Laeocathaica carinifera (H. Adams, 1870)

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 carinifera (H. Adams, 1870)

Figs 2A, C , 9 , 10 , 43G , 45I, J , 50B , 51

Helix (Plectotropis) christinae var. carinifera H. Adams, 1870: 377.

Helix subsimilis Deshayes, 1874: 10, pl. 2, figs 28, 29; - Heude 1882: 22, pl. 20, fig. 18; - Standen 1905: 231.

Helix (Plectotropis) subsimilis - Ancey 1883: 7, pl. 2, figs 28, 29.

Helix christinae var. carinifera - Gredler 1884: 264 (= Helix subsimilis Deshayes, 1873); - Möllendorff 1884: 351.

Helix (Cathaica) christinae var. subsimilis - Pilsbry 1892: 214, pl. 49, figs 29-33.

Helix (Cathaica) subsimilis - Kobelt 1894: 713, pl. 203, figs 1-3.

Laeocathaica christinae carinifera - Richardson 1983: 78.

Laeocathaica (Laeocathaica) subsimilis - Zilch 1968: 175; - Richardson 1983: 79; Wu and Chen 1998: 107.

Cathaica subsimilis - Yen 1938: 456.

Laeocathaica subsimilis subsimilis - Yen 1939: 148, pl. 15, fig. 28.

Laeocathaica subsimilis - Möllendorff 1899: 44; - Wiegmann 1900: 96, pl. 3, figs 84-87; - Yen 1942: 283; - Gude 1902: 6; - Chen and Zhang 2004: 313, fig. 299.

Laeocathaica (Laeocathaica) filippina - Wu 2004: fig. 17.

Laeocathaica carinifera - Páll-Gergely et al. 2022: 50, fig. 12.

Museum material.

Laeocathaica subsimilis (Deshayes, 1874): NHMUK V. W. McAndrew coll. Acc. no. 1563. SMF 95116 , one fms; Szetschwan, Noerdl. der Stadt Juanj-juanj, China; ex Mus. Petersb. 1905 ex Potanin. Det. Möllendorff. SMF 24265 , one fms and one near matured; Yang-dsy-gebiet; ex Heude. Slg. O. v. Möllendorff. SMF 294292, two juvs (the third one, a fms of L. filippina ); Sytshuan, China; Slg. C. Bosch ex H. Rolle. SMF 24261 , two fms; Liu-ba-ting, Shen-hsi, China; ex Potanin 451, Slg. O. v. Möllendorff. SMF 24258 , three fms; O-Sytshuan; ex Möllendorff. Slg. W. Kobelt. SMF 24257 , four fms; O-Sy-tshuan; ex. B. Schmacker, Slg. O. v. Möllendorff. SMF 24262 , four fms; Sy-tshuan, Chung-king, W-M-China; ex Möllendorff, 1890, Slg. O. Böttger. SMF 24255 , four fms; O-Sy-tshuan; ex L. Fuchs, Slg. O. v. Möllendorff. SMF 95712 , one fms; Prov. Sze-chuen; Slg. C. R. Böttger, 1905. SMF 294294, one fms. China; Slg. C. Bosch ex Hermann Rolle. The species of following specimens are not decided: SMF 24260 , one fms (with aperture not full matured), Kwan-juon-hszien (= Guangyuan) [广元县], Prov. Sze-Csuen, China, ex Kormos (Budapest). SMF 24256 , three fms, Zw. Guan-yuan u. Dshau-hoa [昭化], Potanin 275, Slg. O. v. Möllendorff. SMF 24264 , four fms, Lue-feng-kou bei Guan-yuan, ex Potanin 270, Slg. O. v. Möllendorff .

New material.

HBUMM00006, 4 fma, 4 juvs; Xinjianxiang [新建乡], Nanchong [南充市], Sichuan Province, near point (30.963475°N, 105.842133°E); 1964-V-20, coll. Chen, De-Niu. [陈德牛]. HBUMM00228, 1 fma and 2 subadults; Beibei [北碚], Chongqing [重庆], near point (29.809307°N, 106.411375°E); 1980, coll. Chen D.-N. HBUMM01573, 7 fma, 1 dissected; Muping [穆坪镇], Baoxingxian [宝兴县], Sichuan Province; the first hill to the south, between point A (1090 m a.s.l., 30.360333°N 102.7255°E) and point B (1021 m a.s.l., 30.341389°N 102.783722°E); limestone, 2 cm- thick litter layer, typical local vegetation type, seminatural, nearby cultivated with maize; 2003-VII-23, coll. Wu, Min [吴岷]. HBUMM03118, 4 juvs; Changping [长坪], Wanzhou [万州], Chongqing, near point (30.448543°N, 108.260695°E); 2003-VII-16, coll. Yuan, C. etc. HBUMM04064, many fma, 1 dissected; Meizixiang [梅子乡], Fengjiexian [奉节县], Chongqing, near point (430 m a.s.l., 31.178028°N, 109.341583°E); mixed rocks of slate and sandstone, semimature environment, very thin litter layer, nearby with eucalyptus trees; 2004-VII-18, coll. Wu, M. and Wu, Qin [吴琴]; DNA voucher HBUMM05121. HBUMM04162, numerous fma, 1 dissected, Shibaozhai [石宝寨], Zhongxian [忠县], Chongqing, 238 m a.s.l., 30.423083°N, 108.183972°E, shrubs and weed, purple sandy stone, humid, 2004-VII-16, coll. Wu, M., Wu, Q. and Qi, Gang [齐钢]; DNA voucher HBUMM05103. HBUMM04217, 10 fma; Tiejiacun [铁甲村], Jiangnanxiang [江南乡], Fengjiexian, Chongqing, 1005 m a.s.l., 30.956444°N, 109.468306°E; limestone, grass and shrubs, humid; 2004-VII-19, coll. Wu, M. and Wu, Q. HBUMM04219: many fma; same coll. data as HBUMM04064. HBUMM04232, several fma and juvs; near Xinmincun [新民村], Jiangnanxiang , Fengjiexian, Chongqing, 980 m a.s.l., 30.932583°N, 109.455639°E; mixed rocks of sandstone and limestone, grass and shrubs, humid, semi-farmland; 2004-VII-19, coll. Wu, M. and Wu, Q. HBUMM04235, 2 fma and 1 juv, 1 dissected, near town of Wuxixian [巫溪县], Chongqing, 339 m a.s.l., 31.402111°N, 109.634722°E; limestone, grass and shrubs, bare earth without litter layer; 2004-VII-22, coll. Wu, M. ; DNA voucher HBUMM05131). HBUMM08443, 4 fma and 1 subadult, 4 dissected, Baidicheng [白帝城], Fengjie County, Chongqing, two kilometers away from the point (241 m a.s.l., 31.020194°N ,, 109.472833°E), slate and sandy stones, woods with shrubs and trees, humid, 2004-VII-20, coll. Wu, M.

Distribution.

Sichuan: Nanchong, Baoxing. Chongqing: Beibei, Changshouxian [长寿县], Fengduxian [丰都县] (type locality: Fungsiang gorge = Fengxiangxia [风箱峡]), Fengjiexian, Kaixian [开县], Liangpingxian [梁平县], Pengshuixian [彭水县], Shizhuxian [石柱县], Wanzhou, Wushanxian [巫山县], Wuxi, Yunyangxian [云阳县], Zhongxian. Shaanxi: Lueyang [略阳].

Additional information of shell.

On adult shell the protoconch is granulate with thick granules of ~ 40 µm long, each of which looks like a hump in a pit. The obscurity of such granules is caused by erosion or weathering. Spiral grooves are absent throughout the shell. On teleoconch the growth lines are low but distinct.

General anatomy.

Eversible head wart weak. Jaw arcuate, with four or five projecting ribs.

Anatomy of genital organs.

Penial sheath moderately long, covering ~ 1/4-1/3 of penis. Penis somewhat expanded distally. A pair of penial internal pilasters fusing into one Y-shaped fork at proximal 1/3, and another pair fusing into one Y-shaped fork at proximal 1/2 of penis; some other low pilasters variably present. Distally inside penis, numerous fine pilasters merging into 6-8 short but 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 approximately as long as or slightly shorter than 2/3 of dart sac. Dart sac ~ 1/2 of penis. Love dart ~ 2 mm long. Accessory sac spherical, internally almost solid, inserting into dart sac medially, opening to distal dart chamber. Mucous glands six or seven, each simply or complicatedly branched. Proximal accessory sacs two, asymmetrical, left one larger than right one, dorsally separated and ventrally touching, with internal pilasters, each with an opening leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.

Remarks.

See Laeocathaica stenochone .

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 9. Laeocathaica carinifera (H. Adams, 1870) A NHMUK V. W. McAndrew coll. Acc. no. 1563 B SMF 24265 C SMF 95116 D HBUMM 08443 - spec. 1 E HBUMM 08443 - spec. 2.

Figure 10. Genital anatomy of Laeocathaica carinifera (H. Adams, 1870), HBUMM 08443 - spec. 1 A general view B apical view of dart sac apparatus C cross-section of dart sac at the position arrowed in (A). Abbreviations: AS - accessory sac; At - atrium; BC - bursa copulatrix; BCD - bursa copulatrix duct; DS - dart sac; DtC - a chamber containing love dart; Ep - epiphallus; FO - free oviduct; L - left; 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; R - right; Va - vagina; VD - vas deferens.

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 45. SEM micrographs A, B Laeocathaica potanini Moellendorff, 1899, HBUMM 00633 A protoconch B teleoconch C, D L. dityla Moellendorff, 1899, HBUMM 00698 C protoconch D teleoconch E, F L. qingchuanensis Wu, sp. nov., HBUMM 01179 a-spec. 6, paratype E protoconch F teleoconch G, H L. christinae (H. Adams, 1870), HBUMM 01251 a G protoconch H teleoconch I, J L. carinifera (H. Adams, 1870), HBUMM 04162 I protoconch J teleoconch K, L L. polytyla Moellendorff, 1899, HBUMM 05437 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.