Theodoxus altenai Schuett , 1965

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 altenai Schuett, 1965 Figure 4A-G

Theodoxus (Theodoxus) altenai Schütt 1965: 46-49, pl. 1, fig. 4; Schütt and Şeşen 1992: 66; Yıldırım 1999: 885.

Theodoxus altenai : Roth 1987: 75; Kebapçı and Yıldırım 2010: 77; Gürlek et al. 2019: 2993; Glöer 2019: 37, fig. 19.

Type locality.

Lake Kırkgöz, Kırkgöz Kaynaği spring complex, Döşemealtı, Antalya, Turkey.

Type material.

Holotype ( RNL V.56 /1) and paratypes ( RNL V.56) are stored in NMNL. Additional paratypes are stored in ZSM ( ZSM /Mol - 20013211.00) .

Remarks.

The phylogenetic results based on mtDNA and nDNA (Fig. 2 ) suggest T. altenai Schütt, 1965 shares a close sister relationship with T. anatolicus ( Récluz, 1841), where the two species likely diverged in the early Pleistocene ( Sands et al. 2019a; Fig. 2 ). The pseudo-apophysis on the operculum of T. altenai is strongly curved and depressed and lies in the plane of the right adductor, while in T. anatolicus the pseudo-apophysis is more diagonal and less depressed (Figs 4 , 5 ). Schütt (1965) wrote in his original description of T. altenai that there is no pseudo-apophysis but a short strongly curved depressed lamella instead. Furthermore, the periostracum colouration of T. altenai and T. anatolicus can be differentiated by brown and ivory checks as opposed to being uniformly black in the latter (Figs 4 , 5 ). Moreover, the columellar plate extends beyond the shell margin in T. altenai (Figs 4 , 5 ).

Distribution.

All reliable records of this species were attributed to the Kırkgöz Kaynaği spring complex around Döşemealtı, Antalya, Turkey ( Schütt 1965; Geldiay and Bilgin 1969; Bilgin 1980; Yıldırım 1999; Kebapçı and Yıldırım 2010; Gürlek et al. 2019; Fig. 3A ). However, Sands et al. (2019a) indicated the species is also present at the Düden Waterfall, Varsak Karşıyaka, Antalya, Turkey (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.