Laeocathaica amdoana Moellendorff , 1899

Pall-Gergely, Barna, Hunyadi, Andras, Meng, Kaibaryer & Fekete, Judit, 2022, A review of the genus Laeocathaica Moellendorff, 1899 (Gastropoda, Pulmonata, Camaenidae), ZooKeys 1086, pp. 33-76 : 33

Laeocathaica amdoana Moellendorff, 1899

Figures 3 , 4 , 5 , 6 , 7

Laeocathaica amdoana Möllendorff, 1899: 92-93, pl. 5, fig. 5.

Laeocathaica distinguenda Möllendorff, 1899: 93, pl. 5, fig. 6. new synonym

Laeocathaica tropidorhaphe Möllendorff, 1899: 94, pl. 5, fig. 7. new synonym

Laeocathaica amdoana . - Yen 1939: 148, pl. 15, fig. 31.

Laeocathaica distinguenda . - Yen 1939: 149, pl. 15, fig. 32.

Laeocathaica tropidorhaphe . - Yen 1939: 149, pl. 15, fig. 33.

Laeocathaica (Laeocathaica) amdoana . - Zilch 1968: 173.

Laeocathaica (Laeocathaica) distinguenda . - Zilch 1968: 173.

Laeocathaica (Laeocathaica) tropidorhaphe . - Zilch 1968: 175.

Laeocathaica amdoana . - Chen & Zhang 2004: 316, fig. 303.

Laeocathaica distinguenda . - Chen & Zhang, 2004: 318, fig. 305.

Laeocathaica tropidorhaphe . - Chen & Zhang 2004: 319, fig. 307.

Laeocathaica dangchangensis Chen & Zhang, 2004: 339 [Chinese description], 443 [English description], fig. 332. new synonym

Type material.

China (Gansu), Ho-dshi-gou, coll. Möllendorff ex coll. Potanin 853, SMF 8952 (lectotype of Laeocathaica amdoana , Fig. 3A, B ) • China (SO-Gansu), Wen-Hsien, coll. Möllendorff ex coll. Potanin 907, SMF 8953 (paralectotype of Laeocathaica amdoana ) • China (Sy-tshuan): Thal des Pui-hob. Lum-du, coll. Möllendorff ex coll. Potanin 906, SMF 8959 (lectotype of Laeocathaica distinguenda , Fig. 5A ) • SO-Gansu (NW China), coll. C.R. Boettger ex coll. Möllendorff, SMF 95024/1 paralectotype of Laeocathaica distinguenda • SO-Gansu: Nanping, coll. Möllendorff ex coll. Potanin 8, 64, 846, SMF 8958/5 paratypes of Laeocathaica distinguenda • SO-Gansu, Yü-Lin-Guam, u. Wen-hsien, coll. Möllendorff ex coll. Potanin 11, 521, 565, SMF 8955/3 paralectotypes of Laeocathaica distinguenda • China: SO-Gansu, coll. Möllendorff ex coll. Potanin 725a, SMF 8957/3 paralectotypes of Laeocathaica distinguenda • Shy-Pu am Pui-hu, coll. Möllendorff ex coll. Potanin 69, 653, SMF 8956/2 paralectotypes of Laeocathaica distinguenda • SO-Gansu, zw. Li-dshia-pu u. Hsi-gu-tsheng, coll. Möllendorff ex coll. Potanin 923, SMF 9074 (lectotype of L. tropidorhaphe , Fig. 7A ) • Same data, SMF 9077/2 paralectotypes • NW-China, SO-Gansu, coll. C.R. Boettger 1904 ex coll. Möllendorff SMF 95126/1 (paralectotype of L. tropidorhaphe ) • SO-Gansu, Tan-tshang, coll. Möllendorff ex coll. Potanin 545, 623, 808, SMF 9075/4 (paralectotypes of L. tropidorhaphe ) • Gansu: Dshie-dshou, coll. Möllendorff ex coll. Potanin 119, SMF 9076/1 (paralectotype of L. tropidorhaphe ) • IZCAS TM 095895 (labelled as holotype of L. dangchangensis , but its measurements do not match with the ones given in the original description, see also remarks, Fig. 7D ) • IZCAS TM 095843-095894 + IZCAS TM 095896-095950 (103 paratypes of L. dangchangensis in ethanol).

Museum material.

Szechwan, coll. Möllendorff, SMF 42564 /1 • Tan-Tschan, coll. Möllendorff, SMF 24269 /1 (" tropidorhape ") .

New material (typical Laeocathaica tropidorhaphe ).

China • 2 shells; Gansu, Longnan Shi, Dangchang Xian, Chengguan Zhen , Wenchangmiao , temple hill (locality code: 2016/87); 34°02.394'N, 104°23.499'E; 1835 m a.s.l.; 02 June 2016; A. Hunyadi leg.; HA • 2 shells; Gansu, Longnan Shi, Dangchang Xian, Lianghekou Xiang, Lianghekou Cun, rock above the intersection (locality code: 2016/89); 33°41.808'N, 104°29.182'E; 1245 m a.s.l.; 02 June 2016; A. Hunyadi leg.; HA (Fig. 8C ) • 2 shells; Gansu, Longnan Shi, Dangchang Xian, Guanting Zhen, 1.5 km north of Guanting towards Dangchang (locality code: 2016/88); 33°50.803'N, 104°32.470'E; 1815 m a.s.l.; 02 June 2016; A. Hunyadi leg.; HA (Fig. 7B ) • 2 shells; Gansu, Longnan Shi, Zhugqu Xian, 2.5 km west of Suoertou Cun , northern bank of Bailong He (locality code: 2016/91); 33°46.906'N, 104°20.106'E; 1235 m a.s.l.; 03 June 2016; A. Hunyadi leg.; HA • 2 shells; Gansu, Longnan Shi, Wudu Xian, Jiaogong Zhen, 1.5 km west of Chenjiaba Cun , Zhaoyangdong, below the cave (locality code: 2016/95); 33°31.924'N, 104°39.286'E; 1175 m a.s.l.; 04 June 2016; A. Hunyadi leg.; HA .

New material (other morphs).

6 shells; Sichuan, Aba, Jiuzhaigou Xian, Baihe Xiang, southern edge of Taiping Cun, rock wall facing north (locality code: 2016/72); 33°18.026'N, 104°09.500'E; 30 May 2016; A. Hunyadi leg.; HA (Fig. 5B, C ) • 1 shell; Sichuan, Aba, Jiuzhaigou Xian, Baihe Xiang, Taiping Cun, eastern bank of Baishui He (locality code: 2016/73); 33°18.366'N, 104°09.413'E; 30 May 2016; A. Hunyadi leg.; HA • 4 shells; Sichuan, Aba, Jiuzhaigou Xian, Anle Xiang, ca. 1.5 km east of Zhongtianshan Cun towards Jiuzhaigou Shi (locality code: 2016/74); 33°17.279'N, 104°12.702'E; 1445 m a.s.l.; 30 May 2016; A. Hunyadi leg.; HA (Fig. 3C-E ) • 4 shells; Sichuan, Aba, Jiuzhaigou Xian, Guoyuan Xiang, 1 km from Guoyuanyi Cun towards Lengshuishan Cun (locality code: 2016/75); 33°07.681'N, 104°18.489'E; 1220 m a.s.l.; 30 May 2016; A. Hunyadi leg.; HA (Fig. 4 ) • 1 shell; Sichuan, Aba, Jiuzhaigou Xian, Guoyuan Xiang, Guoyuaner Cun, environment of the bridge (locality code: 2016/76); 33°06.922'N, 104°19.617'E; 1200 m a.s.l.; 30 May 2016; A. Hunyadi leg.; HA (Fig. 5D ) • 3 shells; Gansu, Longnan Shi, Wenxian, Buziba Xiang, southern edge of Buziba Cun, western bank of the river (locality code: 2016/77); 33°03.592'N, 104°37.094'E; 1215 m a.s.l.; 31 May 2016; A. Hunyadi leg.; HA • 1 shell; Gansu, Longnan Shi, Wenxian, Buziba Xiang, northern edge of Taojiaba Cun, 200 m towards Buziba (locality code: 2016/78); 33°02.706'N, 104°37.157'E; 1200 m a.s.l.; 31 May 2016; A. Hunyadi leg.; HA • 4 shells; Gansu, Longnan Shi, Wenxian, Buziba Xiang, 1 km south of Taojiaba Cun towards Dongyukou Cun (locality code: 2016/79); 33°01.865'N, 104°37.329'E; 1150 m a.s.l.; 31 May 2016; A. Hunyadi leg.; HA • 4 shells; Gansu, Longnan Shi, Wenxian, Chengguan Zhen, next to a museum (locality code: 2016/64); 32°56.471'N, 104°40.379'E; 960-970 m a.s.l.; 28 May 2016; A. Hunyadi leg.; HA • 3 shells; Gansu, Longnan Shi, Wenxian, Chengguan Zhen, cemetery hill above the city (locality code: 2016/65); 32°57.026'N, 104°40.527'E; 1090 m a.s.l.; 28 May 2016; A. Hunyadi leg.; HA • 4 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 1200 m south of Hekou Cun , eastern bank of Bailong He (locality code: 2016/68); 33°01.703'N, 104°53.602'E; 810 m a.s.l.; 29 May 2016; A. Hunyadi leg.; HA • 2 shells; Gansu, Longnan Shi, eastern edge of Wenxian, northern bank of the river (locality code: 2016/66); 32°56.459'N, 104°41.372'E; 28 May 2016; A. Hunyadi leg.; HA • 4 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, southern edge of Hekou Cun, western bank of Bailong He (locality code: 2016/67); 33°02.014'N, 104°53.478'E; 800 m a.s.l.; 29 May 2016; A. Hunyadi leg.; HA • 3 shells; Gansu, Longnan Shi, Wenxian, Shifang Xiang, 800 m from the northwestern edge of Baiyiba Cun towards Dongyukou Cun, left bank of the river (locality code: 2016/83); 32°58.985'N, 104°37.503'E; 970 m a.s.l.; 31 May 2016; A. Hunyadi leg.; HA • 4 shells; Gansu, Longnan Shi, Wenxian, Shifang Xiang, 1300 m northwest from Baiyiba Cun towards Dongyukou Cun (locality code: 2016/82); 32°59.346'N, 104°37.233'E; 980 m a.s.l.; 31 May 2016; A. Hunyadi leg.; HA • 9 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, western edge of Hekou Cun towards Caojiaba, along road no. 212 (locality code: 2016/69); 33°2.343'N, 104°53.045'E; 29 May 2016; A. Hunyadi leg.; HA (Fig. 6E ) • 8 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 600 m west of Jianshan towards Diaohuya (locality code: 2016/70a); 33°02.559'N, 104°51.254'E; 850 m a.s.l.; 29 May 2016; A. Hunyadi leg.; HA (Fig. 6A, B ) • 4 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 200 m north of Lvjiaba, along road no. 212 (locality code: 2016/71); 33°03.712'N, 104°50.209'E; 29 May 2016; A. Hunyadi leg.; HA (Fig. 6C, D ) .

Distribution.

This species is known from several sites in southwestern Gansu Province and the neighbouring areas in Sichuan (Fig. 8 ).

Remarks.

Laeocathaica amdoana was described from Pass Ho-Dshi-Gou bei Mu-gua-tshi (the exact locality could not be located on the map) near Wenxian at the border with Sichuan province, and characterised by a domed, brown, dorsal side with a relatively broad, sharp, distinctly bordering, white band of the keel. We did not find shells identical to typical Laeocathaica amdoana , but have found similar ones that can be identified as conspecific (Figs 3C, D , 4 ).

Laeocathaica distinguenda is represented in the Senckenberg Museum by several samples. Strangely, the lectotype is the most “untypical” among the lots labelled as L. distinguenda due to its pale caramel colour, the blurry border of light and dark stripes, and the rounded body whorl. Our samples from the vicinity of Wenxian (Fig. 5B-D ) are more similar to the paralectotypes of L. distinguenda .

Laeocathaica tropidorhaphe was described from the north (Tanchang and its vicinity), and is characterised by a large, keeled shell with a flat dorsal side and a thick brown spiral band. The northernmost populations we collected (samples 2016/87, 2016/88, 2016/89, 2016/91, 2016/95) agree with the types of L. tropidorhaphe in size, shape, and colouration, but their spire height is variable. However, some shells from much further south are also similar (i.e., samples 2016/74, 2016/75, see Fig. 4 ). The characteristic colouration of L. tropidorhaphe (brown dorsal side with slender white band on the keel) is also not unique to the northern L. tropidorhaphe populations, but can be found in more southern populations as well (compare Fig. 4a with Fig. 7 ).

Overall, there is a continuous variation across most of the historical and newly collected samples in terms of shape of dorsal side, shape of body whorl, size, colour, and sculpture (see Figs 3B, C ; 6B, C ). Laeocathaica amdoana and the lectotype of L. distinguenda (but not the paralectotypes!) seem to be slightly out of the morphological continuum, but not to a degree that a species-level distinction should be applied.

Colouration can be extremely variable even within a single population (see Fig. 4 ). Therefore, colour is of minor importance in species distinction within this group of Laeocathaica . It is a general trend that towards the southeastern part of the distribution of these “species”, the peripheral keel disappears and the body whorl becomes rounded.

Because of the aforementioned reasons, and until anatomical and DNA sequence data become available, we do not find the names L. distinguenda and L. tropidorhaphe meaningful, and so we provisionally synonymise them with L. amdoana . Table 1 summarises the key traits that are variable across and within newly collected populations.

Laeocathaica dangchangensis Chen & Zhang, 2004 is also a junior synonym of L. amdoana , because it shows the same characteristic conchological features (large shell size, acute, white keel, almost flat dorsal side) as L. tropidorhaphe . Moreover, its type locality (Shawan town, Dangshang county (34°0'N, 104°3'E), Gansu Province, China) is situated close to the known sites of L. tropidorhaphe , whose two closest populations are situated at ca. 31 km and 35 km from the type locality of L. dangchangensis .

According to the original description of Laeocathaica dangchangensis , the holotype has a shell width of 27.22 mm. However, the shell labelled as the holotype is 23 mm wide. Moreover, the number 6 is written on that specimen’s dorsal side, whereas "Holotype: Sp8" is written on the label. We have not found any shells bearing the number 8. Consequently, the shell labelled as the holotype is a paratype, and the real holotype is probably one of the specimens labelled as paratypes, or lost.

Figures

Figure 2. Distribution of the genus Laeocathaica in China A see Figs 8, 11, 17, 24 for details B see Fig. 14 for detail.

Figure 3. Laeocathaica amdoana A, B lectotype (SMF 8952) C, D 2016 / 74, specimen 2 E 2016 / 74, specimen 1. Scale bars: 10 mm. All photographs: B. Pall-Gergely.

Figure 4. Laeocathaica amdoana Moellendorff, 1899, 2016 / 75 (4 different shells of the same sample). Scale bar: 10 mm. All photographs: B. Pall-Gergely.

Figure 5. Laeocathaica amdoana Moellendorff, 1899 A lectotype of L. distinguenda (SMF 8959) B 2016 / 72, specimen 1 C 2016 / 72, specimen 2 D 2016 / 76. Scale bar: 10 mm. All photographs: B. Pall-Gergely.

Figure 6. Laeocathaica amdoana Moellendorff, 1899 A, B 2016 / 70 a C, D 2016 / 71 E 2016 / 69. Scale bars: 10 mm. All photographs: B. Pall-Gergely.

Figure 7. Laeocathaica amdoana Moellendorff, 1899 A lectotype of Laeocathaica tropidorhaphe (SMF 9074) B 2016 / 88 C 2016 / 89 D paratype of Laeocathaica dangchangensis Chen & Zhang, 2004 (labelled as holotype). Scale bar: 10 mm. Photographs: K. Meng (D) and B. Pall-Gergely (A-C).

Figure 8. Distribution of Laeocathaica amdoana Moellendorff, 1899 in China (detail from Fig. 2 A).