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Figure 18 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 18 Theodoxus macri (G.B. Sowerby II, 1849). A–D. Specimen collected close to Harbiye Falls, Harbiye, Antakya, Turkey (UGSB 24179); E–H. Specimen from Lake Yenişehir, Reyhanlı, Turkey (UGSB 24180); I, J. Specimen from Lake Balık, Adalar, Turkey (UGSB 24181). All specimens (A–J) were used in the phylogeny (Fig. 2). Scale bars: 1 mm.

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Figure 22 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 22 Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A–D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E–H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K–M. Specimen from Lake Sapanca, Yeni Eşme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N–P; Q–S; T–U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099/24). Scale bars: 1 mm.

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Figure 19 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 19 Theodoxus major Issel, 1865. A, B. Specimen (T. pallasi-morphotype) collected in the Caspian Sea, offshore from Aktau, Kazakhstan (UGSB 20712); C, D. Specimen (T. pallasi-morphotype) collected in the Aral Sea, Kazakhstan in 1979 (UGSB 19129); E–H. Specimen (T. pallasi-morphotype) from the Shahpol River, Aliabad Askar Khan, Iran (UGSB 18091); I–K. Specimen of T. major collected at the outflow of Lake Akna, Aknalich, Armenia (UGSB 20482); L–N. Specimen (T. astrachanicus-morphotype) collected in Utlyukskij Liman, Ukraine (UGSB 18130); O, P. Specimen of T. major collected in the Hrazdan River at the inflow of the Yerevan Reservoir, Armenia (UGSB 20496); Q–T. Specimen of T. major from Zoeram, North Khorasan, Iran (UGSB 21661); U, V. Specimen of T. schultzii collected in the Caspian Sea, offshore from Aktau, Kazakhstan (UGSB 20714); W–Y. Lectotype of T. pallasi from Caspian Sea, Dagestan, Russia (ZIN 54547/63); Za, Zb. Syntype of N. schultzii from the Caspian Sea (ZIN, no. 5 in systematic catalogue). Specimens E–K and O–T used in the phylogeny (Fig. 2). Scale bars: 1 mm.

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Figure 15 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 15 Theodoxus jordani (G.B. Sowerby I, 1836). A–D. Specimen from the Avakas Gorge, Peyia, Cyprus (UGSB 18321) incorporated into the phylogeny (Fig. 2); E–H. Specimen collected in the Ceyhan River, Eski Misis, Adana, Turkey (UGSB 20777); I–L. Specimen from the Orontes River, Antakya, Hatay, Turkey (UGSB 24175) used in the phylogeny (Fig. 2); M–O. Specimen (T. euphraticus-morphotype) from the Seyed Hosein Park spring, near the Bishapur, Fars, Iran (UGSB 22233); P–R. Topotype of N. jordani from the River Jordan, Israel (UGSB 24177) incorporated into the phylogeny (Fig. 2); S, T. Specimen (T. euphraticus-morphotype) from the Karun River, Ahvaz, Iran (UGSB 21681); U. Specimen (T. niloticus-morphotype) collected in the Nile River, Egypt (UGSB 24178) used in the phylogeny (Fig. 2); V. Specimen (T. euphraticus-morphotype) from the Karun River, Ahvaz, Iran (UGSB 21682). Scale bars: 1 mm.

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Figure 4 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

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

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Figure 11 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 11 Theodoxus gloeri Odabaşı & Arslan, 2015. A–C. Paratype from Balıkdamı Wetland-Sakarya River, Eskişehir, Turkey (coll. Glöer). Scale bar: 1 mm.

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Figure 14 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 14 Radula of Theodoxus gurur Sands & Glöer sp. nov. paratypes (UGSB 20689). A. Portion of the radula showing full sets of teeth; B. Magnified view of the central and lateral teeth; C. Magnified view of the lateral and marginal teeth; D. Magnified view of the first row of marginal teeth; E. Merging point between the first and second rows of marginal teeth; F. Magnified view of the faces of inner marginal teeth belonging to the first row. Scale bars: 100 μm (A–D), 20 μm (E–F).

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Figure 21 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 21 Theodoxus syriacus (Bourguignat, 1852). A–D. Specimen collected at Sultanköyü, Mardin, Turkey (UGSB 24182); E–H. Specimen from the Yanarsu Stream, Alıçlı, Turkey (UGSB 20518). I–L. Specimen collected near Başdeğirmen, Diyarbakır, Turkey (UGSB 24183); M–O; P–R. Two specimens collected by Bourguignat from İskenderun, Turkey (MHNG-MOLL-111712). All UGSB specimens incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm.

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Figure 13 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 13 Theodoxus gurur Sands & Glöer sp. nov. A–D. Holotype from the Suyunbaşı spring complex, Ayrancılar, İzmir, Turkey stored in NMNL (RMNH.MOL.342197); E–G. Paratype from same locality and storage facility (RMNH.MOL.342199). Scale bars: 1 mm.

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Figure 17 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 17 Theodoxus jordani (G.B. Sowerby I, 1836). A–C. Syntype of N. meridionalis var. mesopotamica from near Diyarbakır, Turkey (ZMZ 528914); D, E. Specimen (T. mesopotamicus-morphotype) collected in Şanlıurfa, Turkey (UGSB 23426) and used in the phylogeny (Fig. 2); F–H; K–M. Two syntypes of N. cinctella collected at the source of River Chabur near Ras al-Ayn, Syria (ZMB/Moll 21732); I, J. Topotype of N. cinctella from the same locality (UGSB 18762). Scale bars: 1 mm.

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Figure 10 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 10 Theodoxus fluviatilis (Linnaeus, 1758). A–D. Specimen (conforming to T. sarmaticus) from the Dnieper-Bug estuary, Ukraine (UGSB 18328); E–H. Specimen (T. subthermalis-morphotype) from Novy Afon, Abkhazia, Georgia (UGSB 18402); I–K. Syntype of N. saulcyi from near the Penteli Monastery, Greece (MHNG-MOLL-111736); L. Specimen (T. saulcyi-morphotype) from Stilos, Crete, Greece (UGSB 20519) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm.

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Figure 26 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 26 Type locality of Theodoxus wilkei Sands & Glöer sp. nov. A, B. Çifteler spring, Çifteler, Eskişehir, Turkey, 39.34931°N, 31.05527°E.

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Figure 20 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 20 Theodoxus pallidus (Dunker, 1861). A–D. Specimen from Haji Abad, Fars, Iran (UGSB 22227); E–H. Syntype of N. pallida from southern Iran (ZMB 108.792); I, J; K, L. Two syntypes of N. pallida from southern Iran (ZMB 108.793); M–P. Specimen (T. doriae-morphotype) from a spring near Harat, Yazd, Iran (UGSB 21706); Q, R; S, T; U, V; W–Y. Four syntypes of N. doriae from the thermal springs of Kerman, Iran (MSNG, without coll. no.). Specimens A–D and M–P were used in the phylogeny. Scale bars: 1 mm.

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Figure 12 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 12 Type locality of Theodoxus gurur Sands & Glöer sp. nov. A, B. Suyunbaşı spring complex, Ayrancılar, İzmir, Turkey, 38.24826°N, 27.28117°E.

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Figure 16 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 16 Theodoxus jordani (G.B. Sowerby I, 1836). A, B. Syntype of N. euphratica from the Euphrates River at Samawah, Iraq (ZMZ 528916); C–E. Syntype of N. michonii from Syria (MHNG-MOLL-111706); F–H. Syntype of N. karasuna collected from the Karasu River, south-eastern Turkey (ZMZ 528937); I, J; K, L. Two syntypes of N. ponsoti from Lake Muzayrīb, Syria (MNHN-IM-2000-32754); M. Syntype (labelled as paratype) of N. ponsoti from Lake Muzayrīb (MT820, RBINS); N. Syntype of N. jordani var. turris collected in the Sea of Galilee (ZMZ 528930); O, P. Syntype of N. bellardii from the Litani River, Beqaa valley, Lebanon (ZMZ 528918). Scale bars: 1 mm.

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Figure 1 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 1 Terminology of key morphological characters for Theodoxus. A. Shell: a = apex, ap = aperture (im = inner margin, om = outer margin), cp = columellar plate, p = periostracum, s = spire, sh = shell height, sm = shell margin, sw = shell width, w = whorls; B. Operculum: ap = apophysis, ca = callus, cb = calcareous base, cl = conchioline lamella, la = left adductor, pa = pseudo-apophysis, ra = right adductor, rp = rib-pouch, rs = rib-shield; C. Radula: ac = A-central (c = cusp, r = ridge, tp = tooth plate), bc = B-central, cc = C-central, el = E-lateral (le = lower edge, ue = upper edge), mt1 = first row of marginal teeth (sf = small face of marginal teeth), mt2 = second row of marginal teeth, rc = R-central (ae = anterior edge, f = tooth face).

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Figure 2 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 2 Dated phylogeny of the genus Theodoxus constructed with BEAST, based on existing GenBank and newly incorporated data for COI, 16S and ATPα (Suppl. material 1: Table S1). 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 S3). Dashed dark blue and light blue lines represent the minimum interspecific and maximum intraspecific variation determined by Sands et al. (2019a) to aid Theodoxus spp. delimitation (see see Methods, Phylogenetics and species delimitation and that study for more information on the delimitation methods).

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Figure 28 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 28 Radula of Theodoxus wilkei Sands & Glöer sp. nov. paratypes (UGSB 20687). A. Portion of the radula showing full sets of teeth; B. Magnified view of the central and lateral teeth; C. Magnified view of the lateral and first and second rows of marginal teeth; D. Magnified view of the lateral teeth; E. The first row of marginal teeth; F. Magnified view of the faces of inner marginal teeth belonging to the first row. Scale bars: 100 μm (A–D), 20 μm (E–F).

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Figure 24 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 24 Theodoxus wesselinghi Sands & Glöer sp. nov. A–D. Holotype collected in the Sakarya River, Çayköy, Bilecik, Turkey (RMNH.MOL.342200); E–G. Paratype from the same location as the holotype (RMNH.MOL.342202); H–J. Paratype from Fele, Isparta province, Turkey (RMNH.MOL.342204); K–M. Paratype from Eflatun Pınarı, Konya province, Turkey (RMNH.MOL.342206). All photographed material (A–M) is stored in NMNL. Scale bars: 1 mm.

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Figure 27 from: Sands AF, Glöer P, Gürlek ME, Albrecht C, Neubauer TA (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): 25-66. https://doi.org/10.3897/zse.96.48312

Figure 27 Theodoxus wilkei Sands & Glöer sp. nov. A–D. Holotype (RMNH.MOL.342208); E–G. Paratype (RMNH.MOL.342210); H–J. Paratype (RMNH.MOL.342211). All photographed material is from the Çifteler Spring, Çifteler, Eskişehir, Turkey and stored in NMNL. Scale bars: 1 mm.

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