Phuphania crossei Jirapatrasilp & Tongkerd & Jeratthitikul & Liew & Pholyotha & Sutcharit & Panha, 2021

Jirapatrasilp, Parin, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Liew, Thor-Seng, Pholyotha, Arthit, Sutcharit, Chirasak & Panha, Somsak, 2021, Molecular phylogeny of the limacoid snail family Dyakiidae in Southeast Asia, with the description of a new genus and species, Zoological Journal of the Linnean Society 193 : -

PHUPHANIA CROSSEI ( PFEIFFER, 1862) COMB. NOV.

( FIGS 5C, D, 6B, 7B, 8B)

Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 8701796D-C54F-4A07-B29E-ADCD6351E3B4

Synonymy

Helix crossei Pfeiffer, 1862: 39 , pl. 5, figs 2, 3. Type locality: Siam [ Thailand] .

Helix danae Pfeiffer, 1863 [ 1862]: 268. Type locality: Laos Mountains, Camboja [ Cambodia or Laos]. New Synonym.

Nanina (Xestina) crossei – Kobelt, 1900: 983, 984, pl. 255, fig. 5, pl. 256, figs 3, 4.

Nanina (Xestina) danae – Kobelt, 1902: 1076, pl. 273, figs 4–6.

Euplecta danae – Gude, 1903: 6.

Hemiplecta crossei – Gude, 1903: 7.

Nanina (Hemiplecta) crossei – Fischer & Dautzenberg, 1904: 393.

Nanina (Hemiplecta) danae – Fischer & Dautzenberg, 1904: 393.

Hemiplecta (Hemiplecta) crossei – Hemmen & Hemmen, 2001: 43.

Hemiplecta (Hemiplecta) danae – Hemmen & Hemmen, 2001: 44.

Ariophanta (Cryptozona) crossei – Schileyko, 2011: 29.

Ariophanta (Cryptozona) danae – Schileyko, 2011: 29.

Ariophanta crossei – Inkhavilay et al., 2019: 74 , fig. 34b.

Ariophanta danae – Inkhavilay et al., 2019: 74 , fig. 34c.

Type material examined

S y n t y p e o f H e l i x c r o s s e i P f e i f f e r, 1 8 6 2, MNHN-IM-2000-1869 (one shell; Inkhavilay et al., 2019: fig. 34b). Syntype of Helix danae Pfeiffer, 1863 , NHMUK 20092043 (one shell; Inkhavilay et al., 2019: fig. 34c).

Material examined

Sakunotayan Waterfall, Wang Thong District, Phitsanulok Province: CUMZ 5142 (nine shells), 5145 (two specimens in ethanol). Pa Ma Muang Temple, Ban Mung, Nern Maprang District , Phitsanulok Province: CUMZ 5137 (four shells), CUMZ 5141 (eight shells). Tam Wang Daeng, Nern Maprang District , Phitsanulok Province: CUMZ 5138 (six shells), CUMZ 5140 (one shell), CUMZ 5164 (seven shells). Khao Look Chang, Pakchong District , Nakhon Ratchasrima Province: CUMZ 5139 (two shells), CUMZ 5148 (one specimen in ethanol), CUMZ 5167 (one shell). Tam Yai Nam-nao, Nam-nao District , Phetchabun Province: CUMZ 5144 (one shell). Sap Chomphu Arboretum, Nong Pai District , Phetchabun Province: CUMZ 5146 (one specimen in ethanol), CUMZ 5169 (39 shells). Muak Lek Waterfall, Muak Lek District , Saraburi Province: CUMZ 5143 (one shell). Jed Kot Waterfall, Kaeng Koy District , Saraburi Province: CUMZ 5147 (four specimens in ethanol; Figs 5C, D, 6B, 7B), CUMZ 5168 (seven shells). Wat Tham Phra Phothisat, Kaeng Koy District , Saraburi Province: CUMZ 5272 (four specimens in ethanol; Fig. 8B), CUMZ 5273 (22 shells). Khao Sala, Bau Chet District , Surin Province: CUMZ 5149 (one shell) .

Diagnosis

Shell conic, brownish, with last whorl angular and umbilicus narrow. Penis enlarged and cylindrical. Amatorial organ gland fused lobes with three clusters of amatorial organ ducts and forming a cap over amatorial organ; amatorial organ papilla with corneous spike.

Description

Shell ( Fig. 6B): Shell medium (width ≤ 35.0 mm, height ≤ 24.5 mm), conic to slightly depressed conic, thickened and dull, dextral and narrow perforate. Whorls six to eight, increasing regularly; suture wide and little depressed. Shell colour brownish to yellowish; below periphery with paler colour than upper periphery. Spire convex; apex acute; embryonic shell small and with fine radial ridges. Shell surface coarse, with nodule lines arranged on fine radial ridges; below periphery with nearly smooth surface around umbilicus. Last whorl angular, with weak peripheral keel. Aperture ovate; parietal callus transparent; lip simple, slightly thickened and little expanded near columellar area. Columella slightly dilated. Umbilicus perforate, narrow and deep.

Genitalia ( Fig.5C,D): Atrium very short compared with penis. Penis enlarged, cylindrical and suddenly tapering to small tube. Epiphallus long, almost same length as penis and approximately similar in diameter to penis. Penial retractor muscle thin and attached between penis and epiphallus. Flagellum absent. Vas deferens smaller in diameter than epiphallus and connected at tip of epiphallus. Internal wall of penis with large and nearly smooth-surfaced longitudinal pilasters for entire length of chamber; penial verge absent.

Gametolytic organ (duct and sac) locates very near the end of amatorial organ. Gametolytic duct cylindrical tube; gametolytic sac gradually swollen to the end. Amatorial organ well developed, enlarged cylinder and attached to atrium. Amatorial organ gland fused lobes, covered amatorial organ ducts and forming a cap over amatorial organ gland. Amatorial organ duct consists of three clusters of ducts; tip of amatorial organ with thin connective tissue. Internal wall sculpture of amatorial organ: region close to atrium consists of smooth surface of longitudinal amatorial organ pilasters; area surrounding papilla consists of corrugated wall. Amatorial organ papilla enlarged, smooth surface and obtuse in shape; tipped with dark spike.

Vagina about same length as penis and cylindrical. Oviduct with enlarged lobules; prostate gland bound to oviduct. Internal wall of vagina: area close to atrium sculptured with irregular, undulated pilasters of similar size and arranged in longitudinal rows then with unpattern irregular surface.

Radula ( Fig. 7B): Teeth arranged in very wideangled U-shaped rows, with ~150 (90-(25–27)-1- (23–26)-60+) teeth. Central tooth relatively small, slightly asymmetric bicuspid, and with pointed cusps. Lateral teeth relatively large, asymmetric tricuspid, elongated in shape and inclined towards central tooth. Mesocone large and pointed cusp; endocone and ectocone very small and pointed cusps. Marginal teeth start from tooth 23–27, elongate, unicuspid and sword shaped; outermost teeth shorter, with apically pointed tip.

External features ( Fig. 8B): Living snail typical dyakiid form; body reticulated and yellowish brown. Upper tentacles long and dark grey; lower tentacles shorter and paler in colour. Dorsal part darker, especially near the head. Foot aulacopod and unipartite; foot sole same colour as the body and unspotted.

Remarks

This species has previously been assigned to the family Ariophantidae , for the lack of a prominent keel similar to some ariophantid genera, such as Ariophanta Des Moulins, 1829 and Hemiplecta Albers, 1850 . However, its exact familial or generic assignment could not be resolved owing to the lack of anatomical data. Two nominal species, Helix crossei and H. danae , were described in consecutive years by L. Pfeiffer, with a rather generic type locality of ‘Siam’ and ‘Lao Mountains, Camboja’, respectively. The latter taxon was described based on the Cuming ex. Mouhot collection. Hence, the specimen probably originated from the last journey of H. Mouhot between Bangkok, Thailand and Luang Phrabang, Laos (see Mouhot, 1864; Inkhavilay et al., 2019). We presume that the type specimen of H. danae was collected from the same geographical area as H. crossei . Specifically, the type localities were possibly in north-eastern Thailand, which is within the current distributional range of recently examined specimens. In addition, the syntypes of both nominal species were identical in all shell characters ( Inkhavilay et al., 2019). Thus, we consider these names as synonyms.

Phuphania crossei differs from its congeners in having a stronger peripheral keel, more slender gametolytic organ with a longer and more distinct gametolytic duct, and possessing a corneous spike on the amatorial organ papilla, which is absent in other species. The shell and genitalia of this species are more similar to Phuphania costata and Phuphania carinata than to Phuphania globosa in having a peripheral keel and distinct radial ribs, elongate marginal teeth and the lobes of the amatorial organ gland fusing as a thick sac that contains two to four clusters of internal ducts ( Kongim & Panha, 2013; Tumpeesuwan & Tumpeesuwan, 2014). However, Phuphania crossei has more compact amatorial organ glands and shows a less distinct demarcation between the amatorial organ and its glands. This morphological similarity among Phuphania carinata , Phuphania costata and Phuphania crossei agrees well with the molecular phylogenetic relationship ( Fig. 2).