Euproctus, GENE 1839

Carranza, Salvador & Amat, Felix, 2005, Taxonomy, biogeography and evolution of Euproctus (Amphibia: Salamandridae), with the resurrection of the genus Calotriton and the description of a new endemic species from the Iberian Peninsula, Zoological Journal of the Linnean Society 145 (4), pp. 555-582 : 567-571

GENUS EUPROCTUS GENÈ 1839 NOMEN PROTECTUM

(SEE REMARKS BELOW)

Euproctus Genè, 1839: 281 .

Type species: Euproctus rusconii Genè by monotypy.

Megapterna Savi, 1838: 211 . nomen oblitum (see remarks below). Type species Megapterna montana Savi by monotypy.

Phatnomatorhina Bibron, 1839: 131 . Type species Phatnomatorhina platycephalus Bibron by monotypy. Pelonectes Fitzinger, 1843: 33 . Type species Peleonectes platycephalus Fitzinger by monotypy.

Remarks

Although the genus Euproctus Genè, 1839 was published a year later than Megapterna Savi, 1838 , the latter was only a valid name for two years (synonymized by Bonaparte, 1840 2: 405). Therefore, according to Article 23.9.1 of the International Code of Zoological Nomenclature, we consider Megapterna Savi a nomen oblitum because this name has not been used as a valid name after 1899 (Art. 23.9.1.1 of the ICZN) and we consider Euproctus Genè a nomen protectum because it has been mentioned in at least 25 works in the last 50 years by a minimum of ten different authors (Art. 23.9.1.2 of the ICZN). As a result of this nomenclatorial act, Euproctus should be used to name the Tyrrhenian brook newts E. montanus and E. platycephalus . In compliance with Art. 23.9.1.2 of the ICZN the following list of works where Euproctus Genè has been used in the last 50 years is provided: Accordi et al. (1984: 1); Arnold (2002: 37); Bovero et al. (2003: 1); Brizzi et al. (1995: 1); Caccone et al. (1994: 1, 1997: 1); Clergue-Gazeau (1971: 1); Clergue- Gazeau (1987: 1); Clergue-Gazeau & Bonnet (1980: 1); Clergue-Gazeau & Martínez-Rica (1978: 1); García-París (1985: 113); García-París et al. (2004: 110); Gasser (1975: 1); Guillaume (1999: 1); Guillaume (2002: 1); Hervant, Mathieu & Durand (2000: 1, 2001: 1); Lecis & Norris (2003: 1, 2004: 1); Llorente et al. (1995: 34); Martínez-Rica & Clergue-Gazeau (1977: 1); Mertens & Wermuth (1960: 17); Montori (1988: 20); Montori (1991: 1); Montori & Campeny (1991: 1); Montori et al. (1997: 106); Montori et al. (2002: 48); Montori & Pascual (1981: 1); Read (1998: 1); Salvador & García-París (2001: 37); Salvidio, Sindaco & Emanueli (1999: 1); Sbordoni et al. (1982: 1, 1985: 1); Schlegel (1997: 1); Serra-Cobo et al. (2000a: 1); Serra-Cobo, Marques-Bonet & Martinez-Rica (2000b: 1); Steinfartz et al. (2002: 1); Thiesmeier & Hornberg (1990: 1); Thiesmeier et al. (1997: 1); Thorn (1968: 253); Uiblein et al. (1992: 1); Uiblein, Engelke & Parzefall (1995: 1).

Diagnosis

Small- to medium-sized newts (70–150 mm including tail). Fronto-squamosal arch ligamentous or bony. Cadosacral vertebrae with short transverse processes that are directed obliquely backwards ( Fig. 4J, K ). Head rather large, depressed, longer than broad. Upper jaw slightly or completely overhanging. Paratoid glands more or less distinct. Gular fold absent. Limbs moderate in size, four fingers and five toes, all free. Skin finely tubercular on the flanks and head, smooth beneath. Body relatively flat or slightly rounded in cross-section, no dorsal crest even during the breeding season. Tail about as long as head and body and compressed laterally. Lungs absent or very reduced. Males have spurs on hind legs (see Fig. 9A, B ). Caudal capture of female by male followed by direct transfer of spermatophores.

Species

Euproctus montanus ( Savi, 1838) and E. platycephalus ( Gravenhorst, 1829) .

Distribution

Restricted to the Mediterranean islands of Corsica and Sardinia.

EUPROCTUS MONTANUS ( SAVI, 1838)

( FIGS 4J , 9B, D, F )

Megapterna montana Savi, 1838: 211

Triturus (Euproctus) montanus Boulenger, 1878: 308 Euproctus montanus Giglioli, 1878: 97 ; Despax, 1923: 11; Mertens & Muller, 1928: 10; Mertens & Muller, 1940: 8; Wolterstoff & Herre, 1935: 224; Mertens & Wermuth, 1960: 18; Thorn, 1968: 260

Molge montana Boulenger, 1882: 23

Triton montanus Schultze, 1891: 170 ; Schreiber, 1912: 53

Triton (Euproctus) montanus Wolterstoff, 1900: 6 Diagnosis

Adults usually between 80 and 100 mm including the tail with a maximum of 130 mm. Fronto-squamosal arch ligamentous. Frontal bones with distinct orbital processes. Body relatively flat, no dorsal crest. Dorsal skin smooth in aquatic phase and finely tubercular in terrestrial phase, smooth beneath in both aquatic and terrestrial phases. Brown or olive above, sometimes with lighter yellowish, reddish or greenish markings that may form a thin vertebral line. Venter yellowish, grey or brown, speckled with whitish. Snout rounded, upper jaw slightly overhanging, absence of labial folds and gular fold. Large well-defined paratoid glands present on each side of the neck. Tongue rather large, subcircular, attached along the median line, slightly free behind, protractile. Spurs on hind legs of males laterally flattened and not strongly protruding ( Fig. 9B, D ). Fingers and toes short and depressed ( Fig. 9B, D ). Tail a little shorter than head and body, compressed laterally and pointed; without crests. Cloacal swelling of males conical and projecting backwards. Cloacal chamber containing a pseudopenis, which may protrude from the cloacal cavity during amplexus ( Fig. 9D ). Cloacal swelling hemispherical in females with a vertical slit, the borders of which are scarcely swollen ( Fig. 9F ).

Material examined

BMNH 1882.11.15.51–55; BMNH 1882.4.12.2–3; BMNH 1891.6.15.6–13; BMNH 1928.12.20.378–397; BMNH 1952.1.4.35–47; BMNH 1955.1.1.6–11. Specimens included in the X-ray osteological analysis are listed in Appendix 2.

Distribution

Euproctus montanus is restricted to the Mediterranean island of Corsica ( France), where it is found from sea level up to 2260 m, being more abundant in still or running waters in mountainous areas situated between 600 and 1500 m.

EUPROCTUS PLATYCEPHALUS ( GRAVENHORST, 1829)

( FIGS 4K , 9A, C, E )

Molge platycephala Gravenhorst (part) 1829: 84.

Euproctus rusconni Genè, 1839: 28 , pl. 1, figs 3, 4; Gray, 1858: 139; Duméril, Bibron & Duméril (part) 1854: 158 (under Euproctus rusconi ).

Euproctus platycephalus Bonaparte, 1832 –1841: vol 26; Lataste, 1878: 495; Mertens & Muller, 1928: 10; Wolterstoff & Herre, 1935: 224; Mertens & Muller, 1940: 8; Mertens & Wermuth, 1960: 18; Thorn, 1968: 265.

Phatnomatorhina platycephalus Bibron, 1839 : sign 131.

Pelonectes platycephalus Fitzinger, 1843: 33 .

Triton rusconii Bonaparte, 1832 –1841: 453; Gray, 1850: 24.

Triton platycephalus Strauch, 1870: 50 .

Triturus (Euproctus) platycephalus Boulenger, 1878: 307 .

Triturus (Euproctus) rusconii Boulenger, 1878: 308 ; Wolterstoff, 1900: 5.

Molge rusconii Boulenger, 1882: 24 .

Triton rusconii Schultze, 1891: 170 ; Schreiber, 1912: 58.

Diagnosis

Adult males approximately 127 mm including tail, adult females up to 112 mm with a maximum of 150 mm for both sexes. Fronto-squamosal arch bony. Body relatively flat, absence of dorsal crest. Dorsal skin smooth or finely tubercular above, venter completely smooth. Typically brown or olive above, variegated with greenish, yellowish or light grey with an orange-reddish vertebral stripe. The venter is whitish on the sides and yellowish or reddish along the centre, with irregularly distributed black spots on the belly and throat, especially in males. Head depressed, much longer than broad, snout semi-elliptical, upper jaw overhanging, upper lips with well-developed labial folds. No gular fold. Paratoid glands not very well defined. Tongue small, subelliptical, free at the sides. Spurs on hind legs of males strongly protruding (like a fifth finger), especially in older specimens ( Fig. 9A ). Fingers rather elongate and slender ( Fig. 9A ). Tail about as long as snout–vent length, broad at the base and increasingly compressed laterally, ending in an obtuse point, with small upper and lower crests, the latter beginning half way along the tail. Cloacal swelling conical in both sexes with a circular opening ( Fig. 9C, E ).

Material examined

BMNH 1882.12.15.56–59; BMNH 1886.10.19.2–3; BMNH 1895.4.16.2–10; BMNH 1885.4.16.11 (skeleton); BMNH 1895.5.14.7–9; BMNH 1928.12.20.365– 373; BMNH 1903.1.25.14–16; BMNH 1920.1.20.2055. Specimens included in the X-ray osteological analysis are listed in Appendix 2.

Distribution

Euproctus platycephalus is restricted to the Mediterranean island of Sardinia, Italy, where it inhabits still or more usually running waters in mountainous areas situated between 50 and 1800 m altitude, being more abundant above 600 m.

Figures

Figure 4. X-ray images of several species of newts and pictures of two clear-stained specimens showing a close-up of the caudosacral and caudal vertebrae. Numbers from 1 to 4 correspond to the first caudosacral vertebrae. A, Calotriton arnoldi sp. nov. female from El Montseny (MZB2004-0188). B, C. arnoldi male from El Montseny (MZB 82–8789). C, C. arnoldi male from El Montseny (MZB2004-0187). D, C. arnoldi male from El Montseny (MZB 82-8784). E, C. arnoldi female from El Montseny (MZB2004-0189). F, C. asper male from Baños de Benasque, Huesca, Spain (BMNH, 1970.2448). G, C. asper male from Torrent de Castelmouly, near Bagnères de Bigorre, France (BMNH, 1928.11.18-22). H, C. asper male from Pla de l’Estany, northern slope of the Maladeta, Spain (BMNH, 1928.11.22.13-14). I, C. asper female from Lac d’Oncet, French Pyrenees (BMNH, 1920.1.20.20). J, Euproctus montanus male (BMNH 82.11.15.50-55). K, E. platycephalus male (BMNH, 1947.1.4.4.x6). L, Neurergus kaisseri male (paratype – BMNH, 1952.4.2.85). M, Triturus marmoratus male (BMNH 86.6.29.52-56). N, T. cristatus male (BMNH, 1950.1.4.81-82). O, T. karelinii male (BMNH 96.3.28.18-19). P, ventral view of the caudosacral and caudal vertebrae of a clear-stained male of C. asper from Berga (locality 8 in Fig. 1). Q, dorsal view of the same C. asper specimen as in P. R, ventral view of male from Bergo. S, dorsal view of C. arnoldi from population B1 (locality 1 in Fig. 1).

Figure 9. A, detail of a hind leg of a male Euproctus platycephalus showing the spur (s) that characterizes the Tyrrhenian brook newts. B, detail of hind leg of a male E. montanus showing the spur (s). C, detail of a male cloaca of E. platycephalus; the spur on the right hind leg is also visible. D, detail of a male cloaca of E. montanus showing the pseudopenis (pp) and the spur. E, detail of a female cloaca of E. platycephalus. F, detail of a female cloaca of E. montanus.