{ | = © © bast © © N LL Er LL. N % m 7, LL un > LL a > © © = © O N LL. © ol < Z of =D © n N = n REVUE SUISSE DE ZOOLOGIE TOME 119— FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE Comité de rédaction JACQUES AYER Directeur du Muséum d’histoire naturelle de Genéve ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d’histoire naturelle de Genève Comité de lecture A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger (arthropodes excl. insectes). Le comité soumet chaque manuscrit pour Evaluation a des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, Evolution, morphologie et anatomie comparee. 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, Museum 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 BAY ISON LA iin a NM V f 7 u | \ + tome 119 (JAN IIZ Uld fascicule 4 N LIBRARIES 2012 & 3 2 kl GENEVE DECEMBRE 2012 ISSN 0035 - 418 X SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE REVUE SUISSE DE ZOOLOGIE TOME 119— FASCICULE 4 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE Comité de rédaction JACQUES AYER Directeur du Muséum d’histoire naturelle de Genéve ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d’histoire naturelle de Genéve Comité de lecture A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), M. Ruedi (mammiferes), A. Schmitz (amphibiens, reptiles), P. Schwendinger (arthropodes excl. insectes). Le comité soumet chaque manuscrit pour évaluation a des experts d institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. 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 Genève 6, Suisse REVUE SUISSE DE ZOOLOGIE 119 (4): 409-415; décembre 2012 Description of a new species of the genus Fessonia (Acari: Prostigmata: Smarididae) from Iran Safoura SALARZEHI!, Hamidreza HAJIQANBAR?*, Ali OLYAIE TORSHIZ! & Javad NOEI! l Department of Plant Protection, Jahad Daneshgahi Higher Education Institution of Kashmar, Iran. 2 Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, 14115-336, Tehran, Iran. *Corresponding author, E-mail: hajiqanbar@modares.ac.ir Description of a new species of the genus Fessonia (Acari: Prostigmata: Smarididae) from Iran. - Fessonia torshizica Salarzehi & Hajiqanbar n. sp. is described from larvae collected in the soil of vineyards in northeastern Iran. This is the second species currently known from larvae in the genus Fessonia. Some morphological characters of the new species are compared with its only congener, F! papillosa (Hermann 1804). Keywords: Parasitengona - Trombidiformes - Erythraeoidea - larva - Fessonia torshizica. INTRODUCTION Mites of the cohort Parasitengona (Acari: Trombidiformes) are a large and diverse group of prostigmatic mites distributed worldwide. These mites usually para- sitize invertebrates during their larval stage but the deutonymphs and adults are preda- ceous. The superfamily Erythraeoidea is one of the largest parasitengone superfamilies, consisting of two families: Smarididae and Erythraeidae. The family Smarididae, with 10 genera and 53 species (Zhang ef al., 2011), is a relatively small family usually found in grassland and litter or under the bark of trees (Southcott, 1996; Krantz & Walter, 2009). Mites of the family Smarididae are classified in two subfamilies, Smaridinae and Hirstiosomatinae, the former with five genera including Fessonia von Heyden, 1826. This genus is distributed in Europe, Australia, North America, Asia and Africa, and hitherto encompassed 10 known species, nine of which were described from adults. Previously the larval stage of only one species, Æ papillosa (Hermann 1804) from France, Italy, Croatia, Germany, Greece and Hungary was known (Wohltmann, 2010). During a faunistic survey of mites in vineyards of Kashmar and Torshiz counties (northeastern Iran) in 2010-2011, we found in soil samples larvae of a species of Fessonia which is new to science. This is the first record of the genus Fessonia from Iran. In the following we describe this new species and compare its morphological characters with those of the larva of F papillosa. We consider it very unlikely that the larvae of the new species are conspecific with adults of any of the previously described Fessonia species. Manuscript accepted 20.08.2012 410 S. SALARZEHI ET AL. MATERIALS AND METHODS The mite specimens were collected from litter and soil samples and extracted by Berlese funnel, then cleared in lactophenol and mounted in Hoyer’s medium. The morphology of the specimens was studied with an Olympus BX51 equipped with phase contrast and a drawing tube. All measurements in this description are given in micrometres. In the description, body measurements are given for the holotype and the paratype, the latter in parentheses. Terminology and abbreviations follow those of Grandjean (1947) and Wohltmann (2010). The holotype of the new species is deposited in the Acarological Collection, Zoological Museum, College of Agriculture, University of Tehran, Karaj, Iran. The paratype is deposited at the Natural History Museum of Geneva, Switzerland. The following abbreviations are used in the description: € (famulus) = speciali- zed smooth seta on tarsus of leg I, usually small; © (eupathidium) = specialized smooth seta on legs and palp, in particular on tarsus; z (accessory seta) = smaller seta close to dorsal eupathidium on tarsus of leg I; x (kappa) = microseta on tibia and genu of wal- king legs; o (sigma) = solenidion on genu; @ (phi) = solenidion on tibia; w (omega) = solenidion on tarsus; I = first walking leg; II = second walking leg; III = third walking leg; AL = first pair of non-sensillary setae on scutum in larvae; ASens = first pair of trichobothria on scutum in larvae; as (oral seta) = located anteriorly on gnathosoma; Bf = basifemur; bs (subcapitular seta) = located ventrally on gnathosoma; cs (adoral seta) = usually small smooth pointed setae located dorsally on gnathosoma; Cx (coxa) = epi- meral plate; Fe = femur; fn = number of setae; Ge = genu; IP (index pedal) = sum of all lengths of legs; N = non-specialized seta on palp and legs; PL = second pair of non- sensillary setae on scutum in larvae; PSens = second pair of trichobothria on scutum in larvae; Ta = tarsus; Tf = telofemur; Ti = tibia; Tr = trochanter. SYSTEMATIC ACCOUNT FAMILY SMARIDIDAE VITZTHUM, 1929 Genus Fessonia von Heyden, 1826 Fessonia torshizica Salarzehi & Hajiqanbar n. sp. Figs 1-7 MATERIAL STUDIED: Holotype larva (SS-2010-1a), free-living, collected from soil samples of vineyards in Torshiz, Khorasan Razavi province, Northeastern Iran, 35° 11' N, 58° 72' E, 1215 m. a.s.l. 24-IX-2010, leg. S. Salarzehi. Paratype: 1 larva (SS-2010-1b), same collection data as for holotype. ETYMOLOGY: The specific epithet refers to the type locality, Torshiz. DIAGNOSIS: Tarsus | with 26 serrate setae; genu I without microseta (k); genua I, II, and III each with one solenidion (0); basifemur I and telofemur I with 3 and 4 serrate setae, respectively; tarsus II with 24 serrate setae. DESCRIPTION OF LARVA: Metric data given in Table 1. Idiosoma (Figs 1-2). Length 316 (300), width 249 (221). Dorsal scutum sub- pentagonal, anterior border curved, with two pairs of barbed normal setae (AL, PL), two pairs of trichobothria (ASens, PSens), ASens almost setulose throughout but PSens setulose only in posterior half. Laterally of scutum two pairs of eyes. Remaining part of dorsum carrying 38 barbed setae in rows C-H. Venter with all setae barbed except elc I. Coxa I with setae 1b 52 (49) and la 19 (18), setae la located outside, close to A NEW SPECIES OF FESSONIA 41] TABLE 1. Morphometric data of larvae of Fessonia torshizica n. sp. Character Holotype Paratype Scutum lenght 61 58 Scutum width 65 61 ASens 53 47 ASens-ASens 16 15 PSens 84 14 PSens-PSens 19 18 AL 47 39 AL-AL 44 31 PL 68 61 PL-PL 65 55 Palpal Tr 8 7 Palpal Fe 21 20 Palpal Ge 1] 10 Palpal Ti 14 12 Odontus 10 8 Palpal Ta 8 8 Cx I 29 21 Tel 19 13 Bf I 28 22 Tf I 22 21 Gel 31 26 Til 42 38 Tal 39 31 bes | 208 172 Cx IT 35 23 Tell 22 14 Bf II 26 21 Tf II 22 19 Ge II 29 28 Ti II 38 34 Ta II 34 di Leg II 203 171 Cx HI 36 26 Tr HI 22 19 Bf III 21 22 Tf III 30 25 Ge III 33 32 Ti III 59 dI Ta III 40 35 Leg III 248 215 IP 659 558 margins of coxae I. Setae 2b 21 (19), situated on coxae II; setae 2a 44 (41), situated between coxae II. Setae 3b 35 (32), situated on coxae III; setae 3a 24 (22), situated anteriorly outside coxae III. Three pairs of setae situated posterior to coxae III. Gnathosoma (Fig. 3). Chelicera with cheliceral base 60 (64) long and with smooth movable claw; cheliceral digits short, 5 (4) long. Dorsal gnathosoma with a pair of smooth, pointed adoral setae cs 6 (5) anteriorly and with a pair of short, thick and blunt supracoxal setae e/cp 3 (2) in lateral position. Ventrally a pair of smooth and pointed subcapitular (tritorostral) setae bs 10 (8) and a pair of short, thick and blunt 412 S. SALARZEHI ET AL. Fics 1-3 Fessonia torshizica n. sp., larval holotype. (1) Dorsal view of idiosoma. (2) Ventral view of idio- soma. (3) Dorsal view (right side) and ventral view (left side) of gnathosoma. oral setae as 3 (2). Palpal femur with one barbed dorsal seta, 21 (19) long, palpal genu with one barbed dorsal seta, 11 (9) long. Palpal tibia with one smooth and two barbed setae. Odontus bifid, 9 (7) long. Palpal tarsus with four smooth setae gc 7 (6), ds 5 (4), ic 4 (3), hc 3 (2), one barbed seta nc 15 (14), one solenidion (w) and one prominent distal eupathidium (©). A NEW SPECIES OF FESSONIA 413 50um FIGS 4-7 Fessonia torshizica n. sp., legs of larval holotype. (4) Dorsal view of entire leg I. (5) Ventral view of tarsus of leg I. (6-7) Dorsal view of entire legs II and III. 414 S. SALARZEHI ET AL. TABLE 2. Leg chaetotaxy of larvae of Fessonia torshizica n. sp. and F. papillosa. F. torshizica n. sp. F. papillosa Palpal Tr ON ON Palpal Fe IN IN Palpal Ge IN IN Palpal Ti 3N 3N Palpal Ta SN LC To SN, LC PO Cx I IN, Lele IN, 1 elc Tr I IN IN Bf I 3N 2N Tf I 4N SN Ge I 8N, lo 8N,1o,1« Til 13 N, 3 g, 1 da, 1 «, I tr 13 N, 3 g, 1 da, 1 «, ltr Ta I 20N 26 IZ, le ta IX te 20-22N 26,12 lo, LE ATX 210 Cx ll IN IN Ir il IN IN Bf II 2N 2N Tf II SN SN Ge II 8Nl1lk,lo 8N, Ix,1o TH 14N,2© 14 N,2 © Ta II 24N,1C,1 21-22N,1C,1@ Cx III IN IN ie Il IN IN Bf III IN IN Tf II SN SN Ge III 8N.lo 8 N Ti HI I4N, Io I4N,1@ Ta II DINI 21-23 N, 17 Legs (Figs 4-7). Leg I 208 (172) long, femora I-III divided into basi- and telo- femur; segmentation formula: 7-7-7. Details of leg chaetotaxy in Table 2. Trichobothria present on tibia and tarsus of leg I. Anterior trichobothrial sensilum (ta) on tarsus I for- ming a complex with spatulate seta (x), microsolenidion (£), eupathidium (©) and seta z. Two trichobothria also located in proximal half of tarsus I. Tibia I with one tricho- bothrium in proximal half; shape of all trichobothria on leg I similar, with nude base and setulose upper half. A posterior solenidion on tibia I along with seta da. Solenidion (0) on genu I shorter than 1/2 length of other solenidia on leg I. Leg II 203 (171) long, anterior solenidion on tibia II longer than posterior one. Genu II with only pit of micro- seta K present. Solenidion (0) on genu II subequal to posterior solenidion () of tibia I. Leg HI 248 (215) long, solenidion (0) on genu III a little longer than solenidion (q@) on tibia III. All tarsi I-III with smooth and claw-like empodia; lateral claws pad-like, with small barbs. REMARKS: Fessonia torshizica n. sp. differs from F! papillosa in fnBf I (3 vs 2), fnTf I (4 vs 5), number of microsetae k on genu I (0 vs 1), fnTa I (26 vs 20-22), fnTa II (24 vs 21-22), number of solenidia on genu III (1 vs 0) and fnTa II (23 vs 21-23). All leg chaetotaxy characters of the larvae of Fessonia torshizica n. sp. and F! papil - losa are compared in Table 2. A NEW SPECIES OF FESSONIA 415 ACKNOWLEDGEMENTS We are grateful to Dr Alireza Saboori (Department of Plant Protection, College of Agriculture, University of Tehran, Karaj, Iran) for the confirmation of the new spe- cies and some advice on the figures. We also thank Abdolazım Mortazavı (Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran) for his help during preparation of the figures. Dr Ryszard Haitlinger (University of Wroclaw, Poland) kindly reviewed the manuscript of this paper. REFERENCES GRANDJEAN, F. 1947. Etudes sur les Smarisidae et quelques autres Erythroides (Acariens). Archives de zoologie expérimentale et générale 85(1): 1-126. KRANTZ, G. C. & WALTER, D. E. 2009. A manual of acarology. 3rd edition. Texas Tech University Press, 806 pp. SOUTHCOTT, R. V. 1996. Description of two new larval Smarididae (Acarina) from Australia. Records of the South Australian Museum 30(1): 1-12. WOHLTMANN, A. 2010. Notes on the taxonomy and biology of Smarididae (Acari: Prostigmata: Parasitengona). Annales Zoologici 60(3): 355-381. ZHANG, Z.-Q., FAN, Q. H., PESIC, V., SMITH, H., BOCHKOV, A. V., KHAUSTOV, A. A., BAKER, A., WOHLTMANN, A., WEN, T., AMRINE, J. W., BERON, P., LIN, J., GABRYS, G. & HUSBAND, R. 2011. Order Trombidiformes Reuter, 1909 (pp. 129-138). Jn: ZHANG, Z.-Q. (ed.). Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Magnolia press, Auckland, 237 pp. im REVUE SUISSE DE ZOOLOGIE 119 (4): 417-423; décembre 2012 A new species of Ceratophysella from Peru (Collembola: Hypogastruridae) José G. PALACIOS-VARGAS* & Thalia BOCANEGRA Laboratorio de Ecologia y Sistematica de Microartröpodos, Departamento de Ecologia y Recursos Naturales, Facultad de Ciencias, UNAM, 04510 Mexico, D. F., Mexico. *Corresponding author. E-mail: troglolaphysa@hotmail.com A new species of Ceratophysella from Peru (Collembola: Hypo- gastruridae). - Ceratophysella tupamara sp. nov. from Iquitos, Peru is described and illustrated. It is easy to distinguish from Ceratophysella denticulata (Bagnall, 1941), the only species of the genus previously cited from Peru, by its trilobed apical bulb of antennal segment IV, the number of unguicular teeth and the body chaetotaxy. Keywords: Iquitos - taxonomy - morphology - chaetotaxy. INTRODUCTION The genus Ceratophysella is distributed worldwide, and represented by more than 130 species (Bellinger er al., 2011) mainly in the Holarctic Region. From South America only three taxa are known: C. denticulata (Bagnall, 1941), C. armata (Nicolet, 1841) and C. communis (Folsom, 1897). Collembola samples from the Museum of Natural History of Geneva were provided to us for study. In that important collection there are several genera from many localities in South America (Palacios- Vargas et al., 2011). Among them we found some specimens of Ceratophysella from Peru that belong to a new species which is described in here. MATERIAL AND METHODS The description of the new species is based on material from the collection of the Muséum d’histoire naturelle de Genève (Palacios-Vargas et al., 2011). The spe - cimens were first cleared in 10% KOH, then in lactophenol, and mounted on micro- scopic slides in Hoyer’s solution. Drawings were done with a Carl Zeiss contrast microscope, using a drawing tube. The type material is deposited in the Muséum d’histoire naturelle de Genève (MHNG), and at the senior author’s institution, Facultad de Ciencias, Universidad Nacional Autonoma de México (UNAM). The terminology follows mainly Fjellberg (1998). Abbreviations are: Abd. = abdominal segment; Ant. = antennal segment; ar = anterior row; bm = basimedial; bl = basilateral; plb = proximal basilateral; mr = middle row; Oc. = ocular setae; Th. = thoracic segment; ss = sensilla; PAO = postantenal organ; pr = posterior row; px = proximal. Manuscript accepted 20.07.2012 418 J. G. PALACIOS-VARGAS & T. BOCANEGRA RESULTS Ceratophysella Borner, 1932 TYPE SPECIES: Podura armata Nicolet, 1841 DIAGNOSIS: Well pigmented Hypogastruridae of medium size. Setae differen- tiated in smooth microsetae or barbulate macrosetae. Seta m2 on Th. II usually absent. Chaetotaxy of type “A” or “B”. Anal spines strong and on anal papillae. 8+8 eyes present. PAO with 4 lobes, the anterior lobes larger, about twice the size of one ommatidium. Ant. IV usually with simple apical bulb and 4-8 dorsal sensilla, ventrally with or without sensory file. Between Ant. III and IV often an eversible sac present. The tenent hairs almost always acuminate. Ungues with inner tooth and up to two lateral teeth on each side. Empodial appendix with broad basal lamella and terminal filament reaching half the inner edge of ungues. Ventral tube with 4+4 setae. Retinaculum with 4 + 4 teeth. Furcula well developed, dens with 6-7 posterior setae. Mucro spoon-shaped, with a high triangular flap on outer lamella. Ceratophysella tupamara sp. nov. Figs 1-7 HOLOTYPE: MNHG, without registration number; 1 @ Peru, Department of Loreto, Iquitos, Rio Yanayacù; sample of soil between the bark of a tree and the trunk; 18-X-1980; sample PE-80/8; leg. C. Vaucher. PARATYPES: MNHG, without registration number | juvenile; same locality and date as holotype.- UNAM, without registration number; 1 d, 1 juvenile; same locality and date as holotype. DESCRIPTION: Maximum body length: up to 1.5 mm. Body granules fine and uniform, dorsal of Abd. V with 10-14 granules between pl setae. Color: dark blue. Tergite setae of different lengths, some barbulate macrosetae (50-90 um), others smooth microsetae (22-50 um), besides the sensorial setae (42-50 um) (Fig. 1). Head: Dorsal cephalic chaetotaxy (after Yosii, 1960), see Fig. 1. Differentiation between microsetae and macrosetae clear. Eyes 8+8. Eye patch with 3 setae, Oc, longer than Oc, and Oc;, the last two subequal (Fig. 1). Antennae as long as head. Ant. I with 7 setae. Ant. II with 13 setae. Ant. III organ with 2 short rods in a small integu- mentary fold and 2 guard setae of similar size; eversible antennal sac between Ant. III and Ant. IV present. Ant. IV with trilobed subapical bulb and 7 short, weakly differen- tiated sensilla (Fig. 2), one microsensillum and one subapical organ. Ventral file organ with about 10 setae. Postantennal organ composed of 4 lobes, about two times as large as the nearest eye, with accessory tubercle (Fig. 1). Labrum with 4 distinct round papillae on distal edge; labral setal formula 4/5, 5, 4. Ventral cephalic chaetotaxy (after Fjellberg, 1998) with 5 setae in px, 4 in bm, 5 in bl, and 3 in plb. Maxilla tullberg-type. Outer maxillary lobe with 2 sublobal setae. Dorsal thoracic chaetotaxy as in Fig. 1, Th. I with 3+3 dorsal setae in mr (m2 absent) and one lateral. Th. II with 3 rows of setae, m, absent, m, and py developed as sensorial setae. Th. II with 3 rows of setae: 6+6 setae in ar, 4+4 setae in mr, m, pre- sent, mg developed as sensorial setae; 6+6 setae in pr, py developed as sensilla. Unguis with | small inner tooth at 1/2 distance of its inner edge from base, and one lateral tooth 1/4 from base. Basal unguicular lamella broad, with tip of apical filament reaching 1/2 distance of inner edge of unguis. Tibiotarsi I-III each with 1 acuminate tenent hair NEW CERATOPHYSELLA FROM PERU 419 oe + , LENS Ÿ vy a3 2") . . „ad AR a6 at > — al \ a? HL N ag = Carr ef eS. 2/9 A, om “9 OS, ere A x ù D, de ER : A : ade Li Mat 0 = Fic. 1 Cetatophysella tupamara sp. nov., ° holotype. Dorsal chaetotaxy of head and thorax. (Fig. 3). Leg Ill with 3 setae on subcoxa 1, 3 setae on subcoxa 2, 7 setae on coxa, 7 setae on trochanter, 12 setae on femur, 18 setae on tibiotarsus including the acuminate tenent hair (Fig. 3). Abdominal dorsal chaetotaxy (see Fig. 7) of type “A”, after Thibaud er al. (2004). Abd. I-III with 2 rows of setae, 8+8 setae in ar, and also in pr, ps developed as sensilla. Abd. IV with 3 rows of setae, ps developed as sensilla, ratio between length of ps (s) and length of p6 = 2.0: 1. Abd. V with 2 rows of setae, 5+5 setae in ar and rp, p3 developed as sensilla. Abd. VI with 2 rows of setae, 3+3 setae in ar; pj replaced by two anal spines, p> seta barbulate and long (Fig. 7). Ventral tube short, with 4+4 setae. 420 J. G. PALACIOS-VARGAS & T. BOCANEGRA a ol ) CURE MES M A Ce FIGS 2-6 Cetatophysella tupamara sp. nov., 2 holotype. (2) Dorsal chaetotaxy of Ant. III and IV. (3) Tibiotarsus HI. (4) Tenaculum. (5) Furcula. (6) Genital plate. Tenaculum with 4 teeth on each ramus, no seta on corpus (Fig. 4). Manubrium with 16 setae; dens with 7 thin setae, one basal seta longer than the others. Mucro slightly spoon-like, with outer lamellae, its apex rounded (Fig. 5); ratio between length of dens and length of mucro = 3.2-3.6: 1. Genital plate of female with 3+3 pregenital setae, 15 circumgenital setae and 2 cugenital (Fig. 6), genital plate of male with 3+3 pregenital setae, 13 circumgenital and 4+4 eugenital. Anal lobes with 19 setae each. Two anal spines on Abd. VI short and curved, longer than their basal papillae (Fig. 7). 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SCHATTI ET AL. 450 (TOI) S+h II (TOI) #+£ (LOT) L+9 (COL) F+E LI OI CON OI (801) 6+8 (SOI) 6+8=8 (POT) 6+8=8 pue p+¢ (€01) 6+8 (TSI) S+b=P (LOT) p+€ (671) L+9 El II CI (EST) Stp=r (691) p+£ (671) L+9 (091) S+p=t (SSI) +e (LZI) 8+L Gl EI (671) “Pera pI SI 0918 6661 NHNIN 3U09 (TOI) 6+8 (001) p+e=€ (L6) r+E 61 8I 61 0918 6661 NHNIN (SII) E+7 (S6) 6+8 (06) 6 LI 8I (76) "SA LI 61 (SIT) €+7 (16) 8+Z TTL8 NHNW ‘(aids [e1qovI9A) 's’A “(ASP JO JOQUINU UDAD JO 9S89 UI ‘UOMONPII [EIQILIA) ‘PAT A :SUONBIA9IQqY 'DIEFPJOA UT SMOI UBIPOU ZUIA[OAUI ISP JO JOqUINU oY} ur SaBuBYD 'speyusA Jo sIoqunu omnfosge ur suonisod [eurpnyiduo] Yo] pue (aut Joddn) Jysıy 'ueysıueysjvy ‘Tegepuey JO AWUISIA ayy WO (961) 0918 6661 PUR (seyusA 161) T7L8 NHNIN 141/240 sdaodjnjg 30 wowed uononpaI (ISP) MOI 91898 [esIod ‘7 ATAVL PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 451 Fic. 1 Male Platyceps karelini from Hoseynabad-e Mish Mast (34°27’13”N 51°10°02”E, Esfahan Province), approximately 35 km southeast of Qom, Iran (QDE specimen, see Material and Methods). Longest specimens approximately 700 mm snout-vent length (4, NMW 25446.6, tail incomplete) and about 670 + 212 mm (?, SMF 62940); Khan & Ahmed’s (1987) 680 + 230 mm are from a specimen of unknown gender (see smallprint above). Latifi’s (1991) “107 cm; tail, 25 cm” may rely upon a taxon different from Karelin’s Racer (see Comparison). Smallest individuals 185 mm snout-vent length (CAS 84636, fide Leviton, 1959) and 200 + 65 mm (NMW 25446.3). Tail/body ratio 0.31-0.40 for males and females. Dorsal head colour pattern usually absent but often with a fine line along inter- parietal suture (Figs 1-2, 9A). A dark slanted subocular blotch and a dark oblique streak from the angle of the mouth to the anterior lateral border of the parietals always present. Exterior edge of supraocular diffusely brown or blackish above the eye in 452 B. SCHATTI ET AL. BMNH 1886.9.21.102 (Fig. 2A), MNHN 8722, 1957.60, SMNS 2381, and a QDE Platyceps karelini (Fig. 1, see last paragraph of Comparison section regarding peculia- rities of dorsal head pattern in examined specimens from SW Esfahan, Markazi, and Qom). MNHN 1957.59 (Fig. 2B, see Guibé, 1959) displays conspicuous deep black markings on the pileus made up of ‘eyebrow’ flecks, a broad mid-parietal line with bi- lateral twirls, a hexagonal protrusion at the fronto-parietal border (with a short median line running to a slightly less distinct dark area of similar size on anterior edge of frontal), a half-moon mark behind the parietals, and these scales encroached upon laterally by the temporal streak (also observed in SMNS 2381). The latter extends across the parietal meeting its counterpart mid-dorsally in BMNH 1886.9.21.102, forming an obtuse angle (Fig. 2A). Some specimens (e.g., MNHN 8722, 1957.60) show a roundish, subelliptic or short elongate nape spot. The latter is prominent in MNHN 1957.59 (see above, Fig. 2B) and connected to the post-parietal fleck by a short bar; a unilateral transverse line runs towards a dark mark behind the posterior exterior border of the parietal. A veritable nuchal streak is present in CAS 120714 and NMW 25446.5 (faint) or a QDE male (Fig. 1) and SMNS 2381, which have it fused with the first right nape blotch; in BMNH 1886.9.21.102 the right and left portion of the ante- riormost cross-band are bent cranially and converge into a striking mid-dorsal wedge- shaped extension (Fig. 2A). Body above creamish, pale grey, tan or buff with 41-60 brownish to black cross- bands. They are widest on neck, narrower than interspaces, and distinct along the whole trunk but fading away on tail. Specimens from Afghanistan and Iran as well as AMNH 96220, SMF 62924, and SMF 62940 (Pakistan) have 41 to 54 complete trans- verse blotches. More (58-60) are found in three Pakistani Platyceps karelini (AMNH 96219, PMNH 761-62). On the neck, and sometimes farther behind (e.g., NMW 25446.6), the cross-bands may extend to the venter but they normally do not reach beyond the flanks and alternate, at least posteriorly, with a ventrolateral series of dark spots or bars usually encroaching upon the lateral edges of the ventrals. Underside of neck and venter ivory, ochre, or yellowish, sometimes with a pale orange (e.g., CAS 103785), pinkish, or salmon hue (Fig. 1). The minimum for the number of complete cross-bands (41) is from SMNS 2381 with a vague origin (see Material and Methods) and the maxima for Iran and Afghanistan rely upon CAS 120714 (see below) and SMF 62940 (53). An unexamined QDE specimen has about 52 (Fig. 1, see Comparison). About 42 are present in MNHN 1957.60 (Fig. 9A, see Northern Populations: second smallprint); NHMG 4422 from Tehran [Teheran] Province shows 43 (Nilson & Andrén, 1981). Afghan specimens with pertinent data available (no counts ascertained for CAS 84634-36) have 45-54 or more complete dorsal bands (see Material and Methods); the maximum based upon CAS 120714 (Herat area) requires detailed definition. A potential Platyceps karelini x P. rhodorachis (CAS 120540) from North Afghanistan (Khulm) with approximately 78 comparatively short transverse blotches fits the number of dorsal markings (68-88) observed in two Tadzhik specimens (MHNG 2442.98, MTKD 16095) collected within less than 100 km airline from Khulm (see Northern Populations, Hybrid Racers). A “Coluber karelini” without origin reproduced in Khan (1993: Fig. 13, 1997: Fig. 3, and 2006: PI. 123) is not this species as evidenced by over 70 cross-bands and the neck pattern. Actually, this picture is manipulated; in reality, two entangled snakes are discernible and the almost completely visible specimen is a Platyceps sp. other than Karelin’s Racer, i.e., the taxon revalidated in this paper (see Systematics, Comparison: first smallprint, Conclusions). The report of a melanistic karelini PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 453 FIG. 2 Dorsal head pattern in Platyceps karelini from Afghanistan (A) and Iran (B-C): BMNH 1886.9.21.102 (A, Herat), MNHN 1957.59 (B, Golestan), and USNM 240003 (C, Esfahan, subadult, see Fig. 10A). Courtesy of Ivan Ineich/MNHN. Not to scale. from “Afghanistan” (Brück, 1968, see smallprint above) is probably Natrix tessellata (Laurenti, 1768). Maxillary with 13-15 or 16 (USNM 148631) teeth, anterior series subisodont, diastema usually wide, posterior two teeth enlarged, last offset laterad. Palatine with 8 (9), pterygoid 15-16 (Wall, 1911), and dentary (15) 16-19 teeth. Hemipenis subcylin- drical, spinose at base, apical area calyculate; spines subequal in size except for the fringe of denticules along the sulcus spermaticus; border of calyces serrated (Fig. 3). NORTHERN POPULATIONS Platyceps karelini from Turkmenistan to Kyrgyzstan and Tadzhikistan (Appendix B) show mostly similar variation of head scales (see footnote 7), the number and reduction pattern of dorsal scale rows, body proportions, dentition (in particular maxillary and dentary teeth), and hemipenis ornamentation as observed in southern populations. ZISP 19031.1 displays a loreal which is higher than long (Fig. 4C). The pre - ocular (entire in specimens from Iran to Afghanistan and Pakistan except CAS 120540: Fig. 7B, see Hybrid Racers) is divided in MTKD 13602 (right side), MTKD 16095 (both sides), and ZISP 19031.1 (right, with a faint cleft on the left); a short notch is dis- 454 B. SCHATTI ET AL. FIG. 3 Sulcate view of everted left hemipenis of Platyceps karelini MHNG 2442.99. Scale equals 5 mm. Drawing Heidi Laubscher and Andrea Stutz. cernible in ZISP 17219 (Figs 4C-D). A presubocular is absent in most Platyceps kare- lini from former Republics of the USSR; only SMF 18219, ZISP 17214.1, 19031.1, and ZMB 38816 show a bilateral scale of variable size in front of the anterior subocular (Figs 4B, 4D) ®. ZISP 14741 (right) and 19031.1 (both sides) possess an additional (third) subocular completely excluding the eye from the supralabials (Figs 4A, 4D); this condition is also reported in two syntypes (Strauch, 1873: ZISP 1696, 1698). SMF 18220, ZISP 14741 (left), 17219, and 17682 (right) have ten supralabials (fifth or sixth in contact with eye); the same number occurs on both sides of ZISP 1706 (Strauch, 1873), an unlocated specimen from the lower course of the Murgab (Boettger, 1888: “10-10”), and on the right of an unspecified individual (Strauch, 1873: “als Duplicat 4) A small granular scale between the anterior subocular (“Praeoculare inferius”), the lo- real, and the third and fourth supralabials (Strauch, 1873) is observed in the holotype of Choristodon brachycephalus Severczov, 1873 (ZISP 3581) from Khujand (“Khodzhent” or “Chodshent”, 40°17’N 69°38’E, ca. 320 m) in the Fergana Valley, Tadzhikistan. It has 200 ven- trals, 85 subcaudals (Strauch, 1873, see Northern Populations: first smallprint), and the lowest total body scale count for Platyceps karelini reported in the literature consulted by us. This racer with about (“gegen”) 80 short juxtaposed transverse dorsal bars is also noteworthy for the exceedingly developed cuneiform rostral wedged in deeply between the internasals (“Praefrontalia”) and almost completely separating them, as well as the aberration of its head (right side shorter than left, hence the scientific species name) including an elliptic eye (Strauch, 1873). If not an otherwise aberrant specimen, ZISP 3581 is a malformed P. karelini x P. rhodo - rachis possibly descending from at least one crossbreed parent. The hybrid hypothesis is supported by comparative data for six P. karelini from Tadzhikistan (Appendix B) with 208-217 ventrals, 105-113 subcaudals (n=4), and a much higher sum thereof (315-323). PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 455 A B C D À FIG. 4 Right lateral head scales of Platyceps karelini from Turkmenistan (B-D) and Uzbekistan (A, Karakalpakstan): ZISP 14741 (A), 17214.1 (B), 17219 (C), and 19031.1 (D). Variation of tem- porals as well as uncommon or rare conditions, 1.e., shape of loreal (D), occurrence of a presub- ocular (B, D) or divided preocular (D), ten supralabials (C), and fifth supralabial excluded from contact with eye (A, D). Scale equals 10 mm. Drawings Heidi Laubscher and Andrea Stutz. ausrangirt”, see next smallprint) >). Eight supralabials are found in ZISP 17386 (right, fourth entering orbit) and 19031.1 (left). The upper portion of the right posterior sub- ocular is split off in NMW 25446.1. The posterior subocular and the lower postocular are coalesced in ZISP 1697 (left) and 1698 (Strauch, 1873). Boettger (1888) noted a uni- lateral case of four scales along the posterior border of the eye (“Postocularen [...] 3-4”) in a specimen from the Murgab Valley (see Morphology: first smallprint, footnote 5). ZISP 3647 has the left upper postocular fused with the supraocular plate (Strauch, 1873). We observed eleven (instead of the usual ten) sublabials on one side of MTKD 13436, NMW 25446.2, ZISP 17582 and 19031.2 as well as in ZMB 38816 ©. 5) We located and examined four out of six Turkmen Platyceps karelini reported by Boettger (1888), 1.e., NMW 25446.1-2 (incl. possible hybrid) and SMF 18219-20. Two spe - cimens of unknown gender obtained along the Murgab (“am Murgab”) with 205-211 ventrals and 102-104 subcaudals, respectively, are possibly deposited in the “k. kaukasischen Museum” in Tbilissi (today incorporated into GNM collections) as notified in the introduction (1.c.: 875). 6) Three out of five northern specimens with eleven sublabials (NMW 25446.2, ZISP 17582, ZMB 38816, see Hybrid Racers) originate from Southeast Turkmenistan. A higher than usual number of sublabials is also found in ZMB 38833 and on one side of BMNH 1873.1.7.10 (both Platyceps karelini x P. rhodorachis) as well as a potential hybrid from North Afghanistan (CAS 120540). 456 B. SCHATTI ET AL. Ventrals 201-218 (Sd 202-214, 2 2 203-218, 201-217 in specimens of un- known gender), subcaudals 85-117 (88-117, 92-115, and 85-111, respectively), sum thereof 286-331 (293-331, 296-326, 286-323, resp., Tb. 3). Females of northern groups have slightly more ventrals than males. Northeastern populations (dd, 9 2) have higher means of ventrals and subcaudals than Platyceps karelini from any other area. Males from Southwest Kazakhstan to the Kopetdag (see footnote 8) and Bukhara (Buxoro, Uzbekistan) clearly differ from southern and eastern populations in the number of subcaudals (88-103 versus 102-117) and total body scales (293-310 vs. 308-331). Terentjev & Chernov (1940, 1949) and later authors notified 192-220 ven- trals and 85-117 subcaudals (identical minima are quoted in, e.g., Bannikov et al., 1977). Their lowest ventral count most probably relies on Wall’s (1911, 1923) data for P. karelini from Pakistan (see Morphology: second smallprint). ZISP 1705 and 3646-47 (Strauch, 1873) from “Kenderlinsk” (Kendirli Bay, SW Kazakhstan), “Karatschagly” in the Great Balkhan Range (NW Turkmenistan), and Uzbekistan (“Altes Bett des Oxus”) are classified as females due to their high ventral counts (211-213, 97-100 subcaudals, Tb. 3). The minimum for subcaudals (85) also reported by, for instance, Boulenger (1890, 1893) is from two syntypes (Strauch, 1873: ZISP 1695-96, “Original- exemplar[e]”; type locality possibly in SW Kazakhstan) and the holotype of Choristodon brachycephalus Severczov from the Fergana Valley (see synonymy, Hybrid Racers, footnote 4). The latter (not allowed for in Tb. 3) accounts for the minimum (285) of ventrals and subcaudals combined in Strauch (1873: 273). Our lowest number (286) relies upon ZISP 1696; the tail of this possibly male syntype portrayed in Strauch (1873: Pl. III) appears to be intact. The mor- phological description of Platyceps karelini by this author encompasses data of twenty-one specimens including three that had been exchanged with other museums (the “Verzeichniss der [...] aufgestellten Exemplare” lists only eighteen deposited in the Imperial Academy of Sciences). This explains discrepancies between subcaudal counts (85-101 according to index versus up to 107 as indicated in the body of the text) and the maximum for ventrals (“bei funf Stücken [...] mehr als 210”); to conclude from that latter remark, two out of three specimens then no longer in the St. Petersburg collections had more than 210 ventrals (Strauch, 1873: 113-14, 272-73). Boettger’s (1888) “95” subcaudals for SMF 18220 from Durun is based upon an incomplete tail, and the lowest value (“91”) in the Murgab series is considered incorrect (tip of tail most probably missing, see footnote 5). The maximum for ventrals (220) reported by Terentjev & Chernov (1940, 1949) may originate from one or several specimens collected in the northeastern portion of the distribution range; this count probably comprises one or two pre- ventrals and the actual number may be identical with the maximum for ventrals (218, MHNG 2442.96) observed in this study (Tb. 3). A female from Uzbekistan (ZISP 13110, see next smallprint) shows 17 midbody scale rows; the first reduction (19-17 dsr) occurs at ventral 71 (35%ven) by fusion of lateral rows. MTKD 16095 (6d) with an irregular reduction formula has 17 dsr on a portion of the anterior trunk, 19 msr, and 11 dsr prior to the vent. A fourth fusion (13-11 dsr) involving paravertebral rows at ventral 166 (81% ven) is present in MHNG 13582718), The verified number of dorsal cross-bands of most specimens from Turkmenistan (e.g., CAS 184636, MHNG 1358.27, MTKD 8281, NMW 25446.1-2, see Hybrid Racers: last paragraph) and MTKD 13602 from Southeast Kazakhstan (42 to ca. 55) is virtually identical to the range of Platyceps karelini from Iran and Afghanistan (41 to at least 54, see Morphology: third smallprint). Strauch (1873: incl. Pl. III) reported 40-48 in specimens from Southwest Kazakhstan, Turkmenistan, and ZISP 3647 from the original course ofthe Amu Darja (see smallprint above). However, PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 457 TABLE 3. Ventrals, subcaudals, and total body scales in Platyceps karelini from Iran to Central Asia and Baluchistan (Appendices A-B) including literature records. The Kandahar male sample comprises BMNH 1882.3.20.2, MNHN 8722, and MNHN 1999.8160 (see Tb. 2). Parentheses following the sample size (n) in the last column indicate number of specimens with intact tail. Boldface numbers denote literature data from Iran and Pakistan deemed in need of confirmation (see Morphology: second smallprint, Northern Populations incl. first smallprint, footnotes 2 and 4-5). Group and region Gender Ventrals Subcaudals bates ey Literature source, remark e 202-207 (4) 88-103(4) 293-310 (4) SW Kazakhstan 204.5421 965+66 301.0+7.5 and N i 201-209 (14) 85-101 (12) 286-306 (12) e Turkmenistan to "OWN 205.1427 940453 2989467 Strauch (1873), n=12 (11) Central Uzbekistan 203-214 (13) 92-106 (11) 296-320 (11) u 2 = 209.6438 973447 306.8+8,5 Strauch (1873), n=3 = ae 206-211 (5) 102-116 (5) 308-326 (5) = 208.8+22 108.2+60 317.0+7.7 > ©, SE Turkmenistan 205-213 (4) 102-110(3) 307-323 (3) 3 ine ROME 509 aa ROSIE „315080: ES = 204-216 (2) 2 ne 2100,35 vol) sul) = 207-214(5) 107-117(4) 314-331 (4) = 207- = SE Kazakhstan 99 210.2+29 112.0+42 321.8+72 yigyzstan, 208-217 (3) 105-111 (2) 315-319 (2) Tashkent area (E unknown 2117447 108.0+42 3170428 Uzbekistan), and Tadzhikistan 211-218 (5) 98-115(4) 316-326 (4) £$ 2141233 1063877 3205442 202-208 (7) 97-105 (5) 303-310 (5) dd 204.9+2.0 100.2+33 304.8 + 2.9 199-205 (5) 98-102(3) 300-305 (3) 7 en UNKOWN 590426 1000220 3023425 rauch (1875), n-2 (1) 201-209 (9) 92-106(7) 294-314 (7) 2 99 205.1429 98.6447 3034464 = 216 (1) 104 (1) 320 (1) Nilson & Andrén (1981) 3 = 204-209 (3) DI = Si e an nn © Afghanistan 196-198 (3) 92-100 (3) 288-298 (3) = 1070+10. 67245. 209745] ante È > 207-212 (5) 91-105(4) 300-313 (4) > 2088410. 1013432 3078466 si 33 198-207 (5) 90-96 (4) 290-303 (4) 9 Pakistan (NE 201.6+42 940+27 296.5 +6.0 Baluchistan) unknown 202 ( 1) Fi = Kahn & Ahmed ( 1987) 192-206 92-99 : Wall (1911), n>6 (2) 99 204 (1) 97 (1) 301 (1) ZMB 38816 (2) collected on the Turkmen-Khorasan border (Sarahs) has as much as about 65 transverse blotches, and even more (66-88) seem to be the rule in Kyrgyzstan and Tadzhikistan (MHNG 2442.96-98, MTKD 10450, 11335, 16095, NHMB 21058, ZMB 38591, see also Morphology: third smallprint, footnote 4). Two specimens from the vicinity of Bukhara (MHNG 2443.03, MTKD 13944) have at least 60 cross-bands. In addition, ZMB 38816 differs vis-a-vis female Platyceps karelini from Southwest Kazakhstan, northern Turkmenistan and southeast to Tirmiz (Termez, ZISP 13110, 204 ventrals, 458 B. SCHATTI ET AL. 106 subcaudals) on the Uzbek-Afghan frontier (Balkh Province) in, for instance, a higher num- ber of ventrals (216) as typically observed in females from Kyrgyzstan and Tadzhikistan (Tb. 3, see Hybrid Racers, footnote 7). MNHN 1957.60 (?, Fig. 9A) collected in the immediate vicin- ity (Serakhs, Khorasan-e Razavi) of ZMB 38816 across the Tedzhen (Harirud) River has 203 ventrals, 99 subcaudals, and as few as ca. 42 complete transverse dorsal markings (see Material and Methods, Morphology: third smallprint). DISTRIBUTION Platyceps karelini occurs from the northwestern Central Plateau (Iran) and the northeastern Caspian littoral (Kazakhstan) to the western Tien Shan region, the foothills of the Pamir, and Southwest Pakistan (Baluchistan). The northern limit of dis- tribution is near 47°N latitude north of the former Aral Sea in Kazakhstan (Terentjev & Chernov, 1949: map 29; Bannikov et al., 1977: map 114). Brushko (1983) reports a specimen photographed between Mojyunkum (Furmanovka) and Mount Dzhambul, roughly 100 km airline from the southern end of Lake Balqash (Zhambyl Province, Kazakhstan). In the area under consideration, the species is known from east of the Zagros Mountains and the Atrek River (Golestan) in Iran to North Afghanistan (see below) and south through Yazd, Kerman, northern Sistan-ve Baluchestan (e.g., TMUS 1000; Annandale, 1906; Werner, 1936; Latifi, 1991; see next but one smallprint and Material and Methods regarding the origin of SMNS 2381), and western Afghanistan to Baluchistan (Fig. 5). Mentions from the Arabian Peninsula (e.g., Anderson, 1896; Corkill & Cochrane, 1966; Gasperetti, 1974, 1977; Leviton, 1987; Leviton & Aldrich, 1984) rely on Bedriaga (1879) who erroneously assigned Blanford’s (1876) Zamenis ventrimaculatus [sic] from Ras Musandam (Masandim) to Karelin’s Racer (see Systematics). This specimen (ZMB 10324) from an insular promontory in northernmost Oman (“Cape Massandim, Arabian coast, entrance to Persian Gulf”) belongs to Platyceps cf. rhodorachis (Jan), the only racer species living in that area. Rai’s (1965: map 9, p. 46) indication from Kermanshah in Northwest Iran is incorrect (see Comparison: second smallprint) and “Kerdahan” (MNHN 8722, 1999.8160) is Chahar Dahaneh near Kandahar, Afghanistan. Reports from Khuzestan and East Azarbayjan (Latifi, 2000) are based upon different Platyceps spp. The pur- ported occurrence in “Markazi (Central Prov.)” notified by Firouz (2005: province no. 25) ultimately relies on Latifi’s (1991) P karelini from Kashan in Esfahan (see chresonyms). However, the westernmost verified collecting site (RUZM 11.1) is indeed from Markazi, 1.e., about 20 km roughly southeast of Delijan close to the border with Esfahan. The species is also recorded from southeastern Tehran Province (NW Dasht-e Kavir, Nilson & Andrén, 1981) and adjacent Semnan (Garmsar, Latifi, 1991). Specimens from “Mazandaran” (Latifi, 1991, 2000) including ZMB 6876 were obtained in Golestan, which formed part of the former province until 1997. The reported presence all over Sistan-ve Baluchestan to as far south as the Gulf of Oman littoral (Latifi, 1991, 2000: maps) requires confirmation and, in particular, comparison with P. ventromaculatus (see Systematics). Information by local residents towards the senior author regarding the occurrence of P. karelini in Hormozgan is probably due to confusion with Lycodon striatus bicolor (Nikolskij, 1903) which shows a similar dorsal colour pattern. A recent herpetological investigation of that area by Rajabizadeh et al. (2008) did not provide any evidence for the presence of Karelin’s Racer. PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 459 AFGHANISTAN A’Shiraz" FIG. 5 Distribution records of Platyceps karelini (solid circles) in Afghanistan, Iran, and Pakistan in- cluding five entries (symbol X) based upon two P. karelini x P. rhodorachis from Afghanistan (Herat, Nimruz) and three supposed or potential hybrid racers from Iran (Kerman) and Afghanistan (Arghandab River, Nimruz and vic. Khulm, Balkh). The arrow in Southwest Afghanistan points to Kamran (see Systematics: first smallprint). The question mark in Northeast Baluchistan Province (Pakistan) accentuates reports from near Zhob (Khan, 1997). Triangles show collecting sites of examined Iranian Coluber chesneii Martin except MNHG 1359.12 (extralimital) and published records from Bahrain (various localities), Iraq (Faw Peninsula), Kuwait, Saudi Arabia, and Sir Bani Yas Island, UAE (see Schätti, 2006: 677-78, footnote 2); open symbols denote two intergrades from Fars, Iran (see Conclusions, footnote 10). The encircled area in the Baluchistan Region delimits the assumed distribution range of P mintonorum (Latifi, 1991; Schatti & Stutz, 2005). The stippled line along the Indus Valley and the Makran coast indicates the approximate western distribution limit of P. ventromaculatus (Schatti & Schmitz, 2006: Fig. 3); two specified collecting sites (stars) in the Baluch littoral are Gwadar (25°07’N 62°20’E, BMNH 80.11.10.201) and Rumra (25°23’N 63°44’E, ZSM 222.1989, hoc loco). See text including Material and Methods, Hybrid Racers, and Appendices (A, C) for further explanations. Drawing Andrea Stutz. In Afghanistan, Platyceps karelini is documented from Balkh (relies upon potential hybrid), Helmand, Herat, Jowzjan (Ag Chah), Kandahar, and Nimruz (see Systematics, Hybrid Racers: first smallprints). The species certainly occurs in Farah and two northwestern frontier provinces (Badghis, Faryab), may be found in parts of Ghor and Sar-e Pol (NW Afghanistan), and probably extends as far east as Kunduz. Boulenger’s (1889) “Badghis” record based upon BMNH 1886.9.21.104 is from Herat (see Hybrid Racers). 460 B. SCHATTI ET AL. Reliable Pakistani records are largely confined to Kalat, Khuzdar, Mastung (in- cl. Wali Khan), Pishin (incl. Bostan), Qila Abdullah (Chaman, Gulistan), and Quetta Districts in Northeast Baluchistan. Khan’s (1997) mentions from “Zob” (“Loi Banda” and “Muslim Bazar”), i.e., Zhob (Fort Sandeman, 31°20’N 69°27’E), are in need of confirmation (Fig. 5). Mertens (1969) indicated Platyceps karelini from Northwest and Central Pakistan (“aus dem nordwestlichen bzw. mittleren Teile W-Pakistans”) but his specimens only corroborate the presence at Darzi Chah in Afghanistan (SE Kandahar, “40 mi WNW Nushki”, SMF 64629) close to the border with Pakistan as well as around Quetta and Khuzdar, the southeasternmost record of Karelin’s Racer. The species certainly lives in the Nushki area as evidenced by SMF 64629. Sturdy or trust- worthy reports, and in particular precise collecting sites, are lacking for Northwest Baluchistan Province, 1.e., Chagai and northern Kharan where Karelin’s Racer most probably occurs. Parts of Chagai District were explored by, for example, the Afghan Delimitation Commission (Aitchison, 1889: map 1) and Alcock & Finn (1897). Annandale’s (1904, 1906) de- terminations of ZSI specimens from unspecified localities in the border triangle of Afghanistan, Iran and Pakistan (Northwest Baluchistan) rely on Alcock & Finn (1897) and Arthur Henry McMahon. The inclusion of “Coluber karelini” among reptile species typical of “The Northwest Upland” (i.e., “from the high plains around Kalat and Quetta northeastward through Waziristan into the lower valleys of Swat, Dir, and Chitral”) by Minton (1966: 40, map 5) is misleading; Platyceps karelini is not recorded from beyond Northeast Baluchistan Province. Khan’s (1977) “karalini” [sic] from Darapathar near Rabwah (31°45’N 72°55’E, ca. 170 m a.s.l.), referred to as “an aberrent [sic] race” of “[t]he nominated [sic] taxon” from “arid Punjab” (Khan, 1982), is P. rhodorachis (see Comparison). Seven localities (“Boostan”, Chaman, “Punj Pai”, “Peshin”, Quetta, and in the “Zob” area, see paragraph above) listed by Khan (1997: Appendix I; see also Wall, 1911; Minton, 1966) result in two or possibly three map entries on Pakistani territory along the frontier with Afghanistan; collecting sites as, for instance, Gulistan or Mastung (Wall, 1911; Khan & Ahmed, 1987) are not plotted. Khan (1999: 276, 288) regarded karelini to be a “widely distributed [mountain] species” also enumerated among the “large number of endemics” of the “Chagai-Kharan desert” (“Herpetologically riches [sic] part of Pakistan”). Khan (2002) report- ed P. karelini (as Coluber auct.) only “from Quetta and Pishin area” but three out of four entries on his map lie west of ca. 66°30’E longitude including the vicinity of Nushki (ca. 29°33’N 66°01’E, roughly 1°000 m) and Dalbandin (28°54’N 64°25’E, ca. 850 m), and thus are likely to refer to P mintonorum (see Khan, 2002: 45). Not a single record from east or roughly north of Nushki is shown in Khan (2006: map) where merely three unspecified places are pinpointed; the southernmost and inexplicit locality mapped in Khan (2002) is missing, and the two western col- lecting sites most probably refer to P mintonorum from Dalbandin and Nok Kundi (28°50°N 62°45’E, ca. 680 m) in Chagai (Schätti & Stutz, 2005: Fig. 1). According to Khan (2002: 99), P karelini (type locality “Southwest Asia”, possibly fide Smith, 1943), called the ‘Banded desert racer’ or ‘spotted racer’, “frequents plain deserts between 1500-3000 m of elevation in north- western [sic] Balochistan”; earlier, the same species was denoted as the ‘transversely striped desert racer’ (Khan & Ahmed, 1987). It is further declared that “[s]pecimens have also been col- lected from northwestern [sic] Punjab, from Sulaiman Range” (Khan, 2006); a virtually identi- cal remark (“I collected two specimens from the Sulaiman Range [...] in a rocky area”) is hawked under “Coluber karelini mintonorum” and the putative origin left no mark on the accompanying map (see also Schätti, 2006: 683, 685; Schätti & Schmitz, 2006: smallprint p. 761). The occurrence of Karelin’s Racer above 3’000 m as indicated by Khan (2006: Tb. 10.1) is unsubstantiated. The confirmed altitudinal distribution on the Iranian Central Plateau ranges from approximately 700 m above sea level east of the Tahi Plain (TMUS 1001) in extreme eastern Yazd (Khorasan-e Jonubi border area) to about 1°870 m near Kondor in adjacent South Khorasan (Nikolskij, 1916; see Material and Methods as to USNM PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 461 240003). Similar altitudes (ca. 800 to at least 1°700 m) are attained on the northwestern Plateau. Lower elevations are inhabited around Zabol (ca. 480 m), along the Harirud (Tedzhen) River (Pasgah-e Pol-e Khatun, ca. 400 m; Serakhs, 280 m), and in Golestan where Platyceps karelini lives near sea level, for instance at Aq Qal’eh (Latifi, 1991). Afghan records are from ca. 280 m in Jowzjan (Ag Chah) to at least 1’000 m in the west and south (e.g., “Chinkilok” [Herat] or the vicinity of Kandahar including Chahar Dahaneh). To conclude from collecting sites in the immediate border area (Qila Abdullah District, Pakistan), the species most probably occurs at elevations of at least 1’200 m in Kandahar Province and elsewhere in Afghanistan. Three Pakistani spe- cimens (AMNH 96219-20, SAM 931) were obtained close to 2’000 m “in a cultivated section of Urak Valley” (vic. Hanna, Quetta) and at ca. 17530 m “in similar terrain near Pishin” (Minton, 1966). Altitudes nearing 2°000 m are inhabited around Kalat (PMNH 761). P. karelini from Quetta, Mastung, and Khuzdar (see Appendix A) were collected between ca. 1°200-1°700 m. DISCUSSION SYSTEMATICS It is hardly an exaggeration to assert that Platyceps spp. from the western Sahara and eastern Mediterranean Region to the Himalayas (NW India) represented a taxonomic and systematic disarray for quite a long time. In particular, species from the eastern Caspian area to Pakistan still constitute a difficult problem to solve, confusing generations of zoologists since the days of Gunther (1858). Blanford (1876) synonymized Karelin’s Racer, Zamenis rhodorachis Jan, 1863 and other taxa as, for instance, Coluber chesneii Martin, 1838 (type locality “Euphrates”) with Coluber ventromaculatus Gray, 1834. With this relegation, Chesney’s Racer vanished into the taxonomic muddle of the Platyceps rhodorachis- ventromaculatus complex for 130 years or so to come. Contemporary colleagues (e.g., Boettger, 1880; Murray, 1884) followed Blanford’s (1876) view and thought Zamenis ventrimaculatus [sic] to be distributed from Egypt and the Arabian Peninsula to east of the Caspian Sea and Bengal. Boettger (1888) considered Coluber (Tyria) karelini Brandt, 1838 a colour variety (“Farbenvarietät”) and ranked Karelin’s Racer as a sub- species (“var.”) of Z. ventrimaculatus [sic] auct. Boulenger (1890) re-established spe- cific status for karelinii [sic], rhodorachis (as Z. ladacensis Anderson, 1871), and ventrimaculatus [sic]. Probably the best systematic herpetologist ever, in his monu- mental ‘Catalogue’, even Boulenger (1893) simply seemed to be overwhelmed when it came to the predicament of rhodorachis or ventromaculatus auct., and karelini as well (see Hybrid Racers incl. first smallprint). Blanford’s (1876) sample of Zamenis ventrimaculatus [sic] (see Distribution, Hybrid Racers: first smallprints) may not include any genuine Platyceps karelini. His series of twelve racers assigned to this confusing taxon comprises two specimens (no. “2, 3”) obtained at Kamran (“Zamrän”, 30°53’N 61°47’E, ca. 560 m above sea level) in Nimruz, Southwest Afghanistan, along the border with Iran (Fig. 5: arrow). Just one of them, ZSI 4616 or 8603 (Sclater, 1891: as Z. ladaccensis [sic], i.e., P rhodorachis), has “the sixth supralabial [...] divided, so that there are three postoculars, and only the fifth supralabial enters the orbit.” Its identity is unknown but the specimen undoubtedly may belong to P. karelini (Schatti & Schmitz, 2006: 761); however, the origin and the presence of a posterior subocular do clearly not exclude P mintonorum (see 462 B. SCHATTI ET AL. Comparison). The second Kamran “ventrimaculatus”, apparently with two supralabials in contact with the eye, belongs to a Platyceps sp. other than P. karelini (see Hybrid Racers as to BMNH 1873.1.7.10 and 1886.9.21.101 from Nimruz). Many field (and other) herpetologists remained perplexed regarding the diver- sity of morphological features of Platyceps spp. belonging to the karelini-rhodorachis- ventromaculatus complex within geographically limited areas. To cite a more recent comment (“a complicated genus in S.W. and Central Asia”), we remind the reader that Clark’s (1990) sample of eight “Coluber karelini Brandt” including the living one seen in “the Kabul bazaar” consists of three different species (P. karelini, P. mintonorum, P. rhodorachis) and a potential hybrid racer (see next chapter). Earlier, Leviton (1959) had “pointed out that [...] C. karelini, C. rhodorachis, and C. ventromaculatus, are known from southwestern Asia where their distributions overlap most extensively. They exhibit the same ranges of morphological variation, the same color pattern variations, and are found in similar environmental situations.” Until recently, C. ven- tromaculatus auct., for instance, was by and large considered to be distributed from the Near East to northern India (e.g., Minton, 1966; Leviton er al., 1992; Khan, 1997; Disi et al., 2001: 267). In reality, P. ventromaculatus is only known from the lower parts of Northwest India and Pakistan where it extends along the Makran coast (Fig. 5) to at least as far west as Gwadar (Schatti & Schmitz, 2006: Fig. 3, footnote 2). A specific search would probably provide evidence for the presence of Gray’s Racer in the littoral of Southeast Iran (Sistan-ve Baluchestan). Platyceps cf. ventromaculatus sensu Schatti (2006), with Coluber chesneii Martin as its oldest available name, is a distinct taxon found from the eastern Mediterranean to at least as far east as Fars in Iran (Fig. 5, see Comparison). Chesney’s Racer differs from P. ventromaculatus sensu stricto in, for instance, hemipenis features or molecular data (Schätti et al., 2005: Abb. 8; Schätti, 2006) and is herewith revali- dated. The systematic allocation of C. chesneii M. is addressed in the last chapter of this study (Conclusions). Nagy et al. (2004) sequenced five nucleotides of seven nominal Platyceps spp. including “P rogersi” (Anderson) from the Sinai, P. karelini (SW Turkmenistan), and P. rhodorachis (vic. Ashgabat). They found that the latter is “[b]asally linked to this six-taxon clade” and asserted that their “findings are of a strongly supported sister taxon relationship between P. karelini and the Arabian P. rogersi, thus excluding P rhodorachis.” However, there does not seem to be a reproductive barrier separating karelini and rhodorachis sensu stricto as substantiated by hybrids. Schatti (2006: 675 [abstr.], 684) regarded Zamenis rogersi Anderson, 1893 to be conspecific with Platyceps cf. ventromaculatus. The senior author ranked Rogers’s Racer as a junior synonym of this nomen operandum for unassigned Platyceps populations found in “com- paratively mesic areas from northeast Africa to the Middle East” (Schatti & Schmitz, 2006) and suggested Coluber chesneii Martin as the correct scientific name (“may be a valid northern Saharo-Arabian taxon”, see Conclusions). Based on nucleotide sequences (COI, 12S rDNA), P. cf. ventromaculatus is more closely related to P. cf. rhodorachis from Yemen than to P. ventro- maculatus sensu stricto (Schätti et al., 2005: Abb. 8); incomplete data for P. karelini (only 12S partition) produced a sister taxon relationship with Chesney’s Racer (cf. ventromaculatus) and cf. rhodorachis. In an earlier analysis without karelini, P. rogersi auct. appeared to be closely related to Yemeni cf. rhodorachis (Schätti & Utiger, 2001: Figs 8-9). Amr & Disi (2011) credited the latter authors with the inaccurate quote “that P rogersi is well differentiated from P. ventro- maculatus, while similar to the North African P. karelini [sic]. Their findings strongly supported PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 463 sister taxon relationship between P. karelini and the Arabian P. rogersi, thus excluding P. rhodo - rachis from the clade.” The second assertion is incorrectly ascribed and has to be imputed to Nagy et al. (2004, see above). HYBRID RACERS Bogdanov (1953) was the first to postulate hybridization between Platyceps karelini and P. rhodorachis. This assessment 1s based upon specimens from the Murgab Valley (SE Turkmenistan) combining the diagnostic features of P. karelini (presence of a posterior subocular, 1.e., a single supralabial in contact with eye) and the typical phenotype of P rhodorachis showing a reddish vertebral stripe from the nape all along the trunk. Both character states are observed in ZMB 38833 from the Garagum (Karakum) Desert (Fig. 6A, Tb. 4). This female has two superposed loreals (upper smaller, actually the split off lower lateral edge of the prefrontals) and, on the left side, a divided preocular, a presubocular, a narrow cuneiform scale separating the upper portions of the second and third supralabials, and eleven sublabials; the posterior chin shields are separated by four (cranial, instead of the usual two) to five (caudal) rows of elongate scales. Besides a faint reddish median line on a uniform light brownish dorsum, there are a few tiny scattered dots along the back. Furthermore, we absolutely consider BMNH 1886.9.21.104 (6, Fig. 7A, Tb. 4) from Herat Province (Gulran, see Distribution) with a posterior subocular and a bright orange vertebral stripe to be a hybrid of P karelini and P. rhodorachis (see next but one smallprint) ?. It has a small supplementary dorsolateral head scale, 1.e., the detached posterior outer edge of the internasal. Boulenger (1889), apparently in a quandary, first assigned this “very fine specimen [...] splendidly marked with a bright red broad line down its back” to rhodorachis but referred to it later as karelinii [sic] (Boulenger, 1893: letter h, “2”’). All subsequent reports regarding the presence of a reddish median stripe in karelini from the area under consideration (e.g., Smith, 1943; Mertens, 1969: 59; Clark, 1990), 1.e., except regions within the former Sovjet Union, rely upon this male from northern West Afghanistan. Viable crossbreeds between Platyceps karelini and P. rhodorachis do not appear to be uncommon in certain areas of sympatry, in particular Southeast Turkmenistan (e.g., Bogdanov, 1953, 1962; Czellarius, 1974, 1992; see footnote 8) and adjacent Afghanistan (Figs 6A, 7A). Although the topic needs further investigations (in prep.) including fieldwork and intricate molecular techniques, recombinations involving the hybrid genome may result in a disordered genetic equilibrium reflected in, for example, aberrant morphology. As one of the parent species (P. rhodorachis) features two distinct phenotypes (uniform with longitudinal vertebral stripe versus transversely blotched or spotted dorsal pattern), it seems reasonable to assume that different colour morphs also occur in P. karelini x P. rhodorachis. This is why we speculate that the 7) On the right side, BMNH 1886.9.21.104 shows a split off triangular portion (lower anterior edge) of the seventh supralabial and the comparatively narrow upper anterior temporal extends as far caudal as the lower (Fig. 7A) whereas this scale is distinctly shorter (and much smaller than the lower first temporal) on the left as is often the case in Platyceps karelini (Figs 4, 10A). A long slender upper anterior temporal is also found in TMUS 1001 (W Iran); it is longer than usual in ZMB 38591 (left side, Tadzhikistan) and a Turkmen racer (ZMB 38816, see Northern Populations and last paragraph of this chapter). 464 B. SCHATTI ET AL. FIG. 6 Dorsal view of Platyceps karelini x P. rhodorachis from Turkmenistan (A, ZMB 38833, Garagum Desert) and a supposed Afghan hybrid racer (B, BMNH 1886.9.21.101, Nimruz, see text, Appendix A). Not to scale. holotype of Choristodon brachycephalus Severczov (ZISP 3581) showing a blatant deformation of the head is, if not a teratism, an abortive hybrid (see footnote 4). BMNH 1873.1.7.10 (Fig. 8, juv. 3) has a minuscule left presubocular and a colour pattern reminiscent of Platyceps karelini (Figs 1, 9A, see Comparison: first smallprint). It is remarkable for various head and body scale conditions, e.g., the shape of the loreal (higher than long), orbit completely separated from supralabials by three suboculars (fifth supralabial horizontally divided, lower portion distinctly smaller), number and arrangement of sublabials (ten and only three in contact with anterior chin PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 465 FIG. 7 Rostral shape and right lateral head scales of Platyceps karelini x P. rhodorachis (A, BMNH 1886.9.21.104, Herat) and a potential hybrid between these species (B, CAS 120540, Balkh) from Afghanistan. Courtesy of Philippe Wagneur (A). Not to scale. FIG. 8 Dorsal and left lateral head views of BMNH 1873.1.7.10, a Platyceps karelini x P. rhodorachis from Afghanistan (Nimruz: Qala-i Fateh). 466 B. SCHATTI ET AL. shield on one side, eleven and four, respectively, on other) as well as a much elevated number of ventrals and total body scales compared to males from the whole range and both genders of southern P. karelini, respectively (Tbs 3-4). These character states and meristic data prompt us to classify this racer obtained near the Afghan-Iran border as a hybrid (P. karelini x P. rhodorachis). BMNH 1873.1.7.10 (coll. Major Euan Smith) from “Kila-i-Fath, Sistàn” (Qala-i Fateh, NW Nimruz) has “all the labials below the eye divided”, 216 ventrals, and 107 subcaudals (Tb. 4), pretty close to Blanford’s (1876) counts (“ventrals are 218 in number, subcaudals 108”, see footnote 2). Boulenger’s (1893) data for this specimen (207 and 99, respectively) are incorrect. Two Zamenis ventrimaculatus [sıc] sensu Blanford (1876) are discussed elsewhere (Systematics: first smallprint); BMNH 1873.1.7.10 and probably one of the three Kerman racers (see below; this series includes ZSI 4615, see Sclater, 1891: as Z. ladaccensis [sic], i.e., Platyceps rhodo- rachis) also enumerated by Boulenger (1893) are P. karelini x P. rhodorachis. Four out of seven Z. karelini sensu Boulenger (1893) from Kerman and Afghanistan including BMNH 1886.9.21.104 (see above) and probably 1886.9.21.101 (below) may be hybrids, and there is actually no verified record of genuine P. karelini from Southwest Afghanistan (Fig. 5: arrow). TABLE 4. Body scales (number of ventrals, subcaudals, and sum; extreme tip of tail possibly missing ın ZMB 38833), dorsal scale row (dsr) reductions (longitudinal position in %ven, trans- verse level), and dorsal markings (patterned [transversely blotched or spotted] or striped pheno- type) of Platyceps karelini x P. rhodorachis and three supposed or potential hybrid racers (preceded by an asterisk; see text, Figs 5-8 and Tb. 1 for definitions). Ventrals, subcau- Reduction pattern (dsr) Phenotype Gender and country Specimen an : dals and sum in%ven transv. levels (province) BMNH 1873.1:7.10 216 107 323 59, 60, 75 par-lat-par patterned d, Afghanistan (Nimruz) *BMNH 1874.11.25.10 203 102 305 59, 61, 70 (see text) - par patterned d, Iran (Kerman) *BMNH 1886.9.21.101 203 105 308 61, 62,79 par-lat- par patterned 6, Afghanistan (Nimruz) BMNH 1886.9.21.104, 206 110 316 57,63, 77 par-lat-p+v — striped 3, Afghanistan (Herat) *CAS 120540 207.105 312 56, 63,71 par-lat-par patterned d, Afghanistan (Balkh) ZMB 38833 212 110? 322? 59, 65,71 par-lat-par striped ?, Turkmenistan (Mary) BMNH 1874.11.25.10 from Kerman (see smallprint above, Material and Methods, Schatti & Schmitz, 2006: 761) differs vis-a-vis Iranian Platyceps karelini in its number of complete transverse dorsal blotches, i.e., more than 55 compared to 41-50 except in an unexamined specimen from Qom (Fig. 1, see Comparison). The cross-bands become narrow towards midbody and it seems that the posteriormost portion of the trunk is devoid of any dorsal pattern (or mid-dorsal transverse markings faded away) which is atypical of P. karelini but characteristic of P. rhodorachis. We suppose this racer with a noteworthy dorsal scale row sequence (reduces to 17 dsr on the right side involving low and high levels and then to 15 dsr on the left) and BMNH 1886.9.21.101 from the Arghandab (Helmand, see Material and Methods) in Nimruz to PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 467 be hybrids. The latter, from an area where P. karelini may be syntopic with P minto- norum (Fig. 5, see Comparison), has over 70 consistently narrow transverse bars all along the dorsum (Fig. 6B) compared with 50 or less in specimens from Iran to pro- bably as far west as at least 53°E longitude (see above). Furthermore, CAS 120540 from North Afghanistan with about 78 short cross-bands is considered a P. karelini x P. rhodorachis hybrid. It is outstanding compared to southern (and most northern) P karelini studied by us for its condition of the preocular (divided, Fig. 7B) and a single right anterior temporal (unique among examined material) as well as further unilateral head scale features, 1.e., an elevated number of supralabials (ten) and eleven sublabials. Certain reservations regarding the hybrid nature of this racer arise from the lack of data for genuine P. karelini from North Afghanistan (Fig. 5), a similar number of dorsal markings in contiguous northern populations, and the occurrence of two preoculars in MTKD 16095 from adjacent Tadzhikistan (see Morphology: third smallprint, Northern Populations). The rostral of CAS 120540 is noticeably protruding and it is slightly so in BMNH 1886.9.21.104 (Platyceps karelini x P. rhodorachis, Figs 7A-B). The conspicuously projecting snout of the latter is due to a crushed mandible. Although the muzzle clearly projects beyond the lower jaw in some P. karelini (e.g., BMNH 1886.9.21.102 and USNM 148631, both with a remarkably pointed snout in dorsal, or ventral, view), this condition is subject to variation and does not allow a distinction from, for instance, P rhodorachis (contour of snout in dorsal view inconstant to some degree as well). Moreover, the rostral of BMNH 1886.9.21.104 and, for example, CAS 120540 is deeply wedged in between the internasals (see footnote 4). At least in P. karelini, the shape of the posterior dorsal edge of the rostral is an unstable character, resulting in variable proportions of the anterior snout scales. This is exemplified by SMF 62924 and 62940 (both specimens show a surprisingly blunt anterior edge of the rostral in dorsal view) with distinctly longer internasals (measured along median suture) versus BMNH 1886.9.21.102 (Fig. 2A) or MNHN 1999.8160 which have the rostral wedged in between the internasals and comparatively much longer prefrontals. Except for BMNH 1874.11.25.10 and 1886.9.21.101, specimens discussed in this chapter display peculiarities of head scales, maybe to the point of anomalies (ZISP 3581), and possibly other characters. It cannot be ruled out that our samples of Platyceps karelini (Appendices A-B) comprise a few additional hybrid racers. Possible candidates are ZMB 38816 from the Turkmen-Iran border (Tedzhen) with a much elevated number of cross-bands (ca. 65, see Northern Populations incl. second small- print) and NMW 25446.2 (d, 210 ventrals, 104 subcaudals) from the Murgab Valley with coalesced frontal and parietal shields, eleven left sublabials, and about 55 compa - ratively narrow transverse dorsal bars (posteriormost short) with straight anterior and posterior borders instead of the typical slightly rounded contour as shown by a male P. karelini (NMW 25446.1) from the same area (see footnotes 6-7). COMPARISON With the exception of Platyceps mintonorum (Mertens), Karelin’s Racer differs from most congeneric taxa, and in particular all sympatric species, in the presence of a posterior subocular scale, i.e., a single (usually fifth) supralabial in contact with the eye (rarely none) versus two in Southeast Mediterranean and Saharo-Sindian Platyceps spp. Besides the condition of the suboculars (e.g., Strauch, 1873; Boulenger, 1893; Werner, 1929; Leviton & Anderson, 1970; Khan, 1997), P. karelini (Brandt) is charac- 468 B. SCHATTI ET AL. terized by a conspicuous oblique streak below the eye and another on the temple as well as dark dorsal cross-bands all along the trunk persisting in adults (Minton, 1966; Mertens, 1969; Khan, 1997). These markings separate P. karelini from any taxon dealt with in the present study except Chesney’s Racer. P najadum (Eichwald, 1831), sym- patric with Karelin’s Racer from western Iran to Golestan, northern Khorasan-e Razavi, and the Kopetdag piedmont 9, differs from all discussed species in the number of apical pits (single versus paired) and further morphological characters including the dorsal colour pattern (Schatti et al., 2005). The occurrence of striking dark transverse dorsal blotches in Platyceps spp. other than Karelin’s and Chesney’s Racers is observed in juveniles of, for example, P ventromaculatus (Khan, 2006: Pl. 136B). Regarding the dorsal colour patterns of the taxa discussed in this paper, the reader is referred to the following selection of photographs and other illustrations. Coluber chesneii Martin: Leviton et al., 1992: Pl. 15F; Khan, 1993: Fig. 13, 1997: Fig. 3, and 2006: PI. 123 (see Morphology: third smallprint); Schatti, 2006: Pl. 1; Yıldız, 2011: Figs 2 and 3a; or the excellent engraving of the destroyed MSNM syntype of Zamenis persicus Jan in Jan & Sordelli, 1867: Pl. II.1 (see Conclusions: smallprint, footnote 10). P. karelini: Nilson & Andrén, 1981: Abb. 9; Khan, 2002: Figs 41-43 (see chresonyms); or Strauch, 1873: Pl. II (incl. dorsal view of a syntype) and Latifi, 1991, 2000: Fig. 34 (coloured drawing). P. mintonorum: Minton, 1966: PI. 24.2; Mertens, 1969: Abb. 17 (holotype); Khan, 2002: Fig. 64 (“Coluber rhodorachis”, also published mirror-inverted in Khan, 1993: Fig. 22); Schatti & Stutz, 2005: Pl. 1. P rhodorachis: Minton, 1966: Pl. 25.1 (reproduced in Khan, 2006: Pl. 134); Leviton et al., 1992: Pl. 15E; Khan, 2002: Fig. 66; or Bannikov et al., 1977: Pl. 28.4-4a and Latifi, 1991, 2000: Figs 41-42 (coloured drawings). P. ventromaculatus: Minton, 1966: Pl. 24.1; Leviton er al., 1992: Pl. 15H; Khan, 2002: Fig. 67; Khan, 2006: Pls 136A-B; Schatti & Schmitz, 2006: Fig. 1. Platyceps mintonorum is an endemic racer of Baluchistan (Fig. 5) documented from Afghanistan and Pakistan; according to Latifi (1991, 2000) it also occurs in “Sıstan and Baluchistan Province (Zabol, Zahedan)”. There, as well as in Nimruz, at Chah-i Anjir (Helmand) or near Kandahar, this species and P. karelini may be syntopic (see Systematics: first smallprint, Hybrid Racers). Minton (1966) emphasized that the ‘Variegated Sand Racer’ (i.e., P mintonorum) “is distinct from ventromaculatus, rhodorachis, and karelini in the regular presence of a third preocular [see Morphology: first smallprint], high ventral count, and body pattern’, and that karelini and mintono- rum “are quite different in pattern and in ventral and subcaudal counts.” In fact, Mintons’s Racer differs in various morphological characters including the number of preocular scales (entire in southern and most other karelini, often divided in minto - norum), more ventrals (221-240), subcaudals (110-127), and total number thereof (336-360, Schätti & Stutz, 2005) compared with 196-212, 90-106, and 288-314, respectively, in P. karelini from Iran to Pakistan (Tb. 3, see Morphology: second small- print). Platyceps rhodorachis (Jan) is distributed over most of Iran, Afghanistan and Pakistan except in truly arid country and sandy areas or at high altitudes. The typical phenotype of P rhodorachis can easily be distinguished from P. karelini by its reddish vertebral stripe running all along the dorsum. The condition of the supralabial scales 8) “This lowland snake species [Platyceps karelini] is very rare” along the southern foothills of the Kopetdag in adjacent Turkmenistan (Atajev et al., 1994). In Khorasan-e Razavi, P. n. najadum is recorded from, for example, Darreh Gaz (Dargaz, Latifi, 1991) and the Kuh-e Qamar Ali area (37°27’48”N 58°38°04”E, above 2000 m a.s.1., TMUS specimen). PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 469 beneath the eye (see above) allows instant distinction from the striped and patterned (transversely blotched or spotted) morphs of rhodorachis. Furthermore, the dorsal colour pattern of the latter phenotype fades away behind midbody. Jan’s Cliff Racer usually lacks a posterior subocular (two supralabials enter the orbit). The occurrence of this scale in rhodorachis is definitely a rare condition; a posterior subocular seems to be present in a specimen from Central Punjab assigned to karelini (Khan, 1977) due to a single supralabial in contact with the eye. However, this racer belongs to rhodo - rachis as evidenced by the number of subcaudals (130) and its origin (see Distribution: second smallprint). “The nearest relative to karelini appears to be ventromaculatus; the two differ only in the regular presence of a third postocular in karelini [see Morphology: first smallprint], in head markings, and in the stronger blotched pattern of karelini” (Minton, 1966: 123). Although this observation including the remark regarding the head markings, and in particular the occurrence of a nuchal streak in Platyceps ven- tromaculatus, is correct, there is no substantial evidence of a sister species relationship and the posterior subocular (present or absent) is a variable character in Gray’s Racer (Schatti & Schmitz, 2006). P ventromaculatus, a lowland species from Northwest India and Pakistan, does not extend west of the Indus Valley except along the Baluch coast and is geographically separated from P. karelini (Fig. 5, see Systematics). Platyceps karelini and Coluber chesneii Martin meet on the Central Plateau (Iran), the closest known records being from about 80 km southeast of Esfahan and the eastern foreland of the central Zagros Range (Izad Khvast, Fig. 5, see Conclusions). Chesney’s Racer is confirmed from the Arvand Rud (Shatt Al-Arab, Khuzestan) area and along the frontier with Iraq to at least as far north as Kermanshah (e.g., MHNG 1359.12). It probably occurs in Kurdestan Province and may encroach upon the eastern escarpment of the Zagros (Semirom District in extreme SW Esfahan, Chahar Mahall-ve Bakhtiyari) or extend eastward down the southern Zagros and across the foothills inland from the Gulf littoral. C. chesneii M. covers a considerable altitudinal range from near sea level (Bushehr and Khuzestan) to roughly 2’200 m in northern- most Fars (FMNH 20939). FTHR 15300 was collected in the same general habitat as P. najadum (ssp.), 1.e., rocky terrain with scattered oak forest between 600-1°100 m in North Khuzestan (Torki, 2010: 30-31, Fig. 10, as P. karelini). A Zamenis ventrimaculatus [sic] “Var. A. v. Günther” from “Borazjün” (Borazjan, 29°16’N 51°12’E) in Bushehr Province (Werner, 1917: 211) obtained by Friedrich Carl Andreas could not be located in institutional collections rummaged through by us. Field notes of the col- lector contain three “Z. v. var. D. persica” from “Tangistàn” near Ahram (Bushehr) and a cottage at “Tschäbägh”, possibly in Fars; these specimens were not accessible to Werner (1917) and probably belong to Platyceps rhodorachis (Schatti et al., 2010: 278). Rai’s (1965: map 9) unspecified record of “Coluber karelinii” [sic] from the Kermanshah-Iraq border area is based upon Chesney’s Racer as evidenced by the low number of supralabials (“Labiales supérieures 8 (quelquefois 9)”), as sometimes observed in this taxon (Schätti, 2006), and their condition rela- tive to the eye, i.e., two supralabials in contact with orbit (l.c.: 44-45). Except for FMNH 20939 (Fig. 10B) and NMW 25446.7 (with a right presub- ocular, Fig. 10C), Iranian Coluber chesneii M. examined by us have nine supralabials and usually the fifth and sixth in contact with the eye. In BMNH 1869.8.28.130, the right fifth supralabial is horizontally divided (lower part excluded from orbit) and a 470 B. SCHATTI ET AL. presubocular is present on the left side 9). NHMB 15210 has a narrow upper portion of the anterior subocular split off (the posterior part of this scale is above the lower border of the eye; this condition could be described equally well as a ‘divided preocular’) and a tiny subtriangular presubocular, 1.e., the detached lower posterior edge of the loreal. There are 204-217 ventrals (4 6204-217, 2 2 206-208, 209 in FMNH 20939), 101- 116 subcaudals (102-116, 101, and 110, respectively), and a sum of 307-333 (310-333, 307, 319, resp.). Four males (BMNH 1869.8.28.130, FTHR 15300, MHNG 1359.12, MNHN 7470) have considerably fewer ventrals (204-209) and total body scales (310- 318) than NMW 25466.1 (Fig. 9C) and SMF 61869-70 (8 8: 214-217, 323-333, resp.) from unspecified localities in Iran. The number of maxillary teeth ranges from 13-16 (n=7); the maximum is from the extant syntype of Zamenis persicus Jan (see next smallprint). Apart from the number of supralabials entering the eye and body scales (see above, following paragraph, next chapter, and Tb. 3), Chesney’s and Karelin’s Racers differ in their dorsal colour patterns. Authentic Coluber chesneii M. display charac- teristic markings on the pileus which persist in adults, possess a nuchal streak (Jan & Sordelli, 1867: Pl. IL.1), and have more than 60 transverse bars along the dorsum (see Comparison). Furthermore, this taxon seems to attain larger body size as exemplified by NMW 25446.7 (9, ca. 100 cm total length, Fig. 9B) and, possibly, Latifi’s (1991, 2000) maximum dimension for C. karelini auct. (see Morphology). Besides this, Chesney’s and Karelin’s Racers are very similar regarding their general habitus, scale features, and further morphological characters. They show, for instance, the same range of maxillary teeth (13-16) and virtually identical hemipenes, in particular with respect to the fringe of tiny spines along the sulcus spermaticus (Fig. 3; Schatti, 2006: Fig. 1). Three Platyceps karelini from Markazi (RUZM 11.1, 2, tail broken), Esfahan (USNM 240003, 2, 103 subcaudals), and Qom (unregistered, tail truncated) have 204 ventrals each, which is the minimum ascertained in Chesney’s Racer from Iran (8 à). USNM 240003 possesses the same number of total body scales (307) as NMW 25446.7 (2, see next chapter). Most P. karelini are devoid of dorsal head markings and usually lack a nuchal streak. It is, however, present in a QDE male (Fig. 1) and a mid- dorsal spot behind the parietals occurs in all three examined specimens from Southwest Esfahan, Markazi, and Qom. These four P. karelini show approximately 45-52 cross- bands along the trunk (see Morphology: third smallprint). The posterior edges of the 9) BMNH 1869.8.28.130 from “Bushire” (Bushehr, “Dr. Leith”) unquestionably belongs to Chesney’s Racer. However, this collector obtained his material (e.g., BMNH 1869.8.28.116, 1869.8.28.132 and 134) from “Kurrachee” and “Sind” (Boulenger, 1893: 400, letters n and p-s; Schätti & Schmitz, 2006: 768), and there seems to exist confusion regarding the origin, pro- venance, and published data of BMNH 1869.8.28.130 (Coluber chesneii Martin) and 1879.8.15.27. The former (Boulenger, 1893: letter f) has a truncated tail with 90 subcaudals instead of “92”. BMNH 1879.8.15.27 (Blanford, 1881; Boulenger, 1893: letter g) is a female with 205 ventrals, 110 subcaudals (Schätti, 2006: footnote 1), and 19-19-14 dsr registered as from “Bushire, Persia”. This specimen, purchased from dealers of natural history items (Watkins & Doncaster), lacks dark markings on the pileus, and the transverse dorsal bands (becoming indistinct and confined to the vertebral area towards the tail) show irregularly serrated anterior and posterior borders. This racer belongs to Platyceps ventromaculatus (Gray). PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 471 FIG. 9 Dorsal view of Iranian Platyceps karelini (A, MNHN 1957.60, Serakhs) and Coluber chesneii Martin: NMW 25446.7 (B, “Schiraz” intergrade, see Fig. 10C) and NMW 25466.1 (C, “Persien”). Courtesy of Ivan Ineich/MNHN and Heinz Grillitsch/NMW. Not to scale. frontal are darkened except in RUZM 11.1 (with two indistinct longitudinal dark stripes instead); the latter and USNM 240003 (Fig. 2C) disclose a blurry blotch across the supraocular; RUZM 11.1 exhibits cloudy pigmentation on posterior portion of the parietals. 472 B. SCHATTI ET AL. CONCLUSIONS Morphology (pholidosis, dentition, hemipenis) substantiates that Platyceps cf. ventromaculatus sensu Schatti (2006) from Northeast Africa to Iran, i.e., Coluber ches- neii Martin (see Systematics incl. second smallprint), is conspecific with P. karelini (subspecies). The seemingly parapatric distribution of P k. karelini (Brandt, 1838) and P. karelini chesneii (Martin, 1838) comb. n. as well as the existence of overt intergrades corroborate this new systematic concept. The specific name karelini has priority over chesneii because Brandt’s (1838) descrip- tion was published beginning of February (“Emis le 2 Février 1838”) as notified on the last page (col. 256) of number 16 (“tome troisième”) of the ‘Bulletin scientifique [...]’ whereas Martin’s (1838) report on the “Euphrates expedition’ was delivered in July of that same year (Sclater, 1893). The type series of Coluber chesneii Martin from the “Euphrates” presented to the Geological Society of London (Chesney, 1850) consisted of several individuals but a single adult female syntype (BMNH 1946.1.12.95) is known to be deposited in an institutional collection (Gunther, 1858, 1864; Boulenger, 1893; Schatti, 2006). Apart from the original description, references to C. chesneii M. are found in Günther’s (1858, 1864) ‘Catalogue’ under “Var. A” of Zamenis ventrimaculatus [sic] and ‘The reptiles of British India’ (“A specimen [...] from Mesopotamia”, 1.e., the extant syntype, “agrees completely with the types” of C. ventromacu- latus Gray), as a valid species (Z. chesnei [sic], Giinther, 1868; see also Giinther, 1874) or in the synonymy of Z. ventrimaculatus [sic] auct. (Stoliczka, 1872; Blanford, 1876; Boulenger 1890, 1893; Wall, 1914; again as chesnei [sic]) as well as in Sherborn (1925), Schatti (2005: 170, 172, 174; 2006), Schatti & Schmitz (2006, see Systematics: second smallprint), and in Schatti et al. (2010) who declared the junior synonym Zamenis persicus Jan, 1863 a nomen oblitum. FMNH 20939 and NMW 25446.7 from the Fars-Esfahan border and “Schiraz” 10) are the only specimens out of about 175 “Platyceps cf. ventromaculatus” examined by the senior author (Schatti, 2005, 2006) with a posterior subocular, 1.e., a single (sixth or fifth, respectively) supralabial entering the orbit (Figs 10B-C, see Comparison regarding the condition in BMNH 1869.8.28.130). They were collected in the area of contact with P k. karelini (Fig. 5) and have more than 60 transverse dorsal bars as typically found in P. k. chesneii but the markings on the pileus characteristic of this taxon are absent; the subadult (FMNH 20939) also lacks a nuchal streak whereas NMW 25446.7 (Fig. 9B) shows a stripe along the nape and a black interparietal line (as well as dark posterior edges of the frontal) as often found in k. karelini. Manifestly, these two racers are intergrades with the nominotypical subspecies. They also deviate from the normal number of supralabials (nine) observed in Chesney’s and Karelin’s Racers (eight on right side of unregistered Qom specimen, see remark in Appendix A), 1.e., eight (NMW 25446.7, right) and ten (FMNH 20939, Figs 10B-C). The eastern portion of the species’ range is inhabited by Platyceps k. karelini (see Distribution). P k. chesneii is recorded from Southeast Turkey (e.g., Yildiz, 2011) through the Euphrates Valley to Iran (eastern escarpment of central Zagros) and the northern Arabian Peninsula (NE Saudi Arabia, Kuwait, Bahrain) including Sir Bani Yas 10) “Schiraz” must be interpreted in its broadest sense; specimens with that origin (e.g., destroyed MSNM syntype of Zamenis persicus Jan or NNW 25446.7) were collected during a time when this appellation or “Shiraz” were used virtually synonymous with today’s Fars Province. With reservations, these records are arbitrarily mapped at 29°37’N 52°32’E, ca. 1°520 m a.s.l. (Fig. 5). The collector of NMW 25446.7 is not registered; this is probably the former ‘Indian Museum’ specimen “given to Dr. F. Stoliczka” (Anderson, 1872; see Schätti et al., 2010). PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 473 Fic. 10 Right lateral head view of Iranian Platyceps karelini (A, USNM 240003, see Fig. 2C) and inter- grades with Coluber chesneii Martin (i.e., P. karelini chesneii): FMNH 20939 (B, subadult) and NMW 25446.7 (C, see Fig. 9B) showing a posterior subocular and ten or eight, respectively, supralabials; USNM 240003 and NMW 25446.7 with a presubocular (see text). Courtesy of Alan Resetar/FMNH and Heinz Grillitsch/NMW. Not to scale. Island (UAE, Fig. 5) !D. The actual distribution limits in Iran (see Comparison) and the potential occurrence in Qatar require specific investigations. Populations of Platyceps cf. ventromaculatus sensu Schätti (2006) from southern Syria and Jordan to eastern Libya (Cyrenaica) are referred to P. karelini rogersi (Anderson, 1893) comb. n. The validity of Rogers’s Racer relies entirely upon differences in dorsal colour pattern, i.e., the transverse blotches considerably wider than interspaces (e.g., Disi, 2002: three unnumb. photographs; Disi et al., 2001: Figs 190-191; Schatti, 2005: Fig. 1; Amr & Disi, 2011: Fig. 72) versus cross-bands narrower 11) Various reports of Coluber ventromaculatus auct. from the United Arab Emirates (e.g., Hornby, 1996; Drew et al., 2005; Gardner, 2009) remain unspecified as to the source of in- formation or voucher specimens. We are not aware of any verified record except for Sir Bani Yas Island (24°20’N 52°36’E) documented by, for example, NMW 32192 (Brown, 1991; Tiedemann, 1991: Figs 2-3; Schatti, 2006; see also Gardner, 2005). Most probably, this racer had been introduced into that island by human activity. The identity of the Sir Bani Yas population (i.e., Platyceps chesneii) is further substantiated by a living specimen photographed by Simon Aspinall (picture provided by Andrew Gardner). 474 B. SCHÄTTI ET AL. < KURHHX:”“Xaagy x EISEN = —! z= FIG. 11 Left in situ hemipenis of Platyceps karelini rogersi HUJ 3185 from Jordan. Scale equals 5 mm. Drawing Heidi Laubscher. (or, at best, equal to interspaces) in P. k. chesneii (Figs 9B-C, see Comparison: first smallprint). The hemipenis of P k. rogersi (Fig. 11) coincides in all structural details with the remaining subspecies and the sister taxon relationship of Karelin’s and Rogers’s Racers is supported by molecular data (see Systematics). Due to its prominent dorsal head and nuchal pattern (see Morphology) typical of Platyceps karelini chesneii, Schatti (2006) referred MNHN 1957.59 (Fig. 2B) from the Golestan-Turkmen border region to that taxon (as P. cf. ventromaculatus, see foot- note 8). The reported distribution of Chesney’s Racer to as far north as “the Kopet Dag area” relies upon this specimen and FMNH 109996 (“most probably belongs to the taxon discussed”) from Northwest Khorasan-e Razavi (Quchan, 37°06’N 58°31’E). MNHN 1957.59 (9, 201 ventrals, 96 subcaudals), very similar to P k. karelini FANH 141604 (2) from Golestan (202 ventrals, 92 subcaudals), is reassigned to Karelin’s Racer, confirming its original identification by Guibé (1957). Besides head and body scale features and size (approximately 110 cm total length), the dorsal colour pattern of FMNH 109996 (3, 216 ventrals, 114 subcaudals) in fact recalls the one found in P. k. chesneii. However, the reputed origin is far beyond the distribution range of Chesney’s Racer and the identification of this specimen pends further studies. ACKNOWLEDGEMENTS E. Nicolas Arnold (London), the late Ilja Darevskij (Saint Petersburg), Robert C. Drewes (San Francisco), Tracy Heath (London), Ivan Ineich (Paris), Jeremy Jacobs (Washington), David A. Kizirian (New York), Konrad Klemmer (Frankfurt on Main), the late Eugen Kramer (Basle/Novaggio), Axel Kwet (Stuttgart), Alan E. Leviton (San Francisco), Fritz-Jürgen Obst (Dresden), Jose P. Rosado (Cambridge, Mass.), Franz Tiedemann (Vienna), Jens V. Vindum (San Francisco), Harold C. Voris (Chicago), and Richard G. Zweifel (New York) approved the loans. Mark-Oliver Rödel (Berlin) made this study possible in its present scope. Linda Acker (Frankfurt on Main), Patrick Campbell (London), Heinz Grillitsch (Vienna), Ivan Ineich (Paris), Günther Köhler (Frankfurt on Main), Colin McCarthy (London), and Alan Resetar (Chicago) provided PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 475 additional information on, and photographs of, certain specimens in their custody. Colin McCarthy and Barry Hughes (London) verified some scale counts. Farhang Torki (Nurabad, Lorestan) and Khosro Rajabizadeh (Kerman) sent data and pictures of two Iranian specimens. Frank Tillack (Berlin) shot photographs of a part of the exa- mined material and clarified many details regarding cited references. He and Steven C. Anderson (Stockton) provided collecting data of illustrated specimens from Turkmenistan and Iran, respectively, shown in Khan (2002). Sonia Fisch-Muller (Geneva), Andrew Gardner (Abu Dhabi), Richard Gemel (Vienna), Mahdi Kazemi (Qom), Heidi Laubscher (Diessenhofen), Ksenja Manuylova (Berlin), Nagwa Othman (Geneva), Andrea Stutz (Arroyo Cruz, Oaxaca), and Philippe Wagneur (Geneva) helped with bibliographic search, old cartographic works and itinerary reports of expeditions, translations, photographs, drawings or logistic and technical support. 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Society for the Study of Amphibians and Reptiles, Athens (Ohio), 522 pp. Appendix A Platyceps k. karelini and hybrid racers from Iran to Pakistan examined (see Material and Methods for approximate coordinates and altitudes; specimens preceded by an asterisk were used for the calculation of cephalic indices; only head and body scale counts available in the case of PMNH specimens [2] and limited data set for CAS 84636 and NMW 25446.3; BMNH 1886.9.21.102-103 not individually labelled). P k. karelini. AFGHANISTAN: BMNH *1882.3.20.2 (Kandahar, 31°37’N 65°42’E, ca. 1’000 m, à, pres. Col. Swinhoe), *1886.9.21.102 (Tirpul [Tir Pol], 34°36’N 61°16’E, 768 m, ©, see remark above, Fig. 2A), *1886.9.21.103 (“Kilki”, ca. 34°00°N 61°25’E [Ziarat-e Fateh Mohammad area], ca. 975 m, 2, see remark above); CAS 84634-36 (Chah-i Anjir, 31°40’N 64°19’E, ca. 800 m, ®, juv. à d, see remark above and Tb. 1), 103785 (“Herat to Islam Qala” [Islam Qaleh], ca. 34°30’N 61°17’E [“about 20 miles from the Iranian border”, Kuh-e Chazirak area], ca. 800-950 m [“Elevation 3100 to 2600 feet”: Clark et al., 1969], 2), 120714 (45 km west of Herat, ca. 34°25’N 61°45’E, 880 m [Clark, 1990], 9); MNHN *8722 [juv.] and *1999.8160 [formerly 8722A] (“Ker Dahar” [Chahar Dahaneh], 31°38’N 65°39’E, ca. 17000 m, dd, see Tbs 2-3); SMF *64629 (Darzi Chah, ca. 29°45’N 65°30’E, ca. 915 m, subad. dg, coll. Jeromie A. Anderson). IRAN: FMNH *141604 (“25 mi. N Pahlavi Dezh” [Aq Qal’ch], ca. 37°25’N 54°27’E, nr.s.l., 2); MHNG 2718.11 (Esfandiar [E Yazd], 33°02’N 57°33’E, ca. 1°400 m, gd), 2718.12 (vic. Naseri, 36°10’N 57°33’E, ca. 900 m, 3, coll. Ashgar Afzal Abadi); MNHN 1957.59 (Dashli Borun, 37°38’N 54°49’E, ca. 50 m, 9, Fig. 2B), 1957.60 (Serakhs, 36°32’N 61°10’E, ca. 280 m, 2, Fig. 9A); NMW 25446.3 (vic. Neh [Nehbandan], ca. 31°32’N 60°02’E, ca. 17175 m, juv., see remark above and Tb. 1), *25446.4 482 B. SCHATTI ET AL. (“Zirkukh” [“Germau”, Garmab], 33°53’N 59°42’E, ca. 17155 m, 2), *25446.5 (Gulu Chahak, 31°19’N 59°25’E, 980 m [Gabriel, 1938], 9), *25446.6 (Chah-e Sam, 30°43’N 60°03’E, 1’206 m [Gabriel, 1938], 8); RUZM *11.1 (Kal-e Do Band River, ca. 33°52’N 50°48’E [SE Markazi], 1’700 m, 2); SMNS *2381 (“Baluchistan”, 2, see Material and Methods); TMUS *994 (southern Tandureh National Park, ca. 37°20’N 58°47’E, ca. 1'550 m, 2), *1000 (ca. 50 km north of Nosratabad, ca. 30°18’N 59°57’E, above 1'500 m, d), 1001 (vic. Kavir-e Lut, east of Tahi salt area, 32°07’N 57°53’E, ca. 700 m, juv.), *1002 (Dasht-e Kavir, 35°04’N 55°03’E, below 750 m, 3); USNM *148631 (1 km south of “Isfadeh” [Esfeden], ca. 33°39’N 59°47’E, ca. 17190 m, 2), *240003 (“Esfahan, 80 km SE of”, ca. 32°24’N 52°05’E, ca. 1'600 m, subad. 9, Figs 2C, 10A); ZMB 6876 (“Mazandaran” [Golestan], ¢); unregistered (vic. Navaran, ca. 34°36°N 51°05’E [15-20 km roughly east of Qom], ca. 900 m, coll. H. Bostanji, examined by Khosro Rajabizadeh, colour pattern ascertained from photographs). PAKISTAN: AMNH *96219 (“near Pishin”, ca. 30°35’N 67°00’E, ca. 17535 m, 3), 96220 (2 miles east of Hanna, Urak Valley, ca. 30°15’N 67°10’E, ca. 27000 m, 3); PMNH 761 (Kalat, 29°01’N 66°35’E, 27018 m, d), 762 (Mastung, 29°48’N 66°51’E, ca. 17700 m, 3); SMF *62924 (Khuzdar, 27°48’N 66°37’E, ca. 17215 m [1’140 m fide Mertens, 1969: 88], 3), *62940 (Quetta, 30°12’N 67°01°E, 1’650 m, ®). Pk. karelini x P. rhodorachis. AFGHANISTAN: BMNH 1873.1.7.10 (“Kila-i-Fath, Sistan” [Qala-ı Fateh, Qaleh-e Fath], 30°34’N 61°50°E, 490 m above sea level, juv. à, “Gen. [Frederic J.] Goldsmid” [register entry, coll. Major Euan Smith], Fig. 8), 1886.9.21.104 (“New Gulran”, 35°06’N 61°41’E, ca. 750 m, d, Fig. 7A); ?CAS 120540 (10 km west of “Tashkurgan” [Tashgorghan, Khulm], ca. 36°42’N 67°36’E, ca. 570 m, d, potential hybrid, Fig. 7B). IRAN: BMNH 1874.11.25.10 (Kerman, ca. 30°17’N 57°05’E, ca. 1°760 m, 3, supposed hybrid, see Material and Methods). P k. karelini x Platvceps sp. AFGHANISTAN: BMNH 1886.9.21.101 (“Helmand” [River], ca. 30°17’N 62°03’E [Arghandab Rod near Chahar Burjak], ca. 500 m, juv. d, supposed hybrid with P mintonorum, see Material and Methods, Fig. 6B). Appendix B Northern Platyceps k. karelini and P. k. karelini x P. rhodorachis examined (only limited data, usually number of ventrals and subcaudals, available for specimens marked with an asterisk). P k. karelini. KAZAKHSTAN: MTKD 13602 (Mujunkum [Mojyunkum] Desert, Dzhambul, 3); ZISP 13436 (Mangyshlak [Mangghyshlaq] Peninsula, 9). KYRGYZSTAN: MHNG 2442.96-97 (Bishkek, £ 2); MTKD 10450 (“Frunze” [Bishkek], 9). TADZHIKISTAN: MHNG 2442.98 (Shahrtuz [Shaartuz], subad.); MTKD 11335 (Gissarskaya Dolina [Kafirnigan], 2), 16095 (Shahrtuz, ¢); NHMB 21058 (“Boba Tag” [Mountains], juv.); ZISP *15811 (Leninabad, juv.; ventrals, suboculars); ZMB 38591 (“Tadschikistan”, 9 ). TURKMENISTAN: CAS 184636 (14,2 km southwest of Madau [Madaw], ¢); FMNH 83961 (Krasnovodsk [Tiirkmenbasy], 2); MCZ 109902 (Repetek, d ); MHNG 1358.27 (“Krasnovodsk Plateau”, 3); MTKD 8281, (Ashgabat, 2); NMW 25446.1-2 (Murgab Valley, d d, incl. possible hybrid, see footnote 5); SMF 18219 (Ashgabat, 9), 18220 (Durun, 2); ZISP 17214.1-2 (Kyzyl Arvat, 9, juv.), 17219 (Bajram Ali, 3), 17386 (Kyzyl Arvat, gd), 17582 (Kushka, d), 18625 (Krasnovodsk, 2), 19031.1-2 (vic. Garrygala [Karakala], 2 9); ZMB 38816 (Sarahs [“Seraks”], ®, possible hybrid). UZBEKISTAN: MHNG 2442.99-100 and 2443.01 (vic. Tashkent, d d), 2443.03 (Bukhara [Buxoro], & ); MTKD 13944 (Bukhara, juv.); ZISP *12907 and *13110 [incl. msr] (Tirmiz [Termez], subad., 2), 14741 (Karakalpakstan, ©), *17682 (G’uzor [Guzar], juv.; ventrals, suboculars). ORIGIN UNSPECIFIED: NHMB 7572 (“Transkaspien”, 9). P. k. karelini x P. rhodorachis. TURKMENISTAN: ZMB 38833 (“Kara-kum” [Garagum] Desert, 50 km east of Imambaba, ®, Fig. 6A). Appendix C Coluber chesneii Martin (i.e., Platyceps karelini chesneii) from Iran examined (includes two intergrades with P. k. karelini; no body scale data available for ZFMK 31602; NMW 28932.1 [Ash Shargat, 35°30’N 43°15’E] is from Iraq, not “Iran” [Schätti, 2006: Appendix]): BMNH 1869.8.28.130 (Bushehr, 28°58’N 50°50’E, near sea level, 5, see footnote 9); FMNH 20939 (Izad Khvast [Yezd-e Khast], 31°31’N 52°07’E, ca. 2’200 m a.s.l., subad. intergrade, PLATYCEPS KARELINI (BRANDT, 1838) FROM IRAN TO PAKISTAN 483 Fig. 10B); FTHR 15300 (“Tang-e Sat”, near Nim Istgah-e Kornas railway station, N Andimeshk District, 32°52’N 48°44’E, 600-1°100 m [Torki, 2010: 30], d, examined by Farhang Torki, lateral head scale conditions and colour pattern ascertained from photographs); MHNG 1359.12 (east of Khosravi, ca. 34°23’N 45°29’E, ca. 250-280 m, 3); MNHN 7470 (“en Perse”, & syn- type of Zamenis persicus Jan); NHMB 15210 (Susangerd, 31°34’N 48°11°E, nr.s.l. 9); NMW 25446.7 (“Schiraz”, £ intergrade, see footnote 10, Figs 9B, 10C), 25466.1 (“Persien”, 3, Fig. 9C); SMF 61869-70 (“Iran”, 8 3); ZFMK 31602 (“Shiraz [...] oder [...] Buschähr” [Bushehr; Werner, 1917], juv.). REVUE SUISSE DE ZOOLOGIE 119 (4): 485-500; décembre 2012 Nouvelles données sur la répartition de la souris des moissons (Micromys minutus Pallas, 1771) en Suisse occidentale et implications pour la gestion de son habitat Michel BLANT!, Paul MARCHESI?, Magali DESCOMBES? & Simon CAPT4 l Partenaire Faune Concept, Ch. de Gratte-Semelle 20, CH-2000 Neuchâtel. E-mail: mblant@vtx.ch 2 Partenaire Faune Concept, DROSERA SA, CP 181, CH-1880 Bex. E-mail: Chablais@drosera-vs.ch 3R. du Jura 34, CH-1373 Chavornay. E-mail: magali.descombes@gmail.com 4 Centre suisse de cartographie de la faune, Passage Maximilien-de-Meuron 6, CH-2000 Neuchatel. E-mail: simon.capt@unine.ch New data on the distribution of the Harvest mouse (Micromys minutus) in western Switzerland and conclusions for the management of its habitat. - Knowledge about the distribution of the Harvest mouse (Micromys minutus) in western Switzerland has been completed by search for typical nests in the field. The authors visited formerly known sites and conducted a prospective search in new favourable habitat (sedge commu- nities). Several new populations have been discovered in the canton of Jura and Vaud. Other sites have to be considered as abandoned, such as the region between the lakes of Bienne and Neuchatel. Management measures in favour of the species are proposed. Keywords: Harvest mouse - fragmentation - isolation - nest - western Switzerland. Nouvelles données sur la répartition de la souris des moissons (Micromys minutus) en Suisse occidentale et implications pour la gestion de son habitat. - La répartition de la souris des moissons (Micromys minutus) en Suisse occidentale a été actualisée par la méthode de recherche des nids. Les auteurs ont procédé par contrôle des anciens sites connus et par prospections nouvelles dans les habitats favorables (carigaies). Plusieurs nouvelles populations ont été mises en Evidence dans les cantons du Jura et de Vaud. D'autres sont a considérer comme éteintes, comme dans l'Entre-deux-Lacs neuchatelois. Des mesures de gestion favorables a l'espèce sont proposées. Mots-clés: Souris des moissons - fragmentation - isolement - nids - Suisse occidentale. Manuscrit accepté le 24.07.2012 486 M. BLANT ET AL. INTRODUCTION En Suisse, la souris des moissons (Micromys minutus) est une espèce caracté- ristique des prairies humides hautes, des zones marécageuses, des rives de cours d'eau et roselieres (Rahm, 1995). Elle est intimement liée aux laiches à larges feuilles (Carex sp.) qu'elle utilise pour tresser son nid, mais peut accessoirement utiliser des graminées comme Calamagrostis sp. ou des céréales cultivées. Elle n'a pas été signalée en Suisse dans les milieux cultivés. En Europe, cette espéce a décliné dans de nombreuses régions en raison de la perte ou de l’altération des habitats par les drainages, l’assè- chement des zones humides et l’intensification des pratiques agricoles (Mitchell-Jones et al., 1999). Les champs de blé constituent son habitat principal en Grande-Bretagne, et les rizières dans le nord de l'Italie (Rham, 1995). Plusieurs populations de souris des moissons (Micromys minutus) ont été loca- lisées durant ces dix dernières années dans l'ouest de la Suisse, suite à des recherches ciblées en vue d’une amélioration des connaissances cantonales sur cette espèce et pour actualiser la Liste Rouge des mammiferes terrestres. Ce petit rongeur, classé comme vulnérable dans la premiere Liste Rouge des mammiferes de Suisse (Duelli, 1994), est probablement un des mammiferes terrestres les plus menacés actuellement dans notre pays. Les données a son propos restent lacunaires et pas toujours fiables, les nids pouvant étre confondus avec ceux du muscardin (Muscardinus avellanarius). De méme, les critères de distinction craniens sont ténus, et il y a des risques de confusion avec la souris domestique pour les cranes trouvés dans des pelotes de réjection de rapaces nocturnes (Marchesi ef al., 2008). L’atlas des mammifères de la Suisse (Rahm in Hausser, 1995) signalait l’espèce à Genève, à l’ouest du lac Léman (La Versoix, Coppet), sur la rive sud du lac de Neuchâtel (Grande Cariçaie), dans l'Entre-deux-lacs entre Cressier et Le Landeron, sur la rive nord du lac de Morat et, plus au nord, à Bonfol et dans les environs de Bâle. La plupart de ces données, à l'exemple des cantons de Neuchâtel et du Jura, remontent à plus de 40 ans, soit entre les années 1960 et 1970. Durant les deux dernières décennies, la souris des moissons a été principalement observée dans la Grande Cariçaie. Elle y trouve certainement les conditions d'habitat les plus appropriées, et y développe sans doute la plus importante population du territoire suisse en raison de l’étendue de l’habitat disponible. En dehors de ce grand complexe marécageux, elle a tout d’abord été retrouvée dans le canton du Jura, grâce à l’identification d’un crâne dans une pelote de réjection de chouette effraie (Tyto alba) à Damphreux (Blant et al., 2003). Par la suite, des nids ont pu être observés sur le site des Marais de Damphreux en 2005, à l’occasion de relevés destinés à tester la méthode de mise en évidence de l’espèce, sur mandat du Centre suisse de cartographie de la faune (CSCF) (Blant ef al., 2005). Des relevés plus systématiques y ont encore été effectués par la suite sur mandat de la Fondation des Marais de Damphreux, préalablement à des travaux de réaménagement, suggérant l’existence d’une importante population sur ce site (Blant, 2007). Ces résultats encourageants nous ont incités à vérifier le statut de cette espèce dans un contexte biogéographique élargi, à savoir la Suisse occidentale, afin de vérifier si les populations mentionnées par l'atlas subsistaient encore actuellement. L’objectif de cette publication est de présenter une synthèse des résultats des recherches dans les différents cantons concernés. Les études proposées devaient entre autres fournir des REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 487 indications sur le degré d'isolement des populations résiduelles détectées en dehors du complexe de la Grande Carigaie, qui n'est pas discuté ici. La typologie des milieux occupés par la souris des moissons et leur état général ayant été relevés, l'article propose des recommandations pour un entretien des biotopes favorisant durablement le maintien de cette espèce menacée. MATÉRIEL ET MÉTHODES Les anciennes données (antérieures à 2002) ont été recherchées dans la base de données du CSCF, et par consultation des musées d'histoire naturelle (Musées d'histoire naturelle de Fribourg - MHNF, de Neuchatel, de La Chaux-de-Fonds, Musée de Zoologie de Lausanne, Musée de la Nature de Sion, Musée des sciences naturelles de Porrentruy). Les données du Muséum d'histoire naturelle de Genève - MHNG, du Musée d'histoire naturelle de Berne - NMBE et de celui de Bâle - NMBA étaient déjà connues du CSCF pour l'élaboration de l'atlas. Des naturalistes des régions concernées, actifs dans le domaine des mammifères, ont également été contactés. La mise en évidence des populations récentes (postérieures à 2002) de souris des moissons s’est faite par la recherche de ses nids. Cette méthode a été testée tout d’abord dans des biotopes d'où provenaient des observations (notamment Grande Cariçaie et région bâloise), en prévision de la révision de la Liste Rouge des mammi- fères terrestres (Blant ef al., 2005). Elle a été appliquée ensuite, dans un but essentiel- lement qualitatif (présence/absence), dans des sites parcourus pour leur potentiel d'habitat favorable. Dans quelques sites, une prospection quantitative a été faite de manière à pouvoir calculer un indice de population ou pour des questions relatives à la gestion du milieu. Une limite de surface minimale de 3 ares a été fixée pour la prospection des nouvelles surfaces potentielles. Les fragments de cariçaie inférieurs à cette dimension ont en général été écartés. On peut succinctement résumer la méthode de détection comme suit: - Parcours du site en automne (septembre à novembre) selon un ou plusieurs transects d’échantillonnage déterminés d'abord sur orthophoto, puis sur le terrain en fonction des caractéristiques végétales. - Recherche à vue des nids dans les cariçaies, en écartant les touffes de laîches pour trouver les nids accrochés vers la mi-hauteur des plants. - Recherche durant au moins 1 heure (‘4 h pour les plus petits sites) pour attester la présence/absence. - Recherche dans le Caricion (Carex acutiformis ou autre), ou à défaut dans le Phalaridion, le Phragmidion ou la mégaphorbiaie à Filipendula. - Photographie des nids et vérification de l’attachement au support, caracté - ristique du tressage de Micromys sur le matériel vivant (les nids de Muscardinus sont tressés avec du matériel coupé et sont posés sur un support, sans y être vraiment attachés) (voir Blant et al., 2005) Les prospections se sont limitées aux régions de plaine ou de vallées jusqu’à 700 m d’altitude, en raison de l’affinité de l’espèce pour ces niveaux altitudinaux (Rahm, 1995; Butet & Paillat, 1998). A88 M. BLANT ET AL. RESULTATS Vue d'ensemble La souris des moissons a été localisée dans 21 sites situés du canton de Genéve a celui de Bale-campagne, jusqu'en 1990 (données des musées et CSCF, tableau 1). Par la suite, entre 1996 et 2001, de nombreuses captures ont été réalisées par le Groupe d'étude et de gestion (GEG) de la Grande Caricaie, sur la rive sud du lac de Neuchatel. Les animaux étaient piégés dans des seaux destinés au contrôle de la migration printa- niere des batraciens (A. Gander, comm. pers.). Des données extérieures à la Grande Carigaie font par contre défaut entre 1990 et 2001. Durant nos prospections opérées des 2005, en dehors de la Grande Caricaie, la souris des moissons a été retrouvée (tableau 2) dans 18 sites dans le canton du Jura (17 sites nouveaux + un site ancien), dans deux sites dans le canton de Vaud (nouveaux), et dans un seul site dans le canton de Fribourg (ancien). Dans le canton de Neuchatel, aucun contröle n'a pu permettre de déceler l'espèce, pourtant présente par le passé dans un site au moins. Dans le canton de Genève, aucun site potentiellement favorable n'a permis de retrouver l'espèce. Elle n'existe probablement pas dans le canton du Valais. Le Jura bernois n'a pas été prospecté en raison de l'altitude trop élevée. La figure 1 présente la distribution exhaustive de l'espèce selon les données connues anciennes (jusqu'en 2001) et les découvertes récentes (jusqu'en 2011). La carte mentionne également tous les sites prospectés sans résultat positif. TABLEAU 1. Données anciennes de Micromys minutus enregistrés dans la base de données du CSCF, situation jusqu'en 1990 pour la Suisse occidentale. Sites avec capture ou individu trouvé mort (type = 1), observations visuelles directes (type = 2) ou nids (type = 3), localités de mise en évidence par pelotes de réjection non mentionnées (situation exacte inconnue). Italique: observations douteuses (confusion possible avec Muscardinus). Ct Commune Lieu-dit Année Type Collection GE Genève Av. d'Aire 1945 3 MHNG/CSCF VD Valeyres-sous-Mont _ V.-sous-Mont 1960 CSCF VD Bogis-Bossey Bogis-Bossey 1966 3 MHNG/CSCF VD Bellerive 1974 NMBE/CSCF VD Cudrefin 1976 CSCF VD Noville Les Glariers 1985 3 CSCF VD Noville Gros Brasset 1988 3 CSCF FR Gletterens Gletterens 1957 CSCF FR Bas-Vully Vully 1980 NMBE/CSCF FR Font Vers L'Eglise 1980 CSCF FR Cheyres Cheyres 1981 l CSCF FR Galmiz Le Chablais 2 3 MHNF NE Cressier 1966, 1970 12 MHNG/CSCF JU Bonfol Bonfol 1960 2 CSCF BS Basel Bäumlihof-Hörnli 1909 NMBA/CSCF BS Basel Lange Erlen 1940 2 CSCF BS Basel Rheinhafen 1950 NMBA/CSCF BS Riehen Eisweiher 1980 CSCF BL Allschwil Herzogenmatt 1980 CSCF BL Bottmingen Bruderholzhof 1980 CSCF SO Rodersdorf Rodersdorf 1990 l CSCF REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 489 Fond de carte SWISSTOPO x ha 5 a 2 3, u oO D oO ® FIG. 1 - Carte de répartition de la souris des moissons (Micromys minutus) en Suisse occidentale. Triangles noirs: base de données du CSCF, observations anciennes, avant 2002 (tableau 1). Ronds noirs: observations nouvelles, après 2002. Ronds blancs: sites prospectés sans obser- vations. 490 M. BLANT ET AL. TABLEAU 2. Sites prospectés en Suisse occidentale dés 2005, hors de la Grande Caricaie, avec presence constatee de Micromys minutus. La date mentionnée (avec n = nombre de nids correspondants) est la dernière observation réalisée pour chaque site, certains ayant été suivis sur plusieurs années. Ct Commune Lieu-dit Date n Altitude Coord X Coord Y Marais du Grand- VD Commugny Bataillard 13.09.2010 24 468 500100 132780 VD Salavaux Petit Lac 13.10.2011 16 431 569050 196800 FR Galmiz Le Chablais 24.11.2010 1 431 576300 200180 JU Damphreux Les Coeudres 31.08.2007 28 440 575400 258000 JU Damphreux En Pràtchi 31.08.2007 1 430 575600 258875 JU Bonfol Etangs de Bonfol 23.10.2008 7 440 579800 257150 JU Bonfol Etangs Rougeat 23.10.2008 1 445 579350 256950 JU Bonfol Queue de Chat 23.10.2008 8 445 579400 258700 JU Vendlincourt Etang 23.10.2008 2 450 379350 255700 JU Alle Pisciculture 22.10.2008 6 455 577500 252700 JU Cornol Pre Raisin 22.10.2008 1 530 580350 250900 JU Miécourt Allaine 22.10.2008 2 470 579150 253100 JU Boécourt Les Esserts 24.10.2008 5 490 583850 243700 JU Courfaivre Les Aingles 24.10.2008 1 465 586750 243250 JU Delémont Le Colliard 29.10.2010 1 420 594350 246500 JU Boécourt Etang Bourquard 22.10.2009 I 497 583350 243900 JU Bure Le Teurion 30.10.2009 6 575 568550 254950 JU Courtemaiche Tcherteau 30.10.2009 1 545 569150 256050 JU Courtemaiche Etang 06.11.2009 2 390 570650 257300 JU Lugnez La Vouevre 06.11.2009 1 430 575600 259550 JU Vendlincourt Combe aux Chiens 06.11.2009 1 445 578000 256700 Details par cantons CANTON DE GENEVE: En octobre 2007, 6 sites ont été prospectes dans le canton de Genève. Aucun indice de présence n’a été retrouvé dans ces derniers. Ainsi, une observation de nid au Moulin-de-Vert, faite par un garde de l’environnement il y a une quinzaine d’années, n’a pas pu étre confirmée. Ces différents sites se situent entre 350 et 470 m d’altitude et d’une superficie variant de 2 a 7 ha. Ils sont composés de mosaïques de milieux humides, ne présentant que de petites surfaces de carigaie et phragmitaies-carigaies. La présence de nids et d'une mächoire dans une pelote de réjection est signalée par un observateur (J. Bordon) a l'Etournel (Ain, France) a moins d'un kilometre de la frontiere cantonale (G. Dändliker, comm. pers.). Les autres observations les plus proches le long du Rhône apparaissent dans le Marais de Lavours, situé à une quaran- taine de kilomètres au sud du canton de Genève (J. Gilliéron, comm. pers.). CANTON DE VAUD: En octobre 2007, quatre nids typiques de la souris des moissons ont été retrouvés dans le marais du Grand Bataillard (commune de Commugny, VD). Cette observation confirme la présence encore actuelle de cette espece mise en evidence en 1966 déjà dans ce site par la collecte de trois nids par V. Aellen et P. Strinati (commune de Bogis-Bossey, VD), puis par l’observation d’un individu faite il y a une quinzaine d’année (S. Sachot, comm. pers.). En juin 2008, une série de pièges et tunnels à traces ont été déposés à différents endroits dans ce marais et ses abords pour tester d’autres méthodes de detection de REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 49] cette espèce discrète. Aucune souris des moissons n’a cependant été capturée et aucune observation de traces ou de nids n’a été faite à ce moment la (Marchesi & Descombes, 2008). Afin de confirmer les premières observations réalisées en 2007, un nouveau transect au travers du marais a été effectué en octobre 2008: 39 nids typiques de la souris des moissons, plus ou moins frais ou en construction, ont pu être observés prin- cipalement dans un milieu composé d’une cariçaie à Carex acutiformis et parfois d’un mélange de cariçaie avec du Phalaris. En moyenne, les nids se situaient à 76 cm de hauteur (n= 39; min= 40 cm, max=100 cm). En septembre 2010, un nouveau transect a permis de trouver 24 nids en parcourant une surface représentant environ la moitié du bas-marais. De vastes sur- faces sont malheureusement de plus en plus colonisées uniformément par le solidage (Solidago canadensis), rendant le milieu stérile à toute autre espèce végétale, et non propice à la souris des moissons (Marchesi & Blant, 2010). Au cours de l’automne 2010, des prospections effectuées dans 20 autres sites marécageux du canton de Vaud (hors Grande Cariçaie) n’ont permis de découvrir qu'un seul nouveau site avec des indices d'occupation sûrs de ce rongeur: la roselière de Salavaux située à l’extrémité Ouest du lac de Morat (Petit Lac, embouchure de la Broye). Quatre nids y ont été observés sur une petite bande très étroite entre forêt riveraine et roselière lacustre, en rive gauche de la Broye. Pas moins de 16 nids ont pu y être comptabilisés encore en 2011, tandis qu’un seul autre nid a pu être trouvé de l’autre côté de la Broye, en rive droite de l’embouchure. Dans le marais de Chavornay (Creux de Terre), un seul nid tressé a été trouvé dans la roselière-cariçaie malgré de longues fouilles. Une nouvelle recherche infructueuse effectuée en 2011 nous incite à penser que le nid trouvé en 2010 était celui d’un muscardin (nid tressé non attaché au support). Les différentes recherches effectuées aux Grangettes entre 2002 et 2010 n'ont pas permis de mettre en évidence la souris des moissons malgré la recherche de nid et des essais de piégeage effectués en collaboration avec Peter Vogel de l’Université de Lausanne. Bien qu'une grande surface de milieux favorables ait été prospectée, les seuls nids trouvés ont été attribués au muscardin, aucun n’ayant les caractéristiques du tressage sur plante vivante effectué par Micromys. En 2002, un muscardin hibernait d'ailleurs dans un nid reposant sur une branchette de saule. Aucune souris des moissons n'a jamais été découverte lors des entretiens du site (O. Epars, comm. pers.). Les sites vaudois occupés se situent a moins de 500 m d’altitude et semblent se répartir actuellement en deux sous-populations: la première située dans le Nord vaudois inféodée au bassin du Rhin, la deuxième représentée par la population du Grand Bataillard, dernier site connu actuellement pour cette espèce dans le bassin du Rhône suisse. Aucune observation n’a été faite entre ces deux sous-populations, dans la région des collines du Gros de Vaud. Le contact aurait pourtant pu se faire histo - riquement au niveau de la Sarraz, le seul point bas situé à moins de 500 m d’altitude. CANTON DE FRIBOURG: Un site a été visité dans le canton de Fribourg, sur les bords du lac de Morat où l’espèce nous avait été signalée par André Fasel, directeur du Musée d’histoire naturelle de Fribourg (Marchesi & Blant, 2010). Comme dans la Grande Cariçaie, il existe sur la rive Est du lac une bande de cariçaie plus ou moins 492 M. BLANT ET AL. large, entre la forét riveraine et la roselière lacustre. Cette bande est plus étroite que sur la rive sud du lac de Neuchatel, mais apparait cependant trés favorable. Un seul nid a été trouve lors du contröle, l'essentiel du milieu ayant déjà été fauché. Deux autres sites visites en Gruyere (région de Semsales-Vaulruz, à plus de 800 m d'altitude) a l'occasion de piégeages se sont révélés négatifs, aucun nid de souris des moissons n'ayant pu étre trouvé malgré la présence de surfaces de carigaie jugées favorables dans des marais en bordure de plans d'eau. CANTON DE NEUCHATEL: Sur 19 sites prospectés, aucun n'a été trouvé colonisé par la souris des moissons (Blant & Marchesi, 2009). Ces sites étaient situés entre 430 et 780 m d'altitude. L'étendue des carigaies et phragmitaies-carigaies (les caricaies pures sont rares dans le canton de Neuchatel) était comprise entre plusieurs hectares (rives lacustres) et 3 ares (surfaces résiduelles le long de canaux dans le Val-de-Ruz). Le site ancien de Cressier est aujourd'hui occupé par une raffinerie de pétrole. Dans l'enceinte méme de l'usine et a sa périphérie (réserve naturelle de la Vieille- Thielle) subsistent des milieux de phragmitaie-caricaie résiduels de petite taille. Aucun indice de l'espèce n'a pu y être décelé. Une importante population y vivait pourtant dans les années 1960 (A. Meylan, comm. pers.). Les dernières données en ces lieux provenaient d'animaux découverts lors de la construction de la raffinerie, mise en service en 1966. La population s'était alors réfugiée sur des tas de terre dégrappée (A. Meylan, comm. pers.), au moins jusqu'en 1970. Les souris des moissons ont subsisté encore quelque temps dans ce site (feu A. Schertenleib, comm. pers.) sans qu'il soit possible de déterminer précisément le moment de leur disparition. CANTON DU VALAIS: Aucun nid n’a été trouvé dans les rares milieux humides valaisans présentant quelques surfaces d’habitats potentiellement favorables (cariçaies et roselières peu denses). Cette espèce n’a d’ailleurs jamais été signalée dans ce canton. CANTON DU JURA: Sur 29 sites visités, 17 étaient colonisés par la souris des moissons, avec des populations plus ou moins importantes (entre 1 et 28 nids recensés). Ces sites étaient situés entre 390 et 575 m d'altitude, en Ajoie et dans la Vallée de Delémont (Blant, 2007). L'étendue des cariçaies abritant l'espèce peut être de plusieurs hectares (Etangs de Damphreux), mais aussi de moins de 10 ares (ancienne pisciculture d'Alle, bassin de décantation de Tcherteau à Courtemaiche). Les sites non occupés étaient situés entre 400 et 610 m d'altitude. On trouve dans cette catégorie des objets présentant une surface de cariçaie à priori favorable de plusieurs hectares comme l'étang A16 des Montoies (Courfaivre). Réalisée il y a une quinzaine d'année, cette zone humide pourrait être encore trop récente pour abriter l'espèce, qui semble peu mobile. Aucun des quatre sites de compensation A16 entièrement nouveaux n'a d'ailleurs été colonisé par l'espèce. La découverte d’un crâne dans une pelote de réjection d’effraie dans une région proche de la place d’armes de Bure nous a incités à visiter les quelques zones humides de ce secteur karstique de la Haute Ajoie. L'espèce colonise deux bassins de décan - tation dans le périmètre d'exercices militaires. Ces bassins sont transformés en bas- marais à Typha latifolia. La souris des moissons s'y reproduit malgré une faible couverture en Carex Sp. REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE FIG. 2. Une surface de carıcaie en bordure d'un étang à Damphreux, milieu typique de la souris des moissons (photo M. Blant). Fic. 3. Une surface de bas-marais en bordure de la place d'armes de Bure, milieu secondaire avec Typha et graminées (Agrostis gigantea) (photo M. Blant). 494 M. BLANT ET AL. REGION BALOISE: Deux sites ont été contröles dans cette région, pour tenter de retrouver la souris des moissons, dont la présence ancienne était connue (Fig. 1). Ces deux sites (cantons de Bale-campagne et Soleure) se sont révélés positifs, avec au moins un nid. Au Herzogenmatt (Allschwil BL), il s'agit d'une population introduite dans un site revitalisé, alors qu'à Rodersdorf (SO) sa présence est probablement naturelle. DISCUSSION Typologie des milieux colonises et non colonises La carigaie (Magnocaricion) des bordures d'étangs de faible profondeur (avec en particulier Carex acutiformis) ou des rives de lacs et de cours d'eau lents constitue le milieu le plus colonisé par la souris des moissons, parmi les sites visités. Cependant, l'espèce peut occuper également parfois des phragmitaies avec Carex sp. (rives de l'Allaine, rive du lac de Morat), des bas-marais à 7ypha latifolia (place d'armes de Bure) ou des phalaridaies à Carex le long des cours d'eau, comme au Grand Bataillard. Bohme (1978) cite comme milieux primaires les associations à Calamagrostis, Typha et Carex. La colonisation des phragmitaies peut étre limitée par le diamétre des tiges des vieux roseaux (Rahm, 1995), souvent trop épais pour les petites pattes du rongeur. Sa présence dans les cultures céréalières serait remarquée surtout dans le nord et l’ouest de l'Europe, en particulier en Grande-Bretagne. En Italie, elle fréquente surtout les rizières comme milieu humide de substitution. Sa présence dans d'autres milieux tels les jeunes pépinicres est encore mentionnée dans la littérature (Moore et al., 2003). Une des conditions nécessaires à la colonisation est la hauteur atteinte par les tiges de Carex, le tressage des nids débutant lorsque la végétation atteint environ 1,5 m selon les observations faites à Damphreux (Berret & Goetschi, 2009). De cette manière, tressés à mi-hauteur des tiges, les nids sont suffisamment élevés au-dessus du sol. Ils se situent aussi le plus souvent au-dessus d'un terrain inondable, au moins saisonnièrement. À l'inverse, les roselières terrestres et les cariçaies résiduelles trop sèches, avec une hauteur de végétation limitée (< 50 cm), ou encore les marais à petits Carex (Parvocariçaie) n'abritent pas l'espèce. En l'absence de Carex, le nid peut être tressé avec des tiges de graminées, comme dans le Phalaridion (Phalaris arundinacea) ou les bas-marais à Typha (avec Agrostris gigantea). C'est le cas sur certaines surfaces du Grand Bataillard, ainsi qu'à Bure et au Herzogenmatt. Les colonies de souris des moissons occupent des milieux de taille diverse, de plusieurs hectares a seulement quelques ares. Les surfaces de carigaie colonisées peu- vent même être encore plus petites. Dans ce cas, elles font généralement partie ou sont en contact avec des zones humides plus vastes. Par exemple, une surface de 10 m2 a Bonfol (JU) contenait un nid, au bord d'un étang aux rives dégagées et distant de plus de 800 m d'autres surfaces de caricaie favorable abritant d'autres nids. Aux Esserts a Boécourt (JU), une surface de moins de 100 m? contenait 11 nids de souris des moissons (Schmidlin, 2011) ! Toutefois, il est clair que les colonies occupant de grandes surfaces peuvent étre plus importantes et ont donc plus de chance de subsister a long terme dans un paysage fragmente. REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 495 Fic. 5. Le Grand Bataillard (VD) en novembre 2010, avec la surface fauchée a gauche et la bande préservée en lisière à droite qui reste favorable pour la construction des nids l’année suivante (photo P. Marchesi). 496 M. BLANT ET AL. Dans le canton de Neuchatel, la phragmitaie-carigaie de la rive nord du lac n'est apparemment pas occupée par l'espèce. Elle n'est constituée tantôt que d'une bande mince de Carex pseudocyperus, tantôt plus large (jusqu'à 10-15 m) en retrait de la roselière lacustre. Le substrat minéral (galets) ne laisse cependant se développer que des tiges peu élevées, d'environ 1 m (par exemple La Ramée à Marin). Les quelques zones à végétation plus haute (1,5 à 2 m) sont rares (par exemple port de Bevaix). Dans le canton de Genève, la plupart des sites actuels potentiellement favo- rables à la souris des moissons sont des milieux humides revitalisés, qui étaient souvent recouverts auparavant par la forêt. Un retour naturel de cette espèce dans ces bas- marais est lié à un entretien adéquat favorisant les surfaces de cariçaie non boisées. Urbanisation et fragmentation comme cause de l'absence La disparition de la colonie de souris des moissons de l'Entre-deux-Lacs dans le canton de Neuchâtel a fait suite à la construction de la raffinerie de Cressier. L'industrialisation croissante et les modifications agricoles structurales (remaniements, drainages) ont limité ensuite les possibilités d'existence de l'espèce, qui semble devoir être considérée maintenant comme disparue de la région. Les quelques surfaces de Carex à longues tiges, subsistant en pied de berges de canaux de la Vieille Thielle ou d’un exutoire des drainages de la raffinerie sont de surfaces probablement trop modestes pour abriter encore l’espèce (au plus 20 à 30 m2). Plusieurs sites visités se trouvent actuellement isolés par les surfaces cultivées et construites, ou sont déconnectés du réseau hydrographique, formant souvent les der- nières liaisons biologiques entre les zones humides. Ceci fait suite au développement agricole et urbain et à la mise en tuyau, dans le passé, de tronçons de cours d'eau. C'est le cas par exemple de la petite zone humide de Dozière à Delémont, des anciennes rives du lac de Neuchâtel près de la jonction AS à Auvernier-Colombier ou encore de la zone humide de Belmont, à Boudry. Ces sites n'abritent pas ou plus la souris des moissons. Cependant, la présence de l'espèce sur la place d'armes de Bure, à plus de 1,5 km du réseau hydrographique représenté par l'Allaine, laisse envisager une possi - bilité pour elle de se maintenir ou de coloniser des sites refuges à des distances proches des zones réservoirs, ceci même sans corridor offrant des milieux humides relais. Isolement géographique et altitude L'absence de colonies dans le Val-de-Ruz malgré la présence de milieux humides de surface assez importante (p. ex. La Paulière) pourrait être due à une cause purement géographique. Les gorges du Seyon situées au bas de la vallée constituent un verrou assez serré et peu propice au transit d’une petite espèce de milieu plutôt ouvert. L'absence dans le Val-de-Travers est due probablement au même type de barrière topographique (gorges de l'Areuse). L'absence de l'espèce aux Grangettes peut s'expliquer de manière similaire. Le delta du Rhône s'est transformé progressivement en milieu palustre à la fin du retrait des glaciers, mais cette région est cependant restée plus ou moins isolée du Plateau suisse pour certaines espèces par le lac ainsi que par les versants escarpés et boisés des Préalpes qui tombent abruptement dans le lac au niveau du Haut Léman (Riviera suisse, Chablais français). Plusieurs espèces de REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 497 batraciens comme le crapaud accoucheur (Alytes obstetricans) ou le crapaud calamite (Bufo calamita) n’ont de méme probablement pas pu franchir ces obstacles pour se répandre dans la haute vallée du Rhône située en amont du Léman (Marchesi, obs. pers.). L'altitude elle-méme est probablement aussi un facteur limitant dans la distri- bution de l'espèce, expliquant son absence dans le Gros de Vaud, les vallées neuchäte- loises ou les Préalpes fribourgeoises. Les observations faites en Suisse se situent en effet toutes à moins de 600 m d’altitude, la station la plus basse se situant a 310 m (Herzogenmatt, Allschwil BL), et la plus haute à 575 m (Le Teurion, Bure JU). En France voisine, Michelat et al. (2005) signalent la présence de ce petit rongeur jusqu'à pres de 1000 m a Mouthe (Doubs, France), d'apres des pelotes de réjection et une capture a la Tourbière des Pontets. Il n'est toutefois pas clairement établi que ce site NATURA 2000 abrite une population reproductrice de souris des moissons. Quéré et Louarn (2011) mentionnent en effet que cette espèce dépasse rarement 500 m d'altitude en France. L’existence d’une population viable à 1000 m d’altitude mériterait d’être authentifiée par une preuve de reproduction. Importance respective des populations dans le contexte suisse Les données du Groupe d'étude et de gestion (GEG) confirment une population de souris des moissons occupant de vastes surfaces de la Grande Caricaie, sur la rive sud du lac de Neuchâtel. Il s’agit assurément de la population la plus importante en nombre de cette espèce pour la Suisse. Toutefois, le long des transects de comptage, des fluctuations annuelles du nombre de nids sont constatées (A. Gander, comm. pers.). Il est possible que la fauche des roselières ait une influence négative momentanée sur ce petit rongeur. Nous avons aussi remarqué au Grand Bataillard que les surfaces fauchées ne conviennent pas l’année suivante car elles ne présentent pas encore de roseaux ou de Carex de hauteur suffisante pour la construction des nids. L'Ajoie dans le canton du Jura apparaît comme étant la seconde région possé- dant une population importante de cette espèce en Suisse. Répartie dans un vaste réseau de sites abritant des colonies plus ou moins nombreuses, elle est en relation avec celle du bassin sud-est de la France (Franche-Comté, Alsace du Sud), très riche en étangs et cours d'eau. En 1982-83, une analyse de pelotes de réjection faisait mention d’un nombre relativement important de crânes (68 exemplaires sur 6384) de souris des moissons dans les proies de l'effraie des clochers (Buser, 1984). Il apparaît donc que ce rongeur s'est bien maintenu dans cette région durant les 30 dernières années. Trois autres populations certifiées récemment en Suisse se situent dans le bassin du Rhin: Lac de Morat, Vallée de Delémont, Bâle. Elles sont plus isolées et numé- riquement plus faibles. En ce qui concerne le bassin rhodanien, seuls les marais de la Versoix (Grand Bataillard) abritent encore une population numériquement importante, qui était certainement à l’époque en relation avec d'autres sites de la vallée du Rhône français, bien qu'aucune trace récente n'ait été trouvée entre deux à Genève. Au Tessin, la souris des moissons a vraisemblablement disparu récemment du dernier site connu dans le Mendrisiotto, aucune trace n'ayant pu être retrouvée dans des cariçaies résiduelles (Blant ef al., 2005 ; T. Maddalena, comm. pers.). Cette extinction 498 M. BLANT ET AL. locale est sans aucun doute due aussi à l'urbanisation et à l'implantation de zones industrielles, tout comme dans l'Entre-deux-Lacs neuchätelois. Elements pour une gestion des milieux favorable a l'espèce Dans la Grande Carigaie, l'espèce est retenue comme prioritaire dans les plans de gestion avec comme objectif de suivi l'amélioration des exigences écologiques et de la distribution des espèces de priorité 1 (GEG, 2007). Ce leitmotiv devrait être appliqué à tous les autres sites abritant encore cette espèce très menacée en Suisse, en particulier les réserves naturelles occupées par des populations prosperes (Marais de Damphreux, Grand Bataillard, Les Esserts/Boécourt). Cependant, de nombreux autres sites de plus faible dimension abritent encore l'espèce, mais sont menacés a plus ou moins court terme d'assèchement et/ou d'afforestation. Il est urgent de revitaliser le milieu humide de certains de ces sites, de manière à renforcer la cariçaie. C'est le cas par exemple à Salavaux, vers l'embouchure de la Broye, ou aux Aingles a Courfaivre dans la vallée de Delémont. Les recommandations suivantes sont proposées pour la conservation de l'espèce: 1) Inventorier précisément l’espèce et ses besoins en habitat dans chaque site occupe (surface utilisée, densité et localisation des nids, type d’habitat), et assurer un monitoring annuel prenant en compte les effets de gestion et d’entretien des milieux. 2) Prévoir une fauche alternée sur un rythme d’au moins 3 ans dans les marais concernés, et laisser des bandes en friche, moins souvent entretenues, en lisiere, au bord des bouquets de buissons et, si besoin, une bande non fauchée de 2-3 m de large à travers les grandes surfaces dénudées comme liaison entre les milieux favorables. 3) Effectuer la fauche de préférence d’octobre a mars pour ne pas porter atteinte aux nids occupés et a habitat estival. Eviter de faucher les touradons, car ils servent souvent de refuge a ce rongeur. 4) Laisser dans les lisières et autres surfaces moins intéressantes du marais des tas de branchages et de litiére, ou des andins de roseaux comme refuge. Cette mesure probablement primordiale pour améliorer la survie hivernale de l’espèce est également utile à beaucoup d’autres animaux. 5) Suivant les sites, effectuer des interventions de débroussaillage partiel, utiles pour conserver l’habitat marécageux. Cependant, elles doivent tenir compte du fait que ce rongeur se réfugie souvent dans les boisements clairs et les lisières en hiver. Ces travaux se feront donc de manière fine et ponctuelle, et autant que possible sans grosses machines qui écrasent le sol ou les tas de branchages. Par divers aménagements, les castors peuvent être incités à contribuer naturellement à ce travail d’éclaircie. 6) Creuser des petites mares et fossés ou dégrapper des dépressions humides dans les grandes cariçaies monotones pour raviver la magnocariçaie et favoriser le développement de tiges élevées. Cela permettrait également de diversifier les milieux et offrir un spectre plus large de nourriture durant les saisons (graines, insectes). 7) Lutter contre l'envahissement par le solidage et les autres néophytes dans les sites protégés. REPARTITION DE LA SOURIS DES MOISSONS EN SUISSE 499 REMERCIEMENTS Nous remercions l'office de l'environnement du canton du Jura et les services de la faune, des foréts et de la nature des cantons de Neuchatel, Vaud et Genève pour le soutien financier apporté a ce travail. Des données anciennes ont été fournies amica- lement par S. Sachot, A. Meylan, J.-C. Bouvier, A. Fasel, J.-D. Blant, J.-M. Gisiger, O. Glaizot, J. Gilliéron et M. Ruedi. Nous remercions aussi A. Gander pour ses commentaires et pour sa mise a disposition du plan de gestion de la Grande Caricaie, P. Vogel pour ses conseils avisés ainsi que l'entreprise Petroplus pour nous avoir permis de rechercher l'espèce dans les milieux situés dans l'enceinte de la raffinerie de Cressier. BIBLIOGRAPHIE BLANT, M., OPPLIGER, J. & SCHALLER, J.-C. 2003. Note a propos du régime alimentaire de l'Effraie des clochers 7yto alba en région jurassienne, obtenu par l'analyse de pelotes de rejection. Nos Oiseaux 50: 15-20. BLANT, M., MARCHESI, P. & MADDALENA, T. 2005. Mise en évidence de la souris des moissons (Micromys minutus) par la recherche des nids (Liste Rouge mammiferes). Résultats du test methodologique effectue en 2005. Centre suisse de cartographie de la faune, Neuchatel, rapport Faune Concept (non publié), 6 p. + annexes. BLANT, M. 2007. Recherche extensive de la Souris des moissons (Micromys minutus): distri- bution spatiale des nids, implication pour la gestion. Fondation des Marais de Damphreux, Porrentruy, rapport Faune Concept (non publié), 5 p. + annexes. BLANT, M. & MARCHESI, P. 2009. Localisation des populations résiduelles de la souris des moissons (Micromys minutus). Service des foréts, de la faune et de la nature, Neuchatel, rapport Faune Concept (non publié), 8 p. + annexes. BERRET, J. & GOETSCHI, F. 2009. Etude du rat des moissons ”Micromys minutus” dans son milieu naturel et en captivite. Annales de sciences naturelles en Pays jurassien: 77-103. BOHME, W. 1978. Micromys minutus (Pallas, 1778) — Zwergmaus (pp. 290-304). In: NIETHAMMER J. & KRAPP F. (eds), Handbuch der Säugetiere Europas, Bd 1, Nagetiere I, Akademische Verlagsgesellschaft, Wiesbaden. BUSER, M. 1984. Régime alimentaire de la Chouette effraie, 7yto alba, durant les périodes de nidification 1978 et 1979 en Ajoie (Jura suisse). Nos Oiseaux 37 : 392-394. BUTET, A. & PAILLAT, G. 1998. Insectivores et rongeurs de France: Le Rat des moissons — Micromys minutus (Pallas, 1771). Arvicola X (2) : 29-41. DUELLI, P. 1994. Listes rouges des espèces animales menacées de Suisse. OFEFP, Berne, 97 pp. GOETSCHI, F. 2009. Observation de la Souris des moissons, Micromys minutus, aux environs des etangs de Damphreux (Ju). Travail de maturite, Lycée de Porrentruy, 37 p. GEG (GROUPE D'ETUDE ET DE GESTION) 2007. Plan de gestion des réserves de la Grande Carigaie 2007-2011. Rapport non publié, 141 pp. HAUSSER, J. (ed.) 1995. Mammiferes de la Suisse. Birkhauser, Bale, 501 pp. MARCHESI, P. & DESCOMBES, M. 2008. Confirmation de la présence de la souris des moissons Micromys minutus dans le marais du Grand Bataillard (Commugny). Conservation de la faune du canton de Vaud, St-Sulpice, Rapport Faune Concept (non publié): 9 p. + annexe. MARCHESI, P., BLANT, M. & CAPT, S. 2008. Mammiferes de Suisse — Clés de determination. Fauna Helvetica 21, CSCF & SSBF, Neuchatel: 296 pp. MARCHESI, P. & BLANT, M. 2010. Statut de la souris des moissons Micromys minutus dans le canton de Vaud et recommandations pour sa conservation. Conservation de la faune du canton de Vaud, St-Sulpice, Rapport Faune Concept (non publié): 11 p. + annexe. MICHELAT, D., ROUE, S. & PEPIN, D. 2005. Les mammiferes de la montagne jurassienne. Neo- Typo, Besançon, 183 pp. 500 M. BLANT ET AL. MITCHELL-JONES, A. J. et al. 1999. The Atlas of European Mammals. Societas Europaea Mammalogica, T. & A. D. Poyser publ., London, 484 pp. MOORE, N. P., ASKEW, N. & BISHOP, J. D. 2003. Small mammals in new farm woodlands. Mammal Review 33 : 101-104. QUERE, J.-P. & LOUARN, H. 2011. Les rongeurs de France. Faunistique et biologie. Guide Pratique, Quae, Versailles, 311 pp. RAHM, U. 1995. Micromys minutus (pp. 263-267). In: HAUSSER J. (ed.), Mammiferes de la Suisse. Répartition, biologie, écologie. Société Suisse de Biologie de la Faune, Mémoires de l'Académie Suisse des Sciences naturelles, Vol. 103. Birkhduser, Bâle. SCHMIDLIN, A. 2011. Etude d'une petite population de Souris des moissons, Micromys minutus, dans la Vallée de Delémont et a Damphreux. Travail de maturite, Lycée de Porrentruy, 32 p. REVUE SUISSE DE ZOOLOGIE 119 (4): 501-528; décembre 2012 Civizelotes new genus, and other new or little known Zelotinae (Araneae, Gnaphosidae) Antoine SENGLET Route de Begnins, 19, CH-1267 Vich, Switzerland. E-mail: a.senglet@bluewin.ch Civizelotes new genus, and other new or little known Zelotinae (Ara- neae, Gnaphosidae). - Civizelotes, a new genus of gnaphosid spiders, is established and divided in four species groups. Notes are given on the mating mechanisms of C. medianus (Denis) in the C. civicus-group. First descriptions are given of the male of Z. argoliensis (C. L. Koch, 1839), of the female of Drassyllus villicoides (Giltay, 1932) comb. n., the female of Civizelotes dentatidens (Simon, 1914) comb. n., the female of C. medianus (Denis, 1935) comb. n. C. medianoides sp. n., C. ibericus sp. n., and Heser hispanus sp. n. Transferred to different genera are: C. civicus (Simon, 1878) comb. n., C. caucasius (L. Koch, 1866) comb. n., C. solstitialis (Levy, 1998) comb. n., Heser nilicola (O. P.-Cambridge, 1874) comb. n., A. schmitzi (Kulczynski, 1899) comb. n., H. bernardi (Marinaro, 1967) comb. n., H. bonneti (Marinaro, 1967) comb. n., and H. incisupalpis (Levy, 1998) comb. n. Zelotes balcanicus Deltshev, 2006 is removed from the synonymy of Z. argoliensis. Keywords: Arachnida - taxonomy - zoogeography - mating mechanism. INTRODUCTION A detailed study of the structure of the d pedipalp in Zelotes requires some generic transfers. In Zelotes str. s. the basic structure of the palp is: Strong posterior sclerite of terminal apophysis linked to posterior tegular base, radix or embolus (Fig. 1) and Senglet (2011: figs 13-14, 133). Massive embolar radix with a solid junction to the embolus (Fig. 2). The intercalary sclerite represents an extension of the basal sector of the embolar base, giving a ventral mobile joint to a terminal apophysis with a posterior fixation. All species with a different palp morphology need to be trans- ferred to other genera. Among these species a monophyletic group is here recognised and separated into the new genus Civizelotes gen. n. MATERIAL AND METHODS Except for the 2 syntypes of Zelotes argoliensis and the syntype of Z. medianus (Denis) = Civizelotes dentatidens, all material was collected by myself. Measurements are in millimetres. Vulvae were examined in lactic acid on an excavated microscopic slide. Holotypes and paratypes of the species described in here are deposited in the Natural History Museum of Geneva (MHNG); the other material, if not indicated Manuscript accepted 26.07.2012 502 A. SENGLET otherwise, remains in my private collection. For details on rearing and cryo-fixing of mated spiders see Senglet (2004: 87). The typical leg spination according to Platnick & Shadab (1983) is: Femora; I, II d110, p001; II, IV d110, p011, r011; patella III r010; tibiae: HI pl11, v222, r011; IV pl11, v222, rl11; metatarsi: I, II v200; II p122, v221, r112; IV p122, v220, r122. Only differences from this pattern are given in the text. AME, ALE, PME, PLE and MOQ refer to anterior median, anterior lateral, posterior median, posterior lateral eyes and to the median ocular quadrangle (with eyes included). Terminology of genital structures follows Senglet (2004: 88-90) and Senglet (2011: 514). TAXONOMY AND FAUNISTIC DATA Genus Zelotes Gistel, 1848 TYPE SPECIES: Melanophora subterranea C. L. Koch 1833, subsequent designation. Zelotes fulvopilosus-subgroup of Z. thorelli-group (Senglet, 2011: 540) Zelotes argoliensis (C. L. Koch, 1839) Figs 3-10 Melanophora argoliensis C. L. Koch, 1839: 72, fig. 483 (description of 2). — L. Koch, 1866: 174, pl.7, fig. 113 (2). TYPE MATERIAL: BM1915.3.5.5759, Coll. Koch, Natural History Museum (London); 2 9 syntypes from Greece; received on loan through the courtesy of Mrs Janet Beccaloni. OTHER MATERIAL EXAMINED: GREECE, Peloponnesus, Arcadia, Mt Mainalos, refuge, stones, 37°39'N 22°16'E; 2 Sd, 7 2 (last moults of ¢ 20.07 and 22.08, of 2 2 26.08 and 07.09: 2011); 23.052011; DESCRIPTION: Large dark species. Prosoma dark brown, with black bristles. Opisthosoma black, covered with greyish adpressed hairs and black bristles. Legs dark brown; tarsi feebly lightened. Posterior eye row straight to slightly recurved d : Total length 5.20. Prosoma: 2.25 long, 1.85 wide, 0.90 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.06, ALE 0.12, PME 0.07, PLE 0.12; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.07, ALE-PLE 0.07. MOQ length 0.26, front width 0.40, back width 0.42. Clypeus: 0.10 from AME, 0.08 from ALE. Pedipalp (Figs 3-7): Retrolateral tibial apophysis 2 times dorsal length of tibia. Patella dorsally longer than tibia. Tibia wider than long. Embolar base without connection to embolus, carrying a conical apical projection attached to embolar haema - todocha, and a prolateral guide to embolus. Small hook of median apophysis half concealed by embolus. Very large, one-turn-coiled embolus reaching below proximal quarter of tegulum length. Posterior sclerite of terminal apophysis attached to base of embolar radix (Fig. 3), the latter linked through an intermediary junction to posterior tegular base. Tarsi I, II scopulate, metatarsi I, II only in apical 2/3. Leg spination: Tibiae III, r112; metatarsi III, VI v221. Scutum occupying 1/3 of opisthosoma length. 2: Total length 6.26. Prosoma: 2.5 long, 2.00 wide, 1.08 wide at level of pos - terior eyes. Eye sizes and interdistances: AME 0.07, ALE 0.12, PME 0.07, PLE 0.07; AME-AME 0.03, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.04, ALE-PLE 0.07. MOQ length 0.24, front width 0.36, back width 0.46. Clypeus: 0.10 from AME, 0.08 NEW OR LITTLE KNOWN ZELOTINAE 503 embolar "haematodocha emb. base > projection |“? > radix : radix L N T. apophysis NN # post. sclerite | IN , | VE rina ho post. 33 N, tegular base . term. apoph. ) D EIG T. apophysis post. sclerite 2 Ri BER 3 median apophysis en 1 post. i | er an LE ‘ Le Spermophor intercalary sclerite T. apophysis post. sclerite emb. base projection FIGS 1-10 (1-2) Zelotes apricorum (L. Koch), left male palp, cymbium removed. (1) Ventral view. (2) Dorsal view. (3-10) Z. argoliensis (C. L. Koch). (3-7) Left male palp. (3) Cleared, retrolateral view. (4) Prolateral view. (5) Ventral view. (6) Retrolateral view. (7) Cymbium removed, apical view. (8) Epigynum. (9) Vulva, ventral view. (10) Id., dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. 504 A. SENGLET from ALE. Epigynum (Fig. 8). Vulva (Figs 9-10): Copulatory duct loosely coiled around the straight median duct. Tarsi and metatarsi I and II entirely scopulate. Leg spi- nation: Tibiae III, r011; metatarsı II, v220 ; HI, VI v221. Zelotes petrensis-group (Senglet, 2004: 94, 111) Zelotes balcanicus Deltshev, 2006, removed from the synonymy of Z. argoliensis Figs 11-12 Zelotes balcanicus Deltshev, in Deltshev et al., 2006: 711, figs 2-11 (description of & and ©). Zelotes baram Levy, 2009: 31, figs 67-70 (description of & and ?) syn. n. Zelotes argoliensis. — Chatzaki, 2010: 54, figs 25-30 (2, synonymy of Z. balcanicus) (misiden - tification). MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, Loutra Apollonias, litter, 40°39'N 23°24'E; 1 9; 09.09.2004. — South of Stanos /Arnaia, Pinus litter, 40°32'N 23°35'E; 1 4,1 2; 09.09.2004. — Epiros, Thesprotia, Arghyrotopos, litter, 39°25'N 20°19'E; 1 & (last moult 05.09.2004); 31.08.2004. — Sterea Hellas, Phthiotidas, south-west of Theologos, Pinus litter, 38°39'N 23°11'E; 2 2 (last moults 07. and 18.09.2011); 17.06.2011. — Peloponnesus, Messenia, Kiparissia, leaf litter, 37°15'N 21°40'E; 1 2; 03.06.2011. — Laconia, East of Lira, Pinus litter, 36°39'N 22°58'E; 1 d, 1 9 (with palpus in microvial); 03.10.2004. — Argolida, above Platani, 700 m, 37°48'N 22°30'E; 3 4, 3 2 (last moults of d 22.08.-02.09, of 2 02.08.- 22.09.2011); 21.05.2011. DESCRIPTION: See Deltshev et al. (2006: 711). REMARKS: Contrary to its original placement in the Z. subterraneus-group, Z. balcanicus is a member of the Z. petrensis-group (Senglet, 2004: 94, 111), as can be seen from the structure of the embolar base. A strong mesal apophysis and a posterior sickle-shaped lamina on the embolar base (Figs 11-12) are present. A week direct sinuous link to the embolus is another basic character of the group. Lateral additional pockets are present on the epigynum. The absence of a membranous or sclerotized embolar projection is an indication that this species belongs to a distinct sub-group. Genus Drassyllus Chamberlin, 1922 TYPE SPECIES: Drassyllus fallens Chamberlin, 1922, original designation. Drassyllus villicoides (Giltay, 1932) comb. n. Figs 13-19 Zelotes villicoides Giltay, 1932: 28, fig. 18 (description of 6). — Giltay, 1933: 6, pl. 3, fig. 4(¢). MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, east of Chortiatis, stones in pasture, 40°37'N 23°07'E; 1 & (last moult 02.06.2005); 07.09.2004. — West of Aghios Vassilios, 40°40'N 23°05'E; 1 4,2 ? (last moults of 4 16.06.2005, of 2 08 and 14.05.2005); 08.09.2004. — Aghios Vassilios, 40°40'N 23°07'E; 1 ®; 13.05.1968. — Loutra Apollonias, litter, 40°39'N 23°24'E; 1 6, 1 9 (last moults of d 23.05.2005, of 2 09.07.2005); 09.09.2004. — Sterea Hellas, Phthiotidas, west of Malesina, 38°37'N 23°13'E; 6 4, 4 © (last moults of 4 16.04.-15.05.2005, of 9 16.04.-28.05.2005); 25.09.2004. — Near Malesina, 38°37'N 23°13'E; 1 9; 21.05.1998. — Peloponnesus, Laconia, south-west of Mavrovouni, litter on sand, 36°43'N 22°32'E; 1 ®; 26.05.2011. — Argolida, east of Ligourion, Lentiscus litter, 37°38'N 23°04'E; 1 4, 3 © (last moults of 4 24.04.2005, of 2 17.04.-24.05.2005); 01.10.2004. DESCRIPTION: Prosoma dark brown, with tiny hairs. Opisthosoma black, with short adpressed hairs. Legs dark brown except for tawny-yellow tarsi and metatarsi. Tarsi and metatarsi I, II scopulate. Average size about 3/4 that of D. praeficus. 3: Total length 5.00. Prosoma: 1.95 long, 1.62 wide, 0.77 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.07, ALE 0.08, PME 0.08, PLE 0.07; NEW OR LITTLE KNOWN ZELOTINAE 505 embolar radix — ___, e posterior lamina mesal apoph. embolus term. N mesal apophysis_ 7 term. apophysis intercal.sclerite Fics 11-12 Zelotes balcanicus Deltshev. (11) Left male palp, cymbium removed, anterior-apical view. (12) Id., dorsal view. Scale 0.2 mm. AME-AME 0.03, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.04, ALE-PLE 0.06. MOQ length 0.20, front width 0.37, back width 0.43. Clypeus: 0.11 from AME, 0.07 from ALE. Retrolateral tibial apophysis bent dorsally, shorter than dorsal length of tibia. Pedipalp (Figs 13-16): Tibia short; its dorsal length about equal to lateral width. Patella dorsally longer than tibia. Scutum occupying 1/3 of opisthosoma length. Close to D. praeficus, differing by the thin narrow triangular terminal apophysis and the slender embolar radix apophysis (Figs 14, 16); in D. praeficus (Fig. 20) the latter is stronger and more cone-shaped. 2: Total length 5.66. Prosoma: 2.10 long, 1.58 wide, 0.80 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.07, ALE 0.10, PME 0.08, PLE 0.08; AME-AME 0.08, AME-ALE 0.01, PME-PME 0.02, PME-PLE 0.05, ALE-PLE 0.05. MOQ length 0.24, front width 0.37, back width 0.43. Clypeus: 0.11 from AME, 0.06 from ALE. Epigynum and vulva (Figs 17-19). Anterior anchoring pockets at a distance from the anterior epigynal margins; in D. praeficus (Fig. 21) pocket pressed to the external margin of the epigynum. Modest widening of the lateral fold; median fold reaching the spermatheque in median-anterior position, there covered by cuticle (Figs 18-19); in D. praeficus (Fig. 21) posterior strong widening of the lateral fold; median fold reaching the spermatheque in uncovered lateral-anterior position. BIOLOGY: Adult males can be collected from April to June. Drassyllus praeficus (L. Koch, 1866) Figs 20-21 Melanophora praeficus L. Koch, 1866: 155, pl. 6, figs 97-99 (description of & and ?). For previous synonymy see Platnick (2012). MATERIAL EXAMINED: SWITZERLAND, Vaud, Vich /Nyon; 1 3, 1 9; 01.-30.05.1996. — SPAIN: Lac de Sanabria, 42°06'N 06°41'W and 42°07'N 06°44'W; 1 9; 26.08.1971. — Avila, Road to parking Gredos /Hoyo del Espino, 1470 m, 40°18'N 05°12'W; 1 d (last moult 28.12.2009); 18.06.2009. — Andalusia, Granada, Prado de Zangarrilla (S. Nevada), stones, pas- ture, 2000 m., 37°07'N 03°26'W; 1 2; 29.05.2002. — FRANCE, Provence / Cote d'Azur, Var, Montauroux; 1 9; 07.06.1971. — Languedoc, Pyrénées Orientales, Prats de Mollo, 700 m., 42°24'N 02°25'E; 3 2; 29.06.2001. — Latour de Carol, 42°28'N 01°53'E; 1 2; 06.07.2001. — Above Mas Franco / Enveigt, 42°30'N 01°53'E; 1 2; 06.07.2001. - ANDORRA, La Rabassa, 2000 m, 42°26'N 01°32'E; 2 2 (last moult of 1 9 24.02.2002); 08.07.2001. — West of St Julia 506 A. SENGLET T. apophysis embolar radix apophysis ee embolar base N 3 “embolar base projection 16 Lee a LL “„nunmumnnmn FIGS 13-19 Drassyllus villicoides (Giltay). (13-16) Left male palp. (13) Prolateral view. (14) Ventral view. (15) Retrolateral view. (16) Cymbium discarded, retrolateral view. (17) Epigynum. (18-19) Two vulvae, dorsal view, same population. NEW OR LITTLE KNOWN ZELOTINAE 507 ee e Sa mm FIGS 20-21 D. praeficus. (20) Left male palp, cymbium removed, retrolateral view. (21) Left side of vulva, ventral view. Bold lines indicate epigynal folds. Scale 0.2 mm. de Loria, schist litter, 42°27'N 01°28'E; 3 2 (last moults of 2 2 05 and 08.06.2004); 27.05.2004. — GREECE, Macedonia, Serres, above Oreini, 1200 m, stones, 41°14'N 23°35'E; 1 9; 30.06.2011. — Thessaloniki, west of Aghios Vassilios, 40°40'N 23°05'E; 1 6, 1 £ (last moults of 3 11.06, of 2 09.06.2005); 08.09.2004. — Same; 4 2; 31.05.2008. — East of Aghios Vassilios, litter, 40°40'N 23°07'E; 2 d (last moults 17 and 20.06.2005); 08.09.2004. — Same; 2 8,4 9; 13.05.1968. — Same; 1 2; 28.05.1982. — Same, 40°40'N 23°07'E; 4 9; 14.06.1998. — Thessaloniki, 40°43'N 22°58'E; 1 ®; 13.-18.05.1968. — Kastoria, north of Kastoria (pass), 40°35'N 21°18'E; 1 2; 20.06.1998. — Peloponnesus, Achaia, Kato Alissos, 38°09'N 21°35'E; 3 2 (last moult of 1 £ 10.05.1998); 29.04.1998. — Arla, 38°04'N 21°36'E; 1 2 (last moult of 9 05.05.1998); 30.04.1998. — Argolida, east of Myli / Nauplie, leaf litter, 37°35'N 22°44'E; 1 9 (last moult 03.06.2005); 01.10.2004. — Cyclades, Paros, Sta Maria /Naoussa, (Salicornia), 37°08'N 25°17'E; 1 2; 02.06.1998. — Crete, Chania, Omalos haut plateau, stones, 35°20'N 23°53'E; 2 6,2 © (last moults of d 01.01 and 04.01, of 2 31.12.1999); 07.10.1999. — Episkopi, litter, soil crevices, 35°30'N 23°46'E; 1 @ (last moult 14.01.2000); 10.10.1999. — Rethimnon, Koxare /Spili, irigated vegetation, 35°14'N 24°28'E; 1 6,2 2 (last moults of & 30.12.1999, of 2 08.01.2000); 03.10.1999. — Choumerion / Pérama, 35°21'N 24°44'E; 1 6d (last moult 04.01.2000); 05.10.1999. — Missiria /Rethimnon, leaf litter, soil crevices, 35°22'N 24°31'E; 2 3; 06.10.1999. — Lassithi, Kato Metochi Lassithiou, 35°11'N 25°26'E; 2 d, 2 2 (with palpus in microvial, last moults of & 25 and 30.12.1999, of £ 01.01 and 06.01.2000); 25.09.1999. — ROMANIA, Dobrogea, Constanca, Baneasa; 1 d (with palpus in microvial); 30.05.1972. — Adamclisi; 1 8; 31.05.1972. — Mangalia; 2 2; 02.06.1972. — Techirghiol; 1 2; 03.06.1972. — BULGARIA, Razgrad, Hlebarovo; 1 2; 08.07.1972. — Varna, Slatni Pjasaci; 1 2; 11.07.1972. — Burgas, Obzor; 3 2; 14.07.1972. — Karnobat; 2 2; 15.07.1972. — Plovdiv, Backovo; 1 9; 21.07.1972. — IRAN, Esfahan, Lorestan, Dizgaran, 33°44'N 46°59'E; 3 2; 16.05.1974. — Fars, near Ghaderabad, 30°22'N 53°18'E; 1 2; 11.06.1974. — Khorasan, east of Chaman Bid, 37°26'N 56°37'E; 1 2; 14.07.1974. DESCRIPTION: See Grimm (1985: 267, figs. 317, 319-320). Drassyllus lutetianus (L. Koch, 1866) Melanophora lutetianus L. Koch, 1866: 157, pl. 6, fig. 100 (description of © ). For previous synonymy see Platnick (2012). MATERIAL EXAMINED: GREECE, Peloponnesus, Argolida, East of Myli /Nauplie, leaf litter, 37°35'N 22°44'E; 2 © (last moults 09 and 29.05.2005); 01.10.2004. - FRANCE, Corse, Sud Corse, Solenzara: 1 2; 24.05.1971. - ITALIA, Sardinia, Sassari, Platamona / Porto Torres, damp leaf litter; 2 2; 25.05.1999. — Ozieri, damp vegetation; 2 2; 28.05.1999. - ROMANIA, 508 A. SENGLET Dobrogea, Tulcea, Badabag; 1 2; 05.06.1972. — Murighiol; 1 2; 06.06.1972. — Constan.a, Baneasa; 1 6; 30.05.1972. - BULGARIA, Razgrad, Hlebarovo; 1 2; 08.07.1972. DESCRIPTION: See Grimm (1985: 264, figs 8b, 315, 325-326). Drassyllus pusillus (C. L. Koch, 1833) Melanophora pusillus C. L. Koch, 1833: 120, pl. 22. For previous synonymy see Platnick (2012). MATERIAL EXAMINED: SWITZERLAND, Vaud, Vich /Nyon, 46°26'N 06°15'E; 1 & (with palpus in microvial); 02.05.2004. — Vich /Nyon; 2 2 (with vulva in microvial); 01.05.1996. — GREECE, Macedonia, Serres, Mt Vrondos (Laïlia) 1780 m, stones, 41°16'N 23°36'E; 1 ©; 29.06.2011. — SPAIN, Castilla / Leon, Avila, Parking Gredos / Hoyo del Espino, 1780 m, pasture) 40°16'N 05°14'W; 2 2; 18.06.2009. — Nueva Castilla / La Mancha, Cuenca, Santa Cruz de Moya, 39°57'N 01°13'W; 1 2; 24.06.1971. — Palomera / Cuenca, 40°04'N 02°03'W; 3 &; 27.06.1971. — Andalusia, Granada, Puerto de la Ragua, 2000 m., 37°07'N 03°02'W; 1 9; 17.07.1971. — IRAN, Azerbaidjan, Eastern Azerbaidjan, West of Bostanabad, 1900 m, 37°55'N 46°42'E; 1 3; 07.06.1975. DESCRIPTION: See Grimm (1985: 274, figs 316, 327-328). Genus Civizelotes gen. n. TYPE SPECIES: Prosthesima civica Simon, 1878, designated herewith. ETYMOLOGY: The first four letters of the type species are added as a prefix to Zelotes. DIAGNOSIS: Embolar radix originating from the posterior margin of the tegu- lum, terminating at level of embolus in a large plate of bowl (Figs 47, 53-54, 62). Intercalary sclerite absent. Terminal apophysis without posterior sclerite, or reduced. Long embolus type. Leg spination: Femora IV p001, r001; metatarsi II v220. REMARKS: Zelotes apricorum (Figs 1-2) shows both the embolar radix and the posterior sclerite of the terminal apophysis in Zelotes str. s. KEY TO THE CIVIZELOTES-GROUPS la Embolar base long, with a retrolateral loop (Figs 31, 44). Epigynal plate Narrowed Wits Centte . ir C. civicus-group Ib Embolar base vestigial. Epigynal plate in anterior sector (Fig. 66)..... re ten a arn, be ee eens i C. gracilis-group Ic Embolar base straight (Figs 53, 68). Epigynal plate long or a small au Us IN-anlenor position: Bet teu tet eee tea ee 2 2a Terminal apophysis present; embolar base with a ventral expansion ter- minating in a large hook (Figs 50-52). Epigynal plate long and triangular (Rie: 2 og goes ee oss re ae ae ee es C. dentatidens 2b No terminal apophysis; embolar base without ventral expansion. Epi - gynal plate a small atrium in anterior position ........... C. caucasius-group Civizelotes civicus-group DEFINITION: Light coloured spiders. Male palp: A terminal apophysis present but without posterior sclerite. Strongly developed transverse embolar base forming a NEW OR LITTLE KNOWN ZELOTINAE 509 retrolateral loop (Figs 44, 47). Embolar radix visible prolaterally, with a large dorsal- apical bowl (Figs 31, 43). Epigynal plate narrowed in its centre (Figs 33, 35, 48). PME larger than PLE; posterior eye-row procurved; its eyes separated by less than 40% of their diameter. Of particular interest in this group is the use of the two folds of the em- bolar base to hold the bulbus on the epigynum (Figs 22, 26-27). SPECIES INCLUDED: C. civicus (Simon, 1878) comb. n., C. medianus (Denis, 1935) comb. n., and C. medianoides sp. n. MATING MECHANISM: Civizelotes medianus inserts the left pedipalp into the left side of the epigynum (Figs 22-29). The retrolateral tibial apophysis is inserted into the right anterior anchoring pocket. The erected embolar base (Figs 22-23) is forming two lobes, the apical and retrolateral folds. The wide retrolateral fold is inserted in the left anterior anchoring pocket of the epigynum, and the other below the left lateral epigynal fold (Figs 26-27). KEY TO THE SPECIES OF THE CIVIZELOTES CIVICUS-GROUP la Apical margin of embolar base with distal tooth (Figs 44, 47)........ ee Ee MN à ER 4. C. medianoides sp. n. Ib Apical margin of embolar base convex (Figs 31, 36)................... 2 2a Prolateral-apical part of embolar radix strongly developed (Fig. 31)... eR EUER EEE IE EEE pea C. medianus 2b Prolateral-apical part of embolar radix feebly developed (Fig. 36) . . C. civicus 3a Lateral anterior epigynal folds straight, with a posterior bend (Figs 33, 48). . 4 3b Anterior sector of epigynal folds curved, with visible median ducts (E ARRE ete aie TE C. civicus 4a Lateral anterior epigynal folds nearly parallel (Fig. 33)......... C. medianus 4b Lateral anterior epigynal folds diverging (Fig. 48)...... C. medianoides sp. n. Civizelotes medianus (Denis, 1935) comb. n. Figs 22-34 Zelotes medianus Denis, 1935: 118, figs 4-5 (description of d, not ©, fig. 3 = Z. dentatidens). — Jézéquel, 1962: 606, fig. 31 (2). — Soyer, 1967: 275, figs 5-7 (2 may be the 2 of Z. den- tatidens). MATERIAL EXAMINED: SPAIN, Catalonia, Gerona, Val de Blanya, (4 km south of Capsacosta pass), 42°14'N 02°23'E; 3 Sd, 4 © (last moults of 2 d 06 and 27.06.2004, of 9 01.06.-24.07.2004); 25.05.2004. — Barcelona, Alto de los Bruch, 41°36'N 01°45'E; 1 6; 13.06.1971. — Levant / Murcia, Murcia, Sierra de Espufia / Lorca, evergreen oak leaves, 37°55'N 01°40'W; 4 8,3 % (last moult of 1 2 10.06.2002); 18.05.2002. — Nueva Castilla / La Mancha, Albacete, La Gineta (rio Jùcar), 39°11'N 01°58'W; 1 9; 28.06.1971. — Andalusia, Granada, Puebla de Don Fadrique, Pinus & rosemary litter, 38°00'N 02°27'W; 1 9; 19.05.2002. — La Vidriera /Puerto del Pinar, 38°03'N 02°34'W; 4 4,2 £ (last moults of 2 & 23 and 27.05.2002, of 2 22. and 31.05.2002); 20.05.2002. — La Calahora, pine forest, 37°10'N 03°03'W; 2 d; 23.05.2002. — Road puerto de la Ragua, pine forest, 37°09'N 03°03'W; 2 4,1 9; 24.05.2002. — above Capileira, 1700 m, 36°58'N 03°21'W; 1 8; 27.05.2002. — Collado del Muerto (S. Nevada), 1450 m, 37°08'N 03°28'W; 1 9,1 £ (last moults of d 31.05.2002, 9 31.05.2002); 29.05.2002. — Jaen, Sierra de Cazorla (Fuente del Oso), 37°55'N 02°56'W; 2 3d, 1 2 (last moult of 9 31.05.2002); 21.05.2002. — Same; 1 9 ; 23.07.1971. — Sierra de Cazorla (above Parador), stones, Pinus, 37°55'N 02°57'W; 1 4, 2 2 (last moult of d 25.05.2002, 2 23 and 30.05.2002); 510 A. SENGLET “a subtegulum tibial apophysis embolar radix embolar base D folds embolus embolar base apical fold term. bowl of embolar radix retrolateral tibial apophysis embolar base retrolateral fold hooked into ant. “~~ lateral pocket je". anterior epig. margin zu. .. oe? embolar base apical fold hooked below anterior «7 lateral epigynal fold term. bowl of embolar radix inserted embolus 27 FIGS 22-27 Civizelotes medianus (Denis), left male palp in copula. (22-23) Extracted male palp in anterior- lateral view. (24-25) Id., in apical view. (26) Epigynum, after extraction of palpal organ. (27) Id., palpal organ inserted. Bold lines indicate epigynal folds. Scale 0.2 mm. NEW OR LITTLE KNOWN ZELOTINAE 511 embolar radix terminal bowl FIGS 28-34 Civizelotes medianus (Denis), (28-29) In copula. (28) Posterior view. (29) Lateral view. (30) Palpal organ, retrolateral view. (31) Left male palp, ventral view. (32) Id., retrolateral view. (33) Epigynum. (34) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. 22.05.2002. — Sierra de Cazorla (Linarejas), 37°55'N 02°55'W; 1 3; 22.05.2002. — Canada de las Hazadillas, Pinus, 37°39'N 03°43'W; 2 3; 30.05.2002. — Sevilla, east of Cazalla de la Sierra, deep leaf litter, 37°57'N 05°45'W; 2 Sd, 3 @ (last moult of 1 2 07.06.2009); 04.06.2009. — FRANCE, Languedoc, Aude, Bedos pass / Mouthoumet, 42°58'N 02°34'E; 2 2 (last moult of 1 2 01.06.2004); 30.05.2004. — Eastern Pyrenees, Les Abeilles / Banyuls, 42°28'N 03°04'E; Sil A. SENGLET 1 4,1 2; 28.06.2001. — Cerbère, stones in Cistus, 42°27'N 03°09'E; 3 &, 1 £ (last moult of 1 & 05.06.2004); 22.05.2004. - ANDORRA, St Julia de Loria, oak litter, 42°27'N 01°29'E; 1 9 (last moult of 2 01.07.2004); 25.05.2004. — West of St Julia de Loria, litter, vegetation, 42°27'N 01°28'E; 3 3,1 © (last moult of 2 14.06.2004); 27.05.2004. DESCRIPTION: Prosoma tawny, with short adpressed hair and sparse bristles. Opisthosoma blackish, with short adpressed hairs and short to medium-long bristles. Legs dark tawny to brownish. Leg spination: Femora IV p001, r001; metatarsi II V220. Tarsi I, II scopulate. 3 d from the same locality: Total length 4.12 (3.66). Prosoma: 1.80 (1.53) long, 1.40 (1.21) wide, 0.63 (0.58) wide at level of posterior eyes. Eye sizes and inter- distances: AME 0.07, ALE 0.10, PME 0.07, PLE 0.09; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.03, ALE-PLE 0, 05. MOQ length 0.23, front width 0.33, back width 0.37. Clypeus: 0.07 from AME, 0.04 from ALE. Pedipalp (Figs 30-32): Retrolateral tibial apophysis shorter than tibial dorsal length. Patella dorsally longer than tibia. Superior margin of embolar base rounded in ventral view. Scutum occupying 1/4 of opisthosoma length. 2 2 from the same locality: Total length 4.90 (4.10). Prosoma: 1.92 (1.6) long, 1.48 (1.2) wide, 0.72 (0.64) wide at level of posterior eyes. Eye sizes and interdis - tances: AME 0.08, ALE 0.11, PME 0.11, PLE 0.08; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.03, ALE-PLE 0.04. MOQ length 0.17, front width 0.35, back width 0.38. Clypeus: 0.06 from AME, 0.08 from ALE. Epigynum and vulva (Figs 33-34). Anterior sector of epigynal folds diverging feebly, posterior sector shorter than in Z. medianus. Civizelotes civicus (Simon, 1878) comb. n. Figs 35-39 Prosthesima civica Simon, 1878: 80, pl. 14, fig. 20 (description of d and ©). For previous synonymy see Platnick (2012). MATERIAL EXAMINED: SPAIN, Catalonia, Barcelona, Alto de los Bruch, 41°36'N 01°45'E; 3 gd; 13.06.1971. — Gelida, 41°27'N 01°51'E; 5 2; 14.06.1971. — Levant / Murcia, Valencia, Ayora, 39°03'N 01°03'W; 1 &; 20.06.1971. — Montroy, 39°20'N 00°35'W; 1 9; 22.06.1971. — Requena - Chera, 39°36'N 00°57'W and 39°32'N 01°00'W; 2 ¢, 1 2; 23.06.1971. — Alicante, Elda, 38°30'N 00°47'W; 2 2; 19.06.1971. — Murcia, Bullas, below vegetation, 38°02'N 01°39'W; 2 3; 18.05.2002. — Bullas, 38°02'N 01°39'W; 1 3, 5 9; 04.07.1971. — Caravaca, 38°05'N 01°52'W; 1 2; 05.07.1971. — Lac de Sanabria, 42°07'N 06°44'W and 42°06'N 06°41'W; 1 9; 26.08.1971. — Valladolid, Pefiafiel, 41°35'N 04°08'W; 1 6,1 2; 23.06.2002. — Salamanca, Puente Alagön /Sequeros, 40°30'N 05°57'W; 1 2; 13.08.1971. — Nueva Castilla / La Mancha, Madrid, Navalcarnero, 40°18'N 03°56'W; 1 &; 12.06.1969. — Aldea del Fresno (rio Alberche), 40°19'N 04°13'W; 1 3; 13.06.1969. — Cuenca, Santa Cruz de Moya, 39°57'N 01°13'W; 1 2; 24.06.1971. — Palomera /Cuenca, 40°04'N 02°03'W; 1 8; 27.06.1971. — Toledo, Escalona del Alberche, 40°10'N 04°24'W; 4 d, 2 9; 14.06.1969. — Urda /Consuegra, 39°25'N 03°42'W; 1 2; 12.08.1969. — Albacete, La Gineta (rio Jucar), fine leaf litter, 39°10'N 01°58'W; 1 4,5 2 (last moults of 4 2 20.-31.05.2002); 16.05.2002. — La Gineta (rio Jùcar), 39°11'N 01°58'W; 2 2; 28.06.1971. — Hellin, 38°29'N 01°37'W; 1 9; 29.06.1971. — Ciudad Real, Ruidera, 38°56'N 02°51'W; 5 2; 07.08.1969. — Ojos del Guadiana /Daimiel, 39°08'N 03°34'W; 1 2; 13.08.1969. — Estremadura, Caceres, Jarandilla (rio Tietar), 40°01'N 05°37'W; 3 6, 1 9; 16.06.1969. — Badajoz, south of Monesterio, stones in vegetation, 38°03'N 06°14'W; 3 6,3 9; 05.06.2009. — South of Venta del Culebrin / Monesterio, 37°58'N 06°14'W; 1 9; 19.06.1969. — Rio Sillo (Higuera la Real), 38°06'N 06°41'W; 1 2; 14.06.2009. — Andalusia, Granada, Prado de Zangarrilla (S. Nevada), stones, pasture, 2000 m, 37°07'N 03°26'W; 3 8,5 © (last moults of 9 27.05.-02.06.2002); 29.05.2002. — Puebla de Don Fadrique, 1000-1200 m, 38°00'N 02°27'E; 2 NEW OR LITTLE KNOWN ZELOTINAE S13 embolus term. apoph. embolus radix bowl embolar radix embolar base FIGS 35-39 Civizelotes civicus (Simon). (35) Epigynum. (36) Left male palp, ventral view. (37) Id., slightly expanded, ventral-apical view. (38) Id., cymbium discarded, retrolateral view. (39) Id., prolateral view. Bold lines indicate epigynal folds. Scale 0.2 mm. 3, 6 2; 06.07.1971. — Pampaneira /Orgiva, 1200 m, 36°57'N 03°22'W; 2 9; 10.07.1971. — Caratauna /Orgiva, 36°56'N 03°25'W; 1 4, 6 2 (with palpus in microvial); 13.07.1971. — Capileira, 1500 m, 36°58'N 03°21'W; 1 4,8 2; 10.07.1971. — Trevélez, 37°00'N 03°16'W; 1 9; 14.07.1971. — Huéneja, 37°11'N 02°56'W; 1 3, 6 &; 16.07.1971. — Lacalahora (Ferreira), 37°10'N 03°03'W; 4 2; 17.07.1971. — Guadix (Paulenca de G.), 37°19'N 03°09'W; 6 ®; 18.07.1971. — Jaen, Sierra de Cazorla (Fuente del Oso), 37°55'N 02°56'W; 5 &; 21.05.2002. — Sierra de Cazorla (Fuente del Oso), 37°55'N 02°56'W; 1 2; 23.07.1971. — Sierra de Cazorla (El Sagreo), 37°57'N 03°57'W; 1 2; 22.07.1971. — Sierra de Cazorla (Guadalquivir), 37°58'N 03°55'W; 2 2; 24.07.1971. — Cañada de las Hazadillas, Pinus, 37°39'N 03°43'W; 4 4,1 9 (last moult of 2 02.06.2002); 30.05.2002. — La Aliseda (Sierra de Andujar), 38°22'N 03°49'W; 2 gd (last moult of 1 d 02.06.2009); 29.05.2009. — Hinojares / Cazorla, 37°43'N 02°59'W; 1 9; 19.07.1971. — Cordoba, Los Villares / Cordoba, Eucalyptus litter, 37°58'N 04°49'W; 3 $, 5 © (last moults of 2 2 12. and 19.06.2009); 31.05.2009. — Palma del Rio (Retortillo dam), leaf litter, 37°51'N 05°22'W; 1 3; 03.06.2009. — Malaga, Ronda, 36°46'N 05°13'W; 1 9; 21.07.1969. — Sevilla, La Minilla dam / El Ronquillo, 37°40'N 06°11'W; 2 2; 20.06.1969. — Alcala del Rio, 37°31'N 05°59'W; 1 9; 22.06.1969. — Castilblanco (road C.433), 37°41'N 05°59'W; 1 9: 514 A. SENGLET 23.06.1969. — El Real de la Jara - El Pintado, 37°57'N 06°03'W; 3 2; 03.07.1969. — Huelva, East of Aracena, 37°52'N 06°31'W; 1 3,5 2; 07.06.2009. — Puerto Gil /Aracena, cork-oak leaf litter, 37°53'N 06°29'W; 1 4,2 2; 07.06.2009. — Alajar /Aracena, 37°52'N 06°40'W; 2 8,10 2 (with vulva in microvial); 07.07.1969. — Alajar 2 /Aracena, 37°53'N 06°40'W; 1 3d, 1 2; 09.06.2009. — Fuenteheridos, deep leaf litter, 37°54'N 06°40'W; 2 8,4 2; 12.06.2009. — Molino del rio Alajar, deep leaf litter, 37°52'N 06°40'W; 1 2; 13.06.2009. — Zalamea la Real, 37°41'N 06°39'W; 2 2; 10.07.1969. - PORTUGAL, Beira Alta, Guarda, Maceira / Fornos de Algodres, (Casal do Monte), 40°44'N 07°24'W; 1 9 ; 09.08.1971. — FRANCE, Provence / Cote d'Azur, Var, Cogolin; 2 2; 08.06.1971. — Languedoc, Eastern Pyrenees, Prats de Mollo, 700 m, 42°24'N 02°25'E; 1 9; 29.06.2001. — Arles sur Tech, 42°27'N 02°36'E; 1 2; 01.07.2001. — Latour de Carol, 42°28'N 01°53'E; 1 2; 06.07.2001. ANDORRA, west of St Julia de Loria, 1400 m, schist litter, 42°27'N 01°28'E; 1 2; 08.07.2001. — Same; 2 gd; 29.06.2002. — Same; 2 d,2 © (last moults of 4 31.05 and 27.06, of 1 2 07.06.2004); 27.05.2004. - MOROCCO, Dayet Aaoua, 33°39'N 05°02'W; 1 2; 27.06.1967. DESCRIPTION: See Grimm (1985: 80, pl. 14, Fig. 20). Civizelotes medianoides sp. n. Figs 40-49 HOLOTYPE: SPAIN, Andalusia, Cordoba, Alajar /Aracena, 37°52'N 06°41'W; 3; 07.06.2002. PARATYPES: Same locality as for holotype; 1 d, 1 £ (with vulva in microvial); 07.06.2002. — Alajar 1 /Aracena, cork oak leaf litter, 37°52'N 06°41'W; 5 3, 1 9 (last moult of 2 12.06.2009); 09.06.2009. — SPAIN, Andalusia, Cordoba, Brena dam, evergreen oaks litter, 37°51'N 05°04'W; 1 3d (last moult 10.06.2002); 01.06.2002. — Huelva, Linares de la Sierra /Aracena, 37°54'N 06°37'W; 1 2; 05.07.1969. — Same; 2 4; 05.06.2002. — Huelva, north of La Nava, schist litter, 38°00'N 06°45'W; 1 8,1 © (last moult of 2 10.08.2002); 09.06.2002. OTHER MATERIAL EXAMINED: SPAIN, Cordoba, Alajar /Aracena, 37°52'N 06°40'W; 1 9; 07.07.1969. — Puerto de Alajar, 820 m, evergreen oak leaf litter, 37°53'N 06°40'W; 3 4, 1 £ (last moult of 2 22.06.2009); 10.06.2009. — Molino del rio Alajar, deep leaf litter, 37°52'N 06°40'W; 1 4,3 9; 13.06.2009. — Palma del Rio (Retortillo dam), leaf litter, 37°51'N 05°22'W; 1 à; 03.06.2009. — Road Posada-Villaviciosa 2, stones in Pinus litter, 37°00'N 05°05'W; 1 ©: 02.06.2009. — Estremadura, Badajoz, Rio Sillo (Higuera la Real), 38°06'N 06°41'W; 2 © (last moult of 1 2 16.06.2009); 14.06.2009. ETYMOLOGY: The species name reflects the close relationship with C. medianus (Denis). DESCRIPTION: Prosoma and legs tawny. Opisthosoma grey, with darker hairs. Tarsi I, II scopulate. Leg spination: Femora IV p001, r001; metatarsi II v220. 3: Total length 4.50. Prosoma: 2.00 long, 1.50 wide, 0.69 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.08, ALE 0.10, PME 0.08, PLE 0.08; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.04, ALE-PLE 0.04. MOQ length 0.27, front width 0.36, back width 0.40. Clypeus: 0.08 from AME, 0.06 from ALE. Pedipalp (43-47): Patella and retrolateral tibial apophysis longer than dorsal length of tibia. In ventral view a small conical apophysis visible on superior ventral margin of embolar base. Scutum occupying 1/5 of opisthosoma length. ?: Total length 4.75. Prosoma: 2.22 long, 1.58 wide, 0.81 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.08, ALE 011, PME 0.10, PLE 0.10; AME-AME 0.04, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.04, ALE-PLE 0.04. MOQ length 0.29, front width 0.40, back width 0.43. Clypeus: 0.08 from AME, 0.06 from ALE. Epigynum (Fig. 48): Median narrowing of lateral folds greater than in C. medianus. Vulva (Fig. 49). NEW OR LITTLE KNOWN ZELOTINAE STO 41 42 embolar radix embolar base Apophysis distal tooth embolar radix embolus apophysis—_, emb. base 43 embolar radix embolus term. b DELL LES - > median apoph. carola embolar base term. apoph. Be FIGS 40-49 Civizelotes medianoides sp. n., male. (40) Carapace. (41) Sternum. (42) Male chelicerae and ocular group, anterior view. (43) Left male palp, prolateral view. (44) Id., ventral view. (45) Id., retrolateral view. (46) Id., cymbium discarded, retrolateral view. (47) Id., apical view. (48) Epigynum. (49) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. 516 A. SENGLET Civizelotes dentatidens (Simon, 1914) comb. n. Figs 50-57 Zelotes dentatidens Simon, 1914: 156, 217, fig. 293 (description of 3). Z. medianus Denis, 1935: 118, fig. 3 (description of à , not figs 4-5 = Z. medianus). — Soyer, 1967: 275, figs 1-4 (not figs 5-7 = Z. medianus). MATERIAL EXAMINED: Museum National d'Histoire Naturelle, Paris, 1 slide PM26 (syn- type of Z. medianus Denis), 1 vulva. — SPAIN, Catalonia, Gerona, Cap de Creus, below rose- mary, Cistus, Erica, 42°19'N 03°17'E; 1 d (last moult 08.06.2004); 24.05.2004. — Estremadura, Caceres, Jarandilla (rio Tietar), 40°01'N 05°37'W; 1 9; 16.06.1969. — Andalusia, Granada, La Calahora, pine forest, 37°10'N 03°03'W; 2 © (with 1/2 vulva in microvial); 23.05.2002. DESCRIPTION of d: Femora, patellae and tibiae of legs blackish. Prosoma orange-yellow. Opisthosoma dorsally black, with black hairs, ventrally grey, with very short hairs. Leg spination: Femora IV p001, r001; metatarsi II v220. Total length 4.3. Prosoma 1.74 long, 1.33 wide, 0.61 wide at level of posterior eyes. Eye size and inter- distances: AME 0.07, ALE 0.85, PME 0.10, PLE 0.08; AME-AME 0.04, AME-ALE, PME-PME 0.10, PME-PLE 0.10, ALE-PLE 0.02. MOQ length 0.24, front width 0.36, back width 0.36. Clypeus 0.08 from AME, 0.05 from ALE. Pedipalp (Figs 50-54): Terminal apophysis present. Embolar radix with a large dorsal-apical bowl (Fig. 54). Patella dorsally longer than tibia. Strong tibial apophysis with a tapering tip and a dorsal tooth. Retrolateral margin of cymbium strongly notched at level of embolus. Embolar base with a strong ventral branch terminating in a flattened, conical, retro- laterally directed tip. Embolus emerging from the strong short erected dorsal branch with a conical ventral apophysis. Terminal apophysis without posterior sclerite, straight, with the soft tegular cuticle reaching the second third of its length. Radix forming a large concave bowl at level of embolus (Figs 53-54). Scutum brown, occu- pying 1/3 of opisthosoma. DESCRIPTION of 2: Prosoma and legs tawny. Opisthosoma dorsally grey, with darker hairs, ventrally tawny. Total length 4.25. Prosoma: 2.00 long, 1.43 wide, 0.78 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.09, ALE 0.11, PME 0.11, PLE 0.08; AME-AME 0.07, AME-ALE 0.01, PME-PME 0.02, PME-PLE 0.03, ALE-PLE 0.06. MOQ length 0.28, front width 0.40, back width 0.43. Clypeus: 0.11 from AME, 0.06 from ALE. Epigynum (Fig. 55) with scapus-like anterior anchoring pockets protruding ventrally. Vulva (Figs 56-57). Civizelotes gracilis-group I have not seen C. gracilis and C. pygmaeus, but the detailed drawings of Miller (1967) give good clues. DEFINITION: 6 pedipalp: Embolus emerging from the terminal plate of the embolar radix (“scheibenförmige Basalplatte” in Miller (1967: 269) and “Truncus” in Miller (1967: pl. 4, figs 3-4, 7-8). Radix emitting a pointed apophysis near to the base of embolus (Figs 59, 62) and “S” in Miller, 1967: pl. 4, figs 3-4, 7-8). Female: Epigynal plate in anterior sector (Fig. 66 and Grimm, 1985: figs 236, 238). SPECIES INCLUDED: C. gracilis (Canestrini, 1968) comb. n., C. pygmaeus (Miller, 1943) comb. n., and C. ibericus sp. n. NEW OR LITTLE KNOWN ZELOTINAE 317 embolar radix ‘ ‘ e , LI LU DI LI Li LI e LU LI LI ‘ . * J LI) dan ananne** ” . Coon00*° FIGS 50-57 Civizelotes dentatidens (Simon). (50-54) Left male palp. (50) Prolateral view. (51) Ventral view. (52) Retrolateral view. (53) Prolateral-ventral-apical view. (54) Ventral-apical view. (55) Epigynum. (56) Vulva, external-lateral view. (57) Id., dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. 518 A. SENGLET Civizelotes ibericus sp. n. Figs 58-66 HOLOTYPE: SPAIN, Andalusia, Cordoba, Brefia dam, evergreen oaks litter, 37°51'N 05°04'W; 3; 01.06.2002. PARATYPES: SPAIN, Nueva Castilla / La Mancha, Toledo, Cardiel de los Montes (rio Alberche), 40°02'N 04°39'W; 1 2 (with vulva in microvial); 15.06.1969. — Andalusia, Granada, Pradollano (S. Nevada), stones, dwarf Juniperus, 2350 m., 37°07'N 03°24 2 2 (with vulva in microvial); 28.05.2002. — Huelva, Santa Olalla (rio Cala), 37°55'N 06°11'W; 1 2; 04.07.1969. ETYMOLOGY: The species name reflects the biogeography of the new species. DESCRIPTION of d: Prosoma tawny-brown, with a few posterior bristles. Opisthosoma grey, with medium-long hairs. Legs brown. Total length 4.00. Prosoma: 1.90 long, 1.43 wide, 0.80 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.06, ALE 0.08, PME 0.07, PLE 0.07; AME-AME 0.06, AME-ALE 0.01, PME- PME 0.03, PME-PLE 0.04, ALE-PLE 0.04. MOQ length 0.20, front width 0.34, back width 0.40. Clypeus: 0.08 from AME, 0.05 from ALE. Pedipalp (Figs 58-62): Patella longer than dorsal tibia length. Terminal plate of embolar radix expanded into a posterior branch catching the embolar loop (Fig. 62). Radix expanding into a mem- branous flap along the embolus (Fig. 61). Radix emitting a pointed apophysis at base of embolus. Relict embolar base less reduced than in C. gracilis and C. pygmaeus, coupled to a relict terminal apophysis. Strong median apophysis with a large elevated base. Leg spination: Femora HI r001; IV p001, r001; metatarsi I v000, II v220. Tarsi I, I scopulate. Scutum occupying 1/4 of opisthosoma length. DESCRIPTION of 2: Prosoma dark brown. Opisthosoma black, with medium- long hairs. Legs brown, with tarsi and metatarsi slightly lighter; old specimens much lighter. Total length 3.60. Prosoma: 1.46 long, 1.10 wide, 0.58 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.05, ALE 0.08, PME 0.07, PLE 0.07; AME- AME 0.03, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.03, ALE-PLE 0.04. MOQ length 0.19, front width 0.28, back width 0.31. Clypeus: 0.06 from AME, 0.03 from ALE. Epigynum with sinuous lateral folds (Fig. 66). Anterior median sector filled with a globular epigynal plate containing two slits; lateral pockets situated below the folds. Vulva (Figs 63-65). Leg spination: Femora IV p001, r001; metatarsi I, II v220. Tarsı I, II scopulate. DIAGNOSIS: Civizelotes ibericus differs from the other two species in this group: 1 Absence of a prolateral tegular apophysis ["h” for “Hocker des Tegulum” in Miller (1967: pl. 4, figs 3-4, 7-8)]. 2 Embolar base less vestigial and presence of a relict terminal apophysis at its base. The female paratypes found in central and southern Spain are indicating a wide distribution in Spain, and the probability that they are conspecific with the male holotype is great. Civizelotes caucasius-group DEFINITION: Posterior eye row procurved. d pedipalp: Strong elongated embolar radix (Fig. 67) terminating in a bowl. Embolar base containing the sperm duct (Fig. 68), reaching straight to the radix bowl. Relict terminal apophysis with a posterior sclerite without distinct connection. Embolus long, curled right (clockwise) on left palp. Atrium in anterior epigynal sector. NEW OR LITTLE KNOWN ZELOTINAE 519 emb. radix posterior plate È È embolus emb. radix 7 \: posterir plate embolar base cadi relict median — broken radix apophysis embol. radix \ Be 61 ventral apophyses De ein 62 ant. lateral ‚7 Be | FIGS 58-66 Civizelotes ibericus n. sp. (58-62) Left male palp. (58) Prolateral view. (59) Ventral view. (60) Retrolateral view. (61) Cymbium removed, dorsal view. (62) Id., apical view. (63) Vulva, ventral view. (64) Id., dorsal view. (65) Id., median-lateral view. (66) Epigynum. Bold lines indicate epigynal folds. Scale 0.2 mm. 520 A. SENGLET SPECIES INCLUDED: C. caucasius (L. Koch, 1866), C. sostitialis (Levy, 1998). REMARKS: The structure of the pedipalp (Figs 65-66), with a long embolar radix and a developed embolar base, excludes these species from the Civizelotes gracilis- group. The study of copulatory mechanism (Senglet, 2004: 95, figs 26-31) shows that the embolar base and radix are penetrating to mid-length of the vulva; radix not figured, but radix bowl visible at base of embolus (Senglet, 2004: fig. 31). Civizelotes caucasius (L. Koch, 1866) comb. n. Figs 67-68 Melanophora caucasia L. Koch, 1866: 144, pl. 6, fig. 87 (description of ). For previous synonymy see Platnick (2012). MATERIAL EXAMINED: GREECE, Macedonia, Serres, above Oreini, 1200 m, stones, 41°14'N 23°35'E; 5 3, 3 2; 30.06.2011. — Thessaloniki, west of Aghios Vassilios, 40°41'N 23°05'E; 1 d (last moult 17.06.2008); 31.05.2008. — Aghios Vassilios, 40°40'N 23°07'E; 2 © (last moults of 2 29.07. and 02.08.1998); 14.06.1998. — East of Apollonia, below stones, 40°37'N 23°32'E; 7 6,3 © (last moults of 2 01.07.2008); 01.06.2008. — Loutra /Volvi lake, 40°39'N 23°25'E; 3 6 (last moults of 2 & 16 and 22.06.1998); 14.06.1998. — Kastoria, north of Kastoria (pass), 40°35'N 21°18'E; 5 8,3 ? (last moults of 2 9 21. and 25.06.1998); 20.06.1998. — Thessalia, Larissa, Gonnos, 39°51'N 22°28'E; 1 ¢; 19.05.1968. — Sterea Hellas, Phthiotidas, north of Malesina, below stones, olive trees, 38°38'N 23°14'E; 2 © (last moults of 2 26.07. and 09.08.2008); 19.06.2008. — Near Malesina, 38°37'N 23°13'E; 3 &,5 2; 21.05.1968. — Same; 2 2; 15.06.1968. — Same; 6 6, 3 2 (with vulva in microvial, last moults of & 23.05.-25.06, of 2 02.-23.06.1998); 21.05.1998. — Mt Chlomos, 860 m, stones, 38°36'N 23°10'E; 2 8,5 © (last moults of 4 9 16.06.-01.07.2011); 14.06.2011. — Same, 980 m, stones, 38°36'N 23°01'E; 3 à, 3 ® (last moults of 2 15.06.-01.07.2011); 14.06.2011. — Tràgana, 38°37'N 23°07'E; 1 9; 21.05.1968. — Glyphada / Theologos, 38°39'N 23°14'E; 1 2; 04.06.1978. — Theologos, 38°39'N 23°12'E; 3 8,3 2 (with vulva in microvial, last moults of 2 29.05.-01.06.1998); 20.05.1998. — Eubea, Theologos, 38°29'N 23°47'E; 1 6; 11.06.2008. — Dystos lake, 38°22'N 24°08'E; 1 à; 17.06.1981. — Peloponnesus, Laconia, south-west of Monemvasie, stones, 36°40'N 23°01'E; 1 9 (last moult of 2 25.07.2011); 29.05.2011. — North-east of Geraki, 37°02'N 22°43'E; 1 4,1 9 (last moults of Sd 25.06.1998, of 2 01.09.1998); 13.05.1998. — Arcadia, east of Langadas, 830 m, stones, 37°40'N 22°03'E; 3 4,2 2 (last moults of 4 09.-27.06, of 2 08 and 27.07.2011); 05.06.2011. — Argolida, Palaia Epidauros, Pinus litter, 37°39'N 23°09'E; 1 3; 01.10.2004. — Same 1 8; 05.06.1981. — Same; 2 6,2 2 (last moults of 1 & 17.05, of 2 14.05 and 05.06.1998); 14.05.1998. — Above Platani, 700 m, 37°48'N 22°30'E; 4 3, 1 9 (last moults of & 20.06.-01.07, of 2 15.07.2011); 21.05.2011. — Corinthos, east of Angelokastro, stones, 37°45'N 23°01'E; 1 9 (last moult of 9 07.07.2011); 24.05.2011. — Cyclades, Paros, Sta Maria /Naoussa, Salicornia, 37°08'N 25°17'E; 1 2; 02.06.1998. — Naxos, north-east of Sangri, 37°03'N 25°27'E; 1 ® (last moult of 2 18.06.1998); 05.06.1998. — North of Aghios Prokopios, dry, below stones, 37°05'N 25°21'E; 1 2; 06.06.1998. — Koronido, 650-750 m, 37°09'N 25°37'E; 5 © (last moults 28.06.- 03.07.1998); 08.06.1998. — Crete, Irakleion, Damasta; 1 9 ; 28.06.1970. — Charakas, Eucaliptus bark, 35°01'N 25°08'E; 1 d (last moult 22.03.2000); 01.10.1999. — Lassithi, Kaminakion; 1 d, 1 2; 13.07.1970. — SPAIN, Nueva Castilla / La Mancha, Albacete, La Gineta (rio Jucar), fine leaf litter, 39°10'N 01°58'W; 2 ¢ (last moults 10 and 16.07.2002); 16.05.2002. — Ciudad Real, Fuencaliente, 38°23'N 04°18'W; 3 2; 03.08.1969. — Ruidera, 38°56'N 02°51'W; 1 8; 07.08.1969. — Estremadura, Badajoz, south of Venta del Culebrin / Monesterio, 37°58'N 06°14'W; 1 d (last moult 19.06.1969); 19.06.1969. — Andalusia, Granada, La Vidriera / Pto del Pinar, 38°03'N 02°34'W; 1 ® (last moult of 2 05.07.2002); 20.05.2002. — Road Puerto de la Ragua, pine forest, 37°09'N 03°03'W; 1 do (last moult 10.07.2002); 24.05.2002. — Collado del Muerto (S. Nevada), 1450 m, 37°08'N 03°28'W; 2 & (last moults 13 and 22.07.2002); 29.05.2002. — Cordoba, Almodovar del Rio (Brena dam), 37°50'N 05°04'W; 1 à ; 01.06.2009. — Sevilla, Pintado dam /Cazalla de la Sierra, 37°59'N 05°57'W; 1 4,1 9; 02.07.1969. — Huelva, Santa Olalla (rio Cala), 37°55'N 06°11'W; 1 3; 04.07.1969. -PORTUGAL, Beira Alta, Guarda, Maceira / Fornos de Algodres (Casal do Monte), 40°44'N 07°24'W; 1 d; 09.08.1971. — NEW OR LITTLE KNOWN ZELOTINAE 321 embolar radıx ir RSS embolar radix embolar radix TR term. apophysis È, relict 68 embolar radix bowl FIGS 67-68 Civizelotes caucasius (L. Koch), left male palp, cleared. (67) Apical view. (68) Ventral-prolateral view. BULGARIA, Centre, Pazardzik, Pestera; 1 9; 23.07.1972. — IRAN, Caspian See, Mazanderan, near Dasht, 37°19'N 56°04'E; 1 2; 27.07.1974. DESCRIPTION: See Grimm (1985: 281, figs 231, 234-235). Civizelotes solstitialis (Levy, 1998) comb. n. Zelotes solstitialis Levy, 1998: 139, figs 97-101 (description of & and 9). — Chatzaki, Thaler & Mylonas, 2003: 60, figs 46-47, 50.-51. — Deltshev, Lazarov & Blagoev, 2004: 194, figs 12-14. — Seyyar, Demir & Topcu, 2006: 50, figs 2A-B (d). MATERIAL EXAMINED: GREECE, Sterea Hellas, Phthiotidas, Tragana, 38°37'N 23°07'E; 1 2; 13.07.1968. — Theologos, 38°39'N 23°12'E; 2 6, 1 @ (last moults of 4 15. and 17.08, of 2 07.09.1998); 20.05.1998. — Cyclades, Naxos, Aria /Philoti, 37°02'N 25°29'E; 1 2 (last moult 06.09.1998); 11.06.1998. DESCRIPTION: See Levy (1998: 139, figs 231, 234-235). REMARK: In C. solstitialis half of the radix bowl is concealed by the very large base of the embolus. Genus Heser Tuneva, 2005 TYPE SPECIES: Heser malefactor Tuneva, 2005, original designation REMARK: Tuneva characterized the genus largely by the lack of typical Zelotes organs. In Ze/otinae the terminal apophysis is a ventral branch of the embolar base; in Heser this is not present. The embolar base is strongly reduced. The embolus arises from the prolateral sector. Females have no glandular ducts (blind paramedian ducts). Heser nilicola-group DEFINITION: In these species the apical part of the tegulum is grooved to support the embolus; in the type species the embolus rests on the prolateral part of the median apophysis. Apical grooved sector sometimes partly sclerotized. Long embolus curled 522 A. SENGLET left (conter-clockwise) on the left palp. Palpal tibia and retrolateral apophysis short. Embolar base variably reduced. No intercalary sclerite present; the base of the embolar base (Figs 69, 77, 79) can be mistaken for a intercalary. Median apophysis large, except in A. hispanus. The glandular ducts (blind paramedian ducts) are replaced by scattered glands (Figs 75, 83, 89). SPECIES INCLUDED: Heser nilicola (O. P.-Cambridge, 1874), H. schmitzi (Kulczynski, 1899), H. incisupalpis Levy, 1998, H. bernardi Marinaro, 1967, H. bonneti Marinaro, 1967 and HA. hispanus sp. n. KEY TO SPECIES OF THE HESER NILICOLA-GROUP SAS ee ee I o Res 5 la Median apophysis small (Figs 85-86).................. H. hispanus sp. n. lb Median'apophysis large (Figs (0277). are see 2 2a Embolar base reduced; embolus arising from basal prolateral tegular Secior (PICS MOTO his aes ee ee ee es ee ee es H. schmitzi 2b Embolar base with a projection; embolus arising from dorsal tegular SOCIOL (los Dec H. nilicola 3a Epigynal plate elongated or triangular (Figs 81-82, 87)................. 4 3b Epigynal plate rounded (Figs 73-74) ....................... H. nilicola 4a Epigynal plate triangular, widened almost to the anterior anchoring POCKEIS WICH ATIC Sen ta ig ce ere ee H. hispanus sp. n. Ab Epigynal plate with more or less parallel sides (Figs 81-82)...... H. schmitzi Heser nilicola (O. P.-Cambridge, 1874) Figs 69-75 Prosthesima nilicola O. P.-Cambridge, 1874: 380, pl. 5, fig. 8 (description of 3) For previous synomymy see Platnick (2012). MATERIAL EXAMINED: GREECE, Peloponnesus, Laconia, south-west of Mavrovouni, litter on sand, 36°43'N 22°32'E; 3 d,2 © (last moults of £ 09.06. and 02.07.2011); 26.05.2011. — Cyclades, Paros, Parikia; 3 d, 2 2; 25.06.1968. — Parikia, 37°05'N 25°09'E; 1 3, 1 @ (last moults of 4 29.06, of 2 05.06.1998); 01.06.1998. — Naxos, Kato Potamia, 37°06'N 25°26'E; 1 2; 10.06.1998. — Crete, Chania, Sougia, 35°15'N 23°48'E; 1 d (last moult 22.03.2000); 09.10.1999. — FRANCE, Corsica, Sud Corse, Testa pass /Bonifacio, 69 m; 1 d; 27.05.1971. — Acorane bridge /Sartène; 1 6; 28.05.1971. — Same; 1 2; 19.06.1999. — Portigliolo /Propriano, below vegetation & stones; 2 2; 17.06.1999. — ITALIA, Sardinia, Sassari, Palau (Liscia river); 1 2; 08.09.1968. — Castelsardo; 1 2; 26.05.1999. — Stagno di Calich /Alghero; 1 d, 1 @ (last moults of d 05.06, of 2 01.06.1999); 29.05.1999. — Nuoro, Cala Ginepro /Orosei, below Juncus; 1 ©; 10.06.1999. — Oristano, Iz Arénas / Narbolia, dry leaf litter on sand; 1 3, 1 2; 31.05.1999. — Stagno San Giusta, Eucalyptus bark; 1 5, 1 2; 01.06.1999. — Cagliari, Quartu (laguna); 1 2; 03.06.1999. DESCRIPTION: See Platnick & Shadab (1983: 274, figs 316, 327-328). REMARKS: The cleared d palp shows a relatively well developed embolar base with a projection (Figs 69-72). The base of the embolar base may be mistaken for an intercalary sclerite. 2: The epigynal plate is rounded. The distance between epigynal plate and anterior anchoring pockets is highly variable. Greek form (Fig. 73): Platnick & Shadab (1983: figs 265-266, showing a faulty connection of the median vulval duct); Chatzaki et al. (2003: fig. 85); FitzPatrick (2007: Fig. 95). Occidental form (Fig. 74, NEW OR LITTLE KNOWN ZELOTINAE 523 FIGS 69-75 Heser nilicola (O. P.-Cambridge). (69) Left male palp, prolateral view. (70) Id., ventral view. (71) Id., retrolateral view. (72) Id., cleared, dorsal view. (73) Epigynum, from Greece. (74) Id., from Sardinia. (75) Vulva, dorsal view (Sardinia). Bold lines indicate epigynal folds. Scale 0.2 mm. Sardinia): Cornick et al. (2004: fig. 3a); Melic (1995: figs 1-2, showing a faulty connection of the median vulval ducts): Sardinia, Corsica, south of France and north Spain. Heser schmitzi (Kulczynski, 1899) Figs 76-83 Prosthesima schmitzii Kulczynski, 1899: 359, pl. 6, fig. 32 (description of ). Zelotes schmitzi. — Platnick & Murphy, 1998: 118, figs 1-4 (9, description of à ). MATERIAL EXAMINED: SPAIN, Levant / Murcia, Alicante, Elche, palm grove, 38°17'N 00°42'W; 1 3; 15.05.2002. — Elda, 38°30'N 00°47'W; 1 gd; 19.06.1971. — Murcia, Archena, 38°07'N 01°17'W; 1 3; 17.05.2002. — Andalusia, Almeria, Adra (La Albufera), cultivated dryed pond, 36°46'N 02°58'W; 10 3, 3 2 (last moults of 2 & 27.05 and 15.06, of 2 9 30.05 and 12.06.2002); 25.05.2002. — Malaga, Torre de Mar, 36°44'N 04°07'W; 1 2; 27.07.1969. DESCRIPTION: Prosoma and legs brown, with lighter tarsi. Opisthosoma black, with adpressed hairs and black bristles. Leg spination: Femora IV r001; tibiae III r111, IV p110; metatarsi I v000, IV v220/222. Tarsi I, II scopulate. 524 A. SENGLET embolar base embolus embolar radix embolar base FIGS 76-83 Heser schmitzi (Kulczynski). (76-80) Left male palp. (76) Prolateral view. (77) Ventral view. (78) Retrolateral view. (79) Cleared, prolateral-ventral view. (80) Id., dorsal view. (81-82) Two epigyna, same population. (83) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. NEW OR LITTLE KNOWN ZELOTINAE ILS d : Total length 3.05. Prosoma: 1.36 long, 1.03 wide, 0.54 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.04, ALE 0.07, PME 0.06, PLE 0.06; AME-AME 0.03, AME-ALE 0.01, PME-PME 0.02, PME-PLE 0.02, ALE-PLE 0.03. MOQ length 0.10, front width 0.16, back width 0.18. Clypeus: 0.06 from AME, 0.04 from ALE. Pedipalp (Figs 76-80): Patella dorsally longer than tibia. In cleared bulbus (Figs 79-80) a short embolar radix and a vestigial embolar base visible. Median apophysis large. Scutum occupying 40% of opisthosoma length. 2: Total length 4.70. Prosoma: 1.53 long, 1.14 wide, 0.60 wide at level of pos- terior eyes. Eye sizes and interdistances: AME 0.04, ALE 0.07, PME 0.06, PLE 0.06; AME-AME 0.03, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.03, ALE-PLE. MOQ length 0.14, front width 0.27, back width 0.31. Clypeus: 0.04 from AME, 0.03 from ALE. Epigynum (Figs 81-82), epigynal plate variable, with sides parallel to feebly diverging. Vulva (Fig. 83) with scattered glands on the sides. Heser hispanus sp. n. Figs 84-89 HOLOTYPE: SPAIN, Estremadura, Badajoz, Monesterio, 38°05'N 06°16'W; 6; 19.06.1969. PARATYPES: Same locality as for holotype; 1 2 (with vulva in microvial); 19.06.1969. — Spain, Andalusia, Sevilla, Alanis, 38°02'N 05°11'W; 2 2 (with vulva in microvial); 01.07.1969. ETYMOLOGY: The species name refers to the geographical occurrence of these spiders. DESCRIPTION d HOLOTYPE: Prosoma mid-brown, with medium-sized bristles. Opisthosoma black, covered with medium-long bristles. Tarsı I, II scopulate. Total length 2.76. Prosoma: 1.20 long, 0.87 wide, 0.48 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.03, ALE 0.07, PME 0.06, PLE 0.06; AME-AME 0.03, AME-ALE 0.01, PME-PME 0.02, PME-PLE 0.02, ALE-PLE 0.04. MOQ length 0.12, front width 0.20, back width 0.23. Clypeus: 0.06 from AME, 0.04 from ALE. Pedipalp (Figs 84-86): Patella dorsally longer than the short tibia. Retrolateral tibial apophysis shorter than tibia, dorsally arched. Embolus emerging from backside of tegulum. Median apophysis small. Scutum occupying 35% of opisthosoma length. Leg spination: Femora IV p001, r001; metatarsi I v000; II v210; HI-IV v221. DESCRIPTION ® FROM ALANIS: Prosoma tawny-brown, with median to long bristles. Opisthosoma blackish, covered with dense median light coloured bristles. Legs entirely tawny. Total length 3.40 (3.26). Prosoma: 1.24 (1.22) long, 0.93 wide, 0.48 wide at level of posterior eyes. Eye size and interdistances: AME 0.04, ALE 0.06, PME 0.042, PLE 0.042; AME-AME 0.03, AME-ALE 0.01, PME-PME 0.03, PME- PLE 0.02, ALE-PLE 0.03. MOQ length 0.14, front width 0.20, back width 0.24. Clypeus: 0.04 from AME, 0.04 from ALE. Epigynum (Fig. 87) with triangular epigynai plate, its anterior part almost as wide as anterior anchoring pockets. Vulva (Figs 88-89 from Alanis) with scattered glands on the dorsal sides. Leg spination: Femora IV p001, r001; tibiae IV p110; metatarsi I v000; IV r112. Heser infumatus (O. P.-Cambridge, 1872) Drassus infumatus O. P.-Cambridge, 1872: 238, pl. 15, fig. 16 (description of d and @). Heser infumatus. Tuneva, 2005: 323 (transfer of d and © from Zelotes). For previous synomymy see Platnick (2012). 526 A. SENGLET FIGS 84-89 Heser hispanus sp. n. (84) Left male palp, prolateral view. (85) Id., ventral view. (86) Id., retro- lateral view. (87) Epigynum. (88) Vulva, ventral view. (89) Id., dorsal view. Bold lines indicate epigynal folds. Scale 0.2 mm. MATERIAL EXAMINED: GREECE, Cyclades, Paros, Parikia; 1 2; 25.06.1968. DESCRIPTION: See Levy (1998: 145, figs 112-115). REMARK: Not a member of the Heser nilicola-group; the female of H. infumatus has the anterior epigynal margin ventrally projected like a scapus. NEW OR LITTLE KNOWN ZELOTINAE 527 ACKNOWLEDGMENTS Many thanks to Dr Christine Rollard of the National Museum of Natural History in Paris, and to Mrs Janet Beccaloni of the Natural History Museum in London for making type material available; to Dr Christo Deltshev (Sofia) for reviewing the manuscript; to Dr Peter J. Schwendinger for his constant support and his precious help in editing this paper, and to Dr Andre Piuz (both of the Geneva Museum of Natural History) for producing the Scanning Electron Microscope pictures; to the Geneva Museum of Natural History for covering the charges of these pictures. REFERENCES CAMBRIDGE, O. P. 1872. General list of the spiders of Palestine and Syria, with descriptions of numerous new species, and characters of two new genera. Proceedings of the Zoological Society of London 1871: 212-354. CAMBRIDGE, O. P. 1874. On some new species of Drassides. Proceedings of the Zoological Society of London 1874: 370-419. CANESTRINI, G. 1868. Nuove aracnidi italiani. Annuario della Societa dei Naturalisti e Mate- matic Modena 3: 190-206. CHAMBERLIN, R. V. 1922. The North American spiders of the family Gnaphosidae. Proceedings of the Biological Society of Washington 35: 145-172. CHATZAKI, M., THALER, K. & MYLONAS, M. 2003. Ground spiders (Gnaphosidae; Araneae) from Crete and adjacent areas of Greece. Taxonomy and distribution. III. Zelotes and allied genera. Revue suisse de Zoologie 110(1): 45-89. CHATZAKI, M. 2010. New data on the least known zelotines (Araneae, Gnaphosidae) of Greece and adjacent regions. Zootaxa 2564: 43-61. CORNIC, J.-F., LEDOUX, J.-C. & VILLEPOUX, O. 2004. De araneis Galliae 1.2, Zelotes nilicola O. P.-Cambridge. Revue arachnologique 15: 17. DELTSHEV, C., LAZAROV, S. & BLAGOEV, G. 2004. Spiders (Araneae) from the eastern Rhodopes (Bulgaria and Greece) (pp. 181-198). In: BERON, P. & Popov, A. (eds). Biodiversity of Bulgaria 2. Biodiversity of eastern Rhodopes (Bulgaria and Greece). Pensoft, Sofia. DELTSHEV, C., BOSMANS, R., SPIEGELAERE, W. DE & PROVOOST, L. 2006. Zelotes balcanicus sp. n., a new and widespread species from the Balkan Peninsula (Araneae, Gnaphosidae). Revue suisse de Zoologie 113: 711-716. DENIS, J. 1935. Additions a la faune arachnologique de l'Ile de Port-Cros (Var). Annales de la Société d'Histoire Naturelle de Toulon 19: 114-122. FITZPATRICK, M. J. 2007. A taxonomic revision of the Afrotropical species of Zelotes (Arachnida: Araneae: Gnaphosidae). Bulletin of the British Arachnological Society 14: 97-172. GILTAY, L. 1932. Arachnides recueillis par M. d'Orchymont au cours de ses voyages aux Balkans et en Asie Mineure en 1929, 1930 et 1931. Bulletin du Musée Royal d'Histoire Naturelle de Belgique 8(22): 1-40. GILTAY, L. 1933. Description des arachnides nouveaux recueillis par M. A. d'Orchymont aux Balkans et en Asie Mineure en 1929-31. Acta pro Fauna et Flora Universalis 1(3-5): 1-8. GRIMM, U. 1985. Die Gnaphosidae Mitteleuropas (Arachnida, Araneae). Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg 26: 1-318. JEZEQUEL, J. F. 1962. Contribution a l'étude des Zelotes femelles (Araneidea [sic], Labidognatha, Gnaphosidae) de la faune française (2° note). Bulletin du Museum national d'Histoire naturelle, Paris 33: 594-610. KOCH, C. L. 1833. Arachniden. /n: HERRICH-SCHAFFER, G. A. W (ed). Deutschlands Insekten. Heft 119-121. Kocu, C. L. 1839. Die Arachniden. Fünfter Band, pp. 125-158, Sechster Band, pp. 1-156, Siebenter Band, pp. 1-106. J. L. Lotzbeck, Nürnberg. 528 A. SENGLET KOCH, L. 1866. Die Arachniden-Familie der Drassiden, Hefte 1-6. J. L. Lotzbeck, Nürnberg, 304 pp., 12 pls. KULCZYNSKI, W. 1899. Arachnoidea opera Rev. E. Schmitz collecta in insulis Maderianis et in insulis Selvages dictis. Rozprawy Wydzialu Matematyczno-Przyrodniczego Polskiej Akademii Umiejetnosci Kracow 36: 319-461. Levy, G. 1998. The ground-spider genera Setaphis, Trachyzelotes, Zelotes, and Drassyllus (Araneae: Gnaphosidae) in Israel. /srael Journal of Zoology 44: 93-158. Levy, G. 2009. New ground-spider genera and species with annexed checklist of the Gnaphosidae (Araneae) of Israel. Zootaxa 2066: 1-49. MARINARO, J.-Y. 1967. Les araignées d'Afrique du Nord. I. Sur une collection de Drassidae a peigne metatarsal d'Algérie. Bulletin de la Société Zoologique de France 92: 687-704. MELIc, A. 1995. Un Zelotes nuevo para Europa y otro para la Peninsula Ibérica (Araneae: Gnaphosidae). Zapateri, Revista aragonesa de entomologia 5: 179-181. MILLER, F. 1943. Neue Spinnen aus der Serpentinsteppe bei Mohelno in Mähren. Entomologicke Listy 6: 11-29. MILLER, F. 1967. Studien über die Kopulationsorgane der Spinnengattung Zelotes, Micaria, Robertus und Dipoena nebst Beschreibung einiger neuer oder unvollkommen bekannter Spinnenarten. Prirodovedne Prace ustavu Ceaskoslovenske Akademie Ved v Brno (N. S.) 1; 251-298. PLATNICK, N. I. & MURPHY, J. A. 1998. On the widespread species Zelotes schmitzi (Araneae: Gnaphosidae). Bulletin of the British Arachnological Society 11: 118-119. PLATNICK, N. I. & SHADAB, M. U. 1983. A revision of the American spiders of the genus Zelotes (Araneae, Gnaphosidae). Bulletin of the American Museum of Natural History 174: 271092, PLATNICK, N. I. 2012. The World Spider Catalog, Version 12.5. American Museum of Natural History, online at: http://research. amnh. org/entomology/spiders/catalog.html. SENGLET, A. 2004. Copulatory mechanisms in Zelotes, Drassyllus and Trachyzelotes (Araneae, Gnaphosidae), with additional faunistic and taxonomic data on species from southwest Europe. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 77: 87 119. SENGLET, A. 2011. New species in the Zelotes tenuis-group and new or little known species in other Zelotes groups (Gnaphosidae, Araneae). Revue suisse de Zoologie 118 (3): 513-559. SEYYAR, O., DEMIR, H. & Topcu, A. 2006. A contribution to the spider fauna of Turkey (Araneae: Gnaphosidae). Serket 10: 49-52. SIMON, E. 1878. Les arachnides de France, 4. Roret, Paris, 334 pp. SIMON, E. 1914. Les arachnides de France, 6, lre partie: Synopsis générale et catalogue des espèces françaises de l'ordre des Araneae. Roret, Paris, 308 pp. SOYER, B. 1967. Contribution à l'étude éthologique et écologique des araignées de la Provence occidentale. IX.- Sur quelques araignées du genre Zelotes Gistel. Bulletin de la Société Entomologique de France 72: 275-281. TUNEVA, T. K. 2005. A contribution on the gnaphosid spider fauna (Araneae: Gnaphosidae) of east Kazakhstan (pp. 319-332). In: LOGUNOV, D. V. & PENNEY, D. (eds). European Arachnology 2003. Arthropoda Selecta Special Issue No. 1. REVUE SUISSE DE ZOOLOGIE 119 (4): 529-546; décembre 2012 Leptopsalis foveolata sp. n., a new species of Stylocellidae from Thailand that displays a novel morphological feature in the suborder Cyphophthalmi (Arachnida, Opiliones) Ronald M. CLOUSE!* & Peter J. SCHWENDINGER? ! American Museum of Natural History, Division of Invertebrate Zoology, Central Park West at 79th St., New York City, NY 10024, USA 2 Department of Arthropodology and Entomology I, Natural History Museum of Geneva, P.O. Box 6434, CH-1211 Geneva 6, Switzerland * Corresponding author, E-mail: rclouse@amnh.org Leptopsalis foveolata sp. n., a new species of Stylocellidae from Thailand that displays a novel morphological feature in the suborder Cypho- phthalmi (Arachnida, Opiliones). - Leptopsalis foveolata sp. n., a species of Southeast Asian mite harvestman with an exceptional morphology, is described. Males and females of this species bear rows of deep pits on the dorsal and ventral side of the posterior end of their body. These pits are strongly granular but do not bear obvious pores or especially dense concen- trations of micropores. Also, many of the large granules on the body surface are flattened and digitated, and many around the pits are grooved. This is a new morphological feature in the suborder Cyphophthalmi, and its function is not clear. The species is only known from the type locality, Mt Sankalakheeree in Pattani Province, southern Thailand, formerly known as Bukit Besar. Keywords: Taxonomy - mite harvestmen - opisthosomal pits - Namtok Sai Khao - Bukit Besar. INTRODUCTION Stylocellidae is a family of Cyphophthalmi that can be found from the Eastern Himalayas to New Guinea and out to the Philippines, and which likely arrived in Southeast Asia on the Sibumasu terrane that rifted from the northern Gondwanan coastline in the Paleozoic (Clouse & Giribet, 2010). Cyphophthalmi in general are small harvestmen that favor humid leaf litter and caves, and which, until recently, were poorly understood and not commonly collected. Now that their remarkably poor dispersal abilities have been shown to make them excellent models for vicariance bio- geography (Boyer et al., 2007; Giribet et al., 2012), and as collecting techniques have improved, new collections of Cyphophthalmi have become more routine (Giribet, 2000). These collections have revealed considerable biodiversity, especially in tropical families (Benavides & Giribet, 2007; Clouse & Giribet, 2010). Moreover, not all of these new species differ by subtle body proportions (copulatory organs are still poorly Manuscript accepted 23.08.2012 530 R. M. CLOUSE & P. J. SCHWENDINGER investigated); some present entirely new morphological features, as in the species we describe here. Stylocellid taxonomy has recently been updated after the publication of several new phylogenetic hypotheses (Clouse ef al., 2009; Clouse & Giribet, 2010; Giribet et al., 2012). Twenty-seven of the family’s 34 described species had been placed in the genus Stylocellus, four in Fangensis, and the remaining three distributed one each in Leptopsalis, Miopsalis and Meghalaya (Clouse & Giribet, 2007; Giribet, 2000; Giribet et al., 2007; Schwendinger & Giribet, 2005). Leptopsalis beccarii Thorell, 1882 (the type of that genus) was in the synonymy of Stylocellus sumatranus Westwood, 1874 (see Thorell, 1890) for about a century, but it has been resurrected (informally in Clouse er al., 2009 and in Giribet, 2002; formally after examination of types in Clouse & Giribet, in press) and will soon receive fourteen species transferred from Stylocellus (Clouse, in press; Clouse & Giribet, in press). It is now understood that Leptopsalis encompasses the bulk of the family’s undescribed diversity, including all species on Sulawesi, New Guinea and Java, as well as most species on Sumatra and the Thai- Malay peninsula (Clouse & Giribet, 2010). The species described here is the first to be originally assigned to Leptopsalis in 130 years. The types of the new species were collected at Sai Khao Waterfall, at the foot of Mt Sankalakheeree (or Sankalakhierie) which peaks at ca 1000 m altitude. This is an isolated, still densely forested granite mountain massiv at the border of Pattani, Yala and Songkhla Province, known under the Malay names “Bukit Besar” (= big hill) and “Gunong Negiri” before the Sultanate of Pattani came under Siamese rule in 1909. Bukit Besar was sampled by the “Skeat-Expedition” in May 1899 (Skeat, 1954) and by the “Annandale & Robinson-Expedition” (N. Annandale took part in both expeditions) in April-May and August-September 1901 (Annandale & Robinson, 1903). The disco- very of a velvet-worm (probably Eoperipatus sp.) on Bukit Besar during the “Skeat- Expedition” highlights the presence of typical Malayan faunal elements. Other such elements (of arachnids) await publication. MATERIALS AND METHODS Legs, tarsi, and various details were photographed on an FEI Quanta 200 (www.fei.com) scanning electron microscope (SEM). Color photographs were taken under a Leica MZ 12.5 stereomicroscope (www.leica-microsystems.com) with a mounted JVC KY-F70B digital camera (www.pro.jvc.com), where images were captured at different focal planes and then assembled using the application Auto- Montage Pro Version 5.00.0271 by Syncroscopy (www.syncroscopy.com). We identify specimens using several code numbers, as they have been entered into different databases since their collection. The Natural History Museum of Geneva (MHNG) gave the first code (a sample code), which consists of an area and year abbreviation. The Museum of Comparative Zoology (MCZ) DNA database logs whole collections with identification codes that consist of the prefix “MCZ DNA” and a six- digit number. A Biota database (Colwell, 2004) for Cyphophthalmi in the Giribet Laboratory at the MCZ gives each specimen a different six-digit ID number, with the prefix “SPM.” Moreover, this species described here was also used earlier in phylo- genetic analysis under the moniker “Peninsula sp. 18” (Clouse et al., 2009; Clouse & Giribet, 2010). NEW SPECIES OF STYLOCELLIDAE FROM THAILAND SS] Appendage measurements were not always possible for certain specimens: the first (proximal) cheliceral article must be removed to measure its full length, trochanters can be broken when legs are removed, some individuals have injured or deformed appendages, and in this species in particular, the chelicerae are usually pulled tightly under the overhanging anterior edge of the dorsal scutum, obscuring the dorsal crest. Chelicera depth was measured just proximal to the dorsal crest to a point between the first and second ventral process. The depth of the fourth tarsus was measured on the proximal side of the adenostyle base. TAXONOMY FAMILY STYLOCELLIDAE Leptopsalis foveolata sp. n. Figs 1-34 HOLOTYPE: MHNG sample TH-99/4, MCZ DNA 101093, MCZ Cyphophthalmi database SPM003920; Sai Khao Waterfall, Mt Sankalakheeree, Namtok Sai Khao National Park, ca 20 km NW of Yala city, Pattanı Province, Thailand, 6°39'18.5"N, 101°5'50.5"E, 260 m; male; 22-X-1999, leg. P. Schwendinger. PARATYPES: MHNG sample TH-99/4, MCZ DNA101093, MCZ Cyphophthalmi data - base SPM003917-9 (Figs 1-3); 3 males; same data as for holotype. - MHNG sample TH-99/4, MCZ DNA101093, MCZ Cyphophthalmi database SPM003921-4 (Figs 4-6); 4 females; same data as for holotype. - MHNG sample TH-99/5, MCZ DNA101496, MCZ Cyphophthalmi data- base SPM004415; 1 female; 23-X-1999, 260-350 m, otherwise same data as for holotype. DEPOSITION AND PRESERVATION OF TYPES: All types are deposited in the MHNG except those used for SEM (SPM003918-9), which are deposited in the MCZ Cyphophthalmi SEM collection. Male paratypes SPM003918-9 were mounted for scanning electron microscopy, male paratype SPM003917 and female paratype SPM003922 were dissected to examine genitalia, and female paratypes SPM003921, SPM003922, SPM003924, and SPM004415 had DNA (remaining in MCZ) extracted and sequenced for phylogenetic studies (Clouse et al., 2009; Clouse & Giribet, 2010). DESCRIPTION: Body length (3.27 mm) of male holotype 2.13 times the maximum width (1.53 mm), equal across the ozophores and the second opisthosomal segment. Dorsal scutum (Figs 1, 4, 7): Transverse prosomal-opisthosomal sulcus and opisthosomal sulci distinct; mid-dorsal, longitudinal opisthosomal sulcus absent. Anterior margin of prosoma meeting cheliceral ridge with protruding, weakly bilobed lip, rising gradually; entire scutum weakly arched. Ventral opisthosoma (Figs 2, 5, 8): Transverse sulci shallow and granular, complete after sternite 3, especially shallow between sternites 3 and 4; first three sulci after tergite 3 weakly sinoidal, medial region bending forward about as far as lateral ends. Sutures between tergites 6 and 7, and 7 and 8 gently curved back laterally. Pits: Posterior opisthosomal sulci of males and females with deep, granular pits (Figs 7-12), these not corresponding to either tuberculate or microgranular integument ornamentation described by Murphree (1988). Dorsal pits mostly along sulci between tergites VI and VII and between tergites VII and VIII, numbering five in each row (Figs 7, 12). Sulcus between tergites V and VI granular, with small depressions. Ventral pits more irregular, along antertior edges of sternites 7 and 8, resembling deep, uneven folds along the sutures, forming two rows of five and six pits, respectively (Figs 8, 11). 332 R. M. CLOUSE & P. J. SCHWENDINGER Fics 1-3 Leptopsalis foveolata sp. n., male paratype SPM003917. (1) Dorsal view. (2) Ventral view. (3) Lateral view. NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 333 FIGS 4-6 Leptopsalis foveolata sp. n., female paratype SPM003922. (4) Dorsal view. (5) Ventral view. (6) Lateral view. Tergite IX and sternite 9 with wrinkled, granular edges opposite anal plate (Figs 8, 19). Pits densely covered with large granules, lacking large pores or other structures (Fig. 10). 534 R. M. CLOUSE & P. J. SCHWENDINGER FIGS 7-8 Leptopsalis foveolata sp. n., scanning electron micrographs of male paratypes SPM003919 and SPM003918, respectively. (7) Dorsal view. (8) Ventral view. Ozophores, eyes, and sculpturing: Ozophores lateral, raised above carapace edge (“type 2” of Juberthie, 1970), large, tapered, pointing anteriad (Fig. 14). Eyes distinct, large, with cornea, positioned anterior to ozophores (Fig. 15). Integument uni- formly microgranular but lacking larger tubercules except in particular locations. Large tubercules of two types: raised, smooth and flattened; and rough, digitate. Transitional tubercles also present around pits (Figs 13, 16). Raised tubercules on dorsal, anterior NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 535 FIGS 9-13 Leptopsalis foveolata sp. n., magnified views of pits of male paratypes SPM003919 (9-10, 12-13) and SPM003918 (11). (9) Dorsal pits dorsolaterally on right side. (10) Central pit between tergites VI and VII. (11) Ventral pits and anal region. (12) All dorsal pits. (13) Transition of granule morphology along lower left lip of central pit seen in Fig. 10. edge of carapace; on distal % of ozophores; on central anterior, dorsal prosoma; starting in posterior half and center of tergite VII through tergites VIII and IX; sparsely distributed in center of tergite VI; inside sulcus between tergites V and VI; on anal plate; sparsely distributed on sternite IX; inside, between and lateral to dorsal and ventral pits; inside ventral opisthosomal sulci; on coxae I and extreme distal anterior 536 R. M. CLOUSE & P. J. SCHWENDINGER Fics 14-17 Leptopsalis foveolata sp. n., male paratype SPM003919 (14-15) and male paratype SPM003918 (16-17). (14) Ozophore, dorsal view. (15) Base of chelicera, anterior margin of dorsal scutum, eye and ozophore, lateral view. (16) Transition of granule morphology along edge of ventral pit. (17) Spiracle. of coxae IV; and on all articles of all appendages except third article of chelicerae. Large flattened, digitate granules found in three longitudinal rows on dorsal side of prosoma; anterior to prosomal-opisthosomal sulcus; starting on tergite VI to anterior of tergite VII; and starting laterally on sternite 3, increasing through sternite 9. Ventral prosomal complex (Fig. 18): Coxae IV meeting along midline for dis- tance equal to length of gonostome; coxae HI not meeting, with weak sternum between proximal ends; contact zone between coxae II longer than that between coxae IV, with endites more than twice as large as those on coxae IV. Gonostome wider than long, evenly arched to rectangular anteriorly; first opisthosomal sternite slightly rounded anteriorly, forming weakly concave posterior edge of gonostome; lateral walls formed by distinct, elevated posteroproximal processes of fourth coxae, these most elevated on anterior edge. Anal region (Figs 11, 19): Sternites 7, 8 and 9 and tergite IX free; tergite IX wider than sternite 9, both with densely and coarsely granular, uneven surface opposite NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 537 n TN, N (try A, Hi no si lr ie o me heat Cts FIGS 18-19 Leptopsalis foveolata sp. n., male paratype SPM003918, details of ventral side. (18) Gonostome and ventral prosomal complex. (19) Anal region. 538 R. M. CLOUSE & P. J. SCHWENDINGER FIGS 20-25 Leptopsalis foveolata sp. n., appendages of male paratype SPM003918, lateral views. (20) Chelicera. (21) Palp. (22) Leg I. (23) Leg II. (24) Leg III. (25) Leg IV. anal plate; microgranular, even surface flush along edge of anal plate. Tergite VIII lack- ing gland pore; anal plate with large, raised granules and irregularly convex surface. Chelicera (Fig. 20): Third article (movable finger) about one-fourth length of second article, second article gradually narrowing for distal two-thirds of length. First article with distinct dorsal crest, ridge flattening laterally and visible only as smooth area leading to distinct, moderately-sized second ventral process. Dorsal crest often not visible in specimens, usually pulled under anterior edge of dorsal scutum. First (proximal) ventral cheliceral process distinct, slightly larger than second (distal) ventral process. First article with large, evenly spaced granules laterally and dorsally after dorsal crest. Second article with some distinct granules at curve to proximal joint; irregular, flattened granules laterally in proximal half; remainder of second article and third article smooth. Palp and legs: Palp granular from trochanter to proximal fourth of article IV, without process on trochanter (Fig. 21). Leg I granular to proximal third and dorsal sur- face of tarsus (Fig. 22); leg II granular to proximal half, dorsal and ventral surfaces of NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 539 FIGS 26-29 Leptopsalis foveolata sp. n., leg tarsi of male paratype SPM003918, lateral views. (26) I. (27) II. (28) IT. (29) IV. tarsus (Fig. 23); legs III and IV completely granular (Figs 24-25). Tarsus of leg I with distinct solea extending two-thirds length of tarsus (Fig. 26). Medial dorsal surface of tarsi of legs I and II with solenidia (a row of evenly spaced, curved, blunt-tipped sen- sory hairs; see Willemart & Giribet, 2010), these situated more distally on tarsus I; areas around solenidia generally without granules (Figs 26-27). Tarsus of leg IV even- ly granular; depth even for most of its length; adenostyle located at 30% of tarsus length from the proximal end, its tip pointing distally (Fig. 29). Spermatopositor: Microtrichia formula: 3, 10, 7+7; dorsal microtrichia on two or three different levels, medial three on each side attached distinctly proximally to the lateral four, these about half the length of the former; ventral microtrichia on two levels, medial one near apex of spermatopositor, lateral two more than halfway down to level of most lateral apical microtrichia; four apical microtrichia arranged in two pairs on either side of shallow, apical, medial invagination; remaining six apical micro- trichia evenly spaced along sides to just distal of widest point; microtrichia with few denticles proximal on their shafts. Ventral surface of spermatopositor with numerous denticles (Figs 30-31). Gonopore complex with broad lobus medialis lacking lateral processes; short lobuli laterales present, distinctly proximal, not reaching base of lacinia dorsalis, the latter extending distally to just beyond distal edge of lobus medialis, flanked by curved digiti extending the same length as lacinia dorsalis (Fig. 32). Ovipositor (Figs 33-34): Two apical lobes, each with several setae over whole surface, ending in apical seta; a pair of setae lightly shorter than the apical ones just subapical on each lobe; more subapically a sensitive process containing a dense set of slightly curved, shorter and furcate setae. Distinct pigmented bodies in distal half on prolateral side of each apical lobe. (These pigmented bodies have meanwhile also been found in another Leptopsalis sp. and in Stylocellus globosus Schwendinger & Giribet in Schwendinger et al., 2004, and are thus not apomorphic for Fangensis as postulated 540 R. M. CLOUSE & P. J. SCHWENDINGER 0.10 mm Fics 30-31 Leptopsalis foveolata sp. n., spermatopositor of male paratype SPM003917. (30) Dorsal view. (31) Ventral view. by Schwendinger & Giribet, 2005). Ventroproximal plates narrow, somewhat trian- gular, less wide than apical lobes and half their length, carrying three setae each. A pair of sac-like receptacles near base of each apical lobe. VARIATION: Females are about 9% longer and wider than males, but have the same body proportions. Among males, body size varies by almost 20%, and among females by around 13%. The longest specimen is female paratype SPM004415 (3.68 mm), which is 25% longer than the smallest male and relatively narrower than other specimens (length 2.3 times the width across the ozophores versus average of 2.1 NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 541 0.10 mm FIG. 32 Leptopsalis foveolata sp. n., gonopore complex of male paratype SPM003917, dorsal view. D = digitus, LM = lobus medialis, LL = lobulus lateralis, LD = lacinia dorsalis. for males and 2.2 for other females). The number of opisthosomal pits varies as fol- lows: Two male and two female paratypes have 1-2 small but deep pits in the sulcus between tergites V and VI (Fig. 12), others have none. Males (n=4) have five large pits in the sulcus between tergites VI and VII, and between tergites VII and VIII, females (n=5) have 5-6 large pits in each of these sulci. Males have six pits between sternites 6 and 7 and 4-6 pits between sternites 7 and 8, females have 3-6 pits in each of these sulci. DISTRIBUTION AND HABITAT: The new species is known only from the type locality. The types were collected by sifting and extracting (in Winkler/Moczarski soil extractors) leaf litter and humus from close to the Sai Khao waterfall, in primary lowland rain forest. Soil extractions (by the same method) below the waterfall, along a small side-branch of the stream, in July 2011 yielded no additional specimens. 542 R. M. CLOUSE & P. J. SCHWENDINGER FIGS 33-34 Leptopsalis foveolata sp. n., ovipositor of female paratype SPM003922. (33) Dorsal view. (34) Ventral view. AL = apical lobe, AS = apical seta, PB = pigmented body, R = receptacle, VP = ventroproximal plate. DISCUSSION The literature on Cyphophthalmi systematics has included a rich discussion of their morphological features, and several taxonomic groups have remarkably clear morphological synapomorphies (de Bivort & Giribet, 2004; Giribet & Boyer, 2002; Willemart & Giribet, 2010). Many of these are sexually dimorphic, and of these, most consist of male opisthosomal gland pores and associated modifications such as depressions, apophyses, or “hairy” structures called scopulae. In Stylocellidae, opis- thosomal gland pores occur in some species as a single opening on tergite VIII, which is nearly always associated with a longitudinal strip of modified sculpturing on the anal plate. Also, exclusive to Stylocellidae is another sexually dimorphic gland opening, this one on the ventral surface of the male first tarsus in Fangesis cavernarum Schwendinger & Giribet, 2005 (see Willemart & Giribet, 2010). NEW SPECIES OF STYLOCELLIDAE FROM THAILAND 543 TABLE 1. Lengths, widths (only when specified) and proportions (in brackets) of bodies (= dorsal scuta) and appendage articles of the male holotype and male and female paratype averages. Body proportions are of the dorsal scutum length divided by the corresponding width measurements [across widest part of the body (not counting the ozophores), and across the ozo- phores], and appendage proportions are the lengths divided by the depths. A - male holotype [ratio] Body Leg I Leg Il Leg III Leg IV Chelicera Palp Length yee Trochanter 0.31 [1.10] 0.26 [1.06] 0.23 [0.89] 0:33 [1,15] I (crest) 0.60 [1.68] Trochanter 0.33 [2.30] Width (body) 1.53 [2.14] Femur 0.89 [2.82] 0.76 [2.41] 0.76 [2.52] 0.89 [2.82] I (whole) Femur 0.56.1355] B - male average [ratio] (n = 4) Body Leg I Leg II Leg III Leg IV Chelicera Palp Length 3.18 Trochanter 0.32 [1.02] 0.25 [0.97] 0.23 [1.06] 0321227] I (crest) 0.56 [1.60] Trochanter 0.32 [2.30] Width (body) 151 2.11] Femur 0.83 [2.96] 0.72:]2.58] 0.70 [2.71] 0.86 [2.90] I (whole) 0.94 [2.49] Femur 0.54 [3.77] C - female average [ratio] (n = 5) Body Leg I Leg II Leg III Leg IV Chelicera Palp Length 3.46 Trochanter 0.29 [1.02] 0.26 [0.97] 0.28 [1.06] 0.33 [1.27] I (crest) 0.58 [1.89] Trochanter 0.30 [1.98] Width (body) 1.64 [2.11] Femur 0.87 [2.96] O72 (2.58) 0.74 [2.71] 0.84 [2.90] I (whole) Femur 0.51 [3.44] Width (across ozophores) 1.53 [2.14] Patella 0.51 [1.71] 0.47 [1.43] 0.37 [1:13] 0.50 [1.59] II 1.17 [7.46] Patella 0.36: [2.7] Width (across ozophores) 1.52 [2.10] Patella 0.47 [1.50] 0.42 [1.31] 0.36 [1.23] 0.45 [1.37] II 1.11 [6.45] Patella 0.32 [2.43] Width (across ozophores) 157221] Patella 0.45 [1.50] 0.40 [1.31] 0.37.]1.23] 0.44 [1.37] II 1.10 [6.51] Patella 0331235] II 0.30 [7.50] Tibia 0.41 [4.14] Tibia 0.66 [2.12] 0.54 [1.69] 0.51 [1.55] 0.59 [1.71] III 0.27 [6.28] Tibia 0.38 [3.78] Tibia 0.59 [2.12] 0.49 [1.69] 0.47 [1.55] 0.53 11.71] Ul 02416037] Tibia 0.37 [4.14] Metatarsus 0.27 [1.05] 029112) 0.23 [1.07] 0.26 [1.00] Tarsus 0.34 [3.43] Metatarsus 0.25 [1.09] 0.25 [0.99] 0.23 [1.30] 0.25 [0.98] Tarsus 0.31 [3,02] Metatarsus 0.26 [1.09] 0.23 [0.99] 0.28 [1.30] 0.23 [0.98] Tarsus 0.34 [3.62] Tarsus 0:76:12:31 0.66 [2.70] 0.66 [3.29] 0.73 [2.43] Tarsus 0.71 [2.39] 0.64 [2.84] 0.60 [2.76] 0.70 72.52] Tarsus 0.74 [2.39] 0.66 [2.84] 0.64 [2:70] 0:712.52] 544 R. M. CLOUSE & P. J. SCHWENDINGER Stylocellidae has two other exclusive features that are sexually dimorphic but not readily associated with glandular secretions: Rambla’s organ, which is a modified patch of cuticle on the retrolateral side of the fourth tarsus of males in several species (see Rambla, 1994; Schwendinger & Giribet, 2005), and a row of spines on the ventral surface of the first tarsus of Stylocellus globusus, which is denser in males than females (Willemart & Giribet, 2010). The feature we describe here, the deep pits near the pos- terior end of males and females of L. foveolata sp. n., is also not apparently glandular in function but is additionally not sexually dimorphic. The only indication that these pits are associated with a liquid is the modification of granules along the posterior edge of each pit row, the grooves and digitate surfaces perhaps used to channel and disperse some compound. The one special feature in Cyphophthalmi this most resembles is Hansen’s organ, which is an irregularly grooved raised area of cuticle found on the coxae and more distal leg articles of both males and females in the African genus Ogovea. Like Rambla’s organ, Hansen’s organ has micropores but is not clearly glandular in function (Giribet & Prieto, 2003; Willemart & Giribet, 2010). Sexually dimorphic features are rare in Leptopsalis, the only occurrence being the anal gland pore in a few of the early lineages of the family (Clouse, in press; Clouse et al., 2009). This suggests that there has been a profound behavioral shift in the genus whereby such features are not needed, despite living in a wide variety of habitats and radiating into the largest clade in the family. The pits described here in L. foveolata sp. n. show no difference between males and females, and may indicate an adaptation to some novel selection pressure at the type locality. Living near a waterfall, do these animals experience flooding and need these pits to channel water or trap air bubbles? But why then only at their posterior end? If anything, this novel structure highlights how much more there still is to discover about Cyphophthalmi behavior and ecology. ACKNOWLEDGEMENTS Gonzalo Giribet (MCZ) kindly read an earlier draft of this paper and greatly improved it. Christina Lehmann-Graber produced the ink-drawings from sketches, Florence Marteau (both MHNG) scanned them. The Royal Forest Department of Thailand allowed PJS to collect in protected areas. REFERENCES ANNANDALE, N. & ROBINSON, H. C. 1903. Itinerary in Perak, Selangor, and the Siamese Malay states. Fasciculi Malayenses (supplement) 1: 1-43, 1 map. BENAVIDES, L. & GIRIBET, G. 2007. An illustrated catalogue of the South American species of the cyphophthalmid family Neogoveidae (Arthropoda, Opiliones, Cyphophthalm1) with a report on 37 undescribed species. Zootaxa 1509: 1-15. Boyer, S. 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CLOUSE, R. M., DE BIVORT, B. L. & GIRIBET, G. 2009. A phylogenetic analysis for the South- east Asian mite harvestman family Stylocellidae (Opiliones: Cyphophthalmi) - a combined analysis using morphometric and molecular data. /nvertebrate Systematics 23: 515-529. COLWELL, R. K. 2004. Biota 2: The Biodiversity Database Manager. Sinauer Associates, Sunder- land, MA, available at http://viceroy.eeb.uconn.edu/biota/ (accessed 1.V111.2012). DE BIvORT, B. L. & GIRIBET, G. 2004. A new genus of cyphophthalmid from the Iberian Peninsula with a phylogenetic analysis of the Sironidae (Arachnida: Opiliones: Cyphophthalmi) and a SEM database of external morphology. Invertebrate Systematics 18: 7-52. GIRIBET, G. 2000. Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). Revista Iberica de Aracnologia 2: 49-76. GIRIBET, G. 2002. Stylocellus ramblae, a new stylocellid (Opiliones, Cyphophthalmi) from Singapore, with a discussion of the family Stylocellidae. 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Reminiscences of the Cambridge University Expedition to the north-eastern Malay states, 1899-1900 (pp. 9-147). In: GIBSIB-HILL, C. A. (ed.). The Cambridge University Expedition to the north-eastern Malay states and upper Perak, 1899-1900. Journal of the Malayan Branch of the Royal Asiatic Society 26(4): 3-174. THORELL, T. 1882. Descrizione di alcuni Aracnidi inferiori dell’ Arcipelago Malese. Annali del Museo Civico di Storia Naturale di Genova (series 1) 18: 21-69, pls IV-VI. THORELL, T. 1890. Aracnidi di Nias e di Sumatra raccolti nel 1886 dal Sig. E. Mondigliani. Annali del Museo Civico di Storia Naturale di Genova (series 2a) 10: 5-106. 546 R. M. CLOUSE & P. J. SCHWENDINGER WILLEMART, R. H. & GIRIBET, G. 2010. A scanning electron microscopic survey of the cuticle in Cyphophthalmi (Arachnida, Opiliones) with the description of novel sensory and glandular structures. Zoomorphology 129(3): 175-183. WESTWOOD, J. O. 1874. Thesaurus Entomologicus Oxoniensis; or, illustrations of new, rare, and interesting insects, for the most part contained in the collections presented to the University of Oxford by the Rev. F. W. Hope, M. A., D. C. L., F. R. S., & c. with forty plates from drawings by the author. Clarendon Press, Oxford, 205 pp. REVUE SUISSE DE ZOOLOGIE 119 (4): 547-559; décembre 2012 Ophiotaenia lapata sp. n. (Eucestoda: Proteocephalidea) from Madagascar: a parasite of the endemic snake Madagascarophis colubrinus (Colubridae) Voahirana R. RAMBELOSON]|, Hafaliana C. RANAIVOSON! & Alain de CHAMBRIER?. | Department of Animal Biology, Faculty of Sciences, Antananarivo University, Madagascar. 2 Department of Invertebrates, Natural History Museum, Geneva, Switzerland. Address for correspondence: A. de Chambrier, Muséum d’histoire naturelle, PO. Box 6434, CH-1211 Geneva 6, Switzerland. E-mail: alain.dechambrier@ville-ge.ch Abstract: Ophiotaenia lapata sp. n. (Cestoda: Proteocephalidea) from Madagascar: a parasite of the endemic snake Madagascarophis colu- brinus (Colubridae). - A new proteocephalidean cestode, Ophiotaenia lapata sp. n., 1s described from the gut of the endemic colubrid snake Madagascarophis colubrinus (Schlegel, 1837) (Colubridae) from Madagascar. The new species differs from all but one Ophiotaenia species parasitic in African snakes by the presence of an apical organ. Ophiotaenia lapata differs from O. adiposa Rudin, 1917, which also possesses an apical organ, by the number of testes (89-170 in the new species versus 170-220 in O. adiposa), by the position of the genital pore in relation to the anterior margin (43-53% of proglottis length in O. lapata versus 20-25%) and the scolex width (240-280 um in the former species versus 500-600 um in the latter). The new species possesses, unlike all but one Ophiotaenia species parasitic in African snakes, a three-layered embryophore. The other African species have two-layered embryophore except for Ophiotaenia georgievi de Chambrier, Ammann & Scholz, 2010, which can be distinguished by the absence of an apical organ, by the number of uterine branches on each side (23-28 in O. georgievi versus 41-68 in O. lapata) and by the total length of the strobila (50 mm in O. georgievi and 295 mm in O. lapata). Ophiotaenia lapata is the third proteocephalidean cestode reported from Madagascar. Keywords: Eucestoda - taxonomy - morphology - Serpentes - helminths - Africa. INTRODUCTION Tapeworms of the order Proteocephalidea Mola, 1928 are frequent and cosmo- politan parasites of freshwater fishes, reptiles and amphibians (Rego, 1994). The genus Ophiotaenia La Rue, 1911 comprises ninety-six species parasitic in reptiles and amphibians (for a list of species see Schmidt, 1986; de Chambrier et a/., 2010, 2012). Two specimens of the colubrid snake Madagascarophis colubrinus (Schlegel, 1837) Manuscript accepted 13.09.2012 548 V. R. RAMBELOSON ET AL. endemic to Madagascar were examined in 2011 by the present authors. They harboured proteocephalidean cestodes belonging to Ophiotaenia, which are described herein as a new species. MATERIALS AND METHODS Snakes examined were killed by injection of Nembutal (sodium pentobarbitone) and immediately dissected. Freshly collected tapeworms were rinsed in a saline solu- tion (0.9% in water), placed in a Petri dish with a small quantity of saline and fixed with hot, almost boiling 4% neutral formalin. After 1-3 weeks, the worms were trans- ferred and stored in 75% ethanol solution. The worms used for morphological study were stained with Mayer’s hydrochloric carmine, destained in 75% acidic ethanol (i.e. 75% ethanol with about 2 ml HCI/100 ml), dehydrated in an ethanol series, cleared with eugenol (clove oil) and mounted as permanent preparations in Canada balsam. For histology, pieces of the strobila were embedded in paraffin wax, sectioned transversely at 15 um intervals, stained with Weigert’s haematoxylin and counterstained with 1% eosin B acidic solution (see Scholz & Hanzelova, 1998; de Chambrier, 2001; Oros et al., 2010). Eggs were studied in distilled water. Scolex for scanning electron micro- scopical (SEM) observation was dehydrated in a graded ethanol series (80%, 96%, twice 100%), then transferred to a graded amyl/acetate series, critical point-dried in CO,, sputter coated with gold and examined with a Zeiss 940A electron microscope at the Natural History Museum, Geneva. Microthrix terminology follows Chervy (2009). All measurements are given in micrometres unless otherwise indicated. Abbreviations used in the description are as follows: x = mean; n = number of measu- rements; CV = coefficient of variability (expressed in %); RSO = ratio of the width of the ovary to the width of the proglottis; PGP = position of genital pore (cirrus pore) expressed as percentage of its position to the proglottis length from the anterior mar- gin; RSCS = relative size of the of cirrus-sac expressed as percentage of its length to the width of the proglottis. The worms studied were compared with those collected by G. Brygoo, Madagascar Pasteur Institute, between 1960 and 1970. His collection of tapeworms from snakes and amphibians from Madagascar was originally deposited in the Institute of Zoology in Neuchatel, Switzerland, and then transferred to the Natural History Museum in Geneva. New material studied has been deposited in the helminthological collection of the Natural History Museum, Geneva, Switzerland (acronym MHNG PLAT), in the Department of Animal Biology, Faculty of Sciences, University Antananarivo, department of Animal Biology (acronym UADBA), and in the Institute of Parasitology, Ceské Budéjovice, Czech Republic (acronym IPCAS). RESULTS Ophiotaenia lapata sp. n. Figs 1-14 TYPE MATERIAL: Holotype MHNG-PLAT-79567 (field number Mad 007a) (1 slide) and 7 paratypes: MHNG-PLAT-79568 (Mad 007b), 3 whole mounted slides and 18 cross sections; MHNG-PLAT-82165 (Mad 007p), 1 slide; MHNG-PLAT-82166 (Mad 007x), 1 slide, scolex used for SEM; MHNG-PLAT-82167 (Mad 007z), 1 whole mounted slides and 10 transverse sections; MHNG-PLAT-79570 (Mad 008a), 1 whole mounted slide. UADBA No50001 and 50003, (Mad 007), two specimens, 2 slides. A NEW PROTEOCEPHALIDEA FROM MADAGASCAR 549 i a j | . ee - FIGS 1-7 Ophiotaenia lapata sp. n. Scanning electron micrographs. Paratype (MHNG-PLAT-82166, field number Mad 007HFx). (1) Scolex, apical view (2) Scolex, dorsoventral view. (3) Scolex, lateral view. (4) Capilliform filitriches at level of the apex of the scolex. (5) Capilliform filitriches and small gladiate spinitriches at marginal surface of sucker. (6) Capilliform filitriches and small gladiate spinitriches at non-adherent surface of sucker (7) Gladiate spinitriches at proliferation zone surface. Scale-bars: 1-3 = 50 um; 4-7 = 3 um. OTHER MATERIAL: MHNG-PLAT-82172 (field number Mad 007hf), 2 whole mounted slides, (voucher material used for the study of the eggs). - MHNG-PLAT-82169 (field number Mad 007y), 1 whole mounted slides and 10 transverse sections. - UADBA No50002, 50004 550 V. R. RAMBELOSON ET AL. (field number Mad 007), and 50005 (field number Mad 007c), 3 mounted slides. Some pieces of gravid proglottides were placed in alcohol for DNA analyses. - MHNG-PLAT-79569 (Mad 007c); MHNG-PLAT-82175 (Mad 008hf), and IPCAS C-625, 1 whole mounted slide and 4 slides of serial cross sections (from MHNG-PLAT-82175). - MHNG-PLAT-73222, from Brygoo material; Madagascar, Befandriana S., October 1967, 1 whole mounted slide and 13 transverse sections. TYPE LOCALITY: Ambinda Nord/Beanka (-17.93986°Lat; 44.46822°Long), 18 November 2011. All material listed above is from this locality, except MHNG-PLAT- 719222, DESCRIPTION (based on 8 specimens, 4 complete and 4 incomplete): Proteo- cephalidae, Proteocephalinae. Cestodes up to 295 mm long; maximum width 1.2 mm. Strobila acraspedote, anapolytic. All proglottides longer than wide (length: width ratio 1.03-1.52 to 4.80- 6.00, from immature to gravid). Scolex 120-150 long and 190-280 (n = 3) wide, sligh- tly wider than neck (Figs 1-3, 8). Suckers uniloculate, round, slightly embedded, 85- 115 (n = 12) in diameter, representing 30-48% of scolex width (Figs 1, 2). Apical or- gan 40-45 in diameter, i.e. 15-19% of scolex width, surrounded by cells with finely granular cytoplasm (Fig. 8). Proliferation zone 2.5-3.6 mm long and 140-185 wide. Internal longitudinal musculature weakly developed, anastomosed, formed by numerous tiny muscle fibres (Figs 10, 11). Ventral osmoregulatory canals overlapping testes, reaching laterally vitelline follicles, 5-25 in diameter, with secondary canals di- rected externally; dorsal canal narrow, thick-walled, 5-10 in diameter (Fig. 13). Testes medullary, on one layer, in two narrow lateral bands (poral field separa- ted by terminal genitalia to preporal and postporal groups). Testes rarely reach anterior margin of proglottis, but never reach to ovary, occupying 81-88% of total length of pro- glottis (Figs 10, 13, 14). Testes 89-170 (x = 128, n = 38, CV = 17%) in number, with 47-84 (x = 66) aporal testes, 25-49 (x = 37) preporal testes and 19-38 (x = 29) post- poral testes. Testes spherical, 50-65 (x = 55, n = 22) in diameter, degenerated in gravid proglottides (Fig. 14). Cirrus-sac elongate, thick-walled, 170-250 (x = 210, n = 53) long and 75-90 (x = 80 ,n = 10) wide (Fig. 9); RSCS 19-26 % (x = 22%, n = 53, CV = 8 %). Cirrus length represents about 70% of cirrus-sac length. Vas deferens strongly coiled, situated between proximal part of cirrus-sac and midline of proglottides, but never crossing it. Genital atrium present; genital pores alternating irregularly, more or less equa- torial, PGP = 43-53% (x = 49 %, n= 12, CV = 6%) (Figs 13, 14). Genital ducts passing between osmoregulatory canals. Ovary medullary, bilobed (Figs 11, 13, 14), 520-840 wide, RSO = 68-81% (x = 75%; n = 55; CV = 4%). Mehlis’ glands 50-80 (x = 60, n = 9) in diameter, represen- ting 9-12% of proglottis width (Figs 11, 13, 14). FIGS 8-12 Ophiotaenia lapata sp. n. (8) Holotype, scolex, dorsoventral view (MHNG-PLAT-79567). (9) Paratype, vagina and cirrus-sac region, dorsal view (MHNG-PLAT-79568). (10-11) Transverse sections of a mature proglottis at level of anterior part and ovary, respectively (MHNG-PLAT- 82167). (12) Eggs drawn in distilled water showing the three-layered embryophore (MHNG- PLAT-82172); additional layer marked by an arrow. Abbreviations: ao — apical organ; cg — cells with finely granular cytoplasm; ci — cirrus; cs — cirrus-sac; do — dorsal osmoregulatory canal; em A NEW PROTEOCEPHALIDEA FROM MADAGASCAR 551 LUS Se oh Beat ENFIN, wg Im vo sc ut te vi 500 pm ee CE NEE 11 nl 1%, doses Ae Sh ASA hg ASS ho. AB Aha Agta Mar tad ams ans ni vd vc mg Ov vi / / / Î as j / = fi | S Vi \ i = I} 7 = fi | \ = / | \ È tei | \ 7 | \ LT si N V | \ { 4 N RR Pd S 1 1] NA / \ SP SS 2 Nr eal: —== EN A I ra (/ : i \ / \ Py el ee 4 F f VEE: | = DA / x = Ee — embryophore; Im — internal longitudinal musculature; mg — Mehlis glands; nl — longitudinal nerve; oe — outer envelope; on — oncospheres; ov — ovary; sc — subtegumental cells; so — small outgrowths; te — testes; ud — uterine diverticles; ut — uterus; va — vas deferens; vc — vaginal canal; vd — vitelloduct; vi — vitelline follicles; vo — ventral osmoregulatory canals; vs — vaginal sphincter. Scale-bars: 8 = 100 um; 9 = 250 um; 10-11 = 500 um; 12 = 50 um. 352, V. R. RAMBELOSON ET AL. Vitelline follicles medullary, arranged in two lateral fields near margins of pro- glottides, occupying 90-95% of proglottis length, interrupted at level of cirrus-sac (Figs 13, 14). Vaginal canal forming small seminal receptacle anterodorsal to ovarian isthmus. Terminal part of vaginal canal (pars copulathrix vaginae) surrounded by large vaginal sphincter and chromophilic cells (Fig. 9). Vagina anterior (62%; n = 26) or posterior (38%) to cirrus-sac. Primordium of uterine stem medullary, present in immature proglottides. Development of uterus of type 1 according to de Chambrier et al. (2004): in immature proglottides, uterine stem straight, occupying most length of proglottis but never crossing ovarian isthmus, formed by wide longitudinal band of chromophilic cells situated along midline of proglottides. Lumen of uterus appearing in first mature pro- glottides (Fig. 13); diverticula (lateral branches) formed before first eggs appear in uterine stem. In pregravid proglottides, uterus occupying up to 33% of proglottis width, with 41-68 thin-walled lateral diverticula on each side. In gravid proglottides, diver- ticula occupying up to 80% of proglottis width. Uteroduct enters uterus almost at level of ovary isthmus. Eggs round, with outer envelope 140-165 in diameter (Fig. 12). Embryophore spherical, with thick supplementary spherical layer between outer envelope and onco- sphere, thus forming three-layered embryophore: internal layer 18-20 (n = 8) in diameter, middle layer 29-33 (n = 7) in diameter; external layer 34-39 (n = 8) in diameter; External layer of embryophore covered by small outgrowths 2.5-4 long; oncosphere spherical, 14-15 in diameter (n = 8), with three pairs of hooks, 8-9 long (Fig. 12). Eggs mature very fast in uterus and ripe eggs (oncospheres with hooklets) are present in the first pregravid proglottides. TYPE HOST: Madagascarophis colubrinus (Schlegel, 1837) (Serpentes, Lamprophiidae). SITE OF INFECTION: Intestine. PREVALENCE: 2/2 (100%). ETYMOLOGY: The new species is named after the vernacular local name of the host, 1.e. “lapata”. DIFFERENTIAL DIAGNOSIS: The new species is placed in Ophiotaenia La Rue, 1911 (Proteocephalinae) because of the medullary position of the vitelline follicles, the unarmed scolex with uniloculate suckers and testes forming two separate fields (Schmidt, 1986). Ninety-six species of Ophiotaenia parasitizing reptiles and amphi- bians are currently recognized as valid (Freze, 1965; Schmidt, 1986; Ammann & de Chambrier, 2008; Marsella & de Chambrier, 2008; Coquille & de Chambrier, 2008; de Chambrier & de Chambrier, 2010; de Chambrier et a/., 2010, 2012). Out of these, 64 species are parasites of snakes (Squamata) (see Table 1 in de Chambrier et al., 2010). According to Freze (1965), the species of Ophiotaenia are limited in their dis- tribution to individual continents and/or zoogeographical regions; this assumption has been then supported by other data, like an high degree of isolation determined by the presence of a number of endemic genera such as Marsypocephalus, Sandonella (both DIS A NEW PROTEOCEPHALIDEA FROM MADAGASCAR 0 p ss ove nese Abeer? " en = RER NS = SW ENS CRAN LE A Ya ey ware! n° “a do @ aes dè oy Ot ZN % O O 02% 0,0 0% 0509 00 RT OT NAS LAN AU Hy AVM | Ve XM ! | È | a, U MU à 000$ 3; PEEPS 00 000 © O ci O di PSS Of ODD Oo stelo vive a 41/7 pres even? 4 Sf ats ve sol ud > Ok © 90 09 0009 BE Qp0Lo NER! 00 PATTI AMIN PERD, 79568). -PLAT- 1000 um ventral view (MHNG 14 ’ Fics 13-14 Ophiotaenia lapata sp. n. (13) Paratype, mature proglottis 1000 um. 500 um 13 (14) Paratype, schematic view of a gravid proglottis, ventral view (UADBA No 50001). Abbreviations: cs — cirrus-sac; mg — Mehlis glands; ov — ovary; te — testes; ud — uterine diver - ticles; ut — uterus; va — vagina; vi — vitelline follicles; vo — ventral osmoregulatory canals. Scale- bars: 13 = 500 um; 14 554 V. R. RAMBELOSON ET AL. Africa), and Goezeella (South America) for fish parasites and Rostellotaenia (Africa), Acanthotaenia (Asia) and Kapsulotaenia (Australia, Papua New Guinea) for reptiles parasites (Freze, 1965); furthermore, for species of Ophiotaenia from amphibian hosts (de Chambrier er al., 2006) and from reptilian hosts (Ammann & de Chambrier, 2008), a strict specificity (oioxenous sensu Euzet & Combes, 1980) was observed in all spe- cies of this genus. For this reason, the new species is separable from 14 Ophiotaenia species found in snakes in Africa (for their complete list, see de Chambrier et al. 2010). Ophiotaenia lapata sp. n. differs from all but one Ophiotaenia species parasitic in African snakes by the presence of an apical organ (Table 1), the only African spe- cles possessing an apical organ being O. adiposa Rudin, 1917 described from Bitis arietans from Cameroun. Ophiotaenia lapata differs from O. adiposa by its lower number of testes (89-170 versus 170-220), position of the genital pore (situated at 43- 53% of the proglottis length from the anterior margin, i.e. almost equatorial in O. /a- pata, versus markedly pre-equatorial, i.e. at 20-25% length of the proglottis in O. adi- posa) and smaller scolex (width 240-280 um in the former species versus 500-600 um in O. adiposa) (Table 1). Ophiotaenia lapata n. sp. also differs from all but one Ophiotaenia species parasitic in African snakes in the possession of a third layer of the egg embryophore (Fig. 12). This layer is external to the oncosphere, i.e. it forms the internal envelope of the embryophore. The eggs of all African taxa described until now possess only a two- layered embryophore (Beddard, 1913; Rudin, 1917; Fuhrmann, 1924; Sandground, 1928; Hilmy, 1936; Mettrick, 1960, 1963; Southwell & Lake, 1939; Freze, 1965). A similar structure, i.e. an additional layer of the embryophore, was first observed in some other Proteocephalidea tapeworms (see de Chambrier & Vaucher, 1999; de Chambrier, 2006; Coquille & de Chambrier, 2008; Marsella & de Chambrier, 2008; de Chambrier et al., 2010; de Chambrier & de Chambrier, 2010; de Chambrier et al., 2012): This character is present in a wide range of proteocephalidean genera and geo- graphical areas, such as Proteocephalus (P. hobergi de Chambrier & Vaucher, 1999) ın Paraguay, Kapsulotaenia (K. sandgroundi Carter, 1943) in Indonesia, Cairaella (C. henrii Coquille & de Chambrier, 2008) in Ecuador, Ophiotaenia (O. alessandrae Marsella & de Chambrier, 2008 in Ecuador, O. gallardi (Johnston, 1911) in Australia and O. bungari de Chambrier, Binh & Scholz, 2012 in Vietnam). This additional layer of the embryophore, even if it seems to be a convergence phenomenon, is considered as a good discriminant character. The only African species, the embryophore of which is also three-layered as in the eggs of O. lapata, is O. georgievi de Chambrier, Ammann & Scholz, 2010 described recently from Leioheterodon geayi Mocquard. This species differs from O. lapata, besides being devoid of an apical organ (see above and Table 1), by the number of uterine branches (23-28 in O. georgievi versus 41-68 in O. lapata), and by the total length of the strobila (50 mm in O. georgievi versus 295 mm in O. lapata). ISO A NEW PROTEOCEPHALIDEA FROM MADAGASCAR ‘QIOYASOZUO JO ISYPWEIP - 44 ‘(S961) 9ZSIA UI SIMS] WOAF UONPI - » ‘UISI CUI JOLIQJUE OY} LOI SI} 0]F01d SY} JO YIPIM SY} 0) Yu] SJI JO 98ejuo91od se passaidxs 98S-SNILII9 IyJ JO OZIS SANIOI = SOSY :suonerasiqqy 3 23 89-Ir S6C 087-061 sok jue-ysod Nester “STI OL1-68 "u ‘ds vyndb) ‘O **81 009-055 P>IEUNSO 007 ou ‘que Ayyensn %0S %ST-OC 007-091 re, **31 OV-St 00€ PSUNSS 007 ou isod-jue e%0S-1P = %ST-OC OTE-091 14611941 ‘O Ot cI 98 06 OOS I ou 1sod-que *Y0S570S «%0S-9V Oec-OLI Hie nes DELE ch-0E OLC-OIT = OVIT-O6L ou ysod-jue YOS< = %ST-CT Ocl-Oll xapolydo ‘O SC OV-SC cS sol ou isod-jue «Volt %0C-9I 06-L9 zumogdu ‘O seme lG 07-91 OLI OTE-O0E ou isod-yue #708€ %CC-OC OIc-9LI SIJJ09UBIU ‘O 0€ LS-0S OS xo]oos ou é jue x%IS %II-O1 08 “bur ‘ds 1a1uuou ‘O ST SE 89< uoAlS jou é que Aljensn +%0S YEE 01-98 ‘but ds 1715524 ‘0 SETE SC EC OS SEC-SCC ou jue-ysod %IS-bH CEG6I Or1-Z6 142134023 ‘O 6€ 978€ 08€ < 009-00€ ou 3sod Ajpe1susd x7oTt NOTE OLT-OEI DIIUOGDS ‘O SS-8L OST OOIT-O00T OW jue-jsod #700S "2046 SCI-00I applosdyja 'O 97 8I-SI USAIS JOU 081-091 ou 6 + %oVS «%oV | 86-76 S1H]adoydbioo ‘O SI 0C-SI 08 é é isod-yue »%Lr-br %SC 59 SISU2]08U09 "O **CI OS-OV 00ÿ-00€ 009-008 SOA jsod-jue %ST-OC *%097 Occ-OLI psodipp 'O apis y9e9 910d S339 JO uo soyouRiq (wu) X9[09S uedIO BUISBA OY} [UJIU98 OU} S9]S9] JO Jojo Sun) y}8u9] Apoq vio, = JO Ypim [eoidy JO uonisogq Jo uonIsog SISI JOQUINN sowads (0107 “IP 12 RPLIqwey)y op Wo poıyıpour) voy ul sapydaı wor eruapjoıydo Jo satoads ‘] FTaVL V. R. RAMBELOSON ET AL. 556 CLCEL L ‘ds piuavjo1ydQ ouemmure N SISUAIUAADISDUL pupy L8I Ag pLTEL € ‘ds pIU20101dO 59618 euRlipurjog puiignj0o9 SIYdOIIDISDÉDPDN TESA S9/L6 TITEL 149131093 D1U20:01Y dg 896.7 OALIEUEULJUV 1dpas UOPOABJAYOIT 8LIA OL-8L LOTEL 9 ‘ds pıuonjo1ydo IL6TI eNISULIPUVY pupjuout DJAIUOpOUF OL/IL 6LTEL ¢ ‘ds miuanoiyd() C96L ASBIUBABQUIV SISUOLIDISDBDPDU DIUIZUDS I6T'M £9/807 T8TEL + ‘ds piuanjoiyd¢g ZO6I OI ejuewog 987] sısuojpfpypu sıydoun WTA 79/667 OLTEL pds miuomoiydQ = LI6L OT ATEFEUEIN neayeyd sısuojpfpypu sıydouy GTI LO/EST Seel, € ds piuopgoıydQ = L961 01 eueLipuejog pulignj0o9 SıydoliospöpppW STOH A L9/ISI Tacel nnd viuanjorydg L961'01 euenpuejog DULIGN]OD —s1ydolipospspppy STOA è L9/I81 O9TEL 7 ‘ds pıuonjo1ydo 19617 BUISEWEIPUV snypauyxas siydopyoydoiT 607 A 9/8 7 ds pruomoıydQo 1961 II eur] S1]D49]D] siydopijoydot7 ILTH 19/661 I9TEL [ ‘ds muamo1ydQ 6961 TI euenpuejog snjsapoul uopo1ajayolyT LIGA 69/8€ 18579 | ‘ds pıuamjo1ydo 1961" Ayonog-epoly snjsopou UOPOI2I2Y0IT EIN 19/87 2POIS9D TO6l'II BAUPUBURT snjsopou uopo4ojoyoıT O6TA = EI/LOT LLtS9 - OLYS9 140184008 muomoıydQ 8961" OALIEUEUBJUV 14D28 UOPOA2I2YOIT SEIN 89/LT LLTEL ‘ds p1U2D1ÿ201Q2Q 8961 T PULIOA ‘ds SNYdAIAYI] TEIA L9IL6I SLTEL ‘ds D2S140ÿ200) T6 L unsnöny JS SISUOLIDISDEDPDU uopoupjpy) 6I7U 79/181 -IVId-DNHIN ON SOUSEIVY eq MEO] soroads snuay 1SOH ON (oosAIg ‘][09) Jeosesepepy Wor susıqıydure pue sodo Jo sonsesed sopo]so9 ‘7 ATAV IL A NEW PROTEOCEPHALIDEA FROM MADAGASCAR 537 DISCUSSION Glaw & Vences (1994) and Vences et al. (2009) listed about 300 species of reptiles in Madagascar, most of them (92%) being endemic. Up to now, only two proteocephalidean cestodes, Deblocktaenia ventosaloculata (Deblock, Rose & Broussart, 1962), and Ophiotaenia georgievi de Chambrier, Ammann & Scholz, 2010, were described from these reptiles (Freze, 1965; de Chambrier er al., 2010). Ophiotaenia lapata n. sp. is the third species of proteocephalideans and the second member of Ophiotaenia reported from this country. However, it is evident that the richness of cestode fauna of reptiles in Madagascar is poorly known and it is reasonable to expect that it is much higher than indicated by the available data. Samples of Cestodes from reptiles and amphibians collected by G. Brygoo have been deposited in the Natural History Museum in Geneva. Nine endemic hosts were found to be infected with proteocephalidean cestodes (Table 2). The material collected by G. Brygoo between 1961 and 1970 in Madagascar in reptiles and amphibians is particularly rich. A preliminary evaluation of these specimens has indicated that there are probably as many as seven unnamed species and that each of them occurs in only one host species, thus exhibiting a strict specificity (oioxene type sensu Euzet & Combes, 1981) (see Table 2 and de Chambrier et al., 2010). One host, Madagas - carophis colubrinus, probably harbours two different cestode species, i.e. O. lapata and another species of Ophiotaenia (designated as Ophiotaenia sp. 3 in Table 2), which bears a huge apical organ. The taxonomic evaluation has also shown that only few of these seven poten- tially new Ophiotaenia species can be described because they are not well enough pre- served to enable a morphological description. In addition, the specimens were fixed with formalin and thus are unsuitable for phylogenetic studies using molecular data. It is thus obvious that new material should be collected in order to better describe the still hidden diversity of reptilian parasites in Madagascar and to elucidate their phylo- genetic relationships. ACKNOWLEDGEMENTS The authors are indebted to Jeanne Rasamy, Achille Raselimanana, and Steve Goodman for providing laboratory facilities, for organizing field sampling and for identification of the hosts. We also deeply thank Tomas Scholz for fruitful suggestions, André Piuz for providing SEM photomicrographs, Florence Marteau and Gilles Roth (all Geneva) for finalizing the drawings. This study was supported in part by the National Science Foundation PBI award Nos. 0818696 and 0818823. REFERENCES AMMANN, M. & DE CHAMBRIER, A. 2008. Ophiotaenia gilberti sp. n. (Eucestoda: Proteo - cephalidea), a parasite of Thamnodynastes pallidus (Serpentes: Colubridae) from Paraguay. Revue suisse de Zoologie 115: 541-551. BEDDARD, F. E. 1913. Contributions to the anatomy and systematic arrangement of the Cestoidea. VII. On six species of tapeworms from reptiles, belonging to the genus Ichthyotaenia (s.l.). Proceedings of the Zoological Society London, Part 1, 4-36. DE CHAMBRIER, A. 2001. A new tapeworm from the Amazon, Amazotaenia vvettae n. gen., n. sp., (Eucestoda: Proteocephalidea) from the siluriform fishes Brachyplatystoma filamen - tosum and B. vaillanti (Pimelodidae). Revue suisse de Zoologie 108: 303-316. 558 V. R. RAMBELOSON ET AL. DE CHAMBRIER, A. 2006. Redescription of Kapsulotaenia sandgroundi (Carter, 1943), a parasite of Varanus komodoensis (Varanoidea: Varanidae) from Komodo Island, Indonesia. Systematic Parasitology 63: 83-93. DE CHAMBRIER, A., AMMANN, M. & SCHOLZ, T. 2010. First species of Ophiotaenia (Cestoda: Proteocephalidea) from Madagascar: O. georgievi sp. n., a parasite of the endemic snake Leioheterodon geayi (Colubridae). Folia Parasitologica 57, 197-205. DE CHAMBRIER, A., BINH, T. T. & SCHOLZ, T. 2012. Ophiotaenia bungari n. sp. (Cestoda), a parasite of Bungarus fasciatus (Schneider) (Ophidia: Elapidae) from Vietnam, with comments on relative ovarian size as a new and potentially useful diagnostic character for proteocephalidean tapeworms. Systematic Parasitology 81: 39-50. DE CHAMBRIER, A, COQUILLE, S. C. & BROOKS, D. R. 2006. Ophiotaenia bonneti n. sp. (Eucestoda: Proteocephalidea), a parasite of Rana vaillanti (Anura: Ranidae) in Costa Rica. Folia Parasitologica, 53, 125-133. DE CHAMBRIER, S. & DE CHAMBRIER, A. 2010. Two new genera and two new species of proteo- cephalidean tapeworms (Eucestoda) from reptiles and amphibians in Australia. Folia Parasitologica 57: 263-279. DE CHAMBRIER, A. & VAUCHER, C. 1999. Proteocephalidae et Monticelliidae (Eucestoda: Proteocephalidea) parasites de poissons d’eau douce du Paraguay avec descriptions d’un genre nouveau et de dix espéces nouvelles. Revue suisse de Zoologie 106: 165-240. DE CHAMBRIER, A., ZEHNDER, M., VAUCHER, C. & MARIAUX, J. 2004. The evolution of the Proteocephalidea (Platyhelminthes, Eucestoda) based on an enlarged molecular phylo- geny, with comments on their uterine development. Systematic Parasitology 57: 159-171. CHERVY, L. 2009. Unified terminology for cestodes microtriches: a proposal from the International Workshops on Cestode Systematics in 2002-2008. Folia Parasitologica 56: 199-230. COQUILLE, S. & DE CHAMBRIER, A. 2008. Cairaella henrii gen. n., sp. n., a parasite of Norops trachyderma (Polychrotidae), and Ophiotaenia nicoleae sp. n. (Eucestoda: Proteo - cephalidea), a parasite of Thecadactylus rapicauda (Gekkonidae), in Ecuador. Folia Parasitologica 55: 197-206. EUZET, L. & COMBES, C. 1981. Problémes poses par la spécificité parasitaire des cestodes Proteocephalidea et Pseudophyllidea parasites de poissons. Mémoires de la Société Zoologique francaise 40: 239-285. FREZE, V. I. 1965. [Proteocephalata in Fish, Amphibians and Reptiles]. Essentials of Cestodology. Vol. V. Nauka, Moscow, 538 pp. (In Russian; English translation, Israel Program of Scientific Translation, 1969, Cat. No. 1853, v + 597 pp.). FUHRMANN, O. 1924. Two new species of reptilian cestodes. Annals of Tropical Medecine and Parasitology 18: 505-513. GLAW, F. & VENCES, M. 1994. A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlag, Koln. HILMY, LS. 1936. Parasites from Liberia and French Guinea. Part III. Cestodes from Liberia. Publication of the Egyptian University, Faculty of Medecine 9: 1-72. MARSELLA, C.M.V. & DE CHAMBRIER, A. 2008. Ophiotaenia alessandrae sp. n. (Eucestoda: Proteocephalidea), a parasite of Hyla boans (Anura: Hylidae) from Ecuador. Revue suisse de Zoologie 115: 553-563. METTRICK, D. F. 1960. A new cestode, Ophiotaenia ophiodex, n. sp., from a night-adder, Causus rhombeatus (Licht.), in Southern Rhodesia. Proceedings of the Helminthological Society of Washington 27: 275-278. METTRICK, D. F. 1963. Some cestodes of reptiles and amphibians from the Rhodesias. Proceedings of the Zoological Society London 141: 239-250. Oros, M., SCHOLZ, T., HANZELOVA, V. & MACKIEWICK, J. S. 2010. Scolex morphology of mono- zoic cestodes (Caryophyllidea) from the Palaearctic Region: a useful tool for species identification. Folia Parasitologica 57: 37-46. A NEW PROTEOCEPHALIDEA FROM MADAGASCAR 559 Reco, A. A. 1994. Order Proteocephalidea Mola, 1928. In: L.F. KHALIL, A. JONES & R.A. BRAY (Eds.), Keys to the Cestode Parasites of Vertebrates. CAB International, Wallingford, pp. 257,295: RUDIN, E. 1917. Die Ichthyotaenien der Reptilien. Revue suisse de Zoologie 25: 14-381. SANDGROUND, J. H. 1928. Some new cestodes and nematodes parasites from Tanganyika Territory. Proceedings of the Boston Society of Natural History 39: 131-150. SCHMIDT, G. D. 1986. CRC Handbook of Tapeworm Identification. CRC Press, Boca Raton, Florida, 675 pp. SCHOLZ, T. & HANZELOVA, V. 1998. Tapeworms of the genus Proteocephalus Weinland, 1858: Proteocephalidae, parasites of fishes in Europe. Studie AV CR, 1998 (2), 119 pp. SOUTHWELL, T. & LAKE, F. 1939. On a collection of Cestoda from the Belgian Congo. Annals of Tropical Medecine and Parasitology 33: 107-123. VENCES, M., WOLLENBERG, K. C., VIEITES, D. R. & LEES, D. C. 2009. Madagascar as a model region of species diversification. Trends in Ecology and Evolution 24: 456-465. REVUE SUISSE DE ZOOLOGIE 119 (4): 561-570; décembre 2012 Ophiotaenia oumanskyi sp. n. (Eucestoda: Proteocephalidea), a parasite of Lepidobatrachus laevis Budgett, 1899 (Anura: Leptodactylidae) from Paraguay Alain DE CHAMBRIER! & Alicia GIL DE PERTIERRA? | Muséum d’histoire naturelle, PO. Box 6434, CH-1211 Geneva 6, Switzerland. Corresponding author: alain.dechambrier@ville-ge.ch 2 Laboratorio de Helmintologia, Departamento de Biodivesidad y Biologia Experimental, Facultad de Ciencias Exactas y Naturales, Cuidad Universitaria, Int. Güiraldes 2160, Pabellon II, 4 piso, Universidad de Buenos Aires, C1428EGA-Buenos Aires, Argentina. E-mail: helmintho@bg.fcen.uba.ar Ophiotaenia oumanskyi sp. n. (Eucestoda: Proteocephalidea), a parasite of Lepidobatrachus laevis Budgett, 1899 (Anura: Leptodactylidae) from Paraguay. - A new species of Ophiotaenia, O. oumanskyi sp. n., 1s des- cribed from the intestine of the frog, Lepidobatrachus laevis (Anura: Lepto- dactylidae), from Paraguay. Among the 10 species of Ophiotaenia found in anurans of the Neotropical Region, only O. bonariensis Szidat & Soria, 1954 and O. ecuadoriensis Dyer, 1986 possess an apical organ, whereas it is absent in the 8 other species. O. bonariensis differs from O. oumanskyi by the total length of the strobila (400-500 mm versus 50-96 mm) and by the number of testes (120-140 versus 85-119). O. ecuadoriensis differs of O. oumanskyi by the total lenght of strobila (29 mm versus 50-96 mm), by the position of the vagina to cirrus-sac (posterior versus anterior and posterior) and by the diameter of the embryophore (23-26 versus 30). Proteocephalus bufonis Chandra & Gupta, 2007 becomes Proteocephalus chandrae nom. nov. (to avoid homonymy with Proteocephalus bufonis Vigueras, 1942). Proteocephalus chandrae nom. nov. is transferred to Ophiotaenia and becomes Ophiotaenia chandrae n. comb. Keywords: New species - Proteocephalidae - Ophiotaenia chandrae nom. nov. INTRODUCTION The cestodes of the order Proteocephalidea Mola, 1928 are parasites of fresh- water fishes, amphibians, reptiles and marsupials (Schmidt, 1986; Rego, 1994, Cafieda-Guzman et al., 2001), with the highest species richness in pimelodid fishes in the Neotropical Region (de Chambrier & Vaucher, 1999; Rego et al., 1999, de Chambrier et al., 2006). In contrast, amphibians are scarcely represented as the defini- tive hosts of proteocephalidean cestodes (de Chambrier et al., 2006; Marsella & de Chambrier, 2008). During a herpetological survey in Paraguay between 1979 and 2002 conducted by the Geneva Natural History Museum, proteocephalidean tapeworms Manuscript accepted 13.09.2012 562 A. DE CHAMBRIER & A. GIL DE PERTIERRA belonging to Ophiotaenia La Rue, 1911, were found in a leptodactylid frog Lepidobatrachus laevis. Since this cestode differs from all 24 species of the genus described from amphibians in the world, it is described as a new taxon herein. MATERIALS AND METHODS One specimen of Lepidobatrachus laevis Budgett, 1899 examined was killed by immersion in a 1% MS 222 solution (Methanesulfonate salt, Sigma, No A-5040) and immediately dissected. The digestive tract was fixed with hot 4% neutral formalin and subsequently stored in 75% ethanol. Strobila was stained with Mayer’s hydrochloric carmine, dehydrated in a graded series of ethanol, cleared in eugenol (clove oil), and mounted as permanent preparations in Canada balsam. For histology, pieces of strobila were embedded in paraffin wax, transversely sectioned at 12-15 um intervals, stained with Weigert’s hematoxylin and counterstained with 1% eosin B (acidified with five drops of pure acetic acid for 100 ml solution) following recently updated protocols (see de Chambrier, 2001; Oros et al., 2010). Eggs were studied in distilled water. The spe- cimens have been deposited in the helminthological collection of the Natural History Museum, Geneva, Switzerland (PLAT). All measurements are given in micrometres unless otherwise indicated. For two-dimensional measurements, length is given before width. Amphibian classification and authorities follow Amphibian Species of the World 5.5 (Frost, 2011). Abbreviations used in descriptions are as follows: x, mean; n, number of measurements; RSO, ratio of the width of the ovary to the width of the pro- glottis; PGP, position of genital pore expressed as percentage of its position to the pro- glottis length from the anterior margin; RSCS, relative size of the cirrus-sac expressed as percentage of its length to the width of the proglottis; CV, coefficient of variation. Museum abbreviations used are as follows: MHNG, Geneva Natural History Museum, Invertebrate Collection (PLAT), Geneva, Switzerland. RESULTS Ophiotaenia oumanskyi sp. n. Figs. 1-8 TYPE MATERIAL: Holotype MHNG-PLAT-62560, | whole mounted slide. — Paratype 1, MHNG-PLAT-82004, 1 whole mounted slide. — Paratype 2, MHNG-PLAT-82005, 3 whole mounted slides, 10 cross sections. All material is from the type locality and was collected by Carlo Dlouhy 4.02.2002, field number Py 9044. DESCRIPTION (BASED ON THREE ENTIRE SPECIMENS): Proteocephalidae, Proteo- cephalinae. Large-sized worms, 50-96 mm long, up to 1.23 mm wide, flattened dorso- ventrally, with last proglottides elongated. Strobila acraspedote, anapolytic, with about 150 proglottides; 101-138 (x = 125) immature proglottides (up to appearance of sper- matozoa in vas deferens), 5-9 (x = 7) mature proglottides (up to appearance of eggs in uterus), 2-6 (x = 4) pregravid proglottides (up to appearance of hooks in oncospheres); FIGS 1-6 Ophiotaenia oumanskyi sp. n. from Lepidobatrachus laevis. (1) MHNG-PLAT-62560, holotype 1. Scolex, dorsoventral view. (2) MHNG-PLAT-82004, paratype. Cirrus-sac and vagina, dorsal view; note the presence of a vaginal sphincter. (3) MHNG-PLAT-82005, paratype. Mature proglottis, transverse section at posterior part level. (4) MHNG-PLAT-82005, paratype. Mature proglottis, transverse section at ovarian level. (5) Cross-section of gravid proglottis, at level of anterior part (6) MHNG-PLAT-82005, paratype 2. Eggs drawn in distilled water. A NEW PROTEOCEPHALIDEA FROM PARAGUAY 563 Abbreviations: cg = glandular cells, probably of exocrine type, ci = cirrus, cs = cirrus-sac, do = dorsal osmoregulatory canal, em = embryophore, Im = internal longitudinal musculature, In = longitudinal lateral nerves, oe = outer envelope, on = oncosphere, ov = ovary, sc = subtegumental cells; st = subtegumental muscle fibres, te = testes, tg = tegument, ud = uterine diverticula, ut = uterus, va = vas deferens, ve = vaginal canal, vi = vitelline follicles, vo = ventral osmoregulatory canal, vs = vaginal sphincter. Scale-bars: 1, 5 = 250 um; 2 = 100 um, 3-4 = 500 um, 6 = 20 um. 564 A. DE CHAMBRIER & A. GIL DE PERTIERRA 10-17 (x = 13) gravid proglottides. Proliferation zone, 1000-1400 long. Immature proglottides wider than long; and mature, pregravid, gravid proglottides longer than wide. Last gravid proglottides elongated (length: wide ratio 2.3-5.7). Scolex spherical, 350-410 wide, contains numerous cells with granular inclu- sions in the apical region. Apical organ present, 38-56 (x = 43) x 50-65 (x = 54, n= 3), ratio of the width of the apical organ to the width of the scolex 14-17%. Four small uniloculate suckers, 150-170 in diameter (Fig. 1). Internal longitudinal musculature developed (Figs 3-5), forming small anasto- mosed bundles of muscular fibres. Osmoregulatory canals usually situated between vitellaria and testes. Ventral canal rarely overlapping vitellaria. Ventral canals 25-50 in diameter, with secondary canals ending beneath the tegument; dorsal canals 10-15 in diameter (Figs 1, 3-5). Testes medullary, oval, 60-75 (x = 70) x 35-45 (x = 40, n = 21) in diameter, numbering 85-119 (x = 103, n = 21, CV = 8%), in one or two layers, in two lateral fields between anterior margin and preovarian space, reaching to ovary (Figs 4, 7), degenerated in last gravid proglottides. Occasionally, some testes overlap uterine stem. Vas deferens coiled, thin-walled, reaching to midline of proglottis (Figs 5, 7). Cirrus- sac elongate to pyriform, thick-walled, 180-260 long, representing 20-27% (x = 23%, n= 25, CV = 8%) of proglottis width. Cirrus occupying up to 70% of cirrus-sac length (Figs2). Genital ducts passing between osmoregulatory canals. Genital atrium present. Genital pores irregularly alternating, situated at 35-61% (x = 41%, n = 21, CV = 14%) of proglottis length. Vagina posterior (in 52% of the proglottides) or anterior (in 48% of the pro- glottides, n = 46) to cirrus-sac, in proximal part lined with intensely staining cells. Muscular terminal sphincter present, 35-40 in diameter (Figs 2, 7, 8). Mehlis’ glands 70-105 in diameter, 9-13% of proglottis width. Ovary medullary, bilobed, small, with follicles on ventral side 445-620 wide, occupying 61-70% (x = 66%, n = 29, CV = 4%) of proglottis width (Figs 4, 7, 8). Ovary occupying 6.7% of proglottis surface in mature proglottis and 8.3% of pro - glottis surface in gravid proglottis (see Ammann & de Chambrier, 2008 and de Chambrier et al., 2012 for methodology of measuring the ovarian surface). Vitelline follicles medullary, oval to elongate, in two lateral fields, interrupted porally by vagina and cirrus-sac, reaching almost anterior and posterior margins of proglottides, occupying porally 91-97% and aporally 91-96 % of proglottis length, respectively (Figs 3-5, 7, 8). Anlage of uterus medullary, already present in immature proglottides. Uterus with 18-25 (n = 13) dorsolateral diverticula on each side in gravid proglottides (Figs 5, 6). Formation of uterus of type 1 according to de Chambrier et al. (2004): uterine stem with tubular concentration of numerous intensely stained cells and with lumen in last immature and first mature proglottides (Fig. 7). In mature proglottides, thin-walled lateral diverticula appear, with distal part lined with some intensely staining cells. At this stage, uterus occupying up to 22% of proglottis width. In pregravid proglottides, eggs completely filling uterine stem and diverticula that occupy up to 34% of proglottis width. In gravid proglottides, thin-walled digitate diverticula growing laterally, A NEW PROTEOCEPHALIDEA FROM PARAGUAY 565 go 06 ER Hoyer 9 HR ) |) a FIGS 7, 8 Ophiotaenia oumanskyi sp. n. from Lepidobatrachus laevis. (7) MHNG-PLAT-62560, holotype, mature proglottis, dorsal view. (8) MHNG-PLAT-82004, paratype. Gravid proglottis, ventral view. Scale-bars: 7, 8 = 500 um. occupying up to 72% of proglottis width, opening ventrally by several longitudinal apertures. Eggs spherical, with thin, hyaline outer envelope, up to 55 in diameter; inner envelope consisting in two-layered embryophore, with external thick layer, 23-26 in diameter and nucleate irregular envelope, 18-21 in diameter; oncospheres 11-13 in diameter, with 3 pairs of embryonic hooks, 6-8 long (Fig. 6). TYPE LOCALITY: Loma Plata, Filadelfia, Alto Paraguay Province, Paraguay (22°18'S, 68°18'W). TYPE-HOST: Lepidobatrachus laevis Budgett, 1899 (Leptodactylidae). SITE OF INFECTION: Intestine. PREVALENCE: 1/1. INTENSITY: 3 specimens. 566 A. DE CHAMBRIER & A. GIL DE PERTIERRA ETYMOLOGY: The new species is named in honour of Igor Oumansky, Geneva, who facilitated our field work in South America. DIFFERENTIAL DIAGNOSIS: The new species belongs to the genus Ophiotaenia because of the medullary position of gonads, the presence of four simple unilocular suckers and two lateral field testes (Freze, 1965; Schmidt, 1986; Rego, 1994). Off the approximately 96 currently recognized species of Ophiotaenia parasi- tizing reptiles and amphibians (Schmidt, 1986; de Chambrier et al., 2006; Marsella & de Chambrier, 2008; de Chambrier ef al., 2010, 2012), 25 species of Ophiotaenia parasitize amphibians and 10 of them occur in anurans in the Neotropical Region (Caribbean, southeastern Mexico, Central America and South America; Table 1) (Parodi & Widakowich, 1916; Vigueras, 1942; Wolffhügel, 1948; Szidat & Soria, 1954; Flores-Barroeta, 1955; Dyer & Altig, 1977; Dyer, 1986; Puga & Formas, 2005; de Chambrier et al., 2006; Marsella & de Chambrier, 2008). Among the 10 species of Ophiotaenia found in anurans of the Neotropical Region, only O. bonariensis Szidat & Soria, 1954 and O. ecuadoriensis Dyer, 1986 possess an apical organ, whereas it is absent in O. alessandrae Marsella & de Chambrier, 2008; O. bonneti de Chambrier, Coquille & Brooks, 2006; O. bufonis (Vigueras, 1942); O. calamensis Puga & Formas, 2005; O. ceratophryos (Parodi & Widakowich, 1916); O. hernandezi (Flores-Barroeta, 1955); O. noei Wolffhügel, 1948; and O. olseni Dyer & Altig, 1977. The new species resembles O. ecuadorensis in the size of the scolex (350-410 um and 370-450 um, respectively), but differs in the total length (50-96 mm vs. 29 mm), the position of the vagina to the cirrus-sac (anterior and posterior vs. only posterior) and the diameter of oncospheres (23-26 um vs. 30 um). Ophiotaenia oumanskyi is easily differentiated from O. bonariensis because the latter is very large (400-500 mm versus 50-96 mm), has a higher number of testes (120-140 versus 85-119) and the vagina is always anterior to the cirrus-sac versus anterior and posterior (see Table 1). On the basis of the above differences, specimens found in Lepidobatrachus laevis are considered to represent a new species and the name Ophiotaenia oumanskyi sp. n. is proposed to accommodate it. DISCUSSION Ammann & de Chambrier (2008) used for the first time the relative ovarian size (the ratio of the ovarian size in relation to that of the entire proglottis) as a useful character that discriminates all known species of Ophiotaenia parasitizing reptilian hosts in the New World from species of Palaearctic Proteocephalus parasitizing fresh- water fishes. They found that all species of Ophiotaenia possess a very small ovary, with the relative size of the ovary varying between 1.9 and 5.5%, whereas that of Proteocephalus species is much larger (relative size 13.0-19.7%) (see Table 2 in Ammann & de Chambrier, 2008). Later, de Chambrier et al. (2012) calculated this character for all Ophiotaenia spp. from reptilian hosts (66 species) and for all remaining Proteocephalus spp. from freshwater fish hosts (69 species). 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DE CHAMBRIER & A. GIL DE PERTIERRA TABLE 2. Species of Ophiotaenia from amphibians, with data on the relative size of their ovary Species Host Locality Ovary ratio % Ophiotaenia alessandrae Marsella & de Chambrier, 2008 Hypsiboas boans Ecuador 5.6 O. alternans Riser, 1942 Amphiuma tridactylum U.S.A. 4.8 O. amphiumae (Zeliff, 1932) Amphiuma tridactylum U.S.A. 6.8 O. bonariensis Szidat & Soria, 1954 Leptodactylus latrans Argentina 6.9 O. bonneti de Chambrier, Coquille & Brooks, 2006 Lithobates vaillanti Costa Rica 6.9 O. bufonis (Vigueras, 1942) Bufo peltacephalus Cuba Sh). O. calamensis Pugas & Formas, 2005 Telmatobius dankoi Chile 4.5 O. carpathica (Sharpilo, Kornyushin & Lisitsina, 1979) Triturus cristatus Ukraine 8.8 O. ceratophryos (Parodi & Widakowich, 1916) Ceratophrys ornata Argentina 9 O. chandrae n. comb. Duttaphrynus melanostictus India 8.6 O. cryptobranchi La Rue, 1914 Cryptobranchus alleganiensis U.S.A. 5.6 O. ecuadorensis Dyer, 1986 Hyla geographica Ecuador 2 O. filaroides (La Rue, 1909) Ambystoma tigrinum U.S.A. 10.8 O. gracilis Jones, Cheng & Gillespie, 1958 Rana catesbeiana U.S.A. 1.9 O. hernandezi (Flores-Barroeta, 1955) Rana sp. Mexico 10.3 O. loennbergii (Fuhrmann, 1895) Necturus maculosus U.S.A. 3 O. magna Hannum, 1925 Rana catesbeiana U.S.A. 5.4 O. niuginii (Schmidt, 1975) Rana arfarki Papua New Guinea 8.6 O. noei Wolffhugel, 1948 Calyptocephalella gayi Chile 6.6 O. olor (Ingles, 1936) Rana aurora U.S.A. 7.4 O. olseni Dyer & Altig, 1977 Hyla geographica Ecuador 6.5 O. oumanskyi n. sp. Lepidobatrachus laevis Paraguay 6.7 O. ranae Yamaguti, 1938 Rana nigromaculata Japan 8.9 O. saphena Osler, 1931 Rana clamitans Usa 8.4 O. schultzei (Hungerbühler, 1910) Pyxicephalus adspersus South Africa 212 O. tigrina (Woodland, 1925) Hoplobatrachus tigerinus India 8.8 7.2%) (O. schultzei was not considered because the drawings are not suitable for taking reliable data). In species of Ophiotaenia from reptiles from all parts of the World ex- cept Europe, the relative ovarian size is 1.5-6.7% (x = 3.4%; see table 1 in Ammann & de Chambrier, 2008 and table 2 in de Chambrier et al., 2012). Based on the new data and those of de Chambrier et al. (2012), it is possible to distinguish four groups in all spp. of Ophiotaenia and Proteocephalus (161 species): three for Ophiotaenia spp. and one for Proteocephalus spp: (1) 3 Ophiotaenia species, parasites of reptiles from western part of the Palaearctic region, with relative size of ovary 9.1-12.7% (x = 10.3%); (ii) 63 Ophiotaenia species, parasites of reptiles from all regions except for the Palaearctic Region, with relative size of ovary 1.5-6.7% (x = 3.4%); (iii) 25 Ophiotaenia species parasites of amphibians, with relative size of ovary 4.5%-10.8% (x = 7.2%); (iv) 70 Proteocephalus spp, parasites of teleost fishes from all regions, with relative size of ovary 5.4-20.2 (x = 11.9%) (Proteocephalus midoriensis Shimazu, 1990, with relative size of ovary of 28.8%, is not considered because the ovary illustrated does not seem to be of typical shape — see Shimazu, 1990, Fig. 12) (see table 1 in de Chambrier er al., 2012). A NEW PROTEOCEPHALIDEA FROM PARAGUAY 569 These data show that the relative ovarian size of all known Ophiotaenia spp. from anurans is higher than those of reptilian hosts, but the number of species measured remains low and more information is needed. As observed by de Chambrier et al. (2006), proteocephalidean cestodes are rare parasites of amphibians. In the Neotropical region (Costa Rica, Ecuador and Paraguay), these authors found cestodes in 11 of about 200 species of amphibians and prevalence was only 0.4% to 3.0% (de Chambrier et al., 2006). Proteocephalus bufonis Chandra & Gupta, 2007, a parasite of Bufo melano - stictus 1s preoccupied by Proteocephalus bufonis Vigueras, 1942 (Chandra & Gupta, 2007; Vigueras, 1942). Due to this homonymy, we propose Proteocephalus chandrae nom. nov. for P bufonis Chandra & Gupta, 2007. Furthermore, this species shows the characters of Ophiotaenia, such as mature and gravid proglottides being markedly- longer than wide (see Freze, 1965) and thus is transferred to Ophiotaenia as Ophiotaenia chandrae n. comb. ACKNOWLEDGEMENTS The authors are indebted to Carlo Dlouhy (Asuncion, Paraguay), who collected the specimens, two anonymous referees for fruitful suggestions and to Tomas Scholz (Institute of Parasitology, Ceské Budéjovice, Czech Republic) for helpful discussions. We are also grateful to Florence Marteau and Gilles Roth (Geneva) for their help with drawings. This project was supported in part by the National Science Foundation PBI award Nos. 0818696 and 0818823 and by the Universidad de Buenos Aires, Argentina (Grant UBACyT 20020090200511 and 20020090200529). REFERENCES AMMANN, M. & DE CHAMBRIER, A. 2008. Ophiotaenia gilberti sp. n. (Eucestoda: Proteocephalidea), a parasite of Thamnodynastes pallidus (Serpentes: Colubridae) from Paraguay. 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(Amphibia, Leptodactylidae) de la Republica Argentina. Comunicaciones del Instituto Nacional de Investigacion de las Ciencias Naturales Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia” 2: 189-210. VIGUERAS, I. P. 1942. Proteocephalus bufonis n. sp. (Cestoda), parasitö del intestino de Bufo peltacephalus (Amphibia). Notas helmintologicas 5: 208-221. WOLFFHUGEL, K. 1948. Ophiotaenia noei n. sp. (Cestodae). Biologica 5: 15-27. REVUE SUISSE DE ZOOLOGIE 119 (4): 571; décembre 2012 Corrigendum: Draconectes narinosus, a new genus and species of cave fish from an island of Halong Bay, Vietnam (Teleostei: Nemacheilidae) by M. KOTTELAT, Revue suisse de Zoologie 119 (3): 341-349 Due to a technical error, the figure 1 of the article of M. Kottelat was printed with a too low resolution. The correct photo is reprinted below. The editorial committee apologises for this unfortunate event. Fic. 1 Draconectes narinosus, MHNG 2730.080, 24.7 mm SL; Vietnam: Ha Long Bay: Dao Van Gio island, Dong Duc Tien cave; right side, reversed. Photographs by Tan Heok Hui. REVUE SUISSE DE ZOOLOGIE Tome 119 — Fascicule 4 SALARZEHI, S., HAJIQANBAR, H., OLYAIE TORSHIZ, A. & NOEI, J. Description of a new species of the genus Fessonia (Acari: Prostiemata; Smarididae) from Tran. <....- «6 sends een 0 0 PALACIOS- VARGAS, J. G. & BOCANEGRA, T. A new species of Cerato- physella from Peru (Collembola: Hypogastruridae) ............. ARREDONDO, N. J. & GIL DE PERTIERRA, A. A. A new species of Neo - echinorynchus (Eoacanthocephala: Neoechinorhynchidae) from Pachyurus bonariensis (Perciformes: Sciaenidae) from the Parana River basin in Argentina, with comments on two other species of the CERCA CRT PNR E ae ee ee VL ee at AE SCHATTI, B., KUCHARZEWSKI, C., MASROOR, R. & RASTEGAR POUYANI, E. Platyceps karelini (Brandt, 1838) from Iran to Pakistan and revali- dation of Coluber chesneii Martin, 1838 (Reptilia: Squamata: Folubrinae 29%. Eee ehhh and ew) DWM rei BLANT, M., MARCHESI, P., DESCOMBES, M. & Capt, S. Nouvelles données sur la répartition de la souris des moissons (Micromys minutus) en Suisse occidentale et implications pour la gestion de son habitat . . . SENGLET, A. Civizelotes new genus, and other new or little known Zelotinae (Araneae, Gnaphosidae) .......................... CLOUSE, R. M. & SCHWENDINGER, P. J. Leptopsalis foveolata sp. n. a new species of Stylocellidae from Thailand that displays a novel morpho- logical feature in the suborder Cyphophtalmi (Arachnida, Opiliones) RAMBELOSON, V. R., RANAIVOSON, H. C. & de CHAMBRIER, A. Ophiotenia lapata sp. n. (Eucestoda: Proteocephalidea) from Madagascar: a parasite of the endemic snake Madagascarophis colubrinus (CO ou. otk at RO turk to OS iG ees eta aes nt DE CHAMBRIER, A. & GIL DE PERTIERRA, A. Ophiotaenia oumanskyi sp. n. (Eucestoda: Proteocephalidea), a parasite of Lepidobatrachus laevis Budgett, 1899 (Anura: Leptodactylidae) from Paraguay.......... KOTTELAT, M. Corrigendum: Draconectes narinosus, a new genus and species of cave fish from an island of Halong Bay, Vietnam Cleleoster INémachellidae):..hu ic awa eee asian Pages 409-415 417-423 425-439 441-483 485-500 501-528 529-546 547-559 561-570 371 REVUE SUISSE DE ZOOLOGIE Volume 119 — Number 4 SALARZEHI, S., HAJIQANBAR, H., OLYAIE TORSHIZ, A. & NOEI, J. Description of a new species of the genus Fessonia (Acari: Prostigmata: Smarididae) from Iran. «.. 2... 0s ba eens oe on PALACIOS- VARGAS, J. G. & BOCANEGRA, T. A new species of Cerato - physella from Peru (Collembola: Hypogastruridae)...... ARREDONDO, N. J. & GIL DE PERTIERRA, A. A. A new species of Neo - echinorynchus (Eoacanthocephala: Neoechinorhynchidae) from Pachyurus bonariensis (Perciformes: Sciaenidae) from the Parana River basin in Argentina, with comments on two other species of the GEMS ee a ete © gt oe ak enna nee Oe a SCHATTI, B., KUCHARZEWSKI, C., MASROOR, R. & RASTEGAR POUYANI, E. Platyceps karelini (Brandt, 1838) from Iran to Pakistan and revali - dation of Coluber chesneii Martin, 1838 (Reptilia: Squamata: COUDRE BLANT, M., MARCHESI, P., DESCOMBES, M. & CAPT, S. New data on the distribution of the Harvest mouse (Micromys minutus) in western Switzerland and conclusions for the management of its habitat... .. SENGLET, A. Civizelotes new genus, and other new or little known Zelotinae (Araneae, Gnaphosidae) .......................... CLOUSE, R. M. & SCHWENDINGER, P. J. Leptopsalis foveolata sp. n. a new species of Stylocellidae from Thailand that displays a novel morpho- logical feature in the suborder Cyphophtalmi (Arachnida, Opiliones) RAMBELOSON, V. R., RANAIVOSON, H. C. & de CHAMBRIER, A. Ophiotenia lapata sp. n. (Eucestoda: Proteocephalidea) from Madagascar: a parasite of the endemic snake Madagascarophis colubrinus (COAST cette ae eens ren ree nee ete cen DE CHAMBRIER, A. & GIL DE PERTIERRA, A. Ophiotaenia oumanskyi sp. n. (Eucestoda: Proteocephalidea), a parasite of Lepidobatrachus laevis Budgett, 1899 (Anura: Leptodactylidae) from Paraguay.......... KOTTELAT, M. Corrigendum: Draconectes narinosus, a new genus and species of cave fish from an island of Halong Bay, Vietnam (leleostei: Nemachellidae)en u ze a aaa cane eee an el Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 409-415 417-423 425-439 441-483 485-500 501-528 529-546 547-559 561-570 SH PUBLICATIONS DU MUSEUM D’ HISTOIRE NATURELLE DE GENE VE CATALOGUE DES INVERTEBRES DE LA SUISSE, NOS 1-17 (1908-1926) ........ serie Fr. (prix des fascicules sur demande) REVUE DE PALEOBIOLOGIE ......................... Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris) ....... par fascicule Fr. THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION, WITH KEYS FOR DETERMINATION TINOSER SA De | ee a EEE See rie CLASSIFICATION OF THE DIPLOPODA REA MIN ADI nnt ea Fr. LES OISEAUX NICHEURS DU CANTON DE GENÈVE P. GÉROUDET, C. GUEX & M. MAIRE 3>p.-00mbreuses cartes el Heures 108. ue Ae RR Fr. CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ À LA CONNAISSANCE DES ECHINODERMES ACTUELS NERO 0 D AIS a a Sate Be a ee ae ta como de Fr. RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) Y INET, 1 WUEST oc KR: MAREDA:062/p., 1991 une. nun Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: SHELLS AND RADULAS Y. FINEr, Jo WUBST &K. MAREDA, 1992 usa as das aa an ders Fr. O. SCHMIDT SPONGE CATALOGUE R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 p., 1992 ...................... Es ATLAS DE REPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE Po KELLER, Y OABULEN VE MAHNERT AED 19935 ona tu we tas dete anne Fr, THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST HAINE PG DES ae cuales aan aur Gal ho Sate Dies rs Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE oC ACAIMME Za) POO pipi Si ss de CHa Ss Sime neo eee ces Fr. PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... Fr. 285.— 35.— 35.— 30.— 30.— 45.— 30.— 40.— 30.— 22.— 160.— INSTRUMENTA BIODIVERSITATIS CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) (Instrumenta Biodiversitatis I), 1. LOBL, xii + 190 p., 1997 .................:.... Fr. CATALOGUE SYNONY MIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis II), H. G. DIRICKx, x +187 p., 1998 ................. Fr. A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis III), S. BOWESTEAD, 203 p., 1999 . ........ 2222222... Fr. THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE SOKOTRA ARCHIPELAGO (Instrumenta Biodiversitatis IV), B. SCHÄTTI & A. DESVOIGNES, RP nn Fr. PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY (Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS, xx As 2002 OA Er, REVISION DER PALÄARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) (Instrumenta Biodiversitatis VI), G. SABELLA, CH. BÜCKLE, V. BRACHAT SC BESUCHET; Vi 285.05 2004 ct a ee tee Meee eee ee ee. Bi; PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID FLIES (DIPTERA, TEPHRITOIDEA) Proceedings of the “3rd Tephritoid Taxonomist’s Meeting, Geneva, 19.-24. July 2004” (Instrumenta Biodiversitatis VII). B. MERZ, vi + 274 p., 2006 .................... Fe: LISTE ANNOTEE DES INSECTES (INSECTA) DU CANTON DE GENEVE (Instrumenta Biodiversitatis VIII). (ed. B. MERZ), 532 p., 2012................... Fr. 50.— 50.— 60.— 70.— 180.— 100.— 100.— 85.— Mf Volume 119 - Number 4 - 2012 Revue suisse de Zoologie: Instructions for Authors The Revue suisse de Zoologie publishes original results of zoological systematics and related fields. 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All correspondence should be addressed to: Revue suisse de Zoologie, Museum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: rsz.mhn@ville-ge.ch Home page RSZ: http://www. ville-ge.ch/mhng/publication03.php SALARZEHI, S., HAJIQANBAR, Description of a new Prostigmata: Smarid PALACIOS-VARGAS, J. G. physella from Peru ( River basin in Arg penus SCHATTI, B., KUCHARZE Platyceps karelini | Colubrinas dedi BLANT, M., Muti Sore Proteoce species of cave (Teleostei: Nemache Indexed in CURRENT CONTE N SMITHSONIAN INSTITUTION LIBRARIES 3 9088 01699 5300