Laeocathaica amdoana 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 amdoana Moellendorff, 1899

Figs 2A, D , 5A, B , 6 , 41A, B , 46G, H , 50A , 51 , 52E

Laeocathaica amdoana Möllendorff, 1899: 92, pl. 5, fig. 5; - Gude 1902: 6; - Yen 1939: 148, pl. 15, fig. 31; - Páll-Gergely et al. 2022: 38, fig. 3A, B.

Laeocathaica stenochone amdoana - Wiegmann 1900: 104, pl. 3, figs 91-93.

Laeocathaica (Laeocathaica) amdoana - Zilch 1968: 173; - Richardson 1983: 77.

Museum material.

ZIN RAS No. 2, 1 fma and 1 subadult, " Aegista amdoana Möllendorff.”, Wen-Xian [文县], 1885-IX-8, coll. Potanin, det. Möllendorff. SMF 8952, lectotype, Ho-dshi-gou, Gansu, China . ex Potanin 853. Slg. O. v. Möllendorff. SMF 8953, paratype, Wen-hsien , SO-Gansu, China .

New material.

HBUMM05600, numerous fma and juvs; southern slope of Beishan [北山], Wudu [武都], Longnan [陇南], Gansu Province , limestone hill with sparse shrubs; 2006-X-02, coll. Liu, Jian-Min [刘建民], Zheng , Wei [郑伟]. HBUMM05640, numerous specimens, 2 fma dissected, Jiaogongzhen [角弓镇], Wudu , Gansu Province , (33.57°N, 104.64°E), broken limestone rocks, 2006-X-02, coll. Liu, J.-M. and Zheng, W. HBUMM05639, 4 fma and 1 juv, 1 fma dissected; Jiaogongzhen , Wudu , Gansu Province (33.57°N, 104.64°E), broken limestone rocks, 2006-X-02, coll. Liu, J.-M. and Zheng, W. HBUMM8147, 1 fma dissected: coll. data as HBUMM05640; DNA voucher HBUMM05626. HBUMM08147, 1 fma dissected, Wudu , Gansu Province, 927 m a.l.s., (33.346111°N, 105.044°E), adults on cliff and juvs on deadwood, 2017-VIII-05, coll. Shen, Xue-Fen [盛雪芬], etc.; DNA voucher HBUMM08147a. HBUMM08432, near Shichuanba [石川坝], Wenxian, Gansu Province, near 33.17534°N, 105.019362°E; 2019-X-12, coll. Li, Qi-Ming [李启明]; DNA voucher HBUMM08432a .

Distribution.

Gansu: Wudu, Wenxian (type locality).

Additional information of shell.

Very fine and slim granules (~ 20 µm long) are present on the protoconch. After the fourth whorl, including the umbilical region, irregularly arranged spiral grooves are present. On teleoconch whorls, the growth lines are low and indistinct.

General anatomy.

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

Anatomy of genital organs.

Penial sheath moderately long, covering ~ 1/3 of penis. Penis slightly expanded distally. Inside penis, two penial internal pilasters fusing into Y-shaped fork at proximal 1/4, accompanied with 4-6 low pilasters. Distally inside penis, numerous fine pilasters merging into ~ 4 short but thick folds near opening of epiphallus. Vas deferens narrow throughout. Vagina between atrium and dart sac elongated, ~ 2 times longer than dart sac. Vagina between dart sac and insertion of bursa copulatrix duct short, ~ 3 mm in length. Dart sac ~ 1/3 length of penis. Love dart ~ 1.2 mm long, apically 2-bladed. Accessory sac small, internally solid, inserting into dart sac medially, opening to distal dart chamber. Mucous glands seven or eight, each complicatedly branched. Proximal accessory sacs two, not touching; each proximal accessory sac tiny, ~ 0.5 × 0.5 mm2 (HBUMM05640-spec.1) or smaller (HBUMM05640-spec.2) in size; each with an opening leading to dart sac chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate.

Remarks.

By shell sculpture, including that on the protoconch, this species (Fig. 46G, H ) cannot be immediately distinguished from Laeocathaica tropidorhaphe (Fig. 46I, J ) and L. distinguenda (Fig. 46A, B ). It seems that L. tropidorhaphe forms a continuous variation of shell shape and the conchological delimitation among these three species that are geographically coexistent is not so distinct ( Páll-Gergely et al. 2022). According to our observation, the key morphological features to practically distinguish the three species are:

brownish/chestnut coloration. In Laeocathaica amdoana such coloration is usually (!) obviously darker and in L. distinguenda the coloration is usually very pale (in our Fig. 15C , it is a relatively darker shell). However, in HBUMM05479 (dissected, shell not pictured) the shell is in very pale dirty yellow except peripherally whitish.

peripheral angulation. The level of angulation, in order is Laeocathaica tropidorhaphe > (>: stronger than; >>: much stronger than) L. amdoana >> L. distinguenda , which is also indicated by the geometric morphometric analyses: compared to L. amdoana (Fig. 5A, B ; average shape ‘b’ in Fig. 50A ) and L. tropidorhaphe (Fig. 33 ; average shape ‘c’ in Fig. 50A ), the shell of L. distinguenda appears to be much broader in aperture and the periphery is very bluntly round ( Möllendorff 1899).

the structure of umbilicus. In both Laeocathaica amdoana and L. tropidorhaphe , the umbilicus is funnel-shaped and through which every whorl is visible. However, in Laeocathaica distinguenda , the penultimate whorl is much projecting and makes a suddenly enlarged umbilicus ( Möllendorff 1899; our own observation, e.g., to compare Fig. 15 with Figs 5A, B , 33 ). Laeocathaica amdoana , L. distinguenda , and L. tropidorhaphe are clearly distinguishable based on genital features. Laeocathaica amdoana and L. tropidorhaphe , both of which have a long vagina between dart sac apparatus and atrium that was not seen in L. distinguenda , can be distinguished by the presence of proximal accessory sac in the former species. Laeocathaica tropidorhaphe belongs to the three species in Laeocathaica that have no proximal accessory sac. In Laeocathaica amdoana there are two tiny proximal accessory sacs, and each has an opening leading to the dart sac chamber, which means the proximal accessory sacs could be functional compared to the blind one (i.e., without opening) observed in Pseudiberus tectumsinense pingi ( Zhang et al. 2021). In addition, in Laeocathaica amdoana the dart is apically 2-bladed rather than 4-bladed in L. tropidorhaphe .

The phylogeny based on ITS2 and 16S suggests that Laeocathaica amdoana is sister to L. distinguenda and L. tropidorhaphe (Fig. 51 ). In Chen and Zhang (2004: 316, fig. 303), the species identified as Laeocathaica amdoana is dubious and looks like a L. distinguenda . 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 5. A, B Laeocathaica amdoana Moellendorff, 1899 A SMF 8952, lectotype B HBUMM 05640 - spec. 1 C L. dolani (Pilsbry, 1934), HBUMM 00069 - spec. 1 D L. carinalis Chen & Zhang, 2004, HBUMM 08400 - spec. 1.

Figure 6. Genital anatomy of Laeocathaica amdoana Moellendorff, 1899, HBUMM 05640 - spec. 1 A general view B left part of dart sac apparatus C right view of dart sac apparatus, showing position of proximal accessory sacs D 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; FO - free oviduct; MG - mucous glands; P - penis; PAS - proximal accessory sac; PR - penial retractor muscle; PS - penial sheath; Va - vagina; VD - vas deferens. Asterisk * indicates the opening of proximal accessory sac.

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

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

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.

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.