Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic Author Dufresnes, Christophe Author Strachinis, Ilias Author Tzoras, Elias Author Litvinchuk, Spartak N. Author Denoel, Mathieu text ZooKeys 2019 859 131 158 http://dx.doi.org/10.3897/zookeys.859.33634 journal article http://dx.doi.org/10.3897/zookeys.859.33634 1313-2970-859-131 Pelobates varaldii Pasteur & Bons, 1959 Diagnosis . A smaller version of P. cultripes (Fig. 2) differing by a few phenotypic features. Unlike P. cultripes , the black coloration of the spades is often concentrated on the edges ( Pasteur and Bons 1959 ; Busack et al. 1985 ). The cranial braincase is high and narrow in P. varaldii , while it is low and wide in P. cultripes ( Pasteur and Bons 1959 ; Rocek 1981 ). The background coloration can be yellow, gray, and brown, with dark reticulate patches, and the dorsal surface is abundantly covered by orange dots, most pronounced on the eyelids (usually absent in P. cultripes ; Pasteur and Bons 1959 ; Beukema et al. 2013 ; Fig. 3). Males are usually smaller than females (Fig. 2). Average SVL = 53 mm (range: 36-66 mm) for females (n = 4 populations) and 51 mm ( 33-65 mm) for males (n = 4 populations) (Suppl. material 1, Table S1; Fig. 2). The karyotype includes six large and seven small pairs of two-armed chromosomes. Large centromeric C-bands appears in the pairs 1, 2, 4, 9, and 12; pericentric bands in the short arms of pair 1 and long arm of pair 8; telomeric bands in the long arms of pairs 1, 2, and 11; the short arm of pair 7 is almost heterochromatic ( Herrero and Talavera 1988 ). The nuclear DNA content averages 7.3 pg ( Litvinchuk et al. 2013 ). As shown in Table 1, P. varaldii differs from P. cultripes by 6.0% at mtDNA and 0.40% at nuclear DNA ( Dufresnes et al. 2019b ). Figure 2. Between-population variation of average size ( snout-vent length - SVL) for each Pelobates species, measured separately for females (pink) and males (blue). This compiles average size-data from 82 populations, representing at least 6,004 individuals (Suppl. material 1, Table S1). For P. balcanicus , it only includes populations from the nominal P. b. balcanicus . For P. syriacus , it only includes populations from P. s. boettgeri . Figure 3. Color variation in Pelobates cultripes , P. varaldii , P. fuscus and P. vespertinus . Photo credits and origins as follows a CD ( Herault , France) b, c CD (Algarve, Portugal) d A Sanchez Vialas (Spain) e G Martinez (Kenitra, Morocco) f-h A Sanchez Vialas (Tanger, Morocco) i, j N Suriadna (Ukraine) k CD (Wojewodztwo podkarpackie, Poland) l A Noellert (Burgenland, Austria) m-p N Suriadna (Ukraine). Table 1. Pairwise % of genetic differences between Pelobates taxa (from the data of Dufresnes et al. 2019b ). The estimates below diagonal correspond to mitochondrial DNA (cyt-b + 16S, 1.2 kb); the estimates above diagonal correspond to nuclear DNA (63.5 kb of RAD tags).

P. cultripes	P. varaldii	P. fuscus	P. vespertinus	P. s. syriacus	P. s. boettgeri	P. b. balcanicus	P. b. chloeae
P. cultripes
P. varaldii
P. fuscus
P. vespertinus
P. s. syriacus
P. s. boettgeri
P. b. balcanicus
P. b. chloeae

Taxonomy. The nomen Pelobates varaldii Pasteur & Bons, 1959 is the only one ever proposed for the Moroccan populations of spadefoot toads; holotype: MNHN-RA-1959.1; type locality: Merja Samora, Morocco. The ancient split of P. varaldii , dating back to the Messinian Salinity Crisis (5.3 My), supports its specific distinction from P. cultripes ( Busack et al. 1985 ; Crottini et al. 2007 ). Distribution. It is endemic to north-western Morocco (0-350 m elevation a.s.l.), found along the Atlantic coast, from the south of Tanger to Oualidia, where it is rare ( de Pous et al. 2012 ; Beukema et al. 2013 ; Frost 2019 ). IUCN status: Endangered ( Salvador et al. 2009 ). Diversity. To our knowledge, P. varaldii has not been the focus of any phylogeographic or population genetic work.