Research Article |
Barna Páll-Gergely‡, András Hunyadi§, Kaibaryer Meng|, Judit Fekete¶#
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Corresponding author: Barna Páll-Gergely ( pallgergely2@gmail.com ) Academic editor: Thierry Backeljau
© 2022 Barna Páll-Gergely, András Hunyadi, Kaibaryer Meng, Judit Fekete.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Páll-Gergely B, Hunyadi A, Meng K, Fekete J (2022) A review of the genus Laeocathaica Möllendorff, 1899 (Gastropoda, Pulmonata, Camaenidae). ZooKeys 1086: 33-76. https://doi.org/10.3897/zookeys.1086.77408
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Abstract
In this paper an overview of the Laeocathaica species is provided, and the intraspecific variability of several Laeocathaica species demonstrated on multiple shells. Laeocathaica hisanoi Páll-Gergely, sp. nov. and L. minwui Páll-Gergely, sp. nov. are described based on specimens found in museum collections. Five new synonyms are recognized: L. prionotropis albocincta Möllendorff, 1899 is a new synonym of L. prionotropis Möllendorff, 1899, L. stenochone Möllendorff, 1899 is a new synonym of Laeocathaica carinifera (H. Adams, 1870). Laeocathaica distinguenda Möllendorff, 1899, L. tropidorhaphe Möllendorff, 1899, and L. dangchangensis Chen & Zhang, 2004 are moved to the synonymy of Laeocathaica amdoana Möllendorff, 1899. Furthermore, photos of paratypes of Cathaica bizonalis Chen & Zhang, 2004 are published for the first time.
Keywords
Intraspecific variability, shell, systematics, taxonomy
Introduction
The genus Laeocathaica Möllendorff, 1899 consists of approximately 20 species, and inhabits west China. Most of the species assigned to this genus were reported from the southern part of Gansu Province and the neighbouring Sichuan. A single species, L. filippina (Heude, 1882) is known from Hubei Province, more than 500 km southeast from southern Gansu.
The monophyly of this genus is questionable for several reasons. First, Laeocathaica is defined by the sinistral shell coiling, whereas species with dextral shells of otherwise similar appearance (large, depressed shells with white base colour and brownish spiral bands) are included in other genera such as Cathaica Möllendorff, 1884, Bradybaena Beck, 1837, and Euhadra Pilsbry, 1890 (
In this paper we provide an overview of the genus Laeocathaica after consulting all available types and newly collected samples. We provide precise localities for most species, and photographs of multiple shells showing intraspecific variability for the first time in this genus.
Materials and methods
We counted the whorls of adult shells according to
Abbreviations
D shell diameter;
H shell height.
Depositories
HA Collection András Hunyadi (Budapest, Hungary);
MNHN Muséum National d’Histoire Naturelle (Paris, France);
NHM The Natural History Museum (London, UK);
NHMUK When citing lots deposited in the NHM;
PGB Collection Barna Páll-Gergely (Budapest, Hungary);
Taxonomy and systematics
Family Camaenidae Pilsbry, 1895
Cathaica
Type species
Helix pyrrhozona Philippi, 1845; OD.
Cathaica (Cathaica) bizonalis
Cathaica (Cathaica) bizonalis Chen & Zhang, 2004: 238 [Chinese description], fig. 219 (erroneous figure showing L. carinalis specimen); 439 [English description].
Type locality
陕西洛川县黑木沟. “Hemugou twon [sic!], Luochuan County (35°7'N, 109°04'E), Shaanxi Province”.
Remarks
According to the original description of Laeocathaica carinalis, that species is characterised by a sinistral, strongly depressed, keeled shell with a broad umbilicus. However, the provided photo (
Examining the specimens deposited in
- The shell labelled as the holotype of Cathaica bizonalis is a dextral, juvenile shell.
- A jar labelled as paratypes of
Cathaica bizonalis (
IZCAS TM 006579–006595) contains several probably conspecific juvenile specimens in ethanol. - Another jar labelled as paratypes of
Cathaica bizonalis (
IZCAS TM 097575–097587) contained 13 dry adult shells of a Laeocathaica carinalis (sinistral, strongly keeled species). - Among those shells (
IZCAS TM 097575–097587), specimenIZCAS TM 097578 is exactly the same shell as in fig. 219 ofChen and Zhang (2004) . - A jar labelled as paratypes of
Laeocathaica carinalis (
IZCAS TM 006612–006625) contained several adult specimens of L. carinalis in ethanol. - Another jar labelled as paratypes of
Laeocathaica carinalis (
IZCAS TM 097588–097602) contained several dry adult shells of Cathaica bizonalis (two of them figured in this work, see Fig.1 ). This clearly shows that shells of the above taxa have been confused even before the publication. As a result, not only were the different samples mixed up with the labels in the collection, but also in the original descriptions.
Based on
- Fig. 219 in
Chen and Zhang (2004) shows the holotype of Laeocathaica carinalis Chen & Zhang, 2004 instead of Cathaica bizonalis. - Fig. 334 in
Chen and Zhang (2004) shows an unidentifiable juvenile Laeocathaica species, and not Laeocathaica carinalis. - A holotype was designated in the original description of Cathaica bizonalis, so a lectotype cannot be chosen later. The holotype cannot be located because there is no photograph in the original description, and because the paratypes are of similar size (i.e., the given measurements of the holotype are insufficient to recognize the shell). So, we treat it as lost (perhaps in the lot of paratypes). Since all the paratypes belong to one single species, and are consistent with the original description, we know what the authors intended by the taxon, so designation of a neotype is not needed.
Laeocathaica
Laeocathaica
Möllendorff, 1899: 86;
Type species
Helix (Plectotropis) christinae H. Adams, 1870 (by original designation).
Laeocathaica amdoana
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. –
Laeocathaica distinguenda. –
Laeocathaica tropidorhaphe. –
Laeocathaica (Laeocathaica) amdoana. –
Laeocathaica (Laeocathaica) distinguenda. –
Laeocathaica (Laeocathaica) tropidorhaphe. –
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,
Museum material
New material (typical 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.
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.
Distribution
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
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.
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.
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
Colouration can be extremely variable even within a single population (see Fig.
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
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.
Laeocathaica carinalis
Laeocathaica carinalis Chen & Zhang, 2004: 341 [Chinese description], fig. 334 (erroneous! Shows a juvenile Laeocathaica shell belonging to another species); 444 [English description].
Type locality
“Town of Wenxian County, (33°0'N, 104°6'E), Gansu Province, China”.
Type material
The shell we examined (
New material
China • 4 shells; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 1800 m west of Jianshan towards Diaohuya, right side of road no. 212 (locality code: 2016/70b); 33°2.922'N, 104°50.840'E; 29 May 2016; A. Hunyadi leg.; HA (Fig.
Shell morphological traits of Laeocathaica amdoana Möllendorff, 1899 populations.
| Locality no. | D (in mm) | H (in mm) | Dorsal side | Body whorl |
|---|---|---|---|---|
| 2016/64 | 21.9–27.1 | 12–14.7 | domed | rounded to slightly keeled |
| 2016/65 | 25.5–26.2 | 13.3–14.7 | domed | rounded to slightly keeled |
| 2016/66 | 23.7–24.5 | 11.2–12.9 | domed | rounded |
| 2016/67 | 19.9–23.3 | 8.8–11 | slightly domed | rounded |
| 2016/68 | 24.3–25.4 | 10.4–10.7 | flat to slightly domed | rounded |
| 2016/69 | 27.4–30.7 | 12.2–14.1 | slightly domed | rounded |
| 2016/70a | 30–31.3 | 13.1–14.9 | slightly domed | rounded |
| 2016/71 | 28.5–29.7 | 13–13.6 | slightly domed | rounded |
| 2016/72 | 19.8–24.2 | 9.1–12.2 | domed | slightly keeled |
| 2016/73 | 21.8 | 10 | domed | slightly keeled |
| 2016/74 | 21.4–25.4 | 9–10.7 | flat to slightly domed | strongly to slightly keeled |
| 2016/75 | 25.1–27.4 | 9.8–12.5 | slightly domed to domed | strongly to slightly keeled |
| 2016/76 | 20.8 | 9 | flat | strongly keeled |
| 2016/77 | 20.8–21.6 | 10.5–11.5 | domed | slightly keeled |
| 2016/78 | 22.7 | 11.6 | slightly domed | slightly keeled |
| 2016/79 | 20.7–21.7 | 10–10.3 | slightly domed to domed | strongly to slightly keeled |
| 2016/82 | 20.6–23 | 9.8–11.6 | slightly domed to domed | rounded to slightly keeled |
| 2016/83 | 27.7–28.3 | 13–13.7 | slightly domed | rounded |
| 2016/87 | 23–26.9 | 8.1–11.4 | flat to slightly domed | strongly keeled |
| 2016/88 | 26.6–27.7 | 10.9–11.3 | flat to slightly domed | strongly keeled |
| 2016/89 | 27.9–30.6 | 11.3–12.1 | flat to slightly domed | strongly keeled |
| 2016/91 | 27.5–30 | 10.4–11.5 | flat | strongly keeled |
| 2016/95 | 23.6–27.4 | 9–10.1 | flat | strongly keeled |
Description
Shell sinistral, depressed, strongly keeled, dorsal side with flat, scalariform whorls; ventral side widely conical; dorsal side chocolate brown, ornamented with a white keel on all whorls; ventral side primarily white, below the white keel there is a chocolate brown belt, white part ornamented with greyish radial stripes that sometimes reach the umbilicus, but sometimes thin and stop before umbilicus; umbilicus inside with a chocolate-brown and a white belt; entire shell consists of 7.25–7.75 whorls; protoconch consists of 1.5–1.75 whorls, brownish, seemingly smooth, extremely finely granulose, rather matte, slightly protruding above first whorls of teleoconch; white keels of every whorl slightly elevated from dorsal surface, but dorsal surface flat with usually the last one being scalariform; dorsal side with fine, irregular wrinkles and between the main wrinkles there are very fine radial lines; ventral surface with less prominent wrinkles; umbilicus rather narrow, funnel-shaped, shows all whorls; periumbilical keel absent; aperture oblique to shell axis, semilunar, with pointed incision at the keel; peristome expanded and slightly thickened, but not reflexed; palatal swelling whitish, with a low, blunt basal tooth; parietal callus practically absent, in some old specimens with translucent calcareous layer.
Measurements (in mm): D: 18.6–22.9; H: 6.8–9.8 (n = 13, newly collected shells from multiple samples).
Differential diagnosis
The most similar species is L. pewzowi, which is smaller, paler in colour, has a wider umbilicus, a more domed (and not scalariform) dorsal side, stronger radial sculpture, and a more oblique aperture with a more pointed basal tooth. Furthermore, there is a second broken belt between the main belt and the umbilicus in L. pewzowi, which is not present in any specimens of this species. Laeocathaica potanini has a more scalariform, uniformly light brown shell, and the basal tooth (when present) is situated closer to the columella than in L. carinalis. Laeocathaica amdoana is also similar in colouration, but it is larger, has a blunter keel, a weaker sculpture, and its whorls are never scalariform.
Distribution
Most precise locality data are from the rocky area along the Baishui River, whereas one sample was collected on the bank of the Yangtang River (Fig.
Remarks
We here provide a redescription, an updated differential diagnosis, and notes on the differences between different populations (Table
Laeocathaica carinifera
Helix (Plectotropis) christinae var. carinifera H. Adams, 1870: 377.
Helix subsimilis Deshayes, 1874: 10, pl. 2, figs 28–29.
Helix christinae. –
Helix subsimilis. –
Helix christinae var. carinifera. –
Laeocathaica subsimilis. –
Laeocathaica stenochone
Laeocathaica subsimilis subsimilis. –
Laeocathaica stenochone. –
Laeocathaica (Laeocathaica) stenochone. –
Laeocathaica (Laeocathaica) subsimilis. –
Laeocathaica subsimilis. –
Laeocathaica stenochone. –
Laeocathaica subsimilis subsimilis. –
Type material
China, Woushan, coll. Swinhoe, NHMUK 1870.7.16.7 (3 shells, probably syntypes of Helix christinae var. carinifera, labelled as “christinae var”) (Fig.
Shell morphological traits of Laeocathaica carinalis Chen & Zhang, 2004 populations.
| Locality no. | Shell diameter | Belt below keel | White belt on keel | Dorsal side | Denticle |
|---|---|---|---|---|---|
| 2016/64 | 20.2 | medium | thin | flat/not scalariform | strong |
| 2016/70b | 20.3–21.5 | thin | thick | domed/scalariform | only low thickening |
| 2016/79 | 19.1–20.3 | thick | moderate | flat/scalariform | strong |
| 2016/81 | 19.8–20.4 | thick | moderate | flat/scalariform | 1 out of 5 shells |
| 2016/82 | 18.6–19.8 | medium | thin | flat/scalariform | present |
| 2016/83 | 21.2–22.9 | thick | moderate | moderately domed/slightly scalariform | only low thickening |
Museum material
China, Yangtze-Tal, coll. Jetschin ex coll. Beddome,
New material
China • 3 shells; Gansu, Longnan Shi, Wenxian, Bikou Zhen, above the hydroelectric power plant, northern side of Bailong He (locality code: 2016/63); 32°45.966'N, 105°13.005'E; 28 May 2016; A. Hunyadi leg.; HA • 1 shell; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 1800 m west of Jianshan towards Diaohuya, right side of road no. 212 (locality code: 2016/70b); 33°2.922'N, 104°50.840'E; 29 May 2016; A. Hunyadi leg.; HA • 3 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, 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.
Distribution
The original description Helix christinae var. carinifera was published together with that of the nominotypical form, without any specification of a type locality. However, on one of the boxes of var. carinifera in the NHM, the locality Woushan (probably Wushan, Chongqing at 31°5'N, 109°53'E) was mentioned. The type locality of Helix subsimilis is Moupin (Baoxing, at 30°22'N, 102°49'E) in Tibet. Furthermore,
Remarks
The syntypes of Helix christinae var. carinifera are identical with the types of Helix subsimilis, and thus, the latter is a junior synonym of the former. Although both
Laeocathaica christinae
Helix (Plectotropis) christinae H. Adams, 1870: 377, pl. 27, figs 4, 4a.
Helix christinae. –
Laeocathaica christinae. –
Laeocathaica (Laeocathaica) christinae christinae. – Zilch,1968: 173.
Laeocathaica christinae. –
Type material
China, coll. Swinhoe, NHMUK 1870.7.16.6 (3 shells, probably syntypes, Fig.
Museum material
Asia Centrale, MNHN-IM-2014-7933/3 • Asia Centrale, MNHN-IM-2014-7934/7 • Hupé, China, coll. Staadt, 1969, MNHN-IM-2014-7929/2 + 1 L. carinifera shell (mixed sample, erroneous locality for L. carinifera) • Moupin, leg. Abbé David, coll. Deshayes, 1872, MNHN-IM-2014-7945/2 (probably erroneous locality).
New material
China • 10 shells; Hubei, Enshi Tujiazu Miaozu Zizhizhou, Badong Xian, east of Badong, Bashan Senlin Gongyuan, (next to Xinlingzhen) (locality code: 2010/29); 31°01.472'N, 110°25.284'E; 225 m a.s.l.; 03 November 2010; A. Hunyadi leg.; HA (Fig.
Distribution
The type localities and the newly collected sample suggest that this species lives more upstream in the Yangtze valley than L. filippina. The samples from Moupin (Baoxing County, Sichuan) are probably erroneous. We indicated only the newly collected sample on the map (Fig.
Remarks
We found three samples in the NHM. One of them, containing two shells, was labelled Helix christinae. The other two lots, labelled “Helix christinae var.”, contained three shells each. The latter two samples are probably syntypes of Helix christinae var. carinifera, described in the same publication (
Laeocathaica dejeana
Helix dejeana Heude, 1882: 21, pl. 20, fig. 17.
Cathaica (Campylocathaica) dejeana. –
Laeocathaica dejeana. –
Type material
Museum material
China: Da-tshien-lu am Ya-lung, coll. Möllendorff ex coll. Potanin 380,
Laeocathaica dityla
Laeocathaica dityla Möllendorff, 1899: 99–100, pl. 6, fig. 8.
Laeocathaica dityla. –
Laeocathaica (Laeocathaica) dityla. –
Laeocathaica dityla. –
Type material
SO-Gansu, zw. Li-tshia-pu u. Hsi-gu-tsheng, coll. Möllendorff ex coll. Potanin 776,
New material
China • 4 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.
Laeocathaica dolani
Cathaica (Laeocathaica) dolani Pilsbry, 1934: 16, pl. 3, figs 4, 4a–c.
Laeocathaica dolani. –
Type material
China, Szechuan, Romichengu, Brooke Dolan, W. China Expedition 1931,
Laeocathaica filippina
Helix filippina Heude, 1882: 23, pl. 20, fig. 19.
Helix (Plectopylis) subchristinae Ancey, 1882: 44.
Helix subsimilis var. filippina. –
Laeocathaica filippina. –
Laeocathaica subsimilis filippina. –
Laeocathaica (Laeocathaica) christinae filippina. –
Type material
Patong, Heude type coll.,
Museum material
China, W-Hupei, coll. K. Hashagen,
New material
China • 7 shells; Hubei, Yichang Shi, Changyang Tujiazu Zizhixian, Qingjiang Hualang Fengjingqu, Geheyan Shuiku, Wuluozhougli Shan (locality code: 2010/25); 30°25.805'N, 110°59.254'E; 260 m a.s.l.; 31 October 2010; A. Hunyadi leg.; HA (Fig.
Distribution
The newly collected samples, and the several museum samples from Changyang (at 30°28'N, 111°12'E), suggest that this species lives downstream along the river Yangtze compared to L. christinae (Fig.
Remarks
We did not examine the types of Helix subchristinae Ancey, 1882, and treat it as a synonym of L. christinae following
Laeocathaica hisanoi , sp. nov.
Type material
Holotype
China • S Kansu, China, coll. S. Hisano, 24.05.204,
Diagnosis
A small Laeocathaica species with many (8.5) whorls, conical dorsal side, rounded body whorl and single, small basal tooth that is situated close to the columella.
Description
Shell sinistral, depressed, dorsal side conical with protruding apex, body whorl shouldered; colour chalk white with a single brownish belt below shoulder; entire shell consists of 8.5 whorls, protoconch consists of 1.75–2 whorls, very finely granulose, conspicuously elevated compared to first teleoconch whorls; teleoconch with fine, irregular growth lines, without any notable sculpture, although both examined shells were corroded; last quarter whorl with slight subsutural furrow; aperture semilunar, very strongly oblique to shell axis; peristome sharp, very slightly expanded dorsally, with thickening situated behind peristome edge; basal tooth blunt, elongated, situated ca. at the middle of basal peristome; parietal callus inconspicuous, appears only as thick calcareous layer; umbilicus open, narrow, shows all whorl, with the last half of body whorl extremely widened, resulting in a “9”-shape.
Measurements (in mm): D: 11.5–11.6; H: 5.2–5.3 (n = 2).
Differential diagnosis
The most similar species is L. polytyla, which is usually larger, has one whorl more, has a more elevated spire with a domed dorsal side, a rounded body whorl, and a comparatively smaller basal tooth situated closer to the columella.
Etymology
This new species is named after S. Hisano, who collected the type material.
Distribution
This new species is known from a single museum sample only, consisting of two shells.
Laeocathaica leucorhaphe
Laeocathaica leucorhaphe Möllendorff, 1899: 95–96, pl. 6, fig. 2.
Laeocathaica leucorhaphe. –
Laeocathaica (Laeocathaica) leucorhaphe. –
Laeocathaica leucorhaphe. –
Types examined
N-Sytchuan: am Tung-ho, coll. Möllendorff ex coll. Potanin 312b,
Laeocathaica minwui , sp. nov.
Type material
Holotype
China • China, O. Sy-tshuan, coll. C.R. Boettger ex coll. Möllendorff ex coll. L. Fuchs,
Diagnosis
A rather large Laeocathaica species with a sharp keel, a domed dorsal side, an oval aperture and a marmorated ventral side.
Description
Shell sinistral, depressed, with domed dorsal side, keel strong, situated in the middle of body whorl, whitish; dorsal side latte-coloured, with darker and paler areas alternating as the shell grows; ventral side with white and pale brownish (latte) spiral bands forming a marmorated colour pattern; inner side of umbilicus with brownish spiral band; protoconch light brownish, ca. 1.5 whorls, finely granulose, slightly protrudes above first whorls of teleoconch; entire shell consists of six whorls; dorsal side finely ribbed, ventral side smoother, only with growth lines; umbilicus ca. one third of shell width; shows all whorls; periumbilical keel absent; aperture oblique to shell axis, oval, without incision at the position of keel; peristome white, expanded and slightly thickened, but not reflexed (only in direction of umbilicus); parietal callus practically absent, only with some additional translucent calcareous layer.
Measurements (in mm): D = 23.1, H = 9.1 (holotype).
Differential diagnosis
Laeocathaica minwui sp. nov. has been confused with L. christinae in museum collections, probably due to the lack of examination of the types of L. christinae. However, L. christinae has a flatter dorsal side, a more upper-situated peripheral keel, a darker brown (instead of latte) colour, a more uniformly white ventral side with a brown spiral band inside the umbilicus, and brownish spots. In contrast, in the new species the ventral side is characterised by a marmorated (marbled-like) pattern resulted by the fusing of whitish and pale brown spiral bands. Laeocathaica filippina has a notched aperture at the position of the peripheral keel, a more brownish colour, and a less marmorated ventral side. See also Table
Differences between L. carinifera, L. christinae, L. filippina, and L. minwui sp. nov.
| Species | Dorsal surface | Keel on body whorl | Aperture (palatal part) | Colour of dorsal side | Ventral side |
|---|---|---|---|---|---|
| L. carinifera | domed | acute, in the middle of body whorl | rounded | lighter and darker brownish patches alternate rather abruptly resulting in a mosaic-like structure | umbilicus narrow, colour pale, or light and darker stripes alternate |
| L. christinae | nearly flat/ slightly domed | blunt to acute, upper part of body whorl | rounded | same as in carinifera, just brown colour darker and there is usually a brown and a white spiral band | umbilicus wider, mostly white with slender darker radial stripes and dark dots |
| L. filippina | domed | acute, upper part of body whorl | notched | paler than carinifera and christinae, light and brown patches alternate smoothly | umbilicus wider, similar to minwui sp. nov., but paler and less nicely marmorated |
| L. minwui sp. nov. | domed | acute, upper part of body whorl | rounded | mostly cream/latte-coloured, lighter and darker patches alternate smoothly | umbilicus wider, latte and white spiral bands form a marmorated pattern |
Etymology
This new species is dedicated to and named after Dr. Min Wu, the leading expert of Chinese Camaenidae.
Distribution
This new species is only known from historical samples from the Yangtze valley. Other samples labelled as being collected from Sichuan are not precise enough to understand their geographic origin.
Laeocathaica odophora
Laeocathaica odophora Möllendorff, 1899: 97–98, pl. 6, fig. 6.
Laeocathaica odophora. –
Laeocathaica (Laeocathaica) odophora. –
Laeocathaica odophora. –
Type material
S-Gansu, Dshie-dshou, coll. Möllendorff ex coll. Potanin 254,
New material
China • 1 photographed shell; Gansu, Longnan Shi, Wenxian, Jianshan Xiang, 1800 m west of Jianshan towards Diaohuya, right side of road no. 212 (locality code: 2016/70b); 33°2.922'N, 104°50.840'E; 29 May 2016; A. Hunyadi leg.; HA (Fig.
Description
Shell sinistral, depressed, strongly keeled, dorsal side domed, ventral side conical; shell colour basically brownish, greyish, latte-coloured, with whitish stripes; as a result dorsal surface mosaic-like, ventral side striped; keel always white, there is always a brownish belt just below the keel, periumbilical keel always white; entire shell consists of 9–9.5 whorls; protoconch consists of 1.5 whorls, brownish, seemingly smooth, extremely finely granulose, rather matte; white keels of every whorl slightly elevated from dorsal surface, but dorsal surface almost continuous, suture practically absent; dorsal side with fine, irregular wrinkles (most wrinkles stand alone, but some of them unite to each other); ventral surface with less prominent wrinkles; umbilicus rather narrow, regular funnel-shaped, shows all whorls; periumbilical keel blunt; aperture semilunar, peristome very slightly expanded, but not reflexed or thickened; palatal swelling whitish, with two prominent denticles, situated in some distance from peristome; parietal wall with some whitish thickening in adult shells. Juveniles reverse conical in shape; several apertural barriers are built during lifetime; palatal swelling of juveniles appears as a continuous ridge, although the two denticles recognisable.
Distribution
Known from a few localities in southern Gansu Province (Fig.
Remarks
The single juvenile shell of sample 2016/64 has a narrower, blunter umbilical keel than the holotype, and it is possible that it belongs to another species. However, the juvenile shell of sample 2016/68 is identical with the holotype.
Laeocathaica pewzowi
Laeocathaica pewzowi Möllendorff, 1899: 98, pl. 6, figs 4, 4a.
Laeocathaica pewzowi. –
Laeocathaica (Laeocathaica) pewzowi. –
Laeocathaica pewzowi. –
Laeocathaica pewzowi. –
Type material
S-Gansu, Wen-hsien, coll. Möllendorff ex coll. Potanin 248, 661, 793,
Museum material
Nung-dan b. Wen-Hsien, coll. Möllendorff,
Laeocathaica phaeomphala
Laeocathaica phaeomphala Möllendorff, 1899: 96, pl. 6, fig. 3.
Laeocathaica phaeomphala. –
Laeocathaica (Laeocathaica) phaeomphala. –
Laeocathaica phaeomphala. –
Type material
S-Gansu, Wenhsien, coll. Möllendorff ex coll. Potanin 51b, 72, 741,
New material
China • 6 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.
Laeocathaica polytyla
Laeocathaica polytyla Möllendorff, 1899: 98–99, pl. 6, fig. 7.
Laeocathaica polytyla. –
Laeocathaica (Laeocathaica) polytyla. –
Laeocathaica polytyla. – Chen & Zhang, 2004: 331, fig. 322.
Laeocathaica polytyla. –
Type material
New material
China • 4 shells; 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 (Fig.
Distribution
Known from several precise localities in southern Gansu Province (Fig.
Laeocathaica potanini
Laeocathaica potanini Möllendorff, 1899: 96–97, pl. 6, fig. 5.
Laeocathaica potanini. –
Laeocathaica (Laeocathaica) potanini. –
Laeocathaica potanini. –
Type material
Gansu: Wenhsien, coll. Möllendorff ex coll. Potanin 251, 587, 734,
New material
China • 6 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 • 5 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.
Remarks
The examined shells are identical to the types.
Laeocathaica prionotropis
Laeocathaica prionotropis Möllendorff, 1899: 94–95, pl. 6, figs 1, 1a.
Laeocathaica prionotropis subsp. albocincta Möllendorff, 1899: 95. new synonym
Laeocathaica prionotropis prionotropis. –
Laeocathaica prionotropis albocincta. –
Laeocathaica (Laeocathaica) prionotropis prionotropis. –
Laeocathaica (Laeocathaica) prionotropis albocincta. –
Laeocathaica prionotropis. –
Type material
Zw. Yü-lin-guan u. Wen-hsien, coll. Möllendorff ex coll. Potanin 520a, 908a,
New material
China • 1 shell; Gansu, Longnan Shi, Wenxian, Bikou Zhen, above the hydroelectric power plant, northern side of Bailong He (locality code: 2016/63); 32°45.966'N, 105°13.005'E; 28 May 2016; A. Hunyadi leg.; HA (Fig.
Distribution
Known from two sites in Southern Gansu Province (Fig.
Remarks
Laeocathaica prionotropis subsp. albocincta agrees in size and shell shape with the nominotypical form, and therefore it is here synonymised with Laeocathaica prionotropis.
Discussion
Although we list and publish photographs of all Laeocathaica species in this work, the taxonomy of this group is still far from being solved. Following previous authors, we classify only sinistral species in Laeocathaica. However, it is very probable that the coiling direction has changed multiple times during the evolution of Bradybaeninae inhabiting the arid regions of central China. Furthermore, sinistral species such as Bradybaena micromphala (Möllendorff, 1899) and B. eris (Möllendorff, 1899) also inhabit southern Gansu, and are similar to Laeocathaica species in most traits except for the narrow umbilicus. Future investigations will probably reveal that the latter two species (and maybe some other similar ones from the region) are relatives of Laeocathaica rather than Bradybaena.
One of the main outcomes of the present paper is the clarification of some names that have been incorrectly used in the literature and in museum collections because the types were not examined. One such case is Helix christinae var. carinifera H. Adams, 1870, which resulted in being a senior synonym of Laeocathaica subsimilis (Deshayes, 1874) after examination of both type species. The other case is that of L. minwui sp. nov., which was called L. christinae (H. Adams, 1870) in museum collections, because the types of the “real” L. christinae had not been examined.
The other important outcome of the present study is the recognition of continuous dorsal surface variability from flat to domed, the strongly keeled to rounded body whorl, the colouration, and shell size (D: 19–32 mm) across the taxa Laeocathaica amdoana, L. distinguenda, L. tropidorhaphe, and L. dangchangensis.
| L. amdoana | L. carinalis | L. carinifera | L. odophora | L. phaeomphala | |
|---|---|---|---|---|---|
| L. carinalis | 2016/64, 2016/79, 2016/82, 2016/83 | ||||
| L. carinifera | 2016/67, 2016/68, 2016/70a | 2016/70b | |||
| L. dityla | 2016/88, 2016/95 | 2016/86 | |||
| L. odophora | 2016/64, 2016/68, 2016/69 | 2016/64, 2016/70b | 2016/68, 2016/70b | ||
| L. phaeomphala | 2016/64 | 2016/64 | 2016/64 | ||
| L. polytyla | 2016/65, 2016/72, 2016/73,2016/74, 2016/76, 2016/77, 2016/78, 2016/79 | 2016/79 | 2016/78 | ||
| L. potanini | 2016/65, 2016/66 | ||||
| L. prionotropis | 2016/64 | 2016/64 | 2016/63 | 2016/64 | 2016/64 |
Table
Acknowledgements
We are grateful to Zhe-Yu Chen for his help in locating historical Chinese localities on the map, to Sigrid Hof and Ronald Janssen (
This study was supported by the Bolyai Research Scholarship of the Hungarian Academy of Sciences, the Hungarian Research Fund (OTKA FK 135262) and grants from the SYNTHESYS Project (GB-TAF-2523) to B.P.-G. We are indebted to The Biodiversity Heritage Library for the multitude of rare literature made available to us (www.biodiversitylibrary.org).
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