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Contributions to Zoology, 72 (4) 195-209 (2003) Contributions to Zoology, 72 (4) 195-209 (2003) SPB Academic Publishing bv, The Hague Variation in number of trunk vertebrae and in of costal in count grooves salamanders of the family Hynobiidae S.N. Litvinchuk¹ & L.J. Borkin² Institute Russian 194064 St. 1 of Cytology, Academy of Sciences, Tikhoretsky pr. 4, Petersburg, Russia, e- 2 mail: [email protected]; Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia, e-mail: [email protected] Keywords: Hynobiidae, trunk vertebrae, costal grooves Abstract Introduction Ten from five of the Hynobiidae were modem tailed species genera family Among amphibians, the family Hyno- studied. The number of trunk vertebrae varied between 14 and biidae (about 40 species), distributed predominantly and the of costal from 10 15. Both 21, count grooves ranged to in temperate Asia, is an early diverged branch, which the within-species variation and the within-population varia- retains characters Duellman tion the values of many primitive (e.g., were recorded in some species. In both kinds the coefficient of low. Salamandrella & Larson Based variation were quite In Trueb, 1986; & Dimmick, 1993). the south-eastern differed from keyserlingii, samples markedly on recent studies (Fei & Ye, 2000a; Fu et al., 2001), remaining ones. the the genus Onychodactylus Among hynobiids, ten of the approximately genera hynobiids could number (both species) and Batrachuperus mustersi have higher be The genus is the richest ofvertebrae in the anterior oftrunk and recognized. Hynobius part (5 4, respectively, one and consists of about 22 which versus 3), and, thus, demonstrated a distinct position. The rela- species, may tion between the number of trunk vertebrae and the count of be arranged in three subgenera (Matsui et al., 1992; costal studied. The variation in number of trunk grooves was Borkin, 1999), namely Hynobius (about 14 species), vertebrae across urodelan families was discussed. Pseudosalamandra and Satohius (7 species) (1 spe- cies). The family also includes the genera Liua Content (1 species), Onychodactylus (2 species), Pachy- hynobius (1 species), Protohynobius (1 species), Introduction 195 Pseudohynobius (1 species), Ranodon (2 species), Materials and methods 196 Salamandrella (1 species, which penetrates to east- Results 196 ern Europe), and Batrachuperus sensu lato (about Number of trunk vertebrae 196 9 The taxonomic Patterns of trunk vertebrae composition 199 species). positions of Ranodon, Count of costal 200 and grooves Liua, Pseudohynobius, Batrachuperus were Discussion 201 discussed in recent papers (Fei & Ye, 2000b; Fu et Number oftrunk vertebrae 201 al., 2001; Kuzmin & Thiesmeier, 2001). Within-population variation 201 Based on mitochondrial DNA (Fu et Sexual dimorphisms 202 study al., it found that the Geographic variation 202 2001), was genus Batrachuperus Comparison between the hynobiids and other families 204 be into distinct could split two geographic groups, Patterns of trunk vertebrae composition 204 probably, of generic level. The eastern (Chinese) Count of 205 costal grooves to stricto group belongs Batrachuperus sensu and Correlating number of trunk vertebraewith count of 205 consists of six & Ye, 2000b; et costal grooves species (Fei Song Taxonomic 205 The value al., 2001). western (Middle East) group con- Concluding remarks 206 of three sists two or species. They may be allo- Acknowledgements 206 cated to Paradactylodon. References 206 Hynobiids were and are a favorite model for the Appendix 209 Downloaded from Brill.com10/01/2021 08:55:57AM via free access 196 S.N. Litvinchuk & L.J. Borkin - Variation in Hynobiid salamanders lower study on evolutionary morphology and tetra- Materials and methods pod phytogeny (e.g., Schmalhausen, 1964; Voro- byeva, 1994). Special attention was paid to the study A total of 635 adult and subadult specimens be- of the morphogenesis of vertebral column (Schmal- longing to ten hynobiid species were studied. The hausen, 1957, 1958a, 1958b, 1964; Borchwardt, materials studied are kept at the collections of the 1974). However, as yet, the data about the number Department of Herpetology, Zoological Institute, of in of trunk vertebrae hynobiids are quite scarce. In Russian Academy Sciences, St. Petersburg his comprehensive description of osteology of the (n=265), Department of Vertebrate Zoology, St. Pe- Japanese clawed salamander, Onychodactylus ja- tersburg State University, St. Petersburg (n=171), ponicus, Okajima (1908) mentioned the position and Zoological Museum, Moscow State University, of the sacral vertebra in vertebral column. Later, Moscow (n=199). Hilton (1948) & Teege (1957) provided the infor- The number of trunk vertebrae (or “dorsal ver- mation about vertebral morphology for four and tebrae” according to Mivart, 1870) was accounted three of the number of of species hynobiid salamanders, respec- by the spinal processes by means data of muscles. The tively, including O. japonicus. However, even dissection dorsal position of the based on the same (latter) species were conflict- sacrum was identified by feeling of sacral ribs on ing. Antipcnkova (1994) published a short review the left side of the body. in on number and morphology of vertebrae the Unfortunately, various authors used different tech- Siberian Salamandrella of costal under salamander, keyserlingii. niques grooves counting. Moreover, Some other data were also published by Deynegi the same name they understood different cases. So, (1917), Antipcnkova (1982), Zhang (1985), Nambu Dunn (1923) has applied the name “costal grooves” & Fei & Ye all lateral situated between the (1991), (2000a). to body grooves since the late number of fore- and the Nevertheless, 1950s, hindlimb, probably, including grooves, trunk vertebrae has been using in systematics of which touched the posterior margin of the base of urodelans. devoted hindlimbs. the Some papers were to study geo- According to Highton (1957), cos- in graphic variation various salamanders, mostly tal grooves are such grooves, which are situated plethodontids, proteids and salamandrids (Brodie, between the posterior margin of the head and the 1961; Highton, 1962; Worthington & Wake, 1972; anterior margin of the hindlimb. Misawa (1989) Sket & Arntzen, 1994; Arntzen & Wallis, 1999; has proposed to recognize under the name “costal Litvinchuk lateral & Borkin, 2000). grooves” only grooves between the fore- and As is known, urodelans, at least, at the larval hindlimbs, which didnot touch limb’sposterior and stage, have external body segmentation expressed anteriormargins, respectively. We followed Misawa. the classic by so-called costal grooves. Since mono- However, we counted the number of costal grooves graph about the family Hynobiidae (Dunn, on the left on the body side only (Misawa counted count of such became taxo- both grooves an important on sides). According to Cope (1889), the nomic character used for identification of species grooves contacting the base of forelimbs should (e.g., Chang, 1936; Thorn, 1969; Thorn & Raffaelli, be named “axillary”, whereas that contacting the 2001). The question arises whether a correlation base of hind limbs “inguinal”, respectively. between the number of trunk vertebrae and count ofcostal grooves exists. Some authors have analysed the problem based on plethodontids and Results ambystomatids (e.g., Highton, 1957; Jockusch, 1997). However, hynobiids are still markedly less Number of trunk vertebrae studied. of the is to evaluate the The goal present paper The number of trunk vertebrae in the family Hyno- in of both trunk vertebraeand costal variation amount biidae varied between 14 and 21 (Table 1). The of the grooves among representatives family Hyno- lowest number was recorded in Ranodon sibiricus, biidae. whereas the in highest count was Onychodactylus Downloaded from Brill.com10/01/2021 08:55:57AM via free access Contributions to Zoology, 72 (4) - 2003 197 Table I. Variation in number of trunk vertebrae and in ofcostal in count grooves ten hynobiid species Trunk vertebrae Costal grooves Species n mean SD min max mean SDSD min max Batrachuperus mustersi 6 17.3 0.50.5 17 18 11.811.8 0.40.4 IIii 1212 - - Batrachuperus pinchoniipinchonii 2 15.5 - 15 1616 10.5 - 10 II11 HynobiusHynohius leechii 2 16.0 -- 16 16 11.5 - 111! 1212 - - Hynobius nigrescens 2 15.0 - 15 1515 11.0 - 11 1111 Hynobius nebulosus 1 16.0 -- 16 1616 12.0 -- 12 12 Hynobius naevis 3 16.7 0.6 16 1717 12.0 0 12 12 Onychodactylus fischeri 33 20.2 0.6 19 21 14.2 0.6 13 15 Onychodactylus japonicus 5 17.8 0.4 1717 1818 11.8 0.4 II 12 Ranodon sibiricus 77 15.1 0.3 14 16 10.9 0.4 10 12 Salamandrellakeyserlingiikeyserlingii 504 16.9 0.5 1515 19 12.0 0.5 10 14 “n” is number of specimens examined; “SD” is the standard deviation. ofcostal Table 2. Distribution ofnumber of trunk vertebrae and count grooves in nine hynobiid species Trunk vertebrae Costal (%)(%) grooves (%) Species n 14 15 16 17 18 19 20 , 21 10 11 12 13 14 15 - Batrachuperus mustersi 6 67 33 ' 17 83 Batrachuperus pinchonii 2 50 50 50 50 Hynobius leechii 2 100 50 50 Hynobius nigrescens 2 100 100 Hynobius nebulosus 1 100 100100 Hynobius naevis 3 33 67 100100 Onychodactylus fischeriflscheri 33 9 6767 24 9 67 24 Onychodactylus japonicus 5 20 80 20 80 Ranodon sibiricus 77 1 89 10 13 84 3 Salamandrella keyserlingii 504 2 13 78 6 0.2 1 11 75 13 0.2 The modal classes are in bold. of trunk vertebrae as well. The of fischeri. The modal number was majority species displayed sym- 15 in Ranodon sibiricus (and, probably, Hynobius metrical distributionof numbers of trunk vertebrae, i.e. less and than nigrescens), 16 in Hynobius leechii (and, probably, more the modal class (Table 2). Hynobius nebulosus), 17 in Salamandrella keyser- Geographic variation was studied in two species, lingii, Hynobius naevis and Batrachuperus mustersi, namely Salamandrella keyserlingii and Ranodon Within and 18 in Onychodactylus japonicus. Onychodac- sibiricus. the formerspecies, the salamanders the tylus fischeri was characterized by the higher modal from majority of samples had, as a rule, 17 trunk with the between number, which was equal to 20.
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