Zootaxa, Caudata, Pleurodelesmolevol.cmima.csic.es/carranza/pdf/pleurodeles2.pdf · Carranza and...

488

Accepted by M. Wilkinson: 22 Mar. 2004; published: 13 Apr. 2004 1

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2004 Magnolia Press

Zootaxa 488: 1–24 (2004) www.mapress.com/zootaxa/

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 molecularand morphological evidence. Mitochondrial DNA sequences (396 bp of the cytochrome b and 369of 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 EdoughPeninsula, and another one covering all the rest of its distribution in Algeria and Tunisia. The nameP. poireti (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. poireti originatedapproximately 4.2 Myr ago, probably as a result of the Edough Peninsula being a Pliocene fossilisland, 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 andLarson, 1995). Despite some fossils originally thought to be related to Pleurodeles datingback to the Upper Oligocene of Germany (Palaeopleurodeles Herre 1941), it is believedthat 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, 2003for a critical review of the Palaeontological data). This result is supported by the age of the

CARRANZA & WADE2 © 2004 Magnolia Press

488ZOOTAXA earliest specimens assigned to Pleurodeles, which date from the Upper Miocene or Lower

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 onribs 1 and 2), and sharp rib tips that project through a row of glandular swellings on theflanks. P. waltl is common and widely distributed in the southern two thirds of the IberianPeninsula and occurs with less abundance in northern Morocco, where it is found in thearea delineated by Tangiers, Casablanca and Alhoceima (Bons and Geniez, 1996). Thesecond Pleurodeles species, P. poireti, is smaller (up to about 230 mm in total length), with14 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 thename of Lacerta palustris. Gervais (1835), analysed material of Lacerta palustris fromOran, 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 (=Pleurodelespoireti). However, he already noted the similarity that existed between Triton Poireti andthe 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 PleurodelesWaltl, it was linked to it. Despite Gervais’ (1835) material being collected from widelyseparated 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 (=Euproctusplatycephalus; Gravenhorst, 1829). In the same account, Guichenot (1850) described anew species, Triton nebulosus (=Pleurodeles nebulosus). According to Guichenot, T. neb-ulosus differed from T. Poireti in having longer tail, longer, wider and more depressedhead, shorter teeth slightly curved and less prominent skin tubercles, which appear closertogether. The type material of T. nebulosus is deposited at the MNHNP and includes acomplete 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 theAlgero-Tunisian newt in the genus Triton (T. Poireti) and synonymized T. nebulosus withT. Poireti. Euproctus Rusconii is listed in Gray (1850) as a synonym of the Sardinian

© 2004 Magnolia Press 3REVISION OF PLEURODELES

488ZOOTAXAendemic Euproctus platycephalus. Gervais (1853) showed that T. Poireti from Algeria and

E. Rusconii from Sardinia were different species and argued in favour of Bonaparte’s(1839) proposal of altering the generic name of Triton Poireti to Glossoliga. Guichenot’sTriton nebulosus and E. Rusconii were both listed as synonyms of T. Poireti by Dumeriland Bibron (1954), who changed its generic name from Triton to Euproctus (as E. Poireti).

Lataste (1881), found morphological differences between populations of GlossoligaPoireti. In recognition of such, he described a new morphological variant from Bône(Annaba) as a new species, Glossoliga Hagenmulleri, in honour of Dr. Hagenmüller, whocollected 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 fromthree localities in Algeria, one of which was Bône. Lataste (1881) was in error as to whichof 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 appearsto be based on an illustration (Gervais, 1853; pl. 14 Fig. 9) of a dorsal view of a skull of aspecimen 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 TritonPoireti (Thorn,1968; Frost, 1985; Thireau, 1986).

Lataste’s (1881) type series of Glossoliga Hagenmulleri comprises six specimensfrom Mount Edough, within the Edough Peninsula (northeast Algeria; see Fig. 1): BMNH1946.9.6.77 – 82 (BMNH 1920.1.20.1165 auct.). According to Lataste (1881), G. Hagen-mulleri differed from G. Poireti 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 jawssemi elliptical; limbs, especially fingers and toes, more slender; gular fold less pro-nounced; colour olive-brown above, greyish beneath with more or less distinct darkerspots. These morphological differences between the two species of Algero-Tunisian newtswere later corroborated by Boulenger (1882), who placed them in the genus Molge (M.poireti and M. hagenmulleri). After Boulenger (1882), several authors recognised bothforms 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 Pleurodeleshagenmulleri (Lataste, 1881) with Pleurodeles poireti (Gervais, 1835). He analysed somespecimens of P. poireti and P. hagenmulleri from the Muséum National d’HistoireNaturelle, Paris (MNHNP) for most morphological characters used in the original descrip-tion (Lataste, 1881), and those of Boulenger (1882) and Doumerge (1901). He concludedthat, 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. poireti and P.hagenmulleri. As a result of his work, the only recognised form of Algero-Tunisian Pleu-rodeles today is P. poireti.


488ZOOTAXA In this paper, we revise the taxonomy of P. poireti in North Africa using morphology

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 arepresented in Table 1 and Fig. 1. Species used in the molecular study are a selection ofsome key sequences from a previous phylogeographic study of Pleurodeles published byCarranza and Arnold (2004).

TABLE 1. Details of material and sequences used in the present study. Numbers under CODE

refer to localities shown in Fig 1. All measurements are in mm. BMNH—Natural History Museum,

London; MNHNP—Muséum National d'Histoire Naturelle, Paris; AL—Algeria; Cytoch. b—Cyto-

chrome b.

SPECIES

CODE

ACCESSION

NUMBER

LOCALITY SEX SVL HL HW IOW TL SP3T 3T AL TL/

SVL

HL/

SVL

HW/

SVL

HW/

HL

AL/

SVL

Cytoch. b /

12S rRNA

P.poireti

1 MNHNP 4744

(Syntype)

Bône (AL) F 51.00 12.90 9.60 4.24 56.00 13.50 5.10 13.00 1.098 0.253 0.188 0.744 0.255 -

2 MNHNP 4744A

(Syntype)

Bône (AL) M 43.00 11.20 8.06 3.52 50.00 11.20 4.22 11.80 1.163 0.261 0.187 0.717 0.274 -

3 BMNH

1946.9.6.77

Mount

Edough (AL)

M 49.50 12.90 9.60 3.82 66.60 15.10 6.20 17.00 1.345 0.261 0.194 0.744 0.343 -

4 BMNH

1946.9.6.78

Mount

Edough (AL)

M 46.50 10.90 8.75 3.80 68.00 14.6 5.70 14.50 1.462 0.235 0.188 0.801 0.312 -

5 BMNH

1946.9.6.79

Mount

Edough (AL)

M 47.10 10.80 8.64 3.84 63.00 14.60 5.70 14.50 1.338 0.23 0.183 0.799 0.308 -

6 BMNH

1946.9.6.80

Mount

Edough (AL)

M 43.70 10.80 8.42 3.79 57.20 13.40 5.10 15.50 1.309 0.247 0.193 0.78 0.355 -

7 BMNH

1946.9.6.81

Mount

Edough (AL)

M 46.00 10.90 9.06 4.26 64.00 14.30 5.84 15.70 1.391 0.238 0.197 0.829 0.341 -

8 BMNH

1946.9.6.82

Mount

Edough (AL)

M 44.30 11.00 8.82 3.34 52.10 12.50 5.30 15.60 1.176 0.248 0.199 0.802 0.352 -

9 BMNH

1920.1.20.1369.1

Bône (AL) F 49.30 12.40 9.42 3.70 55.60 13.50 5.74 12.70 1.128 0.251 0.191 0.762 0.258 -

10 BMNH

1920.1.20.1369.2

Bône (AL) F 50.20 13.10 9.65 4.10 55.50 14.30 5.94 14.50 1.106 0.261 0.192 0.737 0.289 AY222508/

AY222464

11 BMNH

1920.1.20.1327.1

Bône (AL) F 59.00 12.00 10.00 4.35 69.50 15.90 6.26 16.20 1.178 0.203 0.169 0.833 0.275 AY222507/

AY222463

12 BMNH

1920.1.20.1327.2

Bône (AL) M 41.50 10.00 8.20 3.41 53.00 10.80 5.50 13.00 1.277 0.242 0.198 0.817 0.313 -

13 BMNH

1920.1.20.3872.1

Hippone

(AL)

M 48.50 11.00 9.10 3.94 59.00 13.90 5.40 13.50 1.216 0.227 0.188 0.827 0.278 -

14 BMNH

1920.1.20.3872.2

Hippone

(AL)

M 45.90 11.80 9.67 3.86 73.00 15.30 6.20 17.00 1.590 0.257 0.211 0.819 0.370 -

15 BMNH

1920.1.20.3872.3

Hippone

(AL)

M 45.00 11.20 8.90 3.60 62.50 14.10 6.40 14.50 1.389 0.250 0.198 0.792 0.322 -

16 BMNH

1920.1.20.3872.4

Hippone

(AL)

M 46.00 12.10 8.16 3.84 65.30 13.10 5.23 16.00 1.420 0.263 0.177 0.674 0.348 -

......continued on the next page


488ZOOTAXATABLE 1 (continued)

SPECIES

CODE

ACCESSION

NUMBER


SVL

HL/

SVL

HW/

SVL

HW/

HL

AL/

SVL

Cytoch. b /

12S rRNA

P.poireti

17 BMNH

1920.1.20.3872.5

Hippone

(AL)

M 49.50 11.80 9.24 3.70 71.50 13.90 5.90 15.00 1.444 0.239 0.187 0.780 0.303 -

18 BMNH

1920.1.20.3872.6

Hippone

(AL)

M 47.50 11.40 8.84 3.66 57.00 13.80 5.68 13.50 1.200 0.240 0.186 0.775 0.284 -

19 BMNH 89.12.7.8 Bône (AL) F 46.50 12.30 9.32 3.74 55.00 12.40 4.82 11.90 1.183 0.265 0.200 0.755 0.256 -

20 BMNH 89.12.7.9 Bône (AL) M 44.00 11.30 8.74 3.40 61.00 13.80 5.58 16.50 1.386 0.257 0.199 0.773 0.375 -

21 BMNH

89.12.7.10

Bône (AL) M 39.80 10.40 8.40 3.58 53.20 11.50 4.90 13.00 1.337 0.261 0.211 0.808 0.327 -

22 BMNH

1920.1.20.3873

Bône (AL) F 46 11.40 9.06 4.11 49.00 12.9 5.70 13.00 1.065 0.248 0.197 0.795 0.283 -

P. nebulosus

23 MNHNP 1442

(syntype)

Algiers (AL) F 71.50 16.70 13.10 5.73 76.00 18.00 6.82 18.00 1.063 0.234 0.183 0.784 0.252 -

24 BMNH

1920.1.20.18

Algiers (AL) M 69 16.1 12.80 5.80 72.00 17.40 6.70 20.00 1.043 0.234 0.186 0.793 0.29 AY222511/

AY222467

25 BMNH

57.10.28.92

Algiers (AL) F 46 12.30 9.30 4.35 48.00 12.60 4.60 13.00 1.043 0.267 0.202 0.756 0.283 -

26 BMNH 1859.? Algiers (AL) F 71.50 18.6 14.10 5.60 74.00 18.60 7.20 17.80 1.035 0.261 0.197 0.756 0.249 -

27 BMNH 88.4.9.2 Algiers (AL) M 60.50 14.9 11.80 5.00 83.80 17.00 6.20 19.50 1.385 0.247 0.195 0.791 0.322 -

28 BMNH 88.4.9.4 Algiers (AL) M 65.00 15.80 12.30 5.12 73.10 16.80 6.36 17.20 1.125 0.243 0.19 0.78 0.265 -

29 BMNH 88.4.9.3 Algiers (AL) F 49.10 13.00 10.30 4.58 54.00 12.90 4.99 14.00 1.10 0.265 0.21 0.794 0.285 -

30 BMNH

1920.1.20.3824

Larba (AL) F 66.00 9.62 8.83 5.76 76.00 17.10 5.92 16.80 1.152 0.146 0.134 0.918 0.255 AY222504/

AY222460

31 BMNH

1920.1.20.1511

Larba (AL) M 56.90 14.50 12.10 5.13 87.00 17.70 7.12 19.80 1.529 0.255 0.213 0.834 0.348 AY222510/

AY222466

32 BMNH

1920.1.20.1529-1

Teniet El-

Had (AL)

F 48.10 13.60 10.30 4.47 57.00 13.30 5.33 15.00 1.185 0.284 0.214 0.755 0.312 AY222544/

AY222500

33 BMNH

1920.1.20.1529-2

Teniet El-

Had (AL)

F 46.50 11.40 9.70 4.60 48.60 12.90 5.24 15.30 1.045 0.244 0.209 0.854 0.329 AY222545/

AY222501

34 BMNH

1920.1.20.1750

Bejaia (AL) M 57.20 14.70 12.00 4.98 90.00 17.30 6.48 19.50 1.573 0.257 0.21 0.816 0.341 AY222505/

AY222461

35 BMNH

19201.20.1504-1

Constantine

(AL)

M 53.00 12.60 10.00 4.28 62.00 14.10 5.34 16.80 1.17 0.237 0.189 0.797 0.317 -

36 BMNH

19201.20.1504-2

Constantine

(AL)

F 55.00 13.50 10.40 4.72 53.00 13.90 5.20 15.00 0.964 0.245 0.19 0.776 0.273 AY222462/

AY222506

37 Tabarca

(TU)

F 62.30 15.30 12.8 4.70 61.30 15.80 5.62 16.00 0.984 0.246 0.205 0.834 0.257 -

38 Tabarca

(TU)

F 54.20 14.00 10.90 4.50 66.30 15 4.80 14.50 1.223 0.258 0.201 0.779 0.268 -

39 Tabarca

(TU)

M 55.00 15.20 11.80 4.73 77.50 17.50 5.20 21.00 1.409 0.276 0.214 0.775 0.382 -

40 Dam Bou-

Heurtma

(TU)

F 82.50 19.80 15.60 5.62 97.00 21.50 5.58 22.00 1.176 0.24 0.189 0.787 0.267 -

41 Dam Bou-

Heurtma

(TU)

M 68.00 17.30 13.60 5.40 90.00 18.00 7.13 24.00 1.324 0.254 0.199 0.784 0.353 -

42 DNA analysis

only

Tabarca

(TU)

- - - - - - - - - - - - - - AY222518/

AY222474

43 DNA analysis

only

Tabarca

(TU)

- - - - - - - - - - - - - - AY222519/

AY222475

......continued on the next page


488ZOOTAXA

Phylogenetic analysis

In the present study we compared 396 bp of the cytochrome b (cytb) and 369 bp of the 12SrRNA mitochondrial genes of representatives of P. poireti from almost all its distributionrange, and some P. waltl from the Iberian Peninsula and Morocco. Salamandra s.morenica, S. s. terrestris and Mertensiella luschani were used as outgroups. Sequenceswere aligned using ClustalX (Thompson et al., 1997) with default parameters. No gapswere necessary to align the cytb sequences and only two were included in order to alignthe three outgroup 12S sequences with the ingroup. Phylogenetic trees were inferred forthe two genes together using maximum-likelihood (ML) with the General Time Reversible(GTR) model of sequence evolution, with gamma distributed rates (G) (selected by MOD-ELTEST, Posada and Crandall 1998). ML analyses were performed in PAUP* 4.0b10(Swofford, 1998) and included heuristic searches involving tree bisection and reconnec-

TABLE 1 (continued)SPECIES

CODE

ACCESSION

NUMBER


SVL

HL/

SVL

HW/

SVL

HW/

HL

AL/

SVL

Cytoch. b /

12S rRNA

P. nebulosus

44 DNA analysis

only

Tabarca

(TU)

- - - - - - - - - - - - - - AY222520/

AY222476

45 DNA analysis

only

5Km of

Nefza (TU)

- - - - - - - - - - - - - - AY222522/

AY222478

46 DNA analysis

only

South of

Tabarca

(TU)

- - - - - - - - - - - - - - AY222543/

AY222499

47 DNA analysis

only

South of

Tabarca

(TU)

- - - - - - - - - - - - - - AY222542/

AY222498

48 DNA analysis

only

Jendouba

(TU)

- - - - - - - - - - - - - - AY222540/

AY222496

49 DNA analysis

only

Near

Fernana

(TU)

- - - - - - - - - - - - - - AY222541/

AY222497

50 DNA analysis

only

Ain Draham

(TU)

- - - - - - - - - - - - - - AY222521/

AY222477

51 DNA analysis

only

Dam Bou-

Heurtma

(TU)

- - - - - - - - - - - - - - AY222535/

AY222491

52 DNA analysis

only

Dam Bou-

Heurtma

(TU)

- - - - - - - - - - - - - - AY222538/

AY222494

53 DNA analysis

only

Dam Bou-

Heurtma

(TU)

- - - - - - - - - - - - - - AY222526/

AY222482

54 DNA analysis

only

Dam Bou-

Heurtma

(TU)

- - - - - - - - - - - - - - AY22251/

AY222473

55 DNA analysis

only

Dam Bou-

Heurtma

(TU)

- - - - - - - - - - - - - - AY222539/

AY222495


488ZOOTAXAtion (TBR) branch swapping with 100 random stepwise additions of taxa. Nodal support

was assessed by bootstrap (Felsenstein, 1985) involving 1000 pseudo-replications.

FIGURE 1. Map of North Africa showing localities of Pleurodeles used in the present study. SeeTable 1 and Fig. 5 for further details. The dashed line delimits the approximate distribution range ofP. poireti. We refer to it in the text as the Edough Peninsula.


488ZOOTAXA

FIGURE 2. Schematic drawing showing all eight morphometric measurements used in the presentstudy. SVL—snout-vent length; TL—tail length; Sp3T—right hind limb shank length; 3T—righthand third toe length; AL—length of the anterior right arm; HW—head width; IOW— interorbitalwidth; HL—head length.

Morphometric analysis

A total of 41 P. poireti were included in the analyses. Eight measurements were takenusing a calliper to the nearest 0.01 mm (see Fig. 2): snout-vent length (SVL), head length


488ZOOTAXA(HL), head width (HW), interorbital width mesured between the bonny margins of the

orbit (IOW), tail length, shank length (SP3T), third toe length (3T), right forearm length(AL). Some specimens of P. poireti were missing the gular fold (see Pasteur, 1881;Lataste, 1958), so for HL measurements we measured from the tip of the snout to the ret-roarticular process in the articulation of the mandible with the quadrate (see Fig. 2). Statis-tica 5.5 was used for statistical analysis. Normality of the data was determined using theShapiro-Wilks’W test implemented in Statistica 5.5. When not normal, data was log-trans-formed in order to meet better the assumption of normality.

In order to assess differences in the vomerine teeth of both species (Lataste, 1881),warmed plasticine was placed in the mouth to the distance of the vomerine tooth bands.Pressure was applied (the jaws closing) leaving the impression of the teeth and the choane.Scaled drawings were made of the impressions using a camera lucida.

Results

The morphological and the molecular analysis clearly indicate that P. poireti consists oftwo full species, one from the Edough Peninsula and the other one throughout the rest ofits range in Algeria and Tunisia. As explained in the introduction, the Edough Peninsulaform is the one to which the name poireti Gervais correctly applies; the earliest availablename for the other form is nebulosus Guichenot. Examples of some specimens of P. poiretiand P. nebulosus are shown in Fig. 3.

Pleurodeles poireti (Gervais, 1835)

Lacerta palustris: Poiret (part), 1789: 290Triton Poireti Gervais (part), 1835: 112. Gray, 1850: 18Glossoliga Poireti: Bonaparte (part), 1839: unnumbered. Gervais, 1853: 312Euproctus Poireti: Duméril & Bibron (part), 1854: 160Glossoliga Hagenmulleri Lataste, 1881: 26Molge hagenmulleri: Boulenger, 1882: 26. 1891: 93Triturus hagenmulleri: Camerano, 1885: 419. Noble, 1924: 304Triton (Pleurodeles) Hagenmulleri: Wolterstorff, 1901: 9Pleurodeles poireti subsp. Hagenmulleri: Wolterstorff, 1905: 263Pleurodeles poireti: Pasteur (part) 1958: 160. Schleich et al. 1996: 94

Diagnosis: A dwarf form of Pleurodeles usually not exceeding 129 mm in total length.Restricted essentially to the Edough Peninsula and lowland surrounding areas (Carranzaand Arnold, 2003; Veith et al, 2004; see Fig. 1). Unique DNA sequences of the cyto-chrome b and 12S rRNA mitochondrial genes (sequences submitted to Genbank for com-parison; see Table 1 for accession numbers). Differentiated from the closely related P.nebulosus on account of the smaller size from the snout to the tip of the tail (t-value = –


488ZOOTAXA 4.50078; P = 0.00006; total size range of P. poireti 93–128.5 mm) and an average uncor-

rected genetic distance of approximately 7.7% for the cytochrome b and 3.7% for the 12SrRNA mitochondrial regions sequenced for the present study. Genetic variability within P.poireti is 0% for the cytochrome b and 0% for the 12S rRNA although only two specimensfrom the same locality have been analysed (see Table 1 and Fig. 1). Males of P. poireti dif-fer from males of P. nebulosus in having smaller snout-vent length (39.8–49.5 mm versus53–69 mm), head length (10.04–12.9 mm versus 14.5–17.3 mm), head with (8.06–9.67mm versus 10–13.56 mm), interorbital width (3.34–3.94 mm versus 4.28–5.8 mm), taillength (50–73 mm versus 62–90 mm), anterior arms length (11.8–17 mm versus 17.2–24

mm), 3rd toe length (4.22 – 6.4 mm versus 5.2–7.13 mm) and length from the elbow to thethird toe (10.76–15.3 mm versus 14.1–18 mm) (see Table 4 for statistics on all measure-ments). Females of P. poireti are differentiated from females of P. nebulosus by the firsthaving shorter interorbital width (3.7–4.35 mm versus 4.35–5.76 mm) and anterior arms(11.9–16.2 mm versus 13–22 mm) (see Table 5 for statistics on all measurements). Somespecimens of P. poireti have clearly a V shaped row of palatine teeth (see Fig. 4). Differen-tiated from P. waltl by its smaller size, in having usually 14 instead of 15 presacral verte-brae, in lacking sharp rib tips that project through a row of glandular swellings on theflanks and a tubercular process on rib 3 and often traces of this structure on ribs 1 and 2. Italso differs from P. waltl in 9.2% of the cytochrome b and 3.7% of the 12S rRNA mito-chondrial regions sequenced for the present study.

The two syntypes were part of a larger series with localities in addition to Bône(Annaba). Therefore, it seems advisable to designate a lectotype. Specimen MNHNP4744A is selected on the grounds of its being complete (see Fig. 3, specimen 9).

Description of lectotype: MNHNP 4744A. Male collected in Bône (Annaba), Algeriaby Marley and Gérard; 93 mm in total length, see Table 1 for other relevant measurements.Preserved-hardened accompanied by some shrinkage. Dorsum and most of the tailstrongly rugose and tuberculate, diminishing in coarseness towards the belly and the head,which anteriorly is almost smooth. Belly slightly rugose and gular fold pronounced. Tes-ticular swelling moderately conspicuous. Anterior humoral baldges present. Black suffu-sion dorsally and dorsolaterally, coalescing into marbeling and ill-defined blotches.

Colour in alcohol: rust with paler tubercles, lateral glandular protuberance and dorsalprotuberances. Throat, belly, underside of the tail and digits pale. Transitions from dark topale on the sides to the belly rather abrupt. Throat lightly spotted, belly very sparsely so.

Description of the paralectotype: MNHNP 4744 (see Fig. 3, specimen 8). Female col-lected in Bône (Annaba), Algeria by Marley and Gérard. Right scapula and forearmremoved. Total length 107 mm. For other measurement see Table 1. Colour and texturesimilar to the lectotype (MNHNP 4744A). The gular fold is virtually absent.


488ZOOTAXA

FIGURE 3. Photograph showing nine specimens of P. pioireti (above) and four P. nebulosus(below). A 23 centimetres scale bar is shown on the left-hand side of the picture; black rectanglesand intermediate white spaces all represent 1 cm. Numbers above the specimens refer to: 1. BMNH1920.1.20.1327.2, largest specimen of P. poireti included in the present study. Female from Bône(Annaba); 2. BMNH 1946.9.6.77, male of P. poireti from Mount Edough; 3. BMNH 1946.9.6.78,male of P. poireti from Mount Edough; 4. BMNH 1946.9.6.79, male of P. poireti from MountEdough; 5. BMNH 1946.9.6.80, male of P. poireti from Mount Edough; 6. BMNH 1946.9.6.81,male of P. poireti from Mount Edough; 7. BMNH 1946.9.6.79, male of P. poireti from MountEdough; 8. MNHNP 4744, female, paralectotype of P. poireti from Bône (Annaba); 9. MNHNP4744A, male, lectotype of P. poireti from Bône (Annaba); 10. BMNH 1.1.3.1.a, largest specimen ofP nebulosus recorded to date. Male from N. Africa; 11. BMNH 88.4.9.3, female of P. nebulosusfrom Algiers; 12. BMNH 88.4.4, male of P. nebulosus from Algiers; 13. MNHNP 1442, female,lectotype of P. nebulosus from Algiers.

Pleurodeles nebulosus (Guichenot, 1850)

Lacerta palustris: Poiret (part), 1789: 290Triton Poireti Gervais (part), 1835: 112. Gray, 1850: 18Glossoliga Poireti: Bonaparte (part), 1839: unnumbered. Gervais, 1853: 312


488ZOOTAXA Euproctus Rusconii: Guichenot (part), 1850: 29

Euproctus Poireti: Duméril & Bibron (part), 1854: 160Glossoliga Poireti Lataste, 1881: 26Molge poireti: Boulenger, 1882: 26. 1891: 93Triturus poireti: Camerano, 1885: 419. Noble, 1924: 304Triton (Pleurodeles) Poireti: Wolterstorff, 1901: 9Pleurodeles Poireti subsp. Poireti: Wolterstorff, 1905: 263Pleurodeles poireti: Pasteur (part) 1958: 160. Schleich et al. 1996: 94

Diagnosis: A medium size Pleurodeles usually up to 180 mm in total length but exception-ally reaching 230 mm (BMNH 1.1.3.1.a. This very large P. nebulosus was discoveredamong P. waltl specimens at the NHM, london. The results of the x-rays clearly showedthat it has 14 presacral vertebrae and that it lacks the typical P. waltl tubercular process onrib 3 and traces of this structure on ribs 1 and 2. Moreover, as in P. nebulosus the specimenalso lacks the typical P. waltl sharp rib tips and a row of glandular swellings on the flanks.It was identified as P. nebulosus and not P. poireti by means of its bigger size). Distributedacross northern Algeria and Tunisia, except in the Edough Peninsula and surrounding low-land areas (Carranza and Arnold, 2003; Veith et al. 2004; see Fig. 1). Unique DNAsequences of the cytochrome b and 12S rRNA mitochondrial genes with low intraspecificvariability (sequences submitted to Genbank for comparison; see Table 1 for accessionnumbers). Differentiated from the closely related P. poireti on account of its bigger totalsize (t-value = –4.50078; P = 0.00006; total size range of P. nebulosus 94 – 230 mm) andan average uncorrected genetic distance of approximately 7.7% for the cytochrome b and3.7% for the 12S rRNA mitochondrial regions sequenced for the present study. Geneticvariability within P. nebulosus is 0.5% for the cytochrome b and 0.8% for the 12S rRNA(21 specimens analysed covering almost the whole distributional range of the species; seeTable 1 and Fig. 1). Males of P. nebulosus differ from males of P. poireti in having largersnout-vent length (53–69 mm versus 39.8–49.5 mm) head with (10–13.56 mm versus 8.06–9.67 mm), interorbital width (4.28–5.8 mm versus 3.34–3.94 mm), tail length (62–90 mm

versus 50–73 mm), anterior arms length (17.2–24 mm versus 11.8–17 mm), 3rd toe length(5.2–7.13 mm versus 4.22–6.4 mm) and length from the elbow to the third toe (14.1–18mm versus 10.76–15.3 mm) (see Table 4 for statistics on all measurements). Females of P.nebulosus are differentiated from females of P. poireti by their larger interorbital width(4.35–5.76 mm versus 3.7–4.35 mm) and anterior arms (13–22 mm versus 11.9–16.2 mm)(see Table 5 for statistics on all measurements). Some specimens of P. nebulosus have aclearly U shaped row of palatine teeth; none of the specimens studied presents clearly Vshaped row of palatine teeth (see Table 1 and Fig. 4). Differentiated from P. waltl by itssmaller size, in having usually 14 instead of 15 presacral vertebrae, in lacking sharp ribtips that project through a row of glandular swellings on the flanks and a tubercular pro-cess on rib 3 and often traces of this structure on ribs 1 and 2. It also differs from P. waltl in9.7% of the cytochrome b and 4.8% of the 12S rRNA mitochondrial regions sequenced forthe present study.


488ZOOTAXA

FIGURE 4. Schematic dia-grams of the palatine teethof P. poireti and P. nebulosusobtained with plasticinecasts (see material andmethods). Numbers insidethe drawings refer to speci-mens codes from Table 1.The position of the choanaewas also obtained in thecasts and it has been high-lighted above the palatineteeth. Every drawing has ascale bar to the left, whichcorresponds to 2 mm.


488ZOOTAXA The type series of P. nebulosus consists of skeletal material and one partially dissected

spirit specimen (MNHNP 1442), which we designate as the lectotype. Description of lectotype: MNHNP 1442 (see Fig. 3; specimen 13). Female collected in

Algiers by Guichenot; lower jaw and throat dissected, tip of the tail missing; 147.5 mm intotal length; head moderately depressed; dorsally and dorsolaterally moderately tostrongly rugose/ turberculate; tubercles tend towards disposition in transverse irregularseries separated by creases becoming progressively smaller and more numerous verte-brally and on the belly; the head is smoother; gular fold present.

Colour in alcohol: dorsum very dark, tubercles little paler (if at all) becoming increas-ingly paler towards the belly; throat and palmar aspects of hands paler of yellowish ochrecoloration; lateral glandular protuberances barely noticeable; dorsal protuberances incon-spicuous anteriorly, slightly evident posteriorly, coalescing at the level of the cloaca to acontinuous line on the tail.

Phylogenetic analysis

The results of the phylogenetic analysis are presented in Fig. 5 and show that the two indi-viduals of P. poireti from Bône (Annaba) region represent an independent phylogeneticlineage sister to P. nebulosus. All three species of Pleurodeles are monophyletic with highbootstrap support. To calibrate the molecular clock we assumed that P. waltl separatedfrom the ancestor of P. poireti and P. nebulosus at the end of the Mesinian Salinity Crisis,some 5.3 Myr ago (Hsu, 1972, 1973; Blondel and Aronson, 1999; Krijgsman et al., 1999;Duggen et al., 2003). This date marks the disappearance of the land bridge that for morethan 600,000 years had connected southern Spain with North Africa, and had facilitatedgreatly the faunal exchange between both continents. Therefore, for many animals that, asthe urodels, do not disperse easily across bodies of salt water, the end of the MessinianSalinity Crisis 5.3 Myr ago was a very important vicariant event, and therefore an excel-lent calibration point (see Carranza and Arnold, 2003 for a discussion on the alternativeclock calibration using the less reliable fossils data available for this group of urodels).According to our calibration, P. poireti and P. nebulosus have been evolving as indepen-dent lineages for approximately 4.2 Myr (see Carranza & Arnold, 2003). Genetic distancesamong all three species of Pleurodeles are very similar for the 12S rRNA gene (3.7% forP. poireti vs P. waltl; 3.7% for P. poireti vs P. nebulosus and 4.8% for P. waltl vs P. nebulo-sus) and not so different for the cytochrome b gene (9.2% for P. poireti vs P. waltl; 7.7%for P. poireti vs P. nebulosus and 9.7% for P. waltl vs P. nebulosus).


488ZOOTAXA

FIGURE 5. ML tree (Log likelihood -2240.36916; GTR+G model of sequence evolution) inferredfrom cytochrome b and 12S rRNA sequences. Bootstrap support is shown by the branches. Num-bers separated by a dash from the species names refer to specimen codes from Fig 1 and Table 1.The estimated date for the separation between P. poireti and P. nebulosus and for the calibrationpoint used in the analysis is marked with a filled circle.

Morphometric analysis

An analysis of sexual dimorphism within P. poireti and P. nebulosus was carried out usinga student-t and indicated that males and females of both species differ statistically in some


488ZOOTAXA of the variables used (see Tables 2 and 3). Our limited sampling suggests sexual dimor-

phism is more accentuated in P. poireti, with males being significantly smaller (SVL) thanfemales, having shorter and narrower heads, shorter interorbital width but having rela-tively longer tails and anterior arms. Sexual dimorphism in P. nebulosus is less pro-nounced than in P. poireti, with males only having absolute and proportionally longer tailsand longer forearms than females. As a result of the positive results of the sexual dimor-phism males and females of both species were analysed separately. The results of the mor-phological comparisons are presented in Tables 4 (males) and 5 (females). Males of P.poireti and P. nebulosus are significantly different in all 8 variables measured. However,they do not differ in any of the 5 body proportions calculated suggesting that there is hasbeen an isometric reduction in size in males of P. poireti. Values for all 8 variables that aresignificantly different between males of both species are always higher for P. nebulosusthan for P. poireti. Females of P. poireti and P. nebulosus are not so different, with P valuesbeing significant only for the difference in the interorbital width and the length of the ante-rior arms. As in males, P. nebulosus females always present higher values than P. poiretifemales in all measurements.

TABLE 2. Analysis of sexual dimorphism between 8 males and 11 females of Pleurodeles nebulo-sus using a student-t test for 13 discrete measurements and body proportions. Significant P valuesat the P < 0.05 level have been highlighted with an asterisk and at the P < 0.01 with two asterisks.

Variable Mean (Males) Mean (Females) t-Value P

SVL 60.575 59.336 0.261 0.796

HL 15.138 14.342 0.684 0.502

HW 12.045 11.391 0.788 0.441

IOW 5.055 4.966 0.362 0.721

TL 79.425 64.654 2.412 0.0274*

SP3T 16.968 15.598 1.253 0.227

T3 6.316 5.572 2.064 0.054

AL 19.725 16.127 3.250 0.0047**

TL/SVL 1.319 1.088 3.575 0.0023**

HL/SVL 0.250 0.244 0.436 0.667

HW/SVL 0.199 0.194 0.641 0.529

HW/HL 0.796 0.799 -0.164 0.871

AL/SVL 0.327 0.275 3.614 0.0021**


488ZOOTAXATABLE 3. Analysis of sexual dimorphism between 16 males and 6 females of Pleurodeles poireti

using a student-t test for 13 discrete measurements and body proportions. Significant P values at theP < 0.05 level have been highlighted with an asterisk and at the P < 0.01 with two asterisks.

Discussion

The molecular and morphological evidence presented here supports Lataste’s (1881)hypothesis that a different species of Pleurodeles lives in the Edough Peninsula in north-east Algeria (see Fig. 1). Our results, are at variance with Pasteur (1958), who recognisedonly one species of Algero-Tunisian Pleurodeles. One of the possible explanations as towhy Pasteur (1958) decided to synonymise both forms is because two MNHNP specimensfrom Tunisia (27.27 and 27.28) labelled as P. hagenmulleri (=P. poireti) included in hisstudy were, in fact, P. nebulosus. According to our analyses, all Algero-Tunisian Pleurode-

les populations outside the Edough Peninsula (Fig. 1) are genetically and morphologicallyP. nebulosus. This includes all 14 Tunisian individuals from 7 different localities (some ofthese very close to the Algerian border). This has important implications, especially sincePasteur (1958) did not include many P. hagenmulleri (=P. poireti) in his work, and thesetwo Tunisian specimens had the morphological characteristics of P. nebulosus. The muchlarger size of the two MNHNP Tunisian specimen 27.27 (135 mm) and 27.28 (179 mm)was used by Pasteur (1958) to seriously question the validity of Lataste’s (1881) new spe-cies. According to our measurements, the maximum total size of P. poireti is 129 mm,while the much bigger P. nebulosus can easily reach 180 mm and exceptionally 230 mm(see Fig. 3, specimen 10). This size difference is statistically significant (t-value = -

Variable Mean (Males) Mean (Females) t-Value P

SVL 45.487 50.333 -3.043 0.0064**

HL 11.233 12.350 -3.494 0.0022**

HW 8.787 9.508 -3.398 0.00284**

IOW 3.710 4.040 -2.838 0.01014*

TL 61.025 56.766 1.285 0.213

SP3T 13.485 13.728 -0.382 0.706

T3 5.553 5.593 -0.155 0.878

AL 14.787 13.550 1.684 0.107

TL/SVL 1.340 1.126 4.351 0.0003**

HL/SVL 0.247 0.246 0.057 0.954

HW/SVL 0.193 0.189 0.855 0.402

HW/HL 0.783 0.771 0.648 0.523

AL/SVL 0.325 0.269 4.138 0.0005**


488ZOOTAXA 4.50078; P = 0.00006) indicating that, as suggested by Lataste (1881), size is one of the

main diagnostic characters that differentiates between both forms of Algero-TunisianPleurodeles (see Tables 4 and 5 and Fig. 3).

Other diagnostic characters listed in Lataste (1881) were the shape of the row ofpalatine teeth (V shaped in the Edough Peninsula species and U shaped in all the rest), andsome subtle differences in the general shape of the head, the snout, and the contour of thejaws (semi-elliptical in P. poireti and semi-circular in P. nebulous). The drawings of thepalatine teeth shown in Fig. 4 indicate that, some Edough Peninsula P. poireti have aclearly V shaped row of palatine teeth (specimens 11 and 9 in Fig. 4). However, anddespite the high level of intraspecific variability, there is not a single specimen of P. nebu-losus in our sample with a clearly V shaped row of palatine teeth (see Fig. 4). Therefore,when present, the character palatine teeth forming a clearly V shaped row can be used todifferentiate between these two forms (Lataste, 1881; Boulenger, 1882; Doumerge, 1901).A drawing of the skull of an individual of each species is shown in Fig. 6 for comparison.Once more, despite the high level of intraspecific variability (Pasteur, 1958, SC and EWpers. obs.), there is a clear difference between both species in the general shape of theskull, snout and contour of the jaws (Lataste, 1881).

FIGURE 6. Ventral aspect of the skull of A.-adult P. poireti, BMNH 1920.1.20.1383 (Bône) and B.adult P. nebulosus, BMNH 130a (Algiers).

The high level of intraspecific morphological variability presented by the EdoughPeninsula P. poireti for most of Lataste’s (1881) diagnostic characters is due to the pro-nounced level of sexual dimorphism existent in this species. One of the consequences ofthis sexual dimorphism (see Table 3), is that the morphological differences between malesof P. poireti and P. nebulosus are greater than the differences between females of both spe-


488ZOOTAXAcies (Tables 4 and 5; Lataste, 1881). Pasteur (1958) analysed his material without any

apparent regard to sex, appearing to overlook sexual dimorphism in P. poireti and, in con-sequence, he was not able to correctly evaluate the level of morphological differencebetween both species. Had he analysed males and females separately, he would have beenless likely to discard Lataste’s (1881) diagnostic characters (see Table 4). Despite beinggenetically very different, the females of P. poireti and P. nebulosus included in our analy-sis are morphologically not so different (see Table 5), and therefore more difficult to dif-ferentiate using only Lataste’s (1881) diagnostic characters. As it is often the case,especially among species of Triturus, the males possess the sexually dimorphic characters,whereas females of closely related species are more uniform and therefore more difficultto differentiate.

TABLE 4. Results of the t-test for 13 morphological characters between males of P. poireti and P.

nebulosus. Significant P values at the P < 0.01 have been highlighted with two asterisks.

Differences in sexual dimorphism between populations or, like in this case, closelyrelated species could be caused by many factors, including differences in climatic condi-tions, resource partitioning with other species, sexual selection, etc. In the Pyrenean Brooknewt (Euproctus asper), for instance, differences in sexual dimorphism observed betweenalpine Central Pyrenean populations and Prepyrenean ones are thought to be caused byincreased selection pressures favouring high reproductive efficiency and reduced intersex-ual competition under alpine, high-mountain climate conditions (Serra-Cobo et al. 2000).Unfortunately, very little is known about the ecology and life history of the populations of

Variable Mean (P. poireti)

Mean (P. nebulosus)

t-Value P

SVL 45.487 60.575 -8.494 0.000**

HL 11.233 15.138 -9.345 0.000**

HW 8.787 12.045 -10.802 0.000**

IOW 3.710 5.055 -9.761 0.000**

TL 61.025 79.425 -5.269 0.000**

SP3T 13.485 16.968 -6.119 0.000**

T3 5.553 6.316 -2.901 0.008**

AL 14.787 19.725 -6.386 0.000**

TL/SVL 1.340 1.319 0.327 0.746

HL/SVL 0.247 0.250 -0.583 0.565

HW/SVL 0.193 0.199 -1.363 0.186

HW/HL 0.783 0.796 -0.807 0.428

AL/SVL 0.325 0.327 -0.134 0.894


488ZOOTAXA P. poireti from the Edough Peninsula and the widespread Algero-Tunisian P. nebulosus.

Until more data is available, it will not be possible to understand which factors are causingthe differences in sexual dimorphism between these two different species.

TABLE 5. Results of the t-test for 13 morphological characters between females of P. poireti and P.nebulosus. Significant P values at the P < 0.05 level have been highlighted with an asterisk and atthe P < 0.01 with two asterisks.

Possible origin of P. poireti

The data presented here indicates that P. poireti is restricted to a relatively small area in theNorth-east of Algeria (see Fig. 1). This region, referred throughout the paper as theEdough Peninsula, is a mountainous area with some peaks reaching up to 850 m above thesea level. Unlike other mountainous parts of the Algero-Tunisian coast, the Edough Penin-sula is connected to the mainland through an area of lowland marshes with an importantfluvial system and a relatively large brackish lake (Lac Fetzara). This area of lowlandmarshes surrounding the Edough Peninsula, probably represents the southern limit of P.poireti, which extends as far as 25 Km to the southeast of Annaba (Carranza and Arnold,2003; Veith et al. 2004; see Fig. 1).

The molecular analysis indicates that speciation between P. poireti and P. nebulosusoccurred approximately 4.2 Myr ago, during the Upper-middle Pliocene. This may beexplained by assuming that, shortly after the land-bridge between Africa and Europe dis-appeared approximately 5.3 Myr ago, a population of the ancestor of P. poireti and P. neb-

Variable Mean (P. poireti)

Mean (P. nebulosus)

t-Value P

SVL 50.333 59.336 -1.709 0.107

HL 12.350 14.342 -1.562 0.139

HW 9.508 11.391 -2.089 0.054

IOW 4.040 4.966 -3.699 0.002**

TL 56.766 64.654 -1.209 0.245

SP3T 13.728 15.598 -1.491 0.156

T3 5.593 5.572 0.055 0.956

AL 13.550 16.127 -2.295 0.036*

TL/SVL 1.126 1.088 1.004 0.330

HL/SVL 0.246 0.244 0.154 0.879

HW/SVL 0.189 0.194 -0.435 0.669

HW/HL 0.771 0.799 -1.212 0.243

AL/SVL 0.269 0.275 -0.535 0.600


488ZOOTAXAulosus became isolated in the Edough Peninsula. During The Pliocene, global changes in

the sea-level, crustal movements and local landform changes affected the planet causingmarine transgressions to inundate many coastal areas all over the planet (Lanza, 1984;Morafka, 1976; and references therein). These Pliocene marine transgression may haveinundated the lowland area surrounding the Edough Peninsula, which would have becomea temporary island. In biogeographical terms, such temporary islands are known as conti-nental fossil islands (Lanza, 1984). The existence of Pliocene fossil islands and their bio-geographic importance has been studied in depth for the Toscana region in Italy (Lanza,1984) and for other regions as far as central coastal California (Santa Lucia and GabilanRanges) (Morkafa and Banta, 1973; Morafka, 1976). In both cases, it was shown that thelevel of endemicity in these regions was much higher than in the surrounding areas. Forinstance, in coastal California, Peabody and Savage (1958) and Morafka and Banta (1973)showed a clear correlation between Pliocene embayment and subspecific boundaries ofherpetofauna in this region. Stebbins and Major (1965) and Jepson (1925) reported thepresence of plant endemic species in ranges that were formerly islands (i.e. Santa Luciaand Franciscan). An endemic rodent species (Diplodomys elephantinus) is known for theGabilan fossil island, in coastal California (Morafka, 1976). In Europe, the level of ende-micity in fossil islands is also high and, among animals, it includes several species of gas-tropods, terrestrial isopodes, coleopterans, a murine rodent and a giant hedgehog (Lanza,1984, Butler, 1980). The presence of the endemic P. poireti and the geology of the areasuggest that the Edough Peninsula might be a Pliocene fossil island. If that were so, wewould expect to find other endemic plants and animals. Among amphibians, good candi-dates to test the fossil island hypothesis are Hyla meridionalis, Salamandra algira, Bufobufo, Bufo mauritanicus, Bufo calamita and Rana saharica. Of all these species, only S.algira has been studied from the genetic point of view (Steinfart et al. 2000). This studyshowed that the Annaba specimens were genetically very different from the Moroccanones. But, apart from those from Annaba, no other samples of S. algira from Algeria wereincluded in the study, so it is not possible to know if there is an endemic Salamandra fromthe Edough Peninsula yet. Future studies should focus in this interesting area of north-eastern Algeria in order to see if there are any other animal or plant endemics and to findany geological clues that may support the fossil island hypothesis.

Acknowledgements

We are indebted to E. N. Arnold, J. Roca and S. Arroyo for their help. We appreciate thecontribution of D. Donaire, R. Bour (MNHNP), Soumia Fahd, R. Fonoll, B. Clark(BMNH), O. Arribas and D. Siebert in several aspects of this work. We are also very grate-ful to the authors of the “Atlas de Andalucia” (J. P. González de la Vega, J. M. Barnstein,D. Donaire and L. García) for all the samples of Pleurodeles from southern Spain used in


488ZOOTAXA this study. Salvador Carranza is supported by a Ramón y Cajal contract from the Ministe-

rio de Educación y Cultura, Spain.

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