Revue Suisse de Zoology V111-1 2004

LU O O o o N LU o \I_ES de la SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSÉUM de la Ville tome D'HISTOIRE NATURELLE de Genève 1 fascicule 2004 > o I I I o o N LU o CD LU O > LU LU O > uu C/5 CD J L GEf JEVE MARS 2004 ISSN 003 - 418 X C/) HÉK REVUE SUISSE DE ZOOLOGIE TOME 111—FASCICULE Publication subventionnée par: Académie suisse des Sciences naturelles ASSN Ville de Genève Société suisse de Zoologie VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève CHARLES LIENHARD Chargé de recherche au Muséum d'histoire naturelle de Genève Comité de lecture Il est constitué Muséum en outre du président de de Genève et la Société suisse de représentants des instituts de Zoologie, du directeur du de zoologie des universités suis- ses Les manuscrits sont soumis des experts d'institutions suisses ou étrangères selon le sujet étudié La préférence graphie, sera donnée aux travaux concernant les domaines suivants: biogéo- systématique, évolution, écologie, éthologie, morphologie et anatomie comparée, physiologie Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE Internet: http://www.ville-ge.ch/musinfo/rnhng/page/rsz.htm Prix de l'abonnement: SUISSE Fr 225.— UNION POSTALE Fr 230.- (en francs suisses) Les demandes d'abonnement doivent être adressées la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P 6434, CH-1211 Genève 6, Suisse Revue suisse de Zoologie 111 3-10; mars (1): 2004 New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt Arayat, Luzon Island, Philippines Samuel W JAMES \ Yong HONG * & Tae Heung KIM Department of Life Sciences, Maharishi University of Management, Fairfield, IA 52557, U.S.A Faculty of Biological Resources Science, College of Agriculture, Jeonbuk National University, Jeonju 561-756, Republic of Korea New earthworms of Pheretima and Pithemera (Oligochaeta: Megafrom Mt Arayat, Luzon Island, Philippines - Three new of Pheretima and two new species of Pithemera are described from scolecidae) species Mt Arayat: Pheretima arayatensis sp n., Pheretima simsi sp n., and Pheretima castilloi sp n., Pithemera rotunda sp n., and Pithemera philippinensis sp n Pheretima arayatensis sp n., Pheretima simsi sp n., and Pheretima castilloi sp n have spermathecae in segment ix, vi, and vi-viii, respectively Pithemera rotunda sp n., and Pithemera philippinensis sp n have spermathecae in segment v-ix The former species has numerous circular genital markings, but the latter Pithemera species has no genital markings Descriptions of the new species are provided, including illustrations of the ventral view, male pore region, and spermathecae Key- words: Earthworms Oligochaeta - Mt Arayat Pheretima - Philippines - - - Pithemera taxonomy - Megascolecidae - INTRODUCTION is an isolated volcanic mountain in the large plains of central Luzon, Its forested upper elevations have always been isolated from other Mt Arayat north of Manila mountain ecosystems, such as the central Cordillera to the north and west, the Caraballo Range to the north, and the southern Sierra Madre to the east The remainwhose habitats would other- ing forests of Mt Arayat are an important refuge for biota wise be destroyed in the region now Its biogeographic importance as an isolated peak accentuated by deforestation and conversion of land to intensive agriculture part of a biotic survey of the from Mt Arayat in is As earthworms of the Philippines, specimens were collected March, 2001 In this paper, we report three new species of Sims & Easton, 1972 James (2004) described 21 new species of Megascolecidae of which 18- were new species of the genus Pheretima from the Mt Kitanglad Range, Mindanao Island He new Pheretima Kinberg, 1867 and two followed the systematics of Sims * To whom & species of Pithemera Easton (1972) These authors assigned species correspondence and reprint requests should be addressed E-mail: geoworm@hanmail.net Manuscript accepted 27.10.2003 W JAMES ETAL S with the intestinal caeca(um) originating in or near xxii to this genus, Pithemera They also defined four species groups within Pithemera, one of which containing the generotype Pithemera bicincta (Perrier, 1875) is the bicincta group, We report new species of the bicincta group herein Museum Holotypes and paratypes are deposited in the National of Annelids of (NMA) Paratypes are deposited in the Field Museum of Natural History (FMNH), and the Museum of Natural History of Geneva (MHNG) the Philippines DESCRIPTIONS & Pheretima arayatensis James Hong Figs sp n 1A-B Material: Holotype: One clitellate (NMA 0003741): Philippines, Luzon Island, Pampanga Province, Mt Arayat (15°12.29'N, 120°43.25'E), 410 m, litter layers in forest, 14 A Castillo colls paratypes: clitellate (NMA 0003746), clitellate March 2001, Y Hong & (MHNG (FMNH 10012), clitellate data as for holotype, clitellate, 43.90'E), 330 m, litter 34807): Same data as for holotype Other material: Same 18 aclitellate specimens; Mt Arayat (15°12.34'N, 120° layers in forest, Etymology: The species is clitellate, named for its semiclitellate, 13 March 2001 type locality Diagnosis: Spermathecal pore in 8/9, brown dorsal pigment, genital marking absent Brown Description: segment x, 7.0-7.2 dorsal pigment Dimensions 88-141 mm at xxx, 5.1-5.5 mm at clitellum, drical in cross-section Setae regularly distributed ing 54 at vii, 87 at xx; AA:AB:YZ:ZZ = 0.3 8/9, mm around segmental equators, number- 5-6 between male pores, size, distance regular; setal formula 2:1:2:4 at xiii First dorsal pore 12/13, furrow mm by 5.8-6.3 mm at segments 96-115; body cylin- Clitellum annular xiv-xvi; setae invisible externally spermathecal pore midventral deep in intersegmental with thickened surrounding tissue, monothecate Female pore single in xiv, oval shape, 0.5 between male pores 2.1 mm mm openings of copulatory bursae paired in Septa 5/6-7/8 thick, 8/9 absent, 9/10 10/11, not to body xviii, distance Genital marking lacking thin, membrane connected only a small to wall, 10/11-13/14 thick, muscular Gizzard large in viii-x, intestine begins in xv, small paired lymph glands from xxviii along dorsal vessel; intestinal caeca simple, originating in xxvii, and extending anteriorly to a finger-shaped sac; typhlosole low simple fold from xxii, each consisting of xxvii Hearts x-xiii esophageal; ix lateral, right side large, left very reduced Ovaries and funnels in xiii, single spermatheca in ix with nephridia on sper- mathecal ducts; spermatheca with small ampulla, ducts shorter than ampulla, diverticula each composed of separate chambers on single stalks or double stalks termi- nating in two chambers, club-shaped in ventral paired sacs in x, xi pseudovesicles xiii, Male sexual system holandric, testes and funnels Seminal vesicles two pairs in xi, xii with dorsal lobes, prostates in xviii, with stout muscular duct entering center of the copulatory bursae without stalked glands Vas deferens muscular xiv-xviii Pads or other structures like penis on inside of copulatory bursae Remarks: The species shares the brown dorsal pigment, dimensions, setae, and monothecate condition with Pheretima ambonensis Cognetti, 1913 from Ambon, Indonesia Pheretima arayatensis has spermathecal diverticula each composed of PHERETIMA AND PITHEMERA FROM MT ARAYAT, PHILIPPINES Fig A-B Pheretima arayatensis sp n A: ventral view; B: spermathecae C-D Pheretima simsi sp n C: ventral view; D: spermathecae E-F Pheretima castilloi sp n E: ventral view; F: spermathecae Scale bars = (A, C, E), (B), (D, F) mm mm mm separate chambers on single stalks or double stalks terminating in P ambonensis has a thick duct and only diverticula on single arayatensis has its spermathecal pore in 8/9 and spermatheca in has the spermathecal pore in 7/8 and spermatheca in Pheretima simsi James & Hong two chambers, but stalks ix, Also Pheretima but P ambonensis vii Figs sp n 1C-D Material: Holotype: One clitellate (NMA 0003742): Philippines, Luzon Island, Pampanga Province, Mt Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 13 March 2001, Y Hong O Castillo colls paratypes: clitellate (NMA 0003747), clitellate & (FMNH (MHNG 10013), clitellate 34808): Same data as for holotype Other material: Same data as for holotype, clitellate, aclitellate specimens; Mt Arayat (15°12.34'N, 120°43.90'E), 330 m, litter worm systematics layers in forest, clitellate, aclitellate, 13 Etymology: Named for Dr R March 2001 W Sims, who made good contributions to earth- Diagnosis: Spermathecal pores in 5/6 Diverticulum, as long as ampulla, chamber and stalk showing spermatozoal iridescence, nephridia covering duct-ampulla junction, part of ampulla Description: segment x, 2.8-3.1 Brown dorsal pigment Dimensions 68-91 mm at xxx, 2.7-3.0 mm at clitellum, mm by 2.6-2.8 mm at segments 82-97; body cylin- S W JAMES ETAL drical in cross-section Setae regularly distributed ing 31-33 39-47 at vii, at xx; mula AA:AB:YZ:ZZ = around segmental equators, number- 6-8 between male pores, size, distance regular; setal for- 1.5:1:2:3 at xiii Clitellum annular xiv-xvi; setae invisible externally pore 11/12, 12/13, spermathecal pores in 5/6, First dorsal ventrum, white color, with slightly protuberant mm oval shape, 0.5 male pores 2.5 mm at lateral Female pore single openings of copulatory bursae paired in xviii, margin of in xiv, 0.4 mm distance between Genital marking lacking 10/11-12/13 thick, muscular, Septa 5/6-8/9 thin, 9/10 absent, Gizzard usual in lips viii, intestine begins in xv, small paired 13/14 thin lymph glands from xxviii along dorsal vessel; intestinal caeca simple, originating in xxvii, and extending anteriorly about to xxvi, each consisting of a finger-shaped sac; typhlosole fold from low simple xxvii Hearts x-xiii esophageal; ix lateral, viii to gizzard, vii lateral, with branches to anterior gizzard wall Ovaries and funnels in xiii, paired spermathecal in vi with nephridia on sper- mathecal ducts; spermatheca with small ampulla, duct very short, diverticulum, as long chamber and as ampulla, stalk showing spermatozoal iridescence, nephridia covering duct-ampulla junction, part of ampulla Male sexual system holandric, testes and funnels in ventral paired sacs in x, xi Seminal vesicles two pairs in xi, xii with narrow dorsal lobes, prostates in xvii- xix, with stout muscular duct entering center of the copulatory bursae without stalked glands Copulatory bursa opening flanked by anterior, posterior pads, male pore within large convoluted opening on roughly conical protrusion from copulatory bursa roof Remarks: Pheretima simsi keys which is composed of two to the urceolata group in Sims species, P urceolata Horst 1893 & Easton (1972), and P baweanensis 1924 Gates (1961) synonymized these two species and P ditheca Michaelsen, Michaelsen, 1928 The species shares the lack of setae in the clitellum, 31 setae per segment differs at vii, from P ing short caeca activity, as and spermathecal pores in 5/6 with P urceolata was The locality of Pheretima simsi the collection site of P and has P close to is urceolata Pheretima kitangladensis (James, 2004), but setal gaps, Pheretima simsi urceolata in having as the ampulla long as the diverticulum, and in hav- human The present disturbance and species is similar to kitangladensis has no dorsal or ventral vertical lamellae in the esophagous Also, P kitangladensis is quite remote from human disturbance at a high elevation site (2250 m), but Pheretima simsi and P urceolata were collected from low and mid- elevation sites (330-1050 m, 1100 m) P urceolata was collected from Sumatura, Kupang, Timor, Indonesia and kitangladensis from Mt Kitanglad Range, Pheretima castilloi James & Hong Mindanao P Island, Philippines Figs 1E-F sp n Material: Holotype: One clitellate (NMA 0003743): Philippines, Luzon Island, Pampanga Province, Mt Arayat (15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, 13 March 2001, Y Hong & A Castillo colls paratypes: clitellate (NMA 0003748), clitellate (FMNH 10014), clitellate (MHNG 34809): Same data as for holotype Other material: Mt Arayat (15°12.41'N, 120°44.45'E), 1050 m, March 2001 litter layers in forest, clitellate, aclitellate, 13 PHERETIMA AND PITHEMERA FROM MT ARAYAT, PHILIPPINES Etymology The species is named after the type's collector, Augusto Castillo Diagnosis: Spermathecal pores in 5/6-7/8, genital marking absent mm by 3.1-3.3 mm mm at xxx, 3.0-3.3 mm at clitellum, segments 64-81 body cylin- Description: Brownish dorsal pigment Dimensions 59-81 at segment x, 3.2-3.3 drical in cross-section Setae regularly distributed ing 26-28 at vii, 46-50 mula AA:AB:YZ:ZZ = around segmental equators, number- 7-12 between male pores, at xx; 1:1.5:2:6 at xiii size, distance regular; setal for- Clitellum annular xiv-xvi; setae invisible externally First dorsal pore 12/13, paired spermathecal pores in 5/6-7/8, of ventrum, distance between spermathecal pores 3.0 0.4 mm mm oval shape, 0.4 between male pores 2.5 Septa 5/6-7/8 mm mm Female at lateral margin pore single in xiv, openings of copulatory bursae paired in xviii, distance Genital marking lacking thin, 8/9 absent, 11/12-13/14 thin Gizzard usual in 9/10 very thin, attached to 10/11, 10/11 thick, viii-x, intestine begins in xv, small paired lymph glands from xxviii along dorsal vessel; intestinal caeca simple, originating in xxvii, and extending anteriorly about to xxiv, each consisting of a small finger-shaped sac; typhlosole very low simple fold from xxvii Hearts x-xiii esophageal; ix lateral, very small viii to gizzard, vii lateral, extra esophageal vessel pharynx to ventral esophageal, then to clitellum area xiii to xiv, xv Ovaries and funnels in xiii, paired spermatheca in vi-viii, viii with nephridia on spermathecal ducts; spermatheca with small ampulla pouch, duct shorter than ampulla, diverticulum golf club-shaped, shorter than ampulla testes and funnels in ventral paired sacs in x, xi with dorsal lobes, prostates in xvi- xviii, Male sexual system holandric, Seminal vesicles two pairs in xi, xii with stout muscular duct entering center of copulatory bursae without stalked glands Copulatory bursa opening flanked by circular pads anterior, posterior, pads long slit backed by glandular chambers, male pore within on side of conical protrusion from copulatory bursa Remarks: In Sims roof & Easton (1972) Pheretima castilloi keys to a couplet offer- ing a choice between one and four thecal segments, corresponding to the urceolata and lejokana (genital marking present), darnleiensis (genital marking absent) species groups, respectively However, the present species has three pairs of spermathecal pores in 5/6-7/8 Therefore Pheretima castilloi Pithemera rotunda James & Hong is unlike all known Pheretima Figs sp n 2A-B Material: Holotype: One duellate (NMA 0003744): Philippines, Luzon Island, Pampanga Province, Mt Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 13 March 2001, Y Hong & A Castillo colls paratypes: clitellate (NMA 0003749), clitellate (MHNG (FMNH 10015), clitellate 34810): Same data as for holotype Other material: Same data as for holotype, clitellate, 15 aclitellate specimens; Mt Arayat (15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, clitellate, aclitellate, 13 March 2001 Etymology: The name rotunda is Latin for circular, referring to the shape of the genital marking Diagnosis: Spermathecal pores in 4/5-8/9, male pores in xviii, on 0.3 mm small hardened slightly protuberant circular porophore Genital markings numerous circular within thickened epidermal area S W JAMES ETAL FlG A: ventral view; B: spermathecae C-D Pithemera philippinensis (A), (B, D) C: ventral view; D: spermathecae Scale bars = (C), A-B Pithemera rotunda sp n mm Description: 2.0-2 body sp n Worm unpigmented mm mm mm (aclitellates 25-32) by mm at clitellum, segments 79-91; Dimensions 31-44 mm at segment x, 2.3-2.6 mm at xxx, 1.9-2.0 cylindrical in cross-section Setae regularly distributed around segmental equa- tors, numbering 49-54 setal formula at vii, 48-61 AA: AB: YZ:ZZ = visible, dorsal at xx; between male pores, size, distance regular; 2:1:1:3 at xiii Clitellum annular xiv-l/2xvi; xvi setae pore invisible externally First dorsal pore 12/13, five pairs of spermathecal pores in 4/5-8/9, very small mm oval shape, mm small hardened slightly protuberant circular porophore, white spot in lateral margin of ventrum Female pores paired in xiv, 0.3 male pores pores 1.9 in xviii, on 0.3 mm apart Circular raised genital markings numerous; on right side xviii, between male pores xviii, presetal in xix, xx, xxi, xxii; genital markings within white thickened epidermal area Septa 5/6 some muscular intestinal thin, 6/7 thick, 7/8 thin, 8/9 absent, 9/10, 10/11 thin, 11/12, caeca simple, originating in xxii, and extending anteriorly about consisting of a small triangle-shaped sac; typhlosole about 1/3 fold from xxii Hearts x-xii esophageal; ix Ovaries and funnels in ducts; 12/13 thickening, 13/14 thin Gizzard usual in viii-ix, intestine begins in xv; xiii, to xxi, each lumen diameter simple lateral, viii absent, vi, vii lateral spermathecae in v-ix; no nephridia on spermathecal spermathecae with small strawberry-shaped ampulla, ducts shorter than ampulla, diverticulum club-shaped, shorter than ampulla Male sexual system holan- U 232 UTIGER & B SCHÄTTI Fig Hierophis cypriensis: Lateral view of right hemipenis of MHNG 2618.99 (roadkill) Photograph by Claude Ratton frogs, i.e., Rana cf bedriagae Camerario (see Plötner et al., 2001) Blosat (1998) observed one specimen that captured a "Lacertide", probably Lacerta laevis troodica Werner, and an incidence of ophiophagy (Natrix natrix cypriaca) The holotype regurgitated a centipede In captivity, H cypriensis also preys on mammals, i.e., small mice (personal observation) Contrary to most congeneric species, Hierophis cypriensis is a placid snake that does not bite upon collecting In one instance, the Cyprus racer was found feigning litt.) Since this was observed on a road, it cannot be ruled out had driven over the specimen before being encountered by the correspondent death (W Teschner, in that a car THE CYPRUS RACER 233 Fig Known examined specimens with prePaphos (see Material and Methods and footnote 3) Open symbols correspond to four localities in the central Troodos (Kakopetria, Platania, and "ca km südlich Troodos") and eastern Paphos District (Stavros tis Psokas) cited in Osenegg (1989: Fig 44) as well as the Xyliatos dam area, an unspecified station near the type locality (Böhme & Wiedl, 1994: Fig 5), and Kampos and Tsakistra (see footnote 2) Elevations from 400 m to 1'200 m hatched, areas over 1'200 m in the central Troodos blank Map partly redrawn from Böhme & Wiedl (1994) distribution of Hierophis cypriensis Solid circles denote cise locality data and MHNG 2618.100 from the upper Livadi River, Zoogeography and relationships Böhme much & Wiedl (1994) presumed that Hierophis cypriensis had been "existing longer on Cyprus than the rest of the herpetofauna." Arguing that "snakes are generally primarily absent from oceanic islands", these authors though that "the case of Coluber cypriensis in particular would corroborate" the assumption of a land bridge This alludes to Haas (1952) who postulated a terrestrial connection across the Golf of Iskenderun in the Pliocene or early Quaternary (see, berger, 1979) However, there period Rather, Miocene e.g., Schmidt, 1960; Spitzen- no geological evidence for a land bridge during that from the surrounding mainland since the Upper of the Mediterranean (Hsii, 1972) as assumed by Schätti & Sigg Cyprus salinity crisis is is isolated (1989a) Hierophis cypriensis is most similar to H gemonensis and H viridiflavus with respect to certain osteological features (Fig 1) as, for instance, the comparatively long palatine processus of the maxilla (also found in H jugularis but not in H caspius) H cypriensis and H gemonensis have remaining species of this genus possess an enlarged spine (hook) more maxillary teeth (16-19 vs 12-16) than the These two species as well as H spinalis at the basis of the hemipenis (in prep.) U 234 UTIGER & B SCHÄTTI Based on morphological evidence and preliminary molecular data sequences), Schätti & "most closely related confirm to congeneric sister species status viridiflavus (mtDNA Utiger (2001: 934, 939) considered Hierophis cypriensis to be European species" ) The present results (Fig 4) of H cypriensis vis-à-vis the western Mediterranean H and H gemonensis, viz the Balkans and European whip snake Hierophis caspius Hierophis schmidti Hierophis jugularis Eirenis modestus Hierophis gemonensis Hierophis viridiflavus carbonarius (Gyaros) Hierophis viridiflavus carbonarius* 32 (Gyaros) 76 Hierophis viridiflavus carbonarius * (Dalmatia) Hierophis viridiflavus carbonarius* 57 (Sicily) 59 100 Hierophis viridiflavus viridiflavus (Sardinia) ' Hierophis viridiflavus viridiflavus — (France) Hierophis cypriensis Hierophis spinalis - Platyceps ventromaculatus Platyceps collaris Platyceps najadum 93 Spalerosophis diadema Hemorrhois hippocrepis -Hemorrhois ravergieri Hemerophis socotrae Coelognathus flavolineatus changes Fig Most parsimonious tree from a weighted MP analysis of two partial gene sequences, COI and 12S rDNA Numbers above branches indicate bootstrap values from an unweighted, those below from a weighted footnote MP analysis Asterisks denote specimens lacking COI sequence data (see also 3) MHNG & Schätti Utiger (2001) examined a fragment of the 12S rDNA subunit of 2618.100 (epidermal skin, see Material and Methods) The sequence shows only small difference (2 out of 592 base pairs) vis-à-vis 2618.99 The specimen was not used for the phylogenetic reconduction because the lack of the COI partition significantly lowered the resolution of the tree topology 3) MHNG THE CYPRUS RACER 235 Molecular data and considerable morphological divergence of Hierophis cypriensis (e.g., dorsal colour pattern, paravertebral reductions, processi of dentigerous bones, general shape of basisphenoid) from congeneric, and in particular the most closely related western Mediterranean, species support the theory of the isolation of Cyprus for the last approx 5,2 my Apart from Macrovipera schweizeri (Werner) from the Cyclades, Hierophis cypriensis is the only endemic Mediterranean insular snake species Hierophis viridiflavus (Lacépède, 1789) Coluber gemonensis gyarosensis Mertens, 1968 syn nov The systematic and status specific allocation of Coluber gemonensis gyaro- sensis Mertens, 1968 have been a matter of controversy (see Introduction) Specific status (Schätti, 1988) was accorded based on morphological differences vis-à-vis two whip snake species {Hierophis caspius, H gemonensis) distributed in the Aegean region Böhme (1993) considered the melanotic taxon from Gyaros to be the rarest European snake species Racers from Gyaros most strikingly resemble the European whip snake, Hierophis viridiflavus, in general appearance, habitus, and morphology (Schätti, 1988) In fact, the insular taxon cannot be distinguished from H viridiflavus and molecular data (Fig 4) clearly demonstrate that Coluber gemonensis gyarosensis Mertens, 1968 synonym of//, viridiflavus (Lacépède, 1789) The western Mediterranean Hierophis viridiflavus has a continuous range from the Pyrenees region (NE Spain) to northern France, Luxembourg, and western Switzerland as well as throughout the Appenine Peninsula, Ticino, and eastward to northern Dalmatia The species is widespread in the Tyrrhenian Sea, found on Sicily is a junior and many surrounding islands and The islets, Malta, Gozo, and in the Adriatic Sea (Krk, Aegean occurrence on Gyaros is most probably due to human introduction in historical times To conclude from our molecular data (Fig 4), H viridiflavus was most likely transported to Gyaros from the northern Adriatic region Gyaros (Gioura, Yioura), nowadays known as Yiaros or Nisos, is a mountainous island (up to 490 m a.s.l in the central part) of ca 20 km at 37°37'N 24°43'E in the Cyclades, roughly 15 km northwest of Syros about halfway between Kea and Tinos From the beginning of our era until the Middle Ages, the island was a haven for Tremiti Islands) isolated marauding corsairs who attacked merchant ships throughout the Aegean During the Roman Imperial epoch, Gyaros served as a place of exile All surrounding larger islands (viz., Andros, Kea, Kythnos, Syros, and Tinos) are inhabited by Hierophis caspius, an eastern Mediterranean species widespread Aegean Sea and found on all 1988; Szczerbak & Böhme, larger Cyclade islands surrounding Gyaros 1993) The absence of Gyaros may have favoured the colonization of the Caspian this island by H in the (e.g., Schätti, whip snake from viridiflavus External morphological characters of Hierophis viridiflavus analysed by Schätti & Vanni (1986) hardly allow a distinction between typical and melanotic populations commonly referred to H v carbonarius (Bonaparte) These authors considered phenotype as a colour morph of the European whip snake, H viridiflavus this Based on U 236 UTIGER & B SCHÄTTI molecular data, our limited sample shows genetic separation of typically coloured specimens (France, Sardinia) and melanotic individuals from Dalmatia (Krk), and Gyaros Pending studies using specimens from further regions, Island Sicily, this result may justify the revalidation of H Italy and Dalmatia, the southern Appenine Peninsula including islands such as Sicily v carbonarius for populations from northeastern and Malta, and the isolated Gyaros population ACKNOWLEDGEMENTS The junior author expresses his gratitude to Nicolas E Arnold (London), Wolfgang Böhme (Bonn), Franz Tiedemann (Vienna), and Yehudah L Werner (Jerusalem) for approving the loans of specimens in their respective care Theodore J Papenfuss (Berkeley, MIZ) and J Robert Macey (Walnut Creek) supplied tissue samples Lara Modolo, Wolfgang Scheffrahn, Claudia Zebib-Brunner (all Zurich), and Jennifer Pastorini (New York) supported the laboratory work, and Vincent Ziswiler and Paul Ward (Zurich) provided the facilities A most special thank you to Corinne Charvet (Geneva) for technical assistance We feel obliged to Walter Teschner (Wuppertal) and "Snake George" Hansjörg Wiedl (Paphos) for field observations on the Cyprus racer Regula Böniger, Claudia Fischer, Barbara Straub (all Zurich) helped with bibliographical research, and Beatrice Naef (Geneva) prepared some of the drawings REFERENCES Blosat, B 1998 Morphologie, Aut- und Populationsökologie einer Reliktpopulation der zypriotischen Ringelnatter, Natrix natrix cypriaca (Hecht, 1930) Bonn, Inaugural-Disseration, 174 pp Böhme, W 1993 Coluber gyarosensis Mertens, 1968 - Gyaros-Pfeilnatter (pp 111-114) In: Böhme, W (ed.) Handbuch der Reptilien und Amphibien Europas, Band 3/1 Wiesbaden, Aula-Verlag, 479 pp & Wœdl, H 1994 Status and zoogeography of the herpetofauna of Cyprus, with taxonomic and natural history notes on selected species (genera Rana, Coluber, Natrix, Vipera) Zoology in the Middle East 10: 31-52 Böhme, W Bonaparte, C L 1834 Coluber leopardinus unnumbered [4 pages] In: Iconografia della Fauna [1832-1841] IL Amfibi [Fase VII, Italica per le quattro classi degli animali vertebrati punt 38] Farris, J S 1989 Roma, The Salvinoci retention index and the rescaled consistency index Cladistics 5: 417-419 Felsenstein, J 1985 Confidence limits on phylogenies: an approach using the bootstrap Evolution 39: 783-791 Gruber, U 1989 Die Schlangen Europas und rund ums Mittelmeer Kosmos-Namrführer Stuttgart, Franckh 'sehe Verlagsbuchhandlung, 248 pp Haas, G 1952 Remarks on the origin of the herpetofauna of Palestine Revue de Sciences de l'Université d'Istanbul (B) 17 Hsü, K 1972 When Lacépède, B G Comte de II cum Paris, Hôtel de Thou, J American 227: 27-36 + 20 +144 + 474 tria naturae, pp secundum characteribus, differentiis, synonymis, locis Vol mata) Holmiae [Stockholm], L Salvii, Lyons-Weiler, 2001 Faculté des 1789 Histoire Naturelle des Quadrupèdes ovipares et des Linnaeus, C 1758 Systema naturae per regna species, la 95-105 the Mediterranean dried up Scientific E., Serpens Vol (2): RASA 3.0 Turbo TV + 823 [+ 1] classes, ordines, genera, I (editio decima, refor- pp for the Macintosh http://bio.uml.edu/LW/RASA.html THE CYPRUS RACER 237 Mertens, R 1968 Eine schwarze Zornnatter von den Cycladen: Coluber gemonensis gyarosensis n subsp Senckenbergiana biologica 49 (3/4): 181-189 Osenegg, K 1989 Die Amphibien und Reptilien der Friedrich-Wilhelms Universität, 200 pp Insel Zypern Bonn, Diplomarbeit, J., Ohst, T., Böhme, W & Schreiber, R 2001 Divergence in mitochondrial DNA of Near Eastern water frogs with special reference to the systematic status of Cypriote and Plötner, Anatolian populations (Anura, Ranidae) Amphibia-Reptilia 22 Sattou, N & Nei, M (4): The neighbor-joining method: a new method Molecular Biology and Evolution 4: 406-425 1987 logenetic trees Schätti, B 1985 Eine neue Zornnatter aus Zypern, Coluber cypriensis Zoologie 92 (2): 471-477 397-412 for reconstructing phy- n sp Revue suisse de Schatti, B 1986 Morphological evidence for a partition of the snake genus Coluber (pp 235238) In: Rocek, Z (ed.) Studies in Herpetology (Proceedings of the European Herpetological Meeting, 1985) Prague, Charles University, XXVIII + 754 pp Schatti, B 1987 The phylogenetic significance of morphological characters in the Holarctic racers of the genus Coluber (Reptilia, Serpentes) Amphibia-Reptilia (4): 401-415 Schatti, B 1988 Systematik und Evolution der Schlangengattung Hierophis Fitzinger, 1843 Zürich, Inaugural-Dissertation, 50 pp Schatti, B 1993 Coluber Linnaeus, 1758 - Zornnattern (pp 69-71) In: Böhme, W (ed.) Handbuch der Reptilien und Amphibien Europas, Band 3/1 Wiesbaden, Aula-Verlag, 479 pp & Schätti, B Sigg, H 1989a Die Herpetofauna der Insel Zypern Die herpetologische Erforschung und Amphibien Herpetofauna (Weinstadt) 11 (61): 9-18 Schätti, B & Sigg, H 1989b Die Herpetofauna der Insel Zypern Schlangen Herpetofauna (Weinstadt) 11 (62): 17-26 Schätti, B & Utiger, U Schildkröten, Echsen und new genus for Zamenis socotrae Günther, and a World racers, whip snakes, and related genera Squamata: Colubrinae) Revue suisse de Zoologie 108 (4): 919-948 2001 Hemerophis, a contribution to the phylogeny of Old (Reptilia: & Vanni, S 1986 Intraspecific variation in Coluber viridiflavus Lacépède, 1979, and validity of its subspecies (Reptilia, Serpentes, Colubridae) Revue suisse de Zoologie 93 (1): 219-232 Schätti, B & Schätti, B Wilson, L D 1986 Coluber Catalogue of American Amphibians and Reptiles 399: 1-4 Schmidt, W F 1960 Zur Struktur und Tektonik der Insel Cypern Geologische Rundschau 50: 375-395 Spitzenberger, F 1979 Die Säugetierfauna Zyperns Teil II: Chiroptera, Lagomorpha, Carnivora und Artiodactyla Annalen des Naturhistorischen Museums Wien 82: 439-465 Swofford, D L 1998 PAUP*: phylogenetic analysis using parsimony (and other methods) Version 4.0b 10 for Mac Sunderland, Sinauer Szczerbak, N N & Böhme, 1993 Coluber caspius Gmelin, 1789 - Kaspische Pfeilnatter oder Springnatter (pp 83-96) In: Böhme, W (ed.) Handbuch der Reptilien und Amphibien Europas, Band 3/1 Wiesbaden, Aula-Verlag, 479 pp W Utiger, U 2002 Molecular phylogeny of the Oriental colubrid genus Coelognathus Fitzinger 1843, and a contribution to the systematics of Holarctic racers and ratsnakes (Reptilia: Squamata: Serpentes) University of Zürich, Ph D thesis [part.], 73 pp Utiger, V U, Helfenberger, N., Schätti, B., Schmidt, C, Ruf, M & Ziswtler, 2002 Molecular systematics and phylogeny of Old and New World ratsnakes, Elaphe auct., and related genera (Reptilia, Squamata, Colubridae) Russian Journal of Herpetology (2): 105-124 U 238 UTIGER & B SCHÄTTI of specimens examined by Schätti & Utiger (2001) used in this Coelognathus flavolineatus (AY039200/AY039162); Eirenis modestus (AY039181/AY039143); Hemerophis socotrae (AY039167/ AY039129); Hemorrhois hippocrepis (AY039194/AY039156), H ravergieri (AY039193/AY039155); Hierophis caspius (AY039164/AY039126), H gemonensis (AY039183/AY039145), H jugularis (AY039188/ AY039150), H schmidti (AY039197/AY039159), H viridiflavus (AY039175/AY039137); Platyceps collaris (AY039195/AY039157), P najadum (AY039179/AY039141), P ventromaculatus (AY039172/AY039134); Spalerosophis diadema (AY039182/AY039144) APPENDIX Genbank numbers study (COI/12S) REVUE SUISSE DE ZOOLOGIE Tome 111 — Fascicule Pages James, Samuel W., Hong, Yong & Kim, Tae Heung New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt Arayat, Luzon 3-10 Island, Philippines Gil de Pertierra, Alicia A Redescription of Monticellia magna (Rego, dos Santos & Silva, 1974) (Eucestoda: Monticelliidae) parasite of Pimelodus spp (Pisces: Siluriformes) from Argentina, and morphological study of microtriches Loureiro, Marcelo, Azpelicueta, Maria de las Mercedes & Garcia, Gracida Austrolebias arachan (Cyprinodontiformes, Rivulidae), a new species of annual fish from northeastern Uruguay Puthz, Volker Edaphus comellinii sp n aus Sri Lanka (Coleoptera: i_ 20 21-30 Sta- phylinidae) 88 Beitrag zur Kenntnis der Euaesthetinen 31-33 & Fresneda, Javier Two new taxa of LeptoLeiodidae, Cholevinae) from the Cantabrian cornice (Asturias, Spain) Biogeographical observations 35-55 Salgado Costas, José Maria dirini (Coleoptera: Papp, Laszló Description of the (Diptera), first apterous genus of Sciomyzidae 57-62 from Nepal Pace, Roberto Specie nuove o poco note di Homalotini, Silusini, Bolitocharini, Diestotini e Autaliini della Cina e della Thailandia (Cole- 63-76 optera, Staphylinidae) Scheller, Ulf Pauropoda (Myriapoda) from Vietnam (Pauropoda and Geneva Museum XIII) Tilbury, Colin R & Mariaux, Jean A re-evaluation of Symphyla of 77-101 the the generic assignment of Bradypodìon spinosum (Matschie, 1892) and some considerations on the genus Rhampholeon Günther, 1874 De Chambrier, Alain, Rego, Amilcar A & Mariaux, 103-110 Jean Redescription of Brooksiella praeputialis and Goezeella siluri (Eucestoda: proteocephalidea), parasites of Cetopsis coecutiens (Siluriformes) from the Amazon and proposition of Goezeella dankbrooksi sp n 111-120 Kottelat, Maurice Salaria economidisi, a new species of freshwater fish from Lake Trichonis, Greece, with comments on variation in S fluviatilis 121-137 (Teleostei: Blenniidae) Schwendinger, Peter J & Martens, Jochen A taxonomic revision of the family Oncopodidae IV The genus Oncopus Thorell (Opiliones, 139-174 Laniatores) Baehr, Martin Colasidia wau, a new leleupidiine species from Papua Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae) Merz, Bernhard Revision of New 175-181 the Minettia fasciata species-group (Diptera, Lauxaniidae) Mirande, Juan Marcos, Aguilera, Gaston & Azpelicueta, Maria de las Mercedes A new species of Astyanax (Characiformes, Characidae) from the upper rio Bermejo basin, Salta, Argentina 183-211 213-223 Utiger, Urs & Schätti, Beat Morphology and phylogenetic relationships of the Cyprus racer, Hierophis cypriensis, and the systematic status of Coluber gemonensis gyarosensis Mertens (Reptilia: Squamata: Colubrinae) 225-238 REVUE SUISSE DE ZOOLOGIE Volume 111 — Number Pages James, Samuel W., Hong, Yong & Kim, Tae Heung New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt Arayat, Luzon 3-10 Island, Philippines Gil de Pertierra, Alicia A Redescription of Monticellia magna (Rego, dos Santos & Silva, 1974) (Eucestoda: Monticelliidae) parasite of Pimelodus spp (Pisces: Siluriformes) from Argentina, and morphological 11-20 study of microtriches Loureiro, Marcelo, Azpelicueta, Maria de las Mercedes & Garcia, Gracida Austrolebias arachan (Cyprinodontiformes, Rivulidae), a new species of annual fish from northeastern Uruguay 21-30 Sri Lanka (Coleoptera: StaKnowledge of Euaesthetinae 31-33 Puthz, Volker Edaphus comellinii from sp n phylinidae) 88th Contribution to the Salgado Costas, José Maria (Coleoptera: Leiodidae, Cholevinae) from the cornice (Asturias, Spain) Biogeographical observations dirini Papp, Laszló Description of the (Diptera), & Fresneda, Javier Two new taxa of Leptofirst Cantabrian 35-55 apterous genus of Sciomyzidae 57-62 from Nepal New or little known species of Homalotini, Silusini, Bolitocharini, Diestotini and Autaliini from China and Thailand (Coleoptera, Staphylinidae) Pace, Roberto 63-76 Scheller, Ulf Pauropoda (Myriapoda) from Vietnam (Pauropoda and Symphyla of the Geneva Museum 77-101 XIII) Tilbury, Colin R & Mariaux, Jean A re-evaluation of the generic assignment of Bradypodion spinosum (Matschie, 1892) and some considerations on the genus Rhampholeon Günther, 1874 De Chambrier, Alain, Rego, Amilcar A & Mariaux, 103-110 Jean Redescription of Brooksiella praeputialis and Goezeella siluri (Eucestoda: proteocephalidea), parasites of Cetopsis coecutiens (Siluriformes) from the Amazon and proposition of Goezeella dankbrooksi sp n 111-120 Kottelat, Maurice Salaria economidisi, a new species of freshwater fish from Lake Trichonis, Greece, with comments on variation in S fluviatilis 121-137 (Teleostei: Blenniidae) Schwendinger, Peter J & Martens, Jochen A taxonomic revision of the family Oncopodidae IV The genus Oncopus Thorell (Opiliones, 139-174 Laniatores) Baehr, Martin Colasidia wau, a new leleupidiine species from Papua Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae) Merz, Bernhard Revision of New 175-181 the Minettia fasciata species-group (Diptera, Lauxaniidae) Mirande, Juan Marcos, Aguilera, Gaston & Azpelicueta, Maria de las Mercedes A new species of Astyanax (Characiformes, Characidae) from the upper rio Bermejo basin, Salta, Argentina 183-211 213-223 Utiger, Urs & Schätti, Beat Morphology and phylogenetic relationships of the Cyprus racer, Hierophis cypriensis, and the systematic status of Coluber gemonensis gyarosensis Mertens (Reptilia: Squamata: Colubrinae) Indexed in Current Contents, Science Citation Index 225-238 — — PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTÉBRÉS DE LA SUISSE, (prix des fascicules sur N°s 1-17 (1908-1926) série Fr Echange ou par fascicule 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