Glis glis ( Linnaeus 1766 )

Kryštufek, Boris, Naderi, Morteza, Janžekovič, Franc, Hutterer, Rainer, Bombek, Dominik & Mahmoudi, Ahmad, 2021, A taxonomic revision of (Rodentia) fat dormice, genus Glis, Mammalia (Warsaw, Poland) 85 (4), pp. 362-378 : 370-373

3.4 Glis glis ( Linnaeus 1766) – Europaean fat dormouse

Sciurus glis Linnaeus 1766: 87 . Type locality ‘‘Habitat in Europa australi [lives in southern Europe]’’; type locality was erroneously restricted to “ Germany ” ( Miller 1912: 577); emended by Violani and Zava (1995: 111) to “Southern Carniola in Slovenia ”. Type locality is further restricted by the neotype (see below) to “above Preserje, Mt. Krim, Slovenia; coordinates 45.924620N 14.439479E ”.

Glis esculentus Blumenbach 1779: 79 . Type locality is ‘‘im südlichen Europa [in southern Europe]’’. Miller (1912: 577) erroneously reported the type locality as “Central Europe”. Blumenbach refers to “Valvassor” (i.e. Valvasor 1689: 437) who quoted the fat dormouse, as “Billich” [German] or “Pouh” [Slovenian] for “Krain [Carniola]”. Therefore, esculentus has the same type locality as S. glis Linnaeus.

Glis vulgaris Oken 1816: 868 . Oken intentionally renamed many species already named; besides, his work has been rejected for nomenclatural purposes ( International Commission on Zoological Nomenclature 1956). Nomen nudum.

Myoxus Giglis Cuvier 1832: 444 . Nomen nudum ( Miller 1912: 577).

Myoxus avellanus Owen 1840: 25 + plate 105. No locality.

Glis italicus Barrett-Hamilton 1898: 424 . Type locality is “Siena”, Italy.

Glis insularis Barrett-Hamilton 1899: 228 . Type locality is “Monte Aspro, near Palermo”, Sicily, Italy.

Myoxus glis orientalis Nehring 1903: 533 . Type locality is “Gebirge Alem-Dagh, nordöstlich von Scutari, in Kleinasien [ Üskudar , Alem Dağı Mts., İstanbul, Turkey in Asia].

Glis glis spoliatus Thomas 1906: 220 . Type locality is “Khotz [Çosandere; KryŠtufek 2010: 196], near Trabizond [Trabzon]. Alt. 100 m ”, Turkey in Asia.

Glis Melonii Thomas 1907: 445 . Type locality is “ Marcurighè , Urzulei, Ogliastra, Sardinia ”, Italy.

Glis glis pyrenaicus Cabrera 1908: 193 . Type locality is “ Navarre Pyrenees, North Spain ” “ All , province of Navarre.”

Glis italicus intermedius Altobello 1920: 22 . Type locality is not specified, the name however was proposed for dormice from the Abruzzi and Molise Province (now two different provinces, the Abruzzo and the Molise), central Italy.

Glis glis subalpinus Burg 1920: 419 . Type locality is “ Münstertal [Val Müstair]”, Canton of Graubünden, Switzerland.

Glis glis tshetshenicus Satunin 1920: 150 . Type locality is “р. Шара- Аргунь [River Shara-Argun’]”, a tributary of the Terek ( Ognev 1947: 465), Chechen Republic, Russian Federation.

Glis glis postus Montagu 1923: 866 . Type locality is “Veliki Dergonel [Veliki Drgomalj; KryŠtufek 2010: 196], the Gorski Kotar, Croatia.”

Glis glis abrutti Altobello 1924: 35 . Type locality (“ Abruzzi e [and] Molise ”) is identical to Glis italicus intermedius Altobello and the two are synonymous.

Glis glis minutus Martino 1930: 60 . Type locality is “Predejane. 30 klm. S. from Leskovac, Serbia ”.

Glis glis vagneri V. E. Martino and E. V. Martino 1941: 9 . Type locality is “ Vrhpolje , Kamnik, KamniŠke Alpe, Slovenia.”

Glis glis intermedius V. E. Martino and E. V. Martino 1941: 9 . Type locality is “Presaća [PesaČa], Donji Milanovac, N. E. Serbia.” Preoccupied by Glis italicus intermedius Altobello.

Glis glis argenteus Zimmermann 1953: 28 . Type locality is “Wälder der Weissen Berge, Kreta [White Mts., Samaria, 1000 m elevation, Island of Crete; Ondrias 1966: 28].”

Glis esculantus : Vietinghhoff-Riesch 1960: 16. Incorrect subsequent spelling of esculentus Blumenbach.

Glis glis abruttii : Vietinghhoff-Riesch 1960: 19. Incorrect subsequent spelling of Glis glis abrutti Altobello.

Glis glis martinoi Mirić 1960: 36 . New name for Glis glis intermedius V. E. Martino and E. V. Martino.

G.[lis] g.[lis] wagneri: Dulić and Tortić 1960: 7. Incorrect subsequent spelling of Glis glis vagneri V. E. Martino and E. V. Martino.

Glis glis pindicus Ondrias 1966: 25 . Type locality is: “ Moni Stomiou , near Konitsa, Epirus, Greece, at an altitude of 1600 m ”.

M. glis martinoi Grekova 1969: 66 . Not Greova ( Gromov and Erbajeva 1995: 174). Type locality is “Limanchik, Abrau- Dyurso, Krasnodarskiy kray”, Russia. Preoccupied by Glis glis martinoi Mirić 1960 .

M. glis germanicus Violani and Zava, 1995: 112 . Type locality is “ Marxheim , Bavaria, Germany ”.

M. glis grekovae Baranova 2003 (in Baranova and Gromov 2003: 27). New name for M. glis martinoi Grekova 1969 .

Designation of a neotype: G. glis , as defined here, is wide-ranging, extending from the Pyrenees as far east as the Caucasus area and the Volga River. Morphological variation among geographic samples is extensive in this species and the list of synonyms involves over 20 names which were in the past tentatively classified into nine subspecies ( Corbet 1978; Ellerman and Morrison-Scott 1951). Nucleotide diversity ( Ahmadi et al. 2018; Hürner et al. 2010; Koren et al. 2015) and extensive morphological variation among populations (this paper) indicate a taxonomic complexity which is not yet understood and requires a comprehensive systematic revision on its own (see also Gippoliti 2013; Gippoliti and Groves 2018). An objective definition of taxa is essential for a stable taxonomy and nomenclature, which induced us to designate a neotype for G. glis as a firm standard for further taxonomic work in the group.

We propose voucher PMS 27369 as the neotype for G. glis . This specimen was collected by Marjan Zavodnik on 10 October 2020 above Preserje , Mt. Krim , Slovenia; coordinates 45.924620 N 14.439479 E. Preserved in ethanol with skull extracted; visceral organs fixed in 10% solution of formaldehyde and subsequently transferred to 75% ethanol; baculum kept in glycerol in a separate vial; tissue sample preserved in non-denaturated 96% ethanol and refrigerated; photographs of glans penis in dorsal, lateral and ventral view deposited in the PMS database. The tissue is also deposited in ZFMK (ZFMK-TIS-54202) .

Dimensions of the neotype: BWt– 282 g; HBL– 195 mm; TL– 152 mm; HfL– 31.4 mm; EL– 16.7 mm; CbL– 40.6 mm; MxT– 7.4 mm; DiL– 10.1 mm; ZgW– 25.4 mm; IoC– 5.2 mm; BcB– 19.8 mm; BcH– 11.5 mm; GpL– 10.42 mm; GpW– 4.67 mm; GpH– 3.58 mm; BaL– 8.44 mm; BaW– 2.75 mm.

An illustration of the neotype skull is to be found in this paper in Figure 9 .

The International Code for Zoological Nomenclature ( International Commission on Zoological Nomenclature 1999; subsequently referred to as the Code) stipulates conditions under which the designation of a neotype is justified. Specifically, “a neotype is not to be designated as an end in itself” (Article 75.2 of the Code) but only when “a name-bearing type is necessary to define the nominal taxon objectively” (Article 75.1). We believe that the neotype of G. glis will remove doubts regarding the taxonomic scope of this species against Glis persicus on the one hand and will facilitate a taxonomic revision within G. glis as it is defined here. As stated earlier on, Gippoliti (2013) and Gippoliti and

Groves (2018) already extracted italicus (with insularis ) from the scope of G. glis , but a thorough taxonomic revision still remains to be done. Furthermore, the taxonomic status of a highly divergent phylogeographic lineage from the Balkans

(Macedonian lineage) was not addressed yet. Therefore, a comprehensive taxonomic revision of G. glis may retrieve a higher species diversity that is still not appreciated.

In addition to the above justification for a valid designation of the neotype we met other qualifying conditions specified by the Code. In detail:

(1) We provide characters that differentiate G. glis from G. persicus (stipulated by Article 75.3.2. of the Code) and demonstrate that the neotype matches the traits which are characteristic for G. glis . Hürner et al. (2010) sequenced five dormice from Mt. Krim which all retrieved a single cyt b haplotype (Hap02) characteristic for the European sublineage of G. glis . Furthermore, the locality of the neotype is well inside the range of the North-Western Balkan microsatellite group of G. glis which occupies Slovenia, North-Eastern Italy and Croatia ( Michaux et al. 2019).

(2) We describe in words and in figures the neotype specimen, hence meeting the provision of Article 75.3.3.

(3) As shown by Violani and Zava (1995), the type of S. glis had not been designated and Linnaeus himself never saw the animal. The description in the 12th edition of Systema Naturae ( Linnaeus 1766) is almost verbatim a summary from the letter sent to Linnaeus on 7th April 1763 by Joannes A. Scopoli, his correspondent from the Austrian province of Carniola (now Slovenia). “Habitat in Europa australi”, which is the Linnean type locality for S. glis , is based on “Carniola, in primis inferiore” in Scopoli’ s letter. Miller (1912: 577) erroneously restricted the type locality to “ Germany ”, possibly a reminiscence of the fact that Carniola was, at the time of correspondence between Linnaeus and Scopoli, part of the Holy Roman Empire which occasionally had an unofficial extension “of the German Nation.” Miller’ s restriction was emended to “Southern Carniola in Slovenia ” ( Violani and Zava 1995: 111).

(4) The neotype comes from Mt. Krim which is located in the southern part of the former Carniola Province and is therefore inside the type locality as validly restricted by Violani and Zava (1995) (Article 75.3.6.). institution”, which “maintains a research collection, with proper facilities for preserving name-bearing types, and … makes them accessible for study” (Article 75.3.7.). In provision with Article 76.3.of the Code (“Type locality determined by the neotype ”), “above Preserje, Mt. Krim, Slovenia ” is the type locality for G. glis .

Diagnosis: Identical to a cluster of five sub-lineages (European, Italian, Sicilian, Macedonian and Greek; Figure 1 ) as retrieved in the phylogenetic analysis of the mitochondrial cyt b gene ( Ahmadi et al. 2018; Hürner et al. 2010). In our dataset, G. glis has unique mutations in comparison with sequences of G. persicus at 43 positions of the cyt b alignment ( Table 1). G. glis has a shorter glans penis (GpL <10.5 mm; GpL> 12.5 mm in persicus ), a shorter and narrower baculum (BaL <10 mm and BaW <3.0 mm vs BaL> 13.5 mm and BaW> 3.5 mm in persicus ), and a narrower posterior extension of the premaxilla ( Figure 6A–C ).

Description: The morphology of G. glis is thoroughly documented in Miller (1912), Ognev (1947), Storch (1978), Rossolimo et al. (2001), and KryŠtufek (2010). Subsequently we list traits which, despite some interspecific overlap, signalize a divergence between G. glis and G. persicus .

(1) The tail is usually only slightly darker than the back in G. glis while it is normally blackish in G. persicus ; dormice from the Italian sub-lineage resemble in the tail colouration persicus rather than glis .

(2) On the hind foot, digit IV is of the same length than digit III in G. persicus , but it is the longest digit in G. glis . (3) The number of abdominal nipples is 1–2 in G. glis and 2–3 in G. persicus .

(4) The maxillary tooth-row is longer in G. persicus (Mxt> 7.0 mm) than in G. glis (MxT <8.1 mm in majority of populations); dormice from the Italian sub-lineage are intermediate in this trait (MxT = 7.3–8.7 mm).

Distribution: G. glis occupies the majority of the genus’ range in Europe, northern Anatolia, and the Caucasus area. The European range was mapped by Storch (1978) and KryŠtufek (1999), the range in Russia, Belarus, Ukraine and Moldova by Likhachev (1972), in Anatolia by KryŠtufek and Vohralík (2005), and in the Caucasus by Shidlovsky (1962). Local updates are summarized in Holden (2005). Along the south-western Caspian coast, the ranges of G. glis and G. persicus are presumably delimited by rivers Kura and Aras (cf. Map 2 in Shidlovsky 1962).

Miscellaneous: Earlier authors noted that italicus ( Ellerman and Morrison-Scott 1951: 547) and also melonii ( Ognev 1947: 470) resemble persicus (or caspius) closer than their counterparts from the rest of Europe. As shown here, the differences are obvious in the tail colouration and in the length of the maxillary tooth-row. Contrary to Gippoliti and

Groves (2018) we are hesitant to propose the elevation of italicus to a species in its own right for the following reasons:

(1) If italicus would be defined to include the Italian and Sicilian sub-lineages, then it would be paraphyletic in the cyt b tree.

(2) Dormice from the Italian region are in two highly divergent cyt b sub-lineages which are sympatric in Sicily ( Hürner et al. 2010). Relationships between these sub-lineages on the island are not known.

(3) Different markers (cyt b and COI) retrieved noncongruent phylogeographic patterning in Italy (cf. Hürner et al. 2010 vs Lo Brutto et al. 2011).

(4) The contact zone between the Italian and the European sub-lineages is not known. Based on morphological variation in a subspecies italicus, Miller (1912: 579) concluded that “northern specimens [from the region of Turin and at Porlezza] are probably best treated as intermediates between glis and italicus ”.

Classification of G. glis as a monotypic species ( Holden-Musser et al. 2016) contradicts the phylogeographic structuring of the species ( Figure 1 ) and the morphological variation ( Ognev 1947; Storch 1978). The pattern of variation is complex and the size classes, upon which the traditional subspecies mainly rely on, do not match the phylogenetic lineages. In this study, the populations from Germany and Macedonia overlapped perfectly in cranial dimensions ( Figure 5 ), although they belong to distant phylogenetic groups. Contrary to this, populations from Germany and Slovenia (both from the European sub-lineage) hardly overlapped in morphospace ( Figure 5 ). An infraspecific revision of G. glis is therefore left unresolved.

Not considered in this review is a single mt haplotype from the Aegean Island of Alonissos which clusters with other G. glis sequences ( Castiglia et al. 2012). Only a single individual is known from the island and we did not see the voucher.

Figures

Figure 1: Geographic range of fat dormice (Glis) with sampling points of museum vouchers used in this study. Circles–Glis glis; triangles– G. persicus; stars–type localities with names of species group taxa. Top insets show distribution of phylogeographic lineages (left) and simplified phylogenetic tree of the genus (right). Acronyms for sublineages: A – Italian; E – European; M – Macedonian; G – Greek; S – Sicilian; 1 – Western Iranian; and 2 – Eastern Iranian (modified from Ahmadi et al. 2018; Hürner et al. 2010;Koren et al. 2015;Lo Brutto et al. 2011; Naderi et al. 2014a, 2016). The range follows KryŠtufek (2010).

Figure 5: Bivariate plot of five Glis populations onto the first two principal components (PC) derived from ordination of seven cranial measurements (transformed to log10). The percentage of variance explained by individual PC is in parentheses. The character vector diagram (left-upper inset) illustrates the relative contribution of the original variables (see text for acronyms) to the principal components. Population identifiers: 1 – Slovenia; 2 – Germany; 3 – North Macedonia; 4 – Italy (including Sardinia); 5 – Iran. Polygon for the Iranian sample is shaded grey and polygon for the Italian sample is shown by an interrupted line.

Figure 6: Dorsal rostrum in dormice showing the difference in relative width of the posterior extension of the nasal (1a) against the posterior extension of the premaxilla (2a; insets C and F). Top (glis lineage): A – Slovenia, Mt. Snežnik (PMS 27319); B – Turkey, Rize, Çat, Çeymakçur Yayla (ZSM 67/29 HMS); C – Italy, Monte Gargano (ZFMK 66.415). Bottom (persicus lineage): D – Iran, Golestan, Kalaleh (PMS 26297);E – Iran, Gilan, Asalem (ZSM 68.443 HMS); F – Azerbaijan, Lerik region (ZIN 71740). Bones: 1 – nasal; 2 – premaxilla; 3 – maxilla; 4 – lachrymal; 5 – frontal. Not to scale; only the right half of the rostrum is shown in insets B, C, E, and F.

Figure 9: Dorsal, ventral and lateral views of the skull and labial side of the mandible in Glis glis (top) and G. persicus (bottom). Based on vouchers PMS 27369 (G. glis; Slovenia, Mt. Krim) and ZSM 68.443 HS (G. persicus; Iran, Gilan, Asalem).