Laeocathaica prionotropis 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 prionotropis Moellendorff, 1899

Figs 2A , 3 , 29 , 30 , 43D , 44E, F , 51

Laeocathaica prionotropis Möllendorff, 1899: 94, pl. 6, fig. 1, 1a; - Wiegmann 1900: 106, pl. 3, figs 94-97; - Gude 1902: 6; - Yen 1939: 149, pl. 15, fig. 34; - Chen and Zhang 2004: 320, fig. 309; - Páll-Gergely et al. 2022: 70, figs 25, 26.

Laeocathaica prionotropis albocincta Möllendorff, 1899: 95; - Gude 1902: 6; - Yen 1939: 149, pl. 15, fig. 35.

Cathaica prionotropis albocincta - Yen 1938: 446.

Laeocathaica (Laeocathaica) prionotropis - Zilch 1968: 174; - Richardson 1983: 79.

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

Museum material.

ZIN RAS No. 7, 2 subadults with soft parts, Wen-Xian, coll. Potanin, 1885-IX-6-8, det. Möllendorff. SMF 9078, lectotype. SMF 9079, 2 paratypes. Laeocathaica prionotropis albocincta Möllendorff, 1899: SMF 9080, lectotype; Tung-ho, W-Sy-tshuan; ex Potanin 312a, Slg. O. v. Möllendorff. SMF 9081, paratype, juv; ex Potanin 312b, Slg. O. v. Möllendorff.

New material.

HBUMM00422, 3 fms; Yanwuba [演武坝], Wenxian, Gansu Province, 1200 m a.s.l., 32.936494°N, 104.557695°E; 1998-IV-23, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM00687, Shanggou [上垢], Shawanxiang [沙湾乡], Dangchangxian, Gansu Province, near 33.625712°N, 104.570162°E; 1998-V-6, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM05165, 3 fma, 2 fms; Yuxushan , Wenxian, Gansu Province, near 32.957259°N, 104.689152°E, shrubs and slate, 1998-V-17, coll. Chen, D.-N. and Zhang, G.-Q. HBUMM05496, numerous fma and juvs, 1 dissected; 5 km away from Hejiawan [何家湾] Bridge , along the road to Bikou , Wenxian, Gansu Province, near point (32.769664°N, 105.22781°E), shrubs, limestones, loess, near farmland, 2006- IX-28, coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05549, many fma and juvs; Hengdan , Wenxian , Gansu Province ; north side of Baishuijiang River, along 212 Guodao, near point (32.864025°N, 104.859517°E); hillside, bushes; 2006-IX-29; coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05559, many fma; HBUMM05777, 7 fma and 2 juvs, 1 dissected; same data as HBUMM05549. HBUMM05756, juvs; Zhouquxian [舟曲县], Gansu Province, near point (33.797295°N, 104.38151°E); limestone, broad-leaved trees and shrubs; 2006-X-5, coll. Zheng, W. and Liu, J. ; 5742. HBUMM05564, many fma and juvs; Hengdan , Wenxian , Gansu Province ; north side of Baishuijiang River, along 212 Guodao, near point (32.864025°N, 104.859517°E); top of hill, bushes; 2006-IX-29; coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05689, 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. HBUMM05765, many fma and juvs; HBUMM05764b, 1 fma and 1 fms: Bikou , along the road from Datang Hydropower Station to Hejiawan Bridge; 2006-IX-28, coll. Liu, J.-M. and Zheng, W. HBUMM05766, many fma; same data as HBUMM05765. HBUMM05774, 5 fma, 4 juvs; same data as HBUMM05765. HBUMM06533, 1 fma; town of Wenxian , Gansu Province, 1024 m a.s.l., 32.941111°N, 104.668889°E, 2011-VIII-08, coll. Wu, M., Xu, Q. and Budha, P. ; DNA voucher HBUMM06534. HBUMM08120, many juvs; Zhuantangzhen [篆塘镇], 2 juv shells, Chongqing, near point (28.898641°N, 106.675179°E); 2014-X-10, coll. Du, Li [杜莉] and Lai , Yitong [来益同]. HBUMM08150, slope near river, Wenxian, Gansu Province, 807 m, 32.847111°N, 104.88775°E; grass and a few shrubs, on branches and rock cliff; 2017-VIII-6, coll. Sheng, X.-F. , etc.; DNA voucher HBUMM08150a. HBUMM08299, 5 fma and 1 juv, 1 dissected; Bikou , Wenxian, Gansu Province; 2019-IV, coll. Li, Q.-S. HBUMM05549, many fma; 1 fma dissected; Hengdan , along national road 212, Wenxian, Gansu Province, near 32.855948°N, 104.865229°E, shrubs on hill slope, 2006-IX-29, coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05469, many fma; 1 fma dissected; bank of Baishuijiang , Wenxian, Gansu Province, 32.946383°N, 104.685343°E, limestone and loess, broad-leaved woods, 2006-IX-27, coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. ; DNA voucher HBUMM05440. HBUMM08366, 1 fma dissected; Gansu Province , 2019, coll. Li, Q.-S. ; DNA voucher HBUMM08366a. HBUMM08423, 1 fma dissected; slope near Wenzhoulu , Wenxian, Gansu Province, near 32.943151°N, 104.692505°E, on slope, 2019-X-12, coll. Li, Q.-M. ; DNA voucher HBUMM08423a. HBUMM08421, 1 fma dissected; slope near X496, Wenxian , Gansu Province, near 32.474345°N, 104.915976°E, on slope, 2019-X-13, coll. Li, Q.-M. ; DNA voucher HBUMM08421a. CZG202008-w4, 2 fms, Yuleixiang [玉垒乡], Wenxian, Gansu Province, near point (32.83335°N, 105.036382°E); 2020-VIII, coll. Chen, Z.-G. CZG202008-w7, 1 subadult, town of Jiuzhaigouxian , Sichuan Province, near point (33.260369°N, 104.248974°E); 2020-VIII, coll. Chen, Z.-G. CZG202008-w8, 1 fms, roadside, from Longnan to Wenxian, Gansu Province, 2020-VIII, coll. Chen, Z.-G.

Distribution.

Chongqing. Gansu: Dangchangxian, Wenxian (type locality), Zhouquxian. Sichuan: Jiuzhaigouxian.

Additional information of shell.

Tiny granules (~ 25 - ~ 40 µm) on the protoconch are regularly and densely arranged. Each granule is like a hump deposited in a shallow socket. On adult shell the protoconch granules are not clear, perhaps because of erosion. On teleoconch, tiny ear-shaped scales are arranged along growth lines. On umbilical side, scale scars are regularly arranged. Spiral grooves are only present on apical side of body whorl. The shell of specimens HBUMM08366 and HBUMM08421 are not typical because the periphery is angular rather than sharply carinate.

General anatomy.

Eversible head wart of normal size. Jaw arcuate, with five projecting ribs (HBUMM05469).

Anatomy of genital organs.

Penial sheath short, proximally covering ~ 1/5 penis. Penis tubular, slightly expanded distally. Inside penis, three or four low pilasters present proximally, two fusing into one Y-shaped fork at proximal 1/2, another two high pilasters fusing into one Y-shaped fork at proximal 1/3 with cross expanded to be a large lump (~ 1 × 1 × 2 mm3). Fine pilasters on distal end of penis merging into more than eight thick but short folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/3-1 length of dart sac. Dart sac ~ 1/2 (HBUMM05469) to 2/3 (HBUMM05549) length of penis. Love dart ~ 8 mm long, apically rhombic in cross-section. Accessory sac small but externally distinguishable, internally solid, inserting into dart sac at distal 1/3, opening to dart chamber. Mucous glands three (HBUMM08423), four (HBUMM05469, HBUMM08421), or eight (HBUMM05549), each simply or complicatedly branched. Proximal accessory sacs two, symmetrical, apically separated and ventrally touching, internally with a few low pilasters, each with a distal opening leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.

Remarks.

The most distinctive character of the genitalia of Laeocathaica prionotropis is the lump formed by the fusing pilasters in the penis.

The undetermined juvenile specimens from Chongqing (HBUMM08120) conchologically resemble Laeocathaica prionotropis . However, their locality is far from the distribution area of L. prionotropis on the South Gansu Plateau [甘南高原].

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 3. Distribution map of Laeocathaica dityla Moellendorff, 1899 (green dots), L. odophora Moellendorff, 1899 (pink dots), L. pewzowi Moellendorff, 1899 (white dots), L. phaeomphala Moellendorff, 1899 (blue dots), L. polytyla Moellendorff, 1899 (yellow dots), L. potanini Moellendorff, 1899 (red dots), and L. prionotropis Moellendorff, 1899 (orange dots). Range of the map is defined by the rectangle in the thumbnail.

Figure 29. Laeocathaica prionotropis Moellendorff, 1899 A SMF 9080, lectotype of L. prionotropis albocincta Moellendorff, 1899 B SMF 9078, lectotype C SMF 9079, paratype D HBUMM 05549 - spec. 1.

Figure 30. Laeocathaica prionotropis Moellendorff, 1899, HBUMM 05549 - spec. 1 A general view B right view of dart sac apparatus C ventral view of dart sac apparatus. 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. Asterisk * indicates the opening of proximal accessory sac.

Figure 43. Internal view of penis A, B Laeocathaica polytyla Moellendorff, 1899, HBUMM 05437 - spec. 1 A general view B proximal penis C L. potanini Moellendorff, 1899, HBUMM 00633 - spec. 1, general view D L. prionotropis Moellendorff, 1899, HBUMM 05549 - spec. 1, general view E, F L. stenochone Moellendorff, 1899, HBUMM 05495 - spec. 1 E general view F proximal penis G L. carinifera (H. Adams, 1870), HBUMM 08443 - spec. 2, general view. Arrows indicate the position where two penial internal pilasters fuse into one distal pilaster.

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 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.