Abstract
Two new Pupinidius species, Pupinidius chenyui sp. nov. and Pupinidius zhengpingi sp. nov. were described from Sichuan Province, China, based on shell and genital characters. Pupinidius chenyui sp. nov. is the first sinistral species in Pupinidius. This discovery further reinforces our understanding of the species diversity of Enidae in the Sichuan.
Introduction
The terrestrial malacofauna of Sichuan Province, China, has been recognized as extremely abundant in early studies (e.g., Hilber 1883, Möllendorff 1901, Pilsbry 1934, Sturany 1899, Wiegmann 1901, Yen 1938, 1939, 1942). Recent research has highlighted that both the families Enidae, Camaenidae, and Cyclophoroidea exhibit high species diversity in this region (Wu 2001, 2002, Wu & Guo 2003, Wu & Zheng 2009, Wu & Gao 2010, Páll-Gergely et al. 2015, 2017, Wu & Asami 2017, Páll-Gergely et al. 2018, Wu 2018, Chen et al. 2022, Chen 2020, Wu et al. 2023, Chen et al. 2024a, 2024b). Pupinidius Möllendorff, 1901 is a group of moderate sized land snail endemic in Western China and Nepal, and characterized by the short cylindrical and mostly keg-like shell, free oviduct (except in Pupinidius obrutschewi contractus (Möllendorff, 1901) almost invisible). It consists of 16 known species (Kuznetsov & Schileyko 1997, 1999, Wu & Zheng 2009, Wu 2018, Chen et al. 2024c).
In this study, two new Pupinidius species, i.e. Pupinidius chenyui sp. nov. and Pupinidius zhengpingi sp. nov., are described from the dry-hot valley in Sichuan Province, China. The new species further enhances our understanding of land snail endemism in western China.
Materials and methods
All the examined specimens were collected alive. The animals were anaesthetized using menthol in water, killed in hot water, fixed and preserved in 70% ethanol, and dissected for the examination of the genitalia. Directions used in the general description of genitalia: proximal = towards the genital atrium; distal = away from the genital atrium.
Shells were photographed using a Canon® 5D Mark IV camera attached with a Laowa® 25 mm f/2.8 2.5-5X Ultra Macro lens. The final high DoF (depth of field) images were stacked from 30 to 50 single photos using Zerene Stacker® 1.04. Shell whorl counting is accurate to 1/8 (= 0.125) whorls, following the criteria by Kerney and Cameron's (1979). Muscle tissues were obtained from specimens HBUMM10064 (sample ID SYS m001018) and HBUMM10066 (sample ID SYS m001019) and then preserved in 95% ethanol. The names of administrative units below provincial level are given in Chinese Pinyin (Shi = city, Xian = county, Zhen = town).
Abbreviations
A-1–A-5 | five sections of penial appendix |
At | atrium |
AR | retractor muscle of appendicular branch |
BC | bursa copulatrix |
BCD | bursa copulatrix duct |
D | diverticle of bursa copulatrix duct |
Ep | epiphallus |
EpC | epiphallic caecum |
Fl | flagellum |
FO | free oviduct |
HBUMM | mollusc collection of the Museum of Hebei University (Baoding, China) |
P | penis |
PR | penis retractor |
Va | vagina |
VD | vas deferens |
Results
Family Enidae B. B. Woodward, 1903 (1880)
Genus Pupinidius Möllendorff, 1901
Type species: Buliminus pupinidius Möllendorff, 1901, by original designation
Diagnosis
Pupinidius zhengpingi sp. nov.
Figures 1A, 2A, 3B
Type material: Holotype: HBUMM10064, Shell height: 17.5 mm, shell breadth:7.8 mm, aperture height: 8.9 mm, aperture breadth: 4.7 mm. Koushan Cun [芤山村], Yanmen Zhen [雁门镇], Wenchuan Xian [汶川县], Ngawa Tibetan and Qiang Autonomous Prefecture [阿坝藏族羌族自治州], Sichuan Province, P.R. China, 103°38′28″N, 31°29′47″E, leg. Zheng-Ping Liu, 15 September 2020. Paratypes: HBUMM08356, One mature shell, same data as holotype; HBUMM10065, Eight mature and two juvenile animals, Koushan Cun (near the locality of holotype), Wenchuan Xian, Ngawa Tibetan and Qiang Autonomous Prefecture, Sichuan Province, P.R. China, leg. Zheng-Ping Liu, 5, July, 2019.
Measurements: Shell height: 13.7–18.0 mm; shell breadth: 6.9–9.3 mm; aperture height: 6.2–8.9 mm; aperture breadth: 4.5–5.8 mm (n = 10).
Diagnosis: Shell small-sized, shuttle-like, dextral. Aperture with weak angular tubercle. Teleoconch with clear and dense growth lines. Most swollen part at penultimate whorls. Penial caecum absent. Epiphallus and bursa copulatrix stalk straight. Epiphallic caecum present.
Description: Shell oval, dextral, yellowish brown, opaque, 7.125–8.250 whorls. Most swollen part at penultimate and body whorls. Last 3 whorls with alternating lighter and darker stripes. Apex prominent. Protoconch smooth, reddish brown. Teleoconch whorls with clear and dense growth lines, not shouldered, without spiral grooves except near umbilicus. Suture shallow, without narrow defined zone on beneath whorl. Last whorl gradually ascending towards aperture. Aperture not adnate, slightly oblique, with shallow angular nodule. Parietal callus conspicuous but thin. Peristome almost on a plane, slightly expanded and reflexed, with a narrow cuff. Columella vertical or slightly bending to umbilical side. Columellar margin reflexed. Umbilicus narrowly open.
Genitalia. Vas deferens slender; entering epiphallus apically with distinct demarcation. Epiphallus long, cylindrical, of uniform thickness. Epiphallic caecum small. Flagellum short and conical. Penis clavate, thin-walled, without penial caecum; internally with more than two fine longitudinal pilasters; with two V-shaped pilasters. V-shaped pilaster with proximal free end approaching at penial retractor insertion, distally not developed into a papilla. Penis retractor absent. Penial appendix long; branching off immediately from penis near atrium; divided into sections including A-1+A-2, A-3 and A-4+A-5. A-1 short. A-1 and A-2 fused. A-2 near A-3 internally with a ring of weak papilla. A-2 and A-3 unfused. A-4 and A-5 without distinct boundary, thin, straight. Atrium short, without atrial retractor muscle. Vagina unpigmented, long, of normal thickness, straight; not lined with loose spongy tissue. Bursa copulatrix duct long; straight. Bursa copulatrix voluminous; without apical ligament; with long neck; distinctly defined. Diverticle normally present; longer than reservoir; straight; terminally slightly expanded. Bursa copulatrix and diverticle distinguishable; forked more distally from their base.
Etymology: This species is named in honour of Mr. Zheng-Ping Liu (Chengdu, China) who collected the specimens.
Vernacular name: 正平蛹巢螺 (zhèng píng yǒng cháo luó).
Distribution and ecology: The new species is only known from the type locality. Like many recently reported new species of Enidae, they also was found on the bushes in the dry-hot valley (Chen et al. 2024c, 2024d).
Pupinidius chenyui sp. nov.
Figures 1B, 2B, 3C
Type material: Holotype: HBUMM10066, Shell height: 14.3 mm, shell breadth:7.3 mm, aperture height: 6.8 mm, aperture breadth: 4.6 mm. Koushan Cun [芤山村], Yanmen Zhen [雁门镇], Wenchuan Xian [汶川县], Ngawa Tibetan and Qiang Autonomous Prefecture [阿坝藏族羌族自治州], Sichuan Province, P.R. China, 103°38′25″N, 31°29′43″E, leg. Zheng-Ping Liu, 15 September 2020. Paratype: HBUMM10067–10068, Two mature shells, same data as holotype.
Measurements: Shell height: 14.3–14.8 mm, shell breadth: 6.8–7.5 mm, aperture height: 6.8–7.5 mm, aperture breadth: 4.6–5.1 mm (n = 3).
Diagnosis: Shell dirty white, sinistral. Aperture with weak angular tubercle. Most swollen part at penultimate whorls. Teleoconch whorls rarely without growth lines. Umbilicus rimate. Penis club-shaped.
Description: Shell oval, sinistral, dirty white, 7.250–7.750 whorls. Apex prominent, darkly tinted. Protoconch smooth, yellowish brown. Teleoconch whorls shortly ascending in front, with indistinct growth lines, not shouldered, without spiral grooves except at umbilical region. Suture shallow, without a narrow defined zone on beneath whorl. Aperture not adnate, truncate ovate. Parietal callus conspicuous but thin. Peristome sharp, slightly reflected but hardly. Columella vertical. Columellar margin reflexed. Aperture with shallow angular nodule. Umbilicus a narrow slit.
Genitalia. Vas deferens slender; entering epiphallus apically with distinct demarcation. Epiphallus long; of uniform thickness, straight on proximal half and forming many loops on distal half. Epiphallic caecum short; nipple-shaped; located on distal 2/3 of epiphallus. Flagellum short; conical. Penis club-shaped; thin-walled; without penial caecum; internally with one fine longitudinal pilaster; with one V-shaped pilaster. V-shaped pilaster with proximal free end approaching at penial retractor insertion; distally not developing into a papilla. Penis retractor long. Penial appendix long, about two times of penis plus epiphallus; branched off from proximal penis; divided into sections including A-1+A-2, A-3 and A-4+A-5. Boundary between A-4 and A-5 indistinct. Vagina unpigmented, not swollen; not lined with loose spongy tissue. Bursa copulatrix duct long; distal 2/3 strongly convoluted. Bursa copulatrix without apical ligament; with short neck; well defined and expanded. Bursa copulatrix and diverticle distinguishable; forked more distally from their base.
Etymology: This species is named after Mr. Yu Chen, a famous paleontological restoration artist with the nickname “sinammonite”, who helps the second author a lot.
Distribution and ecology: This species, inhabiting the bushes in the dry-hot valley, is only known from the type locality.
Vernacular name: 陈瑜蛹巢螺 (chén yú yǒng cháo luó).
Remarks: Conchologically, Pupinidius chenyui sp. nov. is sinistral and pupil-shaped and therefore differs from all the other Pupinidius species. In terms of genital system, the new species is most similar to P. nanpingensis nanpingensis (Möllendorff, 1901). They share the looped distal epiphallus and the looped distal bursa copulatrix duct, the straight A4+A5 (Wu 2018). However, the new species has no penial caecum.
Acknowledgments
We thank Zheng-Ping Liu (Chengdu, China) for providing the Pupinidius specimens. We are grateful to Min Wu (Nanjing University, China) for help polishing the manuscript. Barna Páll-Gergely, Zhong-Guang Chen, Min Wu and one anonymous reviewer for their valuable comments for the manuscript. 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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