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Atti Del Civico Museo Di Storia Naturale Di Trieste ISSN: 0365–1576 ATTI DEL CIVICO MUSEO DI STORIA NATURALE DI TRIESTE VOL. 54 – 2009 TRIESTE 2009 Atti Mus. Civ. Stor. Nat. Trieste 54 2009 3-114 aprile 2010 ISSN: 0365-1576 VERTEBRAL NUMBERS IN THE CAUDATA OF THE WESTERN PALAEARTIC (Amphibia) BENEDETTO LANZA*, JAN WILLEM ARNTZEN** & ELISABETTA GENTILE*** * Museo di Storia Naturale, Sezione Zoologica “La Specola” & Dipartimento di Biologia Animale e Genetica dell’Università degli Studi di Firenze, Via Romana 17, 50125 Firenze, Italy; e-mail: [email protected] ** Natio nal Museum of Natural History, Naturalis, Zoology Research Department, P. O. Box 9517, 2300 RA Leiden, The Netherlands; e-mail: [email protected] ***Istituto per lo Studio e la Prevenzione Oncologica, Viale Alessandro Volta 171, 50131 Firenze, Italy; e-mail: [email protected] Abstract – We present data on the vertebral number in 3877 specimens of 50 Caudata species occurring in the western Palaearctic. Data were collected through de novo X-ray photography and assembled from the literature. Key words: vertebral number, Caudata, western Palaearctic. Riassunto – Il numero di vertebre dei Caudata della Regione Paleartica occidentale (Amphibia) – Si è studiato il numero di vertebre di 50 specie di Anfi bi Urodeli presenti nella parte occidentale della Regione Paleartica. I dati presentati, basati sullo studio radiografi co di 3877 esemplari e su quelli riportati dalla letteratura, sono stati sintetizzati nelle tabelle. Parole chiave: numero vertebrale, Caudata, Regione Paleartica occidentale. 1. – Introduction Our work on vertebral counts in the salamander genus Speleomantes (Lanza et al., 1995) and in the Triturus cristatus superspecies (Lanza et al., 1991; Arntzen & Wallis, 1994, 1999) through radiographic examination yielded interesting patterns of variation. We here extend our survey to all Caudata of the western Palaearctic, the Iberian Calotriton arnoldi Carranza & Amat, 2005 and the Iranian Batrachu- perus gorganensis Clergue-Gazeau & Thorn, 1978 excluded, and report data on preserved material from 21 collections in mostly European museums, as well as on a few living specimens. The evolutionary implications of our fi ndings will be dealt with in a fortcoming paper. 2. – Material and methods The vertebral column of Caudata consists of: 1 atlas, that normally lacks ribs (cervical region), 10 to over 60 trunk vertebrae, all usually provided with ribs but sometimes lacking in the last one (trunk region), typically a single (two in Sirenidae and in rare teratological individuals of other Caudata) rib-bearing sacral vertebra (sacral region), 2-4 usually ribless sacro-caudal vertebrae (caudo-sacral region), and 20 to more than 100 ribless caudal vertebrae (caudal region). This study reports on vertebral numbers in 3877 specimens of salamanders representing 77 taxa and 4 BENEDETTO LANZA, JAN WILLEM ARNTZEN & ELISABETTA GENTILE 50 species from the western Palaearctic. Species considered have at least part of their range in Europe (Europe defi ned to reach 60°E longitude), while some ones come from Turkey, Iran, Lebanon and Israel. The radiographs were made by EG and BL with an «C.G.R. 500 T» apparatus (mammograph, using focus 0.1-0.3 mm, 23-25 kV, 120-200 mA/sec and Kodak De- fi nix or 3 M Type S (for non-screen mammography) plates, at a distance of 55 cm. The radiographs by JWA were taken with an «Elinax 90/20» and an exposure of 0.7 seconds at 38 kV, 4 mA on Agfa-Gevaert D10DW X-ray fi lm. Several radiographs of Salamandra atra aurorae and Batrachuperus persicus were taken in the ZFMK and MVZ, respectively. In the data list presented below, the collection number carried by the particular specimen is followed by the number of trunk and postsacral vertebrae. Unless other- wise stated, the specimens studied are adults. Missing data are indicated by «?». The posterior trunk vertebrae may either have very small ribs that are not easy to detect radiographically or no ribs at all: the trunk vertebrae counts therefore refer to the centra and not to pairs of ribs. Also, ribs more often than centrae show biological anomalies, or are radiographically invisible due to limited ossifi cation. The total vertebral number is obtained from the number of trunk vertebrae plus 2, for the atlas and the sacrum, respectively. Data from the literature have been adjusted to only include the trunk vertebrae. The number of postsacral vertebrae (sacro-caudal plus caudal vertebrae) includes the smallest radiographically detectable apical ele- ments and is to be regarded as an approximation of the real number. Most postsacral vertebrae were counted by BL. The topographical data are shown with maximum known precision. The iden- tifi cation number of caves refer to the regional speleological cadastres. Coordinates for the French caves are that of Lambert. For Italian caves the longitude is that of Monte Mario, Rome, followed by that of Greenwich. The Rome longitudes are particularly useful in combination with the Italian topographic maps (25,000) pu- blished by the Istituto Geografi co Militare of Florence. Distances are «as the crow fl ies», unless stated otherwise. Dates are given in day-month-year format, as e.g. 31.XII.2005. The following abbreviations and collection acronyms have been used: BMNH = The Natural History Museum, London; don. = donavit or donaverunt; KOC = Kurtuluş Olgun Collection (Adnan Menderes University, Biological Department, Aydın, Turkey); leg. = legit or legerunt; m = arithmetic mean; MBC = Marco Bolo- gna Collection (Rome; Italy; presented to the MDBR); MDBR = Museo del Dipar- timento di Biologia Animale e dell’Uomo, Università “La Sapienza” (Rome, Italy); MHNG = Muséum d’Histoire Naturelle (Genève, Switzerland); MNÇN = Museo Nacional de Çiencias Naturales (Madrid, Spain); MSNG = Museo Civico di Storia Naturale “Giacomo Doria” (Genoa, Italy); MSNT = Museo Regionale di Scienze Naturali (Turin, Italy); Mun. = Municipality; MVZ = Museum of Vertebrate Zoolo- gy, University of California at Berkeley (Berkeley, USA); MZUC = Museo Zoolo- gico dell’Università (Cagliari, Italy); MZUF = Museo di Storia Naturale, Sezione Zoologica “La Specola”, Università degli Studi (Florence, Italy); n = number of VERTEBRAL NUMBERS IN THE CAUDATA OF THE WESTERN PALAEARTIC (Amphibia) 5 specimens studied; NHCL = New Herpetological Collection Lanza (Florence, Italy; donated to the MZUF); NMS = Natur-Museum Senckenberg (Frankfurt am Main, Germany); NMW = Naturhistorisches Museum Wien (Vienna, Austria); OBL (so- metimes quoted as DBUL) = Oddelek za Biologijo, Biotehnişka Fakulteta, Univerza v Ljubljani (Department of Biology, Biotechnical Faculty, University of Ljubljana, Slovenia); pov. = number of postsacral vertebrae, i.e. number of sacro-caudal + cau- dal vertebrae, inclusive of the smallest, radiographically detectable apical elements; Prov. = Province; reg. = regenerated tail; RMNH = National Museum of Natural History-Naturalis (formerly Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands); s = standard deviation; spec. = specimen(s); SVC = Stefano Vanni Collection (Florence, Italy; now in the MSNT); TAU = Tel Aviv University (Israel); trv. = number of trunk vertebrae, i.e. number of presacral vertebrae, atlas excluded; uns. = unsexed specimen(s); ZDEU = Zoological Department of the Ege University, Izmir (Turkey); ZFMK = Zoologisches Forschungsinstitut und Museum Alexander Koenig (Bonn, Germany); ZMA = Zoölogisch Museum Amsterdam (Instituut voor Taxonomische Zoölogie, Universiteit van Amsterdam, The Netherlands). Differences in character state distributions between males and females were tested for signifi cance with the G-test of Independence, with Williams’ correction. This was done by splitting the character state distribution observed across the spe- cies in two approximately equally sized groups, with low and high scores respec- tively and cross-tabulate this over males and females. This median G-test has one degree of freedom. As vertebral counts of males and females are sometimes signi- fi cantly distinct, we have pooled the relevant data only when we could not fi nd any signifi cant statistical sexual difference. 3. – Results Cryptobranchoidea Dunn, 1922 Family Hynobiidae Cope, 1860 Genus Batrachuperus Boulenger, 1878 Batrachuperus persicus Eiselt & Steiner, 1970 (4 spec.: 1 larva, 2 ♂♂, 1 ♀) Iran. Prov. Gilan/Rasht. Near Asalem (= Assalem, Navrud; 37°42’N-48°57’E), ca. 800 m. Leg. Steiner H.M.; 25.VII:1968; 1 larva 19435/3 (17-?) NMW. Iran. Prov Mazanderan. Shirabad Cave, 5 km SE of Shirabad and 60 km E of Gorgān (Gorgān = 36°50’N-54°29E), 310 m. Leg. Theodore J.Papenfus; 13.X.1998. 3 spec.: 1 juv. ♂ MVZ 233501 (17-41), 1 ♂ MVZ 241494 (17-44), 1 juv. ♀ MVZ 238502 (16-?). Data from the literature. LANZA et al. (2005c): trv. larvae (n = 1) 17, ♂♂ (n = 6 BENEDETTO LANZA, JAN WILLEM ARNTZEN & ELISABETTA GENTILE 2) 17, ♀♀ (n = 1) 16; larva + ♂♂ + ♀♀ (n = 4) 16 (25%) and 17 (75%); pov. ♂♂ (n = 2) 41 and 44 (both 50%); see also their Table p. 81. Our data: see above and Table I. Genus Salamandrella Dybowsky, 1870 Salamandrella keyserlingii Dybowsky, 1870 (6 spec.: 2 ♂♂, 4 ♀♀) Russia. Jakutia. Yakutsk, 62°13’N-129°49’E, ca. 30 km N of Yakutsk. Leg. Joger U.; IX.1992; 2 ♂♂ 22155 (17-28), 22156 (18-reg.) MZUF (ex 2409-2410 NHCL); 4 ♀♀ 22157 (17-reg.), 22158 (17-25), 22159 (17-reg.), 22160 (17-28) MZUF (ex 2411-2414 NHCL). Data from the literature. ANTIPENKOVA (1995): usually 17 trv. BORKIN (1999): usually 17, sometimes 16 trv. [the lower count could correspond to Sala- mandrella tridactyla, Nikolsky, 1905 (LANZA et al., 2005a)]; 18-32 pov. (the lower count almost surely refer to juveniles and/or animals with reg. tail). LITVINCHUK & BORKIN (2003) (n = 504): 15-19 (m = 16.9 ± 0.5) [the lower count possibly corresponds to Salamandrella tridactyla, Nikolsky, 1905 (LANZA et al., 2005a)]. Our data. Trv. ♂♂ (n = 2) 17 and 18, ♀♀ (n = 4) always 17 (m = 17.2); ♂♂ + ♀♀ (n = 6): 17 (83.3%) and 18 (16.7%); see Table I. Pov. 28 in a ♂, 25 and 28 in 2 ♀♀; ♂♂+♀♀ (n = 3) 25 (33.3%) and 28 (66.7%); see Table I. Salamandroidea Fitzinger, 1826 Family Proteidae Gray, 1825 Genus Proteus Laurenti, 1768 Proteus anguinus Laurenti, 1768 (103 spec.: 47 ♂♂, 37 ♀, 19 uns.) Proteus anguinus anguinus Laurenti, 1768 (99 spec.: 46 ♂♂, 36 ♀♀, 17 uns.) Croatia.
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