Laeocathaica tropidorhaphe Moellendorff , 1899

Wu, Min, Shen, Wang & Chen, Zhong-Guang, 2023, Land snail diversity in central China: revision of Laeocathaica Moellendorff, 1899 (Gastropoda, Camaenidae), with descriptions of seven new species, ZooKeys 1154, pp. 49-147 : 49

Laeocathaica tropidorhaphe Moellendorff, 1899

Figs 2A, D , 33 , 34 , 44A, B , 46I, J , 50A , 51

Laeocathaica tropidorhaphe Möllendorff, 1899: 94, pl. 5, fig. 7; - Gude 1902: 6; - Yen 1939: 149, pl. 15, fig. 33; - Yen 1942: 284; - Chen and Zhang 2004: 319, fig. 307.

Laeocathaica (Laeocathaica) tropidorhaphe - Zilch 1968: 175; - Richardson 1983: 79.

Laeocathaica dangchangensis Chen & Zhang, 2004: 339, 443, fig. 332.

Laeocathaica amdoana - Páll-Gergely et al. 2022: 38, fig. 7A, B, D.

Museum material.

SMF 9074, lectotype; Zw. Li-dshia-pu u. Hsi-gu-tsheng, S. O. Gansu; ex Potanin 923, Slg. O. v. Möllendorff. SMF 9075, paratypes, four fms; Tan-tshang (= Dangchang), SO-Gansu; ex Potanin 545, 623, 808b. SMF 9076, paratype, one fms; Dshie-dshou; ex Potanin 119, Slg. O. v. Möllendorff. SMF 9077, paratypes, two fms; the same data as lectotype. SMF 95126 , paratype, one fms; SO-Gansu, NW-China; Slg. C. R. Böttger 1904 (ex Möllendorff!). SMF 24269 , one fms labeled with " L. stenochone "; Tan-tschan; Slg. O. v. Möllendorff .

New material.

HBUMM00486, 3 fma; Daigusicun , Diebuxian, Gansu Province, 2000 m a.s.l., near 34.004644°N, 103.939651°E; 1998-V-10, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00519, 2 fma, 1 fms; Jinpingxiang [锦屏乡], Wudu, Gansu Province, 33.411281°N, 104.793036°E; 1998-V-11, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00526, 2 fma; Lijiexiang [立节乡], Zhouquxian, Gansu Province, 33.901549°N, 104.076052°E; 1998-V-10, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00527; 2 fma; Shanggou , Shawanxiang , Dangchangxian, Gansu Province, near 33.625712°N, 104.570162°E; 1998-V-6, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00693, 12 fma, 1 fma dissected: Wangguanxiang [望关乡], Kangxian [康县], Gansu Province, near point (33.461266°N, 105.378423°E); 1998-V-5, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM05664, many fma, 1 fma dissected; east bank of Bailongjiang , Lianghekou , Dangchang, Gansu Province, near 33.695553°N, 104.496612°E, limestone, 2006-X-02, coll. Liu, J.-M. and Zheng, W. HBUMM05619, many fma, 1 fma dissected; Foyazhen [佛崖镇], Wudu, Gansu Province, 33.44°N, 105.27°E, limestone hill with thick shrubs and broad-leaved woods, 2006-X-02, coll. Liu, J.-M. and Zheng, W. ; DNA voucher HBUMM05617. HBUMM06621, 2 fma; near Zhangzhazhen , Jiuzhaigouxian, Sichuan Province, 2100 m a.s.l., 33.299167°N, 103.859444°E; 2011-VIII-11, coll. Wu, M., Xu, Q. and Buhda, P. ; DNA voucher HBUMM06620. HBUMM08425, 1 fma dissected; near Yangshanwan No. 2 Bridge , along the national road 212, Gansu Province, on slope, 2019-X-11, coll. Li, Q.-M. ; DNA voucher HBUMM08425a. HBUMM05621, numerous fma, 1 fma dissected; Foyazhen , Wudu, Gansu Province, 33.44°N, 105.27°E, limestone hill with thick shrubs and broad-leaved woods, 2006-X-02, coll. Liu, J.-M. and Zheng, W. HBUMM05719, numerous fma, 1 fma dissected; Dangchang , Gansu Province, 2006-X-04, coll. Liu, J.-M. and Zheng, W. ; DNA voucher HBUMM05716. HBUMM00450 (3 fma), HBUMM05664 (many fma, 1 fma dissected), Wangguanxiang , Kangxian, Gansu Province; 1998-V-1, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00508, 4 fma, 1 dissected; Xinglongcun , Zhongzhaixiang , Wenxian, Gansu Province; 1998-V-19, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00625, 1 fms and 2 shells of subadult, Erlangshan [二郎山], Zhouquxian, Gansu Province; 1998-V-9, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM05166, 2 fma and 2 juvs; Zhongzhaixiang , Wenxian, Gansu Province; 1998-V-19, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM05600, many fma and juvs; southern slope of Beishan , Wudu, Gansu Province, limestone hill with sparse shrubs; 2006-X-02, coll. Liu, J.-M. and Zheng, W. HBUMM05608, numerous fma; northern slope of Beishan , Wudu, Gansu Province, limestone hill with sparse shrubs; 2006-IX-IX-30, coll. Liu, J.-M. and Zheng, W. HBUMM05664, HBUMM05669, HBUMM05684 (many fma, 8 dissected): eastern bank of Bailongjiang , Lianghekou , Dangchangxian, Gansu Province, near point (33.697332°N, 104.493015°E); limestone; 2006-X-02, coll. Zheng, W. and Liu, J. HBUMM05691, HBUMM05692, HBUMM05693b, HBUMM05694: Guantingzhen , Dangchangxian, Gansu Province, near point (33.82428°N, 104.538282°E); limestone hills, along 212 Guodao; 2006-X-3, coll. Zheng, W. and Liu, J.-M. ; DNA voucher HBUMM05688. HBUMM08369, 4 subadults; Gansu Province , 2019, coll. Li, Q.-S. CZG202008-w2, 4 subadults Shangdezhen , Wenxian, Gansu Province, near point (32.907414°N, 104.76994°E), 2020-VIII, coll. Chen, Z.-G. CZG202107-w1, 12 fms and 8 subadults, east of the town of Jiuzhaigouxian , Sichuan Province, near point (33.259523°N, 104.245884°E); 2021-VII, coll. Chen, Z.-G.

Distribution.

Gansu: Dangchangxian (type locality), Diebuxian, Kangxian, Wenxian, Wuduxian, Zhouquxian (type locality). Sichuan: Jiuzhaigouxian.

Additional information of shell.

Protoconch has regularly arranged fine granules (each ~ 30 µm long) and fine radial threads, both of which are usually obscured by erosion or weathering. Apical and umbilical sides of body whorl have densely and regularly arranged spiral grooves.

General anatomy.

Eversible head wart present. Jaw arcuate, with five projecting ribs.

Anatomy of genital organs.

Penial sheath covering ~ 1/6-1/5 of penis. Penis tubular, slightly thick distally. Inside penis, two adjacent pairs of pilasters forming two Y-shaped forks at proximal 1/4. Fine pilasters on distal end of penis merging into ~ 9 more or less thick short folds. Vas deferens narrow throughout. Vagina between atrium and dart sac elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2-1 length of dart sac. Dart sac ~ 1/3 length of penis. Love dart ~ 3 mm long, apically rhombic in cross-section. Accessory sac small, internally empty, inserting into dart sac proximally, opening to dart chamber. Mucous glands 7 (HBUMM05664) - 10 (HBUMM05619), each simply or complicatedly branched. Proximal accessory sacs absent. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.

Remarks.

In terms of shell shape, this species has an obviously depressed spire and more or less sharp carina or angulation above periphery compared to the sympatric (Fig. 2D ) Laeocathaica amdoana and L. distinguenda . Their shell differences are also indicated by the average shapes showed in Fig. 50A . The aperture of Laeocathaica tropidorhaphe , more or less stretched outwards, is narrower than that of L. distinguenda (Fig. 50A ). Laeocathaica dangchangensis Chen & Zhang, 2004 shows a typical shell of L. tropidorhaphe (Fig. 50A ).

The systematic affinity between Laeocathaica amdoana , L. distinguenda and L. tropidorhaphe is supported by the phylogeny resulting from the present analyses (Fig. 51 ), which also indicates that the character state of the presence of proximal accessory sacs may have repeated on different branches of Clade M ( Laeocathaica ) (Fig. 51 ).

For more comments, see Laeocathaica amdoana .

Figures

Figure 2. A distribution map of all known species of Laeocathaica Moellendorff, 1899 except L. anceyi (Moellendorff in Ancey, 1889), L. hisanoi Pall-Gergely, 2022, and L. leucorhaphe Moellendorff, 1899 whose precise localities are not known; white dots: the localities where no Laeocathaica species was ever found B distribution of the new species described in this paper, yellow dots: L. qingchuanensis Wu, sp. nov., green dot: L. zhengpingliui Wu, sp. nov., brown dots: L. parapolytyla Wu, sp. nov., white dot: L. qiminglii Wu, sp. nov., pink dot: L. cheni Wu, sp. nov., blue dots: L. qishilii Wu, sp. nov., orange dots: L. nordsiecki Wu, sp. nov. C distribution of L. carinifera (H. Adams, 1870) (green dots), L. qingchuanensis Wu, sp. nov. (yellow dots), and L. stenochone Moellendorff, 1899 (blue dots) D distribution of L. amdoana Moellendorff, 1899 (green dots), L. distinguenda Moellendorff, 1899 (pink dots), and L. tropidorhaphe Moellendorff, 1899 (blue dots) C equivalent to the larger box in (A), B, D equivalent to the smaller box in (A).

Figure 33. Laeocathaica tropidorhaphe Moellendorff, 1899 A SMF 9074, lectotype B SMF 9075, paratype C SMF 9077, paratype D HBUMM 05664.

Figure 34. Genital anatomy of Laeocathaica tropidorhaphe Moellendorff, 1899 A general view B right view of dart sac apparatus, HBUMM 05664 - spec. 1 C ventral view of dart apparatus A, C HBUMM 05619 - spec. 1. Abbreviations: AS - accessory sac; At - atrium; BC - bursa copulatrix; BCD - bursa copulatrix duct; DS - dart sac; DtC - a chamber containing love dart; Ep - epiphallus; MG - mucous glands; P - penis; PR - penial retractor muscle; PS - penial sheath; Va - vagina; VD - vas deferens.

Figure 44. Internal view of penis A, B Laeocathaica tropidorhaphe Moellendorff, 1899 A general view, HBUMM 05619 - spec. 1 B proximal penis, HBUMM 05664 - spec. 1 C L. qishilii Wu, sp. nov., general view, HBUMM 08298 - spec. 1, holotype D L. zhengpingliui Wu, sp. nov., general view, HBUMM 05553 - spec. 1, holotype E L. qingchuanensis Wu, sp. nov., general view, HBUMM 01179 - spec. 1, holotype F-H L. cheni Wu, sp. nov. F, G HBUMM 05553 b-spec. 1, holotype F general view G proximal penis H HBUMM 08428 - spec. 1, paratype, partial internal view of distal penis. Arrows indicate the position where two penial internal pilasters fuse into one distal pilaster.

Figure 46. SEM micrographs A, B Laeocathaica distinguenda Moellendorff, 1899, HBUMM 05479 A protoconch B teleoconch C, D L. stenochone Moellendorff, 1899, HBUMM 05495 C protoconch D teleoconch E, F L. prionotropis Moellendorff, 1899, HBUMM 05549 E protoconch F teleoconch G, H L. amdoana Moellendorff, 1899, HBUMM 05640 G protoconch H teleoconch I, J L. tropidorhaphe Moellendorff, 1899, HBUMM 05664 I protoconch J teleoconch K, L L. cheni Wu, sp. nov., HBUMM 08368, paratype, juvenile K protoconch L teleoconch.

Figure 50. Scatter plots of canonical variate 1 against canonical variate 2 (Canonical Variate Analysis), showing the shell morphological relationships among A Laeocathaica distinguenda Moellendorff, 1899 (green dots; a, average shape), L. amdoana Moellendorff, 1899 (blue dots; b, average shape) and L. tropidorhaphe Moellendorff, 1899 (orange dots; c, average shape) B L. qingchuanensis Wu, sp. nov. (green dots; a, average shape), L. stenochone Moellendorff, 1899 (orange dots; b, average shape) and L. carinifera (H. Adams, 1870) (blue dots; c, average shape). Encircled dots: 1, SMF 8959, lectotype; 2, HBUMM 05436 - spec. 1; 3, SMF 8952, lectotype; 4, SMF 9074, lectotype; 5, paratype of L. dangchangensis Chen & Zhang, 2004; 6, SMF 9071, lectotype; 7, syntype, NHMUK 1870.7.16.7. Data for 5 from fig. 7 D, 6 and 7 from fig. 12 in Pall-Gergely et al. 2022.

Figure 51. Bayesian phylogram of camaenid species (for detail see in Table 1) based on the concatenated partial mitochondrial 16 S and partial ITS 2 sequences. The tree is rooted with Helix pomatia. Numbers near nodes indicating Bayesian posterior probabilities and Maximum-likelihood bootstrap values are given as BPP ⁄ BP. Black or orange part shows the topology where the result yielded by using Bayesian-Inference method agrees with that by using Maximum-likelihood method or not, respectively. An asterisk indicates the branch is exactly one third shortened in length. Scale bar is for substitutions per site.