Pseudocyclophis, Boettger, 1888

Rajabizadeh, Mahdi, Nagy, Zoltán T., Adriaens, Dominique, Avci, Aziz, Masroor, Rafaqat, Schmidtler, Josef, Nazarov, Roman, Esmaeili, Hamid Reza & Christiaens, Joachim, 2016, Alpine-Himalayan orogeny drove correlated morphological, molecular, and ecological diversification in the Persian dwarf snake (Squamata: Serpentes: Eirenis persicus), Zoological Journal of the Linnean Society 176, pp. 878-913 : 898-899

PSEUDOCYCLOPHIS

The biometric, geometric morphometric, molecular phylogenetic, and habitat analysis data showed significant differences amongst the eastern, western, and nigrofasciatus OTUs. There are also distinctive groupings within the eastern and western OTUs.

The type locality of E. persicus (Anderson, 1987) is located in south-western Iran (the locality specified with circle 29 in Fig. 2 ); hence, specimens belonging to the south-western Iranian sub-OTU are assigned to E. persicus (Anderson, 1987) species. The specimens from western Iran and south-eastern Turkey form a monophyletic lineage, morphologically and ecologically distinguishable from E. persicus (Anderson, 1987) . Given this evidence, we conclude that these populations represent an undescribed species of the subgenus Pseudocyclophis . No specimens with morphological characters intermediate between those of the southwestern Iranian populations and those of the western Iranian and south-eastern Turkish populations were observed. The specimens found around Saveh and Tehran (locality numbers 19 and 20 in Fig. 2 ) are morphologically similar to E. persicus (Anderson, 1987) , genetically closer related to the populations in western Iran and south-eastern Turkey than E. persicus (Anderson, 1987) , and ecologically occur in a habitat that is different from those of both E. persicus (Anderson, 1987) and the new species. We hypothesize that the specimens from northern Iran are isolated and in the process of speciation. As we have few specimens from northern Iran, we could not assess the taxonomy of them properly and therefore leave the exact status of the northern Iran populations for future research.

Specimens of the nigrofasciatus morph are a monophyletic lineage, morphologically distinguishable from both the western and eastern OTUs and ecologically different from the eastern OTU. Given these data, we suggest that the subspecific status of E. persicus nigrofasciatus ( Nikolsky, 1907) should be changed to full species status. The sympatric occurrence of E. nigrofasciatus and E. persicus (Anderson, 1987) in south-western Iran (the locality indicated by circle 25 and triangle 7 in Fig. 2 ; see also Haas & Werner, 1969), as well as the morphological and molecular differences between them support this classification.

Within the eastern OTU, specimens from northwestern Pakistan, as well as those from eastern Iran, Turkmenistan, and southern and western Pakistan, represent two monophyletic lineages, distinguishable from each other morphologically and inhabiting two significantly different habitats. The occurrence of two phenotypes of the subgenus Pseudocyclophis in Pakistan was also reported by Minton (1966). As the eastern and western OTUs are significantly different from each other in their morphological, molecular, and ecological characteristics, we conclude that both groupings within the eastern OTU are representative of two distinct species, differing from typical Eirenis persicus (Anderson, 1987) . Specimens from north-eastern Pakistan agree morphologically with the description of Contia (= Eirenis ) angusticeps Boulenger, 1894 . There are two synonyms available for the specimens from eastern Iran, Turkmenistan, and southern and western Pakistan: Pseudocyclophis (= Eirenis ) walteri Boettger, 1888 , from the Sarakhs region, north-eastern Iran (the locality indicated by squire 2 in Fig. 2 ), and Contia (Eirenis) zebrina Wall, 1923 , from the Bazman region, southeastern Iran (the locality indicated by squire 7 in Fig. 2 ). Based on the principle of priority, Article 23 of International Code of Zoological Nomenclature ( ICZN, 1999), the valid name of a taxon is the oldest available name applied to it. Therefore, we classify the specimens from eastern Iran, Turkmenistan, and southern and western Pakistan as Eirenis walteri Boettger, 1888 .

The biometric and geometric characteristics of the specimens of the novum OTU show no affinities to any of the identified groups within E. persicus . As only two specimens of this colour morph were examined, we were not able to reliably compare them with other populations of the subgenus Pseudocyclophis . As the distribution and number of subcaudal scales of these specimens are similar to those of E. nigrofasciatus , they probably just represent a colour variation of the nigrofasciatus pattern. Further investigations, including molecular studies, are needed to clarify the true nature of this new colour morph.

We did not examine any specimen of the E. persicus species group with 13 anterior dorsal scales that could be referred to Eirenis mcmahoni ( Wall, 1911) . However, based on the data that Wall (1911) presented in his description, and the data of Minton (1966), specimens referred to E. mcmahoni are all located in the Suleiman mountain range in central Pakistan and differ morphologically from any other species in the E. persicus species group by a higher ratio of tail length over total length, ranging from 30.64–32.00 (x = 31.20). Until there is further investigation, we retain this taxon provisionally as E. mcmahoni .

Figures

Figure 2. All available distribution records of the Eirenis persicus species group. Circles indicate the E. persicus specimens, with the persicus morph bearing unicoloured dorsal scales; squires indicate specimens from eastern Iran, southern Turkmenistan, and southern and western Pakistan, having both persicus morph that base of their anterior dorsal scales are darker, and walteri morph; stars indicate specimens referred to Eirenis mcmahoni (Wall, 1911); plus symbols indicate specimens in north-eastern Pakistan, having both persicus morphs that base of their anterior dorsal scales are darker, and with walteri morph; asterisks indicate specimens with the novum pattern; triangles indicate specimens of the nigrofasciatus morph. Circle 29 indicates the type locality of Cyclophis persicus Anderson, 1872; squire 2 indicates the type locality of Pseudocyclophis walteri Boettger, 1888; squire 7 indicates the type locality of Contia zebrina Wall, 1923; triangle 6 indicates the type locality of Contia persica var. nigrofasciata Nikolsky, 1907; star 3 indicates the type locality of Contia angusticeps Boulenger, 1894; plus symbols 2–5 indicate the type series localities of Contia mcmahoni Wall, 1911. For more details, see Appendix 1.