Laeocathaica phaeomphala 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 phaeomphala Moellendorff, 1899

Figs 2A , 3 , 22 , 23 , 42D , 48E, F , 51 , 52D

Laeocathaica phaeomphala Möllendorff, 1899: 96, pl. 6, fig. 3; - Wiegmann 1900: 111, pl. 3, figs 101-103; - Gude 1902: 6; - Yen 1939: 149, pl. 15, fig. 37; - Chen and Zhang 2004: 325, fig. 314; - Páll-Gergely et al. 2022: 67, fig. 18C.

Laeocathaica (Laeocathaica) phaeomphala - Zilch 1968: 174; - Richardson 1983: 78.

Museum material.

SMF 9089, lectotype; Wenhsien, S-Gansu; Potanin 51b, 72, 741, Slg. O. v. Möllendorff. SMF 9090, paratypes, four fms; the same data as lectotype. ZIN RAS No. 4, 1 fma and 1 subadult, Wen-Xian, 1885-IX-8, coll. Potanin, det. Möllendorff.

New material.

HBUMM05433, 2 fma, 1 fms, and many juvs, all fma dissected; Yuxushan , 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. HBUMM05477, numerous fma; Jidushan nearby Baishuijiang , Wenxian, Gansu Province; limestone and broad-leaved woods; 2006-IX-27; coll. Wu, M., Liu, J.-M., Zheng, W. and Gao, L.-H. HBUMM08424, slope near Wenzhoulu [文州路], Wenxian, Gansu Province, near 32.943151°N, 104.692505°E, on slope; 2019-X-12, coll. Li, Q.-M. ; DNA voucher HBUMM08424a. CZG202008-w3, 9 subadults, Shangdezhen , Wenxian, Gansu Province, 2020-VIII, coll. Chen, Z.-G.

Distribution.

Gansu: Wenxian (type locality).

Additional information of shell.

The first 11/4 protoconch whorls are almost smooth, and the subsequent 1/4 whorls have dense radially arranged fine threads. Spiral grooves are regularly present throughout body whorl.

General anatomy.

Head with flat but distinct eversible head wart. Jaw arcuate, with 5-7 projecting ribs.

Anatomy of genital organs.

Penial sheath covering ~ 1/5 of penis. Penis tubular and distally thicker. Inside penis, ~ 7 high parallel pilasters on proximal 1/2, no pilasters fusing into Y-shaped fork. Fine pilasters on distal end of penis merging into 9-12 thin or thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac extremely elongated, ~ 4 × longer than dart sac. Vagina between dart sac and insertion of bursa copulatrix duct approximately as long as dart sac. Dart sac ~ 1/6 length of penis. Love dart ~ 1.5 mm long, apically 2-bladed. Accessory sac tiny, internally empty, inserting into dart sac at middle part, opening to dart chamber. Mucous glands eight, each singly tubular or simply branched. Proximal accessory sac absent. Bursa copulatrix duct equally thick.

Remarks.

This species is one of the three species that do not have proximal accessory sac on dart sac apparatus. However, the shell morphology of Laeocathaica phaeomphala differs greatly from that of the other two species L. polytyla and L. 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 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 22. Laeocathaica phaeomphala Moellendorff, 1899 A SMF 9089, lectotype B SMF 9090, paratype C HBUMM 05433 a-spec. 6.

Figure 23. Genital anatomy of Laeocathaica phaeomphala Moellendorff, 1899, HBUMM 05433 a-spec. 6 A general view B, C ventral and left views of dart sac apparatus. Abbreviations: AS - accessory sac; At - atrium; 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 42. Internal view of penis A, B Laeocathaica dolani (Pilsbry, 1934), HBUMM 00069 - spec. 1 A general view B proximal penis C L. christinae (H. Adams, 1870), HBUMM 01251 a-spec. 1, general view D L. phaeomphala Moellendorff, 1899, HBUMM 05433 - spec. 1 E L. odophora Moellendorff, 1899, HBUMM 08430 - spec. 1 F L. pewzowi Moellendorff, 1899, ZIN RAS No. 4, " Aegista pewzowi Schalf. " G, H L. parapolytyla Wu, sp. nov., HBUMM 06640 - spec. 1, holotype G general view H proximal part. Arrows indicate the position where two penial internal pilasters fuse into one distal pilaster.

Figure 48. SEM micrographs A, B Laeocathaica qishilii Wu, sp. nov., HBUMM 08298, paratype A protoconch B teleoconch C, D L. qiminglii Wu, sp. nov., HBUMM 08448, paratype C protoconch D teleoconch E, F L. phaeomphala Moellendorff, 1899, CZG 202008 - w 3, subadult E protoconch F teleoconch G, H L. filippina (Heude, 1882), HBUMM 04166 G protoconch H teleoconch I, J L. parapolytyla Wu, sp. nov., HBUMM 06640 - spec. 9, paratype I protoconch J teleoconch.

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.

Figure 52. Schematics of terminal genitalia of some dart-sac-bearing camaenids A Cathaica fasciola (Draparnaud, 1801) (stylized pattern from HBUMM 8142 - spec. 1 - 15, 9 fma; Qingyang, Gansu, 35.738 ° N, 107.701 ° E, 1353 m a. s. l.; coll. Sheng, X. - F., 2017 - VII) B " Cathaica " Cathaica pulveratricula (Martens, 1882) (stylized pattern from HBUMM 08208), one proximal accessory sac is present at the left side of dart sac C Laeocathaica spp. with two proximal accessory sacs at both sides of dart sac D L. phaeomphala Moellendorff, 1899, without proximal accessory sac and with elongated vagina above dart sac E L. amdoana Moellendorff, 1899, with two tiny proximal accessory sacs and with elongated vagina above dart sac F L. dolani (Pilsbry, 1934), with a proximal accessory sac at the right side of dart sac. Notes: any fleshy septum inside dart sac chamber is not shown; red dots indicate openings of proximal accessory sac. Abbreviations: AS - accessory sac; DS - dart sac; DSC - dart sac chamber; DtC - a chamber containing love dart; MG - mucous glands; PAS - proximal accessory sac; PR - penial retractor muscle; PS - penial sheath; Va - vagina.