included in these publications. The extensive cladistic analysis of Scorpionoidea by Prendini (2000) gives a precise diagnosis and description for the family, but not for the two species used in that study, i. e. A. lepturus and H. maindroni. Early descrip- tions are very short and not accurate enough, and therefore do not allow species iden- tification. The present paper gives an overview of the hemiscorpiids of Iran, with revised diagnoses and redescriptions of H. gaillardi, H. lepturus and H. persicus. H. enischnochela sp. n. and H. acanthocercus sp. n. are described from the region of Bandar Abbas, Hormozgan Province, south-eastern Iran. This study also allowed to clarify the taxonomic position of the enigmatic genus Habibiella, which is here synonymised with the genus Hemiscorpius. Strong evidence of a close phylogenetic relationship with the liochelids /omachus politus Pocock, 1896, Hadogenes Kraepelin, 1894 and Opisthacanthus (Opisthacanthus) Peters, 1861b is pointed out by the examination of hemispermatophore morphology. Finally a hypothesis on the geolo- gical events that presumably triggered the evolution and present distribution of the family is proposed. MATERIAL AND METHODS Illustrations were produce by using a Wild M5 stereo-microscope with a draw- ing tube. Trichobothrial notations and terminology of metasomal carination follow Vachon (1974), measurements follow Stahnke (1970) and are given in mm. Additional morphological terminology mostly follows Hjelle (1990) and Sissom (1990), termi- nology of carination of pedipalpal chela follows Prendini (2000). Hemispermatophore terminology is modified from the terminology applied by Lamoral (1979) and was used by Monod & Volschenk (2004). The distribution map was generated with ArcView® GIS 3.1, maps and drawings were subsequently edited in Adobe Illustrator® 8.0 and Adobe Photoshop® 6.0. LIST OF ACRONYMS BPBM _ Bernice P. Bishop Museum, Honolulu, Hawaii. CAS Californian Academy of Sciences, San Fransisco, USA. MHNG Muséum d’histoire naturelle, Genéve, Switzerland. MNHN Muséum National d’Histoire Naturelle, Paris, France. NHMW Naturhistorisches Museum Wien, Austria. ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. ZMB Zoologisches Museum, Humboldt Universität, Berlin, Germany. 872 L. MONOD & W. R. LOURENCO SYSTEMATICS HEMISCORPIIDAE Pocock, 1893 Hemiscorpinii Pocock, 1893: 306, 308. Type genus Hemiscorpius Peters, 1861a. Diagnosis (modified from Prendini, 2000). The Hemiscorpiidae can be distin- guished from all other scorpionoid taxa by their trichobothriotaxy: trichobothria ib and it of the pedipalp chela are situated midway on the finger, whereas both are at the base of the fixed finger in all other scorpionoid taxa except Nebinae and ib is situated basally and it distally on the fixed finger in Nebinae, Diplocentridae. Hemiscorpiids also differ from other scorpionoid taxa, except for Heteroscorpionidae and Urodacidae, by their metasomal segments I-IV, which bear a single ventromedian carina equally developed on all segments. Hemiscorpiidae can be distinguished from the Heteroscorpionidae and Urodacidae by the following characters: (1) 3 pairs of lateral ocelli present; (2) median ocular tubercle shallow; (3) movable cheliceral finger with distal external and distal internal teeth approximately equal in size and closely opposed; (4) pedipalp chela fingers with 2 primary rows of denticles often fused at the base; (5) trichobothrium db located on internal surface of fixed finger; (6) trichobothrium Db located basally on dorsal surface of manus; (7) trichobothrium Dr located at proximal end of fixed finger; (8) no accessory trichobothria in v series of chela; (9) telson vesicle of male elongated and laterally compressed; (10) hemispermatophores with a double lamellar hook and a complex median capsular structure. Remarks. Telson vesicles of mature Hemiscorpius leturus males are unusually elongated, laterally compressed, and bear a pair of distal lobes at the base of aculeus (Fig. 1A-B), whereas females possess bulky telsons, which are laterally not compressed and have no distal lobes. Subadult males do not have an extremely elongated metasoma and a modified telson like mature males have, their metasoma and telson are just slight- ly more elongated than in females. These secondary sexual characters appear only in the last developmental instar. In Prendini’s (2000) cladistic interpretation of relationships within the superfamily Scorpionoidea this very unusual feature is considered as a valid character for the whole family Hemiscorpiidae and is mentionned in the diagnosis: tel- sons of H. lepturus and H. maindroni males are described as possessing an extreme elongation and paired distal lobes. Our examination of syntypes of Hemiscorpius main- droni from Mascate also revealed a pronounced sexual dimorphism but not as extreme as in H. lepturus. Indeed H. maindroni males possess telsons (Fig. 1G-H) without any distal lobes or elongation as in A. lepturus. Therefore the species identified as H. main- droni in Prendini’s cladistic revision of the superfamily Scorpionoidea (2000) undoubt- edly belongs to another species. All the other non-Iranian hemiscorpiids examined, i. e. H. socotranus, Hemiscorpius spp. from Somalia, Eritrea and Djibouti possess a simple telson and a less elongated metasoma than in H. maindroni (Fig. 27A-B). The Iranian hemiscorpiids H. acanthocercus sp. n., H. enischnochela sp. n., H. lepturus and pro- bably also A. gaillardi can therefore be placed together in the A. lepturus species complex. However, the Iranian H. persicus does not belong to this group. The mature male holotype studied does not have modified metasoma and telson. Prendini (2000) described the hemispermatophores of hemiscorpiids as possessing a single lamellar hook. In his data matrix (Table 3, p. 6), the character 90 HEMISCORPIIDAE FROM IRAN 873 (hemispermatophore, lamellar hook) is coded 0 (single hook) for both H. lepturus and H. maindroni. Mature males in the material examined have been systematically dis- sected in order to remove and study their hemispermatophores. Several hemisper- matophores in excellent shape of H. lepturus, H. acanthocercus and H. maindroni have been obtained. All of them bear a double lamellar hook, a very unusual feature that was so far only recorded for the liochelid genus Hadogenes. Unfortunately this character cannot be assessed for H. enischnochela sp. n., H. persicus and H. socotranus. The only available mature male of H. enischnochela had been dissected before but the hemispermatophore was not found inside the vial and is probably lost. We were not allowed to dissect the type of H. persicus and no other specimens of this species were available to us. The only available mature male of H. socotranus had poorly preserved paraxial organs from which it was impossible to extract hemispermatophores. Nevertheless the character “double lamellar hook” is used in our diagnosis of the family. In liochelids morphometric proportions of hemispermatophores and shape of the capsular median structure are usually the only features that vary between genera. The single lamellar hook is a constant character within the family, except for Hadogenes, species of which possess hemispermatophores with a double lamellar hook. According to Prendini (2000) Hadogenes and Cheloctonus are the more basal liochelid genera and Hemiscorpiidae is the basal sister-group of Liochelidae. The double lamellar hook can therefore be considered as a plesiomorphy for Hemi- scorpiidae and Liochelidae, and the single lamellar hook is synapomorphic for Liochelidae except Hadogenes. Therefore the probability that the double lamellar hook is present in the whole Hemiscorpiidae is strong. Furthermore a double lamellar hook was assessed for hemiscorpiids from Oman by Dr. Graeme Lowe (pers. comm.) who is currently carrying out a revision of the scorpion fauna of Oman (Lowe, in prep.). This is another argument for including the double lamellar hook into the family diagnosis. Hemiscorpius Peters, 1861 Hemiscorpius Peters, 1861a: 426. Type species by monotypy Hemiscorpius lepturus Peters, 1861a. Habibiella Vachon, 1974: 952, syn. n. Type species Habibiella gaillardi Vachon, 1974. Remarks. Prendini (2000) suggested in his revision of the superfamily Scorpionoidea that Habibiella should be considered as a synonym of Hemiscorpius, because the monophyly of Hemiscorpius was not supported in any of his cladistic analyses. However, he did not formally propose this taxonomic change and listed both Habibiella and Hemiscorpius in the family Hemiscorpiidae. The genus Habibiella is monotypic and only known from a single female. It can be distinguished from Hemiscorpius only on the basis of its trichobotriotaxy, 15 external trichobothria on the pedipal patella instead of 13, and 10-12 ventral trichobothria on the pedipalp patella instead of 3. The study of a mature male from the Natural History Museum of Vienna that belongs to this genus and that is described here under Hemiscorpius enischnochela sp. n. allowed to re-evaluate the taxonomic status of Habibiella. Metasoma and telson of this specimen possess exactly the same morphology encountered in Hemiscorpius lepturus, i. e. extremely elongated metasoma, vesicle strongly elongated and bearing a 874 L. MONOD & W. R. LOURENCO pair of blunt tuberculiform projections at the base of the short and strongly curved aculeus. The African and Arabian hemiscorpiids examined, i. e. H. maindroni and H. socotranus, do not have such features. Species of “Habibiella“ are therefore more closely related to Hemiscorpius lepturus, the type species of Hemiscorpius, than H. lepturus is to the other Hemiscorpius species. Habibiella is therefore considered as a junior synonym of Hemiscorpius. Diagnosis. With Habibiella now in synonymy with Hemiscorpius, the family Hemiscorpiidae becomes monotypic and its diagnosis is also applicable to the genus. Hemiscorpius acanthocercus sp. n. Figs 1C-D, 2-7, 36 Material examined. Holotype: 4 , IRAN (Hormozgan), Abad-Geno, 38 km N Bandar Abbas, 3.1V.1972, K. Bilek, NHMW 4718 (39). Paratypes: 1 4,1 9, no locality specified, H. Löffler, Austrian Iran Expedition 1949-1950, MHNG. 2 ¢ juv., 2 2 juv., no locality speci- fied, H. Löffler, Austrian Iran Expedition 1949-1950, NHMW. 1 à juv., IRAN (Hormozgan), ca. 50 km N Bandar Abbas, 41V.1970, F. Ressl, NHMW 3393. 1 ¢ juv., 1 © juv., IRAN (Hormozgan), ca. 41 km N Bandar Abbas, 7.1V.1972, G. Pretzman & A. Konetschnig, NHMW 3394.1 5 subadult, 2 d juv., 2 2,1 © juv., IRAN (Hormozgan), 115 km E Bandar Abbas, 27.11.1972, K. Bilek, NHMW 4716.1 ¢ juv., 1 2, IRAN (Hormozgan), 22 km N Bandar Abbas, 4.IV.1972, K. Bilek, NHMW 4717.2 2 juv., same data as for holotype, NHMW 4718.1 & juv., 1 2,1 © juv., IRAN (Hormozgan), 40 km N Bandar Abbas, 7.1V.1972, K. Bilek, NHMW 4719. 1 & subadult, IRAN (Hormozgan), 41 km N Bandar Abbas, 20.IV.1974, G. Pretzman, NHMW 4720.1 & subadult, 2 2, IRAN (Hormozgan), 65 km N Bandar Abbas, 30.III.1972, K. Bilek, NHMW 4721.1 ® juv., IRAN (Hormozgan), 22 km N Bandar Abbas, 1974, G. Pretzman, NHMW 21142. 1 3 imm., 1 2, 1 © juv., IRAN (Hormozgan), 38 km N Bandar Abbas, 28.11.1972, K. Bilek, NHMW 21143. 2 2, IRAN (Hormozgan), 65 km N Bandar Abbas, 4.IV.1972, K. Bilek, NHMW 21144. Distribution. Known from the surroundings of Bandar Abbas, Hormozgan Province, eastern Iran (Fig. 36). Etymology. The name acanthocercus is a construct from the Greek words akan- tha [thorn, prickle] and kerkos [tail]. The name is an invariable noun in apposition and refers to the metasomal dorsal carinae of males, which have numerous strong spiniform granules, and to the telson of males which also bears numerous small spiniform gran- ules, especially at the posterior extremity, on the blunt tuberculiform processes at the base of the aculeus. Diagnosis. (1) Carapace longer than wide, shagreened and finely granular, with small smooth patches; lateral margins with small spiniform granules below lateral ocular tubercles; superciliary carinae finely granular; (2) pedipalps stout and bulky, chela fingers slightly shorter than or equal in length to chela manus; (3) internal protu- berance of pedipalpal patella with internodorsal carina bearing 1-2 strong spiniform granules; (4) pedipalp patella orthobothriotaxic, external side with 13 trichobotria (1 est and 2 esb), ventral side with 3 trichobothria; (5) metasoma of males elongated and slender, with dorsal carinae bearing numerous strong spiniform granules, and ventral and ventrolateral carinae of segments IV-V bearing spiniform granules; (7) telson of males strongly elongated, bearing a pair of blunt tuberculiform processes with small spiniform granules at base of aculeus; (8) metasoma of females with dorsal carinae of segments I-IV and ventral and ventrolateral carinae of segment V bearing strong spini- form granules; (9) hemispermatophore with strong double lamellar hook located above distal transverse ridge, very close to it, and pointing anteriad. HEMISCORPIIDAE FROM IRAN Fic. 1 Hemiscorpius spp., males: telsons, ventral face (A, C, E, G) , lateral face (B, D, F, H). A-B, Hemiscorpius lepturus (MNHN-RS 5232); C-D, 875 H. Löffler coll.); E-F, Hemiscorpius enischnochela sp. n. (holotype); G-H, Hemiscorpius main- Hemiscorpius acanthocercus sp. n. (paratype, MHNG, droni, (syntype, MNHN-RS 4328). Scale lines, 1 mm. 876 L. MONOD & W. R. LOURENCO H. acanthocercus sp. n. is closely related to A. lepturus, both of them have stout orthobothriotaxic pedipalps and pronounced sexual dimorphism. However, males of these 2 species can be readily distinguished by (1) carapace granulation, (2) metasoma carination, (3) morphology of telson, (4) pedipalp carination and shape, and (5) hemis- permatophore morphogy. The carapace of H. acanthocercus sp. n. has lateral margins bearing small spiniform granules below the lateral eyes, while in H. lepturus the later- al carapace margins are completely smooth. H. acanthocercus sp. n. has metasomal dorsal carinae bearing numerous strong spiniform granules, while H. lepturus has cari- nae with sparse, much reduced and weaker spiniform granules, especially on the ante- rior segments. The ventral and ventrolateral carinae of metasomal segments IV-V are furnished with spiniform granules in males of H. acanthocercus sp. n., but are almost smooth, with weak spiniform granules (carinae of segment V with more marked spini- form granules in the posterior half) in 7. lepturus. In H. acanthicercus sp.n. the telson of males is less elongated and the aculeus is less curved than in A. lepturus. The later- al surfaces of the telson are fairly smooth, with only small granules, and distal protu- berances bear small spiniform granules in H. acanthocercus, while the lateral surfaces and distal projections are rugose, without distinct granules, in A. lepturus. Furthermore the distal telson projections are bigger in H. lepturus. The pedipalp chela manus in H. acanthocercus sp. n. is slightly thinner than in H. lepturus. H. acanthocercus sp. n. has intero-dorsal carina of the patellal prominence with a single strong spiniform granule (rarely 2), while H. lepturus has carinae with several much reduced, weaker granules. Females of these two species are very similar to each other, but can be distinguished by the superciliary carinae, which are very finely granular in H. acanthocercus sp. n., while smooth, or nearly so, in H. lepturus. Hemispermatophores of H. acanthocercus sp. n. possess a double lamellar hook that is stronger than in A. lepturus. The lamellar hook is located very close to the distal transverse ridge and points anteriad in 7. acan- thocercus sp. n., while it is located more distally and points distad in A. lepturus. Description. MALE (Figs 1C-D, 2A-B, 3A, 4B, 5A-C, E-F, 6A, 7) [description based on the holotype (NHMW 4718)]. Measurements of holotype (in mm) carapace, length 5.2; posterior width 4.9; distance between anterior lateral eyes 2.7, between posterior lateral eyes 3.2, between median eyes 0.2; diameter of median eyes 0.3. Pedipalp, femur length 5.0, width 1.8; patella length 5.0, width 1.7, chela length 9.7; manus length 5.0, width 3.8, depth 2.3; movable finger length 4.6; fixed finger length 3.6. Metasoma, segment I length 5.2, width 1.4; segment V length 7.4, width 1.2, depth 1.4; vesicle length 4.6, width 1.5, depth 1.8, aculeus length 0.9. Total length 60.0. Carapace (Fig. 3A). Colouration mostly uniformly dark orange to brown; me- dian and lateral ocular tubercles black. Carapace dorsoventraly flattened, longer than wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. Carapace shagreened and finely granular, with a few smooth patches; lateral margins with small spiniform granules below lateral ocular tubercles; antero-median furrow narrow, suturiform, anteriorly bifurcated; median longitudinal furrow shallow, conti- nuous from the anterior suture furcation, running through ocular tubercle posteriorly HEMISCORPIIDAE FROM IRAN 877 Fic. 2 Hemiscorpius acanthocercus sp. n. Male paratype (MHNG, H. Löffler coll.): A, dorsal aspect: B, ventral aspect. Female paratype (VHMW 4719): C, dorsal aspect; D, ventral aspect. Scale line, 5 mm. 878 L. MONOD & W. R. LOURENCO into a deep triangular depression; postero-lateral furrow shallow; mesolateral furrow weakly developed, almost indiscernible. Mesosoma. Tergite colouration dark orange to brown anteriorly, becoming lighter posteriorly (dark yellow). Tergites I-III with a shallow median depression divided anteriorly by a weak ridge; tergites IV-VII with median carina reduced to a very weak ridge surrounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae absent on tergites I-VI, but present on the posterior 2/3 of tergite VII as ridges with weak spiniform granules. Surface of tergites shagreened and finely gra- nular; pre-tergites smooth and minutely pitted. Sternite colouration dark yellow. Sternites II-VI smooth and minutely pitted, without granulation or carinae; sternite III with a pair of large, very finely and densely granular depressions, covered by the pectines; sternites IV-VI with a pair of shallow median furrows. Sternite VII with similar surface texture as on preceding sternites; pair of lateral carinae developed as ridges; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII longer than wide. Metasoma (Fig. 4B) very elongated and slender, shagreened, with very few small scattered granules. Colour dark yellow to orange. Segments I-IV with longitu- dinal dorsomedian furrow, dorsal carinae with strong spiniform granules. Segments I-IV with lateral carinae developed as weakly granular ridges, lateral carinae on seg- ments III-IV less granular than those on segments I-II. Segments I-II with ventrolateral carinae as ridges with few weak granules, almost smooth; segments III-IV with ventrolateral carinae as ridges of reduced spiniform granules. Segment I: ventral carina indistinct, reduced to a weak ridge; segments II-III with ventral carina developed as a distinct ridge, almost smooth; segment IV with ventral carina as a ridge, anterior 1/3 smooth and posterior 2/3 with weak spiniform granules. Segment V: longitudinal dorsal furrow present in anterior half, dorsal carinae with strong spiniform granules; lateral carina indistinct, visible as a row of small granules in anterior 2/3 only; ventro- lateral and ventromedian carinae with spiniform granules. Telson (Figs 1C-D, 4B). Vesicle yellow to orange; aculeus darker, tan, due to stronger sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt tuberculiform projection on each side at base of aculeus; ventrolateral furrows absent; ventromedian ridge absent; lateral surfaces smooth, with few weak granules; posterior projections with small spiniform granules; dorsal surface with numerous small spini- form granules and a median shallow depression. Few macrosetae basally, becoming more numerous near base of aculeus. Aculeus short and stout, strongly curved, becoming markedly narrower approximately midway. Chelicerae. Colour dark yellow; teeth of fingers darker orange. Teeth arrange- ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth bifid; movable finger with one subdistal tooth and one basal tooth in external series; distal external tooth smaller than distal internal tooth; cheliceral teeth without secon- dary serrations. Pedipalp stout and bulky. Pedipalp coxa and femur (Fig. 5A, E-F). Inter- noventral margin of coxa with strong spiniform granules. Dorsal surface of femur predominantly dark yellow-orange, carinae darker, orange. Femur stout (length less than 2.5 times its width), pentacarinate, with 4 distinct carinae; internodorsal carina HEMISCORPIIDAE FROM IRAN 879 geet UE ER Fersen ! ore LA Tin GAL Murtas TESINE FIG. 3 Hemiscorpius acanthocercus sp. n., carapace, dorsal aspect: A, male holotype (NHMW 4718); B, female paratype (NHMW 4719). Scale line, 1 mm. ns Lap x Tay De SS TUTO 880 L. MONOD & W. R. LOURENCO developed as a strongly and densely granular ridge, granules spiniform; externodorsal carina forming a ridge with spiniform granules; internoventral carina developed as a granular ridge with spiniform granules; externoventral carina developed as a ridge with strong spiniform granules; ventromedian carina reduced to few granules and confined to base of femur. Dorsal surface shagreened, finely and densely granular, distal end smooth, without granules; internal surface shagreened, with a few strong, scattered, spiniform granules; external surface shagreened; ventral surface shagreened and weakly granular, distal end without granules. A total of 3 trichobothria present on femur; d located externobasally on dorsal surface; i located dorsobasally on internal surface; e located dorsobasally on external surface. Pedipalp patella (Fig. 5A, E-F) stout and bulky (length less than 2.5 times its width). Colour predominantly dark yellow to orange; carinae slightly darker. 7 carinae present, 6 of them distinct; interno- dorsal carina developed as a granular ridge; externodorsal carina developed as a ridge of indistinct granules; internoventral carina developed as a granular ridge; externo- ventral carina developed as a ridge of weak granules; externomedian carina developed as a weakly granular ridge. Dorsal surface shagreened, sparsely and weakly granular; internal and external surfaces shagreened; ventral surface shagreened, sparsely and weakly granular, distal end without granules. Internal protuberance pronounced, bifid (internodorsal and internoventral tubercles separated), internodorsal carina with 1 spiniform granule. A total of 19 trichobothria present on patella, orthobothriotaxy; d, located basally, external to internodorsal carina; d, located midway on patella; d; absent; i in distal half of patella, located dorsally on internal surface. External (e) trichobothrial groups discernible and orthobothriotaxic: eb group composed of 5 tri- chobothria, esb group of 2 trichobothria, em group of 2 trichobothria, et group of 3 tri- chobothria; est very close to ef group. Ventral (v) group orthobothriotaxic, 3 tri- chobothria. Pedipalp chela manus (Fig. SA-C). Colour predominantly orange to red- brown; carinae darker red. Chela stout (length less than 2.5 times its width) with 5 dis- tinct carinae; internodorsal carina continuous, with strong spiniform granules basally and weak granules distally; subdigital carina vestigial, visible basally as a smooth ridge with weak granules; externodorsal carina distinct, visible as a ridge of indistinct fused granules, almost smooth; digital carina a smooth ridge with few weak granules basally; dorsal secondary carina (dorsomedian) a smooth ridge with few strong granules basally and few weak granules distally; internoventral carina continuous, visible as a weak ridge with few weak spiniform granules distally; externoventral carina continuous, forming a granular ridge, becoming almost smooth distally, running parallel to longi- tudinal axis of chela, its distal edge disconnected from external movable finger condyle and directed between external and internal condyles of movable finger; ventromedian carina vestigial, reduced to few coarse granules basally; internal (internomedian) carina vestigial, reduced to few weak spiniform granules distally; external (externo- median) carina indistinct, visible as raised rows of small weak granules. Dorsal surface forming an irregular and subreticulate network of weakly granular ridges surrounding shagreened patches without granules; internal surface shagreened, sparsely granular, with a denser patch of spiniform granules dorsodistally; external surface shagreened, sparsely and weakly granular; ventral surface with few weak granules surrounding shagreened patches, distal end smooth and pitted. A total of 15 trichobothria present on chela; Db trichobothria located externobasally on dorsal surface; Eb group (3 tri- HEMISCORPIIDAE FROM IRAN 881 Fic. 4 Hemiscorpius acanthocercus sp. n., metasoma, lateral aspect: A, female paratype (VHMW 4717), B, male paratype (MHNG, H. Löffler coll.). Scale line, 2 mm. 882 L. MONOD & W. R. LOURENCO o Ya vw = + Q = (SI 5.) Se — n v Q (Hy de) u ul u il © 53 3 et 1 est esb 1 e Fic. 5 Hemiscorpius acanthocercus sp. n. Male paratype (MHNG, H. Löffler coll.): A, pedipalp with trichobothrial pattern, dorsal aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur and patella, external aspect; F, idem, ventral aspect. Female paratype (NHMW 4719): D, pedi- palp, dorsal aspect. Scale line, 2 mm. chobothria) orthobothriotaxic, located basally on external surface; Esb very close to Eb group: Em absent; Est located distally, very close to Et group; Et group composed of 5 trichobothria, Er; located ventrally; V group comprising 4 trichobothria, V3 and V4 lo- cated in basal half of manus, V/ and V2 located very distally. Fingers of pedipalpal chela (Fig. SA-C). Basally reddish tan, becoming gradually lighter distally, tips of fin- ee HEMISCORPIIDAE FROM IRAN 883 Fic. 6 Hemiscorpius acanthocercus sp. n., pectines and genital opercula, ventral aspect: A, male holo- type; B, female paratype (VHMW 21143). Scale line, 1 mm. 884 L. MONOD & W. R. LOURENCO gers lighter orange. Dorsal and ventral surfaces predominantly smooth and minutely pitted, slightly shagreened basally. Fingers slightly shorter than chelal manus. Fixed finger with weak basal concavity, movable finger with corresponding lobe weakly de- veloped. Tips of fingers with pronounced terminal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of denticles becoming fused basally, interrupted at regular intervals by stronger granules, each of these coupled with an accessory gran- ule; rows fused into a single row above the concavity on the fixed finger and on notch of movable finger, single basal row running to base of fingers, in some males absent on movable finger. A total of 11 trichobothria present on fingers; Dt located very basally on dorsal surface; db located dorsally on internal surface, in basal half of fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb located dorsally on exter- nal surface, in basal 1/3 of fixed finger, opposite db; esb, est and et in distal half of fixed finger, opposite dsb, dst and dt respectively; it and ib located in median 1/3 of fixed finger. Coxosternal sclerites. Dark yellow to orange; smooth and minutely pitted. Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, penta- gonal, slightly shagreened; median furrow deep, more pronounced in posterior half; posterior pit absent. Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal surfaces of trochanter and femur finely and weakly granular. Tarsus with a ventro- median row of microspinules and with 2 rows of rigid “spinoid” ventral macrosetae, tarsi I-II: prolateral row with 4/5 macrosetae, retrolateral row with 5/6 macrosetae; tarsi I-IV: prolateral row with 5/6 macrosetae, retrolateral row with 6 macrosetae; setae acuminate; tarsal claws of equal length. Pectines and genital operculum (Fig. 6A). Colour dark yellow; genital oper- culum composed of 2 subtriangular plates; genital papillae short, not protruding from beneath operculum; pectines with 14/15 teeth. Hemispermatophore (Fig. 7) lamelliform, with complex capsule; distal lamellum slender, basally curved, with a strong double hook pointing antero-distad and located very basally, flagellum more than 1.5 times longer than basal part; distal crest absent. Double lamellar hook (Dh) located very slightly above distal transverse ridge (Tr); distal transverse ridge costate, distally strongly curved towards anterior margin, continuous from posterior to anterior margins, almost merging with lamellar hook, reaching anterior margin of distal lamella slightly below its hook. Capsule lamella (La) broad, with a deep longitudinal furrow, forming an acute angle with longitudinal axis of capsule, bearing a strong hook on its external surface and an accessory lobe (Lac) on its posterior margin; ventral margin straight. Distal lobe (Ld) forming a strong hook pointing anteriad; basal margin straight. Basal lobe (Lb) strongly reduced. Posterior lamella (Lp) costate. FEMALE (Figs 2C-D, 3B, 4A, 5D, 6B). Measurements of paratype (NHMW 4719). Carapace, length 5.9, posterior width 5.0; distance between anterior lateral eyes 3.1, between posterior lateral eyes 3.6, between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, femur length 5.0, width 1.9; patella length 5.2, width 1.8, chela length 10.1; manus length 5.2, width 4.0, depth 2.7; movable finger length 5.2; fixed finger length 3.9. Metasoma, segment I length 3.3, width 1.9; segment V length 5.1, HEMISCORPIIDAE FROM IRAN 885 17 a qa a 1504 FIG. 7 Hemiscorpius acanthocercus sp. n., male holotype, hemispermatophore: A, whole hemisper- matophore, arrows indicating its post-extrusion orientation, Ant (anterior), Bas (basal), Dist (distal), Post (posterior), Tr (transverse ridge); B-C, detail of the capsular region, internal aspect, Dh (double hook), La (lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (pos- terior lobe). Scale lines, 0.25 mm. 886 L. MONOD & W. R. LOURENCO width 1.4, depth 1.7; vesicle length 3.7, width 2.0, depth 2.1, aculeus length 1.2. Total length 53.0. Same characters as in males, except as follows. Carapace (Fig. 3B) shagreened and finely granular, each anterior lobe with a smooth, shiny and minutely pitted median area between median and lateral ocular tubercles; median longitudinal furrow shallow, continuous from the anterior suture furcation, ending after the median ocular tubercle and not running posteriorly into a deep triangular depression. Mesosoma. Median carina absent from tergites I-III. Surface of tergites smooth, shiny; median area finely granu- lar; median carina on tergites IV-IV smooth and pitted posteriorly. Sternites III-VI smooth, without granulation or carinae. Sternite VII smooth, with a pair of median cari- nae developed as very weak ridges, almost indiscernible. Tergites and sternites VII as long as wide. Metasoma (Fig. 4A). Short and stout, shagreened, without granules. Segments I-II with ventrolateral carinae forming almost smooth ridges. Segment I without ventral carina; segments II with ventral carina developed as a ridge, smooth in anterior half, with weak granules in posterior half; segment III with ventral carina form- ing a ridge of weak granules; segment IV with ventral carina developed as a ridge of weak spiniform granules. Segment V: dorsal carinae with spiniform granules; lateral carina vestigial, barely visible in anterior 2/3, composed of few very weak scattered granules; ventromedian carina with strong spiniform granules. Telson (Fig. 4A). Vesicle globular, very bulky; lateral surfaces smooth, with few extremely weak granules (almost indistinct); dorsal surface smooth, median shallow depression indistinct, only visible anteriorly. Aculeus without tubercles at its base, narrowing slightly less marked than in male. Pedipalp patella (Fig. 5D). Internal protuberance with internodorsal carina carrying 1-2 spiniform granules, less developed than in male. Pedipalpal chela (Fig. 5D) slightly more bulky than in male, chela fingers equal in length to chela manus or slightly shorter. Carinae of chelal manus generally less granular and less distinct than in males. Interno-dorsal carina vestigial, basally visible as few strong spiniform granules; sub- digital carina vestigial; externodorsal carina distinct, visible as a smooth ridge; digital carina developed as a weak ridge with few weak granules basally; dorsal secondary carina (dorsomedian) forming a weak, smooth ridge with 1-2 strong granules basally; internoventral carina continuous, visible as a weak ridge with few weak granules dis- tally; externoventral carina continuous, a ridge of indistinct granules, becoming almost smooth distally. Ventral surface smooth and pitted with shagreened patches. Pectines and genital operculum (Fig. 6B). Genital operculum composed of a single posteriorly truncated subtriangular plate with a median longitudinal furrow and a relatively deep posterior notch; pectines with 10/10 teeth. Intraspecific variation. Females have pectines with 8 to 11 teeth, usually 9/9 or 10/10, males have pectines with 13 to 16 teeth, usually 14/14 or 15/15. Juveniles (males and females) have slender pedipalps, which are morphologically similar to the pedi- palps of mature males. Juveniles and subadult males do not possess an extremely elon- gated metasoma, they apparently acquire this feature only in the last developmental stage. Hemiscorpius enischnochela sp. n. Figs 1E-F, 8-12, 26A-B, 27 C-D, 36 Material examined. Holotype: d, IRAN (Hormozgan), ca. 115 km E Bandar Abbas, VII.1974, G. Pretzman, NHMW (52). Paratypes: 2 2, IRAN (Khuzestan), S of Masdjed-e- HEMISCORPIIDAE FROM IRAN 887 Fic. 8 Hemiscorpius enischnochela sp. n. Male holotype: dorsal aspect. Scale line, 5 mm. Soleyman, 1.1970, R. Farzanpay ?, MHNG (SF 0057-1/2). 1 @ juv., same data as for holotype, NHMW (53).1 2 subadult, IRAN (Hormozgan), 38 km N Bandar Abbas, 28.III.1972, K. Bilek, NHMW (54). 1 2 juv., IRAN (Hormozgan), 38 km N Bandar Abbas, 28.III.1972, K. Bilek, NHMW (58). 1 © juv., IRAN (Hormozgan), 22 km N Bandar Abbas, VII.1974, G. Pretzman ?, NHMW (60). 1 2 juv., IRAN (Hormozgan), 65 km N Bandar Abbas, 4.IV.1972, K. Bilek, 888 L. MONOD & W. R. LOURENCO NHMW (64). 1 3 juv., IRAN (Hormozgan), 65 km N Bandar Abbas, 4.IV.1972, K. Bilek, NHMW (67). 1 2, IRAN (Hormozgan), Kol River bridge on a road near Lar, 32 km W Bandar Abbas, 1972, K. Bilek, NHMW (76). 2 3 subadults, IRAN (Hormozgan), Bandar Abbas, VI.1972, G. Pretzman, NHMW (80-81). 1 2 juv., IRAN (Hormozgan), ca. 41 km N Bandar Abbas, 7.1V.1972, G. Pretzman & A. Konetschnig, NHMW 3395. Distribution. Known from surroundings of Bandar Abbas, Hormozgan Province, eastern Iran; S of Masdjed-e-Soleyman, Khuzestan Province (doubtful record, see below) (Fig. 36). Remark. The species occurs around Bandar-Abbas in Eastern Iran. Among the material studied by Vachon, there was a female found far from this area, near Masdjed- e-Soleyman. There was no indication about the collector, who could possibly be R. Farzanpay. Since much information about Vachon’s material was lost and mixed up after his death, it could be a case of mislabelling. Etymology. The name enischnochela is a construct from the Greek words enischnos [slender, thin] and chele [claw]. The name is an invariable noun in appo- sition and refers to the pedipalp chela, which is slender and with long fingers. Diagnosis. (1) Carapace longer than wide, shagreened and sparsely granular, with coarse granules on anterior margin between lateral ocular tubercles; lateral mar- gins finely granular, especially below lateral ocular tubercles; superciliary carinae weak and finely granular; (2) pedipalps elongated, chela fingers slightly longer than chela manus; (3) internal protuberance of pedipalp patella with internodorsal carina bearing 3-4 strong spiniform granules; (4) pedipalp patella neobothriotaxic, external side with 14 trichobotria (1 est and 3 esb), ventral side with 10-12 trichobothria; (5) males with elongated and slender metasoma, with dorsal carinae of segments I-V and ventral and ventrolateral carinae of segments IV-V bearing small spiniform granules; (7) telson strongly elongated, bearing a pair of blunt tuberculiform processes at base of aculeus, surface irregular, rugose, without distinct granules; (8) metasoma of females with dorsal carinae of segments I-IV and ventral and ventrolateral carinae of segments IV-V bearing spiniform granules. H. enischnochela sp. n. is closely related to A. gaillardi, both of them have elongated neobothriotaxic pedipalps. They can be readily distinguished by (1) their tri- chotriotaxy (H. gaillardi has an additional est trichobothry on the external face of the patella) and by (2) their pedipalp carination (H. enischnochela sp. n. has patellar processes with intero-dorsal carinae bearing 3 strong spiniform granules, while H. gail- lardi has patellar processes with carinae bearing 6 spiniform granules, which are slightly smaller than those in 7. enischnochela sp.n.). Description. MALE (Figs 1E-F, 8, 10A-C, E-F, 11A, 12,26A,27D) [description based on the holotype (NHMW 52)]. Measurements of holotype (in mm). Carapace, length 7.6; posterior width 7.0; distance between anterior lateral eyes 4.0, between pos- terior lateral eyes 4.9, between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, femur length 8.8, width 2.7; patella length 8.6, width 2.6, chela length 15,2; manus length 7.4, width 4.6, depth 3.2; movable finger length 8.2; fixed finger length 7.2. Metasoma, segment I length 9.6, width 2.5; segment V length 12.3, width 1.6, depth 2.0; vesicle length 7.4, width 2.3, depth 1.9, aculeus length 1.0. Total length 93.0. Carapace (Fig. 26A). Colouration mostly uniformly dark yellow to orange; median and lateral ocular tubercle black. Carapace dorsoventraly flattened, longer than HEMISCORPIIDAE FROM IRAN 889 Fic. 9 Hemiscorpius enischnochela sp. n. Female paratype (VHMW 76, Kol River): A, dorsal aspect; B, ventral aspect. Scale line, 5 mm. 890 L. MONOD & W. R. LOURENCO Fic. 10 Hemiscorpius enischnochela sp. n. Male holotype: A, pedipalp with trichobothrial pattern, dor- sal aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur and patella, external as- pect; F, idem, ventral aspect. Female paratype (MHNG, Masdjed-e-Soleyman): D, pedipalp, dor- sal aspect. Scale line, 2 mm. wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; lat- eral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. Carapace shagreened, with sparse granules; stronger granules on the anterior margin between lateral ocular tubercles; edges granular, with stronger spiniform granules be- low lateral ocular tubercles; anteromedian furrow narrow, suturiform, anteriorly bifur- cated; median longitudinal furrow shallow, continuous from the anterior suture furca- tion, running through ocular tubercle posteriorly into a deep triangular depression; pos- terolateral furrow shallow; mesolateral furrow weakly developed, almost indiscernible. HEMISCORPIIDAE FROM IRAN 891 FIG. 11 Hemiscorpius enischnochela sp. n., pectines and genital opercula, ventral aspect: A, male holo- type; B, female paratype (MHNG, Masdjed-e-Soleyman). Scale line, 1 mm. 892 L. MONOD & W. R. LOURENCO Mesosoma. Tergite colouration dark orange to brown, becoming lighter poste- riorly (dark yellow), tergite VII with a larger yellow posterior zone than on other ter- gites. Tergites I-III with a shallow median depression divided anteriorly by a weak ridge, tergites IV-VII with median carina reduced to a weak non-granular ridge sur- rounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae ab- sent on tergites I-VI, but present on posterior 2/3 of tergite VII, visible as strong ridges with spiniform granules. Surface of tergites shagreened, with scattered granules; pre- tergites smooth, shiny and minutely pitted. Sternite colouration pale yellow. Sternites II-VI smooth, shiny and minutely pitted, without granulation or carinae; sternite III with a pair of large, very finely and densely granular depressions, covered by the pectines; sternites IV-VI with a pair of shallow median furrows. Sternite VII with sim- ilar surface texture as on preceding sternites; a pair of lateral, slightly granular carinae present; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII longer than wide. Metasoma (Fig. 27D). Very elongated and slender, shagreened, with scattered granules. Colour dark yellow to orange. Segments I-IV with longitudinal dorsomedian furrow and with dorsal carinae composed of spiniform granules. Segments I-II with lat- eral carinae developed as granular ridges; segments III-IV with lateral carinae as ridges of indistinct fused granules, almost smooth. Segments I-II with ventrolateral carinae as granular ridges; segments III-IV with ventrolateral carinae as ridges of reduced spini- form granules. Segment I: ventral carina almost absent, reduced to a weak ridge; seg- ments II-III with ventral carina forming a ridge of fused granules, almost smooth; seg- ment IV with ventral carina forming a ridge of fused granules, almost smooth in the anterior half, and with more defined spiniform granules posteriorly. Segment V: longi- tudinal dorsal furrow present in anterior half, dorsal carinae with reduced spiniform granules; lateral carina indistinct, only visible in anterior half as a ridge of fused gran- ules; ventrolateral carinae with reduced spiniform granules; ventromedian carina with spiniform granules. Telson (Fig. 1E-F, 27D). Vesicle orange; aculeus darker, tan, due to stronger sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt tuberculiform projection on each side at base of aculeus; ventrolateral furrows absent; ventromedian ridge absent; lateral surfaces rugose, irregular, with scattered weak granules; dorsal surface with numerous small spiniform granules and a median shallow depression. Macrosetae very sparse basally, becoming more numerous near base of aculeus. Aculeus short and stout, strongly curved, becoming markedly narrower approximately midway. Chelicerae. Colour pale yellow; teeth of fingers darker orange. Teeth arrange- ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth bifid; movable finger with one subdistal tooth and one basal tooth in external series; distal external tooth smaller than distal internal tooth; cheliceral teeth without second- ary serrations. Pedipalp slender and elongated. Pedipalp coxa and femur (Fig. 10A, E-F). Internoventral margin of coxa with strong spiniform granules. Dorsal surface of femur predominantly dark yellow-orange, internodorsal, internoventral and externoventral carinae darker orange. Femur slender, elongate (equal to or longer than 3 times its HEMISCORPIIDAE FROM IRAN 893 — Ce / OT ef Li cer MSA | SS “ = SU dr Ai sia Li (ji i n x « È DL 44 7 & AU] RATE si Min : EN mS LI aa ANY Al A SENET LORI RIVESTITO È AMEN o cpu! Fie Hemiscorpius enischnochela sp. n., male holotype: left tarsus IV, ventral aspect. Scale line, 0.5 mm. width), pentacarinate, with 4 distinct carinae; internodorsal carina developed as a strongly and densely granular ridge, granules spiniform; externodorsal carina a strong granular ridge; internoventral carina developed as a granular ridge with spiniform granules; externoventral carina developed as a ridge with strong spiniform granules; ventromedian carina reduced to few granules and confined to base of femur. Dorsal surface shagreened, finely and densely granular, distal end smooth, without granules; internal surface shagreened, with few strong scattered spiniform granules; external sur- face shagreened; ventral surface shagreened and sparsely granular, distal end without 894 L. MONOD & W. R. LOURENCO granules. A total of 3 trichobothria present on femur; d located externobasally on dor- sal surface; i located dorsobasally on internal surface; e located dorsobasally on exter- nal surface. Pedipalp patella (Figs 10A, E-F) slender (length less than 2.5 times its width). Colour predominantly dark yellow to orange; carinae slightly darker. 7 carinae present, 6 of them distinct; internodorsal carina developed as a ridge of coarse gran- ules; externodorsal carina forming a ridge of indistinct coarse granules; internoventral carina developed as a strongly granular ridge (spiniform granules); externoventral ca- rina as a ridge of weak granules; externomedian carina as a granular ridge. Dorsal and internal surfaces shagreened, sparsely and weakly granular; external surface sha- greened, with a few scattered granules; ventral surface shagreened, sparsely and weak- ly granular, distal end without granules. Internal protuberance pronounced, bifid (in- ternodorsal and internoventral tubercles separated), internodorsal carina with 3-4 strong spiniform granules. A total of 28 trichobothria present on patella, neobothrio- taxy major; d, located basally, external to internodorsal carina; d> situated in distal half of patella; d; absent; i in distal half of patella, located dorsally on internal surface. External (e) trichobothrial groups discernible and neobothriotaxic major: eb group composed of 5 trichobothria, esb group of 3 trichobothria, em group of 2 trichobothria, et group of 3 trichobothria; est very close to et group. Ventral (v) group neobothrio- taxic, 11 trichobothria. Pedipalp chela manus (Figs 10A-C). Colour predominantly orange to red-brown; carinae darker red. Chela slender (length more than 3 times its width), with 5 distinct carinae; internodorsal carina continuous, with spiniform granules; subdigital carina vestigial, visible basally as a smooth ridge with weak granules; externodorsal carina distinct, visible as a ridge of indistinct fused granules, almost smooth; digital carina a smooth ridge with few weak granules basally; dorsal secondary carina (dorsomedian) a smooth ridge with few strong granules basally and few weak granules distally; internoventral carina continuous, visible as a weak ridge with few spiniform granules; externoventral carina continuous, forming a granular ridge, becoming almost smooth distally, running parallel to longitudinal axis of chela, its distal edge disconnected from external movable finger condyle and directed between external and internal condyles of movable finger; ventromedian carina ves- tigial, reduced to few coarse granules basally; internal (internomedian) carina visible as a weak ridge with few spiniform granules; external (externomedian) carina hardly discernible, visible as raised rows of small indistinct granules. Dorsal surface sha- greened, sparsely and weakly granular; internal surface shagreened, sparsely granular, with a denser patch of spiniform granules dorsodistally; external surface shagreened, sparsely and weakly granular; ventral surface shagreened, sparsely and weakly granu- lar, distal end without granules. A total of 15 trichobothria present on chelal manus; Db trichobothria situated externobasally on dorsal surface; Eb group (3 trichobothria) or- thobothriotaxic, located basally on external surface; Esb very close to Eb group; Em absent; Est located distally, very close to Et group; Et group composed of 5 tri- chobothria, Et, located ventrally; V group comprising 4 trichobothria, V3 and V4 situ- ated in basal third of manus, V/ and V2 located in distal quarter. Pedipalp chela fingers (Figs 10A-C). Basally reddish tan, becoming gradually lighter distally, tips of fingers yellow. Dorsal and ventral surfaces predominantly smooth, slightly shagreened basally. Fingers longer than chela manus. Fixed finger with a weak basal concavity, movable HEMISCORPIIDAE FROM IRAN 895 finger with corresponding lobe weakly developed. Tips of fingers with pronounced distal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of den- ticles becoming fused basally, interrupted at regular intervals by stronger granules, each of them coupled with an accessory granule; rows fused into a single row above concavity on fixed finger and on notch of movable finger, single basal row running towards base of fingers. A total of 11 trichobothria present on fingers; Dr located on dorsal surface, in basal third of fixed finger; db located dorsally on internal surface, ap- proximately midway on fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb located dorsally on external surface, approximately midway on fixed finger, opposite db; esb, est and et in distal half of fixed finger; esb and et opposite dsb and dt, respectively; it and ib located in distal half of fixed finger. Coxosternal sclerites. Dark yellow to orange, smooth and minutely pitted. Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentago- nal, shagreened; median furrow deep, more pronounced in posterior half; posterior pit absent. Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal surfaces of trochanter and femur finely granular. Tarsus (Fig. 12) with a ventromedian row of microspinules and with 2 rows of rigid “spinoid” ventral macrosetae; tarsi I-II: prolateral row with 5 or 6 macrosetae, retrolateral row with 7 macrosetae; tarsi III-IV: prolateral row with 7 macrosetae, retrolateral row with 7/8 macrosetae; setae acumi- nate; tarsal claws of equal length. Pectines and genital operculum (Fig. 11A). Colour pale yellow; genital opercu- lum composed of 2 ovoid plates; genital papillae short, not protruding from beneath operculum; pectines with 17/17 teeth. Hemispermatophore. Unknown. The specimen was already dissected and the hemispermatophores were probably lost. FEMALE (Figs 9, 10D, 11B, 26B, 27C). Measurements of paratype (MNHN Masdjed-e Soleyman). Carapace, length 7.4, posterior width 7.0; distance between an- terior lateral eyes 4.0, between posterior lateral eyes 4.7, between median eyes 0.2; di- ameter of median eyes 0.2. Pedipalp, femur length 7.5, width 2.4; patella length 7.2, width 2.6, chela length 13.6; manus length 6.8, width 4.6, depth 3.3; movable finger length 6.9; fixed finger length 6.2. Metasoma, segment I length 4.8, width 2.2; segment V length 6.9, width 1.6, depth 2.0; vesicle length 5.2, width 2.8, depth 2.7, aculeus length 1.3. Total length 64.0. Same characters as in males, except as follows. Carapace (Fig. 26B) shagreened and sparsely granular, each anterior lobe with a reduced central smooth area between median and lateral ocular tubercles. Mesosoma. Surface of tergites shagreened, with few smooth and pitted patches and scattered granules. Tergites and sternites VII almost as long as wide. Metasoma (Fig. 27C). Short and stout. Segments I-II with ventral carina developed as a smooth ridge; segments III with ventral carina as a ridge of weak spiniform granules; segment IV with ventral carina as a ridge of spiniform granules. Segment V: lateral carina indistinct; ventromedian carina with spiniform granules. Telson (Fig. 27C). Vesicle elliptical or ovate, short and bulky; lateral surfaces smooth and irregular; dorsal surface smooth and irregular, with a shallow median depression 896 L. MONOD & W. R. LOURENCO posteriorly. Aculeus without tubercles at its base, narrowing slightly less marked than in male. Pedipalp coxa and femur (Fig. 10D). Pedipalp slightly less slender and elon- gated than in males. Femur: externoventral carina developed as a ridge with spiniform granules, these weaker, more scattered and less numerous than in male. Pedipalp patel- la (Fig. 10D). Less elongated than in males. Externodorsal carina developed as a ridge of weak granules, almost smooth. Internal and external surface shagreened, not granu- lar. Pedipalp chela manus (Fig. 10D). Chela more bulky than in males. Pectines and genital operculum (Fig. 11B). Genital operculum composed of a hexagonal ovoid plate divided by a moderately deep longitudinal furrow; posterior notch weakly pronounced; pectines with 8-10 teeth. Intraspecific variation. The specimens examined possess 10 to 12 trichobothria on the ventral side of the patella. Females have pectines with 8 to 10 teeth, usually 8- 8, males have pectines with 14 to 17 teeth. The same remark about pedipalps and meta- soma of males can be given as for H. acanthocercus. Hemiscorpius gaillardi (Vachon, 1974) comb. n. Figs 13-15, 26C Habibiella gaillardi Vachon, 1974: 952, fig. 105. Material examined. Holotype: 1 9 , IRAN, East, no locatity specified, MNHN-RS 4328. Distribution. Known from eastern Iran, no locatity specified. Diagnosis. (1) Carapace almost as wide as long, shagreened and sparsely granular, with few smooth patches and with coarse granules on anterior margin be- tween lateral ocular tubercles; lateral margins with small spiniform granules below lat- eral ocular tubercles; superciliary carinae weak, finely granular; (2) pedipalps very elongated, chela fingers longer than chela manus; (3) internal protuberance of patella with internodorsal carina bearing 6 strong spiniform granules; (4) pedipalp patella neothobothriotaxic, external side with 15 trichobotria (2 est and 3 esb), ventral side with 10-12 trichobothria; (5) metasoma of females with dorsal carinae of segments I-IV and ventral and ventrolateral carinae of segments V bearing weak small spiniform granules. The male of H. gaillardi remains unknown. For more details see also diagnosis of H. enischnochela sp.n., to which H. gaillardi is closely related. Description. FEMALE. Measurements of holotype (MNHN-RS 4328). Carapace, length 6.6, posterior width 6.7; distance between anterior lateral eyes 3.6, between posterior lateral eyes 4.4, between median eyes 0.2; diameter of median eyes 0.3. Pedipalp, femur length 7.7, width 2.3; patella length 7.5, width 2.4, chela length 14.2; manus length 6.2, width 3.8, depth 2.7; movable finger length 7.7; fixed finger length 6.7. Metasoma, segment I length 4.5, width 2.2; segment V length 6.5, width 1.7, depth 1.9; vesicle length 4.2, width 2.3, depth 2.3, aculeus length 1.5. Total length 58.0. Carapace (Fig. 26C). Colouration mostly uniformly pale yellow to orange; median and lateral ocular tubercle black. Carapace dorsoventraly flattened, almost as wide as long, with sides nearly parallel, slightly convergent; median ocular tubercle weakly developed, low, distinctly situated anteriorly, with superciliary carinae weak but finely granular. Frontal concavity or notch well-developed; anterior lobes truncate; HEMISCORPIIDAE FROM IRAN 897 Fic. 13 Hemiscorpius gaillardi, female holotype: A, dorsal aspect; B, ventral aspect. Scale line, 5 mm. 898 L. MONOD & W. R. LOURENCO = nana csrl ER „A S fan > 1 È 6 È h = i # 8 à « > RUE Nr ING = RY, # Ni add x © NN es pe eee § ry È est 2 1 À ; | È WS em 1-2 : N esb 1-2 VI esb 3 NI & a NEC I\ EX Fic. 14 Hemiscorpius gaillardi, female holotype, pedipalp with trichobothrial pattern: A, chela, dorsal aspect; B, idem, external aspect; C, idem, ventral aspect; D, femur and patella, dorsal aspect; E, idem, external aspect; F, idem, ventral aspect. Scale line, 2 mm. en nn rn HEMISCORPIIDAE FROM IRAN 899 Fic. 15 Hemiscorpius gaillardi, female holotype: A, metasoma, lateral aspect (scale line, 2 mm); B, pectines and genital opercula, ventral aspect (scale line, 1 mm). 900 L. MONOD & W. R. LOURENCO lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. Carapace shagreened and sparsely granular, with few smooth patches and with coarse granules on anterior margin between lateral ocular tubercles, each lobe with a smooth central area between median and lateral ocular tubercles; edges granular below lateral ocular tubercles; anteromedian furrow indistinct, very shallow, suturiform, anteriorly bifurcated; median longitudinal furrow indistinct, almost absent, very shallow, conti- nuous from the anterior suture furcation, running through ocular tubercle posteriorly into a deep triangular depression; posterolateral furrow shallow; mesolateral furrow weakly developed, almost indiscernible. Mesosoma. Tergite colouration dark yellow, with lighter (pale yellow) posteri- or margins. Tergites I-III with a shallow median depression divided anteriorly by a weak ridge, tergites IV-VII with median carina reduced to a weak, non-granular ridge surrounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae absent on tergites I-VI, but present in posterior 2/3 of tergite VII, developed as weak ridges with weak, spiniform granules. Surface of tergites shagreened, with smooth patches; pre-tergites smooth, shiny and minutely pitted. Sternite colouration pale yel- low. Sternites III-VI smooth, shiny and minutely pitted, without granulation or carinae; sternite II with a pair of large, very finely and densely granular depressions, situated under the pectines; sternites IV-VI with a pair of shallow median furrows. Sternite VII shagreened, very finely granular and with very few smooth patches; pair of lateral cari- nae, weak, smooth; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII almost as long as wide. Metasoma (Fig. 15A). Slender, predominantly shagreened. Colour dark yellow. Segments I-IV with longitudinal dorsomedian furrow, their dorsal carinae with spini- form granules. Segments I-II with lateral carinae developed as ridges with weak spini- form granules, almost smooth; segments IV with lateral carinae as almost smooth ridges. Ventrolateral carinae developed as almost smooth ridges on segments I-II; as smooth ridge with weak spiniform granules in posterior half of segment III; as ridge with weak spiniform granules on segment IV. Segment I: ventral carina almost indis- cernible, reduced to a very weak ridge; segments II-IV with ventral carina a smooth ridge with weak spiniform granules on segment IV. Segment V: longitudinal dorsal furrow present in anterior half, dorsal carinae weakly marked, with small, very weak granules; lateral carina absent; ventrolateral carinae with weak spiniform granules; ventromedian carina with small spiniform granules. Telson (Fig. 15A). Vesicle yellow to orange; aculeus darker, tan, due to stronger sclerotisation; vesicle elliptical or ovate, very bulky; ventrolateral furrows absent; ventromedian ridge absent; lateral surfaces smooth; dorsal surface smooth, with a median shallow depression anteriorly. Macrosetae very sparse basally, becoming more numerous near base of aculeus. Aculeus short, strongly curved, becoming markedly narrower approximately halfway. Chelicerae. Colour pale yellow; teeth of fingers darker, orange. Teeth arrange- ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth bifid; movable finger with one subdistal tooth and one basal tooth in external series; distal external tooth smaller than distal internal tooth; cheliceral teeth without secondary serrations. HEMISCORPIIDAE FROM IRAN 901 Pedipalp slender and elongated. Pedipalp coxa and femur (Fig. 14D-F). Inter- noventral margin of coxa with few strong spiniform granules. Dorsal surface of femur predominantly yellow, internodorsal, internoventral and externoventral carinae darker, orange. Femur slender, elongate (longer than 3 times its width), pentacarinate, with 4 distinct carinae; internodorsal carina developed as a strongly and densely granular ridge, granules spiniform; externodorsal carina as a granular ridge; internoventral carina as a ridge with spiniform granules; externoventral carina as a ridge with strong spiniform granules; ventromedian carina reduced to few scattered granules and con- fined to base of femur. Dorsal surface shagreened, finely and densely granular, distal end smooth, without granules; internal surface shagreened, with few strong scattered spiniform granules; external surface shagreened; ventral surface shagreened and sparsely granular, distal end without granules. A total of 3 trichobothria present on femur; d located externobasally on dorsal surface; i located dorsobasally on internal surface; e located dorsobasally on external surface. Pedipalp patella (Fig. 14D-F) slender (length more than 2.5 times its width). Colour predominantly yellow; carinae slightly darker. 7 carinae present, 6 of them distinct; internodorsal carina developed as a ridge of coarse granules; externodorsal carina as a ridge of indistinct coarse granules (almost smooth); internoventral carina as a strongly granular ridge (spiniform granules); externoventral carina as a ridge of weak granules; externomedian carina as a granular ridge. Dorsal surface shagreened, sparsely and weakly granular; internal, external and ventral surfaces shagreened, without granules. Internal protuberance pro- nounced, bifid (internodorsal and internoventral tubercles separated), internodorsal carina with 6 strong spiniform granules. A total of 29-31 trichobothria present on patella, neobothriotaxy major; d) located basally, external to internodorsal carina; d; located in distal third of patella; d3 absent; i in distal third of patella, located dorsally on internal surface. External (e) trichobothrial groups discernible and neobothriotaxic: eb group composed of 5 trichobothria, esb group of 3 trichobothria, em group of 2 tri- chobothria, est group of 2 trichobothria, et group of 3 trichobothria. Ventral (v) group neobothriotaxic, composed of 10 to 12 trichobothria. Pedipalp chela manus (Fig. 14A- C). Colour predominantly yellow; carinae, darker orange. Chela slender (length more than 3 times its width), with 5 distinct carinae; internodorsal carina continuous, with spiniform granules; subdigital carina vestigial, visible basally as a smooth ridge; ex- ternodorsal carina distinct, visible as a ridge of indistinct fused granules, almost smooth; digital carina a smooth ridge with few weak granules basally; dorsal secondary carina (dorsomedian) a smooth ridge with few strong granules basally and few weak granules distally; internoventral carina continuous, a weak ridge with spini- form granules; externoventral carina continuous, a granular ridge, running parallel to longitudinal axis of chela, its distal edge disconnected from external movable finger condyle and directed between external and internal movable finger condyles; ventro- median carina vestigial, reduced to few coarse granules basally; internal (interno- median) carina visible as a weak ridge with few spiniform granules; external (externo- median) carina weakly pronounced, visible as raised rows of small weak granules. Dorsal surface of chela shagreened, sparsely and weakly granular; internal surface sha- greened, sparsely granular, with a denser patch of spiniform granules dorsodistally; external surface shagreened, sparsely and weakly granular; ventral surface shagreened, 902 L. MONOD & W. R. LOURENCO sparsely and weakly granular, distal end without granules. A total of 15 trichobothria present on chelal manus; Db trichobothria located externo-basally on dorsal surface; Eb group (3 trichobothria) orthobothriotaxic, located basally on external surface; Esb very close to Eb group; Em absent; Est located distally, very close to Et group; Et group composed of 5 trichobothria, Er, located ventrally; V group comprising 4 trichobothria, V3 and V4 located in the basal half of manus, V/ and V2 situated very distally. Pedipalp chela fingers (Fig. 14A-C). Dark reddish tan, with tips yellow-orange. Dorsal and ven- tral surfaces predominantly smooth, slightly shagreened and minutely pitted basally. Fingers longer than chela manus. Fixed finger with a very weak basal concavity, movable finger with corresponding lobe very weakly developed. Tips of fingers with pronounced distal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of denticles becoming fused basally, interrupted at regular intervals by stronger granules, each of them coupled with an accessory granule; rows fused into a single row above concavity on fixed finger and on notch of movable finger, single basal row run- ning towards base of fingers. A total of 11 trichobothria present on fingers; Dt located basally on dorsal surface of fixed finger; db located dorsally on internal surface, ap- proximately midway on the fixed finger; dsb, dst and dt on dorsal surface, in distal third of finger; eb located dorsally on external surface, midway on fixed finger, opposite db; esb, est and et in distal third of fixed finger; esb, est and et, opposite dsb, dst and dt, re- spectively; it and ib located in distal half of fixed finger. Coxosternal sclerites. Dark yellow to orange; smooth. Anterior margin of coxapohysis I smooth, with few weak granules, expanded but not sub-triangular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentagonal, sha- greened; median furrow deep, more pronounced in posterior half; posterior pit absent. Legs. Pale yellow. Predominantly smooth, dorsal surfaces of trochanter and femur finely and weakly granular. Tarsus with a ventromedian row of microspinules and with 2 rows of rigid “spinoid” ventral macrosetae; tarsi I-II: prolateral row with 5-6 macrosetae, retrolateral row with 7 macrosetae; tarsi III-IV: prolateral row with 6 macrosetae, retrolateral row with 7 macrosetae; setae acuminate; tarsal claws of equal length. Pectines and genital operculum (Fig. 15B). Colour pale yellow; genital oper- culum composed of a single subtriangular plate, posterior extremity truncate, longitu- dinal median furrow relatively deep, posterior notch pronounced; pectines with 9/9 teeth. Hemiscorpius lepturus Peters, 1861 Figs 1A-B, 16-21, 27E-F, 36 Hemiscorpius lepturus Peters, 1861a: 426-427, 8 figs. Syntypes (not examined). 1 & (ZMB 43a), 1 9 (ZMB 43b), IRAQ, Baghdad; 2 specimens (ZMB 43), IRAQ, Mendeli; YEMEN, Aden. Material examined. 1 © (7408), IRAN (Kohkiluyeh), Charam road, 22.V.1974, A. Senglet, MHNG. 1 © juv. (7412b), IRAN (Kohkiluyeh), Yasudj road, 25.V.1974, A. Senglet, MHNG. 1 ¢ (2002/27), IRAN (Ghilan), 2 km W Cham-Khaled (Sea Star, Hotel Resort), nsl, 37°14.922’N, 50°15.037’E, 10.V.2002, B. Schätti, MHNG.2 5,2 2, IRAN (Khuzestan), Ahwaz region, 21.XI.1995, B. Masihpour, MHNG. 1 ¢, IRAN, ? Lali, VI.1961, M. Vasserot, MNHN- RS 4332.1 © juv., IRAN, 28.11.1961, M. Vasserot, MNHN-RS 4333.1 3,1 9, without locality, MNHN-RS 5232 (122-123).2 8,2 2,4 & subadult, IRAN (Khuzestan), Masched-Soleyman, R. Farzanpay ?, MHNG (SF 0006/13-20).1 3 juv.,1 2,2 © juv., IRAN (Fars), ca. 90 km W Shiraz, HEMISCORPIIDAE FROM IRAN 903 Fic. 16 Hemiscorpius lepturus. Male (MNHN-RS 5232): A, dorsal aspect; B, ventral aspect. Female paratype (MNHN, SF0006/17): C, dorsal aspect; D, ventral aspect. Scale line, 5 mm. 904 L. MONOD & W. R. LOURENCO 14.1V.1970, G. Pretzman, K. Bilek and F. Ressl, NHMW 3390.1 3 subadult, 3 © , IRAN (Fars), ca. 5 km NE Persepolis, 20.1V.1970, G. Pretzmann, K. Bilek and F. Ressl, NHMW 3391.1 3 juv., 1 9,1 © juv., IRAN (Fars), ca. 18 km SE Abadeh, 20.IV.1970, G. Pretzman, K. Bilek and F. Ressl, NHMW 3392. Distribution (completed with data from the literature). Esfahan, Fars, Hamadan, Kohkiluyeh, Khuzestan and Lorestan Provinces, south western Iran, eastern Iraq (Fig. 36). The distribution area of Hemiscorpius lepturus mentioned by Fet (2000) also in- cludes the west of Pakistan. However, this is not confirmed by the present study. The species seems to be restricted to Iraq and the western and south western regions of Iran. Diagnosis. (1) Carapace longer than wide, very finely granular with anterior area smooth and minutely pitted, and area around median ocular tubercle very finely granular; lateral margins smooth, without granules; superciliary carinae weak, smooth; (2) pedipalps stout and bulky, with weaker and less distinct granular carinae than in any other Iranian Hemiscorpiidae, chela fingers slightly shorter than chela manus; (3) internodorsal carina of patellar protuberance smooth, with 1-2 weak granules; (4) pedi- palp patella orthobothriotaxic, external side with 13 trichobotria (1 est and 2 esb), ven- tral side with 3 trichobothria; (5) metasoma of males elongated and slender, with dor- sal carinae bearing sparse spiniform granules (weaker granules on anterior segments), and ventral and ventrolateral carinae of segments IV-V developed as ridges with weak spiniform granules (carinae of segment V with more distinct spiniform granules in pos- terior half); (6) telson of males strongly elongated, bearing a pair of blunt tuberculi- form processes at base of aculeus, lateral surfaces rugose, without distinct granules; (7) metasoma of females with dorsal carinae of segments I-IV and ventral and ventrolat- eral carinae of segment V bearing strong spiniform granules; (8) hemispermatophore with double lamellar hook located above distal transverse ridge and pointing distally. See also diagnosis of H. acanthocercus sp. n., to which A. lepturus is closely related. Description. MALE (Figs 1A-B, 16A-B, 17A, 18A-C, E-F, 19A, 20, 21, 27F). Measurements of specimen MNHN-RS 5232. Carapace, length 6.4, posterior width 5.8; distance between anterior lateral eyes 3.6, between posterior lateral eyes 4.0, between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, femur length 6.0, width 2.2; patella length 5.8, width 2.2, chela length 10.8; manus length 6.0, width 4.4, depth 2.7; movable finger length 5.1; fixed finger length 3.8. Metasoma, segment I length 8.0, width 2.2; segment V length 10.4, width 1.6, depth 1.9; vesicle length 7.2, width 1.7, depth 2.0, aculeus length 0.8. Total length 85.0. Carapace (Fig. 17A). Colouration yellow to light orange; median and lateral ocular tubercle black. Carapace dorsodistally flattened, longer than wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle weakly devel- oped, very low, distinctly situated anteriorly, with superciliary carinae weak and smooth; frontal concavity or notch well-developed; anterior lobes truncate; lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. Carapace very finely granular, with few smooth patches; anterior area between median ocular tubercle and anterior margin smooth and minutely pitted, area around median ocular tubercle very finely granular; edges usually smooth, sometimes with very few weak granules below lateral ocular tubercles: anteromedian furrow narrow, suturiform, ante- riorly bifurcated; median longitudinal furrow shallow, continuous from anterior suture PE ii HEMISCORPIIDAE FROM IRAN 905 Fic. 17 Hemiscorpius lepturus, carapace, dorsal aspect (MNHN-RS 5232): A, male; B, female. Scale line, Imm. 906 L. MONOD & W. R. LOURENCO NL Rog Ren AN N \\ SÙ E Fic. 18 Hemiscorpius lepturus, male (MNHN-RS 5232): A, pedipalp with trichobothrial pattern, dorsal aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur and patella, external aspect; F, idem, ventral aspect. Female (MNHN-RS 5232): D, pedipalp, dorsal aspect. Scale line, 2 mm. furcation, running through ocular tubercle posteriorly into a deep triangular de- pression; postero-lateral furrow shallow; mesolateral furrow weekly developed, almost indiscernible. Mesosoma. Tergite colouration pale yellow. Tergite VII longer than wide. Tergite I without any depressions or carinae, tergites II-III with a shallow median depression divided anteriorly by a weak ridge, tergites IV-VII with median carina we mn ee eee HEMISCORPIIDAE FROM IRAN 907 reduced to a weak, non-granular ridge surrounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae absent on tergites I-VI, but present in posterior 2/3 of tergite VII, developed as ridges with indistinct granules, almost smooth. Surface of tergites finely granular; pre-tergites smooth, shiny and minutely pitted. Sternite colouration pale yellow. Sternite III smooth, shiny and minutely pitted, with a large, rough, very finely granular depression under each pectine; sternite IV-VI smooth, shiny and minutely pitted, without granulation or carinae, with a pair of shallow median furrows. Sternite VII smooth, shiny, minutely pitted medially and fine- ly granular laterally and posteriorly; a pair of weakly developed, smooth lateral carinae in posterior half; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII longer than wide. Metasoma (Fig. 27F). Very elongated and slender, slightly shagreened, with fine scattered granules. Colour yellow to light orange. Segments I-IV with longitudinal dor- somedian furrow, dorsal carinae developed as ridges with scattered spiniform granules becoming more numerous on posterior segments, lateral carinae as smooth ridges without granules. Segments I-II with ventrolateral carinae developed as smooth ridges; segments III-IV with ventrolateral carinae as ridges of indistinct granules, almost smooth. Segment I with ventral carina almost absent, reduced to a weak ridge; segments II-III with ventral carina a smooth ridge; segment IV with ventral carina a ridge of fused granules, almost smooth. Segment V: longitudinal dorsal furrow present in anterior half; dorsal carinae with small spiniform granules posteriorly; lateral carina developed as a weak ridge; ventrolateral and ventromedian carinae as ridges of indis- tinct granules, almost smooth anteriorly and with reduced spiniform granules in pos- terior half. Telson (Figs 1A-B, 27F). Vesicle yellow; aculeus darker, tan, due to stronger sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt tuberculiform projection on each side at base of aculeus; ventrolateral furrows and ventromedian ridge absent; lateral surfaces irregular, with numerous weak granules; dorsal surface with numerous small spiniform granules and a median shallow depression. Macrosetae very sparse basally, becoming more numerous near base of aculeus. Aculeus short and stout, strongly curved, becoming markedly narrower approximately midway. Chelicerae. Colour pale yellow; teeth of fingers darker, orange. Teeth arrange- ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth bifid; movable finger with one subdistal tooth and one basal tooth in external series; distal external tooth smaller than distal internal tooth; cheliceral teeth without secondary serrations. Pedipalp short and stout. Pedipalp coxa and femur (Fig. 18A, E-F). Coxa with internoventral margin mostly smooth, with only few strong spiniform granules. Dorsal surface of femur yellow, internodorsal, internoventral and externoventral carinae darker, orange. Femur short, stout (length less than 2.5 times its width), pentacarinate, with 4 distinct carinae; internodorsal carina developed as a strongly granular ridge; externodorsal carina reduced to a slightly raised row of granules; internoventral carina developed as a strongly granular ridge; externoventral carina as a weak granular ridge; ventromedian carina reduced to few granules and confined to base of femur. Dorsal surface shagreened, finely and densely granular, distal end smooth, without granules; 908 L. MONOD & W. R. LOURENCO Fic. 19 Hemiscorpius lepturus, pectines and genital opercula, ventral aspect: A, male (MNHN-RS5232); B, female (MHNG, SF 0006). Scale line, 1 mm. HEMISCORPIIDAE FROM IRAN 909 Fic. 20 Hemiscorpius lepturus, hemispermatophores of two males, external aspect: A, male from Masched-Soleyman (MHNG, SF 0006); B, male from Iran, without precise locality (MNHN-RS 4332). Scale line, 0.5 mm. 910 L. MONOD & W. R. LOURENCO internal surface shagreened, with few strong scattered spiniform granules; external surface slightly shagreened; ventral surface finely and densely granular, with shagreened patches, distal end without granules. A total of 3 trichobothria present on femur; d located externobasally on dorsal surface; i located dorsobasally on internal surface; e located dorsobasally on external surface. Pedipalp patella (Figs 18A, E-F) short and stout (length less than 2.5 times its width). Colour predominantly yellow; carinae slightly darker, orange. 7 carinae present, 6 of them distinct; internodorsal carina developed as a ridge of indistinct coarse granules; externodorsal carina as a weak ridge of indistinct granules, almost smooth; internoventral carina as a ridge of indistinct granules, almost smooth; externoventral carina as a smooth weak ridge, almost indiscernible; externomedian carina as a ridge of weak coarse granulaes, almost smooth. Dorsal surface finely and weakly granular, with small shagreened patches; internal surface shagreened, with few small scattered granules; external surface minutely pitted, with shagreened patches; ventral surface weakly and finely granular, with shagreened patches, distal end smooth and minutely pitted. Internal protuberance weak, bifid (internodorsal and internoventral tubercles separated), dorsal carina smooth, with 1-3 weak granules. A total of 19 trichobothria present on patella, orthobothriotaxy; d; located basally, external to internodorsal carina; d, located sub- medially on patella; d3 absent; i in distal half of patella, located dorsally on internal surface. External (e) trichobothrial groups discernible and orthobothriotaxic: eb group composed of 5 trichobothria, esb group of 2 trichobothria, em group of 2 trichobothria, et group of 3 trichobothria; est very close to er group. Ventral (v) group orthobothrio- taxic, 3 trichobothria. Pedipalp chela manus (Figs 18A-C). Colour predominantly yellow; carinae and fingers darker, orange. Chela short and stout (length less than 2.5 times its width), with 5 distinct carinae; internodorsal carina weakly developed, visible as a row of few granules, highlighted by a slightly darker line; subdigital carina vestigial, a smooth, weak ridge, more distinct basally; externodorsal carina vestigial, visible as a ridge of indistinct granules, almost smooth; digital carina vestigial, visible as a smooth, weak ridge more distinct basally; dorsal secondary carina (dorsomedian) vestigial, visible as a very weak, smooth ridge with a few weak granules basally; internoventral carina not continuous, developed as a weak, smooth ridge, with a few weak granules distally, less distinct basally; externoventral carina continuous, devel- oped as a ridge of weak fused granules, smooth distally, running parallel to longitudinal axis of chela, its distal edge disconnected from external movable finger condyle and directed between external and internal movable finger condyles; ventromedian carina reduced to 1-2 coarse granules basally; internal (internomedian) carina vestigial, visible as a row of very few sparse granules; external (externomedian) carina weakly developed, visible as ridge of small indistinct granules. Dorsal surface smooth and minutely pitted, with patches of a subreticulate network of weak granular ridges and shallow shagreened dimples; internal surface sparsely and finely granular, with small shagreened patches, with a patch of stronger spiniform granules dorsodistally; external surface minutely pitted, forming an irregular and subreticulate network of weakly and finely granular ridges and shallow shagreened dimples; ventral surface smooth and minutely pitted, with few small shagreened patches. A total of 15 trichobothria present on chela manus; Db trichobothria located externobasally on dorsal surface; Eb group HEMISCORPIIDAE FROM IRAN 911 TOLER = CAE) a enne; N Lac Lp Fic. 21 Hemiscorpius lepturus, hemispermatophores of two males, detail of capsular region, internal as- pect: A-B, male from Iran, without precise locality (MNHN-RS 4332), Dh (double hook), La (lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (posterior lobe), Tr (trans- verse ridge); C, male from Masched-Soleyman (MHNG, SF 0006). Scale lines, 0.5 mm. (3 trichobothria) orthobothriotaxic, located basally on external surface; Esb very close to Eb group; Em absent; Est located distally, very close to Et group; Et group composed of 5 trichobothria, Et; located ventrally; V group comprising 4 trichobothria, V3 and V4 located in the basal half of manus, V/ and V2 located very distally. Pedipalp chela fingers (Figs 18A-C). Dark orange to reddish tan, smooth, minutely pitted basally. DI L. MONOD & W. R. LOURENCO Fingers slightly shorter than chela manus. Fixed finger with a weak basal concavity, movable finger with corresponding lobe weakly developed. Tips of fingers with distal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of denticles becoming fused basally, interrupted at regular intervals by stronger granules, each of these coupled with an accessory granule; rows fused into a single row above concavity on fixed finger and on notch of movable finger, single basal row running towards base of fingers. A total of 11 trichobothria present on chelal fingers; Dt located very basally on dorsal surface of fixed finger; db located dorsally on internal surface, in basal half of fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb located dor- sally on external surface in basal half of fixed finger, opposite db; esb, est and et in distal half of fixed finger; esb, est and et, opposite dsb, dst and dr, respectively; it and ib located midway on fixed finger. Coxosternal sclerites. Yellow; smooth and minutely pitted. Anterior margin of coxapohysis I smooth, with few weak granules, expanded but not sub-triangular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentagonal, smooth and minutely pitted; median furrow deep, more pronounced in posterior half; posterior pit absent. Legs. Pale yellow. Predominantly shagreened; dorsal surfaces of trochanter and femur finely and densely granular; ventral surface smooth and minutely pitted. Tarsus with a ventromedian row of microspinules and with 2 rows of rigid “spinoid” ventral macrosetae; tarsi I-II: prolateral row with 5-6, retrolateral row with 7 macrosetae; tarsi I-IV: prolateral and retrolateral rows with 7 macrosetae; tarsal claws of equal length. Pectines and genital operculum (Fig. 19A). Pale yellow; genital operculum composed of 2 ovoid plates; genital papillae short, not protruding from beneath operculum; pectines with 14-16 teeth, ususally 15/15. Hemispermatophore (Figs 20-21). Lamelliform, with complex capsule; distal lamella slender, basally curved, with a double hook pointing distad and located basally, flagellum approximately 1.5 times longer than capsule region; distal crest absent. Double lamellar hook (Dh) located above transverse distal ridge (Tr), the latter costate, distally strongly curved towards anterior margin, continuous from posterior to anterior margins, not merging with lamellar hook, reaching anterior margin of distal lamella below its hook. Capsule lamella (La) broad, with a deep longitudinal furrow and forming an acute angle to longitudinal axis of capsule, bearing a strong hook on its external surface and an accessory lobe (Lac) on its posterior margin; ventral margin straight. Distal lobe (Ld) forming a strong hook pointing anteriad; basal margin straight. Basal lobe (Lb) absent. Posterior lamella (Lp) costate. FEMALE (Figs 16C-D, 17B, 18D, 19B, 27E). Measurements of specimen MNHN-SF0006/18. Carapace, length 6.0, posterior width 5.6; distance between anterior lateral eyes 3.4, between posterior lateral eyes 3.8, between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, femur length 5.7, width 2.1; patella length 5.6; width 2.1, chela length 10.7; manus length 6.0, width 4.5, depth 2.5; movable finger length 4.9; fixed finger length 3.6. Metasoma, segment I length 3.9, width 2.3; segment V length 5.7, width 1.6, depth 2.0; vesicle length 3.9, width 2.3, depth 2.3, aculeus length 1.1. Total length 56.0. Same characters as in males, except as follows. Carapace (Fig. 17B) less elon- gated than in male, almost as wide as long. Carapace finely granular, with larger HEMISCORPIIDAE FROM IRAN 913 longitudinal median area smooth and minutely pitted; area around median ocular tubercle finely and densely granular. Mesosoma. Tergite VII as wide as long. Surface of tergites smooth, shiny and minutely pitted, finely granular laterally; pre-tergites smooth, shiny and minutely pitted, tergites VI-VII more granular than others. Sternite VII as wide as long. Metasoma (Fig. 27E). Short and relatively stout, slightly shagreened, with fine scattered granules. Carinae generally more granular than in males, spiniform granules stronger. Segments I-IV, dorsal carinae developed as ridges with spiniform granules (denser than in males), lateral carinae as weakly granular ridges. Segments I-II with ventrolateral carinae developed as weakly granular ridges, segment III-IV with ventrolateral carinae bearing weak spiniform granules. Segments I-II with ventral carina reduced to a smooth ridge; segments III-IV with ventral carina a ridge of weak spiniform granules. Segment V: dorsal carinae with weak spiniform granules; lateral carina vestigial, reduced to a row of scattered granules, ventrolateral carinae with spiniform granules; ventromedian carina with spiniform granules. Telson (Fig. 27E). Vesicle ovoid, globular and bulky; lateral surfaces smooth, slightly irre- gular. Aculeus without tubercles at its base, narrowing slightly less marked than in male. Pedipalp (Figs 18D). Slightly shorter and stouter than in males. Pedipalp chela fingers (Figs 18D). Fixed and movable fingers with slightly weaker basal concavity and lobe, respectively. Pectines and genital operculum (Fig. 19B). Genital operculum composed of a single subtriangular plate, posterior extremity truncate, longitudinal median furrow relatively deep, posterior notch pronounced; pectines with 9-11 teeth, usually 10/10. Intraspecific variation. The size of males can vary greatly from 52 mm up to 85 mm body length, females display less variation in size. Females have pectines with 8-11 teeth, usually 10-10, males haves pectines with 14-16 teeth. The same remark about pedipalps and metasoma of males can be given as for H. acanthocercus. Hemiscorpius persicus Birula, 1903 Figs 22-25, 36 Hemiscorpion persicum Birula, 1903: 77-80. Material examined. Syntypes: 1 6, IRAN (Sistän va Balüchestan), “Province Ssarbas”, village Riss, 24.11.1901, N. Zarudny, ZISP 1120; 1 3 subadult, IRAN (Sistän va Balüchestan), “Province Ssarbas”, on the road between village Riss and locality Kaptegin-Dukan, 24- 26.11.1901, N. Zarudny, ZISP 1120. Distribution. Known only from Sistän va Balüchestan Province, Eastern Iran (Fig. 36). Diagnosis. (1) Carapace longer than wide, shagreened and finely granular, with small smooth patches; lateral margins with small spiniform granules below lateral oc- ular tubercles; superciliary carinae finely granular; (2) pedipalps relatively stout and bulky, chela fingers slightly shorter or equal in size to chela manus; (3) internal protu- berance of pedipalpal patella with internodorsal carina bearing 1-2 strong spiniform granules, usually 2; (4) pedipalp patella orthobothriotaxic, external side with 13 tri- chobotria (1 est and 2 esb), ventral side with 3 trichobothria; (5) metasoma of males relatively elongated and slender, with dorsal carinae of segments I-V and ventral and ventrolateral carinae of segments IV-V bearing spiniform granules; (7) telson of males not strongly elongated, without blunt tuberculiform processes at base of aculeus. 914 L. MONOD & W. R. LOURENCO Fic. 22 Hemiscorpius persicus, male syntype (ZISP 1120): A, dorsal aspect; B, ventral aspect. Scale line, 2.5 mm. HEMISCORPIIDAE FROM IRAN 915 ihn. Fic. 23 Hemiscorpius persicus, male syntype (ZISP 1120): A, carapace, dorsal aspect; B, metasoma, lat- eral aspect. Scale lines, 1 mm. L. MONOD & W. R. LOURENCO 916 br aa: "EN (ag Sy e. RS a SS IT SARA SSE SS 7 we Lt ho Mb yy Fic. 24 Hemiscorpius persicus, male syntype (ZISP 1120), pedipalp with trichobothrial pattern: A, chela, dorsal aspect; B, idem, external aspect; C, idem, ventral aspect; D, femur and patella, dorsal as- pect; E, idem, external aspect; F, idem, ventral aspect. Scale line, 1 mm. HEMISCORPIIDAE FROM IRAN 917 Fic. 25 Hemiscorpius persicus, male syntype (ZISP 1120): pectines and genital opercula, ventral aspect. Scale line, 0.5 mm. H. persicus is very different from all the others Iranian Hemiscorpius. Despite possessing a relatively slender metasoma, the male does not show the extreme sexual dimorphism of metasoma and telson observed in the other Iranian species and can therefore be easily distinguish from them. Description. MALE (description based on the syntype, ZISP 1120). Measurements of syntype (ZISP 1120). Carapace, length 4.0; posterior width 3.4; distance between anterior lateral eyes 2.0, between posterior lateral eyes 2.4, between median eyes 0.1; diameter of median eyes 0.1. Pedipalp, femur length 4.2, width 1.5; patella length 4.2, width 1.4, chela length 8.0; manus length 4.2, width 2.8, depth 1.7; movable finger length 4.1; fixed finger length 3.2. Metasoma, segment I length 3.2, width 1.4; segment V length 4.6, width 1.1, depth 1.3; vesicle length 2.8, width 1.1, depth 1.4, aculeus length 0.9. Total length 39.0. Carapace (Fig. 23A). Colouration mostly uniformly dark yellow to orange; median and lateral ocular tubercles black. Carapace dorsodistally flattened, longer than wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- 918 L. MONOD & W. R. LOURENCO ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. Carapace shagreened, finely and densely granular, with a few smooth patches; larger smooth patches in median area of anterior lobes between median and lateral ocular tubercles; lateral margins with small spiniform granules below lateral ocular tubercles; anteromedian furrow narrow, suturiform, anteriorly bifurcated; median longitudinal furrow shallow, continuous from anterior suture furcation, running through ocular tubercle, becoming wider posteriorly before merging with a deep triangular depression; posterolateral furrow shallow; mesolateral furrow weakly developed, almost indis- cernible. Mesosoma. Tergite colouration dark yellow to orange anteriorly, becoming lighter (yellow) posteriorly. Tergites I-III with a very shallow median depression divided anteriorly by a very weak ridge, tergites IV-VII with median carina reduced to a very weak ridge surrounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae absent on tergites I-VI, but present in posterior 2/3 of tergite VII, developed as ridges bearing strong spiniform granules. Surface of tergites shagreened, finely and densely granular; pre-tergites smooth. Sternite colouration dark yellow. Sternites III-VI smooth, slightly granular laterally, without carinae; sternite III with a pair of large, very finely and densely granular depressions covered by the pectines; sternites IV-VI with a pair of shallow median furrows. Sternite VII shagreened, finely and densely granular; pair of lateral carinae developed as granular ridges; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII longer than wide. Metasoma (Fig. 23B). Relatively elongated and slender, shagreened, with very few scattered small granules. Colour dark yellow to orange, carinae darker. Segments I-IV with longitudinal dorsomedian furrow and with dorsal carinae composed of strong spiniform granules. Segments I-IV with lateral carinae developed as ridges bearing sparse spiniform granules, lateral carinae on segments III-IV less granular than those on segments I-II. Segments I-II with ventrolateral carinae developed as ridges with weak granules; segments III-IV with ventrolateral carinae as ridges of spiniform granules. Segment I-II with ventral carina a ridge with few very weak granules, almost smooth; segments III-IV with ventral carina a ridge with more distinct spiniform granules. Segment V: longitudinal dorsal furrow present in anterior half, dorsal carinae with strong spiniform granules; lateral carina vestigial, only visible in anterior 2/3 as a row of small scattered granules; ventrolateral and ventromedian carinae with spiniform granules. Telson (Fig. 23B). Vesicle yellow to orange; aculeus darker, tan, due to stronger sclerotisation; vesicle elliptical or ovate, without blunt tuberculiform projections at base of aculeus; ventrolateral furrows and ventromedian ridge absent; lateral surfaces smooth, with sparse weak granules; dorsal surface with very small spiniform granules and a median shallow depression anteriorly. Macrosetae very sparse basally, becoming more numerous near base of aculeus. Aculeus short and stout, strongly curved, becoming markedly narrower approximately midway. Chelicerae. Colour dark yellow; teeth of fingers darker, orange. Teeth arrange- ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth HEMISCORPIDAE FROM IRAN 919 bifid; movable finger with one subdistal tooth and one basal tooth in external series; distal external tooth smaller than distal internal tooth; cheliceral teeth without secondary serrations. Pedipalp relatively elongated and slender. Pedipalp coxa and femur (Fig. 24D- F). Internoventral margin of coxa with few strong spiniform granules. Dorsal surface of femur predominantly dark yellow-orange, carinae darker, orange. Femur relatively short and stout (length less than 2.5 times its width), pentacarinate, with 4 distinct carinae; internodorsal carina developed as a strongly and densely granular ridge, spini- form granules strong; externodorsal carina developed as a ridge with spiniform granules; internoventral carina as a granular ridge with strong spiniform granules; externoventral carina as a ridge with spiniform granules; ventromedian carina reduced to few spiniform granules and confined to base of femur. Dorsal surface shagreened, finely and densely granular, distal end smooth, without granules; internal surface shagreened, with a few strong scattered spiniform granules; external surface shagreened; ventral surface shagreened and weakly granular, distal end without granules. A total of 3 trichobothria present on femur; d located externobasally on dorsal surface; i located dorsobasally on internal surface; e located dorsobasally on external surface. Pedipalp patella (Fig. 24D-F) relatively stout (length less than 2.5 times its width). Colour predominantly dark yellow to orange; carinae slightly darker. 7 carinae present, 6 of them distinct; internodorsal carina developed as a granular ridge; externo- dorsal carina as a ridge of indistinct granules; internoventral carina as a strongly gran- ular ridge with spiniform granules; externoventral carina as a ridge of weak granules; externomedian carina as a weakly granular ridge. Dorsal surface shagreened, sparsely and weakly granular; internal and external surfaces shagreened; ventral surface shagreened, sparsely and weakly granular, distal end without granules. Internal protu- berance pronounced, bifid (internodorsal and internoventral tubercles separated), internodorsal carina with 2 strong spiniform granules. A total of 19 trichobothria present on patella, orthobothriotaxy; d, located basally, external to internodorsal carina; d, located midway on patella; d3 absent; i in distal half of patella, located dorsally on internal surface. External (e) trichobothrial groups discernible and orthobothriotaxic: eb group composed of 5 trichobothria, esb group of 2 trichobothria, em group of 2 trichobothria, et group of 3 trichobothria; est midway between em to et groups. Ventral (v) group of 3 trichobothria (orthobothriotaxic). Pedipalp chela manus (Fig. 24A-C). Colour predominantly orange to red-brown; carinae reddish brown to almost black. Chela relatively stout (length less than 2.5 times its width) with 5 distinct carinae; internodorsal carina continuous, with spiniform granules, stronger basally; subdigital carina vestigial, visible basally as a smooth ridge with weak granules; externodorsal carina distinct, visible as a ridge of indistinct fused granules, almost smooth; digital carina as a smooth ridge with few weak granules basally; dorsal secondary carina (dorsomedian) as a smooth ridge with very few strong granules basally and few weak granules distally; internoventral carina continuous, visible as a weak ridge with very weak granules; externoventral carina continuous, developed as a granular ridge, becoming almost smooth distally, running parallel to longitudinal axis of chela, its distal edge disconnected from external movable finger condyle and direc- ted between external and internal movable finger condyles; ventromedian carina 920 L. MONOD & W. R. LOURENCO vestigial, reduced to few coarse granules basally; internal (internomedian) carina vestigial, reduced to few weak spiniform granules in distal half; external (externome- dian) carina indistinct, visible as raised rows of granules. Dorsal surface shagreened and sparsely granular; internal surface shagreened, sparsely granular, with a denser patch of bigger spiniform granules dorsodistally; external surface shagreened, sparse- ly and weakly granular; ventral surface shagreened and sparsely granular, distal end smooth and pitted. A total of 15 trichobothria present on chela manus; Db trichobothria located externobasally on dorsal surface; Eb group (3 trichobothria) orthobothriotaxic, located basally on external surface; Esb very close to Eb group; Em absent; Est located distally, very close to Et group: Et group composed of 5 trichobothria, Er, located ven- trally; V group comprising 4 trichobothria, V3 and V4 located in basal third of manus, VI and V2 located very far distally. Pedipalp chela fingers (Fig. 24A-C). Basally red- dish tan, becoming gradually lighter distally, tips of fingers yellow, edges black. Dorsal and ventral surfaces predominantly smooth and pitted, slightly shagreened basally. Fingers slightly shorter than chela manus. Fixed finger with weak basal concavity, movable finger with corresponding lobe weakly developed. Tips of fingers with pro- nounced terminal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of denticles becoming fused basally, interrupted at regular intervals by stronger granules, each of them coupled with an accessory granule; rows fused into a single row above concavity on fixed finger and on notch of movable finger, single basal row run- ning towards base of fingers. A total of 11 trichobothria present on chelal fingers; Dt located very basally on dorsal surface; db located dorsally on internal surface, in the basal half of fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb located dorsally on external surface, in basal 1/3 of fixed finger, opposite db; esb, est and et in distal half of fixed finger, opposite dsb, dst and dr, respectively; it and ib located in median 1/3 of fixed finger. Coxosternal sclerites. Dark yellow to orange; smooth, with shagreened patches. Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, penta- gonal, slightly shagreened; median furrow deep, more pronounced in posterior half; posterior pit absent. Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal surfaces of trochanter and femur finely and weakly granular. Tarsus with a ventro- median row of microspinules and with 2 rows of ventral rigid “spinoid” macrosetae; tarsi I-II: prolateral row with 4/5 macrosetae, retrolateral row with 5/6 macrosetae; tarsi I-IV: prolateral row with 5/6 macrosetae, retrolateral row with 6 macrosetae; setae acuminate; tarsal claws of equal length. Pectines and genital operculum (Fig. 25). Dark yellow; genital operculum composed of 2 subtriangular plates; genital papillae short, not protruding from beneath operculum; pectines with 13/12 teeth. Hemispermatophore. The presence of fully developed paraxial organs was assessed by cutting a small slit into the lateral pleural membrane between tergites and sternites. We did not have the permission to dissect the specimen. Intraspecific variation. The other specimen examined, a subadult male, has pectines with 15-15 teeth. HEMISCORPIIDAE FROM IRAN 921 Remarks. Apparently more male and female specimens of H. persicus from the type locality are lodged in the collections of the Zoological Institute of Saint Petersburg (Fet, pers. comm.) but unfortunately we could not get this material on loan. DISCUSSION TAXONOMIC CHARACTERS Sexual dimorphism. Apart from H. persicus, all Iranian species of the genus Hemiscorpius show a pronounced sexual dimorphism: mature males have a distinctly more elongated metasoma and telson than females (subadult males do not exhibit this sexual dimorphism) (Figs 4, 27). In H. acanthocercus sp. n., H. enischnochela sp. n., H. lepturus and probably also in H. gaillardi this dimorphism reaches extreme levels. Among scorpions extreme elongation of the metasoma in males is fairly common. It is encounter in the buthid genera Centruroides, Compsobuthus and Isometrus, as well as in genera belonging to families that are closely related to Hemiscorpiidae, i. e. Hadogenes Kraepelin, 1894 (Liochelidae), Heteroscorpion Birula, 1903 (Hetero- scorpionidae) and Urodacus Peters, 1862 (Urodacidae). Hadogenes spp. are ultralithophilous scorpions occurring in the Afrotropical region. They inhabit weathered rock outcrops where their extremely flattened mor- phology allows them to creep into narrow crevices and fissures. The very long tails of the males of most Hadogenes species are an essential adaptation to this specialized en- vironment. During courtship the male usually stings the female in the pleural mem- brane between the sternites and tergites. This behaviour is known as sexual sting. In very narrow cracks where courtship usually takes place, the male must reach around the side of the female’s body to sting her (Leeming, 2003). The endemic Malagasy genus Heteroscorpion is closely related to the Urodacidae. As in Hadogenes, the four known Heteroscorpion species are well adapted to live in rocky habitats and occur in narrow fissures between layers of splintered but still standing rocks. The metasoma of males is also very elongated compared to females (Lourenco & Goodman, 2002, 2004; Lourenco et al., 2004). However, these scorpions do not show an extreme dorsoventral depression of the body as in Hadogenes. On the other hand, scorpions of the genus Urodacus are mostly burrowers, therefore they are usually very stocky. However, males of four species, i. e. U. elon- gatus L.E. Koch, 1977, U. megamastigus L.E. Koch, 1977, U. mckenziei Volschenk, Smith & Harvey, 2000 and U. varians Glauert, 1963, show an extreme elongation of their metasoma (L. E. Koch, 1977; Volschenk et al., 2000) but they are not strictly lithophilous. U. varians is a burrowing species and the others excavate shallow scrapes under stones and are sometimes found in rock crevices (Volschenk, pers. com.). Ecomorphological adaptations to rocky habitats also occur in the scorpionid genus Opisthophthalmus C.L. Koch, 1837, which is mostly composed of burrowing species (Prendini, 2001). Because of soil hardness in their habitats, O. austerus Karsch, 1879, O. karooensis Purcell, 1898 and O. pattisoni Purcell, 1899 have abandoned burrow construction in favour of a semi-litophilous existence under slabs of rock. All these species display morphological adaptations similar to those in Hadogenes, 1. e. dorsoventral depression of the body and greatly elongated pedipalp chelae. L. MONOD & W. R. LOURENCO 922 "UU | ‘our 9[89S ‘odAJojou afewaf “I :1p40]]108 *H ‘(94 MWHN) >dAyered seuss ‘gq :>dAjojoy apew Sy su ‘ds pfoysouyasmua snidsoosnuay ‘yoodse jesiop ‘ooedereo “dds snidsoosnuayy 9¢ “SIA Di 1/27 HEMISCORPIIDAE FROM IRAN 923 Fic. 27 Hemiscorpius spp., metasoma, lateral aspect. Hemiscorpius maindroni: A, female syntype; B, male syntype. Hemiscorpius enischnochela sp. n.: C, female paratype (MNHG, Masdyed- Soleyman); D, male holotype. Hemiscorpius lepturus: E, female (MNHN, Ahwaz); F, male (MNHN-RS 5232). Scale lines, 2 mm. 924 L. MONOD & W. R. LOURENCO Fig. 28 Rhopalurus agamemnon (C. L. Koch, 1839), male, ventral view of central part of body with left pecten removed showing the granular depressed region of the third sternite III (from Lourengo & Cloudsley-Thompson, 1995). There is no information on the labels of the specimens studied referring to col- lecting methods, ecology, or habitat. However, the rugged, mountainous topography of their environment and their morphology, i. e. strongly elongated metasoma and dorsoventral depression of the body, indicate that Iranian Hemiscorpiidae are lithophilous species. Lithophilous scorpions adapted to live in narrow cracks and crevices of rocks are characterized by the following features: (1) dorsoventral depres- sion of the body; (2) elongation of the metasoma and/or pedipalps; (3) spiniform macrosetae on the telotarsi, operating in conjunction with highly curved telotarsal ungues to provide grip on rock surfaces (Prendini, 2001). All these adaptations are present in hemiscorpiids. HEMISCORPIIDAE FROM IRAN 925 Fic. 29 Hemiscorpius maindroni, male syntype (MNHN-RS 4328), hemispermatophore: A, hemisper- matophore in toto; B, detail of the capsular region, internal aspect. Scale lines, 0.25 mm. L. MONOD & W. R. LOURENCO A | ee OT ATV RN 926 Fic. 30 Jomachus politus, hemispermatophores of two males, external aspect: A, male from German East Africa, now TANZANIA (NHMW 1733); B, male from Handeni, Tanganyika, TANZANIA (CAS). Scale line, 1 mm. HEMISCORPIIDAE FROM IRAN 927 + Fic. 31 Jomachus politus, hemispermatophores of two male, detail of the capsular region, internal aspect, h (hook), La (lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (pos- terior lobe), Tr (transverse ridge): A, male from Handeni, Tanganyika, TANZANIA (CAS); B-D, male from German East Africa, now TANZANIA (NHMW 1733). Scale line, 0.5 mm. 928 L. MONOD & W. R. LOURENCO FIG. 32 Opisthacanthus lecomtei, male (MNHN-RS 6264, Belinga, GABON), hemispermatophore: A, hemispermatophore in toto (scale line, 1 mm); B, detail of the capsular region, internal aspect (scale line, 0.5 mm). HEMISCORPIIDAE FROM IRAN 929 FIG:53 Opisthacanthus cayaporum Vellard, 1932, male (MHNG, Campos de Cayapos, BRASIL), hemispermatophore: A, hemispermatophore in toto; B, detail of the capsular region, internal aspect. Scale lines, 1 mm. 930 L. MONOD & W. R. LOURENCO = DIS E CINZIA SUNT. ) J P Y 46 jj 4 HIT, 4°; ea, 47 Fic. 34 Hadogenes troglodytes (Peters, 1861b), male (NHMW 1703, South Rhodesia, now ZIMBABWE), hemispermatophore: A, hemispermatophore with in toto; B, detail of the capsular region, internal aspect, Dh (double hook), La (lamella), Lb (basal lobe), Ld (distal lobe), Lp (posterior lobe), Tr (transverse ridge). Scale line, 1 mm. «e. HEMISCORPIIDAE FROM IRAN 931 Abu), NL TUE S AR ee ALU [ | ij G \ | es [NE \ IN ı Fic. 35 Liocheles australasiae (Fabricius, 1775), male (BPBM, Tanna Island, VANUATU), hemisper- matophore: A, hemispermatophore in toto; B, detail of the capsular region, internal aspect, h (hook), La (lamella), Lb (basal lobe), Ld (distal lobe), Lp (posterior lobe), Tr (transverse ridge). Scale lines, 0.5 mm. In male hemiscorpiids the genital operculum is composed of two overlapping plates, whereas in females it is a single undivided plate. Like in other scorpions, males have bigger pectines with a higher pectinal teeth count than in females. In the A. lep- furus species group the carapace is usually more elongated in males (longer than wide) 932 L. MONOD & W. R. LOURENCO than in females (almost as wide as long) (Fig. 26). Likewise, tergites and sternites VII are longer than wide in males, while as wide as long in females. Like the dimorphism in metasoma length, this difference is present only in the final instar. Immature males and females look much the same and can usually be distinguished only through exam- ination of their genital operculum and pectines. Granular depressions on sternites III. In males and females of Hemiscorpius acanthocercus sp. n., H. enischnochela sp. n., H. gaillardi, H. lepturus and H. persi- cus, the foremost sternite exhibits a pair of large, rough, very finely granular depres- sions. This feature is morphologically very similar to the stridulatory apparatus of species belonging to the neotropical buthid genus Rhopalurus (Fig. 28). The Rhopalurus stridulatory apparatus consists of large granular areas on the third sternite, which are located under the pectines and are used as a rasp. Pectinal teeth with stria- tions on their dorsal surface are rubbed against these areas in order to produce sound. Stridulation have been reported for all Rhopalurus species (in both males and females), but only the stridulatory sounds of larger species are audible to humans (Lourenço & Cloudsley-Thompson, 1995; Lourengo, Huber & Cloudsley-Thompson, 2000). Production of stridulatory sound has never been reported for any Hemiscorpius species, but this can be attributed to the small or medium size of these scorpions (regardless of the length of the metasoma), which probably produce barely audible sounds. The dorsal surfaces of their pectinal teeth seem to be finely granular. However, this character could not be assessed accurately because of the very reduced size of the structures. More thorough investigations by means of a stereo-electron microscope are needed. Hemispermatophores. The hemispermatophores of Hemiscorpiidae (Figs 7, 20, 21, 29) share morphological similarities with hemispermatophores of Hadogenes species (Fig. 34) and of a liochelid group composed of Jomachus politus (Fig. 30-31) and Opisthacanthus (Opisthacanthus) species (Figs 32, 33). The general shape of hemispermatophores is the same in these taxa, 1. e. distal lamella longer than the basal part, lamellar hook situated above the distal transverse ridge and not merging with it. However, Hemiscorpiidae have hemispermatophores with the distal lamella bearing a double hook, whereas all the other liochelid genera except Hadogenes have distal lamellae with a single hook. The capsular median structure of hemiscopiid hemispermatophores is complex (Figs 7B-C, 21A-C, 29B), composed of 4 distinct lobes/lamellae: (1) capsule lamella (La) complex, bearing a strong hook on his external surface and an accessory lobe (Lac) pointing to the posterior side, (2) distal lobe (Ld) forming a strong hook pointing anteriad, (3) basal lobe (Lb) very reduced, (4) posterior lobe (Lp) costate. Jomachus politus and Opisthacanthus (Opisthacanthus) species (both African and neotropical representatives) possess hemispermatophores (Figs 30-33) very similar to those encounter in hemiscorpiids, i. e. with distal lobe with a strong hook pointing anteriorly and with a complicated capsular lamella bearing an internal hook and an additional basal lobe. In Hadogenes (Fig. 34) and other liochelid genera the capsule lamella is much more simple, without any hook or accessory lobe. In Liocheles Sundevall, 1833 HEMISCORPIIDAE FROM IRAN 933 the distal lobe is not modified into a large hook and the basal lobe is much more de- veloped (Fig. 35). Based on this analysis Hadogenes and the group composed of /omachus politus and Opisthacanthus (Opisthacanthus) spp. are considered to be liochelid taxa closely related to hemiscorpiids. The double lamellar hook can therefore be considered as a plesiomorphy for Hemiscorpiidae and Liochelidae, and the single lamellar hook is apo- morphic for all Liochelidae except Hadogenes. On the other hand, the complex capsule lamella can also be considered as a plesiomorphy for Hemiscorpiidae and Liochelidae. However, the derived, more simple structure examined in Hadogenes and other liochelid genera (e. g., Liocheles) is assumed to have evolved independently in these two groups. BIOGEOGRAPHICAL AND GEOLOGICAL IMPLICATIONS Iomachus politus occurs in East Africa (Ethiopia, Kenya Tanzania, Uganda) and the only African representative of the subgenus Opisthacanthus, O. (O.) lecomtei (Lucas, 1858), was recorded from approximately the same latitude in West Africa (Cameroon, Gabon). The neotropical species of Opisthacanthus (O.) are found in Central and South America and Hadogenes is a South African genus. Hemiscorpius 1s present all over the Middle East (Iran, Iraq, Oman, Pakistan, Saudi Arabia, United Arab Emirates and Yemen), but has also some representatives in north-eastern Africa (Djibouti, Eritrea, Somalia), not far from the distribution area of Jomachus politus. The presence of Hemiscorpiidae in Djibouti, Eritrea, Somalia, and on the island of Socotra indicates that the ancestor of Hemiscorpius was probably present on Gondwanaland. Since Hemiscorpiidae is the sister group of Liochelidae and Hemiscorpius shares characters with certain African liochelids, i. e. Jomachus politus and Opisthacanthus (O.) lecomtei, the occurrence of Hemiscorpius in Africa is probably due to a Gondwanian origin and not a recent invasion from the Middle East (Laurasia). However, the Iranian Hemiscorpius, i. e. the H. lepturus species group, are very divergent from the Hemiscorpius species from Africa, Oman, Saudi Arabia, United Arab Emirates and Yemen. The African species of Hemiscorpius do not show the highly derived features that their Iranian relatives possess, i. e. extreme sexual dimor- phism and cytotoxic venom. Iranian Hemiscorpius have probably been separated from the other hemiscorpiids for quite a long time. H. persicus represents an exception among Iranian Hemiscorpius; this species is more similar morphologically to African and Arabian representatives of the genus than to the Iranian ones. This could be the result of a more recent invasion from the Arabian peninsula. An old separation of the Iranian hemiscorpiids can be explained by the geolo- gical history of the region. In the Precambrian and Palaeozoic the Cimmerian terrane, which included Iran, was part of Gondwana (Fig. 37, 38A). The common ancestor of Liochelidae and Hemiscorpiidae was probably widespread on this super-continent. Approximately 250 Ma BP (Fig. 38B) the Neotethys ocean started to open (Golonka, 2000; Scotese, 2000; Stampfli & Borel, 2002), inducing the rift of the Cimmerian super terrane off the northern margin of Gondwana and its northward drift toward Laurasia. Iran was separated from Gondwana in the Middle Triassic. The ancestor population of the Middle East hemiscorpiids was probably isolated on this rafting continent and diverged from its relatives. L. MONOD & W. R. LOURENCO 934 0) ME: mm = | 009 OSP 00€ OGI SZ 0 snaisiad snidioosiway} (aimesayy) snınyday snidioosiwaH snın]daj snıdiossiwaH BJ9y90Uyasıus SNILI09SILI9H SN918904}UE2E snidioosiway puaba] 3.0,0,09 3.0.0,09 3.0,0,0% FIG. 36 Distribution map of Hemiscorpius spp. In Iran and Iraq. 935 HEMISCORPIIDAE FROM IRAN ‘0007 ‘eYUOOH Wor POUIPOIN "wueMpuoH) Wo oURLEVEdNg URLOWTUNTD ay) Jo Suns oy) Suronpur ‘uodo 0) payes sktAOON UL “Surpeys A918 Aq poyeorpur ase sossewpur] “(qq EN 8PT-697) uelunag Se] OY SuLINP YEAH oy} JO UONINISUOIII orydeis0a3 00K [eg Le Old VIHABIS L. MONOD & W. R. LOURENCO 936 da VIN 022 (nnogiia) 040 x UBHON Ajsez ‘ds sniduoosiwapy (idol) Aalen OWE ‘ds sniduoosiwiay (eidoryyg) een 1061 ‘21089 Jura) snıdioaswuag (usWeA) 21090S 6681 ‘90904 SNUE11090$ SNIdIOISIWUIH da EN 057 Asepunogd 2ISSEI1] -UeIWI8d (uewo) ayeosey| (006, ‘uedee:y) ospwew Ssniduoosiwmay usway ‘Sa}21ILT paun "eigeiy Ipnes (6681 190904) snaiqese snidioosiway (suoi5a1 1940) sepiidsoosiwmay EN 087 uelued AE] ‘ueweuyes uoqe! 'Uoolsweg (8681 ‘seon7) 18141099 snyjueseysido eAu9y ‘BIUEZUE] 9681 ‘90004 snyyod snyaeuloy LA aepl|ayoory £06L ‘eng snaisuad sniduoosjway snunjdaj snıdıoosnuay (pZ6| ‘uouoe\) Ipseyeb sniduoosiway ‘u ‘ds ejay2ouyasiua snidioosiway "u ‘ds sna1s2oyjuese snidioasiusy © (NVHI) sepııdıossiuaH FIG. 38 geographic reconstructions of the Tethys Oceans and surrounding landmasses 280, 250 Palaeo and 220 Ma BP. The opening of the Neotethys separated the Cimmerian Superterrane from Gondwana inducing the Cimmerian Terrane to drift toward Eurasia. Its accretion to Laurasia 3 occurred 220 Ma BP when the oceanic crust of the Palaeotethys was completely subducted to the north. Landmasses are indicated by grey shading and current distribution areas of Hemiscorpiidae and related Liochelidae are indicated by pictograms. Positions of equator and tropics are also shown. Modified from Stampfli & Borel, 2002. HEMISCORPIIDAE FROM IRAN 937 The Hemiscorpiidae lineage was supposedly separated from the Liochelidae lineage before the rifting of the Cimmerian terrane, probably by climatic changes on Gondwana that occurred during the massive Permian global warming (290-250 Ma BP; Rees, 2002). Four main terrestrial climate zones (biomes) have been determined from fossil floras and climate-sensitive sedimentary rocks for the Late Carboniferous- Early Permian (300-280 Ma BP) (Gibbs er al., 2002; Rees et al., 1999; Rees et al., 2002; Scotese, 2000; Willis & McElwain, 2002). In both hemispheres, climates gradually changed from the equator to the poles as follow: (1) tropical everwet (equa- torial region), (2) subtropical desert, (3) cool temperate and (4) cold (polar region). At that time Gondwana was situated south of the equator and was therefore under mostly lush and green vegetation (cool temperate), except for its north-eastern part where arid conditions prevailed (subtropical desert). Throughout the Permian, the climate change from icehouse to hothouse conditions (Rees, 2002). With global warming and increasing aridity, tropical forests disappeared and cool temperate biotopes were segregated in the southern part of Gondwana as deserts spread across central Pangea; tropical everwet biomes were only well-developed on the Chinese microcontinent (Rees, 2002; Scotese, 2000). Therefore the hemiiscorpiid ancestor adapted to an envi- ronment (in Africa and Arabia) that became more and more arid during the Permian, while liochelids remained in the more humid and cool ecosystems prevailing in the southern part of Gondwana (Scotese, 2000; Willis & McElwain, 2002). By the Late Triassic-Early Jurassic (210-200 Ma BP), large subtropical deserts formed a climatic barrier (Rees, 2002; Rees et al., 1999) between the liochelid ancestor in warm tem- perate forests of Southern Gondwana and the African hemiscorpiid ancestor in the Equatorial summerwet biotopes of the Arabian region, and interrupted gene flow between these two lineages. Most of the modern scorpion families probably emerged at the end of the Palaeozoic Era, when changing biomes, high rates of extinction and the resulting isolation of surviving populations in scattered “biotopes islands” were favourable for diversification. After being detached from North and South America, Africa began to rotate counterclockwise toward Eurasia and closed the western part of the formerly extensive Neotethys sea (Brown & Lomolino, 1998; Golonka, 2000). A bridge between Asia and Africa was formed by Arabia, following their collision in the middle Tertiary (35 Ma BP) that created the Zagros mountains of Iran. Then hemiscorpiids could have easily established new colonies through dispersal from the Arabian peninsula to Eurasia. Another hypothesis is the drift of microcontinents from the northern margin of Gondwana towards the southern margin of Laurasia during the last 200 million years, scorpions could have been carried on them to Eurasia. ACKNOWLEDGEMENTS We express our sincere gratitude to all individuals and institutions that assisted in the realization of the present study: the Natural History Museum of Geneva for providing working space and access to the collection; Dr Peter Schwendinger for kindly reviewing the manuscript and for his extensive help, constructive comments and suggestions; Dr Jiirgen Gruber from the Natural History Museum of Vienna for 938 L. MONOD & W. R. LOURENCO arranging loans, providing information on collectors and help in deciphering old labels; Dr Victor Fet and Dr Graeme Lowe for kindly allowing the examination of the type specimens of Hemiscorpius persicus and for constructive criticism on the manuscript; Dr Erich Volschenk for further discussion; to M. Claude Ratton for taking the habitus photographs. REFERENCES ANDERSON, P. C. 1998. Missouri Brown Recluse Spider: a review and update. Missouri Medicine 95: 318-322. BIRULA, A. A. 1903. 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S., SMITH, G. T. & HARVEY, M. S. 2000. A new species of Urodacus from Western Australia, with additional descriptive notes for Urodacus megamastigus (Scorpiones). Records of the Western Australian Museum 20: 57-67. WALCKENAER, C. A. 1847. Araneae (pp. 365-564). In: WALCKENAER, C. A. & GERVAIS, P. (eds). Histoire naturelles des insectes apteres 4. Librairie encyclopédique de Roret, Paris. WILLIS, K. J. & Mc ELWAIN, J. C. 2002. The evolution of plants. Oxford University Press, Oxford. ao + ca Lay and palo hard, Maw! rah ad A eu A] LIT aies m = > REVUE SUISSE DE ZOOLOGIE 112 (4): 943-961; décembre 2005 New genera and species of nematode parasites (Drilonematoidea: Ungellidae) from coelomic cavity of Neotropic acanthodrilids deposited in the Natural History Museum of Geneva, Switzerland Elena S. IVANOVA! & Claude VAUCHER? ! Institute of Parasitology, Russian Academy of Science, 33 Leninskii pr., Moscow, 119071, Russia. 2Muséum d’histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland. New genera and species of nematode parasites (Drilonematoidea: Ungellidae) from coelomic cavity of Neotropic acanthodrilids deposited in the Natural History Museum of Geneva, Switzerland. - Two new genera and two new species of nematodes (Ungellidae, Drilonematoidea) parasitic in Neotropic acanthodrilids from the collection of the Natural History Museum of Geneva, Switzerland, are described. Yagansiella longi- collis gen. n., sp. n. and Ungella chileana sp. n. are parasitic in Yagansia spatulifera, whereas Ungella micronychium sp. n. in Y. diversicolor and Patagoniella capitoporus gen. n., sp. n. in Y. papillosa. Yagansiella longi- collis gen. n., sp. n. resembles Ungella in general morphology but differs by anterior position of ovary top cell and posterior vulva position, charac- teristic body and caudal fimbriate organs shape. U. chileana sp. n. is distin- guished by large circular fimbriate organs in tail, female body shape and arrangement of female genital tube and spicule shape. Ungella micro- nychium sp. n. differs by minute cephalic hooks and shape of fimbriate organs and spicules and gubernaculum. Patagoniella capitoporus gen. n., Sp. n. is distinguished by anteriormost position of excretory pore, anterior vulva position and shape of fimbriate organs and spicules and guberna- culum. Keywords: Earthworms, nematode parasites - Ungellidae - Drilonema- toidea - Yagansiella longicollis gen. n., sp. n. - Ungella chileana sp. n. - Ungella micronychium sp. n - Patagoniella capitoporus gen. n., sp. n. - Yagansia spatulifera - Yagansia diversicolor - Yagansia papillosa. INTRODUCTION Earthworm collection deposited in the Natural History Museum of Geneva contains European lumbricids as well as annelids from around the globe. From 1 to 6 specimens of 142 species of earthworms from museum collection were studied on the presence of nematodes (Drilonematoidea, Rhabditida) parasitic in coelomic cavity of earthworms. The bulk of museum collection consists of Neotropic glossoscolecids and acanthodrilids. So far, the only drilonematid parasite had been recorded from these Manuscript accepted 07.04.2005 944 S. IVANOVA & C. VAUCHER earthworms. A new genus and the new Ungella species from coelomic cavity of Yagansia spatulifera, the new Ungella species from Y. diversicolor and the new genus from Y. papillosa all belonged to Ungellidae family are described below. MATERIAL AND METHODS Earthworms were stored in ethanol after formaldehyde fixation which found did not affect the nematode morphology. Earthworms were dissected at anterior end and body cavity content was rinsed out and nematode parasites removed. Four specimens of Yagansia spatulifera, six ones of Y. diversicolor and three of Y. papillosa were dissected. Three females and three males of Yagansiella longicollis gen. n., sp. n. and a male and a female of Ungella chileana sp. n. were obtained from coelomic cavity of single specimen of Yagansia sparulifera. Two males of U. micronychium sp. n. were recovered from a specimen of Y. diversicolor and nine males, five females, three juve- niles and fragments of Patagoniella capitoporum gen. n., sp. n. from a specimen of Y. papillosa. Nematodes were processed into glycerol using slow evaporation technique (Seinhorst, 1959) and mounted on slides. Drawings and measurements were made using “Jenaval” microscope. De Manian indices and absolute measurements are given, where D is maximal diameter (if not indicated), Oes is oesophagus length, Cd is tail length and Ex is distance from base of cephalic hooks to excretory pore. A name “fimbriate organs” was used when describing caudal structures of sensory nature (phasmids?) instead of “suckers” (Ivanova & Hope, 2004). DESCRIPTIONS Yagansiella gen. n. DIAGNOSIS. Rhabditida, Drilonematoidea, Ungellidae. Very long and thin neck region and swollen body. Spacious pseudocoel. Paired blade-like cephalic hooks present. Cuticular ridges situated posteriorly to mouth. Amphids elliptical. Stoma lacking; oesophagus with corpus, isthmus and bulb. Nerve ring situated around isthmus. Excretory pore at isthmus level. Large paired deep transversely oriented fimbriate organs in caudal region with prominent lip-like basal rim. Females mono- delphic, prodelphic, spermatheca set-off, distal tip of ovary near oesophageal base, vulva posterior with oblique vagina. Males monorchic; spicules paired and curved; thick gubernaculum with dorso-caudal apophysis and crurae, bursa absent. TYPE AND ONLY SPECIES. Yagansiella longicollis sp. n. ETYMOLOGY. Generic name refers to annelid host of the species. Yagansiella longicollis sp. n. Figs 1-2 MEASUREMENTS. Holotype female. L = 1216 um; D = 153 um; Oes = 213 um; Ex = 129 um; NR = 105 um; Cd = 123 um; Ovum = 54 x 27 um; V% = 89 44%; a= 7295 AES — 9:89: Paratype females (n= 2). L= 1185-1458 um; D = 108-140 um; Oes = 215 um; Ex = 131-135 um; NR = 81-90 wm; Ova = 52-54 x 25-27 um; Cd = 112-127 um; V% = 90.55-90.81; a = 10.41-10.97; b = 5.51-6.78; c = 10.58-11.48. NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 945 FIG. 1 Yagansiella longicollis gen. n., sp. n. Female. A — paratype, entire body; B — paratype, head end; C — holotype, anterior end; D — holotype, posterior end; E — holotype, spermatheca region. All in lateral position. Scale bars in um. Paratype males (n = 3). L = 1093.67+142.1 (930-1185) um; D = 83+25.2 (60- 110) um; Oes = 188.33+9.5 (179-198) um; Ex = 118.33+6.7 (114-126) um; NR = 99 .33+7.8 (93-108) um; Cd = 200.67+8.1 (195-210) um; Sp (arc)= 104.33+5.5 (99- 110) um; Sp (chord)= 6745.6 (62-73) um; Gub = 30.33+2.1 (28-32) um; a = 14.32+5.7 (8.45-19.75); b = 5.8+0.6 (5.2-6.3); c = 5.47+0.9 (4.43-6.02). 946 S. IVANOVA & C. VAUCHER DESCRIPTION. Adults. Very long and thin neck region and swollen body. Body starts to widen gradually behind the base of esophagus and turns uniformly wide from the point of where genital tube forms flexure. Terminal portion of tail filamentous. Epidermis thick. Spacious pseudocoel. Cuticle 1 thick, annulated at both ends, with annules 1.5 um thick. Head bluntly rounded. No cephalic sensilla present. Head with paired, independently-moveable, submedian, claw-like cephalic hooks situated on anterior surface of head; base of each hook consists from pair of thick equal pointed diverged processes 2-3 um long; longer pointed blade directed dorsad. Minute oral aperture shifted slightly dorsad near distal tips of blades. Stoma absent. Cuticular ridges on dorsal side just beneath mouth. Amphid faint, situated close to hooks base; its aperture transversely elongated. Anteriormost portion of esophagus slightly expanded. Slightest constriction of corpus at 10-12 um from head end. Oesophagus slender, muscular, from very long corpus 6-8 um wide, thinner long isthmus and bulb with granular secretion of dorsal oesophageal gland. Bulb elongated with nearly rectangular base, 30-40 um long and 10-11 um wide. One large nucleus of dorsal gland at posterior of bulb and two smaller ones of subventral glands situated more anteriorly. Nerve ring wide, encircling anterior part of isthmus. Excretory pore in 20-40 um behind nerve ring, 2 um wide. Excretory duct strongly cuticularized, 2 um wide and 105-153 um long. Excretory cell large, with granular content, observed untill mid- body. Cardia from two elongated cells. Intestine thick-walled, contains debris. Large paired symmetrically placed deep transversely oriented fimbriate organs in caudal region with prominent lip-like basal rim. Females. Blades of head hooks 7.67+0.6 (7-8) um long, hook base 5.33+1.2 (4- 6) um wide. Amphidial aperture 3 x 2 um. Oesophagus 9.33+0.6 (9-10) um wide at anterior, corpus 8 um and isthmus 6 um wide. Bulb 37+2.6 (35-40) um long and 10.67+0.6 (10-11) um wide. Genital tube starts in 150-207 um behind the oesophageal base, sometimes extending prior to spermatheca, which situated in 388.33+30.4 (363- 422) um from anterior. Tube thick, runs posteriorly to fimbriate organs, then turns back by convoluted course and forms rounded or elongated off-set spermatheca 74.67+17.5 (60-94) x 50+26.5 (30-80) um with thick walls filled with spherical or bean-like sper- matozoa 1.5-2 um in diameter. Descending and ascending branches of genital tube spirally twisted. Spermatheca joined with thick-walled oviduct by thin tube. Thin- walled muscular uterus obscured by twists of ovary. Muscular vagina 36.33+4 (32-40) um long. Vulva at posterior. No post-uterine sack present. Anus in 43.33+7.6 (35-50) um posteriorly to vulva. Up to 6 eggs with thin finely punctated shells. Fimbriate organs with deep chamber, 39+5.3 (35-45) um long and 6645.3 (60-70) um wide with prominent lip-like muscular rim 5-7 um thick. A channel inside tail end runs through fimbriate organs. Terminal portion of tail 50.67+4 (47-55) long and 5.33+.6 (5-6) um wide. Males. Resembles females in body shape but a little shorter and less pro- minently expanded behind neck region. Amphidial aperture 4.67+1 .2 (4-6) x 2.33+0.6 (2-3) um. Testis reflexes at 333 .67494.2 (255-430) um from oesophageal base, flexure curved, 180.67+19.9 (159-198) um long. Developing spermatocytes in 7-8 rows, small, spherical (2 um in diameter). Testis very wide, ejaculatory duct muscular, sepa- rated from vas deferens by constriction. Spermia in vas deferens spherical or bean-like, NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 947 FIG. 2 Yagansiella longicollis gen. n., sp. n. Male. A — paratype, entire body; B — paratype, head end; C — paratype, anterior end; D — paratype, spicules and gubernaculum; E — paratype, tail. All in lateral position. Scale bars in um. resemble those from female spermatheca. Very long strongly cuticularized sickle-shaped paired spicules, with elongated manubria 12.33+1.2 (11-13) wm long and 10.33+1.5 (9-12) um wide. Spicule shafts 5.66+0.6 (5-6) um wide at mid-length tapering gradually to rounded tips and bearing two front ridges. Gubernaculum massive, dark-brown in colour, with dorso-caudal apophysis 12.33+3.1 (9-15) um long and crurae as long as gubernaculum body. A slit in gubernaculum body parallel to spicules. Nine pairs of subventral precloacal and four (five) pairs of postcloacal papilli- form sensilla. First pair of postcloacal sensilla subventral, situated between anus and 948 S. IVANOVA & C. VAUCHER fimbriate organ; second one dorsal in front of fimbriate organ; third subventral between fimbriate organ and terminal portion of tail; fourth subdorsal close to the latter and fifth lateral symmetrically to the fourth in a paratype specimen. Copulatory muscles pro- minent but bursa lacking. Fimbriate organs smaller than in females: 24.67+3.1 (22-28) um long and 22.33+2.3 (21-25) um wide with basal rim 4-5 um thick. Thick channel ends blindly posterior to fimbriate organ. Filamentous portion of tail 7143.6 (68-75) um long and 4 um thick. TYPE HOST AND LOCALITY. Yagansia spatulifera (Mich.) AF 4262, Chile, Villarica Vulcan, 1200 m, 14.02.01, coll. and identified by C. Czudi. TYPE HABITAT. Coelomic cavity at mid-body region. TYPE SPECIMENS. Holotype female MHNG 35445 INVE and paratype female and two paratype males MHNG 35446 INVE deposited in Natural History Museum (Geneva, Switzerland). DIFFERENTIAL DIAGNOSIS. The present species is closely related to Ungella Cobb, 1928. From the latter genus, it can be distinguished by position of extremity of ovary which is anterior in Yagansiella and posterior in Ungella and vulva position which places in front of anus whereas it is typically post-median. It differs from Ungella as well by having swollen body posterior and large and broad fimbriate organs of different shape. From Plesioungella Yeates et al., 1998, which it resembles by swollen posterior, it differs by having paired spicules. From Onychonema Baylis, 1943 it differs by presence of prominent fimbriate organs which lacked in the former. From Acanthungella (Ivanova & Hope, 2004) it differs in lack of somatic sensilla. ETYMOLOGY. The species name derived from Latin words longus (long) and collum (neck) and reflects appearance of the nematodes. Ungella chileana sp. n. Figs 3-4 MEASUREMENTS. Holotype male. L = 1205 um; D = 48 um; Oes = 182 um; Ex = 149 um; NR = 105 um; Cd = 183 um; Sp (arc) = 70 um; Sp (chord) = 51 um; Gub HS mA — 25Hle bh) —16)03; € = 10,58. Paratype female. L= 1218 um; D = 90 um; Oes = 215 um; Ex = 170 um; NR = 130 um; Cd = 163 um; V% = 64.5; a= 13.53; b = 5.67; c = 7.42. DESCRIPTION. Adults. Anterior of body thin. Terminal portion of tail filamentous. Cuticle 1 um thick, annulated. Epidermis well-developed. Head truncate, with paired, independently-moveable, submedian cephalic hooks situated on anterior surface of head. Each hook from thick bone head-like base and pointed outwardly-curved blade and directed dorsad. Mouth shifted slightly dorsad near distal tips of blades. Stoma absent. Amphid pocket-like, situated close to hooks base; amphidial aperture trans- versely elongated. Anteriormost portion of esophagus slightly expanded. No head sensilla visible. Oesophagus straight, slender, from long finely muscular corpus, no distinct isthmus and narrow bulb with 3 nuclei of oesophageal glands. Base of oesophagus not rounded. Nerve ring situated a little farther back from mid-oesophagus. Excretory pore opposite anterior of bulb, 1.5 um thick. Excretory duct 1.5-2 um wide 90-100 um long. Excretory cell with fine granular content visible till mid-body. Cardia prominent from two large oval cells. Intestine thin. NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 949 Fic. 3 Ungella chileana sp. n. Male holotype. A - entire body, lateral view; B - head end, apical view; C - tail, lateral view; D - spicules and gubernaculum, lateral view. Scale bars in pm. 950 S. IVANOVA & C. VAUCHER Male. Body slender, tapering to both ends and slightly expanded in testis region. Cephalic hooks with blades 8 um long and base 5 um wide. Amphidial aperture 6 x 2 um. Oeasophageal corpus 9 ym wide at anterior, isthmus 8 um wide, bulb 13 um wide, 32 um long. Intestine collapsed. Testis reflexed in 298 um from anterior, flexure 95 um long. Developing spermatocytes numerous, in 4, then 8 rows, spherical, 1.5-2 um in diameter. Vas deferens separated from ejaculatory duct by constriction. Spermia spherical, 1-1.5 um in diameter. Spicular muscles and levator ani muscle prominent. Copulatory muscles present preanally. Bursa absent. Spicules paired, light brown in colour, cephalated, highly sclerotized, bent at one third length from distal part. Manu- bria elongated, 7 um wide and 14 um long, shafts distally 6 um wide, then gradually tapering to tips which curved anteriad and bluntly rounded, 2 um thick. Gubernaculum darker than spicules, massive, parallel to spicules, with crurae 13 um wide embracing spicules and thick dorso-caudal apophysis 16 um long. Four pairs of preanal papilliform ventral sensilla. Two (?) pairs of postanal lateral papilla around left-side fimbriate organ. Fimbriate organs asymmetrically disposed, large, circular, rised, with fibrous basal rim 2 um thick, internal chamber covered by transparent membrane 11 um and aperture 4 um in diameter. Filamentous portion of tail 46 um long and 3 um thick. Female. Body stout, very slightly tapering posterior to vulva. Tail abruptly changes into bluntly rounded terminus 8 um thick 98 um long. Cephalic hooks with blades 10 um long and base 7 um wide. Minute oral aperture 1 um wide. Amphidial aperture oval, 4 x 2 um, pouch 4 um wide 5 um long. Corpus 10 um wide, isthmus 8 um wide and bulb 14 um wide 37 um long very slightly displaced dorsally. Cuticu- larized meandering channel 5 um thick observed subventrally in vulva region running through gland with numerous nuclei (excretory gland?). Distal tip of ovary in 110 um posterior to vulva. Ovary runs back nearly to end of expanded part of tail, then returns to the same distance, then runs back to start and turns to anterior where reflexes in 408 um from oesophageal base and forms spermatheca. Spermatheca 95 um long 35 um wide packed with spherical spermatozoa resembled those from male testis. Thick- walled uterus with two smooth-shelled eggs 60 x 25 um. Vagina straight, muscular, 38 um long, with slightly protruded lips. Vulva post-median. Anus in 15 um before right- side fimbriate organ. Fimbriate organs situated slightly asymmetrically near extremity of expanded part of tail (distance between centres of these 20 um), circular, raised, with basal rim 7 um thick 34 um in diameter and aperture 23 um in diameter covered with transparent membrane. Channel 4 um thick runs through fimbriate organs. TYPE HOST AND LOCALITY. Yagansia spatulifera (Mich.) AF 4262, Chile, Villarica Vulcan, 1200 m, 14.02.01, coll. and identified by C. Czudi. TYPE HABITAT. Coelomic cavity at mid-body region. TYPE SPECIMENS. Holotype male MHNG 35447 and paratype female MHNG 35448 deposited in Natural History Museum (Geneva, Switzerland). DIFFERENTIAL DIAGNOSIS. The genus Ungella comprises 7 species: U. secta Cobb, 1928, U. astrida Spiridonov, 2001, U. burmensis Ivanova & Hope, 2000, U. ituriensis Spiridonov, 2001, U. kivuensis Spiridonov, 2001, U. mexicana Ivanova & Hope, 2000 and U. sucofera Timm, 1962. Ungella chileana sp.n. is characterised by having swollen posterior of females and filamentous portion of tail; not-embedded cephalic hooks with characteristic head 3.0 mm) has only 2 hyaline spots in r2+3, remaining pattern as in other specimens. ABDOMEN MALE. Strongly reddish-brown, subshining; basal two tergites mainly white setulose, other tergites covered with short black setulae. Epandrium reddish; structure not studied. ABDOMEN FEMALE. Preabdomen as male; oviscape black, shining, pale brown setulose, about as long as length of 2.5 preceding tergites combined; eversible mem- brane with numerous denticles; aculeus evenly narrowed, apically sharply pointed. Distribution. Eastern Afrotropical Region (Kenya, Tanzania) and Arabian Peninsula (Saudi Arabia, Yemen). Host plants. Unknown. The specimens from the Arabian Peninsula were col- lected in Malaise traps. Pest status. Not a pest species. 998 B. MERZ & H. A. DAWAH Remarks. The generic position of this species may be subject to discussions. In the key of Munro (1947) it runs with some problems to Platensina Enderlein because of the rather thick, ash-grey microtrichose mesonotum and the chaetotaxy (couplet 21 in Munro, 1947). However, some species of Dicheniotes Munro, such as D. polyspila (Bezzi), have also an ash-grey microtrichose mesonotum and a very similar type of wing pattern and head shape (some 30 specimens of both sexes from Kenya, along road from Nairobi to Mombasa between Hunters Lodge and Mtito Andei studied, MHNG). Its inclusion in Dicheniotes (3 frontal setae, 2 orbital setae, 2 pairs of scutellar setae, postocular setae mixed white and black, wing of Tephrella type of Munro, 1947) seems therefore justified. Within Dicheniotes the new species differs from all congeners in wing pattern with 3 hyaline spots in cell rl, of which the apical spot is smaller than the two basal spots, but still clearly present, and the more numerous hyaline spots on the remaining wing surface. Because of the presence of a hyaline spot in r2+3 below tip of R2+3 D. multipunctatus runs to couplet 9 in Munro (1947, p. 178) next to D. polyspila which is morphologically similar with the same reddish abdomen and the thick, ash-grey microtrichose mesonotum and thus differing from the much more subshining mesono- tum and shining black abdomen of other species, such as D. angulicornis, D. erosa (Bezzi) or D. katonae (Bezzi). The new species differs from D. polyspila by the follo- wing set of characters in addition to those given above: wing with less conspicuous di- morphism (very prominent in D. polyspila: male with basal half of rl and r2+3, and cell br entirely hyaline, and r2+3 at tip with two broadly confluent hyaline spots; fe- male with more isolated spots), hyaline spots often not sharply separated from black area (usually very sharply separated in D. polyspila), and oviscape much shorter than preabdomen (in D. polyspila about as long as preabdomen). Gymnaciura austeni (Munro, 1935) Material. 2: 19. Distribution. Entire Afrotropical Region, Yemen (Merz et al., in press). New for Saudi Arabia. Host plants. Unknown. Pest status. Not a pest species. Metasphenisca hazelae (Munro, 1947) Material. 2:18,32. Distribution. Eastern Africa. New for Saudi Arabia. Host plant. Barleria rehmanni (Acanthaceae) (Munro, 1947). Pest status. Not a pest species. Remarks. Hancock (in litt.) found another specimen from Saudi Arabia in the unsorted material of the NHML: 1d , Dhamar, 3.IX.62 (or 42), G. Popov (NHML). Metasphenisca negeviana (Freidberg, 1974) Material. 2: 16. Distribution. Near East. Recorded from Saudi Arabia by Freidberg & Kugler (1989). FRUIT FLIES FROM SAUDI ARABIA 999 Host plant. Blepharis attenuata (Acanthaceae) (Freidberg & Kugler, 1989). Pest status. Not a pest species. Oxyaciura tibialis (Robineau-Desvoidy, 1830) Material. 2: 299. Distribution. Western Palaearctic Region to Afghanistan, Ethiopia. New for Saudi Arabia. Host plants. Lavandula spp., Nepeta septemcrenata (Lamiaceae) (Freidberg & Kugler, 1989; Hendel, 1927; Merz, 1992). Pest status. Not a pest species. Paraspheniscoides binarius (Loew, 1861) Material. 2: 19. Distribution. Afrotropical Region and islands of the Indian Ocean, Yemen (Merz et al., in press). New for Saudi Arabia. Host plants. Lippia spp. and Lantana spp. (Verbenaceae) (Munro, 1947). Pest status. Not a pest species. Paraspheniscus debskii (Efflatoun, 1924) Material. 2:12. Distribution. Described from Egypt. New for Saudi Arabia. Host plant. Stachys aegyptiaca (Lamiaceae) (Efflatourn, 1924; Freidberg & Kugler, 1989; Hendel, 1927). Pest status. Not a pest species. Tribe TEPHRITINI Campiglossa genus group Campiglossa ignobilis (Loew, 1861) Material. 2: 346 3,419 2,1 without abdomen. Distribution. Eastern Afrotropical Region, Yemen. New for Saudia Arabia. Host plant. Sonchus oleraceus (Asteraceae) (Munro, 1957a). Pest status. Because its host is a weed in some parts of the world, this species may be useful in the natural regulation of S. oleraceus. Desmella sp. nr myiopitoides (Bezzi, 1908) Material. 2:29 9. Distribution. D. myiopitoides is known from the Eastern Afrotropical Region. Recently, a specimen of the D. myiopitoides-group was also found in Oman (Jebel Shams, 2500m, 7.V.1989, leg. M. J. Ebejer, MHNG, new record). New for Saudi Arabia. Host plant. D. myiopitoides was reared in South Africa from flowers of Chrysocoma tenuifolia (Asteraceae) (Munro, 1925). Pest status. Not a pest species. Remarks. As Munro (1957a) pointed out this genus is in bad need of a revision. No further progress has been carried out since then. The two females at hand key out as D. myiopitoides (Bezzi), but this may be a complex of different species (Munro, 1000 B. MERZ & H. A. DAWAH 1957a). One female has a wing pattern similar to Fig. 137, and the other is similar to Fig. 139 in Munro (1957a). No males are available which may have good characters (Munro, 1957a). For these reasons the specimens from Saudi Arabia cannot be iden- tified unambiguously. The genus is newly recorded for Saudi Arabia. Dioxyna sororcula (Wiedemann, 1830) Material. 2: 599. Distribution. Tropics and Subtropics of the Old World, Australia, introduced to Hawaii. New for Saudi Arabia. Host plants. Bidens spp., Guizotia sp., Lactuca sativa (Asteraceae) (Freidberg & Kugler, 1989; Merz, 1992; White & Elson-Harris, 1992). Pest status. A potential or minor pest on some cultivated plants (Lactuca in the Philippines). On the other hand it may be important in the natural regulation of Bidens spp. which are weeds in some parts of the world (White & Elson-Harris, 1992). Tanaica Munro, 1957 Type species: Ensina hyalipennis Bezzi, 1924 Diagnosis. Within the Campiglossa group of genera Tanaica (Munro, 1957a) may be recognized by the flattened head (Fig. 24) which is longer than high in profile and the narrow gena; by the long labella which are longer than heigth of head and which are projecting distinctly beyond anterior vibrissal corner; by the wing, which is either entirely hyaline or which has a faint reticulate pattern (Figs 15, 25); by the legs, with the femora black except for the apical quarter; and by the male terminalia: preglans with two setulose areas and glans with two sclerotized plates of uneven length emerging from acrophallus, with the longer plate strongly concave (Figs 29-30). Redescription. HEAD (Figs 14, 24). In profile flattened, longer than high; gena narrow, less than 0.8 times as wide as width of postpedicel; heigth of gena less than 0.15 times the heigth of compound eye; frons bare; frontal stripe (as defined by Munro, 1957a) indistinct, almost invisible; face concave; vibrissal corner distinctly projecting beyond frontofacial corner; mouthparts strongly geniculate with labella longer than heigth of head and projecting beyond vibrissal corner; palpus thin, parallel-sided, as long as labella; scape and pedicel black setulose; postpedicel about 1.5 times as long as wide, dorsoapically rounded; arista almost bare. Chaetotaxy: 2 black frontal setae; 2 orbital setae, anterior black, posterior short, white; ocellar seta black; medial vertical seta black; lateral vertical seta and postocellar seta white; paravertical seta absent or short and white; medial postocellar seta absent; postocular setae mixed white and black. THORAX. Scutum white setulose, more or less uniformly grey microtrichose, without distinct brown longitudinal stripes; scutellum apically with small yellow patch. Chaetotaxy (all setae black): 1 postpronotal seta; 2 notopleural setae, the posterior shorter; 1 presutural supra-alar seta; 1 postsutural supra-alar seta; 1 intra-alar seta; 1 postalar seta; 1 dorsocentral seta distinctly anterior to level of postsutural supra-alar seta; 1 acrostichal seta; 2 (pairs of) scutellar setae, the apical less than half as long as the basal; 1 anepisternal seta and 1 katepisternal seta, both black; 1 white, lanceolate anepimeral seta. FRUIT FLIES FROM SAUDI ARABIA 1001 LEGS. Mat yellow, but basal 0.7-0.8 of all femora black; coxae and trochanters black or mixed black and yellow; midtibia with 1 apicoventral black seta. WING (Figs 15, 25). Hyaline or with faint reticulate pattern; pterostigma hyaline or brown with round hyaline spot medially; posterodistal extension of cell bcu short, but distinct; R-M in distal third of cell dm; distance between crossveins equal to length of DM-Cu; veins bare, only R1 with usual dorsal row of setulae, which is interrupted opposite level of tip of Sc, and ventrally with 0-5 setulae in distal half of pterostigma. Calypteres yellow, lower, ventral calypter strip-like, much smaller than upper, dorsal; halter yellow. ABDOMEN MALE. Shining black or grey microtrichose and then with indistinct black paired spots on tergites; sternite 5 (Figs 16, 26) smoothly concave apically; epan- drium (Fig. 18) ovoid, with distinct epandrial process; medial surstylus with 2 prensi- setae of very unequal length, medial prensiseta much larger (Fig. 19); preglans area with two areas of fine setulae (Fig. 20); glans with strongly sclerotized, tube-like acro- phallus; vesica twice as long, with two sclerotized plates, the longer twice as long as the shorter, concave; shorter plate distally truncate. ABDOMEN FEMALE. Preabdomen as male; oviscape black, fine black setulose; at most as long as preceding 3 tergites combined; aculeus (Figs 21-22) evenly pointed apically; two spherical, pyrifrom spermathecae with small papillae (Fig. 23). Remarks. Tanaica was described by Munro (1957a) in the Ensina series of genera for a species with entirely hyaline wings and a shining black abdomen. As it was already admitted by Munro (1957a), the genera placed in this group do not belong to a homogenous group, but are merely an assemblage of taxa which cannot be placed elsewhere. In fact, Ensina Robineau-Desvoidy is now placed in the tribe Noeetini, whereas the other genera in the tribe Tephritini (Norrbom er al., 1999a). Ptosanthus Munro, Sphenella Robineau-Desvoidy and Telaletes Munro belong to the Sphenella genus group (Freidberg, 1987; Norrbom et al., 1999a), whereas the relationships of the two monotypic genera Deroparia Munro and Namwambina Munro are still not satis- factorily known. Tanaica, the last of these genera, was correctly associated with the Campiglossa genus group (Norrbom et al., 1999a) because of the presence of setulose preglans area (absent only in Antoxya Munro and Dioxyna Frey), the flattened head and the mostly black setae on head and thorax. The only species included in Tanaica by Munro (1957a) exhibits two unusual characters for the Campiglossa genus group and may have influenced the placement of the genus in the Ensina series: the entirely hyaline wing and the shining black abdomen. However, with the inclusion of the two species described below the concept of the genus is modified, including also species with faint reticulate wing pattern and with grey microtrichose abdomen. Superficially they resemble species of Desmella Munro and Dioxyna. Whereas the latter genus is somewhat isolated by the lack of setulae on the preglans area, it is possible to separate safely Desmella from Tanaica only by the structure of the glans. More careful comparative studies of all included species are, however, needed to establish whether both genera may be maintained or eventually be synonymized. In order to place Tanaica in the Campiglossa genus group, a key to the Afrotropical and Near Eastern genera is provided. Excluded are Homoeotricha Hering 1002 B. MERZ & H. A. DAWAH (6 species in Central and Eastern Palaearctic, to the west to Kazakhstan and Kyrghyzia) and Oxyparna Korneyev (2 species in mountains between Kyrghyzia and China). 1 One pair of frontal setae present (rarely on one side 2-3 frontal setae present, in these cases lunule very high and scape white setulose)......... 2 hel At least 2 pairs of subequal frontal setae present; scape always black SECU OSS hay RI o en oOo mouse kenne a ee 00 0 00000000 3 2 Scape black setulose; frons setulose anteriorly; lunule normal; posterior notopleural seta black; postpedicel about twice as long as wide ei KERN Pilone cri A eten ca | resa ge era Antoxya Munro [1 Afrotropical species; A. oxynoides (Bezzi)] DE Scape white setulose; frons bare; lunule very high, a semicircle; poste- rior notopleural seta usually white; postpedicel barely longer than wide 3 0 DR) ABI NO NES ERS N ARI AA PS PAL eRe Oxyna Robineau-Desvoidy [23 species in Nearctic and Palaearctic Regions] 3 Apical scutellar setae at least 0.75 times as long as basal scutellar setae ..... 4 35 Apical scutellar setae at most 0.6 times as long as basal scutellar setae, orCntirelrabsente 4e. e oh de ee A E 5 4 Labella long geniculate, longer than heigth of head (as in Fig. 24) REA RO ee MI ee i on TEE Id PETE DE! Mesoclanis Munro [8 Afrotropical and 1 Oriental species; 1 species introduced to Australia] 4* Labella short geniculate, much shorter than heigth of head . . . . Scedella Munro [15 Afrotropical and 2 Oriental/Australasian species] 5 Apical. scutellar‘setae entirely absent) ....2 0.1... a a eee 6 She Apical scutellar setae present, though sometimes only little longer than Surrounding. setulae... 26s. 0 as AiG we eee) Gee ae SR ee U 6 Head in profile higher than long; labella about 0.8 times as long as heigth of head; gena 0.2 times as high as eye and about as high as width of postpedicel; abdominal tergites dull, with indistinct paired spots on tersitesB Ss: ymalespreslanszareassetuloser Serra Lethyna Munro [7 Afrotropical species, 1 unidentified species in Yemen (Merz et al., in press)] 6* Head in profile wider than long; labella longer than heigth of head; ge- na at most 0.15 times as high as eye; about 0.7 times as high as width of postpedicel; paired spots on abdominal tergites distinct; male: preglans ATE AN DANS rer CORE Dioxyna Frey, part [10 species, worldwide distribution] 7 Rostenorn notopleuralisetaswhiter = re Campiglossa Rondani, part [almost 200 species worldwide, most diverse in Afrotropical, Nearctic and Palaearctic Regions] The Posterior notopleural.seta black.......< Lt u... „ug. RESA 8 8 Dorsocentral seta distinctly posterior to level of postsutural supra-alar setaJheadktlattened. BR 2 2 eins ee ee Campiglossa Rondani, part [here only C. martii (Becker) from the Canary Islands and Southern Spain] 8* Dorsocentral seta closer to suture than to postsutural supra-alar seta; Shape of head variable... Le INR ee eee B FRUIT FLIES FROM SAUDI ARABIA 1003 9 Head in profile higher than long; labella usually shorter than heigth of head; wing pattern usually reticulate, rarely banded; male: preglans area SetulOSe! ik). Ak See COLORE stds IE Campiglossa Rondani, part [see couplet 7] IE Head in profile flattened, longer than high; labella longer than heigth of head, usually projecting beyond vibrissal corner; other characters variable . . 10 10 Wing pattern strong, usually of banded appearance, with clear contrast between brown and hyaline areas; pterostigma uniformly brown without central hyaline spot; femora at least in basal half black; male: preglans ATCIISCIMIOSCERIONI I m N Sa: RP TRS Desmella Munro [3 Afrotropical and 2 Palaearctic species] 10* Wing pattern weak, reticulate, or wing entirely hyaline; border between brown and hyaline areas indistinct; pterostigma uniformly hyaline or brown with central hyaline spot; femora yellow or partly black; male: preslanstarea»setulosesorbare.. RE, ee fut 11 At least midfemur mostly yellow. Male: preglans area bare . Dioxyna Frey, part [here only the Palaearctic species D. bidentis (Robineau-Desvoidy) with short apical scutellar setae] 11* All femora at least in basal 0.7 black. Male: preglans area setulose (Figs DODO WAS MAREN RER ELMALEH EN) LTR IONS Yad eh Pa Tanaica Munro [3 Afrotropical species, see below] Key to species of Tanaica Munro, 1957 1 Wing with faint, but distinct reticulate pattern (Fig. 15); abdomen grey mierotrichosesnotshininsne un. nenn: T. maculata sp.n. 1 Wing entirely hyaline, at most pterostigma yellowish (Fig. 25); abdomen variable an... sy) el PE RT 2 2 Abdomen strongly microtrichose, mat .....:.......... T. pollinosa sp. n. 2 Abdomen shining black, only very thinly microtrichose RIE ZERO: AED T. hyalipennis (Bezzi, 1924) Tanaica hyalipennis (Bezzi, 1924) Ensina hyalipennis Bezzi, 1924: 549. Holotype 9: SOUTH AFRICA: Cape, Cedarbergen, Clanwilliam, 4000-5000 ft, IX. 1923, leg. K. H. Barnard (SAMCT) (not examined). Material. 13,29 9, NAMIBIA: Keetmanshoop, 12.1X.2003, leg. A. Freidberg (TAU). Remarks. The species was extensively described by Bezzi (1924), and redes- cribed by Munro (1929, 1957a) by adding new characters. Munro (1929) stated that the abdomen is shining black in the holotype, whereas it was described as «black, grey dusted» by Bezzi (1924). Because it was not possible to study the holotype we assume here that Munro correctly interpreted the colour of the abdomen. On the other hand, the illustration of the glans by Munro (1957a) is misleading because the distal two sclero- tizations were not illustrated nor mentioned in the description. It is possible that he studied a teneral specimen with the sclerotization not yet fully developed. The above 1004 B. MERZ & H. A. DAWAH mentioned male was dissected and it was found that the terminalia are the same as in T. pollinosa (Figs 27-30). According to available information (Munro, 1957a; Norrbom et al., 1999b) T. hyalipennis has a rather narrow distribution in the Western part of South Africa and Namibia. This species was reared from Chrysanthemoides monilife- ra (Asteraceae) (Munro, 1957a). Tanaica maculata sp. n. Figs 14-23 Material. Holotype ¢: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 22.II.-25.V.2002, H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 28 6, 32 2, same data as holotype (CHD, MHNG, NMWC). ETHIOPIA: 156 8,92 2, Mojo, 75 km SE Addis Abeba, 20.XII.1989, A. Freidberg & F. Kaplan (MHNG, TAU). KENYA: 16, W-Nariobi (ILRAD), 1900 m, 1.111.1993, leg. B. Merz; 944,599, same data, but 2.1II.1993; 1 2, same data, but 3.III.1993; 1 © , same data, but 5.III.1993; 14,299, same data, but 8.III.1993; 14,29 2, same data, but 16.11.1993; 26 gd, 40 km SE Nairobi, [on road] A 109, 1650 m, 17.11.1993, leg. B. Merz; 36 8,75 km SE Nairobi (A 109), 1700 m, 17.11.1993; 12, 135 km SE Nairobi (A 109), 1200 m, 17.11.1993, leg. B. Merz (MHNG); 1d, Rt. 104, 15 km SE Nairobi, 29.IV.-15.V.1991, A. Freidberg & F. Kaplan (TAU). The holotype is double-mounted on a minutien pin on a polyporus block and is in good condition with the right postocular seta and the right dorsocentral seta broken off and scutum slightly sunken medially. Etymology. The name refers to the patterned wing, which is unique among known species of the genus. Diagnosis. Within Tanaica this species is easily recognizable by the presence of a reticulate, weak wing pattern (Fig. 15). Cell rl with 3 hyaline spots. The other two species of the genus have an entirely hyaline wing. The abdomen is microtrichose as in T. pollinosa. Description. WING LENGTH. Male: 2.1-3.4 mm; female: 2.0-3.5 mm. HEAD (Fig. 14). Ground colour mat yellow; face, gena and parafacial whitish yellow; frons and antennae orange-yellow. In profile slightly wider than high, with swollen occiput; gena about 1/8 as high as eye, the latter about 1.3 times as high as wi- de; vibrissal corner strongly projecting beyond anterior base of antennae; frons bare, sides slightly converging towards lunule; interfrontal stripe indistinct, only visible as whitish line under certain angle; paravertical seta white, short, not longer than setulae on ventral half of occiput. THORAX. Mesonotum black, densely ash-grey to silvery grey microtrichose, with traces of 3 brown longitudinal stripes over lines of dorsocentral setae and in midd- le; postpronotum and tip of scutellum yellowish; scutum covered with very dense, whi- te setulae; setulae on pleura less numerous; prosternum white setulose. Chaetotaxy as described for the genus. LEGS. As described for the genus. WING (Fig. 15). With a weak reticulate pattern; cell c often with a pale brown mark in middle; pterostigma brown with a hyaline spot in middle or uniformly yellow; rl with 3 hyaline spots; preapically on level of tip of R2+3 with 4 large spots in one line; apex of wing with small hyaline spot; remaining pattern faint, especially in anal lobe. FRUIT FLIES FROM SAUDI ARABIA 1005 SAL Mae Mi 21 Fics 14-23 Tanaica maculata sp. n. (16-20, male; 21-23, female, paratypes from Abha): 14, head, lateral view; 15, wing; 16, abdominal sternites 4 and 5; 17, epandrium, lateral view; 18, same, caudal view; 19, medial surstylus with prensisetae enlarged; 20, aculeus; 22, tip of aculeus; 23, sper- matheca (only one illustrated). ABDOMEN MALE. Preabdomen as in 7. pollinosa. Terminalia (Figs 16-20). Epandrium (Fig. 18) ovoid, black, but epandrial process brown; the latter with serrate border, visible in lateral view (Fig. 17); medial surstylus with two unequal prensisetae present on prominent plate, the lateral prensiseta much larger than the medial (Fig. 19); preglans area with 2 rows of short setulae; glans (Fig. 20) with well sclerotized basal 1006 B. MERZ & H. A. DAWAH acrophallus and with 2 sclerites in vesica, of which one is long, apically pointed and medially curved and the other sclerite is short, widening apically with concave distal margin. ABDOMEN FEMALE. In all characters, including terminalia (Figs 21-23) as in T. pollinosa (see below). Distribution. Ethiopia, Kenya, Saudi Arabia. Host plants. Unknown. The specimens from Kenya were swept from various herbs in grassland areas, whereas the specimens from Saudi Arabia were collected in a Malaise trap. Pest status. Not a pest species. Remarks. This species is easy to recognize within Tanaica by the reticulate wing pattern. The species is somewhat variable in wing pattern. Specimens from Ethiopia and Kenya tend to have a weaker pattern, with notably cell c entirely hyaline (without central darker spot) and with pterostigma uniformly yellow (in Saudi Arabian spe- cimens usually brown with central hyaline spot). However, male and female termina- lia are the same for populations from Kenya and Saudi Arabia, and few specimens from Kenya have the same wing pattern as those from Saudi Arabia. At the present moment it is not possible to decide whether these populations represent different species. Further studies, in particular the knowledge of host plants, would give further eviden- ce about their status. T. maculata has the same type of wing pattern as do some Dioxyna Frey. However, all Dioxyna lack the two rows of setulae on the preglans area (present in Tanaica). Moreover, they lack, except for D. bidentis, the apical scutellar setae. The latter species differs from 7. maculata by the almost entirely yellow legs (femora blackish in 7. maculata). Tanaica pollinosa sp. n. Figs 24-33 Material. Holotype d: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 22.II.-25.V.2002, H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 14,299 ,sa- me data as holotype (CHD, MHNG). SOUTH AFRICA: 36 8,2? 9, N Transvaal, Sautpansberg, 1500 m, 15km NW Louis Trichardt, 25.XII.1994, A. Freidberg. The 222 with additional label: ex ? seeds Tripteris auriculata, 1.1.1995 (TAU, MHNG). The holotype is double-mounted on a minutien pin on a polyporus block and is in good condition with few setae on head and thorax broken off. Etymology. The name refers to the grey microtrichose abdomen. Diagnosis. This species differs from the other species from Saudi Arabia by its entirely hyaline wing without traces of a pattern (Fig. 25). The abdomen is densely grey microtrichose; tergites 2-5 each with a pair of indistinct brown spots. Description. WING LENGTH (both sexes). 2.4-2.6 mm. HEAD (Fig. 24). Ground colour mat yellow, but occiput and ocellar triangle black; face and gena whitish yellow; frons and antenna orange-yellow. Head in profile wider than high, with prominent occiput; gena narrow, less than one sixth as high as compound eye; the latter ovoid, about 1.25 times as high as wide; vibrissal corner pro- jecting beyond level of antennal base; frons bare; interfrontal stripe indistinct, narrow, almost white; paravertical seta white, shorter than posterior orbital seta. FRUIT FLIES FROM SAUDI ARABIA 1007 Fics 24-33 Tanaica pollinosa sp. n. (24-30, male; 31-33, female, paratypes from Abha): 24, head, lateral view; 25, wing; 26, abdominal sternites; 27, epandrium, lateral view; 28, same, caudal view; 29, glans, lateral view; 30, glans, frontal view; 31, aculeus; 32, tip of aculeus; 33, spermathecae. 1008 B. MERZ & H. A. DAWAH THORAX. Mesonotum black, densely silvery grey microtrichose; postpronotal lobe and posterior half of scutellum yellow; scutum densely covered with white, lanceolate setulae; pleura slightly darker grey, setulae on anepisternum, katepisternum and anepimeron less numerous and more acuminate; prosternum grey, white setulose. Chaetotaxy as described for the genus. LEGS. As described for the genus. WING (Fig. 25). Entirely hyaline, only stigma faintly yellowish. ABDOMEN MALE. Tergites darker grey than mesonotum, but still distinctly golden-grey microtrichose; each tergite with a pair of almost parallel-sided spots which are forming a more or less uniform band from base to tip of abdomen; setulae short, dense, lanceolate; sternites (Fig. 26) darker grey, sparsely setulose; sternite 5 shallowly concave with smooth apicolateral corners; epandrium (Figs 27-28) black, but epandrial process brown; the latter prominent and distinctly projecting, with serrate border; inner terminalia as usual in tribe; aedeagus (Figs. 29-30) with 2 rows of setulae on preglans area; glans with strong basal sclerotization and 2 curved, unevenly long plates forming an open tube; the longer apically pointed and strongly curved medially; the shorter plate distinctly widening distally, forming an open rim; vesica as broad as acrophallus, distally truncate, rather small. ABDOMEN FEMALE (Figs 31-33). Coloration and chaetotaxy of pregenital segments as male; oviscape black, fine setulose, without white setae, as long as pre- ceding 3 tergites combined; aculeus apically evenly pointed, without step; 2 almost spherical spermathecae, covered with short papillae; aculeus length: 0.77 mm (n = 1). Distribution. Saudi Arabia, South Africa. Host plant. Tripteris auriculata (Asteraceae). Pest status. Not a pest species. Remarks. This species differs from T. hyalipennis (Bezzi), the other species with entirely hyaline wings, only in the colouration of the abdomen. It is shining black in the latter, but grey microtrichose in T. pollinosa. All other external characters and the terminalia of both sexes are the same in both species. Spathulina genus group Spathulina acroleuca (Schiner, 1868) Material. 2:29 °. Distribution. Widespread in the Afrotropical, Southern Palaearctic, Oriental and Australasian Regions. Recorded from Yemen by Merz et al. (in press). New for Saudi Arabia. Host plants. Various Asteraceae, such as Bidens, Ceruana, Matricaria and Spilanthes (Munro, 1938). Pest status. The species was recorded as minor pest on Mexican sunflower (Tithonia diversifolia) in the Philippines (White & Elson-Harris, 1992). Sphenella genus group Sphenella marginata (Fallén, 1814) Material. 2: 18,19. Distribution. Palaearctic and Afrotropical Regions. New for Saudi Arabia. FRUIT FLIES FROM SAUDI ARABIA 1009 Host plants. Senecio spp. (Asteraceae) (Freidberg & Kugler, 1989; Merz, 1994). Pest status. May be important in the natural regulation of those Senecio spp. which are weeds in some parts of the world (White & Elson-Harris, 1992). Remarks. After the study of large series from the Palaearctic and Afrotropical Regions we conclude that the differences given to separate S. marginata s.str. (Palaearctic subspecies) from S$. marginata austrina Munro, 1957 (Afrotropical sub- species) (Munro, 1957b) are unreliable with many intermediate specimens known. The two taxa are considered here to belong to the same species and they are herewith synonymized (syn. n.). Sphenella setosa sp. n. Figs 34-41 Material. Holotype d: «SAUDI ARABIA: Aseer, Abha, Farm Centre, 14.IV. 2001, H. A. Dawah»; «Tephritis sp. d , lower angle anal cell more prominent than any sp. photo in Die Fliegen» [J. C. Deeming’s handwriting] (MHNG). Paratype: 14, SAUDI ARABIA: Aseer, Abha, Farm Centre, 14.1V.2001, H. A. Dawah (MHNG). The holotype is glued laterally on a card point, and is in good condition with left arista and few setae missing (left ocellar seta, few postocular setulae, left postpronotal seta, both left notopleural setae, left presutural supra-alar seta; left postsutural supra-alar seta; both right scutellar setae). Etymology. The name refers to the strongly setulose legs. Diagnosis. This species differs from all described Sphenella by the strongly setulose hindfemur and hindtibia (Figs 36-37), the wing (Fig. 35) with the ill-defined pattern of paler and darker areas on the entire surface, the strongly modified sternite 5 with 2 medial tubercles (Fig. 38), and the very large, black, globose epandrium (Fig. 40). The wing pattern resembles superficially species of Paratephritis Shiraki, Tela- letes Munro and Orotava Frey, but the combination of bare R4+5, flat scutellum, presence of anteroventral setae on hindfemur and presence of only one long, pointed prensiseta on the medial surstylus are typical for Sphenella. Other diagnostic characters of this species are the mixed white and black postocular setulae, the yellow legs with only hindfemur black in basal half, the yellow postpedicel, and the sclerotization of the glans. Description (male only). WING LENGTH. 3.50-3.75 mm. HEAD (Fig. 34). Mat yellow, but ocellar triangle and occiput grey microtricho- se. In profile 1.25 times as high as long, gena 0.15 times as high as compound eye; fronto-facial angle about 110°; frons white setulose anteriorly; frontal plate rather densely white setulose; face distinctly concave, without carina; vibrissal corner white setulose; gena dorsally black and ventrally white setulose. Antenna yellow, scape with white setulae, pedicel with black setulae; postpedicel apicodorsally slightly pointed; arista virtually bare. Mouthparts short geniculate, labella fleshy, not extending beyond vibrissal corner. Chaetotaxy: 2 black frontal setae; 1 black anterior and | white, shor- ter posterior orbital seta; 1 black ocellar seta; 1 black medial vertical seta; 1 white la- teral vertical seta; 1 white postocellar seta; 1 white paravertical seta; postocular setu- lae mixed white and black. THORAX. Scutum densely golden grey microtrichose, in dorsal and posterior view with 3 faint brown longitudinal stripes on lines of dorsocentral setae and in 1010 B. MERZ & H. A. DAWAH SSSA AGG RN EN AN Ting al DS Fics 34-41 Sphenella setosa sp. n. (male paratype): 34, head, lateral view; 35, wing; 36, hind leg, anterior view; 37, hind leg, posterior view; 38, abdominal sternites 4 and 5; 39, epandrium, lateral view; 40, same, caudal view; 41, glans. middle; entire surface covered densely with white setulae; scutellum flat, mat yellow, white setulose laterally; pleura slightly more ash-grey than scutum; anepisternum and anepimeron white setulose; prosternum densely white setulose. The following setae are black: 1 postpronotal seta; 2 notopleural setae (posterior notopleural seta about half as FRUIT FLIES FROM SAUDI ARABIA 1011 long as anterior notopleural seta); 1 dorsocentral seta, which is inserted well posterior of suture, about at level of anterior wing base; 1 presutural supra-alar seta; 1 post- sutural supra-alar seta; 1 intra-alar seta; 1 postalar seta; 2 (pairs of) scutellar setae which are of same length; 1 katepisternal seta. Anepisternal seta and anepimeral seta white. LEGS (Figs 36-37). Ground colour yellow, forecoxa whitish yellow, other parts brownish yellow; midfemur posteriorly at base with black stripe; hindfemur in anterior half black. All femora rather stout, conspicuously setulose; hindfemur with a row of long, black setae anteroventrally (Fig. 36), one strong, black anterodorsal seta, and with numerous soft, whitish, very long setulae (about as long as diameter of femur) posteroventrally in basal third (Fig. 37). Hindtibia anterodorsally and posterodorsally each with a row of conspicuous, outstanding, black setulae. WING (Fig. 35). R4+5 bare on both sides; R1 setulose dorsally, with gap on level of subcostal break; ventrally with few setulae in distal half of pterostigma. Pattern brown to yellow brown, indistinctly striped, with darker areas along C, at wing tip, and around crossveins; other parts with a pattern of ill-defined subhyaline spots on pale brown surface. ABDOMEN. Mat ash-grey microtrichose with yellow hind margins of tergites 2-5; tergite 5 about twice as long as tergite 4; all tergites white setulose, only tergite 5 posteriorly with a row of black setulae. MALE TERMINALIA. Sternite 5 (Fig. 38) greatly modified, much larger than pre- ceding sternites; apically with two medial bulbous projections, which are separated by a cleft and which bear apically about 12 stout, black setulae; other setulae rather soft, pale. Epandrium (Figs 39-40) very large, globose, black; epandrial process small, on- ly little projecting; lateral surstylus separated from epandrium by a fold, strongly bent medially and pointed apically; medial surstylus with one very long, spine-like black prensiseta, which is directed posteriorly; hypandrium as in other species of the genus; glans (Fig. 41) rather narrow, tapered apically and truncated, with a strongly sclero- tized ductus. Distribution. Saudi Arabia. Host plants. Unknown. Pest status. Not a pest species. Remarks. This species keys out readily in Munro (1957b) in couplet 4 as Sphenella with the setulose hindfemur anteroventrally, the bare vein R4+5, and the flat scutellum. In addition, only one long, pointed prensiseta is developed. Sternite 5 carries apicomedially two tubercles as in other species of Sphenella but they are much more developed and they are covered with more conspicuous black setae posteriorly. Wing pattern, however, is very different, as other species of Sphenella have large hyaline areas, and the black areas are usually clearly delimited. In this respect S. seto- sa resembles some species of Paratephritis Shiraki, Telaletes Munro, and Orotava Frey, which have the same indistinct reticulation with the ill-defined hyaline spots and the brown stripe over DM-Cu. Within Sphenella the new species exhibits, in addition to the wing pattern and the structure of sternite 5, a number of unique characters: hind- femur with row of black setulae anteroventrally on entire length (in other species of Sphenella only near apex with few setulae) with white setulae in basal third postero- 1012 B. MERZ & H. A. DAWAH ventrally (not present in other Sphenella); epandrium shining black and globose (much more elongated and less conspicuous in other Sphenella). Telaletes ochraceus (Loew, 1861) Material 1:26 6; 21,19. Distribution. East Africa. New to Saudi Arabia. Host plants. Unknown. Pest status. Not a pest species. Tephritis genus group Arabodesis gen. n. Gen. nr. Trupanodesis Merz - Merz et al. (in press). Type species. Arabodesis reductiseta sp. n. Etymology. The name refers to the distribution in Arabia and its resemblance to Trupanodesis Merz and Tephrodesis Merz. Gender is feminine. Diagnosis. The new genus belongs to the Tephritis group as defined by Merz (1999). The absence of medial postocellar setae (Fig. 43), the presence of only white postocular setae (Fig. 42), two (pairs of) subequal scutellar setae (Fig. 45) and the rather simple structure of the glans with short vesica (Fig. 50) place Arabodesis close to Tephrodesis and Trupanodesis. It differs from these genera by the black posterior notopleural seta, the dorsocentral seta situated well behind the transverse suture at le- vel of postsutural supra-alar seta (Fig. 45), the convex lower calypter, the rather conspi- cuous posterodistal extension of cell bcu (Fig. 46), and the aculeus with an apical step (19399) Description. HEAD (Figs. 42-44). In profile at least 1.5 times as high as long, gena and parafacial narrow; vibrissal corner projecting anteriorly; frons parallel-sided, narrow, longer than wide, with few tiny, fine, white setulae on interfrontal stripe; ocel- lar triangle with few white setulae between ocelli; scape white setulose and pedicel black setulose; postpedicel (Fig. 44) about twice as long as wide; arista almost bare; proboscis capitate; palpus not projecting beyond anterior margin of vibrissal corner. Chaetotaxy: 1-2 frontal seta, 1 orbital seta, 1 ocellar seta and 1 long medial vertical se- ta black, acuminate; postocellar seta, lateral vertical seta and row of postocular setae white, lanceolate; medial postocellar seta absent. THORAX (Fig. 45). Shape as in other genera of the Tephritis-group; scutum covered with white setulae. Chaetotaxy: scapular setae absent; setae on thorax black, including posterior notopleural seta, only anepimeral seta white; 1 postpronotal seta; 2 notopleural setae, the posterior shorter; 1 presutural supra-alar seta; 1 postsutural supra-alar seta; 1 intra-alar seta; 1 postalar seta; 1 acrostichal seta; 1 dorsocentral seta well behind transverse suture, about at level of postsutural supra-alar seta; 2 (pairs of) subequal, strong scutellar setae; 1 anepisternal seta; 1 katepisternal seta; 1 anepimeral seta; prosternum white setulose. LEGS. Hindfemur anteroventrally with 1-2 outstanding, longer setulae at tip; midtibia apicoventrally with one black seta. WING (Fig. 46). Stigma about twice as long as wide; R-M crossvein situated in distal third of cell dm; distance between crossveins about 0.8 times as long as DM-Cu; FRUIT FLIES FROM SAUDI ARABIA 1013 posterodistal extension of bcu distinct; veins bare, but RI dorsally setulose except for gap on level of tip of Sc, ventrally with 0-2 setulae at tip of pterostigma. Both calypters with convex distal border, of subequal width. ABDOMEN MALE. Densely white setulose, without longer setae; sternite 5 (Fig. 47) over 1.3 times as wide as sternite 4, distally smoothly concave; epandrium (Figs 48-49) ventrally very wide, bulging, fused with lateral surstyli, the latter unmodified; epandrial process present but not very prominent; medial surstylus with 2 subequal prensisetae and 2-3 setulae on plate; hypandrium and associated structures as usual for the Tephritis group, with unpaired hypandrial arms; distiphallus long, coiled, preglans area bare; glans (Fig. 50) with rather simple sclerotization; sclerotized ductus apically with few small spines; vesica short. ABDOMEN FEMALE. Preabdomen as male; tergites 5 and 6 subequal; oviscape (Fig. 51) white setulose, at distal margin with numerous soft setulae; aculeus (Figs 52- 53) with apical step; 2 spermathecae, covered with small papillae (Fig. 54). Remarks. Arabodesis is a puzzling genus which exhibits an unusual combi- nation of characters rendering its placement within the family difficult. Lack of scapular setae, the presence of only white postocular setae, and the gap of setulae on RI opposite tip of Sc show that it belongs to the Tephritini. Based on the revisions of some genus groups within the tribe by Freidberg (1987, Sphenella group), Korneyev (1990, Campiglossa group) and Merz (1999, Tephritis group) it can be concluded that the new genus shares the synapomorphies of the Tephritis group as defined by Merz (1999): glans rather simple, weakly sclerotized and the presence of capitate mouthparts. Within the Tephritis group, Arabodesis exhibits a number of plesio- morphic character states, such as (1) the absence of medial postocellar setae (apomor- phic state (= A): medial postocellar setae present), (2) the black posterior notopleural seta (A: white), (3) the position of the dorsocentral seta well behind the suture (A: almost on level of suture), (4) the rather large lower calypter with convex margin (A: small, stripe-like lower calypter with straight margin), and (5) the apical step of the aculeus (A: aculeus evenly pointed), although the polarity of the latter character is unresolved. A black posterior notopleural seta is known in the Tephritis group only from few Tephritis (arnicae (Linnaeus), conura (Loew)). The position of the dorso- central seta is comparable only with Euarestella Hendel, which belongs to the Goniurellia clade (medial postocellar setae present; in addition, this genus has 2+1 frontal and 2 orbital setae). The large lower calypter is only known from Tephritis Latreille (but this genus has 2 frontal and 2 orbital setae; head shape is different, and the dorsocentral seta is almost on level of suture). The apical step on the aculeus is developed in the Tephritis group only in some Tephritis (pulchra group) but an apical step is widespread in many genera of Tephritini. Arabodesis shares with Tephrodesis Merz a similar type of glans of aedeagus, which is evidence for a sister-group relationship and this character may be a good synapomorphy for these two genera. However, the position of Tephrodesis within the Tephritis group is not well understood, as its sister-group relationships are unknown (Merz, 1999). Tephrodesis differs from Arabodesis by the development of the apo- morphic character states of characters (2) to (5) mentioned above, but also by the patterned wing, the 2+1 frontal and 2 orbital setae, and a black oviscape. Further, it 1014 B. MERZ & H. A. DAWAH lacks the small spinules on the inner side of the ductus of the glans which are charac- teristic for Arabodesis. Autapomorphies of Arabodesis are probably the presence of (a) only 1 orbital seta (in Tephritini very rare, only known from Actinoptera Rondani, but this genus differs in addition to the apomorphic character states (2) — (5) listed above by the very small pterostigma and a patterned wing), (b) the very high head (otherwise only in Hyalotephritis Freidberg, but this genus has the apomorphic character states (1) — (5), and different male terminalia) and (c) the large posterodistal extension of cell bcu (less conspicuous in other species of the Tephritis group). In the key of Merz (1999) the new genus runs to couplet 11 which needs to be modified as follows: 11 One black orbital seta present; posterior notopleural seta black; dorso- central seta almost at level of postsutural supra-alar seta; wing entirely MY aM Peep SITR IE Arabodesis gen. n. 11* Two orbital setae present; posterior notopleural seta white; dorsocentral seta closer to suture than to postsutural supra-alar seta; wing usually with pattern, though sometimes weak ~~... 5.2.2 TE lla ita “Two concolourous frontal setae present... i i ee 12 11a* Three frontal setae, the anteriormost seta sometimes white .............. 15 This genus was first mentioned, but not formally described from Yemen based on three badly preserved specimens belonging to two species differing chiefly in the chaetotaxy of the head (Merz er al., in press). Arabodesis reductiseta sp. n. Figs 42-54 Gen. nr. Trupanodesis, sp. 1 - Merz et al. (in press). Material. Holotype 2: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 22.11.-25.V.2002, H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 56 46,1522, same data as holotype (CHD, MHNG, NMWC, TAU). YEMEN: 14,19, San’a, light trap, 3.1V.1999, leg. A. von Harten (MHNG, ZMUA); 19, San’a, ca. 7900 ft., 10.- 15.X.1937, D. Carl Rathjens, B.M. 1939-396, from Lucerne (NHML). The holotype is glued laterally on a cardpoint, and is in good condition but both wings folded over each other. Etymology. The species is named for the small number of setae on the head. Diagnosis. Easy to recognize by the presence of only | frontal and 1 orbital seta, both short, fine and black (Fig. 42); ocellar seta short; postpedicel with distinct dorso- apical point (Fig. 44); wing entirely hyaline (Fig. 45); male: epandrium medially strongly convex (Fig. 49), epandrial process in lateral view hardly visible (Fig. 48); glans with few small papillae near tip of ductus (Fig. 50); female: aculeus with apical step (Fig. 53). Description. WING LENGTH. Male: 2.05-2.30 mm; female: 2.20-2.60 mm. HEAD (Figs 42-43). Mat yellow, but ocellar triangle and occiput dorsal to occi- pital foramen ash grey microtrichose; head in profile 1.5-1.6 times as high as long; compound eye about 1.75 times as high as wide; gena about 1/6 as high as compound eye; only one weak frontal seta almost at level of antennal bases; 1 weak, reclinate FRUIT FLIES FROM SAUDI ARABIA 1015 42 3 N Fics 42-46 Arabodesis reductiseta sp. n. (paratype from Abha): 42, head, lateral view; 43, head, frontal view; 44, antenna; 45, thorax, dorsal view; 46, wing. orbital seta very high on head near vertex; all other setae also weak except for long medial vertical seta; postocular setae all white, short, in one row; pedicel without strong dorsal seta; postpedicel about twice as long as wide, with conspicuous dorso- apical point (Fig. 44). THORAX (Fig. 45). Scutum, pleura and subscutellum mostly ash grey micro- trichose; only postpronotal lobe, dorsal part of anepisternum, posterior half of noto- pleuron, most of prosternum and entire scutellum mat yellow; scutum covered with short, weak, white setulae; scutellum bare, but white setulose along margin; chaetotaxy as described for genus; halter yellow; both calypters yellow. LEGS. Yellow, but hindfemur ventrally in middle with grey patch. WING (Fig. 46). Entirely hyaline, only pterostigma slightly yellowish. ABDOMEN MALE (Figs. 47-50). Mat ash grey microtrichose; posterior margin of tergites with yellow transversal stripes, broader on syntergite 1+2 than on posterior 1016 B. MERZ & H. A. DAWAH tergites; sternites yellow, each with a pair of grey spots (Fig. 47); sclerotization of glans compact (Fig. 50); acrophallus with two internal ducts, which are crossing in distal half; near tip with few small papillae on inner side; vesica small, apically broadly rounded. ABDOMEN FEMALE (Figs 51-54). Oviscape about as long as preceding 1-1.5 tergites combined, about 0.5 mm long; orange on surface, black at both sides and along a medial longitudinal stripe: aculeus with slightly concave margin in basal half and abruptly narrowing in apical third, tip with apical step; length about 0.45 mm. Distribution. Saudi Arabia, Yemen. Host plants. Unknown. All specimens were collected in Malaise traps. Pest status. Not a pest species. Remarks. At first glance A. reductiseta resembles species of Hyalotephritis, which is the only genus of the Tephritis group with species having entirely hyaline wings and a similar head shape. Chaetotaxy (especially absence of medial postocellar setae), position of the dorsocentral seta, colour of posterior notopleural seta, and presence of 1-2 anteroventral setulae on hindfemur near tip indicate immediately that the new species does not belong to the Goniurellia clade in the Tephritis-group (see also above). A second species of Arabodesis is known from Yemen (Merz et al., in press). It differs from A. reductiseta by the stronger setae on the head, the presence of 2 frontal setae, and the postpedicel being dorso-apically rounded. As only one female is available, which is slightly shrivelled, this species is not formally described here. Capitites augur (Frauenfeld, 1857) Material. 2: 28 à. Distribution. Egypt, Israel, Sudan. Recorded from Saudi Arabia by Al-Ahmadi & Salem (1999, as Trupanea augur) and by Norrbom er al. (1999b, as Paradesis au- gur). It should be noted that this species was misidentified in the past, and its status was resolved by Freidberg & Kugler (1977). Therefore, it may be possible that old records refer to other species (Goniurellia spp.). Host plant. Pulicaria crispa (Asteraceae) (Freidberg & Kugler, 1989, as Dectodesis augur). Pest status. Not a pest species. Dectodesis auguralis (Bezzi, 1908) Material. 2:28 8,1%. Distribution. Eastern Afrotropical Region from Eritrea to Zimbabwe, Yemen (Merz et al., in press). New for Saudi Arabia. Host plants. Unknown, but other species of the genus live in flowerheads of Erigeron, Helichrysum and Gnaphalium (Asteraceae). Pest status. Not a pest species. Dectodesis sp. 1 Material. 2: 14. Remarks. This male lacks the head and is also otherwise in bad condition. It is very similar to the previous species, but the apical scutellar setae are absent. Judging from wing pattern and colour of mesonotum this species may belong to D. confluens FRUIT FLIES FROM SAUDI ARABIA 1017 Fics 47-54 Arabodesis reductiseta sp. n. (47-50, male; 51-54, female, paratypes from Abha): 47, abdominal sternites 4 and 5; 48, epandrium, lateral view; 49, same, caudal view; 50, glans; 51, oviscape; 52, aculeus; 53, tip of aculeus; 54, spermatheca (only one illustrated). 1018 B. MERZ & H. A. DAWAH (Wiedemann, 1830), a very widespread and abundant species in Eastern and Southern Africa. Because of the poor condition of the specimen we are not naming it here. Freidbergia mirabilis Merz, 1999 Material. 2: 19. Distribution. Described from Kenya and Ethiopia (Merz, 1999). New for Saudi Arabia. Host plants. Pluchea spp. (Asteraceae) (Merz, 1999). Pest status. Not a pest species. Goniurellia persignata Freidberg, 1980 Material. Vs VG; 2: 116,22 2. Distribution. From Sri Lanka and China in the East through Middle Asia and the Southern Mediterranean Region to Morocco. Also known from Ethiopia. New for Saudi Arabia. Host plant. Pulicaria arabica (Asteraceae) (Freidberg, 1980). Pest status. Not a pest species. Goniurellia spinifera Freidberg, 1980 Material. 2: 16,19. Distribution. Near East and North Eastern Afrotropical Region. Recorded from Saudi Arabia by Freidberg (1980). Host plant. Pulicaria desertorum (Asteraceae) (Freidberg, 1980). Pest status. Not a pest species. Hyalotephritis Freidberg, 1979 This genus was proposed by Freidberg (1979) for two species which are morphologically very similar: H. complanata (Munro) and H. planiscutellata (Becker). He provided a key and illustrated the main differences between the two species (head shape, structure of glans, aculeus tip). Larvae of both species live in flowerheads of Conyza dioscoridis (Asteraceae). The comparison of the 35 specimens from Saudi Arabia with numerous specimens of H. planiscutellata from Israel, Kenya and Oman, and two pairs of H. complanata from South Africa showed that both species are present in Saudi Arabia. It was observed that the structure of the glans is the only reliable character separating the two species, whereas head and aculeus shape are more variable. The glans of the Saudi Arabian specimens of H. planiscutellata fits the illus- tration of Freidberg (1979), but a notable difference could be found in the sclerotiza- tion of the glans in all specimens of H. complanata (Fig. 55), including one male from South Africa, compared with the illustration of Freidberg (1979, Fig. 13): The distal part of the sclerotized tube is longer and stronger, undulating, ending in a small apical sclerite which is darker than the previous part. i Hyalotephritis complanata (Munro, 1929) Fig. 55 Material 2723'S 332 2; INTO Distribution. South and East Africa. New for Saudi Arabia. Host plant. Pluchea dioscoridis (Asteraceae) (Freidberg, 1979). FRUIT FLIES FROM SAUDI ARABIA 1019 Pest status. Not a pest species. Remarks. For the structure of the glans of the aedeagus see above (Fig. 55). Hyalotephritis planiscutellata (Becker, 1903) Material. 3: 118 6,699. Distribution. Near East south to Ethiopia, Yemen (Merz et al., in press). New for Saudi Arabia. Host plant. Pluchea dioscoridis (Asteraceae) (Freidberg & Kugler, 1989). Pest status. Not a pest species. Tephritomyia despoliata (Hering, 1956) Figs 56-58 Acanthiophilus (Tephritomyia) despoliatus Hering, 1956: 88. Holotype d : «IRAN (Chorassan), Birdjant, 23.VII.-2.VIII.1954, Richter & Schäuffele», «Acanthiophilus despoliatus m. Type, det. M. Hering 1955» (SMNS). Paratype d, same data as holotype (SMNS) (examined). Material. 2:73 8,1022. Distribution. So far only known from Iran. New for Saudi Arabia. Host plants. Unknown. All other species of the genus with known host plants live in flowerheads of Echinops spp. (Cardueae, Asteraceae) without causing galls. Pest status. Not a pest species. Remarks. Tephritomyia is a difficult Palaearctic and Afrotropical genus of morphologically similar species and is in need of revision. Based on the original description the specimens from Saudi Arabia resemble Acanthiophilus despoliatus Hering, which was described from 3 specimens from Iran (Hering, 1956). The compa- rison of the holotype and a paratype of the latter and the specimens from Saudi Arabia shows a general resemblance. The differences are: 1. Wing length: The two specimens from Iran have longer wings (4.60-4.81 mm, compared to 3.6-4.2 mm in Saudi Arabian specimens), but the specimens from Saudi Arabia were collected in alcohol and dry mounted, and have the wings usually slightly folded. The real length may be longer. 2. Leg colour: The types from Iran have entirely yellow legs, whereas the colour of the Saudi Arabian population is more variable, with specimens having entirely yellow legs to others with black marks or a black ring in basal third of one to all femora. T. despoliatus is morphologically quite similar to the widely distributed Palae- arctic species Tephritomyia lauta (Loew) (compare head shape, Fig. 56 with Fig. 133 in Freidberg & Kugler, 1989, and male terminalia, compare Fig. 58 with Fig. 24.12.C in Merz, 1999, for T. lauta). The two species differ by the much paler wing pattern without clearly delimited hyaline spots in T. despoliatus (Fig. 57). In T. lauta, the pattern is stronger with the hyaline spots more regular and entirely surrounded by the dark pattern (see Fig. 24.8.A in Merz, 1999). In general, T. despoliatus is much paler, with the scutellum either entirely yellow or only grey at base (usually entirely grey in T. lauta). The abdominal tergites have a more extensive yellow pattern (usually almost entirely grey in 7. lauta), but intermediate specimens occur in both species. The length of the oviscape is more variable in 7. lauta (as long as preceding 2.5-4 tergites combined, but at most as long as preceding 3 tergites combined in T. despoliatus). 1020 B. MERZ & H. A. DAWAH Fics 55-58 Hyalotephritis complanata (Munro, 1929, specimen from Abha) (55) and Tephritomyia despo- liata (Hering, 1956, specimen from Abha) (56-58): 55, 58, male, glans; 56, head; 57, wing. Hering (1956) considered the shape of the postpedicel as the best discriminating cha- racter between T. despoliatus and T. lauta. The study of numerous specimens of T. lau- ta from Israel, Cypus and Crete shows, however, that there is no clearcut gap between the two species for this character. It seems that the shape of the postpedicel in the dif- ferent specimens is the result of the drying process. At the present state of knowledge we prefer to consider the two populations from Iran and Saudi Arabia as conspecific. Their status should be revised, however, as soon as more specimens become available, and/or biological data about the host plants are known. Trupanea amoena (Frauenfeld, 1857) Material. 2: 383 6,319 2. Distribution. Southern Palaearctic Region, East Africa, Oriental Region, Australia. Recorded from Saudi Arabia by Al-Ahmadi & Salem (1999). Host plants. This is one of the most polyphagous Tephritinae, attacking capitu- la of species of various Asteraceae (Achillea, Carthamus, Ethulia, Lactuca, Launaea, Leontodon, Picris, Sonchus and others) (Freidberg & Kugler, 1989; Merz, 1994; Munro, 1964). FRUIT FLIES FROM SAUDI ARABIA 1021 Pest status. Recorded as minor pest on Calendula officinalis (Asteraceae) in India (White & Elson-Harris, 1992). Trupanea pseudoamoena Freidberg, 1974 Material. 2:29 9. Distribution. Near East. Recorded from Saudi Arabia by Freidberg & Kugler (1989). Host plant. Pulicaria crispa (Asteraceae) (Freidberg & Kugler, 1989). Pest status. Not a pest species. Trupanea repleta Bezzi, 1918 Figs 59-69 Trypanea aucta var. repleta Bezzi, 1918: 45. Holotype à : [ERITREA] «Ghinda, Mochi, VI.16» (handwritten, white paper), «repleta» (in Bezzi’s handwriting, white paper), «Holotypus d , Trupanea aucta var repleta Bezzi 1918, desig. B. Merz 2004» (handwritten, red label) (MCSNM) (examined). Material. 2: 1086, 2099. Further specimens studied. CAPE VERDE ISLANDS: 36 4, S. Tiago, Lagoa, 15.11.1954, Lindberg; 46 8, 12, Sol, Terra Boa, 21.1.1954, Lindberg; 16, Maio, Pedro Vaz, 3.11.1954, Lindberg; 14 , Maio, Morrinho, 3.1.1954, Lindberg; 16, S. Vicente, Rib. Juliäo, 9.-11.III.1954, Lindberg; 14, Nicolau, Rib. Brava, 6.-19.XII.1953, Lindberg (all FMNH). YEMEN: see Merz et al. (in press). Diagnosis. Within Trupanea this is a typical species of the former Urelliosoma, with crescent hyaline apical wing pattern (Figs 60-63) and with a pale brown abdomen in both sexes. No sexual dimorphism in wing pattern; wing with narrow basal cross- band with an obtuse angle on level of M; this band often broken around M; small hyaline spots in r2+3 and r4+5 sometimes present; head with slightly produced vibrissal corner (Fig. 59); glans with typical sclerotized hook (Fig. 67). Redescription. WING LENGTH. Male: 2.5-3.05 mm; female: 2.5-2.85 mm. HEAD (Fig. 59). Mat yellow, upper frons, ocellar triangle and occiput partly thin grey microtrichose; compound eye 1.3-1.5 times as high as long, gena 1/10 as high as eye; frons about as long as wide, anteriorly with few pale setulae; lunule rather high, conspicuous; face slightly concave; vibrissal corner projecting in profile slightly beyond base of antennae; antennae yellow, scape white setulose, pedicel with black setulae; postpedicel dorsally concave, dorso-apically slightly pointed; mouthparts yellow, capi- tate; palpus not projecting beyond vibrissal corner. Chaetotaxy: frontal setae, anterior orbital seta, ocellar seta and medial vertical seta pale brown, other setae yellow to whi- te; 3 frontal setae, 2 orbital setae, 1 ocellar seta, 1 medial and 1 lateral vertical seta, 1 postocellar seta; row of postocular setae white; no medial postocellar seta. THORAX. Scutum, scutellum and pleura ash-grey microtrichose; ground colour of notopleura, postpronotum and pleura partly yellow; scutum covered with short, white setulae; all setae pale brown, only posterior notopleural seta white; dorsocentral seta almost at level of suture; chaetotaxy as usual in the genus. Less. Yellow; foretarsus without modifications or modified setulae in both Sexes. WING (Figs 60-63). R4+5 bare on both sides; posterodistal extension of cell bcu indistinct; pattern brown, but basal crossband often paler, almost yellow; no bulla; 1022 B. MERZ & H. A. DAWAH basal crossband narrow, stripe-like, sometimes interrupted on one or both sides of obtuse angle on level of M; both crossveins bordered by brown ray; hyaline spot at tip of R2+3 present or absent; small hyaline spots in dark patch of r2+3 and r4+5 some- times present (Fig. 61); hyaline spot in r4+5 just anterior to DM-Cu always present; at tip of R4+5 sometimes with small dark, isolated spot; brown ray in dm just distal to of R-M reaching CuAl (Figs 62-63) or reduced (Fig. 61), rarely absent. No sexual dimorphism in wing pattern. Halter and calypter yellow. ABDOMEN MALE. Mat orange-brown, in middle rarely grey microtrichose. Sternite 5 (Fig. 64) concave distally, with 1 lateroapical seta; epandrium yellow, ovoid (Figs 65-66); prensisetae subequal, claw-shaped; glans (Fig. 67) with sclerotized hook dorsally and small sclerotization basally; vesica small. ABDOMEN FEMALE. Oviscape as long as last two tergites of preabdomen combined, black, at base with white setulae; aculeus as in Figs 68-69, apically rounded, without step. Distribution. Cape Verde Islands, Eritrea, Yemen. New for Saudi Arabia. Host plants. They are unknown for T. repleta. Taking into account the very limited host plant range of species of the group of species formerly placed in the genus Urelliosoma which is restricted to Launaea (Asteraceae, Cichorioideae) it is assumed that 7. repleta attacks the same plant genus. According to Kilian (in litt., 17.V.2004) a possible host is Launaea intybacea, whose distribution matches the distribution of the fly. This plant forms a monophyletic species group with Launaea nudicaulis, which is the only known host plant of 7. pulcherrima, probably the sister species of T. repleta (see below). Pest status. Not a pest species. Remarks. The specimens from Saudi Arabia are superficially very similar to specimens of Trupanea pulcherrima (Efflatoun, 1924) from Israel, but they key out as T. repleta in Munro (1964). The latter 1s a very poorly known species. The original description and illustration of Bezzi (1918) are insufficient and misleading because they are based on one male only and compared with an Indian species which belongs to another species group. Frey (1958) reports on the variability of the wing pattern in specimens from the Cape Verde Islands, and Munro (1964) who did not see the holo- type, did not add further characters. Judging from the illustration of the wing in Bezzi (1918) it seems reasonable to assume that 7. repleta belongs to the group of species for- merly placed in Urelliosoma Hendel, 1927, a genus which was synonymized with Trupanea by Merz (1999) and which includes also T. pulcherrima. In order to clarify the status of the two taxa, the holotype of T. aucta var repleta Bezzi and the specimens mentioned by Frey (1958) were studied, as well as numerous specimens of T. pulcher- rima (see below). It is concluded here that both taxa represent distinct species which differ in morphology, distribution, and probably also in their biology. Trupanea pulcherrima (Efflatoun, 1924) Tephritis pulcherrima Efflatoun, 1924: 100. Syntypes: 75 8,22 2, EGYPT: Wadi Hoff; 7th Tour, Suez Road; Ezbet-el-Naghl (Plant Protection Department, Ministry of Agriculture, Dokki, Cairo, Egypt; Entomological Society of Egypt, Cairo; for details see Norrbom et al., 1999a) (not examined). FRUIT FLIES FROM SAUDI ARABIA 1023 Fics 59-63 Trupanea repleta Bezzi, 1918: 59, head, lateral view; 60, wing male (holotype); 61 wing male (Abha); 62, wing female (Abha); 63, wing male (Abha). Material. ISRAEL: 26 4, 19, Hameshar, 16.VI.1986, A. Freidberg; 19, Mishor Paran, 70 km N Elat, 17.11.1995, Merz; 48 4, 29 9, Nahal Hiyyon, 13.1V.1992, Merz & Freidberg; 16, Nahal Zofar, 11.1V.1992, Merz & Freidberg; 16, 19, Dead Sea, Qalya (= Kallia), 7.VI.1996, Merz & Freidberg (all MHNG). TUNISIA: 19, 24 km S Gabes, 33.42N/10.00E, 8.VI.2000, Schmid-Egger; 19,5 km W Douz, 33.29N/8.59E, 21.III.2001, Schmid-Egger (both MHNG). Distribution. Iran, Tunisia, Egypt, and Israel. Recorded from Saudi Arabia by Freidberg & Kugler (1989). This last record should be revised because of its confusion with 7. repleta. Host plants. Launaea nudicaulis (Asteraceae) (Freidberg & Kugler, 1989). The distribution of the plant corresponds perfectly with the known distribution of 7. pul- cherrima. Remarks. The descriptions of Efflatoun (1924), Hendel (1927) and Freidberg & Kugler (1989) are very comprehensive and the species is therefore not redescribed here. Colouration and structure of head, thorax, abdomen and terminalia of both sexes are the same for 7. pulcherrima and T. repleta. Differences between the two species concern the wing pattern. In 7. repleta the basal dark crossband (from the tip of Sc to the tip of Al+CuA2) is narrow, often paler than the remaining pattern, and in some 1024 B. MERZ & H. A. DAWAH i ots Ste IDS AT y | FIGS 64-69 Trupanea repleta Bezzi, 1918 (64-67, male; 68-69, female, specimens from Abha): 64, abdo- minal sternites 4 and 5; 65, epandrium, lateral view; 66, same, caudal view; 67, glans; 68, aculeus; 69, tip of aculeus. specimens broken in the middle. Usually this band is oblique and directed towards R- M until M, forming a very obtuse angle and parallel to DM-Cu until wing margin. In T. pulcherrima this basal crossband is much wider and its basal border is almost straight from C to the hind margin of the wing without distinct angle at level of M. Moreover the enitre pattern is uniformly brown, or only slightly paler in the stigma. Trupanea sp. 1 Material. 2: 388,299. Remarks. Trupanea is one of the largest genera of Tephritidae and is known from all major biogeographical regions, with 40 species known from the Afrotropical Region (Munro, 1964) and about 12 species from the Palaearctic Region (Norrbom ef al., 1999b). The 5 specimens belong probably to a species close to the T. stellata sub- group as defined by Munro (1964). However, they do not fit the description of any species in this monograph nor any other description available to us. Taking into account the large size of the genus, the small differences between species and the unknown variability in many species we are not describing this species here. FRUIT FLIES FROM SAUDI ARABIA 1025 Trupanodesis sp. | Material.2:1¢. Remarks. The genus was proposed by Merz (1999) for Trupanea aurea Bezzi, 1924 (type species) and an undescribed species from Kenya based on a cladistic ana- lysis. The specimen from Saudi Arabia may be the same species as the undescribed species from Kenya. However, as only one specimen is available, this identification is tentative. Distribution and host plants. This is an Afrotropical genus with both species assigned to it living in flowerheads of Vernonia (Asteraceae, Vernonieae) (Merz, 1999). DISCUSSION In the course of a survey of the insect fauna of the Aseer (= ‘Asir ) area in south- western Saudi Arabia 51 species of Tephritidae were found, of which 42 species could be named (Tab. 1). Six of them are new to science, and 9 species could not be identi- fied safely to species level. Previously, 22 species were recorded from this country, and the presence of half of them could be confirmed in this study. For 11 species we could not find specimens in the Malaise traps during this survey. In total, 62 species of Tephritidae are currently known to occur in Saudi Arabia. This diversity matches well with other countries of the Near and Middle East except for Israel which is much better known through the collecting efforts of A. Freidberg in the last 35 years in all parts of the country (Freidberg & Kugler, 1989). No comparative effort has been conducted for other countries in the region, and it is thus likely that their real species number is much higher. For instance, 17 species recorded from the southernmost part of Israel and the Sinai mountains in Egypt have not yet been found in Saudi Arabia, but most of them should also occur in the mountains next to the Gulf of Aqaba in the northwestern part of Saudi Arabia. Examples are Capparimyia savastani (Martell), Euarestella kugleri Freidberg and E. pninae Freidberg, Goniurellia lacerata (Becker) and G. longicauda Freidberg, Notomma mutilum (Bezzi) and Trupanea desertorum (Efflatoun). The present study is interesting from a biogeographical point of view: Over two thirds of the species of the Aseer area are of Afrotropical origin. This means that these species have a predominantly Afrotropical distribution, or most species of the genus are Afrotropical. Four species are typical for desert areas in the Near and Middle East (e. g., Metasphenisca negeviana, Trupanea pseudoamoena), and only one species, Ensina sonchi, is a typical Palaearctic species. Four species are widespread in the Old World (e. g., Dioxyna sororcula, Spathulina acroleuca), and one species, Bactrocera zonata, 1s a recent introduction from the Oriental Region. On the other hand, typical Oriental species are not present in the Aseer area nor elsewhere in Saudi Arabia. Interestingly enough, some genera and tribes which are widespread in the Palaearctic Region are not yet known in Saudi Arabia, such as Terelliini, Myopitini, or the large genus Tephritis. They are all known from desert localities in Israel, Jordan and Syria (Freidberg & Kugler, 1989; Korneyev & Dirlbek 2000), and their occurrence in the North of Saudi Arabia may be expected. In contrast, Israel has a higher proportion of Palaearctic species, with only about 20% of Afrotropical elements (Freidberg, 1988). Although a remarkable diversity of Tephritidae was collected by a Malaise trap it is generally known that traps are not the best collecting method for Tephritidae. 1026 B. MERZ & H. A. DAWAH Sweeping over potential host plants and rearing Tephritids from infested plants yield usually a much higher number of species. Consequently, it may be assumed that the real number of species in Saudi Arabia may be doubled by using better collecting techniques and by collecting in the northern, palaearctic part of the country. It can be seen from Tab. 1 that 10 species recorded from Saudi Arabia are pest species according to White & Elson-Harris (1992) and require special attention in fruit plantations. Only 4 of these species were found in the present survey: Carpomyia incompleta, Bactrocera zonata, Dacus frontalis, and D. vertebratus, and all of them only in small number of specimens. No new pest species was discovered. But other pest species attacking Cucurbitaceae, such as Bactrocera cucurbitae or Dacus ciliatus, should also occur in the Aseer area. Probably this is again a question of the collecting method. Monitoring those species needs specific collecting methods (e. g., traps with cue lure or methyl eugenol for Dacini; see Munro, 1984). ACKNOWLEDGEMENTS Our sincerest thanks go to John Deeming (Cardiff, Wales) who initiated this study, established the contact between the authors, arranged the loans of specimens to BM, gave important information, and reviewed the manuscript. The generous help of Amnon Freidberg (Tel-Aviv, Israel) is thankfully acknowledged. He was loaning and donating numerous specimens, helped us with fruitful discussions and was reviewing an earlier version of the paper. For help with identifications and useful comments we are indebted to Marc De Meyer (Tervuren, Belgium), David Hancock (Cairns, Australia), and Ian White (London, United Kingdom). For loan of specimens we express our sincerest thanks to Fabrizio Rigato (Milano, Italy), Gunilla Stahls-Mäkäla (Helsinki, Finnland) and Hans-Peter Tschorsnig (Stuttgart, Germany). A special thank is sent to Norbert Kilian (Botanical Garden and Botanical Museum, Free University, Berlin, Germany) for information about the plant genus Launaea. Finally, we would like to thank Florence Marteau (Geneva) for technical help. REFERENCES AL-AHMADI, A. Z. & SALEM, M. M. 1999. Entomofauna of Saudi Arabia (Part 1). King Saud University, Academic Publishing & Press, 240 pp. ANONYMOUS 2002. 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An analytical study in Bio-Taxonomy. Department of Agricultural Technical Services, Republic of South Africa, Entomology Memoirs 8: 1-101. Munro, H. K. 1984. A taxonomic treatise on the Dacidae (Tephritoidea, Diptera) of Africa. Department of Agriculture and Water Supply, Republic of South Africa, Entomology Memoir 61: 1-313. NORRBOM, A. L., CARROLL, L. E. & FREIDBERG, A. 1999a. Status of Knowledge (pp. 9-47). In: THOMPSON, F. C. (ed.). Fruit Fly Expert Identification System and Systematic Information Database. Myia 9: 1-524. NORRBOM, A. L., CARROLL, L. E., THOMPSON, F. C., WHITE, I. M. & FREIDBERG, A. 1999b. Systematic Database of Names (pp. 65-299). In: THOMPSON, F. C. (ed.). Fruit Fly Expert Identification System and Systematic Information Database. Myia 9: 1-524. SHALABY, F. 1961. A preliminary survey of the insect fauna of Saudi Arabia. Bulletin de la Société Entomologique d’Egypte 14: 211-228. STUCKENBERG, B. R. 1999. Antennal evolution in the Brachycera (Diptera), with a reassessment of terminology relating to the flagellum. Studia Dipterologica 6 (1): 33-48. WHITE, I. M. & ELSON-HARRIS, M. M. 1992. Fruit Flies of Economic Significance: Their Identification and Bionomics. CAB International and ACIAR, London, XII & 601 pp. WHITE, I. M., HEADRICK, D. H., NORRBOM, A. L. & CARROLL, L. E. 1999. 33. Glossary (pp. 881- 924). In: ALUJA, M. & NORRBOM, A. L. (eds). Fruit Flies (Tephritidae): Phylogeny and Evolution of Behaviour. CRC Press, Boca Raton, London, New York, Washington D. C., 944 pp. REVUE SUISSE DE ZOOLOGIE 112 (4): 1029-1044; décembre 2005 Proxiandrena subgen. nov. und Revision der west- und zentralpaläarktischen Arten der Andrena proxima-Gruppe (Hymenoptera, Apidae) Christian SCHMID-EGGER Kirchstr. 1, D-82211 Herrsching, Deutschland. E-mail: christian@schmid-egger.de Proxiandrena subgen. nov. and revision of the Andrena proxima group in the western and central Palaearctic Region (Hymenoptera, Apidae). - The former Andrena proxima group is revised for the western and central Palaearctic Region. Proxiandrena subgen. nov. is described for the here treated species with Andrena proxima as type species. Proxiandrena includes 5 valid species, which had been treated as subspecies of Andrena proxima s.1. before: Andrena ampla Warncke in Morocco, Spain, southern France, the western Alpes, and — separated from this area — in Central Asia north of Turkmenistan; Andrena alutacea Stoeckhert with a range from southeast Germany and eastern Switzerland to the eastern Mediterranean area and Turkmenistan; Andrena aspericollis Pérez from Algeria; Andrena bernicla Warncke from Turkey; Andrena proxima (Kirby) from Central Europe and Italy. A key to the species and diagnostic characters are given and distribution of the species is discussed. The female of A. bernicla is unknown and its status as a species remains doubtfull. Keywords: Apidae - Andrena proxima - new subgenus - taxonomy - revision - faunistics - systematics. EINLEITUNG Andrena proxima (Kirby, 1802) ist eine in Mitteleuropa weitverbreitete und stellenweise häufige Sandbienenart. Stoeckhert (1942) beschrieb die sehr ähnliche alu- tacea aus Mittel- und Südosteuropa. Neben morphologischen Unterschieden berief sich Stoeckhert dabei hauptsächlich auf die unterschiedliche Flugzeit der beiden Arten. Ihr Status ist seither jedoch umstritten, da alutacea in der Folge von verschiedenen Autoren nicht anerkannt wurde (Warncke, verschiedene Arbeiten; Dylewska 1987, Gusenleitner & Schwarz 2002). Erst Schmid-Egger & Scheuchl (1997) führten die Art wieder in ihrem Bestimmungsschlüssel der mitteleuropäischen Andrena-Arten auf. Die Autoren wiesen außerdem auf ein drittes Taxon der Artengruppe aus den Westalpen hin, welches sie ebenfalls für eine eigenständige Art hielten. In Südeuropa sind drei weitere Formen aus der proxima-Gruppe beschrieben, nämlich proxima ssp. bernicla Warncke, 1975 aus der Türkei, proxima ssp. ampla Warncke, 1967 aus Spanien und proxima ssp. aspericollis Perez, 1895 aus Algerien. Eine weitere Form, die von Warncke als Unterart ausgezeichnet ist, aber nicht beschrieben wurde, befindet sich in seiner Sammlung im Oberösterreichischen Landes- museum Linz. Manuskript angenommen am 02.03.2005 1030 C. SCHMID-EGGER Ziel der vorliegenden Revision ist es, die genannten Formen zu untersuchen und ihren Status abzuklären. So war insbesondere die Frage zu klären, ob die beschriebe- nen Taxa Unterarten, Arten oder infrasubspezifische Formen sind. Gleichzeitig erwies es sich als notwendig, fiir die Artengruppe um Andrena proxima eine neue Untergattung zu beschreiben, da die bisherige Lösung, die Arten in die Untergattung Micrandrena Ashmead, 1988 zu stellen, mehr als unbefriedigend ist. Die Klärung der verwandschaftlichen Beziehungen innerhalb der untersuchten Artengruppe war nicht möglich, da die sichtbaren Unterscheidungsmerkmale dafür nicht ausreichend sind. UNTERSUCHTES MATERIAL Das untersuchte Material stammt aus den folgenden Sammlungen (mit den im Text verwendeten Abkürzungen): ETH Sammlung der Eidgenössischen Technischen Hochschule in Zürich, Schweiz GRU coll Griinwaldt, in der Zoologischen Staatssammlung Miinchen, Deutschland Kraus Privatsammlung Dr. M. Kraus, Niirnberg, Deutschland MHNG Muséum d’histoire naturelle, Geneve, Schweiz OLL coll. Öberösterreichisches Landesmuseum Linz (vor allem coll. Warncke), Osterreich ORT Privatsammlung Sabine Oertli, Ziirich, Schweiz Scheuchl Privatsammlung Erwin Scheuchl, Velden, Deutschland SE Privatsammlung Dr. Christian Schmid-Egger, Herrsching, Deutschland ZSM Zoologische Staatssammlung Miinchen, Deutschland Für die Schweiz werden die Kantone gemäss den Autokennzeichen folgender- massen abgekürzt: BE = Bern; GR = Graubünden; TI = Tessin; VS = Wallis. ERGEBNISSE PROXIANDRENA SUBGEN.N. TYPUSART. Melitta proxima Kirby, 1802 (= Andrena proxima). DIAGNOSE UND BESCHREIBUNG: Diagnostische Merkmale sind hervorgehoben: WEIBCHEN: Behaarung und Körpergröße: 9-12 mm. Körperfarbe schwarz, Tarsen manchmal rötlich aufgehellt. Körperbehaarung grau, weißlich oder gelblich, Tergitendbinden schneeweiß, grau oder schwach gelblich. Kopf und Thorax struppig abstehend behaart, Haare im Mittel doppelt so lang wie Scapus. Kopf: Labrum doppelt so breit wie lang, unten seicht ausgerandet. Clypeus auf gesamten Oberfläche dicht und grob quer gerunzelt. Gesicht zwischen Ocellen und Clypeus längs gerunzelt, bzw. wabenartig skulpturiert. Foveae schmal, auf Höhe der Fühlereinlenkung etwa 0,28 x so breit wie halbe Gesichtsbreite (Gesichtsbreite ohne Auge gemessen). Fühlerglied IV halb so lang wie Fühlerglied III. Thorax: Mesonotum im vorderen Teil quer runzlig, auf der übrigen Oberfläche grob und dicht punktiert (Punktabstand = 0,5-1 Punkdurchmesser), Punktzwischen- räume glänzend. Mesopleuren mit grob wabenartiger Skulptur. Oberseite des Prodo- PROXIANDRENA SUBGEN. NOV. 1031 deum grob netzartig skulpturiert, das dreieckige Mittelfeld ist mehr oder weniger deutlich mit einer feinen Linie von den ansonsten gleich skulpturierten Seitenfeldern abgegrenzt. Propodeumseitenfelder bedeckt mit sternformig zusammenlaufenden Runzeln, die in der Mitte ein Haar tragen. Oberkante und Hinterkante der Seitenfelder mit mittellanger Sammelfranse, die Haare sind kaum gebogen. Hintertibia schlank, mit grauer Sammelfranse. Die Haare auf der Außenseite sind etwa doppelt so lang wie der Durchmesser der Tibia. Flügeladerung dunkelbraun, Flügel leicht graubraun getönt. Abdomen: Tergite spiegelglatt oder fein chagriniert, matt oder glänzend, manch- mal mit öligem Schein. Tergite II-IV mit Endbinden, die durchgehend oder breit unter- brochen sind. Endfranse gelblich oder graubraun, von weißen oder gelblichen längeren Haaren überdeckt. Sternitscheiben grob skulpturiert und punktiert, Sternitdepressionen fein skulpturiert. Sternite II-[V mit abstehenden Endbinden (Winkel von ca. 40 Grad). MANNCHEN: Behaarung und Körpergröße: 8-11 mm, ansonsten wie Weibchen. Kopf: Labrum quer rechteckig, unten seicht ausgerandet. Clypeus dicht und grob quer gerunzelt. Der übrige Kopf grob gerunzelt. Fühlergeißel dunkel oder unten rötlich aufgehellt. Fühlerglied II etwa 1,3x so lang wie Fühlerglied IV. Thorax: Mesonotum vollständig gerunzelt oder punktiert und mit glänzenden Punktzwischenräumen. Mesopleuren mit grob wabenartiger Skulptur, Propodeum mit sternförmigen Runzeln (ähnlich wie beim Weibchen). Abdomen: Tergite glänzend, punktiert oder mit feiner Runzelung. Tergite bis auf eine Art mit grauweißen oder gelblichen Endbinden, die unterbrochen oder durch- gehend sind. Sternite mit undeutlichen Endfransen, Sternit VIII schmal, am Ende verbreitert, seitlich mit langen Haaren, die länger als der Durchmesser des Sternites sind und sich zum Ende zu verkürzen. Genital. Genital von einfachem Bau, Penisvalve sehr schmal, Gonostylus schmal, ohne Besonderheiten. DISKUSSION: Andrena proxima wurde von Stoeckhert (1942) in die minutula- Gruppe und von Warncke (1968) in die entsprechende Untergattung Micrandrena Ashmead, 1899 gestellt. Dylewska (1987) schloß sich dieser Ansicht nicht an, sondern plazierte proxima aufgrund der besonderen Skulptur der Mesopleuren in eine eigene Gruppe, die Andrena proxima-Gruppe. Dieser Ansicht schließen sich auch Dubitzky & Schönitzer (2001) an und berufen sich dabei vor allem auf die sternförmig zusammen- laufenden Runzeln der Propodeumseiten. Dieses Merkmal betrachten sie als Autapo- morphie der Artengruppe. Eine nähere Verwandtschaft sehen die Autoren vor allem zur labialis Gruppe (subg. Holandrena Pérez 1890). Da sich meine Kenntnisse der Gattung Andrena lediglich auf die mitteleuropäischen Arten beziehen, kann ich diesen Fall nicht selbst abschliessend beurteilen, sondern schließe mich dieser Meinung an. Ich stimme mit Dylweska (1987) und Dubitzky & Schönitzer (2001) darin über- ein, daß proxima und ihre Verwandten eine eigene monophyletische Artengruppe innerhalb der Gattung Andrena bilden. Ihre Stellung rechtfertigt die Beschreibung einer eigenen Untergattung: Proxiandrena subgen. n. Stoeckherts und Warnckes An- sicht, daß proxima zur Untergattung Micrandrena gehört, kann ich nicht folgen. Die Übereinstimmung ist eher oberflächlicher Natur (schwarze Arten mit grauer Körper- behaarung und teilweise glänzenden, bzw. punktierten Körperoberflächen, weiße Ter- gitendbinden). Zwei wesentliche Merkmale von Micrandrena, die geringe Körper- größe (< 8 mm) und die dem Stigma stark angenährte Cubitalader I finden sich bei 1032 C. SCHMID-EGGER proxima nicht. Auf der anderen Seite sind die beiden charakteristischen Merkmale von proxima, die typische Skulptur der Mesopleuren und des Propodeums, nirgends bei Micrandrena ausgepragt. BESTIMMUNGSSCHLUSSEL ZU DEN ANDRENA-ARTEN DER UNTERGATTUNG PROXIANDRENA Weibchen 1 Tergite ohne weiße Endbinden. Verbreitung Algerien...... aspericollis Perez - Tergite mit deutlichen weißen, grauen oder gelblichen Endbinden. Ver- breitung2 Marokko, Europa, Äsien..............2...02 Si E 2 2 Mindestens Scheiben und Depressionen der Tergite II-II, für Tiere aus Mitteleuropa auch von Tergit IV gleichmäßig gerunzelt und chagriniert, ohne Glanz. Tergite unpunktiert oder bei manchen Tieren Tergite H-IV seitlich punktiert. Tergitendbinde IV durchgehend (nur bei frischen Tieren sichtbar!). Thorax (in lateraler Sicht) mit einer kurzen schwarzen Unterbehaarung zwischen den langen hellen Haaren. Von Süd- deutschland, Ostschweiz, Südosteuropa, Türkei bis Zentralasien. In der Türkei auch bernicla, die im weiblichen Geschlecht nicht von alutacea ZINCO CRISI RO alutacea Stoeckhert - Mindestens die Tergitbeulen und Teile der hinteren Tergite glänzend. Tergite häufig punktiert, mindestens auf Tergit II und IV seitlich. Tergitendbinde IV bei proxima auch bei frischen Tieren in der Mitte breit unterbrochen, bei ampla können die Tergitbinden durchgehend sein. Mit oder ohne schwarze Unterbehaarung auf dem Thorax ........... 3, 3 Tergitscheiben und Tergitdepressionen fein gerunzelt und mehr oder we- niger glänzend, zum Teil mit öligem Schein. Tergitscheibe H-IV ohne deutliche Punktierung. Tergitbeulen spiegelglatt, glanzend. Thorax (in lateraler Sicht) meist ohne kurze schwarze Unterbehaarung, manchmal ist diese schwach ausgebildet. Dreieckiges Mittelfeld des Propodeum undeutlich zu den Seitenfeldern abgegrenzt. Binden gelblich. Kleiner, 9-10 mm. Verbreitung: Europa außer Iberische Halbinsel, fehlt an- scheinend in den zentralen und westlichen Alpentälern........ proxima (Kirby) - Tergitdepression spiegelglatt, Tergitscheiben H-IV mit deutlicher Punktierung, die der Tergite III und IV gröber als die von Tergit II. Tergitscheiben zwischen den Punkten ebenfalls spiegelglatt, im basalen Teil mit schwachen Runzeln. Thorax (in lateraler Sicht) zwischen der langen hellen Behaarung mit dichter und kurzer schwarzer Unterbe- haarung. Dreieckiges Mittelfeld des Propodeum bei den meisten Tieren mit deutlichen, aber feinem Kiel zu den Seitenfeldern zu abgegrenzt. Binden schneeweiß. Größer, 10-12 mm. Zentrale und westliche Alpen: Aostatal, Südschweiz, Durancetal, Iberische Halbinsel (Verbreitung in Südfrankreich noch unklar), Zentralasien... CRE ampla Warncke Männchen l Tergite ohne weiße Endbinden. Mesonotum auf der ganzen Fläche grob gerunzelt, dazwischen chagriniert. Verbreitung: Algerien . . . . aspericollis Perez PROXIANDRENA SUBGEN. NOV. 1033 Tergite mit deutlichen weiBen, grauen oder gelblichen Endbinden. Tiere aus Marokko: Mesonotum mit glänzenden Punktzwischenräumen. Ver- breitung=) Marokko Europa AS ENT I RE PE anal ae 2 Gesicht dunkelbraun behaart. Türkei ................... bernicla Warncke Gesichthellbehaante os se a A RAI 3 Mesonotum vollständig chagriniert und wabenartig gerunzelt, keine ein- gestochenen Punkte oder glänzende Punktzwischenräume erkennbar. Tergite I-IV: Basis fein gerunzelt, Mittelteil glänzend mit zerstreuten Mikropunkten, Depression fast punktfrei, im apikalen Teil sehr feine Mikropunkte. Tergitendbinden II-IV in der Mitte breit unterbrochen. Körperbehaarung gelblich. Europa außer Iberische Halbinsel, Zentral- asien, fehlt anscheinend in den zentralen und westlichen Alpentälern es Ale I ren POESIA SI ES E proxima (Kirby) Mesonotum mit glänzenden Punktzwischenräumen, die im hinteren Teil 0,5-1 Punktdurchmesser betragen (bei ampla aus der Schweiz vereinzelt stärker chagriniert). Tergite I-IV: Basis und Mittelteil glänzend mit deut- lich eingestochenen Punkten, die im direkten Vergleich mindestens dop- pelt so groß wie bei proxima sind. Depression unterschiedlich. Ter- gitendbinden unterschiedlich. Körperbehaarung weißlich. Die beiden folgenden Arten sind sehr ähnlich und ohne zugehörige Weibchen nur aufgrund ihrer geografischen Verbreitung sicher zu trennen... 4 Verbreitung: Süddeutschland, Ostschweiz, Südosteuropa, Türkei. Tergite im Mittel weniger dicht punktiert, Punkte im direkten Vergleich kleiner als bei ampla. Tergitendbinden III und VI bei frischen Tieren durchgehend u area ra N tar alutacea Stoeckhert Verbreitung: Zentrale und westliche Alpen: Aostatal, Südschweiz, Durancetal, Spanien (Verbreitung in Südfrankreich noch unklar, in der Schweiz Überschneidung mit dem Areal von alutacea), Zentralasien. Tergitendbinden in der Mitte breit unterbrochen, selten Binde IV durch- schendilbeisasiatischenlieren) sa a re ee eos ampla Warncke VERGLEICHENDE BESCHREIBUNG DER ANDRENA-ARTEN DER UNTERGATTUNG PROXIANDRENA Nachfolgend werden die für die Artunterscheidung wichtigen Merkmale be- schrieben und die Arten dabei gegenübergestellt. Bei der nachfolgenden Besprechung der Arten wird lediglich eine kurze Diagnose der Arten gegeben. Behaarung des Mesonotum, Weibchen alutacea Weißlich, in der Länge unregelmäßig, in der Tendenz etwas kürzer als bei proxima. Auf der ganzen Länge des Tergites mit feiner, kurzer dunkler Unterbehaarung. ampla Gelblich, bei spanischen Tieren auch weißlich, die übrigen Merk- male wie bei alutacea. aspericollis Gelblich, lang, ähnlich proxima. proxima Gelblich, in der Länge gleichmäßig, im Mittel etwas länger als bei alutacea und ampla. 1034 Tergitskulptur, Weibchen alutacea / Mitteleuropa alutacea / Tiirkei ampla aspericollis proxima C. SCHMID-EGGER Tergite gleichmäßig fein gerunzelt (chagriniert), matt. Tergitbeulen leicht glänzend, ebenfalls gerunzelt. Glanz nimmt auf Tergit IV leicht zu. Auf den Tergitscheiben undeutliche Punkte erkennbar. Tergitdepressionen in der Tendenz schmaler und weniger stark bo- genförmig als bei proxima, außerdem wirkt der Niveauunterschied zwischen Scheibe und Depression niedriger als bei proxima. Die feine Runzelung der Tergite verschwindet nahezu, Tergite wir- ken glatt. Scheiben und Tergitdepressionen mit identischer Skulptur. Auf den Tergiten I-II ist noch eine feine Chagrinierung erkennbar, die auf den Endtergiten nahezu verschwindet. Tergitscheiben II und IV sehr schwach, klein und zerstreut punktiert. Tergite spiegelglatt und stark glänzend, auf den Tergitscheiben I und III leichte Chagrinierung erkennbar. Tergitscheiben II-IV mit deut- lich eingestochener und teilweise grober Punktierung, Punkte im Mittel 2-3 Punktdurchmesser voneinander entfernt. Bei den spani- schen Tieren kann die Punktierung weniger stark und die Chagri- nierung stärker ausgeprägt sein. An der Tergitpunktierung sind solche Tiere aber immer von proxima und alutacea zu unterscheiden. Tergitskulptur entspricht der von alutacea. Tergite chagriniert, mehr oder weniger stark glänzend, besonders auf den hinteren Tergiten. Die Tergitdepressionen stärker glänzend als die Scheiben, Depression IV manchmal spiegelglatt. Tergitbeulen fast immer spiegelglatt. Tergitendbinden auf den Tergiten II-IV, Weibchen alutacea ampla aspericollis proxima Schneeweiß, anliegend. Sie entspringen etwas oberhalb des Tergitendrandes, so dass sie den Tergitendrand um ihre halbe Länge oder 2/3 ihrer Länge überragen. Binde auf Tergit IV durchgehend, auf Tergit II und III breit unterbrochen. Unterbrechung auf Tergit III entspricht etwa der Breite der Binde (Breite: vom Tergitseitenrand Richtung Tergitmitte), auf Tergit II etwa der doppelten Bindenbreite. Schneeweiss, etwa im Winkel von 30° abstehend (bei den asiatischen Tieren anliegend, und nach hinten gerichtet). Die Binde entspringt etwas tiefer als bei alutacea, aber weiter zur Tergitbasis als bei pro- xima. Binde auf Tergit IV schmale (Ozellenbreit) bis breiter (Länge des Fühlergliedes III) unterbrochen, die anderen Binden breit unter- brochen. Bei asiatischen Tieren Binde auf Tergit IV durchgehend. Keine Binden vorhanden. Gelblich, etwa im Winkel von 30 abstehend. Die Binde entspringt fast unmittelbar am Tergitendrand. Binde IV breit unterbrochen (etwa um Bindenbreite), die übrigen Binden sehr kurz (etwa ein Viertel der Tergitbreite). Skulptur des Mesonotum, Männchen alutacea ampla Deutlich eingestochen punktiert, mit glänzenden Punktzwischen- räumen, die 0,2-1 Punktdurchmesser betragen. Basal zum Teil auch chagriniert, bzw. ohne erkennbare Punkte. Wie alutacea, allerdings mit großer Variationsbreite. Tiere aus dem Aostatal sind sehr deutlich punktiert mit glänzenden Zwischen- räumen, bei Tieren aus der Schweiz ist das Mesonotum manchmal vollständig chagriniert, ohne erkennbare Punkte. aspericollis bernicla proxima PROXIANDRENA SUBGEN. NOV. 1035 Chagriniert, mit körnigen Erhabenheiten oder Runzeln, keine Punkte oder glänzende Flächen erkennbar. Wie aspericollis. Wie aspericollis. Skulptur der Tergite II-IV (Punktzwischenräume bei allen Arten glänzend), Männchen alutacea ampla aspericollis bernicla proxima Tergitscheiben fein eingestochen punktiert, Punktdurchmesser ca. 0,3-0,5 x wie Punktdurchmesser auf Mesonotum, Punktabstand 1-2 Punktdurchmesser. Tergitbasis fein quer chagriniert. Tergitdepres- sion mit zerstreuten, sehr feinen Punkten, die vordere Hälfte der Tergitdepression teilweise punktfrei. Wie alutacea, Punktdurchmesser im Mittel größer (ca. 0,5-0,8 x Punktdurchmesser auf Mesonotum), meist bis zur Tergitbasis glän- zend. Tergitscheibe mit sehr undeutlichen und kleinen Punkten, basal fein quergerunzelt. Tergitdepression nahezu glatt. Tergitscheiben mit feinen Punkten (in der Größe ähnlich alutacea), Punktabstände 1-3 Punktdurchmesser, Tergitdepression mit sehr wenigen zerstreuten Punkten. Tergitscheiben mit sehr feinen und zerstreuten Punkten, Punkt- abstände 2-5 Punktdurchmesser, Tergitdepressionen punktlos oder mit vereinzelten sehr feinen Punkten. Endbinden Tergite II-V Männchen (kann nur bei frischen Tieren beurteilt werden) alutacea ampla aspericollis bernicla proxima Körpergröße alutacea ampla aspericollis bernicla proxima Verbreitung alutacea ampla Tergit III und IV mit (fast) durchgehender Endbinde. Tergit V mit lockerer Endbinde, die aus langen Haaren besteht. Binden schnee- weiß. Tergite [I-IV mit breit unterbrochenen Binden, Binde auf Tergit IV kann jedoch auch durchgehend sein (bei zentralasiatischen Tieren). Tergit V mit undeutlicher durchgehender Endbinde. Binden schnee- weiß, dichter als bei proxima. Die beiden untersuchten Männchen ohne erkennbare Binden. Dies mag jedoch vielleicht auf den Erhaltungszustand der Tiere zurück- zuführen sein. Wie bei proxima, Binden jedoch weiß. Tergitendbinden II-IV immer breit unterbrochen (mindestens auf 1/3 der Tergitbreite). Tergit V apikal ohne erkennbare Endbinde. Binden gelblich. Weibchen Männchen 10-11 mm 9-11 mm 10-12 mm 9,5-11 mm 11-12 mm 8 mm - 8-9 mm 9-10 mm 8-10 mm südöstliches Deutschland, Ostschweiz bis Südosteuropa, Lettland, Türkei, Iran und Turkmenistan. Marokko, Iberische Halbinsel, Frankreich, Südfrankreich, Durancetal; Südschweiz, Italien: Aostatal, Aserbeidschan bis Zentralasien. 1036 C. SCHMID-EGGER aspericollis Algerien. bernicla Türkei (nördliche Hälfte?). proxima Mitteleuropa, Großbritannien, Italien, Südosteuropa bis Griechen- land. Andrena alutacea Stoeckhert, 1942 Andrena alutacea Stoeckhert, 1942: 236-252 (Erstbeschreibung d und ®, Holotypus 2, Typusfundort: Eining/Donau, ZSM, untersucht) — Pittioni & Schmidt, 1943: 33 (Fauna von Österreich, Bulgarien) — Kocourek, 1966: 69 (Fauna Tschechoslowakei, nur in Mähren) — Schmid-Egger & Scheuchl, 1997 67-68, 142 (Schlüssel für Mitteleuropa, Angaben zur Verbreitung und Phänologie) — Mandery, 2001: 116 (Fauna für Franken, aktuelle Funde). Andrena proxima: Warncke, 1967: 209, 250 (synonym mit proxima) — Warncke, 1986: 45 (syn- onym mit proxima) — Westrich, 1984: 23 (synonym mit proxima, mit Paratypus ver- glichen) — Dylewska, 1987: 547 (synonym mit proxima) — Schwarz et al., 1996 50 (synonym mit proxima: „A. alutacea ist entgegen bisweilen geäußerter Vermutungen keine eigene Art“) — Gusenleitner & Schwarz, 2002: 610-611 (Literaturzitate, zu pro- xima gestellt, bzw. als forma von dieser behandelt). UNTERSUCHTES MATERIAL (PT = Paratypus): Holotypus: © mit drei Etiketten: [Eining a.D 24.6.1927 leg. Stoeckhert]; [Type]; [Andrena alutacea ©, det. E. Stoeckhert] (ZSM), Fundort: Eining an der Donau, Bayern, Deutschland. ASERBEIDSCHAN: ® 21.6.1957 Nachitschewan, Chrusa, 2000 m. (OLL). BULGARIEN: ® Pirin Geb. 1000-1800 m, Banska 25.6.1938, PT (ZSM). 2 dg 2 © 30.5.1989 Slancev Briag (OLL). DEUTSCHLAND: Bayern: 2 9 24.06.1932 Eining (1 2 PT); 2 24.6.1946 Markt Schwaben (GRU); 2 © 6.8.1933, 27.7.1933 Pappenheim, PT; 9 21.6.1883 Allach (ZSM); Eining 1.7.1938 (PT); 9 3.8.97 Ergersheim, Eschenau, Weinbergbrache, Daucus carota (Kraus); 2 17.7.2004 Altmühltal, 12 km NEE Eichstätt, Gundoldinger Heide (SE), Thüringen: 9 20.6.98 Kyffhäuser/Auleben/Solberg, Kalk- magerrasen auf Achillea sp; 19 23.6.01 Wanderslebener Gleiche b. Gotha, Apiaceae gestreift; Sd 20.6.98 Kyffhäuser/Badra/Feldrain, Apiaceae gestreift (Burger, die beiden letzten Tiere det. Burger); Baden Württemberg: ? 2.7.1995 Baar/Südostschwarzwald, 2 km N Geisingen, NSG Klause; 3 10.6.1996 Hegau, Singen, Hohentwiel (SE). FÜRSTENTUM LIECHTENSTEIN: 2 13.6.1997 Balzers, Elltal, ein frisches Tier, zusammen mit einer abgeflogenen proxima (ETH). GRIECHENLAND: ® 9.05.1973 Mistras; 9 29.06.1970 Vytina; 13.6.1956 Olymp, Prioni, 1000 m; ® 19.05.1973 Mistria; Mazedonia, Treskaschlucht; 2 8.6.1973 Lehau (GRU). IRAN: © 20.5.1975 Kermanshahan, 80 km W Kermanshah, Paweh (GRU). ITALIEN: 16 2 23.7.- 3.8.1933; 19.8.1926 Rovereto; 9 25.8.1913 Ritten (Südtirol) (ETH); 2 9 16.7.1925 Bozen, PT; bei Bozen, leg Kohl (MHNG). KROATIEN: 2 2 19.7.1971, 2.8.1972 Istria, Ucka (GRU). LETTLAND (Lativa): 2 9.08.1931 Meiten (GRU). MACEDONIEN: ® Cakor-Pass 1969 (ZSM). ÖSTERREICH: © 26.6.1976 Dürnstein, Wachau (GRU), 4 © 22.7.1920 Innsbruck (PT, ZSM); 2 © 24.7.1998 Burgenland, Mühlgraben; 9 24.7.1998 Burgenland, Oberdrosen-Bergen; 2 28.7.1999 Tauka; © 27.7.1996 Steiermark, Katzelsdorf; 9 28.7.1996 Steiermark, Reith bei Hartmannsdorf; © 9.7.1998 Tirol, Igls S. Innsbruck; 2 13.7.1991 Oberösterreich, St. Georgen/Gusen (OLL). SCHWEIZ: 29 26.7.1926 GR, Sent; 2 31.7.-3.8. GR, Brusio; 4 9 23.7.1997, 8.8.1997 GR, Haldenstein, Böfel; © Juli 1915 TI, Lugano, Maroggia, PT (ZSM); © 29.7.1999 TI, Indemini, Paese (ETH) — 3 2 1 g TI, Valle Maggia, Lodano, 340 m, an Anthriscus; 2 1 TI, Valle Maggia, Gordevivo, 310 m (SE). TÜRKEI: ® 8.8.1982 Pr. Hakkari, Sai Dag, Shagulut, SW Yüksekova 1700 m; 9 12.06.1978 Konya, Sille, © 31.5.1959 Amasya, 2 12.6.1976 Mus, 2 4.6.1972 Gürün, 2 20.05.1975 Karakurt, Arastal (GRU), 2 5.6.1945 Gürün; 2 12.6.1978 Konya; 2 31.5.1959 Amaxya, © 12.6.1976 Mut, 9 22.5.1975 Karakurt, Arastal; 3 © Horosan, Arastal; 3 2 1.6.1980 35 km SW Hakkari, d Kars, 20 km W Karakurt 1600 mNN, & 29.5.1980 Agri, 20 km N Patnos, d Nevshehir, Ürgüp 21.8.1971 (OLL). TURK- MENISTAN: 5 2 16.5.1975 Bacharden (GRU). PROXIANDRENA SUBGEN. NOV. 1037 DIAGNOSE: Weibchen: Andrena alutacea ist anhand der gleichförmigen Tergitchagrinierung, den matten Tergiten, der schwarzen Unterbehaarung auf dem Mesonotum und der durchgehenden Tergitendbinde IV von den verwandten Arten zu unterscheiden. Männchen: Das Männchen ist im direkten Vergleich kaum von ampla zu trennen. Von den anderen Arten läßt sich alutacea anhand der glänzenden Punktzwischenräume auf dem Mesonotum und der weißlichen Körperbehaarung sowie den — bei frischen Tieren — durchgehenden hinteren Tergitendbinden abtrennen. In Mitteleuropa läßt sich alutacea ebenfalls anhand der späten Flugzeit von pro- xima abgrenzen. Männchen treten in der Regel ab Mitte Juni, Weibchen ab Anfang Juli auf. Um diese Zeit gibt es praktisch keine frischen proxima mehr. VARIATION: Weibchen aus der Türkei weisen teilweise eine viel schwächer aus- geprägte Tergitchagrinierung auf als die mitteleuropäischen Tiere. Sie erscheinen dadurch stärker glänzend und sehr viel feiner skulpturiert. Allerdings ist die Variations- breite bei türkischen Tieren relativ hoch. Diese Merkmalsausprägung ist auch bei ein- zelnen Tieren aus Griechenland zu beobachten, so dass hier wahrscheinlich eine kontinuierliche Merkmalsverschiebung stattfindet, die der Variationsbreite der Art zuzurechnen ist. Allerdings sind Weibchen aus Turkmenistan wiederum stärker chagri- niert und weisen außerdem zum Teil eine relativ grobe, fast schon körnige Tergit- punktierung auf den Tergiten II-IV seitlich auf. DISKUSSION: Andrena alutacea wurde von Stoeckhert (1942) unter besonderem Hinweis auf die unterschiedliche Flugzeit von proxima abgetrennt und als eigene Art beschrieben. Dennoch zog Warncke (1967) die Art ohne jede Begründung ein, weitere Autoren folgten ihm (Dylewska, 1987; Gusenleitner & Schwarz, 2002). Erst Schmid- Egger & Scheuchl (1997) betrachteten alutacea mit Hinweis auf die Stoeckhertsche Beschreibung und aktuellem Material wieder als selbständige Art. Eine genaue Prüfung des vorliegenden Materials zeigt, daß sich alle Weibchen, die in Mitteleuropa im Juli und August gefangen wurden, auch morphologisch gut von Tieren unterscheiden, die aus dem Mai und Juni stammen. Das bestätigt meiner Meinung nach die Ansicht von Stoe ckhert, daß alutacea eine valide Art ist. Die Unter- scheidungsschwierigkeiten zu proxima liegen im Bereich dessen, was bei vielen ‚schwierigen’ Andrena Artengruppen (zZ. B. Micrandrena) üblich ist. Probleme können bei abgeflogenen Tieren in Griechenland auftreten, da dort die Variation von alutacea beträchtlich zunimmt. Auch wenn man die bestehende Ähnlichkeit der beiden Arten in Betracht zieht, würde eine Synonymisierung von alutacea mit proxima die unterschiedliche Phänologie keinesfalls auf befriedigende Weise erklären. Für eine partielle zweite Generation ist der zeitliche Abstand beider Taxa sehr kurz (sie überlappen sich in der Flugzeit um 1-2 Wochen), außerdem müssten dann beispielsweise im wärmebegün- stigten und gut besammelten Baden-Württemberg mehr Tiere von alutacea auftreten, was aber nicht der Fall ist (eigene Beobachtung, Doczkal mündl.). Daher erscheint das Konzept zweier valider Arten die wahrscheinlichere Lösung für den vorliegenden Fall. Anders verhält sich die Situation in Südosteuropa und vor allem in der Türkei. Während in Griechenland alutacea etwas später als proxima fliegt, wurden alle alu- tacea aus der Türkei, aber auch in Südosteuropa, jeweils im Mai und Juni gefangen (aus der Türkei liegen keine Tiere vor, die proxima entsprechen). Einzige Ausnahme 1038 C. SCHMID-EGGER bilden ein Männchen und ein Weibchen aus der Türkei, die aus dem August stammen. Das Männchen aus Ürgüp besitzt eine sehr weitläufige Mesonotumpunktierung, die viel weiter als bei allen anderen Männchen ist (Punktabstände zum Teil mehrere Punktdurchmesser groß). Das Weibchen aus der Provinz Hakkari ist völlig frisch, besitzt eine vollständige Endbinde auf Tergit IV und hat eine insgesamt etwas kürzere und wie geschoren wirkende Körperbehaarung. Ansonsten stimmt es vollständig mit alutacea überein. Ob hier eine partielle zweite Generation vorliegt oder sich hinter alu- tacea noch weitere Arten verbergen, muss derzeit offen bleiben. VERBREITUNG: Andrena alutacea ist von Mitteleuropa über Südosteuropa bis in die Türkei, den Iran und Turkmenistan verbreitet. In Deutschland kommt sie in Bayern und dem Osten von Baden-Württemberg vor, die meisten Funde stammen dabei aus dem Einzugsbereich der Donau. Ein Einzelfund stammt aus dem südlichen Rheintal (Grißheim; Stoeckhert, 1942). Außerdem liegen verschiedene Funde aus Thüringen vor. In der Schweiz ist alutacea im Tessin sehr häufig, weitere Funden stammen aus der Ostschweiz (Graubünden) und aus Liechtenstein. Der nördlichste Nachweis stammt aus Lettland. Das Verbreitungsgebiet von alutacea überschneidet sich in weiten Teilen mit proxima, an mehreren Orten konnten beide Arten sogar zusammen gefangen werden. Stoeckhert (1942) meldet sie aus Deutschland: Bayern; Thüringen: Jena, Erfurt; Baden-Württemberg: Grißheim in Südbaden. Österreich: viele Funde. Polen: Krakau, Lemberg (Galizien). Schweiz: Lugano. Italien: Bozen, Ligurische Alpen, Turin; Trient, Fiume. Kroatien. Rumänien: Szankesd (Siebenbürgen). Bulgarien. Griechenland: Parnass. Georgien: Kaukasus, Borsholm bei Tiflis (dieser letzte Fund bezieht sich höchstwahrscheinlich auf bernicla). ÖKOLOGIE: Nach bisherigen Erkenntnissen ist alutacea wie ihre Schwesterart proxima oligolektisch auf Doldenblütler spezialisiert (Westrich 1989, der die Oligolektie von proxima feststellte, trennt nicht zwischen den beiden Taxa). Andrena alutacea besiedelt in Deutschland vielfach trockenwarme Magerwiesen. Im Tessin (Schweiz) fing ich die Art auf einer typischen ‚Fettwiese’ auf Anthriscus sylvestris und im Altmühltal (Bayern) am Rande einer Wachholderheide auf Daucus carota. Auf- grund der wenigen Funde und der Spezialisierung auf trockenwarme Offenstandorte müßte die Art in Deutschland in den betreffenden Bundesländern in die Rote Liste aufgenommen werden. Andrena ampla Warncke, 1967, stat. nov. Andrena proxima ssp. ampla Warncke, 1967: 229, 250: Erstbeschreibung d und 9 , Holotypus d; Typenfundort: Spanien, Alberche, OLL, untersucht. — Warncke, 1974a: 11 (Fauna Marokko) — Warncke, 1976: 151: (Fauna Iberien) — Erlandsson, 1979: 125 (Fauna Spanien) — Gusenleitner & Schwarz, 2002: 611-612 (Katalog, S. 1145 Verbreitungskarte von ampla). UNTERSUCHTES MATERIAL: Holotypus: © mit drei Etiketten: [Alberche V 1909 Arias], [Andrena proxima ssp. ampla War. det. Dr. Warncke], mit rotem Karton: [Holotype], in coll. OLL. ASERBEIDSCHAN: © 21.6.1957 Nachitachawan, Chrusa (GRU). FRANKREICH: © 29.6.1978 Provence, Mt. de Lure, Cruis, ca. 500 m (GRÜ); 4? 23.8.1998 Ht. Alpes, Aiguilles, le Lombard 1800 m; 12 © 6.6.1997 Ht. Alpes, 18 km S Briancon, Gouffre 900 m, 9 6.6.1997 Ht. Alpes, 6 km SW Briancon, Prelles 1200 m (CS). ITALIEN: Westalpen, Valle d’Aosta: 22 9 8 6 7.7.1995; 29 16.7.1995; 2 2 19 d 27.6.1996, 4 2 2.8.1997 Cognetal, Lillaz, 1700-1800 m, an Lasiopitium aff. siler L.; 3 16.5.1996, St. Pierre 8000 m; 19 10 km E Aosta, Trois Villes, PROXIANDRENA SUBGEN. NOV. 1039 1400 m NN, 16.5.1998 (CS). KASACHSTAN: 2 16.6.79 Alma Ata, Alataugeb. 1900m (Kraus) - Axaj bei Alma Ata, 19.6.1981 (OLL) (Etikettiert als Andrena kocoureka ssp. n., det Warncke Paratypus]. KIRGISIEN: 9 © 15.8.1998 Alaj, G.K. Gultscha Tal, Ak-Bosogo (GRU) — 2 d 1 9 7.6.1998 Sandalach-Gebirge, Kain-Su Schlucht. 2000 m NN (Scheuchl). MAROKKO: % 13.5.1965 Azrou (proxima det Warncke) (OLL). SCHWEIZ: ® 29-31.7.1933 GR, Braggio-S- Marina, Calanca, 1000-1600 m leg. Nadig; © 24.6.1934 GR, Rabius; 9 24.6.1885 VS, Sierre; 3d 19.8.1943 VS, Follatéres; 2 26.7.1991 VS, Visperterminen 1380 m; ® 1.6.1991 VS, Hohtenn; (ETH) — 65 2 47 © , Fangjahre 2001-2002 VS, Erschmatt: Undere Chastler, 1210 m; Oberu Castler 1270 m; Oberi Zaig 1450 m; Chritzmatte 1220 m; Rossuwald 1370 m; Rüemetschbodu 1530 m; Chritzmatte 1120 m, Ritin 1190 m; Platten 890 m, 4 15.5-2.6., 2 16.5.-27.6. (ORT) — 9 21.6.1998 VS, Jeizinen 1500 mNN, felsige Bergwiese (Burger, Herrman); 4 2, VS: Sierre, Martigny, St. Luc, Niouc, alle vor 1900 (MHNG). SPANIEN: © Sierra Guadarrama, Dusmet; 4 Alberche V 1909 (leg) Arias; Montarco (leg) Dusmet, alles Paratypen det. Warncke (OLL) - 2 Villaviciosa, 2 2 d Escorial (alle det K. Warncke als proxima ssp. am- pla), 2 11.7.1977 Pr. Guadalajara, Tierzo bei Molina, 1100 m; ? 10.7.1975 Malaga, Ronda (GRU). TADSCHIKISTAN: © Dusambe, 40 km vych. Javros, 24.6.1981 (OLL) (Etikettiert als Andrena kocoureka ssp. n., det Warncke Holotypus]. DIAGNOSE: Andrena ampla ist im weiblichen Geschlecht wie folgt charakteri- siert: Tergite überwiegend spiegelglatt, Tergitscheiben II-IV deutlich und grob punk- tiert, Punkte gleichmäßig über die Tergitscheibe verteilt. Mit alutacea teilt die Art die schwarze kurze Unterbehaarung des Mesonotums. Andrena ampla ist durchschnittlich 1 Millimeter länger als alutacea (11-12 mm). Andrena alutacea Weibchen aus Turkmenistan, die auf den hinteren Tergiten seitlich ebenfalls grob punktiert sind, unter- scheiden sich auch hier durch die feine Chagrinierung auf den Tergiten. Die Männchen sind sehr ähnlich wie alutacea. Von dieser unterscheiden sie sich durch eine geringfügig dichtere und stärkere Tergitpunktierung sowie Tergitendbinden, die auf den Tergiten III und IV in fast allen Fällen unterbrochen sind (bei alutacea durchgehend). Andrena pro- xima läßt sich im männlichen Geschlecht leicht aufgrund des chagrinierten Mesonotums und der nur sehr fein und zerstreut punktierten Tergite unterscheiden. VARIABILITÄT: Die Tiere aus Zentralasien stimmen mit den europäischen Indivi- duen fast vollkommen überein. Die Tergitendbinden sind bei zwei vollständig erhal- tenen Weibchen nach hinten gerichtet und anliegend, während sie bei den europäischen Individuen abstehend und nach außen gerichtet sind. Die Tergitendbinde IV ist bei den asiatischen Tieren durchgehend, während sie bei den europäischen Individuen fast immer unterbrochen ist. Außerdem ist die dichte Tergitpunktierung bei einigen der asiatischen Weibchen körnig-dicht, die Punkte sind nur von schräg hinten zu erkennen. DISKUSSION: Andrena ampla wurde von Warncke als Unterart zu proxima von der iberischen Halbinsel beschrieben. Laut seiner Verbreitungskarte (publiziert in Gusenleitner & Schwarz, 2002) stellen die Pyrenäen die Verbreitungsgrenze zwischen der ssp. ampla und der ssp. proxima dar. Durch die aktuellen Funde von ampla in den Alpen wird diese Einteilung hinfällig. Außerdem ist das Taxon von den beiden Arten alutacea und proxima morphologisch gut unterschieden. In der Flugzeit liegt ampla etwa zwischen proxima und alutacea. Somit halte ich seinen Status als Unterart zu proxima für unwahrscheinlich und betrachte ampla als valide Art. Andrena ampla ist außerhalb der iberischen Halbinsel nur aus den trocken- warmen großen Binnentälern der Süd- und Südwestalpen nachgewiesen (Wallis, Graubünden, Tessin, Aostatal, Durancetal, bei Genf). Weiterhin liegt ein Fund aus der südlichen Provence vor. Es ist zu vermuten, dass die Art in Südfrankreich weiter verbreitet ist. Wie sie sich dort zu proxima abgrenzt, ist noch unbekannt. 1040 C. SCHMID-EGGER In der Schweiz überlappen sich die Areale der drei mitteleuropäischen Arten der Untergattung Proxiandrena Allerdings sind auch hier Unterschiede in der Verbreitung feststellbar. So ist das Wallis (wie das ummittelbar südlich daran angrenzende italie- nische Aostatal) nach den bisherigen Daten nur von ampla besiedelt. Diese dringt jedoch auch noch weiter nach Südosten bis Graubünden und dem Tessin vor und findet sich auch in der Umgebung von Genf. Andrena alutacea besiedelt vor allem die Ostschweiz, bzw. Lichtenstein, ihr Verbreitungsgebiet erstreckt sich dabei vom Rheintal bis ins Tessin. Andrena proxima hingegen ist vor allem auf die Nordschweiz beschränkt, kommt aber stellenweise auch in den südlichen Landesteilen vor. Eine ähnliche Verbreitung wie ampla weist Andrena afrensis Warncke, 1967, auf, die in Spanien, dem Wallis (Westrich er al., 1997) und im Aostatal (Schmid-Egger, unpubl.) vorkommt. Die Schwesterart von afrensis, Andrena agilissima Scopoli 1770, die in Mitteleuropa weit verbreitet ist, scheint zumindest im Wallis zu fehlen, während im Aostatal beide Arten zum Teil syntop fliegen. Interessant ist weiterhin das Vorkommen von ampla in West- und Zentralasien, welches auf ein disjunktes Areal hinweist. Das Verbreitungsgebiet überschneidet sich dort nach den vorliegenen Daten weder mit proxima noch mit alutacea. Die asiatischen Tiere sind geringfügig von den europäischen unterschieden (runzelig dichte Punktierung auf den Tergiten III und IV lateral, durchgehende Endbinde auf Tergit IV), allerdings rechtfertigen diese Unterschiede keine Abtrennung der östlichen Populationen als Subspecies. ÖKOLOGIE: Im Aostatal wurde ampla auf trockenen, blütenpflanzenreichen, süd- exponierten Almwiesen in Höhen zwischen 1700 und 1800 m ü. NN angetroffen. Die Art flog dort zahlreich auf Laserpitium cf. siler und war unter anderem vergesellschaf- tet mit Osmia dalmatica Morawitz (Apidae), Bembix tarsata Latreille (Crabronidae) oder Parnassius apollo Linné (Lepidoptera). Im Durancetal flog sie in 800-1000 m ii. NN an Wegrändern und auf Wiesen mit magerer, trockenheitsliebender Vegetation. Im Wallis konnte Sabine Oertli die Art ebenfalls in Anzahl auf südexponierten mageren Wiesen und Weiden antreffen. Sie stellte Blütenbesuch auf Anthriscus sylvestris, in geringerem Mass auch auf Heracleum sphondylium und Pimpinella major fest. Auf den Weiden blühte Laserpitium siler (Oertli in litt.). Somit scheint die Art wie auch proxima oligolektisch an Doldenblütlern (Apiaceae) gebunden zu sein. VERBREITUNG: Disjunkt in Europa und Asien. Europa: Südliche Schweiz; Norditalien: Aostatal; Südwestfrankreich: oberes Durancetal; Südfrankreich: Provence; Spanien, Marokko. Nach Warncke (1976) in Spanien und Portugal weit ver- breitet. Asien: Aserbeidschan, Kasachstan, Kirgisien, Tadschikistan. Andrena aspericollis Pérez, 1895, stat. rest. Andrena aspericollis Perez 1895: 37 (Beschreibung ©, d, keinen Holotypus festgelegt) - Paralectotypus untersucht. Andrena proxima ssp. aspericollis: Warncke 1967: 183, 252 (Lectotypus 2 festgelegt, Teniet, Algerien, coll. Perez/Paris, stat. nov. ) — Warncke 1974a: 11 (Fauna Algerien, Flugzeit Mai bis Juni) — Gusenleitner & Schwarz 2002: 610-611, 1145 (Katalog). UNTERSUCHTES MATERIAL: ALGERIEN: © mit unleserlichem Etikett, 2 d mit gedruk- ktem Etikett ‚Algerien’, alle det Warncke als ,proxima ssp. aspericollis’, das 9 zusätzlich mit einem roten Etikett .Paratype’ (OLL) — © mit unleserlichem Etikett und verschiedenen PROXIANDRENA SUBGEN. NOV. 1041 Etiketten: Rosa mit schwarzem Rand [Cotypus], [Andrena aspericollis det. E. Stoeckhert]. [Andrena proxima det K. Warncke] (ZSM). DIAGNOSE: Andrena aspericollis ist laut Warncke (1967) dunkler gefärbt als proxima und besitzt keine Binden auf den Tergiten. Die Art ist in ihrer Verbreitung auf Algerien beschränkt. Die Weibchen sind auf den Tergiten sehr gleichförmig und fein chagriniert und zerstreut punktiert, das Mesonotum weist eine dichte grobe Punktierung mit kleinen, glänzenden Punktzwischenräumen auf (ampla mit glänzenden und auf dem Mesonotum größeren Punktzwischenräumen). Die Männchen besitzen wie proxima ein durchgehend chagriniertes Mesonotum und unterscheiden sich dadurch von der ebenfalls in Nordafrika vorkommenden ampla, deren Punkt- zwischenräume auf dem Mesonotum glänzen. DISKUSSION: Ich konnte nur zwei mangelhaft etikettierte Pärchen untersuchen, die vermutlich alle aus Algerien stammen und wahrscheinlich über 100 Jahre alt sind. Alle diese Tiere sind ohne Tergitbinden. Ob das von Pérez und Warncke beschriebene Hauptmerkmal der Art, die fehlenden Tergitbinden, auf den schlechten Erhaltungs- zustand der Tiere zurückzuführen oder natürlich ist, Kann ich nicht beurteilen. Da sich die Art aber skulpturell sowohl von ampla als auch von den europäischen Arten unter- scheidet, habe ich keine Zweifel, daß es sich hier um eine eigene Art handelt. Somit stelle ich den ehemaligen Status von aspericollis als eigene Art wieder her: Andrena aspericollis stat. rest. VERBREITUNG: Algerien. Andrena bernicla Warncke, 1974, stat. nov. Andrena proxima ssp. bernicla Warncke 1974b: 96. Auflistung. -— Warncke 1975: 50 (Teil B der Arbeit von 1974): Beschreibung d, Holotypus 3: Türkei, Ankara. (OLL) - Holotypus untersucht, dieser stammt nicht wie bei Warncke 1975 angegeben von Ankara, sondern von Sereflikochisar. Allerdings liegen Paratypen von Ankara vor. Andrena alutacea: Stoeckhert 1942: 236 ff. Ein Männchen mit schwarzer Gesichtsbehaarung aus Tiflis/Georgien, bezieht sich höchstwahrscheinlich auf bernicla. UNTERSUCHTES MATERIAL. Holotypus: é [TR Sereflikochisar 17.5.1970 leg. Warncke] [Andrena proxima ssp. bernicla War. det. Dr. Warncke ]; rotes Etikett [Holotypus] (OLL). TURKEI. 4 2 2 21.5.1972 Ankara leg. Warncke (alle Paratypen, det Warncke, OLL). Die Weibchen unterscheiden sich nicht von alutacea und werden zu dieser gerechnet. DIAGNOSE: Die Männchen unterscheiden sich von allen anderen Arten der Untergattung durch die schwarzbraune Gesichtsbehaarung (bei den übrigen Arten hell). Das Mesonotum ist ähnlich wie bei proxima skulpturiert. Das Weibchen ist nicht bekannt, bzw. läßt sich nicht von alutacea unterscheiden. DISKUSSION: Warncke (1974b) basiert die Beschreibung seiner Unterart bernicla auf die schwarze Gesichtsbehaarung des Männchen, die laut dem Autor allerdings nur in der ersten Generation vorkommen soll. Weiterhin ging Warncke in einer Ver- breitungskarte (publiziert in Gusenleitner & Schwarz 2002) davon aus, daß in der Türkei nur ein Taxon aus der proxima-Gruppe vorkommt, nämlich die ssp. bernicla. Wie das vorliegende Material zeigt, ist die Hypothese einer ersten dunkelgefärbten Generation jedoch nicht zu halten, da ich verschiedene türkische Männchen mit heller Gesichtsbehaarung untersuchen konnte, die wie bernicla ebenfalls im Mai gefangen wurden (zum Teil von Warncke selbst gesammelt). Da sich diese auch skulpturell von 1042 C. SCHMID-EGGER bernicla unterscheiden, gehe ich vielmehr davon aus, daß in der Türkei mindestens zwei Taxa aus der proxima-Gruppe, nämlich alutacea und bernicla, nebeneinander vorkommen. Andrena proxima wurde bisher in der Türkei nicht nachgewiesen. Allerdings ist die Situation um die beiden Taxa damit noch längst nicht befrie- digend gelöst. In der coll. Warncke (OLL) befinden sich zwei Weibchen von Ankara, bzw. Sille bei Konya, die Warncke als bernicla-Paratypen auszeichnete (ohne daß diese in der Literatur Erwähnung fanden). Beide Tiere - ein Weibchen wurde zeitgleich und am selben Ort mit dem männlichen Paratypus gefangen - sind völlig identisch mit alu- tacea-Weibchen aus Mitteleuropa und aus der übrigen Türkei. Entweder lassen sich die beiden Taxa im weiblichen Geschlecht nicht unterscheiden oder das Weibchen von bernicla ist noch unbekannt. Ein weiteres Problem besteht außerdem bei den Männchen. Stoeckhert (1942) erwähnt in der Originalbeschreibung von alutacea, daß die meisten Männchen ge- mischte schwarze und weiße Gesichtshaare mit Übergängen zu ganz weiß oder ganz schwarz besitzen sollen. Im von mir selbst untersuchten Material gab es nur entweder weiße oder schwarz gefärbte Tiere. Ich konnte keine einzige Mischform finden. Es bleibt offen, von wo Stoeckhert solche Männchen untersucht hat, da er keine explizi- ten Fundortangaben dafür macht. Ingesamt basiert er die Beschreibung von alutacea auf nur 7 Männchen und über 80 Weibchen, was einen Hinweis auf die Seltenheit der Männchen gibt. Stoeckhert erwähnt jedoch ausdrücklich ein Männchen aus Tiflis/ Georgien mit vollständig dunkler Gesichtsbehaarung, was höchstwahrscheinlich eine bernicla ist. Vorerst schließe ich aufgrund der morphologischen Unterschiede zwischen beiden Taxa und dem zeitlich übereinstimmenden Auftreten aus, daß bernicla eine schwarz gefärbte Form von alutacea ist und betrachte das Taxon als eigenständige Art: Andrena bernicla stat. nov. Die Betrachtung des Taxon als Form von alutacea wäre nicht plausibel, da mir solche lokalen Formen bei Andrena nicht bekannt sind. Die Gesichtsbehaarung ist vielmehr zumindest bei den mitteleuropäischen Arten ein konstantes Artmerkmal. Um eine partielle erste Generation von alutacea Kann es sich auch nicht handeln, da eine solche eher in einer wärmeren Region der Türkei als gerade Ankara zu erwarten wäre. Die Frage der Weibchen von bernicla bleibt allerdings ungeklärt. VERBREITUNG: Türkei, Georgien (Stoeckhert 1942, als alutacea). Andrena proxima (Kirby, 1802) Melitta proxima Kirby, 1802: 146-147. Andrena proxima wird sehr häufig in der taxonomischen und faunistischen Literatur erwähnt. Die einzelnen Meldungen und alte Synonyme sind hier nicht einzeln aufgeführt. Siehe dazu Gusenleitner & Schwarz, 2002: 610-611. UNTERSUCHTES MATERIAL: Tiere aus Deutschland und Österreich werden nicht einzeln aufgeführt. Hier standen etwa 200 Tiere aus den Sammlungen CS, GRU, ZSM u.a. zur Verfügung, außerdem untersuchte D. Doczkal sein mehrere hundert Tiere umfassendes Material aus Baden-Württemberg auf mögliche alutacea. FÜRSTENTUM LIECHTENSTEIN: 2 13.6.1997 Balzers, Elltal, ein abgeflogenes Tier, zusammen mit einer frischen alutacea; 9 24.4.1997 Vaduz, Möhliholzröfi (ETH). ITALIEN: 2 20.04.1971 Sizilien, Taormina; 2 06.1920 Liguria, Vittoria (GRU) — © 20.5.1922 Sizilien Taormina, 2 Puglia, 2 31.5.1993 Mt. Gargano, St. Giovanni, 2 16.5.1992 Mt. Gargano, S. Marco (ETH). GRIECHENLAND: 11.05.1968 PROXIANDRENA SUBGEN. NOV. 1043 Florina; 2 2 13.6.1959 Olymp, Prioni, 1000 m (sehr abgeflogene Tiere, zusammen mit einer frischen alutacea (GRU) — 29 9.5.1976 Nea Agathoupolis / Makedonien (Kraus). UNGARN: 9 28.05.1959 Tatatovaros, 2 04.06.1957 Roros, Ipoly; 2 8.06.1954 Bükk hegys; Hosszuberc; Tihany u.a. Fundorte (GRU). SCHWEIZ: 2 © 4.6.1997 BE, Lauperswil, Bluttengrat 900 m; 9 12.4.1942 GR, Haldenstein; 9 6.6.1936 GR, Felsberg; © 8.6.1924 GR, Mundaun; 2 27.05.1925 (ETH); TI, Lugano, Mt. Areo (GRU) DIAGNOSE: Andrena proxima ist im weiblichen Geschlecht charakterisiert durch die teilweise glänzenden Tergitflächen, die zumindest seitlich auch deutlich punktiert sein können. Alle Tergitendbinden sind in der Mitte breit unterbrochen, das Meso- notum zeigt keine schwarze kurze Unterbehaarung, oder diese ist nur im hinteren Teil schwach ausgeprägt. Siehe auch die Diskussion bei alutacea. Die Männchen lassen sich anhand des vollständig chagrinierten Mesonotums, der nur sehr fein und zerstreu- ten Tergitpunktierung und wie beim Weibchen an den breit unterbrochenen Tergitendbinden erkennen. VERBREITUNG: Andrena proxima ist in Mitteleuropa verbreitet und stellenweise häufig (Verbreitungskarte bei Gusenleitner & Schwarz, 2002). Im Norden erreicht sie Schleswig Holstein und das südliche Dänemark. Außerdem besiedelt sie den Süden von Großbritannien. Ihre Arealausdehnung im Osten ist unbekannt, allerdings scheint sie in Asien (auch in der Türkei) zu fehlen. Ein Tier in der Sammlung GRÜ aus Lettland erwies sich zu alutacea gehörend. Im Südwesten erreicht sie Mittel- und wahrscheinlich auch Südfrankreich, allerdings muss hier die Artzugehörigkeit von Belegmaterial zu ampla überprüft werden. Italien ist bis Sizilien von proxima besie- delt, im Südosten erreicht sie Griechenland und kommt dort zum Teil syntop mit alu- tacea Vor. OKOLOGIE: Andrena proxima besiedelt vor allem mageres Griinland, aber auch andere trockenwarme Offenstandorte. Die Art ist auf Doldenbliitler (Apiaceae) als Pollenquelle spezialisiert (Westrich, 1989). Die Art ist in Deutschland nicht gefährdet. DANKSAGUNG Fiir die Bereitstellung von Sammlungsmaterial bedanke ich mich bei den fol- genden Personen: Frank Burger (Weimar), Dr. Andreas Miiller und Sabine Oertli (Ziirich), Dr. Bernard Merz (Genf), Johannes Schuberth und Dr. Stefan Schmidt (München), Dr. Manfred Kraus (Nürnberg), Fritz Gusenleitner (Linz), Erwin Scheuchl (Velden) und Dr. Mike Herrmann (Konstanz). Erwin Scheuchl und Prof. Dr. Klaus Schönitzer gaben wertvolle Hinweise zum Manuskript. Erwin Scheuchl stellte mir außerdem freundlicherweise ausführliche Literaturauswertungen zur Verfügung. LITERATUR DUBITZKY, A. & SCHONITZER, K. 2001. The propodeal corbicula of Andrena proxima and allied species (Hymenoptera, Andrenidae). Apidologie 32: 429-434. DyLEWSKA, M. 1987. Die Gattung Andrena Fabricius (Andrenidae, Apoidea) in Nord- und Mitteleuropa. Acta Zoolocia cracovil 30: 359-708. ERLANDSSON, S. 1979. Hymenoptera Aculeata from the European Part of the Mediterranean countries II. Acta Entomologica 15: 125. 1044 C. SCHMID-EGGER GUSENLEITNER, F. & SCHWARZ, M. 2002. Weltweite Checkliste der Bienengattung Andrena mit Bemerkungen und Ergänzungen zu paläarktischen Arten (Hymenoptera, Apidae, Andreninae, Andrena). Entomofauna, Supplement 12: 1280 Seiten. KirBy W. 1802. Monographia Apum Angliae II. Ipswich: 146-147. KOCOUREK, M. 1966. Prodromus der Hymenopteren der Tschechoslowakei, part 9: Apoidea, 1. Acta Faunistica Entomologica Musei Nationalis Pragae, Suppl. 2, 12: 69. MANDERY, K. 2001. Die Bienen und Wespen Frankens. Bund Naturschutz Forschung 5: 285 Seiten. PEREZ, J. 1895. Espèces nouvelles de Melliferes de Barbarie (Diagnoses préliminaires). Bordeaux. 37. PITTINOI, B. & Schmidt, R. 1943. Die Bienen des südöstlichen Niederdonau. 2. Andrenidae und isoliert stehende Gattungen. Niederdonau /Natur und Kultur 24: 33. SCHMID-EGGER, C. & SCHEUCHL, E. 1997. Illustrierte Bestimmungstabellen der Wildbienen Deutschlands und Osterreichs unter Beriicksichtigung der Arten der Schweiz. Band III. Andrenidae. Eigenverlag, Velden, 180 Seiten. SCHWARZ, M., GUSENLEITNER, F., WESTRICH, P. & DATHE, H.H.. 1996. Katalog der Bienen Osterreichs, Deutschlands und der Schweiz (Hymenoptera, Apidae). Entomofauna, Supplement 8: 398 Seiten. STOECKHERT, E. 1942. Andrena proxima K. und ihr bisher unbekannter «Doppelganger», Andrena alutacea n.sp. (Hym. Apid.). Mitteilungen der Münchener Entomologischen Gesellschaft 32: 236-252. WARNCKE, K. 1967. Beitrag zur Klärung paläarktischer Andrena-Arten. Eos 43: 209, 250. WARNCKE, K. 1968. Die Untergattungen der westpaläarktischen Bienengattung Andrena F. Memoires Estudes Museo Zooogico Universitade Coimbra 307: 1-111. WARNCKE, K. 1974a. Beitrag zur Kenntnis und Verbreitung der Sandbienen in Nordafrika (Hymenoptera, Apoidae, Andrena). Mitteilungen des Zoologischen Museums in Berlin 50: 3-53. WARNCKE, K. 1974b. Die Sandbienen der Türkei, Teil A. Mitteilungen der Münchener Entomo- logischen Gesellschaft 64: 96. WARNCKE, K. 1975. Die Sandbienen der Türkei (Hymenoptera, Apoidea, Andrena). Teil B. Mitteilungen der Münchener Entomologischen Gesellschaft 65: 29-102. WARNCKE, K. 1976. Die Bienengattung Andrena F. 1775, in Iberien. Teil B. Eos 50. 151. WARNCKE, K. 1986. Die Wildbienen Mitteleuropas, ihre gültigen Namen und ihre Verbreitung. Entomofauna, Supplement 3: 45. WESTRICH, P. 1984. Kritisches Verzeichnis der Bienen der Bundesrepublik Deutschland (Hymen- optera, Apoidea). Courier Forschung Institut Senckenberg 66: 1-86. WESTRICH, P. 1989. Die Wildbienen Baden-Württembergs. Stuttgart, 972 Seiten. WESTRICH, P., GUSENLEITNER, F. & AMIET, F. 1997. Andrena afrensis Warncke 1967, eine für Mitteleuropa neue Bienen-Art (Hymenoptera, Apidae). Linzer biologische Beiträge 29: 1167-1174. REVUE SUISSE DE ZOOLOGIE 112 (4): 1045-1069; décembre 2005 African biodiversity hotspots: the reptiles of Mt Nlonako, Cameroon Hans-Werner HERRMANN! 2, Wolfgang BOHME?, Oliver EUSKIRCHENY, Patricia A. HERRMANN? & Andreas SCHMITZ> 1 Center for Reproduction of Endangered Species (C.R.E.S.), Zoological Society of San Diego, P.O. Box 120551, San Diego, CA 92112-055, U.S.A. E-mail: hwherrmann@sandiegozoo.org 2P.O. Box 3055, Messa, Yaounde, Cameroon. 3 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. 4 Bismarckstrasse 10, D-56626 Andernach, Germany. 5 Department of Herpetology and Ichthyology, Museum of Natural History, C.P. 6434, CH-1211 Geneva 6, Switzerland. African biodiversity hotspots: the reptiles of Mt Nlonako, Cameroon. - The reptiles of Mt Nlonako, a mountain at the southeastern edge of the Cameroon mountain range (“Dorsale camerounaise”), were inventoried continually over a six year period from 1998 to 2004. This area encom- passes 150 km? of lowland, submontane and montane rainforest with an elevation up to 1,825 m. Accounts of 89 species are provided based on collected and photo-documented material. This inventory proved Mt Nlonako to be the most species rich single-locality area in reptilian fauna in Africa. With 63 snake species Mt Nlonako exhibits the greatest number of species in Africa and possibly worldwide. Analysis showed the reptilian species composition to be most similar within Cameroon with that of Korup National Park followed by the Dja Faunal Reserve in the south. Relative to the snake composition the Korup NP and the Dimonika region in Congo- Brazzaville show the highest resemblance. In an African context the reptile fauna of Mt Nlonako is characterized by species which occur in both West and Central Africa as opposed to the mountain’s amphibian species which more closely resemble Central African fauna. The high species richness and endemicity is discussed from a paleoclimatic perspective. Conservation status and threats to the reptiles are noted. Keywords: Amphisbaenia - Crocodylia - Chelonia - Sauria - Serpentes - species richness - endemicity - biogeography - conservation. INTRODUCTION Despite efforts in surveying the reptile (and amphibian) fauna of the rain forests of tropical central and west Africa (Perret, 1959, 1960, 1961; Leston & Hughes, 1968; Böhme, 1975; Joger, 1982; Trape, 1985; Böhme & Schneider, 1987; Ota et al., 1987; Manuscript accepted 15.12.2004 1046 H.-W. HERRMANN ET AL. Trape & Roux-Esteve, 1990; Lawson, 1993; Böhme, 1994a, b; Rödel et al., 1995; Schmitz, 1998; Rodel et al., 1999; Böhme, 2000; Euskirchen et al., 2000; Schmitz et al., 2000; Gossmann et al., 2002; Branch & Rédel, 2003) the knowledge on African rainforest reptile faunas falls behind that of its equivalents in tropical Middle and South America and in tropical Asia (Dunn, 1949; Inger & Colwell, 1977; Murphy et al., 1994). Within the rainforest zones of central and west Africa however, Cameroon, Ghana and Ivory Coast have the best studied reptile faunas (Hughes, 1983; Rödel er al., 1995; LeBreton, 1999; Rödel et al., 1999). Lawson (1993) lists 83 reptile species for the Korup National Park (Korup NP), which makes this one of the most species rich areas for reptiles on the continent. Trape (1975) records 45 snake species from the Dinamika forest site in the People’s Democratic Republik of Congo (Congo- Brazzaville). In Cameroon, the Western and Southwestern Cameroon highlands also known as “Dorsale camerounaise” extending from Mt Cameroon in the south to Tchabal Mbabo in the north are characterized by a high amphibian and reptile species richness and an extraordinary high proportion of endemic species; a veritable hotspot of African herpetofauna diversity (Duellman, 1999; Poynton, 1999; Herrmann et al., 2005). Contrary to the high biodiversity value of such areas, especially in West and Central Africa, most have no formal conservation status protection (STUART ef al., 1990) and are endangered by habitat destruction, mainly by logging activities and human encroachment. Additionally, the natural history of most reptile species from tropical forests in Africa remains unknown. Quantitative ecological studies are entirely lacking with only the first steps taken towards quantitative studies of chameleons (Herrmann & Herrmann, 2005). Ultimately, only a comprehensive understanding of the reptiles of these areas and their natural history will provide the fundamental knowledge for their future survival. This paper presents a comprehensive reptile species list for Mt Nlonako, which was derived over a period of six years, in which we compile the largest number of reptile species (89), in particular snake species (63), for any single-locality on the African continent. STUDY SITES We surveyed the Mt Nlonako area (Fig. 1) which extends roughly from 4°49’- 4°56’N and from 9°56°-10°01°E and encompasses ca. 15,000 ha. The western and northern flanks face the town of Nkongsamba, and the busy road between Douala and Bamenda. The slopes on this side are heavily cultivated with the forest destroyed up to an elevation of approximately 1,100 m. To the south and east however, the forest slopes are much less influenced by human activities. A vast lowland rainforest, encompassing several thousand km?, extends from the foothills of Mt Nlonako reaching past Nkondjok in the East and past Yabassi in the South. This area is divided by some unpaved roads and settlements. Logging is or has been carried out in many places within this area. Although hunting pressure is imminent, forest elephants, gorillas, chimpanzees, drills and other large mammals persist. Mt Nlonako itself raises from about 400 m elevation on the southern side to 1,825 m on its peak. The highest and central part of the mountain forms a cuvette, ca. 1.5 km in diameter, with much THE REPTILES OF MT NLONAKO, CAMEROON 1047 A= study site Cameroon! Nlona ko = N û QEkambeng Nkebe waterfall a G Badjong {ph |Ekomtolo O km Fic. 1 Survey locations (a) in Cameroon and (b) at Mt Nlonako. Shaded areas are rural communities. 1048 H.-W. HERRMANN ET AL. grass/bracken in its center and with forested rims at 1,600 m on the north, east and south sides and the peak on the western side (Dowsett-Lemaire & Dowsett, 1999). The forest above 1,100 m is pristine with a tall canopy (25-30 m). The forest here seems to be drier and warmer than forests on the close-by Manengouba and Bakossi mountains or Mt Kupe at comparable altitudes. Botanically, Guttiferae (Allanblackia sp.) and Burseraceae (Santiria trimera) are especially common. Caesal- piniaceae (incl. Tessmannia anomala), Ebenaceae (Diospyros), Meliaceae, Mimo- saceae (Albizia), Moraceae, Olacaceae (Strombosia), Sapotaceae (incl. Chrysophyllum albidum), Steruliaceae (Cola) and Apocynaceae (Tabernaemontana sp.) are recorded (Dowsett-Lemaire & Dowsett, 1999). Above 1,450 m some rare montane species such as Polyscias fulva can be found locally. Many small to medium-sized creeks, often fast flowing and rocky, can be found in the forest. Swamps and pools are rare. The climate is warm and humid. Over a period of 34 years Nkongsamba (882 m elevation) received an average of 2,762 mm rainfall per year (table 1, Amiet, 1975). During that period the peak dry season extended from December to February with less than 50 mm precipitation per month. The peak rainy season extended from July to September with up to 482 mm precipitation per month. For detailed data on climate and rainfall see Herrmann et al. (2005). Fieldwork initiated in November 1998 and continual sampling extended to June 2004. Sampling occurred in all seasons. Our survey efforts concentrated on the northern, eastern and southern slopes of Mt Nlonako with the following localities being the main points of collecting: - Ekambeng (EKA): village at the foot of the northern slopes in the vicinity of Nkongsamba, many coffee plantations, no primary forest - Mekoum (MEK): village on northern side of the mountain, many coffee plan- tations, no primary forest - Nguéngué (NGU): village on the rim between northwestern and southeastern slopes, N 4°55°02”, E 9°59’21”, 1,140 m elevation, some coffee plantations, second- ary and predominately primary forest - Summit (SUM): eastern side of the cuvette, N 4°54°47”, E 9°57’93” , 1,660 m elevation, rock outcrops, primary montane forest, some areas with grass and bushes - Eyimba (EYI): very small village on the eastern side of the mountain, N 4°52°92”, E 9°59’19”, 710 m elevation, small cultivated areas, much primary forest - Nkebe waterfall (NWF): area between the villages Ekomtolo and Badjong (but also including those two villages), N 4°49’83” , E 9°55°49”, 470 m elevation, coffee, oil palm and food crop plantations, predominately secondary forest, area with previous and current logging. METHODS SAMPLING METHODS We used Y-shaped drift fence/pitfall trap arrays (Corn, 1994) with segments of 5 m length during the initial phase of the project at several localities. Catching success was very low to nil. This method was abandoned after some weeks. Initially, gluetraps (Bauer & Sadlier, 1992; Glor et al., 2000) which were fixed on tree trunks at about 1.5 m height, were used to sample geckos and other arboreal THE REPTILES OF MT NLONAKO, CAMEROON 1049 reptiles. Only a few skinks (Trachylepis) were caught using this method. The abundant ants tended to attack trapped reptiles immediately. This method was abandoned after the first two weeks. We applied quadrat sampling with 8 x 8 m quadrats (Jaeger & Inger, 1994) at several localities at various elevations. Quadrats worked satisfactorily to gain quanti- tative data but are work intensive. For the quantitative results see Herrmann et al. (2000). Visual encounter surveys along transects (Crump & Scott, 1994) or opportu- nistic searches during the day and at night were the dominant methods used. The number of persons surveying varied, but was usually two or three; this method yielded the best success and was carried out throughout the study periods. Acoustic monitoring was also applied but only species represented by at least one voucher specimen are included in the species list. Our data was complemented by donated specimens which were encountered by villagers during their daily movements. Voucher specimens are deposited in the herpetological collection of the Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn (ZFMK) and in the collection of the senior author (HWH) in the U.S. The latter collection will be transferred to the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). All specimens were either fixed in 10% buffered formal- dehyde or 75% ethanol and consecutively preserved in 75% ethanol. BIOGEOGRAPHIC ANALYSIS To compare the species composition of Mt Nlonako with other areas in West and Central Africa we calculated the “coefficient of biogeographic resemblence” (CBR) after Duellman (1990) with the formula CBR = 2C / (N,+N)) in which C is the number of shared taxa (here species) in two compared areas, Nj is the number of taxa (here species) in area one and N, is the number of taxa (here species) in area two. A CBR value of 0 would mean that Mt Nlonako shares no rep- tilian species with the compared area, a CBR value of 1 would mean that the species inventory in both areas are identical (Jansen & Köhler, 2002). Jansen and Köhler (2002) excluded ubiquitous species from their analysis arguing that those species are not primary inhabitants of the mountain forest habitats they compared and thus do not bear any information on the biogeographic relationships of those areas. We do not follow this approach for (1) we do not know if ubiquitous species are or are not pri- mary inhabitants of the areas in question, (2) possible “noise” by such species should be similar over the areas analyzed and (3) they are low in number and thus have a limited effect on the analysis. RESULTS SPECIES ACCOUNT In the following we present a systematic list of reptile species encountered during our survey. We list localities and voucher specimens for each taxon. Nomen- clature follows Uetz et al. (2004), unless otherwise noted. For voucher specimens with no exact locality data we state the locality as Mt Nlonako (MtNL). Otherwise col- 1050 H.-W. HERRMANN ET AL. lecting localities are Nkebe waterfall area (NWF), Ekambeng (EKA), Mekoum (MEK), Eyimba (EYI), Nguéngué (NGU) and the Mt Nlonako summit area (SUM). For habitat we differentiate between forest (F) and farmbush (FB). Farmbush are the ecotones between forests and cultivated areas as well as the vicinities of villages and the dirt roads. We also include degenerated secondary forest patches in this category. AMPHISBAENIA AMPHISBAENIDAE Monopeltis sp. (Duméril, 1859) Locality: EYI. Voucher specimen: ZFMK 68960. Habitat: FB. Remarks: Only two species, Monopeltis jugularis Peters, 1880 and M. galeata (Hallowell, 1852) are known from Cameroon. This is the first record of an am- phisbaenid from Western Cameroon. Since this specimen is a roadkill in a rather bad conservation state, an identification to the species-level was not possible. CROCODYLIA CROCODYLIDAE Osteolaemus tetraspis Cope, 1861 Locality: NWF, EYI. Voucher specimen: none. Habitat: F. Remarks: Regularly seen in streams and pools. This species is commonly found as bushmeat in local markets such as in Nkongsamba (see Akani et al., 1998 for the situation in Nigeria). Although under severe hunting pressure, this species seems to occur in stagnant river arms and pools at lower elevations throughout the study area. CITES appendix I. CHELONIA TESTUDINIDAE Kinixys erosa (Schweiger, 1812) Locality: NGU. Voucher specimen: ZFMK 68982. Habitat: F. Remarks: Shells of this species can regularly be found in villages where this species is appreciated as bushmeat (Lawson, 2000). CITES appendix II. Kinixys homeana Bell, 1827 Locality: NWF. Voucher specimen: none. Habitat: F. Remarks: Shells of two specimens of this species are kept in the compound of the chief of Ekomtolo. Like K. erosa, this species is commonly eaten by local people. CITES appendix II. SAURIA AGAMIDAE Agama agama (Linnaeus, 1758) Locality: NGU. Voucher specimens: ZFMK 69017, 75376. Habitat: FB. Remarks: This species was found in all settlements of the study area where it is ‘very abundant in villages and along forest roads. It was never observed in the forest. THE REPTILES OF MT NLONAKO, CAMEROON 1051 CHAMAELEONIDAE Chamaeleo camerunensis Miiller, 1909 Locality: NWF. Voucher specimen: HWH 20. Habitat: F. Remarks: Only one 2 of this species was found and is treated in detail in Harbort & Herrmann (2002). CITES appendix I. Chamaeleo cristatus Stutchbury, 1837 Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 68961, 68980- 1, 73063-6, 75146, 75394, HWH 31, 202-4, 617, 816-29, 893-6. Habitat: F, FB. Remarks: CITES appendix II. Chamaeleo montium Buchholz, 1874 Locality: MtNL, NGU. Voucher specimens: ZFMK 69117-20, 72855, 73067- 74, 73491-5, 75388-93, HWH 208-12, 830-44. Habitat: F, FB. Remarks: CITES appendix II. Chamaeleo oweni Gray, 1831 Locality: MtNL, NWF. Voucher specimens: HWH 205-7. Habitat: F, FB. Remarks: CITES appendix II. Chamaeleo pfefferi Tornier, 1900 Locality: SUM. Voucher specimens: ZFMK 73490, 74336, HWH 1013. Habitat: F. Remarks: CITES appendix II. Rhampholeon spectrum (Buchholz, 1874) Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 69101-16, 72853- 4, 73075-8, 73496, 75287-95, HWH 16, 194-200, 618, 845-85. Habitat: F. GEKKONIDAE Hemidactylus intestinalis Werner, 1897 Locality: NWF. Voucher specimens: ZFMK 78684, 81713, HWH 213-4, 602. Habitat: F, FB. Remarks: Henle & Böhme (2003) provide a detailed species account, though using the name Hemidactylus ansorgii Boulenger, 1901, which has been shown to be a junior synonym of H. intestinalis Werner, 1897 by Perret (1975), and this synonymy has also been mentioned in Kluge (1991). Hemidactylus echinus O’ Shaughnessy, 1875 Locality: NGU. Voucher specimen: ZFMK 69040. Habitat: F, FB. Hemidactylus fasciatus Gray, 1842 Locality: NWF. Voucher specimen: ZFMK 77893. Habitat: F. Hemidactylus mabouia (Moreau de Jonnès, 1818) Locality: NGU. Voucher specimens: ZFMK 69026-39, 75377-9, HWH 1056-7. Habitat: FB. Remarks: Like Agama agama, this species can be found in all human settlement in the study area. It is especially abundant on houses. 1052 H.-W. HERRMANN ET AL. Hemidactylus sp. indet. Locality: NWF. Voucher specimen: ZFMK. Habitat: F, FB. Remarks: Juvenile specimen which we could identify to species level. Lygodactylus conraui Tornier, 1902 Locality: EYI. Voucher specimen: ZFMK 68966. Habitat: F, FB. LACERTIDAE Poromera fordii (Hallowell, 1857) Localities: NWF, NGU. Voucher specimens: ZFMK 69021-5, 78660, 81715, HWH 1058. Habitat: F, FB. SCINCIDAE Trachylepis affinis (Gray, 1838) Localities: EYI, NGU. Voucher specimens: ZFMK 68962-4, 69062-73, 75383- 7. Habitat: FB. Remarks: Listed as Mabuya affinis in Uetz et al. (2004). We follow the nomen- clature proposed by Bauer (2003) and use the genus name Trachylepis. Common in villages, farms and along roads throughout the study area. Trachylepis maculilabris (Gray, 1845) Localities: NGU, SUM. Voucher specimens: 69041-61, 69434, 75576-7. Habitat: F, FB. Remarks: Listed as Mabuya maculilabris in Uetz et al. (2004). We follow the nomenclature proposed by Bauer (2003) and use the genus name Trachylepis. Found in human settlements but also in secondary forest and on rock faces at the summit of Mt Nlonako. Locally abundant. Lygosoma fernandi (Burton, 1836) Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 68959, 69018, 73497, 78026-7, 78814. Habitat: F, FB. Remarks: In Uetz er al. (2004) as Mochlus fernandi. We prefer the genus name Lygosoma as molecular analyses reveal a closer relationship with this genus (A. S. unpublished results). Panaspis breviceps (Peters, 1873) Localities: NWF, NGU. Voucher specimens: ZFMK 69074-85, 72856, 75380, 77889-92, 78700, 78711, HWH 601. Habitat: F. Lacertaspis reichenowi (Peters, 1874) Locality: EYI. Voucher specimen: ZFMK 68965. Habitat: F. Remarks: Listed as Panaspis reichenowi in Uetz et al. (2004). We follow the nomenclature proposed by Schmitz er al. (2005) and use the genus name Lacertaspis. Lacertaspis rohdei (Miiller, 1910) Locality: NGU. Voucher specimens: ZFMK69086-99, 75382, HWH 1052-5. Habitat: F. THE REPTILES OF MT NLONAKO, CAMEROON 1053 Remarks: Listed as Panaspis rohdei in Uetz et al. (2004). We follow the nomen- clature proposed by Schmitz et al. (2005) and use the genus name Lacertaspis. Leptosiaphos vigintiserierum (Sjöstedt, 1897) Localities: NGU, SUM. Voucher specimens: ZFMK 69426-32, 69549. Habitat: E. Leptosiaphos spp. Locality: NGU. Voucher specimens: ZFMK 69529-31, 69550-4, 75381. Habitat: F. Remarks: Our material of hitherto undescribed or unidentifiable Leptosiaphos includes five morphologically distinct groups. One groups resembles L. pau- liani (Angel, 1940) (Fig. 2) from the Bamboutos Mtns, Cameroon, another L. vigintiserierum (Fig. 3). VARANIDAE Varanus ornatus (Daudin, 1803) Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 69016, HWH 18, 982. Habitat: F, FB. Remarks: A common species which is often found as bushmeat in villages and local markets. Large specimens sometimes are encountered in snare traps. CITES appendix II. SERPENTES TYPHLOPIDAE Typhlops angolensis (Bocage, 1866) Localities: EKA, NGU. Voucher specimens: ZFMK 75126, 75178-9. Habitat: EB} Typhlops congestus (Duméril & Bibron, 1844) Locality: MtNL. Voucher specimens: ZFMK 77642, 78132-6, 78819-24, HWH 587. Habitat: FB. Typhlops lineolatus Jan, 1864 Locality: MtNL. Voucher specimens: HWH 588, 695. Habitat: FB. Typhlops punctatus (Leach in Bowdich, 1819) Locality: MtNI. Voucher specimens: ZFMK 78815-8. Habitat: FB. BOIDAE Calabaria reinhardtii (Schlegel, 1848) Localities: MtNL, NGU. Voucher specimens: ZFMK 75780, HWH 690-1, 808- 15, 897-901, 1009. Habitat: F, FB. Remarks: This is one of the most abundant snakes in the area. Often found cros- sing roads or dead on the road (DOR). The species was observed in many dif- ferent localities in the study area. CITES appendix II. Python sebae (Gmelin, 1789) Localitiy: MtNL. Voucher specimens: HWH 890-1. Habitat: F, FB. 1054 H.-W. HERRMANN ET AL. FIGs 2-3 2. Leptosiaphos sp. cf. pauliani from the forest above Nguéngué. 3. Leptosiaphos sp. cf. viginti- serierum from the forest above Nguéngué. THE REPTILES OF MT NLONAKO, CAMEROON 1055 Remarks: This species is probably restricted to the lower elevations of the study area. People of NGU for example did not confirm the existence of the species in their area. On one occasion, villagers of Ekomtolo were observed preparing a medium sized P. sebae for consumption. The species is well known to the peo- ple of Ekomtolo and Badjong. It is not common. CITES appendix II. COLUBRIDAE Afronatrix anoscopus (Cope, 1861) Locality: MtNL, NWF. Voucher specimens: ZFMK 81716, HWH 400, 753. Habitat: F, FB. Boiga blandingii (Hallowell, 1844) Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 68987, 75119, 75175-7, 75857-8, HWH 681, 749-51. Habitat: F, FB. Boiga pulverulenta (Fischer, 1856) Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75155-7, 77913-4, 78085, 78852-6, 81717, HWH 423-32. Habitat: F, FB. Bothrophthalmus brunneus (Ginther, 1863) Locality: MtNL, NWF. Voucher specimens: ZFMK 75121, 78839, HWH 418, 450. Habitat: F. Remarks: The former B. lineatus subspecies brunneus is considered to be a full species by Hughes (2000) and Gossmann et al. (2002). Chamaelycus fasciatus (Ginther, 1858) Localities: MtNL, NWF, EKA, MEK, EYI, NGU, SUM. Voucher specimens: ZFMK 69002, 69433, 75224-32, 77631, 77663, 78103, 78845-6, HWH 407-17, 633. Habitat: F. Dasypeltis fasciata Smith, 1849 Locality: MtNL. Voucher specimens: HWH 404-5. Habitat: F. FB. Dasypeltis scabra (Linnaeus, 1758) Localities: MtNL, NGU. Voucher specimens: ZFMK 78118-9, 78864-7. Habitat: F, FB. Dipsadoboa duchesnei (Boulenger, 1901) Locality: MtNL, NWF. Voucher specimens: ZFMK 81718, HWH 435-7, 715. Habitat: F. Dipsadoboa underwoodi Rasmussen, 1993 Locality: NWF. Voucher specimen: ZFMK 81719. Habitat: F. Dipsadoboa unicolor Giinther, 1858 Localities; MtNL, NGU. Voucher specimens: ZFMK 75266-7, HWH 438, 716. Habitat: F. Dipsadoboa viridis (Peters, 1869) Locality: MtNL. Voucher specimens: ZFMK 78862-3, 79031, HWH 433-4, 717, 759-64. Habitat: F. Dipsadoboa weileri (Lindholm, 1905) Localities: MtNL, NGU. Voucher specimens: ZFMK 75340, 78121-2, HWH 439-40. Habitat: F. 1056 H.-W. HERRMANN ET AL. Gastropyxis smaragdina (Schlegel, 1837) Localities: MtNL, NWF, EKA, MEK, EYI, NGU. Voucher specimens: ZFMK 68975, 68989-91 , 75134, 75152-4, 75262-4, 77657-60, 77906, 78077, 78093-4, 79029-30, HWH 399, 626, 802. Habitat: F, FB. Gonionotophis brussauxi (Mocquard, 1889) Localities: MtNL, NWF, EYI. Voucher specimens: ZFMK 77636, 77915, 78850-1. Habitat: F. Grayia caesar (Giinther, 1863) Locality: NGU. Voucher specimen: ZFMK 78120. Habitat: F. Grayia smythii (Leach, 1818) Locality: NWF. Voucher specimen: HWH 1208. Habitat: F, FB. Hapsidophrys lineatus Fischer, 1856 Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 75125, 75265, 75373, 78078-9, 79022-8, HWH 387-93, 803-6. Habitat: F, FB. Hormonotus modestus (Duméril, Bibron & Duméril, 1854) Locality: MtNL. Voucher specimens: ZFMK 78859-61. Habitat: F. Hydraethiops melanogaster Giinther, 1872 Locality: MtNL. Voucher specimens: ZFMK 72729, HWH 694, 752. Habitat: F. Lamprophis fuliginosus-lineatus-complex Thorpe & McCarthy (1978) state, that the two species L. fuliginosus and L. lin- eatus are difficult to differentiate in some areas. Our material allowed the as- signment of specimens to either of the two species. Lamprophis fuliginosus (Boie, 1827) Locality: NGU. Voucher specimens: ZFMK 69010-5, 78219. Habitat: F, FB. Lamprophis lineatus Duméril, Bibron & Duméril, 1854 Localities: MtNL, NWF, NGU. Voucher specimens: HWH 441-9, 628-30, 702- 9, 778-94, 965-74, 976, 1008, 1163. Habitat: F, FB. Lamprophis olivaceus (Duméril, 1856) Localities: MtNL, NWF, MEK, NGU. Voucher specimens: ZFMK 68976, 75122, 75240, 75343, 77920, 79037-9, HWH 590, 981. Habitat: F, FB. Lamprophis virgatus (Hallowell, 1854) Localities: MtNL, NWF, EKA, MEK, EYI, NGU. Voucher specimens: ZFMK 68977, 69007-9, 75116-8, 75138-45, 75147-51, 75344-71, 77637-8, 77662, 77921-2, 78203-18, 79040-66. Lycophidion laterale Hallowell, 1857 Localities: MtNL, NGU. Voucher specimens: ZFMK 69398, 75233-5, 75374, 78080-1, 78843-4, HWH 406, 754-5. Habitat: F. Mehelya guirali (Mocquard, 1887) Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 69001, 75169- 74, 75270, 77909, 78082-4, 78092, 78830-2. Habitat: F, FB. Mehelya poensis (Smith, 1847) Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75268, 77910- 2, 78825, 78827-9, HWH 419, 616, 713, 756-8. Habitat: F, FB. THE REPTILES OF MT NLONAKO, CAMEROON 1057 Mehelya riggenbachi (Sternfeld, 1910) Localities: MtNL, NWF. Voucher specimens: HWH 420, 631, 718. Habitat: F. Mehelya stenophthalmus (Mocquard, 1887) Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 75269, 78131, 78826, HWH 632, 984. Habitat: F, FB. Natriciteres fuliginoides (Giinther, 1858) Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: 75220, 77643- 56, 77916-7, 78104, 78178-202, 78985-79003, HWH 451-74, 610-1, 613-4, 696-8, 701, 767-74, 942-58, 1010. Habitat: F, FB. Natriciteres olivacea (Peters, 1854) Localities: MtNL, MEK, NGU. Voucher specimens: ZFMK 68974, 68992, 68997-9, 75201-19, 75341-2, 75372, 78979. Habitat: F, FB. Natriciteres variegata (Peters, 1861) Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 68993-6, 75180- 200, 77918-9, 78095, 78173-7, 78980-4, HWH 475-89, 698-700, 776-7, 959- 64. Habitat: F, FB. Philothamnus carinatus (Andersson, 1901) Localities: MtNL, MEK, NGU. Voucher specimens: ZFMK 75236-9, 75339, 78127, 79032-6, HWH 634-6, 712, 798, 985. Habitat: F. Remarks: In Uetz et al. (2004) not listed for Cameroon and in LeBreton (1999) as a subspecies of heterodermus. Philothamnus heterodermus (Hallowell, 1857) Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 77632-5, 78123-6, HWH 710-1, 795-7, 807. Habitat: F, FB. Philothamnus nitidus (Günther, 1863) Locality: MtNL. Voucher specimen: HWH 637. Habitat: F. Poecilipholis cameronesis Boulenger, 1903 Locality: MINI. Voucher specimen: HWH 589. Habitat: F. Rhamnophis aethiopissa Günther, 1862 Locality: NGU. Voucher specimen: ZFMK 68979. Habitat: F, FB. Rhamnophis batesii (Boulenger, 1908) Localities: MtNL, NWF, EKA, NGU. Voucher specimens: ZFMK 75334, 77661, 78105-10, 78833-8, 79067, HWH 401-2, 627. Habitat: F. Thelotornis kirtlandii (Hallowell, 1844) Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68988, 75271-5, 77664, 78073-4, 78871-9, HWH 421-2, 615, 625, 680, 975, 1006-7. Habitat: F, FB. Thrasops flavigularis (Hallowell, 1852) Localities: MtNL, NGU. Voucher specimens: ZFMK 78032-3, 78840-2, HWH 403. Habitat: F, FB. 1058 H.-W. HERRMANN ET AL. ATRACTASPIDIDAE Aparallactus modestus (Giinther, 1859) Localities: NWF, EYI, MEK, NGU. Voucher specimens: ZFMK 68978, 69003- 6, 75123-4, 75241-60, 75335, 75337-8, 77640-1, 77907-8, 78150-72, 79006- 21, 79221-2, HWH 640-9, 977-80, 1011-2. Habitat: F, FB. Aparallactus niger Boulenger, 1897 Locality: MtNL. Voucher specimen: ZFMK 79005. Habitat: F. Remarks: The specimen possesses a single prefrontal with the nasals in broad contact. This specimen represents the first record of this species for Cameroon. Atractaspis aterrima Günther, 1863 Locality: MtNL. Voucher specimen: HWH 766. Habitat: F, FB. Atractaspis boulengeri Mocquard, 1897 Locality: EKA. Voucher specimen: ZFMK 75136. Habitat: F. Atractaspis irregularis (Reinhardt, 1843) Locality: EKA. Voucher specimen: ZFMK 75137. Habitat: F, FB. Atractaspis reticulata Sjöstedt, 1896 Localities: MtNL, NGU. Voucher specimens: ZFMK75261, HWH 639, 714, 765. Habitat: F. Polemon collaris (Peters, 1881) Localities: MtNL, NGU. Voucher specimens: ZFMK 69000, 75221-3, 78137- 49, 78847-9, HWH 364-5, 693. Habitat: F. Polemon gabonensis (Duméril, 1856) Locality: NWF. Voucher specimen: ZFMK 75120. Habitat: F. Polemon gracilis (Boulenger, 1911) Locality: NGU. Voucher specimen: ZFMK 78129. Habitat: F. Remarks: This is the first voucher specimen from Cameroon. Polemon notatus (Peters, 1882) Localities: NWF, EYI, NGU. Voucher specimens: ZFMK 77905, 78128, HWH 17. Habitat: F. ELAPIDAE Dendroaspis jamesoni jamesoni (Traill, 1843) Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK68957, 75276, 75336, 77672-4, 77898, 78096-102, 78887-95, HWH 380-2, 719-24, 996-8. Habitat: F, FB. Naja melanoleuca Hallowell, 1857 Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68985- 6, 69019-20, 75127-33, 75158-62, 75277-84, 75793, 77630, 77667-71, 77900- 4, 78034-53, 78947-78, 79004, HWH 384-6, 604-5, 612, 619-22, 676-9, 727- 36, 889, 925-41, 1000-5. Habitat: F, FB. Remarks: In the study area this is the most abundant snake in villages where specimens are often attracted by poultry. THE REPTILES OF MT NLONAKO, CAMEROON 1059 Pseudohaje goldii (Boulenger, 1895) Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68984, 69598, 75163, 77629, 77665-6, 77897, 78028-31, 78880-6, HWH 370-9, 682- 4, 725-6, 999. Habitat: F, FB. VIPERIDAE Atheris squamigera (Hallowell, 1854) Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 75285-6, 77622-3, 78111-7, 78868-70, HWH 394-5, 692, 1183. Habitat: F. Causus lichtensteinii (Jan, 1859) Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 68959, 75164, 77624-8, 78075-6, 78896-901, HWH 396-8, 607-9. Habitat: F, FB. Causus maculatus (Hallowell, 1842) Localities: MtNL, NWF, EKA. Voucher specimens: ZFMK 75135, 77899, 78902, HWH 687-9, 740-8. Habitat: F, FB. Bitis gabonica (Duméril, Bibron & Duméril, 1854) Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 68983, 75168, 75331-3, 77894-6, 78070, 78086-91, 78903-9, HWH 367-9, 888, 902, 993-5. Habitat: F, FB. Bitis nasicornis (Shaw, 1802) Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75165-7, 75296-330, 75375, 77617-21, 78054-69, 78071-2, 78130, 78910-46, HWH 366, 606, 623-4, 685-6, 737-9, 903-24, 987-92. Habitat: F, FB. DISCUSSION We recorded a total of 89 reptile species from Mt Nlonako (table 1 and 3). Thirty-six percent of all 249 reptile species (excluding sea turtles) and 42 percent of the 151 snake species recorded for Cameroon (LeBreton, 1999) occur at Mt Nlonako. Thus Mt Nlonako hosts the most species rich single-locality reptile and especially snake fauna in Africa (Böhme, 2000). For snakes Mt Nlonako at this stage possibly represents the most species rich single-locality in the word. The subsequent area of rep- tile species richness is represented by Korup NP approximately 130 km W of Mt Nlonako with 83 reptile species (Lawson, 1993). This is followed by the Dja Faunal Reserve in Cameroon (80 species, LeBreton, 1999). Relative to snakes Mt Nlonako is again followed by the Korup NP (55 species), Mt Nimba in Guinea (52 species, Ineich, 2003), the Dja Faunal Reserve (49 species, LeBreton, 1999), the Dimonika forest site in Congo-Brazaville (45 species, Trape, 1975), the Comoe NP in Ivory Coast (44 species, Rödel 1995 and 1999), the Ziama Forest in Guinea (41 species, Böhme, 2000) and 40 species found in the Tai NP in Ivory Coast (Rödel & Mahsberg, 2000; Ernst & Rödel, 2002). Four snake species assemblages, reported from Ivory Coast, Ghana, Gabon and the Central African Republic, ranged between 35 and 39 species (Böhme, 2000). These results may not only represent the actual pattern of snake species richness in the rainforest areas of West and Central Africa but may also indicate survey activi- ties and gaps as many areas with a very high potential for exceptional species richness have not been sampled adequately. 1060 H.-W. HERRMANN ET AL. TABLE 1. Reptile species richness of three different rainforest areas in southwestern and southern Cameroon. Numbers in parentheses are the number of species directly recorded by the authors for the areas. NP = National Park; FR = Faunal Reserve. From LeBreton’s list only species found or mentioned from the Dja FR or adjacent areas were included in this analysis. Mt Nlonako Korup NP Dja FR area in km? 150 1,240 8,400 reptile species 89 83 (69) 80 (14) reptile species/km? 0.59 0.07 0.01 snake species 63 55 (41) 49 (10) snake species/km? 0.42 0.04 0.01 source this study Lawson, 1993 LeBreton, 1999 TABLE 2. West and Central African rainforest reptile and snake faunas and their relationships with the Mt Nlonako reptile and snake fauna. country locality reptile CBR snake CBR source species species Cameroon Korup National Park 83 0.5 55 0.72 Lawson, 1993 Congo (RC) Dimonika region - - 45 0.70 Trape, 1985 Cameroon Dja Faunal Reserve 80 0.60 49 0.66 LeBreton, 1999 Guinea, Ivory Coast, Mt Nimba 69° 0.51 52 0:57 Ineiche2003 Liberia Gabon Mounts de Cristal 48 0.54 32 0.55 Pauwels et al., 2002 Gabon Massif du Chaillu 46 047 29 0.50 Pauwels et al., 2002 Guinea Zima forest 46 043 41 0.50 Böhme, 2000 Ivory Coast Tai NP - - 40 0.50 Rödel & Mahsberg, 2000, Ernst & Rédel, 2002 Cameroon Mt Kupe & Bakossi 38 047 18 0.35 Schmitz, 1998 Mtns Ivory Coast Haute Dodo 17040:23 9 0.19 Branch & Rödel, 2003 Within Cameroon, our data shows not only the highest species richness for a single locality, but also the highest number of species per area (km?) as presented in table 1. Of the 89 species recorded for Mt Nlonako, one species is an amphisbaenid (1%), one is a crocodile (1%) and two are tortoises (2%). Of the 22 lizard species (25%) one is an agamid (5%), one a lacertid (5%), one a varanid (5%), five are geckos (23%), six are chamelons (26%), and eight skinks (36%). The 63 snake species (71%) include two boids (3%), three elapids (5%), four typhlopids (6%), five vipers (8%), ten atractaspidids (16%), and 39 colubrids (62 %). The lizard fauna appears depauperated compared to lizard faunas in arid or semiarid African environments. However, similar findings are presented by Böhme (2000) for the Zima Forest in Guinea and Branch & Rödel (2003) for areas in western Ivory Coast. 1061 THE REPTILES OF MT NLONAKO, CAMEROON + Di ++ Ipupusaf vuOSOSKT] + +++ SLAQD]YNIDU S1dajKYIVAT, AP iti +++ smuiffp sıdojky2DA] AVGIGNIO§ sli == Tap Npsof D19W0104 AVOLLITOVT È + + INDAUOD SNIAJIDPOBAT + +++ pınogpu SN]{2DPIUIH + de + snjp19spf SN]{I2DPIWIH + + snulyra SN]{1J9DprudH + ++ SI]DUNSAJUI SN]{1J9DprutdH AVAINOHHAO) + +++ wun11294S uoajoydupyy + ++ 112ffofd oajapwpy) > + IU2MO 02]2PUDUyJ fn, +++ wmmuou 02]2DUDY) + +++ SNIDISIAD OaJADUUDYD + + SISUIUNAIUDI 02]2PUDYJ AVCAINOTTIVINVHI + +++ DUDSD DUDÈY AVCAINVOY VIANVS + 4 + DUDIUIOY SAXIULY + + + DS0.49 SAXIUIY AVOINICALSAT, VINOTIHI) Ar aa + sıdsp.1101 SNWAV]OIISE AVAITACODOND VITAGODOND di + ‘ds s1]adouoyy AVCINAVASIHANY VINHVASIHAINY VSS VO VA 49 WOOL | WOOT | WOOL WOOS gUOHNgLNSIP o1ydess093002 zUONNQLISIP [eurpnynye joourpunge UX} "OXBUOIN IAL Worf sorsods apndos Jo voy ur Uonnquasip 91yde18098007 pur OYEUOIN IN IV UONNQLYSIp jeumpmmpe sourpunqy ‘€ MIAVL H.-W. HERRMANN ET AL. 1062 +++ + + + + + + +++ + + + + + + + + + + ++++++ + + + + + + + + é é é é VSS VO VA gUONNALISIP o1yde18098007 49 WOOL I WOOT" I WOOL wos zUOHNqINSIP euipmnje ib G ++ é é ar ++ +++ — — —— ++ 9 ouepunge ADSADI DUAL) ixnpssniq siydojouolu0y DUIPIDADWS SIXKdOAISDH) 119]12M Dogopvsdiq sıpıaa Dogopvsdiq 4ojoo1un pogopvsdiq 1poomaapun pogoppsdic] lausayonp pogoppsdic] DAQDIS SYJaddsvg DIDDSD[ su]jadasp(q SHIDIOSDS snoakjapuvy yD snauunig snujpoyjydoiyjog pJuamnada]jnd vsiog 13upun]jq Dd10g sndoos0uD X1AJDUOALY AVALHAINTOD) apgas UOYIKd 11]PADYU1dYA DIAPGD]DI avalog snypjound sdo]yd«a], snpjoaun sdojydäL sn]s28uo9 sdoyydA] sisuajosup SdojydK] AVOIdOTHdA L SHLNAdHAS SHIDULO SNUDADA AVGINVUVA ‘dds soydpisojdaT UNANAISHUTTIA SOYdDISO1d27] 19PYyoA4 S1d$D}4990] IMOUBYIIAA SIASDJABIVDT Sd291491q Sidspuvd exe] 1063 THE REPTILES OF MT NLONAKO, CAMEROON S14D]]09 UOW2]04 AVAIQIASVLOV' ALT sisuaod DAJ2Y2IN 1D4InS DA]QYAW NYIAUS DIADAD + +++ + + DID] NINA SIdSDIID.AYV + + + s11D]n82441 SIUSDIIDAY + + 14asuajnog S1dsDJIDANV + + i é i + DUNAAIID SIUSDIIDALY + é Ö it + 4231U snjopjyjpavdy + + +++ snısapow snjapjjpavdy + + ++ sııpjnSiapyf sdospay J de ats tte HDUDIEEIESIUACIOICUT a ++ nsopq siydoumpyy + + + pssidoiyjap siydouupyy + i if i + SISUIUOABUDI SIOYAIIIIOg + a é é é ar SAP SHULD YONG ti + + sati SNUAIPOLIIAY SNUWDIYIO]IYK + + +++ SNIDULADI SNUWDYIONYT + = +++ DIDBAIADA SAAAPIIAION + tr DIIDAIO SA2NI1PN + + +++ saplouls yn S24INIIAIDN + + = = at snuypyiydouas vAjayapy + + 1Y2DqU25811 DAJAYAW + + ++ + + ++ ni a ++ 210.210] uoıpıydo9AT + + +++ SHIDSMA siydosduvT + + ++ sna9pano siydosduvT + + — — +++ snpaun SıydoAadıwmT + ++ snsouis1jnf sıydo4dumT + À è é + daspsounjau SdoyjavApaAy] + + i A } + snısopow SNIOUOW.IOH + + —_ — +++ snpaun staydopisdvy Jf + + VSS VO VM 49 WOOL'T WOOT'T WOOL woos gUOHNGLHSIP °1yde18098007 zUONNAINSIP Jeurpnynfe poourpunqe UX) H.-W. HERRMANN ET AL. 1064 "POLY ULIEUES-QNS = VSS ‘POLY [RNID = VO ‘POLY ISOM = VM ‘oruopuo OUR) = FI ‘7007 "Te 19 ZION UO posegg SAIS Apms 1olewu oy) 0) Spuods>1109 SII UL UONEA9]I ‘SUOTEAIISQO pue suouttoads JoyoNOA UO paseqz Jurpunge = +++ Jurpunge A[or.I9PoW = ++ “DEI = + ‘SUOTRAJOSGO pur SUdLUTdOds 19y9noA UO poseg] ++ + + + + + + + + + + VSS VO VA gUOHNALNSIP ITyde1309300Z qo WOOL WOOT I wool „uonngensip [eUIpNyHye +++ SIUAODISDU SHG bat DIAUOGDS SIG ++ SNID|NIDU SNSNDI) ++ NUIAISUAJYIT] SNSNDY ++ DAISNUDNDS STAIYIV AVONIAdI A +++ 1pjos alpyopnasd hte pan>2joupjau DID +++ luosaupl mosaupl sidspospuaq AVAIdV 19 + SNIDJOU UOW2]0 cf SIPIDAS UOWA]Od + SISUIUOGDS UOWA]Od WOQOS poourpunge exe) THE REPTILES OF MT NLONAKO, CAMEROON 1065 The distribution of species along an elevational gradient shows that most species are found at lower elevations (table 3). Only a few species can be considered purely montane such as Chamaeleo pfefferi and three skink species (Lacertaspis rohdei, Leptosiaphos vigintiserierum, Leptosiaphos spp.). In a broader biogeographic context, 51 of the reptile species of Mt Nlonako can be considered West / Central African in distribution (table 3). Only two species have a pure West African distribution, of which one, Aparallactus niger is recorded here for the first time in Cameroon and Central Africa and had its eastern distribution border in Ivory Coast. Twenty-one are distributed in Central Africa only. Ten of the recorded species are endemic to Cameroon. Eight species, Agama agama, Hemidactylus mabouia, Trachylepis maculilabris, Dasypeltis scabra, Lamprophis fuliginosus , Natri- citeres olivaceus and N. variegata, and Naja melanoleuca have a wider distribution in Sub-Saharan Africa. Overall, Mt Nlonako is dominated by a West / Central African reptile fauna. This is supported by the relatively high coefficient of biogeographic resemblence (CBR) as demonstrated in table 2. This is different to the biogeographic relationships of the amphibian fauna which showed closer relations to the Central African amphi- bian fauna as compared to the west African amphibian fauna (Herrmann et al., 2005). Here, the reptile faunas of the southwest Cameroonian Korup NP and the south Cameroonian Dja Faunal Reserve show a high degree of resemblance. As for snakes the CBR’s exhibited by the Korup NP, the Dimonika region in Congo-Brazzaville, the Dja Faunal Reserve and Mt Nimba in West Africa indicate the highest resemblance. The exceptional species richness and high degree of endemicity in lizards of Mt Nlonako and the West Cameroonian mountain range can be explained by palaeogeo- graphic events. Historically this area has served as a refuge during drastic climate fluc- tuations. During the Pleistocene circa 20,000 years ago, African wet forests were restricted to a few isolated areas (Livingstone, 1982). Those fluctuations and refuges played an important role in the evolution of the high number of (endemic) reptile species as Lawson (1993) also describes for the Korup NP. Osteolaemus tetraspis, Kinixys erosa and K. homeana, all five Chamaeleo species, Varanus ornatus, Calabaria reinhardtii and Python sebae are listed in CITES appendices for which international trade is regulated. The only species listed as threatened is Osteolaemus tetraspis (vulnerable, [UCN 2003). Akani et al. (1998) pro- vide further insight into the local trade and consumption of this species in Nigeria; the situation in Cameroon can be considered very similar. The two tortoise species Kinixys erosa and homeana are listed as data deficient by the IUCN (2003). Lawson (2000) provides data on their conservation status in Cameroon. Osteolaemus tetraspis, Python sebae and Naja melanoleuca are protected by Cameroonian law. Threats to the Mt Nlonako reptile fauna are (1) habitat destruction by logging in the eastern and south- ern parts of the area, (2) habitat destruction by human encroachment as notable on the northern and western slopes of Mt Nlonako adjacent to the town Nkongsamba and (3) the hunting and consumption as food of species like Osteolaemus tetraspis, the two Kinixys species and Python sebae. 1066 H.-W. HERRMANN ET AL. ACKNOWLEDGEMENTS We thank ALSCO (American Linen Supply Company) Germany, especially Horst Nobis, for their generous financial support during the initial phase of this project and the continuation thereafter. The Zoological Garden Cologne, Germany, supported the work of H.-W. H.. The Cameroon Ministry of Scientific and Technical Research (MINREST) issued research permits and the Ministry of Environment and Forestry (MINEF) issued collecting and export permits. The Bundesamt fiir Naturschutz, Bonn granted import permits. We thank the traditional authorities of Mt Nlonako for per- mitting us to work in their tribal areas. The Worldwide Fund for Nature (WWF) Cameroon and the WWF Mount Kupe Forest Project staff helped with logistics, literature and information on local issues. In the field we acknowledge the efforts of Mirco Plath, Otto Mesumbe, and many other Cameroonian assistants; their knowledge of their forests and the animals therein helped make this survey possible. REFERENCES AKANI, G.C., ANGELICI, F.M. & POLITANO, E. 1998. Bushmen and herpetofauna: notes on am- phibians and reptiles traded in bush-meat markets of local people in the Niger delta (Port Harcourt, Rivers State, Nigeria). Anthropozoologica 27: 21-26. AMIET, J.-L. 1975. Ecologie et distribution des amphibiens anoures de la région de Nkongsamba (Cameroun). Annales de la Faculté des Sciences. Université de Yaoundé, Cameroun, 20: 33-107. BÖHME, W. 1975. Zur Herpetofaunistik Kameruns, mit Beschreibung eines neuen Scinciden. Bonner zoologische Beitrdge 26: 2-48. BÖHME, W. 1994a. Frösche und Skinke aus dem Regenwaldgebiet Südost-Guineas; Westafrika. I. Einleitung; Pipidae, Arthroleptidae, Bufonidae. Herpetofauna 16 (92): 11-19. BÖHME, W. 1994b. Frösche und Skinke aus dem Regenwaldgebiet Südost-Guineas; Westafrika. II. Ranidae, Hyperoliidae, Scincidae; faunistisch-6kologische Bewertung. Herpetofauna 16 (93): 6-16. BÖHME, W. 2000. Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes) (pp. 69-78). In: RHEINWALD, G. (ed). Isolated Vertebrate Communities in the Tropics. Proceedings of the 4" International Symposium, Bonner zoologische Monographien 46. BOHME, W. & SCHNEIDER, B. 1987. Zur Herpetofaunistik Kameruns (III) mit Beschreibung einer neuen Cardioglossa (Anura: Arthroleptidae). Bonner zoologische Beitrdge 38: 241-263. BRANCH, W.R. & RODEL, M.-O. 2003. 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Heidelberg. http://www.embl-heidelberg .de/~uetz/livingreptiles html REVUE SUISSE DE ZOOLOGIE Tome 112 — Fascicule 4 CERNY, Milo’. Additional notes on the fauna of Agromyzidae (Diptera) in SWACZET AN di NA RO O ne conc Tae En GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae in Paraguay, with descriptions of five new species (Diplopoda, Poly- AES OE ee oe EEA YOSHIZAWA, K., LIENHARD, C. & IDRIS, A. B. On the systematic position of Podopterocus Banks and Dinopsocus Banks, with a revised diagnosis of the genus Sigmatoneura Enderlein (Psocodea: IEsocopterazPsocidae) mann. ta he ren re EE ME es YANG, Ding & MERZ, Bernhard. Review of the species of Platypalpus Macquart from Guangxi, China (Diptera, Hybotidae, Tachydro- o ne a ERI A I I TAN e a ARS EBEJER, Martin J. A contribution to the knowledge of the Chyromyidae (Diptera) of Italy with description of a new species of Apha- MEQSOMAIBECCKEI REI OO ASI Monop, Lionel & LOURENÇO, Wilson R. Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new species and notes on biogeography and phylogenetic relationships ................. IVANOVA, Elena S. & VAUCHER, Claude. New genera and species of nematode parasites (Drilonematoidea: Ungellidae) from coelomic cavity of Neotropic acanthodrilids deposited in the Natural History MuseumonGeneva; Switzerland CREER RR SALGADO COSTAS, José Marfa. New species of Leiodidae (Coleoptera) and mnewarecords trom the Neotropical: Region ee eee MERZ, Bernhard & DAWAH, Hassan A. Fruit flies (Diptera, Tephritidae) from Saudi Arabia, with descriptions of a new genus and six new SDÉCIES RTE SII re OT ster oh OU. cele ants, Soa le ces SCHMID-EGGER, Christian. Proxiandrena subgen. nov. und Revision der west- und zentralpaläarktischen Arten der Andrena proxima- Gruppe (Hymenoptera, Apidae)... 0 0 ee HERRMANN, Hans-Werner, BOHME, Wolfgang, EUSKIRCHEN, Oliver, HERRMANN, Patricia A. & SCHMITZ, Andreas. African biodiversity hotspots: the reptiles of Mt Nlonako, Cameroon .............. Pages 771-805 807-830 831-848 849-857 859-867 869-941 943-961 963-982 983-1028 1029-1044 1045-1069 REVUE SUISSE DE ZOOLOGIE Volume 112 — Number 4 Pages CERNY, Milo’. Additional notes on the fauna of Agromyzidae (Diptera) in Switzerland Amen sms Ale RE 771-805 GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae in Paraguay, with descriptions of five new species (Diplopoda, Poly- CEE) eter else sete lO 807-830 YOSHIZAWA, K., LIENHARD, C. & IDRIS, A. B. On the systematic position of Podopterocus Banks and Dinopsocus Banks, with a revised diagnosis of the genus Sigmatoneura Enderlein (Psocodea: HPSOcopiérai: Bsocidae).. =. u... er ee 831-848 YANG, Ding & MERZ, Bernhard. Review of the species of Platypalpus Macquart from Guangxi, China (Diptera, Hybotidae, Tachydro- MIO I en Bunn MRIS IO 849-857 EBEJER, Martin J. A contribution to the knowledge of the Chyromyidae (Diptera) of Italy with description of a new species of Apha- MOSOMUMBECKET unsre. ee RE 859-867 Monop, Lionel & LOURENÇO, Wilson R. Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new species and notes on biogeography and phylogenetic relationships ................ 869-941 IVANOVA, Elena S. & VAUCHER, Claude. New genera and species of nematode parasites (Drilonematoidea: Ungellidae) from coelomic cavity of Neotropic acanthodrilids deposited in the Natural History Museumof Geneva, Switzerland." MIRTO ET ee 943-961 SALGADO COSTAS, José Marfa. New species of Leiodidae (Coleoptera) and new records from the: Neotropical-Revion tt Ie 963-982 MERZ, Bernhard & DAwAH, Hassan A. Fruit flies (Diptera, Tephritidae) from Saudi Arabia, with descriptions of a new genus and six new PISCINA ves en cet thors ee anal A ee ct a _ 983-1028 SCHMID-EGGER, Christian. Proxiandrena subgen. nov. and revision of the Andrena proxima group in the western and central Palaearctic Region! (Hymenoptera, Apidae) 2.2)... una eee 1029-1044 HERRMANN, Hans-Werner, BOHME, Wolfgang, EUSKIRCHEN, Oliver, HERRMANN, Patricia A. & SCHMITZ, Andreas. African biodiversity hotspots: the reptiles of Mt Nlonako, Cameroon .............. 1045-1069 Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Volume 112 - Number 4 - 2005 Revue suisse de Zoologie: Instructions to Authors The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies that it has been approved by all named authors, that it reports their unpublished work and that it is not being considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of colour plates and large manuscripts. 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