Northern Spectacled Salamander
Species Description: Nascetti G, Zangari F, Canestrelli D 2005 The spectacled salamanders, Salamandrina terdigitata (Lacepede, 1788) and S. perspicillata (Savi, 1821): 1) Genetic differentiation and evolutionary history. Rend. Fis. Ace. Linei s. 9, v. 16:159-169
© 2013 Wouter Beukema (1 of 32)
Salamandrina perspicillata is a tiny salamander of about 35 mm SVL and 85 mm total length, respectively (Romano et al. in press; Zuffi 1999). Total length up to 92 mm in largest males (Vanni, 1980) and up to 130 mm in largest females or little more (Romano and Mattoccia 2005; Bovero et al. 2006). On the Lepini mountains (Latium, Central Italy) populations are characterised by large size salamanders (Romano and Mattoccia 2005 and references therein; Bovero et al. 2006; Angelini et al. 2008). The body is dorsoventrally flattened, with clearly visible ribs, giving the salamander a very skinny appearance. No parotoids. Four toes on both the front and the hind feet. Salamandrina is usually deep brown or grey-blackish on the dorsal side of body and tail. Tail is also partially reddish dorsally. The underside of the tail and the feet, and frequently the distal part of the belly, are bright red. The rest of the ventral region is white, whitish or greyish, with dark grey to black spots. On the head a V–shaped, more or less evident, whitish or yellowish spot between the eyes, forms a sort of “spectacles” which give rise to both the common name (Spectacled Salamander) and scientific name ( perspicillata is a Latin neologism which means “with spectacles”). Variation in dorsal coloration includes semialbinism, completely red back, or a yellowish spotted pattern (see Lanza and Canestrelli 2002, for a review).
While the sexes do not show any variation in external body features, there is some sexual dimorphism. Males are smaller than females, slightly different in the ratio of tail length to body length, have relatively more developed feet, larger head and eyes, and more distant nostrils (Vanni, 1980). Unfortunately, because the biometric parameters of males and females overlap each other partially (Vanni, 1980), they cannot be used to distinguish the sexes (Lanza 1983; Zuffi 1999). In living males the cloaca seems just slightly more prominent than in females (Vanni, 1980; Lanza, 1983), however this character is not unambiguously discriminant.
Salamandrina perspicillata can be distinguished from S. terdigitata on the basis of mtDNA haplotypes and allozyme profiles (Mattoccia et al. 2005; Nascetti et al. 2005; Canestrelli et al. 2006), but they differ also in body size and dorsal coloration (Romano et al. in press) and in the ventral pattern (Costa et al. 2008). Salamandrina perspicillata has a larger size and less extended red coloration on the back tail, and a median reddish dorsal line is very rare in this species (Romano et al. in press). At the moment, the only way to definitively distinguish between the two species is on the basis of mtDNA haplotypes and allozyme profiles because any distinction on the basis of body size and dorsal coloration is only statistical and is not definitive for any given specimen. In other words, to attribute individuals to either of the two species, morphometry, dorsal and ventral patterns can be very useful, but not determinative. However the percentage of correct classification using these non-genetic approaches is very good (> 90% in both methods).
Distribution and Habitat
Salamandrina perspicillata is endemic to central and northern peninsular Italy, prevalently in the Apennine Mountains and other hilly areas, in the whole of eastern side of Apennine and on the western side until the province of Caserta (Campania region, southern Italy. South of this it is replaced by Salamandrina terdigitata (Barbieri and Pellegrini, 2006; Romano et al. in press). It is usually found at elevations between 200 and 900m asl, but might occur between near sea level (Ruffo and Stoch 2005; Romano et al. in press) and 1,900m asl. (Barbieri, 1998)
The northern limit of this species is near Brallo di Pregola, Lombardia region (Bonini et al. 2004) and the southern limit is in the municipality of Caserta, Campania (Romano et al. in press). The western limit is near Bolzaneto, Liguria region, province of Genoa (Barbieri 1994; Barbieri & Pellegrini 2006), although Bedriaga (1897) reported records also for Cogoleto, which is about 25 km further to the west. The eastern limit of S. perspicillata is in the northern Apulia (Romano et al. in press).
It is mainly found in forests with dense undergrowth in hilly and mountainous areas, often in north-facing mountain valleys. Only females of this species are aquatic during the short oviposition phase for which well-oxygenated waters, slow running streams usually with rocky beds, drinking troughs, residual and vernal ponds (sometime of very small dimensions) and, rarely, caves are used (e.g. Lanza 1983; Corsetti 1999a; Angelini et al. 2006; Romano et al. 2007; Corsetti and Romano 2007). It appears to avoid seriously modified habitats. During summer and winter the Spectacled Salamander has been found between 20-135 cm underground (Bruno 1973, Vanni 1980, Lanza 1983), but on average there is scanty available information on terrestrial shelter used by S. perspicillata . Furthermore in spring and summer some salamanders have been found between 130-170 cm from the ground in the clefts of vertical rocks near breeding sites (cf. Romano and Ruggiero 2008).
Life History, Abundance, Activity, and Special Behaviors
The terrestrial activity of Salamandrina perspicillata is mainly, but not exclusively, crepuscular and nocturnal. The species is active mostly between 6:00-12:00 pm and there is a positive correlation between time of activity and average ambient air temperature (Utzeri et al. 2004).
Very scanty information is available on the reproductive behaviour of S. perspicillata. Courting Salamandrina have been observed only once in captivity, in December, by Strotgen (1927) who is considered by Lanza (1988) as the only author who has made reliable observation of the animals in heat. Recently, Houck and Arnold (2003) described first-hand observations of courtship behaviour in the field which agree substantially with that reported by Strotgen (1927). However both reports describing courtship behaviour are considered lacking in detail and not validated under natural conditions by authors who have carried out research on courtship patterns and phylogeny of the Salamandridae (Salthe, 1967; Steinfartz et al., 2007 and references therein). Salamandrina exhibit a variant of ventral amplexus (cf. Houck and Arnold, 2003) in which, on the ground, the pair face head-to-head and circle but without the forelimbs interlocked. The male deposits a 2 mm high spermatophore at the end of courtship. Sperm storage is among the longest in the Salamandridae and it can reach six months (Brizzi et al. 1995).
Oviposition usually occurs mainly from March to April but, according to the year and locality, can begin at the end of December and last until August (Lanza 1983; Zuffi 1999). However in the Latium there is a mountain chain and an adjacent volcano complex where spawning activity has been recorded in some sites also from late September (Corsetti 1999a, 1999b; Angelini et al. 2006; Angelini et al. 2007). Salamandrina perspicillata show high fidelity to the spawning site (Vanni, 1980; Della Rocca et al. 2005; Angelini et al. 2008) and larger females oviposit earlier than the smaller ones (Angelini et al. 2001; Della Rocca et al. 2005; Angelini et al. 2006). A female generally spawns 30-60 eggs (Vanni 1980, Zuffi 1999). The eggs are deposited singularly on suitable submerged objects, each egg glued to the substrate by a peduncle. Egg masses are the result of adhesion of closely eggs deposited by the same female or by several females. The diameter of the yolk is 1.5-3.7 mm (Angelini et al. 2007).
Development takes 20-22 days at 14ºC, and the larvae hatch at a length of 7-13 mm (Angelini et al. 2007). They superficially resemble the larvae of Salamandra salamandra, although S. perspicillata larvae are much smaller and more slender. The time to metamorphosis varies between 2 and 5 months (Lanza 1983, Corsetti 1999b), depending mainly on water temperature, oxygen availability and food resources. Metamorphosis occurs at a length of 20-35 mm (Angelini et al. 2007). The newly metamorphosed salamanders take on adult coloration within a few months.
Sexual maturity is reached in females at 4-5 years (Bovero et al. 2006). Longevity was recorded in captivity to be 10 to 12 years (Boehme et al 1999) and up to 12 years in the wild (Bovero et al. 2006).
These salamanders feed on the ground, on all sorts of small invertebrates, especially arthropods such as Arachnida, Isopoda, Emyptera, Collembola, Coleoptera, Orhtoptera, Hymenoptera although preferred prey seem to be Acarina (see Utzeri et al. 2004 and references therein). Salamanders feed also on worms and snails (Utzeri et al. 2004). Larvae consume small aquatic invertebrates and especially on Chironomidae larvae (Corsetti 1994).
The only documented cases of predation on adult salamanders concerns a bird, Lanius sp. (Angelini et al. 2007), and the crustaceans Potamon fluviatile and Austropotamobius pallipes (Vanni 1980; Della Rocca et al. 2005). The slow-worm Anguis fragilis and the common toad Bufo bufo are also thought to be predators (Vanni 1980). Salamandrina larvae are preyed upon by other urodeles such as Triturus carnifex (Romano pers. obs.) and invertebrates such as Notonecta (Corsetti 1994), Odonata (Romano pers. obs.), Crustacea (Lanza 1983) and probably many others. Salamandrina eggs are preyed upon by a variety of aquatic organisms, such as caddisfly larvae (Vignoli et al. 2001; Romano et al. 2008a), leeches (Romano and Di Cerbo 2007; Romano et al. 2008a), tadpoles or Rana italica (Laghi and Pastorelli 2006).
When threatened, the Spectacled Salamander displays a typical defensive posture reminiscent of the "unken-reflex" (e.g. Lanza, 1967; Corsetti 1994; Utzeri et al. 2005) which can be held for few minutes. The salamander curls its body dorsally to a variable degree as far as forming a loop. In exhibiting this behaviour, salamanders partially display the bright red colour of the underside of the tail and limbs and a small part of the ventral pattern of their belly and throat (which are lack and white patched). As a defense mechanism, these salamanders are also capable of appearing to be dead while otherwise alive (thanatosis), lying with the back on the ground and displaying the ventral coloration (Utzeri 2005; Romano pers. obs.). A stand-up behavior has been described by Utzeri et al. (2005): on the ground, some salamanders standing up on their limbs, apparently supported by the tail. However the meaning of the stand-up behavior is completely unknown.
Preliminary work shows that extraocular photoreceptors are involved in the migration of females towards spawning sites (Romano and Diego-Rasilla 2008). However, olfaction could represent the main orientation system in the proximity of a given water body, where the finest spatial discrimination is needed. It was found that females of Northern Spectacled Salamanders were able to recognize their home breeding water when a multiple-choice experimental device was used (Romano et al. 2008b). Perhaps scent trails are also used in the return towards terrestrial winter shelters (Romano and Ruggiero 2008). Detection of the scent of conspecifics in a terrestrial environment could be considered as one of the mechanisms adopted by these salamanders to facilitate the migration towards their winter refuges (Romano and Ruggiero 2008).
This species is widespread in the Apennine Mountains (Barbieri and Pellegrini, 2006; Romano et al. in press).
Trends and Threats
This species is relatively common in some parts of its range, but habitat alteration, fragmentation and pollution have resulted in the decline or disappearance of some populations. Considering that the species often breeds in artificial water bodies, the adequate management of these sites appears a crucial prerequisite for effective conservation of the populations in habitats (Mediterranean and sub-Mediterranean) where the water availability can be very scanty (Corsetti and Romano 2007; Romano et al. 2007). Also deforestation and clear-cutting is considered a possible threat for natural populations (Romano et al. 2008). Finally, introduction of predator fishes can damage natural populations. Salamandrina is currently protected by law in a few Italian regions (see the legislative references in Angelini et al. 2007).
Salamandrina (Lacépède, 1788) was previously considered a monotypic genus (with Salamandrina terdigitata as the sole species: see also the taxonomic notes on this species). Subsequently this genus has been split into two species on the basis of analysis using both mitochondrial and nuclear genetic markers (Mattoccia et al. 2005; Nascetti et al. 2005; Canestrelli et al. 2006). Salamandrina perspicillata (Savi 1821) was considered a junior synonym of S. terdigitata. However, after the split into two species, the name S. perspicillata has been upgraded because the two type localities of S. terdigitata and S. perspicillata are different and S. perspicillata (type locality: Mugello, Tuscany, Central Italy) is the first description of the central-northern species.
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Written by Antonio Romano (antonioromano71 AT gmail.com), Università di Roma, Tor Vergata
First submitted 2008-11-13
Edited by Kellie Whittaker (2014-03-05)
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