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Zootaxa, Caudata, Pleurodeles Zootaxa 488: 1–24 (2004) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 488 Copyright © 2004 Magnolia Press ISSN 1175-5334 (online edition) Taxonomic revision of Algero-Tunisian Pleurodeles (Caudata: Salamandridae) using molecular and morphological data. Revalidation of the taxon Pleurodeles nebulosus (Guichenot, 1850) SALVADOR CARRANZA1* & EDWARD WADE2 1. Department of Zoology, The Natural History Museum, London, SW7 5BD ([email protected]) * Present address: Departament de Biologia Animal, Universitat de Barcelona, Av. Diagonal 645, E-08028 Barcelona, Spain ([email protected]) 2. Middlesex University, Cat Hill, Barnet, Hertfordshire, EN4 8HT ([email protected]) Abstract The taxonomic status of Algero-Tunisian Pleurodeles was reanalysed in the light of new molecular and morphological evidence. Mitochondrial DNA sequences (396 bp of the cytochrome b and 369 of the 12S rRNA) and the results of the morphometric analysis, indicate that Algero-Tunisian P. poireti consists of two genetically and morphologically distinct forms. One restricted to the Edough Peninsula, and another one covering all the rest of its distribution in Algeria and Tunisia. The name P. p oire ti (Gervais, 1835) is restricted to the population of the Edough Peninsula, while P. nebulous (Guichenot, 1850) correctly applies to all other populations in the distribution. P. po ire ti originated approximately 4.2 Myr ago, probably as a result of the Edough Peninsula being a Pliocene fossil island, allowing both forms of Algero-Tunisian Pleurodeles to diverge both genetically and mor- phologically. Key words: Pleurodeles, Algeria, taxonomy, mitochondrial DNA, 12S rRNA, cytochrome b, mor- phology, Pliocene fossil island Introduction The genus Pleurodeles currently consists of two species. P. waltl Michaelles and P. poireti (Gervais, 1835). Morphology and mitochondrial DNA sequences indicate that, among liv- ing forms, the sister taxon of Pleurodeles is Tylototriton from southeast Asia (Titus and Larson, 1995). Despite some fossils originally thought to be related to Pleurodeles dating back to the Upper Oligocene of Germany (Palaeopleurodeles Herre 1941), it is believed that the origin of Pleurodeles is much more recent, having split from its sister taxon (Tylo- totriton) during the Middle Miocene, some 10 Myr ago (see Carranza and Arnold, 2003 for a critical review of the Palaeontological data). This result is supported by the age of the Accepted by M. Wilkinson: 22 Mar. 2004; published: 13 Apr. 2004 1 ZOOTAXA earliest specimens assigned to Pleurodeles, which date from the Upper Miocene or Lower 488 Pliocene of Spain (Sanchiz, 1977). Pleurodeles waltl is large (up to 300 mm in total length—Pasteur, 1958), with usually 15 presacral vertebrae, a tubercular process on rib 3 (and often traces of this structure on ribs 1 and 2), and sharp rib tips that project through a row of glandular swellings on the flanks. P. waltl is common and widely distributed in the southern two thirds of the Iberian Peninsula and occurs with less abundance in northern Morocco, where it is found in the area delineated by Tangiers, Casablanca and Alhoceima (Bons and Geniez, 1996). The second Pleurodeles species, P. poireti, is smaller (up to about 230 mm in total length), with 14 presacral vertebrae, no obvious tubercular processes on the ribs, which are not sharp, and no glandular swellings on the flanks. It occurs in northern Tunisia and Algeria extend- ing westwards as far as Oran. Nomenclatural history The first person to mention the presence of P. poireti in northern Algeria was Poiret (1789) in his Voyage en Barbarie. In there, he gives a brief description of this species under the name of Lacerta palustris. Gervais (1835), analysed material of Lacerta palustris from Oran, Algiers and Bône from Marly and Gerard and recognised it represented a new spe- cies distinct from L. palustris of Linné 1758, which he named Triton Poireti (=Pleurodeles poireti). However, he already noted the similarity that existed between Triton Poireti and the Ibero-Moroccan Pleurodeles Waltl, especially in the shape of the head and tail. He con- cluded that, although T. Poireti lacked the exposed long ribs characteristic of Pleurodeles Waltl, it was linked to it. Despite Gervais’ (1835) material being collected from widely separated localities, the extant type series of T. Poireti actually consists of only two speci- mens, both from Bône (= Annaba) (MNHNP 4744 and MNHNP 4744A—Thireau, 1986), within the general area to which we subsequently refer herein to as the Edough Peninsula (see Fig. 1). Bône, therefore should be considered the type locality (Thireau, 1986). Bonaparte (1839) assigned T. Poireti to the genus Glossoliga. Latter Guichenot (1850) synonymized T. Poireti with the Sardinian Euproctus Rusconii Gené, 1838 (=Euproctus platycephalus; Gravenhorst, 1829). In the same account, Guichenot (1850) described a new species, Triton nebulosus (=Pleurodeles nebulosus). According to Guichenot, T. neb- ulosus differed from T. Poireti in having longer tail, longer, wider and more depressed head, shorter teeth slightly curved and less prominent skin tubercles, which appear closer together. The type material of T. nebulosus is deposited at the MNHNP and includes a complete female specimen from Algiers collected by Guichenot himself (MNHNP 1442), and two other specimens, which were dissected about 1867 to produce a complete skeleton (MNHNP CD 8) and a skull (MNHNP CB 21) (Thireau, 1986). Gray (1850) placed the Algero-Tunisian newt in the genus Triton (T. Poireti) and synonymized T. nebulosus with T. Poireti. Euproctus Rusconii is listed in Gray (1850) as a synonym of the Sardinian 2 © 2004 Magnolia Press CARRANZA & WADE endemic Euproctus platycephalus. Gervais (1853) showed that T. Poireti from Algeria and ZOOTAXA E. Rusconii from Sardinia were different species and argued in favour of Bonaparte’s 488 (1839) proposal of altering the generic name of Triton Poireti to Glossoliga. Guichenot’s Triton nebulosus and E. Rusconii were both listed as synonyms of T. Poireti by Dumeril and Bibron (1954), who changed its generic name from Triton to Euproctus (as E. Poireti). Lataste (1881), found morphological differences between populations of Glossoliga Poireti. In recognition of such, he described a new morphological variant from Bône (Annaba) as a new species, Glossoliga Hagenmulleri, in honour of Dr. Hagenmüller, who collected all of the 32 specimens (26 males and 6 females) used in the original description (Lataste, 1881). But in fact, Gervais’ (1835) type material of T. Poireti originated from three localities in Algeria, one of which was Bône. Lataste (1881) was in error as to which of the three localities should apply Gervais’ (1835) T. Poireti. He decided that the speci- mens from d’Alger in his account (= Algiers) should carry the name Poireti. This appears to be based on an illustration (Gervais, 1853; pl. 14 Fig. 9) of a dorsal view of a skull of a specimen clearly stated to be d’Algérie i.e. Algeria, therefore too imprecise to be applica- ble. Moreover, the only types that appear to have survived are from Bône (Thireau, 1986), which should therefore be considered most appropriate as the type locality of Triton Poireti (Thorn,1968; Frost, 1985; Thireau, 1986). Lataste’s (1881) type series of Glossoliga Hagenmulleri comprises six specimens from Mount Edough, within the Edough Peninsula (northeast Algeria; see Fig. 1): BMNH 1946.9.6.77 – 82 (BMNH 1920.1.20.1165 auct.). According to Lataste (1881), G. Hagen- mulleri differed from G. P o iret i in having the palatine teeth row forming a V; smaller size; head longer than broad; tongue small and less free behind; snout narrower; contour of jaws semi elliptical; limbs, especially fingers and toes, more slender; gular fold less pro- nounced; colour olive-brown above, greyish beneath with more or less distinct darker spots. These morphological differences between the two species of Algero-Tunisian newts were later corroborated by Boulenger (1882), who placed them in the genus Molge (M. poireti and M. hagenmulleri). After Boulenger (1882), several authors recognised both forms at the specific or subspecific levels (Camerano, 1885; Woltersttorff, 1905; Noble, 1924) but reassigned their generic name to Triton (Camerano, 1885), to Pleurodeles (Woltersttorff, 1905), and again to Triton (Noble, 1924). In a revision of Lataste’s (1881) work, Pasteur (1958) synonymized Pleurodeles hagenmulleri (Lataste, 1881) with Pleurodeles poireti (Gervais, 1835). He analysed some specimens of P. poireti and P. hagenmulleri from the Muséum National d’Histoire Naturelle, Paris (MNHNP) for most morphological characters used in the original descrip- tion (Lataste, 1881), and those of Boulenger (1882) and Doumerge (1901). He concluded that, with the only exception of MNHNP 00.158, which perfectly fitted Lataste’s (1881) description of P. hagenmulleri, he could not clearly differentiate between P. po i re ti and P. hagenmulleri. As a result of his work, the only recognised form of Algero-Tunisian Pleu- rodeles today is P. p oi re ti. REVISION OF PLEURODELES © 2004 Magnolia Press 3 ZOOTAXA In this paper, we revise the taxonomy of P. poireti in North Africa using morphology 488 and cytochrome b (cytb) and 12S rRNA (12S) mitochondrial sequences. Materials and Methods Relevant data for all the specimens used in the molecular and morphological analyses are presented in Table 1 and Fig. 1. Species used in the molecular study are a selection of some key sequences from a previous phylogeographic
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