Laeocathaica distinguenda 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 distinguenda Moellendorff, 1899

Figs 2A, D , 15 , 16 , 41D, E , 46A, B , 50A , 51

Laeocathaica distinguenda Möllendorff, 1899: 93, pl. 5, fig. 6; - Gude 1902: 6; - Yen 1939: 149, pl. 15, fig. 32; - Yen 1942: 283; - Chen and Zhang 2004: 317, fig. 305.

Laeocathaica subsimilis distinguenda - Wiegmann 1900: 99, pl. 3, fig. 88.

Laeocathaica (Laeocathaica) distinguenda - Zilch 1968: 173; - Richardson 1983: 78.

Laeocathaica amdoana - Páll-Gergely et al. 2022: 38, fig. 5A.

Museum material.

SMF 8959, lectotype; Thal des Pui-ho (= Baishuijiang) [白水江] b. Lum-du , Sy-tshuan, China; ex Potanin 906, Slg. O. v. Möllendorff. SMF 95024 , paratype; SO-Gansu, NW-China; Slg. C. R. Böttger 1904 (ex Möllendorff!). " Aegista distinguenda ", ZIN RAS No. 7, 3 fms. ZIN RAS No. 22, 1 fma and 1 subadult, between Nanping [南坪 = Jiuzhaigou 九寨沟县] and Sung-Pan (= Songpan), coll. Beresowskij, 1894, det. Möllendorff.

New material.

HBUMM00674, numerous fma; Jianshanxiang [尖山乡], Wenxian, Gansu Province, 840 m a.s.l., 33.043291°N, 104.869226°E; 1998-IV-29, coll. Chen, D.-N. HBUMM05473, many fms, Jidushan [基督山] near Baishuijiang , Wenxian, Gansu Province, near point (32.94936°N, 104.699043°E); limestone and broad-leaved woods; 2006-IX-27; coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, Lin-Hui [高林辉]. HBUMM05434, HBUMM05436, 2 fma dissected, Yuxushan [玉虚山], Wenxian, Gansu Province, 32.957259°N, 104.689152°E, shrubs and slate, 2006-IX-27, coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05415, 2 fma; hill foot of Yuxushan , Wenxian, Gansu Province; slate and shrubs, dry; 2006-IX-27, coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM05417, numerous fma, 3 fma dissected, same data as HBUMM05436; DNA voucher HBUMM05407. HBUMM05434, 2 fma; same data as HBUMM05436. HBUMM05479, 2 fma; 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 HBUMM05442. HBUMM05554, 1 fma, 4 subadults. HBUMM05565a, many fma. HBUMM05571, many fma and subadults. HBUMM05576b, 1 fma: Hengdan [横丹], 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. HBUMM06536, not dissected; town of Wenxian , Gansu Province, 1024 m a.s.l., 32.941111°N, 104.668889°E, 2011-VIII-08, coll. Wu, M., Xu, Qin [徐沁], Budha , Prem ; DNA voucher HBUMM06535. HBUMM06579, 2 fma, 4 fms and 1 juv (DNA voucher HBUMM06578). HBUMM06556, 2 fms: Fengchengsi [风成寺], Jiuzhaigouxian, Sichuan Province, 1515 m a.s.l., 33.256280°N, 104.238386°E; 2011-VIII-14, coll. Wu, M., Xu, Q. and Buhda, P. ; DNA voucher HBUMM06555. HBUMM06646, 1 fma; Wenxian , Gansu Province, 1269 m a.s.l., 33.091944°N, 104.360833°E; 2011-VIII-9, coll. Wu, M., Xu, Q. and Buhda, P. HBUMM06736, 1 subadult; Eastern bank of Baishuijiang , Lihuacun [梨花村], Jiuzhaigouxian, Sichuan Province, 1425 m a.s.l., 33.267222°N, 104.234722°E; 2011-VI-14, coll. Wu, M., Xu, Q. and Buhda, P. HBUMM08148, 1 fma and several subadults; 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. HBUMM08429, 1 fma dissected; toward Danpuzhen [丹堡镇] on national road 212, Wenxian, Gansu Province, near 32.872636°N, 104.784372°E, on slope, 2019-X-13, coll. Li, Q.-M. ; DNA voucher HBUMM08429a. CZG202008-w5, 8 fms, Zhangzhazhen [彰扎镇], Jiuzhaigouxian, Sichuan Province, near point (33.30674°N, 103.877345°E); 2020-VIII, coll. Chen, Z.-G. CZG202008-w9, Heihexiang [黑河乡], Jiuzhaigouxian, Sichuan Province, near (33.54941°N, 104.041691°E); 2020-VIII, coll. Chen, Z.-G. CZG202008-w10, 5 fms, whitish shells; Shijiba [石鸡坝], Wenxian, Gansu Province, near point (33.067936°N, 104.456784°E); 2020-VIII, coll. Chen, Z.-G.

Distribution.

Gansu: Wenxian (type locality), Wudu (type locality), Zhouquxian (type locality); Sichuan: Jiuzhaigouxian (type locality).

Additional information of shell.

The protoconch has the widespread granules each of which is ~ 20 - ~ 40 µm long and looks like a hump embedded in a shallow socket. All granules are present on densely arranged fine radial threads. Spiral grooves are shallowly present. The growth lines on teleoconch are more or less distinct but are irregularly arranged.

General anatomy.

Eversible head wart present. Jaw arcuate, with 4-7 projecting ribs.

Anatomy of genital organs.

Penial sheath moderately long, covering ~ 1/6-1/4 of penis. Penis slightly expanded distally. Inside penis, two penial pilasters forming one Y-shaped fork at proximal 1/3 penis (only indistinct in HBUMM05417); a rather thick pilaster that is made up of numerous thread-like longitudinal folds occupying proximal 1/3-2/3 of penis. Pilasters of median to distal penis weaving into a delicate net that bears regularly arranged diamond-shaped papillae, which are usually lost perhaps because of bad specimen condition. Distally inside penis, fine pilasters merging into 6-9 short and more or less thick folds near opening of epiphallus. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/4 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 9 mm long, apically 2-bladed. Accessory sac small but distinguishable from outside, internally solid, inserting into dart sac at middle part, opening to distal dart chamber. Mucous glands 4 (HBUMM08429, each complicatedly branched) - 12 (HBUMM05479, simply branched). Proximal accessory sacs two, symmetrical or the right one slightly larger, separated dorsally and ventrally, with a few internal pilasters, each with an opening leading to dart chamber near opening of dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate.

Remarks.

The sculpture on protoconch and teleoconch of this species (Fig. 46A, B ) resembles that of Laeocathaica amdoana (Fig. 46G, H ). However, in Laeocathaica distinguenda , as in the most congeners of Laeocathaica , the part of the vagina between atrium and dart sac apparatus is not elongated.

For more comments, see Laeocathaica tropidorhaphe .

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 15. Laeocathaica distinguenda Moellendorff, 1899 A SMF 8959, lectotype B SMF 95024 C HBUMM 054361.

Figure 16. Genital anatomy of Laeocathaica distinguenda Moellendorff, 1899 A, B HBUMM 05436 C, D HBUMM 05417 A general view B-D ventral, left, and apical views of dart sac apparatus. Abbreviations: AO - opening of accessory sac leading to dart chamber; 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; PAS - proximal accessory sac; PO - opening of proximal accessory sac leading to dart chamber; PR - penial retractor muscle; PS - penial sheath; Va - vagina; VD - vas deferens. Asterisk * indicates the opening of proximal accessory sac.

Figure 41. Internal view of penis A, B Laeocathaica amdoana Moellendorff, 1899, HBUMM 05640 - spec. 1 A general view B proximal penis C L. filippina (Heude, 1882), HBUMM 04166 - spec. 1, general view D, E L. distinguenda Moellendorff, 1899, HBUMM 05436 - spec. 1 D general view E proximal penis F L. carinalis Chen & Zhang, 2004, HBUMM 08300 - spec. 1, general view G, H L. dityla Moellendorff, 1899, HBUMM 00698 - spec. 1 G general view H proximal 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.