Notes from the Anatolian underground: two new mole taxa from Eastern Turkey, together with a revised phylogeny of the genus Talpa (Mammalia: Eulipotyphla: Talpidae) Author Gündüz, İslam Department of Biology, Faculty of Sciences, Ondokuz Mayıs University, Samsun, Turkey gunduzi@omu.edu.tr Author Demirtaş, Sadık Department of Biology, Faculty of Sciences, Ondokuz Mayıs University, Samsun, Turkey Author Silsüpür, Metin Department of Biology, Faculty of Sciences, Ondokuz Mayıs University, Samsun, Turkey Author Özmen, Medine Department of Biology, Faculty of Sciences, Ondokuz Mayıs University, Samsun, Turkey Author Polly, P. David Department of Earth & Atmospheric Sciences, Indiana University, Bloomington, IN, USA Author Bilton, David T. School of Biological and Marine Sciences, University of Plymouth, Drake Circus, Plymouth PL 4 8 AA, Devon, UK & Department of Zoology, University of Johannesburg, PO Box 524, Auckland Park, Johannesburg 2006, South Africa text Zoological Journal of the Linnean Society 2023 2023-07-18 199 3 567 593 http://dx.doi.org/10.1093/zoolinnean/zlad049 journal article 10.1093/zoolinnean/zlad049 0024-4082 10469936 14A0E12E-1A69-4725-8F99-4747F284C738 Talpa hakkariensis sp.nov. Zoobank registration: urn:lsid:zoobank.org:act: 58C51143- 81D5-45C3-BC16-381429ADCE74 . Type locality: Turkey , Hakkari Province , Cemşililan yaylası c . 8 km NNE of Durankaya , 37˚38ʹN, 43˚37ʹE, 2700–2980 m ( Figs 1 , 5A ; Table 1 ) . Holotype : Adult female, specimen no. 1533 , stuffed skin labelled ‘13/08/21-1533 ♀ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası, Hakkari// 160-31-21 = 73.84 g .// 13, August, 2021’. Body frozen at –50˚C, internal organs in ethanol, skull extracted and labelled with specimen number ( Fig. 6A ). Collected by İ. Gündüz and S. Demirtaş . Sequences available from GenBank: ( Cytb haplotype: Hap.15, accession number OQ695523 ; BRCA2 haplotype: Hap.7, accession number OQ695507 ); standard voucher specimens (skin, frozen body, skull, and various tissues in ethanol) are deposited in the Department of Biology, Faculty of Sciences, Ondokuz Mayıs University ( OMU-IG/SD ), Samsun, Turkey. Paratypes (6): One male , specimen no. 1487, stuffed skin labelled ‘10/06/21-1487 ♂ IG/SD// Talpa hakkariensis sp. nov. // Bilmizit yaylası-Hakkari, 37˚36ʹN 43˚38ʹE// 164- 28-20 = 73.05 g .// 10, June, 2021 ’; one male , specimen no. 1488, stuffed skin labelled ‘10/06/21-1488 ♂ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası-Hakkari// 166- 30-20 = 77.38 g .// 10, June, 2021 ’; one male , specimen no. 1489, stuffed skin labelled ‘10/06/21-1489 ♂ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası-Hakkari// 170- 31-21 = 71.25 g .// 10, June, 2021 ’; one male , specimen no. 1490, stuffed skin labelled ‘10/6/21-1490 ♂ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası-Hakkari// 165-29- 21 = 67.96 g .// 10, June, 2021 ’; one female , specimen no. 1531, stuffed skin labelled ‘11/08/21-1531 ♀ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası-Hakkari// 147-33- 21 = 60.34 g .// 11, August, 2021 ’; one female , specimen no. 1532, stuffed skin labelled ‘12/08/21-1532 ♀ IG/SD// Talpa hakkariensis sp. nov. // Cemşililan yaylası-Hakkari// 154-30- 21 = 67.39 g .// 12. August, 2021 ’. Bodies frozen at –50˚C, internal organs in ethanol, skulls extracted and labelled with specimen numbers. Sequences available from GenBank: ( Cytb haplotypes: Hap.11–14, accession numbers OQ695519 – OQ695522 ; BRCA2 haplotypes: Hap.7–8, accession numbers OQ695507 – OQ695508 ); standard voucher specimens (skins, frozen body, skulls, and various tissues in ethanol) are deposited in the Department of Biology, Faculty of Sciences, Ondokuz Mayıs University ( OMU-IG/SD ), Samsun, Turkey . Measurements of holotype Body mass 73.84 g , head and body length 160 mm , tail length 31 mm , hindfoot length 21 mm , condylobasal length of skull 33.40 mm , maxillary tooth row 14.93 mm , breadth of braincase 16.90 mm , height of braincase 9.29 mm , breadth of rostrum over canines 5.02 mm , breadth of rostrum over molars 9.30 mm (Supporting Information, Table S3). Measurements of paratypes Means ± standard deviation (minimum–maximum). Linear measurements are in millimetres; body mass in grams. Sample size = 6 for all measurements. Body mass 69.56 ± 5.30 (60.34– 77.38), head and body length 161.00 ± 7.920 (147.0–170.0), tail length 30.17 ± 1.570 (28.0–33.0), hindfoot 20.67 ± 0.471 (20.0–21.0), condylobasal length of skull 32.88 ± 0.421 (32.10– 33.24), maxillary tooth row 14.15 ± 0.142 (13.88–14.31), breadth of braincase 16.30 ± 0.348 (15.8–16.88), height of braincase 9.31 ± 0.102 (9.20–9.48), breadth of rostrum over canines 5.24 ± 0.279 (4.88–5.59), breadth of rostrum over molars 9.86 ± 0.442 (9.24–10.4) (Supporting Information, Table S3). Diagnosis Very large mole; skull large with a stout, broad rostrum; dental formula of C3/3 I1/1 P4/4 M3/3. A member of the subgenus Talpa , closest to T. davidiana and T. streetorum morphologically and T. davidiana and T. talyschensis genetically ( Figs 2 , 3 , 6 ). Distinguished from both T. davidiana and T. talyschensis by the larger body size [ T. hakkariensis sp. nov. body mass 60.34– 77.38 g , head and body length 147–170 mm ; T. davidiana body mass 34.12–52.87 g , head and body length 128–153 mm ; T. talyschensis body mass 31–49 g , head and body length 104–114 mm ; ( Fig. 6A )]. Condylobasal length of skull in T. hakkariensis sp. nov. also consistently greater, 32.10–33.40 mm vs. 29.04–31.27 in T. davidiana studied here, and a mean of 31.10 mm in T. talyschensis (Zaitsev et al . 2014) . The new species also differs from T. davidiana davidiana on: the broader, blunter rostrum ( 9.24–10.40 mm over the molars in T. hakkariensis sp. nov. vs. 8.35–9.87 mm in T. davidiana ; Figs 7–10 ); typically four rather than three premolars in each maxilla ( Figs 7–10 ; only one of the nine T. davidiana davidiana specimens we examined had four premolars; see also: Kryštufek et al . 2001); 1st upper molar with small, reduced parastyle, always much more prominent in T. davidiana davidiana ( Fig. 11A ); larger hands and feet (see Fig. 12 ); and longer (28–33 vs. 16–18 mm ) and paler tail, with most tail hairs of more uniform length (compare Fig. 12D, E ). The broad snout of the new species resembles that of T. davidiana tatvanensis ssp. nov., but this taxon is much smaller than T. hakkariensis sp. nov. (see below) and has a relatively prominent parastyle on the first upper molar, as in T. davidiana davidiana . Distinguished from T. streetorum by the latter’s larger cranium (see above), in having the posteriormost mandibular foramen positioned more anteriorly, and having a coronoid process that is narrower in anterior–posterior breadth and inclined more anteriorly (Supporting Information, Fig. S6B ). The net p -distance in Cytb sequence between T. davidiana and T. hakkariensis sp. nov. is 10.50%; raw distance 11.33%. The net and raw K2P distances of 11.79% and 12.64%, respectively, between these taxa is in the range observed between other species of moles (e.g. within Talpa , interspecific net and raw K2P distances average c . 11.47% and 12.61%, respectively). In fact, the net genetic distance in Cytb sequence T . davidiana and T . hakkariensis sp. nov. is much higher than the raw (total) genetic distance between several recognized sister-species in Talpa (e.g. T . aquitania vs. T . occidentalis , T . caucasica vs. T . ognevi , and T . levantis vs. T . transcaucasica ) (Supporting Information, Tables S4, S5). Description A relatively large, heavy mole ( Figs 5B , 6A ), with a medium tail; 17.07–22.45% of head and body length. Tail ( Fig. 12D ) parallel-sided, with sparse, stiff, sub-erect, overlapping bristles; individual bristles mostly pale greyish, almost translucent, up to 25% blackish or with black cores; bristles relatively even in length; tail tip tapered and rounded, visible when bristles are moved. Overall appearance of tail paler than body. Dorsal and ventral pelage greyish brown throughout, with marked silvery sheen; individual hairs typically brownish at tips, greyish below. Body elongated. Palm of manus ( Fig. 12A ) large, broad, relatively flat; claws at ends of all five phalanges, relatively broad and flattened dorsoventrally, with convex upper and concave lower margins, flattened oval in cross-section. Feet ( Fig. 12D ) large, approximately two-thirds tail length. Muzzle ( Figs 5B , 6A ) elongated, with hairs, including longer sensory bristles, paler than on head. Snout pinkish with close, fine, small, pale hairs. Skull ( Fig. 7 ) large, with a stout, broad rostrum. Width across canines 15.0–17.05%, and width across molars 27.84–31.92% condylobasal length. Maxillary tooth row equals 42.36–44.7% of the condylobasal length. The braincase is broad and deep, flattened above; height of neurocranium 27.71–29.12% of condylobasal length. The posterior margin of palatine is usually slightly posterior to the imaginary line connecting the posterior alveolar margins of 3rd upper molars, and the anterior border of the infraorbital foramen is above the 2nd upper molar. Upper incisors of decreasing size; 3rd incisor approximately half the size of the 1st. Upper molars robust, decreasing in size 1–3. Fourth upper premolar largest; 1st intermediate and 2nd and 3rd small ( Fig. 7 ). First upper molar with very small, almost imperceptible parastyle ( Fig. 11A ), visible in holotype and some paratypes , but lost in specimens with substantial tooth wear; protocone much larger than paracone; mesostyle relatively flattened at apex, with clear mesal furrow. Second upper molar with well-developed parastyle, somewhat bilobed at apex in holotype and most paratypes ; protocone only slightly larger than paracone; mesostyle well developed and clearly bilobed at apex. Third upper molar with well-developed parastyle, slightly more prominent than mesostyle; mesostyle clearly bilobed at apex; protocone only slightly larger than paracone. Mandible Fig. 13A . Lower incisors subequal. First lower premolar large, with prominent distal cusp; 4th lower premolar subequal to 1st; 2nd and 3rd lower premolars much smaller. First and 2nd lower molars large; 3rd smaller; all with prominent parastyles. Etymology The new species is named in reference to Hakkari Province , Turkey where the type locality is situated (adjective) . Distribution and ecology Known from a handful of geographically close localities in the mountains of Hakkari province , Turkey . Moles were collected from moist soils beside a mountain stream at Cemşililan yaylası ( Fig. 5A ), in Eastern Anatolian Montane Steppe subject to nomadic grazing, but otherwise with little human influence. Most of the precipitation in this region falls as winter snows. At Bilmizit yaylası, specimen was collected in a similar habitat, but in this case without surface water. Comment This new mole probably corresponds to the specimens from Hakkari referred to T. davidiana by Kryštufek et al . (2001), T. streetorum by Doğramacı (1989) , and T . d. streetorum by Kryštufek and Vohralik (2005) . Our genetic, morphological, and morphometric data indicate that Hakkari populations represent a distinct species of broad-rostrum mole, rather than a subspecies of T. davidiana , and our morphometric analysis indicates that it is also distinct from T. streetorum (see above). At the time of writing, we have been unable to sequence the type material of T. streetorum or to obtain new material from Iran , but consider the morphometric results from our comparison of skulls conclusive. Kryštufek and Vohralik (2005) noted the similarities between the moles from Hakkari and a fossil mole from the Late Pleistocene of Israel , T. chthonia Bate, 1937 , which they speculated could be conspecific. Because we consider T. hakkariensis sp. nov. and T. streetorum to be distinct, it is not clear without further analysis which, if either, would be associated with this fossil but, if conspecific with either, the earlier name T. chthonia would have priority.