What should we call the Levant mole? Unravelling the systematics and demography of Talpa levantis Thomas, 1906 sensu lato (Mammalia: Talpidae)

Abstract

Turkey hosts 5 of the 11 species of Talpa described to date and Anatolia, in particular, appears to be an important centre of diversity for this genus. Of these taxa, the Levant mole, Talpa levantis Thomas, 1906 has been found to consist of two genetically divergent sublineages, which may represent separate species. Here, we use a combination of mitochondrial and nuclear DNA sequences from specimens of T. levantis s.lat., collected across the species’ geographical range, to explore the systematics and demographic history of Levant moles. Both mitochondrial and nuclear markers confirm the existence of distinct eastern and western sublineages, which apparently diverged from each other in the early Pleistocene. Given the degree of cytochrome-b divergence between these (7.28%), we consider them to represent independent cryptic species. By including topotypic specimens of T. levantis s. str. in our study, we are able to show that this name applies to the western sublineage, distributed across most of the Anatolian Black Sea coastal region, from the vicinity of Trabzon in the east, westwards to Marmara. The earliest name available for the eastern taxon, found in Transcaucasia and adjacent parts of north-eastern Anatolia, is T. transcaucasica Dahl, 1944. Cytochrome-b haplotype diversity in T. levantis is relatively high and demographic analyses suggest that the species may have survived in multiple, separate, refugial areas during the Pleistocene. Our work brings the total number of named mole species recognized in Turkey to six, emphasising the importance of this region as a global centre of mole diversification.

Appendix: Additional GenBank accession numbers (not listed in Table 1) for sequences of Talpa species used in phylogenetic analyses

Species	Cyt-b			BRCA2
	Locality	Accession numbers	References	Locality	Accession numbers	References
T. europaea	Germany, Lübeck	AB037601	Tsuchiya et al. (2000)	Russia, Northern Urals, Pechora-Ilych NR	KP717098	Bannikova et al. (2015)
	France, Semilly-Escamps	KU189429	Nicolas et al. (2017b)	Turkey, Kuleli Köyü, Babaeski, Kırklareli	KP717101	Bannikova et al. (2015)
	Russia, Garevka	KF801564	Feuda et al. (2015)	Ukraine, Cherkassk region, Kanevskiy Nature Reserve	KP717103	Bannikova et al. (2015)
	Russia, Moscow	KF801558	Feuda et al. (2015)	Italy, Assisi, Casa Casella, Perugia	KP717102	Bannikova et al. (2015)
	Sweden, Dalby	Y19192	Mouchaty et al. (2000)
	France, Trieves	FN640550	Colangelo et al. (2010)
	Turkey, Kuleli Köyü, Babaeski, Kırklareli	KP717322	Bannikova et al. (2015)
	Ukraine, Cherkassk region, Kanevskiy Nature Reserve	KF801572	Feuda et al. (2015)
	Ukraine, Kiev region, Brovary	KF801575	Feuda et al. (2015)
	Ukraine, Kiev	KF801573	Feuda et al. (2015)
	Ukraine, Kosovo	KF801574	Feuda et al. (2015)
	Italy, Valfabbrica, Monte Villano, Perugia	KP717320	Bannikova et al. (2015)
	Italy, Assisi, Casa Casella, Perugia	KP717321	Bannikova et al. (2015)
	Italy, Belvedere, Perugia	FN640551	Colangelo et al. (2010)
	Italy, Valtellina, Sondalo	KP717324	Bannikova et al. (2015)
	Italy, Modena	FN640549	Colangelo et al. (2010)
	Italy, Valtellina, Sondalo	KF801528	Feuda et al. (2015)
T. aquitania	Spain, Montseny	KF801509	Feuda et al. (2015)
	France, Pyrenees-Orientales	KU189596	Nicolas et al. (2017b)
	Spain, Sant Pere De Vilamajor	KF801507	Feuda et al. (2015)
	France, Nantheuil	KU189691	Nicolas et al. (2017b)
	France, Avensan	KU189615	Nicolas et al. (2017b)
	France, Gaillan en Médoc	KU189696	Nicolas et al. (2017b)
	France, Creste	KU189628	Nicolas et al. (2017b)
	France, Nieul-les-Saintes	KU189674	Nicolas et al. (2017b)
	Spain, Hijas	KU189588	Nicolas et al. (2017b)
	Spain, Aizarnazabal	KU189589	Nicolas et al. (2017b)
T. occidentalis	Spain, Oles	KU189707	Nicolas et al. (2017b)	Spain, Granada	KP717106	Bannikova et al. (2015)
	Spain, Priesca	KU189704	Nicolas et al. (2017b)
	Spain, Vegadali	KU189709	Nicolas et al. (2017b)
	Spain, La Fajera	KU189721	Nicolas et al. (2017b)
T. martinorum	Bulgaria, Strandzha Mt.	MH093591	Kryštufek et al. (2018)
T. romana	Italy, Pizzone, Isernia	FN640565	Colangelo et al. (2010)	Italy, Assisi, Croce di Mora, Perugia	KP717109	Bannikova et al. (2015)
	Italy, Acri	HQ233519	Canestrelli et al. (2010)
	Italy, Assisi, Croce di Mora, Perugia	KP717329	Bannikova et al. (2015)
	Italy, Formello, Roma	FN640564	Colangelo et al. (2010)
	Italy, Lagonegro	HQ233503	Canestrelli et al. (2010)
	Italy, Melia	HQ233603	Canestrelli et al. (2010)
	Italy, Decollatura, Catanzaro	FN640562	Colangelo et al. (2010)
	Italy, Decollatura, Catanzaro	FN640563	Colangelo et al. (2010)
	Italy, Borgia	HQ233562	Canestrelli et al. (2010)
	Italy, Arcavacata	HQ233529	Canestrelli et al. (2010)
	Italy, Gambarie	HQ233595	Canestrelli et al. (2010)
T. stankovici	Greece, western Macedonia	FN640566	Colangelo et al. (2010)	Macedonia, Bistra	KP717123	Bannikova et al. (2015)
	Greece, Axiopulus	FN640567	Colangelo et al. (2010)
	Macedonia, Pelister	KP717347	Bannikova et al. (2015)
	Macedonia, Bistra	KP717348	Bannikova et al. (2015)
T. caeca	Italy, Capracotta	KP717333	Bannikova et al. (2015)	Italy, Majelletta	KP717113	Bannikova et al. (2015)
	Italy, Majelletta	KP717331	Bannikova et al. (2015)
	Italy, Prati di Tivo	FN640558	Colangelo et al. (2010)
	Italy, Prati di Tivo, Gran Sasso	KP717332	Bannikova et al. (2015)
	Switzerland, Bellinzone	DQ630425	Dubey et al. (2007b)
	Italy, Monte Cucco, Umbria	FN640559	Colangelo et al. (2010)
T. davidiana	Turkey, Çataklı Köyü, Tatvan, Bitlis	KP717363	Bannikova et al. (2015)	Turkey, Çataklı Köyü, Tatvan, Bitlis	KP717132	Bannikova et al. (2015)
	Turkey, Çataklı Köyü, Tatvan, Bitlis	KP717364	Bannikova et al. (2015)
T. talyschensis	Azerbaijan, Astara, Vazezamin	KP717368	Bannikova et al. (2015)	Azerbaijan, Astara, Vazezamin	KP717137	Bannikova et al. (2015)
				Azerbaijan, Astara, Vazezamin	KP717138	Bannikova et al. (2015)
T. caucasica	Russia, Nalchik vicinity	KP717350	Bannikova et al. (2015)	Russia, Nalchik region, Khasanya	KP717125	Bannikova et al. (2015)
	Russia, Adygea, Kisha river	KP717349	Bannikova et al. (2015)	Russia, Adygea, Kisha river	KP717124	Bannikova et al. (2015)
	Russia, Adygea, Kisha river	KP717351	Bannikova et al. (2015)	Russia, Krasnodar Krai, Krasnaya Polyana	KP717128	Bannikova et al. (2015)
	Russia, Krasnodar Krai, Krasnaya Polyana	KP717352	Bannikova et al. (2015)
T. ognevi	Turkey, Cumhuriyet Mahallesi, Hopa, Artvin	KP717354	Bannikova et al. (2015)	Turkey, Cumhuriyet Mahallesi, Hopa, Artvin	KP717129	Bannikova et al. (2015)
	Turkey, Cumhuriyet Mahallesi, Hopa, Artvin	KP717355	Bannikova et al. (2015)	Turkey, Cumhuriyet Mahallesi, Hopa, Artvin	KP717131	Bannikova et al. (2015)
T. altaica	Russia, Krasnoyarsk region, Middle Yenisei	KP717371	Bannikova et al. (2015)	Russia, Krasnoyarsk region, Middle Yenisei	KP717141	Bannikova et al. (2015)
	Russia, Krasnoyarsk region, Middle Yenisei	KP717372	Bannikova et al. (2015)	Russia, Altai, Teletskoye Lake	KP717139	Bannikova et al. (2015)
	Russia, Krasnoyarsk region, Middle Yenisei	KP717373	Bannikova et al. (2015)	Russia, Altai, Teletskoye Lake	KP717140	Bannikova et al. (2015)
	Russia, Novosibirsk	AB037602	Tsuchiya et al. (2000)	Unknown	JN414313	Meredith et al. (2011)
	Russia, Altai, Teletskoye Lake	FN640580	Colangelo et al. (2010)
Mogera robusta	Russia, Ussuri Nature Reserve	KC481327	Zemlemerova et al. (2013)	Russia, Ussuri Nature Reserve	KP717143	Bannikova et al. (2015)
Parascaptor leucura	China, Yunnan	HG737879	He et al. (2014)	China, Yunnan	KX755213	He et al. (2017)
Euroscaptor mizura	Japan, Ashiu, Kyoto	AB037604	Tsuchiya et al. (2000)	Japan, Aomori	KX755199	He et al. (2017)
Euroscaptor longirostris	Vietnam, province Laokai, Sa Pa	KC481347	Zemlemerova et al. (2013)	Vietnam, Vinh Phuuc	KX755203	He et al. (2017)