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This family includes the species regarded as typical triton), on its dorsal surface (as in the American species and . It is a diverse family, including of and ), by its tail ( both large- and small-bodied, terrestrial and – at least and ), or make no physical contact with the partly – aquatic species. Some are strictly terrestrial female at all (Cynops, , Parameso triton, Lao- (for instance Salamandra in Europe and the Middle East, triton, , , , Ichthyo- in China and Japan) and some are perma- saura, Echinotriton, and Salamandrina). nently aquatic (such as Pachytriton in China). The largely The presence of well-developed lungs is ancestral for terrestrial salamanders (Chioglossa, Lyciasalamandra, salamandrids with fi ve evolutionarily independent Mertensiella, Salamandra) are smooth-skinned while reductions or losses of lungs in the genera Calotriton, most of the other genera have a rough skin. Most Chioglossa, Euproctus, Pachytriton, and Salamandrina. species have aquatic larvae except for some viviparous The family Salaman dridae contains some 93 species salamanders that give birth to fully metamorphosed assigned to the following genera: Calotriton (2 species), offspring (for instance Lyciasalamandra). No single Chioglossa (1 species), Cynops (9 species), Echinotriton feature uniquely characterises all salamandrids as a (2 species), Euproctus (2 species), monophyletic group, but a combined analysis of mor- (1 species), Laotriton (1 species), Lissotriton (5 species), phological and molecular characters strongly supports Lyciasalamandra (7 species), Mertensiella (1 species), the monophyletic view (Titus & Larson, 1995; Zhang Neurergus (4 species), Ommatotriton (2 species), Pachy- et al., 2008). The family has a holarctic triton (7 species), (12 species), Pleuro- distribution and is represented in Europe, north-west- deles (3 species), Salamandra (6 species), Salamandrina ern Africa, Asia, and North America. (2 species), Triturus (8 species) and (18 species). For a detailed review of the biology, repro- Fertilisation is internal. A male inseminates a female duction and diversity of salamandrids, see Griffi ths using a spermatophore that he deposits in front of her. (1996), Zhao et al. (1988) and Larson et al. (2007). Stein- The female picks up the sperm with her cloaca. The eggs fartz et al. (2007), Weisrock et al. (2006) and Zhang et al. are fertilised before they are laid. Courtship in salaman- (2008) have analysed the phylogenetic relationships drids can be classifi ed into four general types based on within the family Salamandridae. the degree of contact between the male and female (Houck & Arnold, 2003). Male salamandrids may actively References Griffi ths (1996), Houck & Arnold (2003), Larson capture the female on its ventral surface (Mertensiella, et al. (2007), Steinfartz et al. (2007), Titus & Larson (1995), Lyciasalamandra, Salamandra, , and Tyloto- Weisrock et al. (2006), Zhang et al. (2008), Zhao et al. (1988).

Salamandridae Goldfuss, 1820

Salamandra corsica. Vizzavona. Photo: Max Sparreboom.

170 Salamanders of the Old World | Salamandridae Calotriton Gray, 1858

This was resurrected to comprise two species of Pyrenean Their distribution is restricted to the Pyrenees. The genus mountain brook newts formerly included in the genus Euproctus Calotriton is sister to Triturus (Pyron & Wiens, 2011). (Carranza & Amat, 2005; Frost, 2013). They are small­ to medium­ sized newts. Head depressed and fairly robust. Prominent Calotriton arnoldi Carranza & Amat, 2005 swellings on posterior sides of the head. Paratoid glands absent. Calotriton asper (Dugès, 1852) Gular fold present. Skin covered with tubercles bearing horny tips. Tips of fi ngers and toes covered by a black corneous, nail­ References Carranza & Amat (2005); Frost (2013); Pyron & Wiens like sheath. No dorsal and caudal crest during breeding season. (2011).

Calotriton arnoldi Carranza & Amat, 2005 | Montseny

Description A rough­skinned newt without paratoid glands. Venter translucid, light ochre­brown, with dark markings most Head relatively fl at, longer than wide, snout more or less round­ developed at the sides (Carranza & Amat, 2005). ed and slightly truncated; upper jaw reaching over the edge of lower jaw; eyes small. Labial folds present. Gular fold present. Males are not signifi cantly smaller than females. Cloaca Tail ending in obtuse point and shorter than snout­vent length. rounded and with a vertical slit in the male, and cylindrical in First three to four caudosacral vertebrae with short transverse the female, relatively narrow and of a contrasting bright red­ processes, not elongate as in Calotriton asper. Four fi ngers, fi ve dish­orange colour at the tip. toes, all depressed and free, with tips covered by black claw­like sheath. Skin rough, but much smoother than in C. asper. Total length ca. 10 cm.

The colour is chocolate brown, sometimes with light, silvery­ Diagnosis Similar to Calotriton asper, but differs in its gold stippling on the sides. Adult and young specimens mitochondrial DNA and various other features: fi rst caudosacral uniform dorsally, with a light and very thin brownish­orange vertebrae have short transverse processes directed obliquely stripe from the base to the tip of the tail. Complete absence in backwards; dorsal skin is smoother; no tubercles on underside; both adults and young of the broad light­coloured vertebral smaller than C. asper; lack of vertebral stripe (Carranza & Amat, stripe common in C. asper. Throat very light, largely unspotted. 2005).

Calotriton arnoldi, Montseny. Photo: Frank Pasmans. Calotriton arnoldi, Montseny. Photo: Frank Pasmans.

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