Corresponding author: Min Wu (minwu1969@aliyun.com)
Academic editor: Eike Neubert
Central China harbors the native dart-sac-bearing camaenids
Wu M, Shen W, Chen Z-G (2023) Land snail diversity in central China: revision of
This paper examines the shell morphology and genital anatomy of
Animals in most cases were relaxed by drowning in water before being transferred to 70% ethanol for fixation, which was replaced with ethanol of the same concentration after 3 days. To observe the animals with evaginated dart sac apparatus, before fixation the animal was drowned in water with 5% ethanol, 10% ethanol and 15% ethanol, in each solution for ~ 60 min. Photographs for illustrations (by MW) and geometric morphometric analyses (by WS) were taken using a Canon camera. The shells were measured with digital vernier calipers to the nearest 0.1 mm. Whorls were counted following
Distribution maps were created using ArcGIS Desktop 10.8 under GCS_WGS_1984 coordinate system. In addition, mapping was carried out to map both the distribution limit and the distribution pattern of
Geometric morphometric methods (
The material used in the
Whole genomic DNA was extracted from a piece of the foot muscle using the Animal Genome Quick Extraction Kit (
Chromatographs and sequences were first studied and compiled in sequencer 4.5. DNA sequences of both genes were aligned by T-Coffee with standard parameters (
The generic classification of species adopted in this study mainly follows MolluscaBase (2021a–p; 2022a–c) (Table
The most complicated structure in the genital system of dart-sac-bearing camaenid species (here, we use “dart-sac-bearing camaenids” to replace the former subfamily
The so-called
The proximal accessory sac, whose opening leads to the outside of the body (if is not blind) (Fig.
Distribution map of
In
Some character states of the terminal genitalia in
Characters\Species | Vagina elongated above dart sac (+) or not (–) | Number of mucous glands | Branching of mucous gland duct complicated (+) or simple (–) | Blades of apical dart | Number of |
Number of |
||
---|---|---|---|---|---|---|---|---|
+ | 8 | + | 2 | 2 (tiny) | + | 2 | – | |
– | 4 | – | / | 2 | + | 2 | + | |
– | 6–7 | +/– | / | 2 | – | 2 | + | |
– | 4–5 | + | 2 | 2 | + | 2 | – | |
– | 4–12 | +/– | 2 | 2 | + | 2 | – | |
– | 2–3 | – | ? | 2 | + | 2 | + | |
– | 2–3 | + | 4 | 1 | – | 1 | / | |
– | 5–7 | + | ? | 2 | + | 2 | + | |
– | 5 | – | 2 | 2 | – | 2 | + | |
– | 4 | – | / | 2 | + | 2 | + | |
+ | 8 | – | / | 0 | / | / | / | |
– | 5 | – | 2 | 0 | / | / | / | |
– | 2–3 | + | 4 | 2 | + | 2 | + | |
– | 4–8 | – | 4 | 2 | + | 2 | + | |
– | 5 | + | 2 | 2 | + | 2 | + | |
+ | 7–10 | + | 4 | 0 | / | / | / | |
– | 4–5 | – | / | 0 | / | / | / | |
– | 2 | + | 2 | 2 | + | 2 | + | |
/ | / | / | / | / | / | / | / | |
+(somewhat) | 6 | – | 0 | 2 | + | 2 | – | |
– | 4 | – | 2 | 2 | + | 2 | + | |
+ | 6–8 | – | / | 2 | + | 2 | + |
Terminal genitalia in the evaginated pouch in
Unlike the structurally closed proximal accessory sac observed only in
The accessory sac in some
The penis of
Shell sinistral, strongly depressed to broadly conic, moderately solid, of 5–10.5 almost flat whorls. Last whorl abruptly descending in front, angulated to strongly keeled; rarely rounded. Coloration consists of whitish, corneous, or chestnut background and mostly with one or two dark sub-peripheral bands; besides, usually there are several fulvous, diffuse radial streaks. Protoconch usually with fine radial threads and/or fine granules that each is formed by a low hump deposited in a shallow socket. Sculpture of teleoconch whorls varying from fine, silky radial striation to rather strong ribbing; on basal surface below keel or angle this sculpture becomes much weaker. Aperture rounded to peach-shaped, oblique, margins usually more or less reflexed. Within aperture a ring-like thickening present. Apertural teeth absent or with one tuberculiform basal tooth and sometimes with another one on palatal wall. During postembryogenesis several sets of teeth, different from those developed at adult stage in shape and/or number, present and remained to adult stage. Umbilicus moderate to very wide, ratio of umbilicus diameter to maximum diameter 0.21–0.50. Height 3.5–14 mm, maximum diameter 10.0–29.5 mm.
On left and right side of mantle edge, no leaf-shaped appendage present. Head wart between ommatophores present, weak, or developed. Jaw arcuate, with 3–8 more or less projecting ribs.
Slender vas deferens entering epiphallus at penial retractor muscle insertion. Penis generally clavate, rarely subcylindrical. Penis internally divided into three regions: the proximal part with narrow or thick longitudinal pilasters, among which one pair or two pairs of adjacent pilasters fuse into one Y-shaped fork or two Y-shaped forks (not in
China: S Gansu, W Hubei, W Shaanxi, Chongqing, Sichuan.
HBUMM05600, numerous
Gansu: Wudu, Wenxian (type locality).
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.
Eversible head wart lowly present. Jaw arcuate, with five projecting ribs.
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.
By shell sculpture, including that on the protoconch, this species (Fig.
brownish/chestnut coloration. In
peripheral angulation. The level of angulation, in order is
the structure of umbilicus. In both
The phylogeny based on ITS2 and 16S suggests that
Genital anatomy of
Genital anatomy of
None.
Sichuan.
HBUMM08300, 3
Genital anatomy of
In juveniles, regularly arranged fine threads and elongate granules are present on the protoconch, where in adults such sculpture is difficult to be observed because of erosion. Spiral grooves are present after the fourth whorl or only on apical body whorl, but are absent in umbilical side. The teleoconch has distinct and regularly arranged ribs, among which there is no fine threads.
Gansu: Wenxian (type locality).
Eversible head wart not prominent, perhaps due to specimen of not full maturity. Jaw arcuate, with five projecting ribs.
Penial sheath moderately long, covering ~ 1/4 of penis. Penis slightly expanded distally. Accompanied with two or three lower pilasters, at proximal 1/2 of penis two thin penial pilasters fusing into one Y-shaped fork. Fine pilasters of distal 1/2 penis weaving into delicate net. Near epiphallic opening numerous fine pilasters merging into ~ 4 short but thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 2/3 length of penis. Information of love dart unknown. Accessory sac small. Mucous glands ~ 4, each complicatedly branched. Proximal accessory sacs two, symmetrical on dart sac, dorsally separated and ventrally touching, internally smooth (without pilasters), each with an opening leading to dart chamber near dart chamber opening. Bursa copulatrix duct of even diameter.
HBUMM00006, 4
Genital anatomy of
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].
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.
Eversible head wart weak. Jaw arcuate, with four or five projecting ribs.
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.
See
HBUMM01251a, many
Chongqing: Fengjie (type locality: Fungsiang gorge (= Fengxiangxia)); Hubei: Badong.
Tiny granules (~ 5 – ~ 50 µm) are densely and radially arranged throughout the protoconch. The erosion or weathering of sculpture on adult shell is not observed. Spiral grooves are absent throughout the shell. On teleoconch the growth lines are usually indistinct.
Eversible head wart weakly present. Jaw arcuate, with four projecting ribs.
Penial sheath covering ~ 1/3 of penis. Penis tubular and equally thick. Inside penis, 5–6 narrow or thick pilasters present, Y-shaped fork formed by adjacent pilasters absent. Fine pilasters on distal end of penis merging into seven or eight thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac slightly shorter than penis. Love dart ~ 5 mm long, apically 2-bladed. Accessory sac small but externally distinct, internally empty but spatially narrow, inserting into dart sac at middle part, opening to distal dart chamber. Mucous glands four (HBUMM04204-spec.14, spec.16) or five (HBUMM04204-spec.13, HBUMM01251a-spec.1), each complicatedly branched. Proximal accessory sacs two, symmetrical or the left one slightly larger (HBUMM04204-spec.14), dorsally and ventrally separated, each with a distal opening leading to dart chamber near its opening. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.
Genital anatomy of
This species is not known from NW Sichuan, so the specimens distributed in “Wentschun” [Wenchuan 汶川] identified as
Sichuan: western region (type locality) including Kangding [康定].
Protoconch has small granules. Spiral grooves are absent throughout the shell. Shell surface has tiny scales.
Penis as long as dart sac, distally expanded. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Love dart ~ 2 mm long, apically 2-bladed. Accessory sac small but distinguishable from outside, inserting into dart sac at middle part, opening to distal dart chamber. Mucous glands two, each complicatedly branched. Bursa copulatrix duct of equal thickness, as long as dart sac. Bursa copulatrix ovate (
HBUMM00674, numerous
Gansu: Wenxian (type locality), Wudu (type locality), Zhouquxian (type locality); Sichuan: Jiuzhaigouxian (type locality).
The protoconch has the widespread granules each of which is ~ 20 – ~ 40 µm long and looks like a hump embedded in a shallow socket. All granules are present on densely arranged fine radial threads. Spiral grooves are shallowly present. The growth lines on teleoconch are more or less distinct but are irregularly arranged.
Eversible head wart present. Jaw arcuate, with 4–7 projecting ribs.
Penial sheath moderately long, covering ~ 1/6–1/4 of penis. Penis slightly expanded distally. Inside penis, two penial pilasters forming one Y-shaped fork at proximal 1/3 penis (only indistinct in HBUMM05417); a rather thick pilaster that is made up of numerous thread-like longitudinal folds occupying proximal 1/3–2/3 of penis. Pilasters of median to distal penis weaving into a delicate net that bears regularly arranged diamond-shaped papillae, which are usually lost perhaps because of bad specimen condition. Distally inside penis, fine pilasters merging into 6–9 short and more or less 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 ~ 1/4 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 9 mm long, apically 2-bladed. Accessory sac small but distinguishable from outside, internally solid, inserting into dart sac at middle part, opening to distal dart chamber. Mucous glands 4 (HBUMM08429, each complicatedly branched) – 12 (HBUMM05479, simply branched). Proximal accessory sacs two, symmetrical or the right one slightly larger, separated dorsally and ventrally, with a few internal pilasters, each with an opening leading to dart chamber near opening of dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate.
The sculpture on protoconch and teleoconch of this species (Fig.
For more comments, see
HBUMM00532, 2 juvs; Xinglongcun [兴隆村],Zhongzhaixiang [中寨乡], Wenxian, Gansu Province, near point (
Genital anatomy of
Gansu: Wenxian, Diebuxian, Dangchangxian, Zhouquxian (type locality).
Tiny, low and sparsely arranged granules (~ 10 µm) on the smooth protoconch are present but difficult to be observed because of erosion or weathering. Spiral grooves are absent throughout the shell. On teleoconch the growth lines are not observed except on the part just following the protoconch.
Eversible head wart present but not prominent. Jaw arcuate, with four or five projecting ribs.
Penial sheath rather short, covering ~ 1/8 of penis. Penis tubular and equally thick. Inside penis, two penial internal pilasters forming one Y-shaped fork at proximal 1/2 of penis, accompanied with another two thicker pilasters. Distal 1/2 of penis occupied by numerous isolated minute diamond-shaped papillae. 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 dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 4 mm long, apically 2-bladed and then rounded. Accessory sac small but distinguishable from outside, internally empty but spatially narrow, inserting into dart sac medially, opening to distal dart chamber. Mucous glands two (HBUMM00698-spec.2, spec.3) or three (HBUMM00698-spec.1), each a single tube. Proximal accessory sacs two, dorsally and ventrally touching, without internal pilasters, each with an opening leading to dart chamber near its opening. The right proximal accessory sac somewhat larger and with thicker wall than the left one. Bursa copulatrix duct basally slightly expanded. Bursa copulatrix ovate.
This species has a very unique shell, but the anatomy of the terminal genitalia coincides with those of the congeners.
HBUMM00069, 6
Genital anatomy of
Sichuan: Kangding, Danbaxian (type locality: Rumichangu).
The shell is covered with radially arranged low scales of variable length (~ 25 – ~ 300 µm long) everywhere. The size (esp. length) of scales increase from protoconch to body whorl. Spiral grooves are absent throughout the shell.
Eversible head wart present but not prominent. Jaw arcuate, with 3–6 projecting ribs.
Penial sheath covering ~ 1/4–1/3 of penis. Penis short, thick, distally expanded. Inside penis, two thick and prominent pilasters fusing into one Y-shaped fork at proximal 1/3–1/2 penis, accompanied with ~ 6 narrower or equally thick pilasters nearby. Pilasters on distal end of penis not merging into thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac approximately as long as penis. Love dart ~ 5 mm long, apically with cross-section rhombic. Accessory sac small, internally empty, inserting into dart sac at middle part, opening to distal dart chamber. Mucous glands two (HBUMM00069-spec.1–3), each complicatedly branched. Proximal accessory sac one, large, on right side of dart sac, with an opening leading to dart chamber near its opening. Bursa copulatrix duct basally slightly expanded. Bursa copulatrix elongate, ovate.
In
HBUMM01256b, Fuhusi, Emei [峨嵋], Sichuan; sandstone, humid; 2003-VII-26, coll. Wu, M.; DNA voucher HBUMM01256b. HBUMM04166, numerous
Genital anatomy of
Hubei: Badong (type locality), Yichang (Changyang); Sichuan (Emei); Chongqing: Fengjie.
Sculpture on the protoconch was present but indistinct because of erosion or weathering. Spiral grooves are absent throughout the shell.
Eversible head wart low. Jaw arcuate, with four or five projecting ribs.
Penial sheath moderately long, covering ~ 1/4–1/3 of penis. Penis slightly expanded distally. A pair of penial internal pilasters fusing into one Y-shaped fork at proximal 1/3; ~ 2 larger pilasters present nearby. Distally inside penis, numerous fine pilasters merging into 6 to 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 equal to or slightly shorter than 2/3 length of dart sac. Dart sac ~ 1/2 length 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 5–7, each complicatedly branched. Proximal accessory sacs two, symmetrical, dorsally separated and ventrally touching, with a few internal pilasters, each with an opening leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.
This species has a spire of variable height, from flatness to dome shape. The umbilicus of this species is obviously narrower than that of
None.
S Gansu (type locality).
W Sichuan (type locality: Daduhe River).
The protoconch is finely granulate. Growth lines are indistinct. Spiral grooves are only more distinct on umbilicus side near aperture. Bottom-umbilicus transition changes gently. Peristome is somewhat sinuate.
Sichuan (type locality); Hubei (type locality).
The protoconch is finely granulate. Spiral grooves are very weakly present. Bottom-umbilicus transition changes gently. Peristome is somewhat sinuate.
HBUMM00566a, 5 juvs; Luotuoxiang [骆驼巷], Bikou [碧口], Wenxian, Gansu Province,
Genital anatomy of
Gansu: Wudu (type locality), Wenxian.
The protoconch is smooth except some fine threads at the beginning and the end of the whorls. Spiral grooves are absent. On teleoconch between two adjacent ribs there are several indistinct fine threads.
Eversible head wart present but not prominent. Jaw arcuate, with seven projecting ribs.
Penial sheath short, covering ~ 1/6 of penis. Penis tubular and equally thick. Inside penis, ~ 5 low pilasters present proximally, two high pilasters fusing into one Y-shaped fork at proximal 1/4. Fine pilasters on distal end of penis not merging into thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 4.5 mm long, apically 2-bladed. Accessory sac small but externally distinguishable, internally solid, inserting into dart sac at distal 1/3, opening to dart chamber. Mucous glands 5 (HBUMM08430-spec.1), each singly tubular or simply branched. Proximal accessory sacs two, asymmetrical with the right one distinctly larger, apically separated and ventrally touching, each centrally with an opening (rounded, diameter ~ 0.2 mm) leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.
This species has a pair of asymmetrical and ventrally touching proximal accessory sacs that is rarely observed in the genus. In comparison, its conchologically similar species
For more comments, see
Genital anatomy of
HBUMM08452, 1
Gansu: Wenxian (type locality).
Protoconch is smooth and has no granules, perhaps caused by erosion or weathering. Teleoconch has rough ribs between which fine threads are present. Spiral grooves are absent throughout.
Head with flat but distinct eversible head wart. Jaw arcuate, with four projecting ribs.
Penial sheath covering ~ 1/9 of penis. Penis tubular and distally thicker. Inside penis, ~ 4 high parallel pilasters on proximal 2/3, at proximal ~ 1/4 two weak pilasters fusing into one Y-shaped fork. Fine pilasters on distal end of penis merging into ~ 7 thin or thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 2/5 length of penis. Accessory sac tiny, internally solid, inserting into dart sac at middle part, opening to dart chamber. Mucous glands four, each singly tubular or simply branched. Proximal accessory sacs two, of similar size and symmetrical, apically separated and ventrally touching, each distally with an opening leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix long-ovate.
The species has the most common pattern of the dart sac apparatus of the genus, i.e., two equally- sized and ventrally touching proximal accessory sacs, each with an opening leading to the dart chamber; this has been observed in ten of the 21 species anatomically studied herein. This species can be immediately recognized by its particular shell morphology.
HBUMM05433, 2
Genital anatomy of
Gansu: Wenxian (type locality).
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.
Head with flat but distinct eversible head wart. Jaw arcuate, with 5–7 projecting ribs.
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.
This species is one of the three species that do not have proximal accessory sac on dart sac apparatus. However, the shell morphology of
HBUMM00506, 13
Gansu: Wenxian. Sichuan: Jiuzhaigouxian (type locality).
Tiny elongate granules on protoconch, which can be only observed in juvenile shells, are densely and radially arranged. The granules on protoconch are absent in adult shell because of erosion or weathering. On teleoconch the growth lines are indistinct. Spiral grooves are absent throughout.
Eversible head wart prominent. Jaw arcuate, with three projecting ribs.
Penial sheath covering ~ 1/4 of penis. Penis proximately tubular and distally expanded. Inside penis, two pilasters fusing into one Y-shaped fork at proximal 1/5, two more pilasters of similar thickness also present. Fine pilasters on distal end of penis merging into ~ 8 thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 1 mm long, apically 2-bladed. Accessory sac small but externally distinguishable, internally solid, inserting into dart sac at distal 1/3, opening to dart sac chamber. Mucous glands four or five, each singly tubular or simply branched. Proximal accessory sac absent. Bursa copulatrix duct equally thick.
Genital anatomy of
See
HBUMM00633, many
Genital anatomy of
Gansu: Wenxian (type locality).
Granules on protoconch can only be unclearly and partially observed near suture because of erosion or weathering in adults. Granules are short (~ 20 µm long) on fine threads. Spiral grooves are absent throughout. On teleoconch whorls, fine threads are present between every two adjacent ribs.
Genital anatomy of
Eversible head wart present. Jaw arcuate, with five projecting ribs.
Penial sheath short, covering ~ 1/6 of penis. Penis tubular, somewhat expanded distally. Inside penis, ~ 5 low pilasters present proximally, another two high pilasters fusing into one Y-shaped fork at proximal 1/2. Fine pilasters on distal end of penis merging into ~ 4 thick folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 6 mm long, apically rhombic in cross-section, subsequently 2-bladed. Accessory sac small but externally distinguishable, internally solid, inserting into dart sac at middle part, opening to dart chamber. Mucous glands two (
HBUMM00422, 3
Chongqing. Gansu: Dangchangxian, Wenxian (type locality), Zhouquxian. Sichuan: Jiuzhaigouxian.
Tiny granules (~ 25 – ~ 40 µm) on the protoconch are regularly and densely arranged. Each granule is like a hump deposited in a shallow socket. On adult shell the protoconch granules are not clear, perhaps because of erosion. On teleoconch, tiny ear-shaped scales are arranged along growth lines. On umbilical side, scale scars are regularly arranged. Spiral grooves are only present on apical side of body whorl. The shell of specimens HBUMM08366 and HBUMM08421 are not typical because the periphery is angular rather than sharply carinate.
Eversible head wart of normal size. Jaw arcuate, with five projecting ribs (HBUMM05469).
Penial sheath short, proximally covering ~ 1/5 penis. Penis tubular, slightly expanded distally. Inside penis, three or four low pilasters present proximally, two fusing into one Y-shaped fork at proximal 1/2, another two high pilasters fusing into one Y-shaped fork at proximal 1/3 with cross expanded to be a large lump (~ 1 × 1 × 2 mm3). Fine pilasters on distal end of penis merging into more than eight thick but short folds. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/3–1 length of dart sac. Dart sac ~ 1/2 (HBUMM05469) to 2/3 (HBUMM05549) length of penis. Love dart ~ 8 mm long, apically rhombic in cross-section. Accessory sac small but externally distinguishable, internally solid, inserting into dart sac at distal 1/3, opening to dart chamber. Mucous glands three (HBUMM08423), four (HBUMM05469, HBUMM08421), or eight (HBUMM05549), each simply or complicatedly branched. Proximal accessory sacs two, symmetrical, apically separated and ventrally touching, internally with a few low pilasters, each with a distal opening leading to proximal dart chamber. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.
The most distinctive character of the genitalia of
The undetermined juvenile specimens from Chongqing (HBUMM08120) conchologically resemble
HBUMM05495, many
Genital anatomy of
Gansu: Huixian [徽县], Wenxian, Wudu, Zhouqu (type locality).
The protoconch is radially and regularly covered with dense thick granules (~ 25 – ~ 55 µm long) on the first 11/4 whorls. On the remaining protoconch whorls granules are indistinct and replaced by crowded radial threads. Spiral grooves are only indistinctly present on the apical side of body whorl.
Eversible head wart indistinct. Jaw arcuate, with three projecting ribs.
Penial sheath very short, covering ~ 1/10 of penis. Penis tubular, equally thick. Inside penis, two high pilasters forming one Y-shaped fork at proximal 1/5, besides with two pilasters parallelly merging into one thickest pilaster. Fine pilasters on distal end of penis merging into ~ 6 thick folds, among which one is thicker than the others. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 1/2 length of penis. Love dart ~ 9 mm long, apically 2-bladed, subsequently rounded. Accessory sac small, internally solid, inserting into dart sac at middle part, opening to dart chamber. Mucous glands ~ 5, each complicatedly branched. Proximal accessory sacs two, symmetrical, separated apically and touching ventrally, each with an opening leading to proximal dart chamber near dart chamber opening. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, very small.
Geographically, the distribution of
Compared to
The specimens HBUMM08433 (8
HBUMM00486, 3
Genital anatomy of
Gansu: Dangchangxian (type locality), Diebuxian, Kangxian, Wenxian, Wuduxian, Zhouquxian (type locality). Sichuan: Jiuzhaigouxian.
Protoconch has regularly arranged fine granules (each ~ 30 µm long) and fine radial threads, both of which are usually obscured by erosion or weathering. Apical and umbilical sides of body whorl have densely and regularly arranged spiral grooves.
Eversible head wart present. Jaw arcuate, with five projecting ribs.
Penial sheath covering ~ 1/6–1/5 of penis. Penis tubular, slightly thick distally. Inside penis, two adjacent pairs of pilasters forming two Y-shaped forks at proximal 1/4. Fine pilasters on distal end of penis merging into ~ 9 more or less thick short folds. Vas deferens narrow throughout. Vagina between atrium and dart sac elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2–1 length of dart sac. Dart sac ~ 1/3 length of penis. Love dart ~ 3 mm long, apically rhombic in cross-section. Accessory sac small, internally empty, inserting into dart sac proximally, opening to dart chamber. Mucous glands 7 (HBUMM05664) – 10 (HBUMM05619), each simply or complicatedly branched. Proximal accessory sacs absent. Bursa copulatrix duct of even diameter. Bursa copulatrix ovate, small.
In terms of shell shape, this species has an obviously depressed spire and more or less sharp carina or angulation above periphery compared to the sympatric (Fig.
The systematic affinity between
For more comments, see
Shell height 6.9 mm, maximum diameter 15.6 mm, aperture height 3.9 mm, aperture breadth 5.8 mm, umbilicus diameter 5.0 mm, protoconch whorls 11/2, whorls 87/8.
Protoconch with dense radially-arranged threads, visible through umbilicus. Beneath carina a clear chestnut band present. Mucous glands four or five. Penis with two pairs of pilasters fusing into two Y-shaped forks. Vagina between atrium and dart sac not elongated. Proximal accessory sac absent.
Sinistral, depressed, thin but somewhat solid. Shell with 81/8–93/8 fairly flat whorls. Suture impressed. Protoconch 11/2–13/4 whorls, with densely arranged radial threads that may be invisible because of weathering or erosion. Periphery distinctly angulate. Growth lines indistinct. Spiral grooves are absent throughout. Aperture oblique, roundly square, descending in front. Peristome not expanded and indistinctly reflexed at lower part. Within aperture a ring-like thickening present, basally with a flat tooth. Columella oblique. Umbilicus abruptly broadened after penultimate whorl, ~ 1/4 of maximum diameter. Protoconch visible through umbilicus. Shell apically in intermittent yellowish white and pale chestnut patches except white carina. In umbilical view shell distinctly paler in greyish yellow, just beneath carina a clear chestnut band present, umbilical region brownish.
Shell measurements (range, mean ± s.d.) of the new species of
Species |
|
Shell height | Maximum diameter | Aperture height | Aperture breadth | Umbilicus diameter | Protoconch whorls | Whorls |
---|---|---|---|---|---|---|---|---|
8 | 5.5–8.4 7.2±1.05 | 11.4–17.2 15.1±2.23 | 2.8–4.6 3.9±0.66 | 4.2–6.8 5.8±0.95 | 3.0–5.0 4.1±0.65 | 1.500–1.750 1.594±0.1108 | 8.125–9.375 8.875±0.4432 | |
2 | 5.2–5.3 5.2±0.03 | 12.6–13.2 12.9±0.44 | 2.0–2.4 2.2±0.28 | 3.9–4.2 4.0±0.24 | 5.2–5.4 5.3±0.13 | 1.500 | 7.625–7.750 7.688±0.0883 | |
10 | 7.9–10.0 9.2±0.77 | 17.2–20.1 19.0±1.1 | 4.6–6.8 5.8±0.61 | 5.9–8.1 7.2±0.67 | 6.0–7.9 6.7±0.56 | 1.500–1.750 1.600±0.0986 | 8.625–9.250 8.838±0.2361 | |
9 | 8.6–10.4 9.3±0.56 | 16.5–19.7 18.7±1.05 | 5.0–6.5 6.0±0.48 | 6.0–8.1 7.3±0.62 | 4.8–7.1 6.2±0.72 | 1.500–1.625 1.609±0.0442 | 9.000–9.625 9.219±0.0.2086 | |
6 | 10.7–13.9 12.3±1.19 | 20.7–27.5 24.5±2.10 | 6.2–8.5 7.6±0.78 | 9.0–11.9 10.7±0.99 | 4.1–6.4 5.1±0.59 | 1.500–1.625 1.510±0.0347 | 6.125–7.000 6.500±0.2447 | |
8 | 7.4–8.2 7.9±0.31 | 21.3–23.7 22.8±0.79 | 5.6–6.9 6.2±0.49 | 9.8–11.4 10.5±0.47 | 5.0–6.1 5.6±0.35 | 1.500–1.625 1.516±0.0442 | 5.500–5.875 5.703±0.1325 | |
5 | 3.6–3.8 3.7±0.08 | 10.3–10.8 10.5±0.18 | 2.2–3.0 2.7±0.31 | 2.9–3.3 3.1±0.16 | 4.6–5.0 4.8±0.13 | 1.500–1.625 1.600±0.0560 | 5.000–5.125 5.100±0.0559 |
Eversible head wart prominent. Jaw arcuate, with four projecting ribs.
Penial sheath covering ~ 1/5 of penis. Penis distally fairly expanded. Inside penis, two high pilasters fusing into one Y-shaped fork at proximal 1/4, another pair of pilasters fusing into one Y-shaped fork at middle part. Fine pilasters on distal end of penis merging into ~ 7 thick and short folds. Epiphallic papilla absent. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Vagina between dart sac and insertion of bursa copulatrix duct ~ 1/2 length of dart sac. Dart sac ~ 1/2 length of penis. Accessory sac small but externally distinguishable, internally solid, ventrally inserting into dart sac at distal 1/3, together with mucous glands opening to dart chamber. Mucous glands four or five, each singly tubular or bifurcated. Proximal accessory sac absent. Bursa copulatrix duct equally thick.
The name of this new species is made up of
On rocks of local hill.
Only known from the type locality.
The new species exhibits a large intraspecific change in size (Fig.
Shell height 8.1 mm, maximum diameter 23.7 mm, aperture height 5.6 mm, aperture breadth 10.6 mm, umbilicus diameter 5.6 mm, protoconch whorls 11/2, whorls 55/8.
Genital anatomy of
Protoconch with decussate radial and spiral threads. Umbilicus broad and deep, through which protoconch is visible. A bright band present between carina and suture. Beneath carina a dark chestnut band present. Mucous glands two. Penis with two proximal thick internal pilasters fusing into a Y-shaped fork at proximal 1/3. Vagina between atrium and dart sac not elongated. Two proximal accessory sacs on both sides of dart sac, with two pores leading to opening of dart chamber.
Sinistral, depressed, thin but somewhat solid. Shell with 51/2–57/8 fairly flat whorls. Suture impressed. Protoconch 11/2–15/8 whorls, with decussate radial and spiral threads, on the first whorl of which may be invisible because of weathering or erosion. Growth lines fine, more or less clear. Aperture oblique, peach-shaped, descending. Peristome expanded and reflexed at lower part. Columella oblique. Umbilicus broad, ~ 1/3 of maximum diameter. Protoconch visible through umbilicus. Shell apically in chestnut except white carina, after the first three or four whorls a bright band present between carina and adjacent suture. In umbilical view shell distinctly paler in greyish yellow and just beneath carina a chestnut band present.
Eversible head wart present.
Penial sheath short but well developed. Penis of equal thickness, externally simple. Inside penis, two very thick longitudinal pilasters fusing into a Y-shaped fork at proximal 1/3, accompanied with another two thick pilasters, these plasters then change into numerous fine pilasters that distally merge into three short but thick folds near opening of epiphallus. Epiphallic papilla absent. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Accessory sac spherical, solid, inserting into dart sac medially, opening to distal dart chamber. Mucous glands two, each complicatedly branched. Proximal accessory sacs two, symmetrical, dorsally separated and ventrally touching, each with a pore leading to proximal dart chamber. Love dart ~ 6 mm long, apically 2-bladed, medially round-hexagonal. Bursa copulatrix duct equally narrow. Bursa copulatrix pear-shaped.
This new species is named after Mr Li, Qi-Shi, who made this work possible with his field work.
This species is found under rotten wood.
Only known from the type locality.
This new species has a unique protoconch on which decussate radial and spiral threads are present compared to the granulation on this part in the other
Shell height 5.2 mm, maximum diameter 13.2 mm, aperture height 2.0 mm, aperture breadth 4.2 mm, umbilicus diameter 5.4 mm, protoconch whorls 11/2, whorls 75/8.
Protoconch without granules. Umbilicus extremely broad, ~ 1/2 of maximum diameter. Shell evenly pale brown with umbilicus side paler.
Sinistral, fairly depressed, solid. Shell with 75/8–73/4 fairly flat whorls. Suture impressed. Spire depressed-cone-shaped. Protoconch 11/2 whorls, smooth on the first whorl where sculpture may be erased by weathering or erosion, followed by sparse radial threads. Protoconch visible through umbilicus. Growth lines unclear. Spiral grooves absent. After ~ 21/2 whorls, with regularly spaced thick ribs between which are many fine threads. Aperture oblique, peach-shaped, descending in front. Body whorl sharply carinate above periphery. Peristome seldom expanded and only slightly reflexed at lower part. Columella very oblique. Umbilicus very broad, ~ 1/2 of maximum diameter. Shell evenly pale brown with umbilicus side paler.
Eversible head wart small but prominent.
Penial sheath present. Vagina between atrium and dart sac not elongate. Mucous glands four (observations of the genitalia of this species are only based on the paratype HBUMM08422-spec.2).
This new species is named after the collector Mr. Qiming Li.
In October, this species was found in the crevices of broken stones on dry slope.
Only known from the type locality.
This species has the strongest ribs on the shell surface and the broadest umbilicus of all species of
Shell height 10.0 mm, maximum diameter 19.6 mm, aperture height 5.6 mm, aperture breadth 7.5 mm, umbilicus diameter 7.0 mm, protoconch whorls 15/8, whorls 91/4.
Genital anatomy of
Protoconch without granules. Umbilicus more than 1/3 maximum diameter, through which protoconch is visible. Beneath carina a chestnut band present. Palatal with two blunt teeth. Mucous glands six. Proximal 4/5 of penis with ~ 6 thick internal pilasters. Two adjacent pairs of penial pilasters fusing into two Y-shaped forks at distal 2/3 of penis. Vagina between atrium and dart sac moderately elongated. Proximal accessory sacs two, separate, symmetrical, each with a pore leading to opening of accessory sac.
Sinistral, depressed, thin but somewhat solid. Shell with 85/8–91/4 fairly flat whorls. Suture impressed. Protoconch 11/2–13/4 whorls, with very fine axial striae which may be invisible on the first whorl possibly by weathering or erosion. Growth lines thick or rib-like above but fine beneath carina. Above periphery a sharp whitish carina present. Aperture oblique, peach-shaped, slightly descending in front. On ring-like thickening within aperture, a blunt tooth present near columella and another one near carina. Peristome almost not expanded, just minutely reflexed at lower part. Columella oblique. Umbilicus with a tint of pale brown, broadly conical, more than 1/3 of maximum diameter. Protoconch visible through umbilicus. Whorls apically in yellowish white with intermittent brownish patches. In umbilical view shell yellowish white with several brownish patches, and just beneath carina a chestnut band present.
Eversible head wart lowly present.
Penial sheath very short. Penis distally slightly expanded, externally simple. Penis of proximal 4/5 internally with ~ 6 thick longitudinal pilasters, two adjacent pairs of which fuse into two Y-shaped forks at distal 2/3; pilasters then branching into fine pilasters that are connected to form network. Epiphallic papilla absent. Vas deferens narrow throughout. Vagina between atrium and dart sac somewhat elongated. Accessory sac spherical, empty, inserting into dart sac at middle part, opening near the opening of dart chamber. Mucous glands six, each a single tube or simply branched. Proximal accessory sacs two, separate, symmetrical, each with a pore leading to opening of accessory sac/dart chamber. Love dart ~ 5 mm long, rounded and bladeless throughout. Bursa copulatrix duct equally narrow.
This new species is named after Mr. Liu, Zheng-Ping [刘正平], an amateur Chinese conchologist.
This species is found on exposed slate rocks of hill side.
This species is only known from the type locality.
The new species is conchologically close to
Genital anatomy of
Shell height 9.0 mm, maximum diameter 19.3 mm, aperture height 5.6 mm, aperture breadth 7.3 mm, umbilicus diameter 7.1 mm, protoconch whorls 15/8, whorls 91/8.
Protoconch with elongated granules. Umbilicus suddenly narrowed from penultimate whorl, more than 1/3 maximum diameter, through which protoconch is visible. Beneath carina a chestnut band present. Mucous glands four. Proximal 1/2 of penis with ~ 5 thin internal pilasters, two of them fusing into a Y-shaped fork at middle part. Distal region of penis with tongue-shaped papillae. Vagina between atrium and dart sac not elongated. Proximal accessory sacs two, symmetrical, each with a pore leading to dart chamber.
Sinistral, depressed, somewhat solid. Shell with 9–95/8 fairly flat whorls. Suture impressed. Protoconch 11/2–15/8 whorls, with fine granules (each ~ 20 µm long) almost invisible because of weathering or erosion. Growth lines thick or rib-like above but fine below carina. Above periphery a whitish carina present. Aperture oblique, peach-shaped, descending. On ring-like thickening within aperture, a very blunt tooth present near columella. Peristome almost not expanded, just minutely reflexed at lower part. Columella oblique. Umbilicus brownish, more than 1/3 of maximum diameter, suddenly narrowed from penultimate whorl. Protoconch visible through umbilicus. Whorls apically yellowish brown with intermittent darker brown patches. In umbilical view shell yellowish white with several brownish patches, and just beneath carina a thick chestnut band present.
Eversible head wart weakly present.
Penial sheath short but well developed. Penis distally expanded, externally simple. Inside penis, proximal 1/2 with ~ 5 internal pilasters, two of which fuse into one Y-shaped fork at middle of penis; distal 1/2 with tongue-shaped (HBUMM05553b) or diamond-shaped papillae (HBUMM08428) that are erect or inclined towards atrium. Fine pilasters on distal end of penis merging into ~ 6 more or less thick short folds. Epiphallic papilla absent. Vas deferens narrow throughout. Vagina between atrium and dart sac not elongated. Accessory sac tiny, internally solid, inserting into dart sac medially, opening to dart chamber. Mucous glands four, each a single tube or simply branched. Proximal accessory sacs two, symmetrical, dorsally separated and ventrally touching each other, each with a ventral pore leading to proximal dart chamber. Love dart ~ 7 mm long, apically 2-bladed or rhombic (HBUMM08428-spec.1) then rounded. Bursa copulatrix duct equally narrow.
This new species is named in honor of Prof Chen, De-Niu, who works on land mollusks in the Institute of Zoology, Chinese Academy of Sciences, Beijing.
This species is found on exposed slate rocks of the hill side.
Only known from the type locality.
The new species is conchologically close to
Voucher information and the GenBank accession numbers of the species in the phylogenetic study. An asterisk (*) indicates obtained from NCBI. The generic assignment for species follows MolluscaBase (2021a–p; 2022a–c).
Taxa | GenBank accession numbers 16S/ITS2 | Museum voucher | Voucher information |
---|---|---|---|
|
|||
HBUMM04882_2 | Fukuoka, Japan; coll. Guo, J.-Y. [郭建英], 2004-X-12 | ||
HBUMM04882_3 | |||
HBUMM05089c2 | Haikou, Hainan; coll. Wu, M., 2005-VIII-15 | ||
HBUMM05089c4 | |||
|
|||
HBUMM06480 | Zhenjiang, Jiangsu; coll. Wu, M., Xu, Q., 2011-VI-12 | ||
|
|||
HBUMM05104_2 | Zhongxian, Chongqing; coll. Wu, M., 2004-VII-16 | ||
HBUMM05122_2 | Fengjie, Chongqing; coll. Wu, M., 2004-VII-18 | ||
HBUMM05122_3 | |||
HBUMM05112_2 | Guangan [广安], Sichuan; coll. Wu, M. | ||
HBUMM01031_2 | Maoxian, Sichuan; coll. Wu, M., 2001-X-11 | ||
HBUMM01031_3 | |||
HBUMM05493_3 | Wenxian, Gansu; coll. Wu, M., 2006-IX-28 | ||
HBUMM06900_1/HBUMM06900_2 | Qixiashan [栖霞山], Jiangsu; coll. Wu, M. | ||
/ | Koehler, unpublished data; |
||
HBUMM05406_2 | Wenxian, Gansu; coll. Wu, M., 2006-IX-27 | ||
HBUMM05406_3 | |||
HBUMM05446_2 | Wenxian, Gansu; coll. Wu, M., 2006-IX-27 | ||
HBUMM05446_3 | |||
HBUMM06603 | Jiuzhaigouxian, Sichuan; coll. Wu, M., 2011-VIII-11 | ||
HBUMM08276a1 | Changbaishan [长白山], Jilin; coll. Shen, X.-F., 2018-VIII-18 | ||
HBUMM08276a2 | |||
|
|||
HBUMM06678 | Wenxian, Gansu; coll. Wu, M., 2011-VIII-9 | ||
HBUMM06676 | Wenxian, Gansu; coll. Wu, M., 2011-VIII-9 | ||
|
|||
HBUMM05089a_2 | Haikou [海口], Hainan; coll. Wu, M., 2005-VIII-15 | ||
HBUMM05089a_3 | |||
HBUMM05089a_4 | |||
|
|||
HBUMM08358a1 | Wenchuanxian, Sichuan; coll., Liu, Z.-P., 2019-VII-5 | ||
HBUMM08358a3 | |||
HBUMM05168_1 | Haidian [海淀], Beijing; coll. Wu, M., 2006-VII | ||
HBUMM05368_3 | Qingdao [青岛], Shandong; coll. Wu, M., 2006-X-17 | ||
HBUMM05602cb | Wudu, Gansu; coll. Liu, J.-M. and Zheng, W., 2006-IX-30 | ||
HBUMM05655 | Dangchang, Gansu; coll. Liu, J.-M. and Zheng, W., 2006-X-2 | ||
HBUMM06560 | Jiuzhaigouxian, Sichuan; coll. Wu, M., 2011-VIII-14 | ||
HBUMM08208a | Dingxi, Gansu; coll. Shen, X.-F., 2017-VIII-4 | ||
|
|||
HBUMM05933 | forest between Vladivostok and Artion cities, Russia; coll. Sayenko, E.M., 2005-VI-26 | ||
|
|||
/ | Europe | ||
|
|||
HBUMM08432a1 | See in the text | ||
HBUMM08432a2 | |||
HBUMM05103_2 | See in the text | ||
HBUMM05103_3 | |||
HBUMM05131_2 | See in the text | ||
HBUMM05131_3 | |||
HBUMM08368a | See in the text | ||
HBUMM08368a3 | |||
HBUMM08368a4 | |||
HBUMM08428a2 | See in the text | ||
HBUMM08428a4 | |||
HBUMM05407_3 | See in the text | ||
HBUMM01256b_2 | See in the text | ||
HBUMM01256b_3 | |||
HBUMM05097_2 | See in the text | ||
HBUMM08430a1 | See in the text | ||
HBUMM08424a2 | See in the text | ||
HBUMM05411_4 | See in the text | ||
HBUMM05409_2 | See in the text | ||
HBUMM05409_3 | |||
HBUMM05409_4 | |||
HBUMM05440_3 | See in the text | ||
HBUMM08421a1 | See in the text | ||
HBUMM08421a3 | |||
HBUMM08421a4 | |||
HBUMM08423a1 | See in the text | ||
HBUMM08423a2 | |||
HBUMM08422a1 | See in the text | ||
HBUMM08431a1 | See in the text | ||
HBUMM08431a2 | |||
HBUMM08431a3 | |||
HBUMM05688 | See in the text | ||
HBUMM05617 | See in the text | ||
HBUMM05716 | See in the text | ||
HBUMM08425a1 | See in the text | ||
|
|||
HBUMM05164_2 | Quyang [曲阳], Hebei; coll. Wu, M., Wu, Q., 2006-V-4 | ||
HBUMM05164_3 | |||
HBUMM05164_4 | |||
|
|||
HBUMM06796 | Hangzhou [杭州], Zhejiang; coll. Wu, M., Xu, Q., 2012-V-21 | ||
|
|||
HBUMM04909_2 | Badong, Hubei; coll. Wu, M., Wu, Q., Qi, G., 2004-VII-31 | ||
HBUMM04909_3 | |||
|
|||
HBUMM06586b | Jiuzhaigouxian, Sichuan; coll. Wu, M., 2011-VIII-14 | ||
|
|||
HBUMM05412_2 | Wenxian, Gansu; coll. Wu, M., 2006-IX-27 | ||
HBUMM06720 | Jiuzhaigouxian, Sichuan; coll. Wu, M., 2011-VIII-12 | ||
|
|||
HBUMM08239a_1 | Shaoguan [韶关], Guangdong; coll. Yu, Di [余迪] | ||
HBUMM08239a_2 | |||
|
|||
HBUMM05451 | Wenxian, Gansu; coll. Wu, M., 2006-IX-27 | ||
HBUMM05439_2 | Wenxian, Gansu; coll. Wu, M., 2006-IX-27 | ||
HBUMM05491c_2 | Wenxian, Gansu; coll. Wu, M., 2006-IX-28 | ||
HBUMM05528 | Wenxian, Gansu; coll. Wu, M., 2006-IX-29 | ||
|
|||
HBUMM08302a | Wuhan, Hubei; coll. Chen, Zhe-Yu [陈哲宇], 2019-III-20 | ||
|
|||
HBUMM01221_3 | Meitan, Guizhou; coll. Wu, M., 2003-VIII-2 | ||
HBUMM04904_2 | Badong, Hubei; coll. Wu, M., 2004-VIII | ||
HBUMM04904_3 | |||
HBUMM05095_2 | Zigui, Hubei; coll. Wu, M., 2004-VIII-2 | ||
HBUMM05095_3 | |||
HBUMM05123_2 | Fengjie, Chongqing; coll. Wu, M., 2004-VII-18 | ||
HBUMM05132_2 | Wuxi, Chongqing; coll. Wu, M., 2004-VII-22 |
Genital anatomy of
Shell height 10.8 mm, maximum diameter 21.7 mm, aperture height 6.1 mm, aperture breadth 9.2 mm, umbilicus diameter 5.1 mm, protoconch whorls 11/2, whorls 61/4.
Internal view of penis
Protoconch with fine granules. Umbilicus moderately broad, through which protoconch is visible. Bluntly carinate slightly above periphery. Beneath carina a brown band present. Mucous glands 6–8. Approximately 1/2 of penis with three proximal thick internal pilasters, Y-shaped fork formed by adjacent pilasters absent. Vagina between atrium and dart sac moderately elongated. Proximal accessory sacs two, dorsally separated and ventrally touching, each with a pore leading to dart sac chamber near entrance of dart chamber.
Internal view of penis
Sinistral, depressed, thin but somewhat solid. Shell with 61/8–7 slightly convex whorls. Suture impressed. Protoconch 11/2–15/8 whorls, fine granules (each ~ 30 – ~ 50 µm long) distinctly present. Growth lines fine and unclear. Aperture oblique, peach-shaped, descending. Within aperture with a white thickening. Peristome expanded and reflexed at lower part. Columella oblique. Umbilicus moderately broad, ~ 1/4 of maximum diameter. Protoconch visible through umbilicus. Shell apically in pale brown with some intermittent darker patches. Carina slightly above periphery, blunt, white; beneath which a brown band present. Whorls apically in yellowish white with intermittent brownish patches. In umbilical view shell yellowish white with several white radial striations.
Internal view of penis
Eversible head wart lowly present.
Penial sheath short. Penis distally swollen, externally simple. Proximal 1/2 penis with four thick internal pilasters which do not fusing, pilasters then branching into numerous fine pilasters that merge into ~ 6 short but thick (thickest in the genus) folds near opening of epiphallus. Vas deferens narrow throughout. Vagina between atrium and dart sac moderately elongated. Accessory sac spherical, internally with high pilasters and fairly solid inside, inserting into dart sac medially, opening to distal dart chamber. Mucous glands 6–8, each single tube or simply branched.
This new species is named after name of the type locality Qingchuan, Sichuan Province.
This species is found on slate rocks covered with mosses in a humid mountainous environment.
Only known from the type locality.
Internal view of penis
Shell height 3.7 mm, maximum diameter 10.8 mm, aperture height 3.0 mm, aperture breadth 3.3 mm, umbilicus diameter 5.0 mm, protoconch whorls 11/2, whorls 51/8.
SEM micrographs
Protoconch with dense fine granules. Umbilicus very broad, ~ 1/2 of maximum diameter. Body whorl shouldered above periphery. On shoulder with a white band. Peristome continuous. Shell strongly glossy as in
SEM micrographs
Sinistral, depressed, thin but somewhat solid. Shell with 5–51/8 convex whorls. Suture impressed. Protoconch 11/2–15/8 whorls, densely with tiny granules (each ~ 10 µm long) which are obscured by erosion or weathering. Growth lines indistinct. Shell in pale brown, strongly glossy, almost transparent. Body whorl shouldered above periphery, with a white band on shoulder. Aperture oblique, round, abruptly descending in front. A white thickening within aperture and thickened callus forming a continuous peristome. Peristome expanded and slightly reflexed at lower part. Columella oblique. Umbilicus very broad, ~ 1/2 of maximum diameter. Protoconch visible through umbilicus.
SEM micrographs
This cute new species is named in memory of Hartmut Nordsieck, a German malacologist who showed strong interest in the snails of the South Gansu Plateau and was a good friend of the first author.
SEM micrographs
This species was found on bare earth with a few broken rocks.
Habitats (excluding
Only known from two localities where the types were found.
Scatter plots of canonical variate 1 against canonical variate 2 (Canonical Variate Analysis), showing the shell morphological relationships among
The new species is the smallest species in
Bayesian phylogram of camaenid species (for detail see in Table
To investigate the systematic position of the putative
The phylograms produced by both Maximum Likelihood and Bayesian Inference based on partial 16S and partial ITS2 sequences are topologically identical in the major branches with the exception of different positions of two nodes (orange parts in Fig.
The indigenous dart-sac-bearing camaenids aggregated in Central China, which are grouped into the genera “
the monophyly of
Both “
“
More than 200 species belonging to 28 genera were recorded as bradybaenine or believed to be dart-sac-bearing camaenids in Chinese Mainland (
The work of
Until now, the systematic position of
The present phylogenetic inference is consistent with the phylogram of
Clade F (Fig.
Schematics of terminal genitalia of some dart-sac-bearing camaenids
Special thanks are due to Pavel Kijashko (