Speleomantes Dubois, 1984

Class: Amphibia > Order: Caudata > Family: Plethodontidae > Subfamily: Plethodontinae > Genus: Speleomantes
8 species

Atylodes Gistel, 1868, Die Lurche Europas: 158. Type species: Salamandra genei Schegel, by monotypy. A nomen oblitum when used for the same taxon as Speleomantes Dubois, 1984, according to Crochet, 2007, Amphibia-Reptilia, 28: 170-172, who discussed the nomenclatural issues involved.

Speleomantes Dubois, 1984, Alytes, 3: 103-110. Type species: Spelerpes italicus Dunn, 1923, by original designation. Coined as a subgenus of Hydromantoides.

SpeleomantesLanza, 1986, in Camard et al. (eds.), Ambiente Nat. Sardegna: 310; Lanza, Pastorelli, Laghi, and Cimmaruta, 2006 "2005", Atti Mus. Civ. Stor. Nat. Trieste, 52 (Suppl.): 5-135. Treatment as a genus.

AtylodesWake, Salvador, and Alonso-Zarazaga, 2005, Amphibia-Reptilia, 26: 543–548. Treatment as a subgenus of Hydromantes.

AtylodesVieites, Min, and Wake, 2007, Proc. Natl. Acad. Sci. USA, 104: 19904 (and electronic supplemental data). Treatment as a genus.

English Names

None noted. 

Distribution

Sardinia and montane central to northwestern Italy and southeastern France.

Comment

Lanza and Vanni, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 117–121, believed that the American and European components of Hydromantes (sensu lato) were convergently derived. They therefore erected a genus Hydromantoides for the New World species and restricted the name Hydromantes for the European species. Taxonomy of this genus passed through a period where this systematic controversy was complicated by serious problems of nomenclature regarding whether the name Hydromantes was even available for the species to which it had traditionally been applied. This discussion culminated in a ruling by the Commission on Zoological Nomenclature to change the type species of Hydromantes to a North American species, Hydromantes platycephalus Camp, 1916, thereby "saving" the name Hydromantes for its North American usage and rendering Hydromantoides Lanza and Vanni, 1981, an objective synonym of Hydromantes. However, in the course of the nomenclatural discussion the name Speleomantes Dubois, 1984, had been coined to cover the monophyletic European component of Hydromantes sensu lato, which is the sister taxon of the American component (in the sense of including Atylodes). See Anonymous, 1997, Bull. Zool. Nomencl., 54: 72–74, for a summary of the controversy. Subsequently American authors (e.g., Wake, 1993, Herpetologica, 49: 229–237) retained Hydromantes for both American and European components because of their supposed evolutionary propinquity, while Europeans recognized Speleomantes (e.g., Lanza, Pastorelli, Laghi, and Cimmaruta, 2006 "2005", Atti Mus. Civ. Stor. Nat. Trieste, 52 (Suppl.): 5–135) for the European component. That the European and American groups of species are sister taxa is not controversial so this is not a scientific controversy, but one of aesthetics. Wake, Salvador, and Alonso-Zarazaga, 2005, Amphibia-Reptilia, 26: 543–548, noted that while Hydromantes (sensu stricto) is the sister taxon of Speleomantes (in the pervasive sense of European authors), they preferred a taxonomy in which Hydromantes was construed to be a genus with three subgenera: Hydromantes (for former Hydromantes, sensu stricto), Atylodes (for Speleomantes genei, which they suggested was the sister taxon of the remaining species of former Speleomantes), and Speleomantes (for the species of former Speleomantes other than Hydromantes genei). Lanza, Pastorelli, Laghi, and Cimmaruta, 2006 "2005", Atti Mus. Civ. Stor. Nat. Trieste, 52 (Suppl.): 5–135, treated Speleomantes as a genus and implied that Atylodes should be treated as a subgenus of that genus. Crochet, 2007, Amphibia-Reptilia, 28: 170–172, discussed the nomenclature of this group, noting that if the two, reciprocally monophyletic groups of Californian and European species were to be considered two genera (an arrangement consistent with the phylogenetic tree of Wake, Salvador, and Alonso-Zarazaga, 2005, Amphibia-Reptilia, 26: 543–548), the generic names would be Hydromantes and Speleomantes. Speybroeck and Crochet, 2007, Podarcis, 8: 10, formally suggested this arrangement. Vieites, Min, and Wake, 2007, Proc. Natl. Acad. Sci. USA, 104: 19903–19907, considered Atylodes, Hydromantes (sensu stricto), and Speleomantes as genera.  Nardi, 1991, in Green and Sessions (eds.), Amph. Cytogenet. Evol.: 131–156, reported on evidence of phylogeny based on karyology. Macey, 2005, Cladistics, 21: 194–202, considered Aneides to be the sister taxon of Hydromantes (in the sense of including Speleomantes). Carranza, Romano, Arnold, and Sotgiu, 2008, J. Biogeograph., 35: 724–739, discussed the phylogenetics and biogeography of Hydromantes and Speleomantes. Vieites, Min, and Wake, 2007, Proc. Natl. Acad. Sci. USA, 104: 19904 (and electronic supplemental data) treated Atylodes, Hydromantes, and Speleomantes as genera due to the genetic distances between them and as a means to standardize usage across Europe and North America; these authors apparently reconsidered this arrangement and Vieites, Nieto-Roman, Wake, and Wake, 2011, Mol. Phylogenet. Evol., 59: 623–635, considered all three taxa to be part of a single, larger Hydromates. van der Meijden, Chiari, Mucedda, Carranza, Corti, and Veith, 2008, in Corti (ed.), Herpetol. Sardiniae: 367–369, commented on the molecular phylogenetics of the genus. Atylodes was considered a synonym of Hydromantes (sensu lato) by Mertens, 1936, Senckenb. Biol., 18: 76, prior to its recognition as a subgenus by Wake, Salvador, and Alonso-Zarazaga, 2005, Amphibia-Reptilia, 26: 543–548, and recognition as a genus by Vieites, Min, and Wake, 2007, Proc. Natl. Acad. Sci. USA, 104: 19903–19907. van der Meijden, Chiari, Mucedda, Carranza, Corti, and Veith, 2009, Mol. Phylogenet. Evol., 51: 399–404, continued the use of Atylodes as a subgenus of Hydromantes, but did not address the arrangement suggested by Vieites, Min, and Wake, 2007, Proc. Natl. Acad. Sci. USA, 104: 19903–19907 (suppl. material), which is therefore retained here. Crochet, 2007, Amphibia-Reptilia, 28: 170–172, noted that Atylodes Gistel, 1868, must be treated as junior to Speleomantes Dubois, 1984, when they compete nomenclaturally. Speybroeck, Beukema, and Crochet, 2010, Zootaxa, 2492: 5, discussed why they a synonym of Hydromantes. Vieites, Nieto-Roman, Wake, and Wake, 2011, Mol. Phylogenet. Evol., 59: 623–635, considered Hydromantes, sensu lato, to be in their Hydromantini; they also treated Atylodes as a subgenus of Hydromantes, thereby rejecting their 2007 position that it should be considered a genus on the basis of antiquity. Sardinian species and their interrelationships discussed by Lanza, Nascetti, and Bullini, 1986, Boll. Mus. Reg. Sci. Nat. Torino, 4: 261–289. Nascetti, Cimmaruta, Lanza, and Bullini, 1996, J. Herpetol., 30: 161–183, and Nardi, Batistoni, Marracci, and Lanza, 1999, Herpetologica, 55: 131–139, discussed the molecular relationships of the species. Lanza, Caputo, Nascetti, and Bullini, 1995, Monogr. Mus. Reg. Sci. Nat. Torino, 16, provided a key and accounts. Thorn and Raffaëlli, 2000, Salamand. Ancien Monde: 368–377, provided accounts for the European species of Hydromantes (sensu lato) as Speleomantes. Lanza, 1999, in Grossenbacher and Thiesmeier (eds.), Handbuch Rept. Amph. Eur., 4(1): 81–204, discussed all of the European species, provided accounts, and a key, as well as the nomenclatural history of Speleomantes. Nascetti, Cimmaruta, Lanza, and Bullini, 1996, J. Herpetol., 30: 161–183, provided a discussion of molecular phylogenetics of the European species. Nardi, 1991, in Green and Sessions (eds.), Amph. Cytogenet. Evol.: 131–156, reported on evidence of phylogeny based on karyology. Lanza, Pastorelli, Laghi, and Cimmaruta, 2006 "2005", Atti Mus. Civ. Stor. Nat. Trieste, 52 (Suppl.): 5–135, reviewed Speleomantes. Lanza, Pastorelli, Laghi, and Cimmaruta, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 142–174, provided a key and detailed accounts for the Italian species. Wake, 2013, Amphibia-Reptilia, 34: 323–326, discussed the taxonomic history of the genus. The phylogeny, particularly of the generic-group, is reasonably well understood. What is going on now is that David B. Wake thinks it is best to retain all three units (Hydromantes, Atylodes, and Speleomantes) in one genus (Hydromantes), while Europeans (e.g., Vieites, Raffaelli, Dubois) prefer to place the North American monophyletic group in one genus (Hydromantes) and the European monophyetic group in another genus (Speleomantes) with two subgenera (Atylodes and Speleomantes). Rather than go back and forth with each series of publication, I am adopting the European point of view, simply because where there is a tradition of treating monophyletic groups at different ranks in the genus-group, I have uniformly gone with the more fine-grained approached simply as a methodological principle. Raffaëlli, 2013, Urodeles du Monde, 2nd ed.: 436–442, provided a discussion of the taxonomic controversy and provided brief apecies accounts, photographs, and a range map. Chiari, van der Meijden, Mucedda, Lourenço, Hochkirch, and Veith, 2012, PLoS One, 7(3: e32332): 1–15, reported on biogeography among Sardinian populations and species and provided a tree for all species in the genus. Sparreboom, 2014, Salamanders Old World: 148–150 (Atylodes), 153–165 (Speleomantes), reviewed the biology, characteristics, distribution, reproduction, and conservation of the species. Raffaëlli, 2022, Salamanders & Newts of the World: 1016–1033, provided species accounts summarizing systematics, morphology, life history, population status, and distribution (including a polygon map).

Contained taxa (8 sp.):

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