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J.W. Arntzen and M. García-París

Three competing phylogenetic hypotheses for the genus Alytes (midwife toads) are evaluated. Based on quantitative coding of protein characters the most parsimonious solution shows a sister taxon relationship for Alytes dickhilleni and A. muletensis. The alternatives in which A. obstetricans has its sister group in either A. dickhilleni or A. muletensis lack support. Using calibrations derived from protein evolutionary rates, the vicariant events giving rise to A. obstetricans and the lineage leading to the A. muletensis and A. dickhilleni clade and the subsequent splitting between A. muletensis and A. dickhilleni cannot be placed much earlier than the Miocene-Pliocene boundary. Biogeographical scenarios invoking an earlier time of divergence should be rejected.

J.W. Arntzen and M. García-París

Allozyme variation in 31 to 50 presumptive loci of 12 populations of European midwife toads of the genus Alytes show appreciable genetic divergences (D Nei from 0.29 to 0.72) among four groups. These groups correspond to A. cisternasii Boscá, 1879, A. obstetricans (Laurenti, 1768), A. muletensis (Sanchiz & Adrover, 1979), and a new species from the Betic mountains described here as A. dickhilleni n. sp. Smaller divergences among geographic groups of populations of A. obstetricans (up to D Nei = 0.17) support the existence of three geographic units in Europe corresponding to the previously recognized subspecies A. o. obstetricans and A. o. boscai Lataste, 1879, plus a third one described here under the designation of A. o. almogavarii n. ssp. The analysis of morphological variation among taxa using principal component and discriminant analysis shows a remarkable similarity between A. dickhilleni and A. obstetricans; these are cryptic species. A phylogenetic analysis of allozyme data using A. cisternasii as the outgroup supports a sister taxon relationship between A. muletensis and A. dickhilleni, with A. obstetricans the sister taxon to this clade. The historical biogeography of the genus is discussed.

M. Garcia-Paris, J.W. Arntzen, B. Arano and P. Herrero


Allozyme and C-banding studies on Triturus alpestris cyreni from north and central Spain and of T. a. alpestris from west and central Europe show a distinct differentiation between the taxa. Within the Iberian subspecies a marked genetic differentiation is detected among those populations that are geographically isolated, with the exception of that from central Spain. One of the populations from northern Spain stands out as distinct on account of both cytogenetic and allozyme analyses.

M. García-París, P. Herrero, C. Martín, J. Dorda, M. Esteban and B. Arano

Triturus marmoratus pygmaeus, a problematicsubspecies of the Marbled Newt from the southern part of the Iberian Peninsula, is redescribed using specimens collected in the “typical” area. Diagnostic external morphological features are provided to permit the accurate determination of the specimens belonging either to T. m. marmoratus or to T. m. pygmaeus. These diagnostic features were applied to individuals both from the field and from museum collections. The results indicate a larger distributional area for to T. m. pygmaeusto than was previously recognized. The distribution of to T. m. marmoratusto ranges over the northern half of the Iberian Peninsula and most of France; to T. m. pygmaeusto occupies the southwestern part of the Iberian Peninsula. The contact area between the two subspecies seemsto be located alongthe Central Range Mountains (Sistema Central) in Portugal and Spain. to T. m. marmoratusto extends southwards beyond this borderline in three areas: Serra da Estrela (Portugal), Sierra de Gata (Spain) and Sierra de Guadarrama (Spain). The only point at which to T. m. pygmaeusto reaches northwards beyond the Central System is near Puerto de Malagón (Madrid Province, Spain). No cases of strict sympatry, nor individuals with intermediate morphologicalfeatures have been observed. The results of an extensive cytogenetical analysis do not show any differences between to T. m. pygmaeusto and to T. m. marmoratusto . Interestingly, however, the to T. m. pygmaeusto populations from Doñana (Huelva Province, Spain) showed an exclusive, though little differentiated, C-banding pattern.