Theodoxus anatolicus ( Recluz , 1841)

Sands, Arthur F, Gloeer, Peter, Guerlek, Mustafa E, Albrecht, Christian & Neubauer, Thomas A, 2020, A revision of the extant species of Theodoxus (Gastropoda, Neritidae) in Asia, with the description of three new species, Zoosystematics and Evolution 96 (1), pp. 25-66 : 25

Theodoxus anatolicus ( Recluz, 1841) Figure 5A-G

Nerita Anatolica Récluz 1841: 342-343, pl. 1, fig. 3 (partim, only regarding material from İzmir).

Neritina belladonna Mousson 1874: 16.

Neritina anatolica : Martens 1879: 86-88, pl. 3, figs 4, 5, pl. 13, figs 17-19, 25-29; Kobelt 1899: 3-4, pl. 211, figs 1321-1324.

Theodoxus (Neritaea) anatolicus : Bilgin 1980: 37-38; Schütt and Şeşen 1992: 65-66.

Theodoxus anatolicus : Yıldırım 1999: 885; Kebapçı and Yıldırım 2010: 77; Gürlek et al. 2019: 2993; Glöer 2019: 38, fig. 20.

Type locality.

Smyrna (= İzmir), Turkey; Aleppo, Syria; Sidon, Lebanon; Scio (= Chios), Greece.

Type material.

A set of nine syntypes of N. anatolica from İzmir is stored in MHNG (coll. no. MHNG-MOLL-15028), seven syntypes from Chios are stored in MNHN (coll. no. MNHN-IM-2000-32519) (also see Kabat and Finet 1992) . The syntypes from İzmir and Chios differ considerably and might even represent different species. In order to bring stability to the taxonomy of this species, we designate one of the syntypes from İzmir (MHNG-MOLL-15028) as the lectotype (Fig. 5E-G ) . Twenty-three syntypes of N. belladonna from İskenderun ( “Alexandretta”), Turkey, are stored in ZMZ (coll. no. 528908-528909) .

Remarks.

Schütt and Şeşen (1989b) indicated a close relationship among Theodoxus that contained a pseudo-apophysis on the operculum such as T. anatolicus , T. cinctellus (Martens, 1874) [= T. jordani ], T. euphraticus [= T. jordani ], and T. jordani , where they suggested these species could still be differentiated based on structural details of the operculum. Bunje and Lindberg (2007), using mtDNA, could not find a distinct T. anatolicus clade, and their specimens either grouped with T. jordani or T. baeticus (Lamarck, 1822). It is possible that the specimens used in that study may have been misidentified, as Sands et al. (2019a), using mtDNA and nDNA of material conforming to T. anatolicus from near İzmir, found an independent monophyletic clade. Their clade shares a close sister relationship with T. altenai and is distinct from both T. jordani and T. baeticus (Sands et al. 2019; also see Fig. 2 ). According to our analyses, the divergence of T. altenai and T. anatolicus likely occurred during the early Pleistocene ( Sands et al. 2019a; Fig. 2 ). Theodoxus anatolicus and T. altenai also depict noticeable morphological differences. The periostracum patterning is more uniformly black in T. anatolicus as opposed to checkered ivory and brown in T. altenai (Figs 4 , 5 ). Moreover, the columellar plate does not extend past the shell margin in T. anatolicus as it does in T. altenai (Figs 4 , 5 ). Neritina belladonna was considered a synonym of T. anatolicus by Martens (1879), which we follow herein.

Distribution.

Martens (1874) suggested T. anatolicus to have a very wide distribution range across Anatolia, Mesopotamia, and some of the Turkish and Greek Aegean islands. Roth (1987) revised this view and indicated T. anatolicus to be restricted to southern parts of western Anatolia and referred eastern Anatolian occurrences to T. jordani . Schütt and Şeşen (1989b, 1992) followed a more intermediate approach, suggesting that T. anatolicus occurred north-west of Hatay province in southern Turkey. Bank (2006) indicated the presence of this species on the Greek Aegean Islands, once again expanding the distribution range in support of Martens (1874); however, comprehensive sampling by Sands et al. (2019a) only found genetic evidence for T. anatolicus from south-western Anatolia, corroborating Roth (1987) (Fig. 3A ).

Figures

Figure 2. Dated phylogeny of the genus Theodoxus constructed with BEAST, based on existing GenBank and newly incorporated data for COI, 16 S and ATPα (Suppl. material 1: Table S 1). Morphospecies where associations are queried are highlighted by a question mark after the species name (see the " Systematic checklist " for more information). Small green (PP ≥ 0.95) and red (PP <0.95) dots on nodes indicate the relative posterior probability support of divergence events. Node labels (1 - 32) correspond to divergence dates and HPD intervals in millions of years ago (Mya) for selected nodes (see Suppl. material 1: Table S 3). Dashed dark blue and light blue lines represent the minimum interspecific and maximum intraspecific variation determined by Sands et al. (2019 a) to aid Theodoxus spp. delimitation (see see Methods, Phylogenetics and species delimitation and that study for more information on the delimitation methods).

Figure 3. Occurrence map for Theodoxus spp. in Asia. All points conform to specimens used in the phylogenetic analyses (Suppl. material 1: Table S 1), except for T. gloeri for which no molecular data are available and only the type and paratype localities are indicated. Species are partitioned across four identical maps (A - D) to limit overlap of points as far as possible. Barring T. gloeri, closer related species are grouped together (Fig. 2).

Figure 4. Theodoxus altenai Schuett, 1965. A - C. Holotype from Lake Kirkgoez, Doesemealti, Antalya, Turkey (RNL V. 56 / 1, NMNL). D - G. Topotype (UGSB 24167) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm.

Figure 5. Theodoxus anatolicus (Recluz, 1841). A - D. Specimen collected at Isikli, Denizli, Turkey (UGSB 24168) incorporated into the phylogeny (Fig. 2); E - G. Lectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028). H, I. Paralectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028); J, K. Paralectotype of N. anatolica from Chios, Greece (MNHN-IM- 2000 - 32519); L, M. Syntype of N. belladonna from Izmir, Turkey (ZMZ 528908). Scale bars: 1 mm.