Zootaxa,A new Satsuma species (Pulmonata ...

Zootaxa 1608: 59–68 (2007) www.mapress.com / zootaxa/

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ZOOTAXA

A new Satsuma species (Pulmonata: Camaenidae) endemic to Taiwan SHU-PING WU1, YAO-SUNG LIN1 & CHUNG-CHI HWANG 2,3 1

Institute of Ecology and Evolutionary Biology, National Taiwan University, 1, Roosevelt Rd, Sec. 4, Taipei, 10617, Taiwan. Department of Bioresources, Da-yeh University, No. 112 Shanjiao Rd, Dacun, Changhua, 51591, Taiwan. E-mail: [email protected] 3 Corresponding author 2

Abstract A new species of camaenid land snail, Satsuma longkiauwensis sp. nov. from southern Taiwan is established. This large terrestrial and herbivorous snail inhabits the lowland forests with a narrow geographical distribution. The species is characterized by having a large shell, roundly angulated peripheries adjacent to the peristome, an open umbilicus, a robust flagellum, a weak expansion on male genitalia instead of a penial caecum externally and a hemispherical verge instead of an elongated pilaster internally. A key is provided for the first time to identify camaenids from Taiwan. Key words: Camaenidae, Satsuma, new species, taxonomy, anatomy, identification key

Introduction More than thirty species of camaenid land snails from Taiwan have been described, but the diversity of this family is still far from being completely assessed (Wu & Wu 1998). Currently, 23 species and subspecies of Satsuma-like camaenids were recorded and confirmed; nevertheless the detailed species richness of this island requires further investigations (Hsieh et al. 2006). The genus Satsuma A. Adams, 1868 is a group of varied sized snails distributed in East Asia having depressed to high conic shells variable in color ranging from yellowish to dark brown with polymorphic banding patterns. In this study, a new species of Satsuma is described on the basis of shell and anatomical characters.

Material and methods Twelve specimens of the new species, which include three live snails, one decaying specimen and eight empty adult shells, were collected from Shuangliu and Mudan in the north of Hengchun Peninsula, southern Taiwan (Fig. 1). One individual was dissected instead of the holotype, as the soft body of holotype was decaying when collected. The live specimen was anesthetized using menthol in water, killed in hot water and then fixed and preserved in 70% ethanol. The shell of this specimen was broken and the soft body was dissected for the examination of the reproductive system. The dissections were performed using a stereo microscope (Leica MZ7.5). Drawings were made with a camera lucida attachment. Shell measurements were taken following Kerney and Cameron (1979), using a digital caliper (Kanon EMS-8) with an accuracy of 0.1 mm. The systematics followed Vaught (1989). Two individuals were examined for radula and jaw. The preparation for scanning electron microscopy (= SEM) followed the procedure given by Wu and Huang (1989). Subsequent examination was carried out using a FEI QUANTA 200 SEM. Accepted by B. Ruthensteiner: 5 Sep. 2007; published: 5 Oct. 2007

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The following abbreviations are used: National Taiwan Museum, Taiwan: TMMT; National Museum of Natural Science, Taiwan: NMNS; Natural History Museum, London, United Kingdom: BMNH; Academy of Natural Sciences of Philadelphia, United States of America: ANSP.

FIGURE 1. Map of Taiwan showing the distribution of Satsuma longkiauwensis sp. nov. Solid asterisk, type locality; empty asterisk, locality of paratypes NMNS 5415-001–003 and TMMT 0702–0703; grey area around asterisks, distribution of Satsuma longkiauwensis; bold black line, Central Range; thin line, administrative boundary.

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Systematics Order Stylommatophora Family Camaenidae Pilsbry, 1895 Satsuma A. Adams, 1868 Type species: Helix japonica Pfeiffer, 1847, by original designation

Satsuma longkiauwensis sp. nov. (Figs. 2–5) Type material. Type locality: TAIWAN: Pingtung County, Shuangliu, 22º13’7.3”N 120º47’35.9”E, alt. 280 m; TMMT 0701 (holotype): adult with decaying soft body in ethanol, collected by S.-P. Wu, 20 Oct. 2005; TMMT 0705 (paratype): adult dissected for genitalia, radula and jaw, remaining soft parts in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0704 (paratype): sub-adult dissected for radula and jaw, remaining soft parts in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0706 (paratype): sub-adult soft body in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0707 (paratype): adult shell, collected by S.-P. Wu, 21 Feb. 2004; BMNH 20060795 and ANSP 413696 (paratypes): adult shell, collected by S.-P. Wu, 20 Oct. 2005. Other locality: TAIWAN: Pingtung County, Mudan, 22º07’55.4”N 120º47’18.5”E, alt. 225 m; empty adult shells: NMNS 5415-001–003 (three paratypes) and TMMT 0702–0703 (two paratypes): collected by C.-C. Hwang, 26–27 Aug. 1997. Diagnosis. A large-sized, dextral Satsuma with light brown to olive background with depressed base and open umbilicus; Reproductive system with robust flagellum, swollen distal vagina, truncated oval verge, thick and muscular sheath tying male genitalia and expansive base of spermathecal stalk. Etymology. Longkiauw (Langjiao) is the ancient name of nowadays Hengchun Peninsula and originates from the aboriginal race Paiwan. Description. Shell (Fig. 2): large, dextral, semi-spherical, thick, rigid; height 24–33 mm; width 34–41 mm; 5 7/8 –6 1/8 whorls. Apex obtuse. Whorls strongly inflated. Periphery bluntly angulated. Base of shell bulged outwards and flattened behind peristome; Surface of periostracum smooth, glossy, with faint oblique and spiral striations. Color of shell light brown to olive with red-brown stain. Peripheral band thin, sharp, redbrown. Surface above peripheral band brown, gradually lighter towards suture; area immediately below peripheral band yellowish white; basal band light brown colored, broad, gradually smear towards umbilicus. Umbilicus uncolored. Aperture diagonal, ovate. Peristome in lateral view concave. Lip thick, expanded; reflected at outer and inferior parts. Columellar lip oblique, straight; dilation of columellar lip not covering umbilicus. Lip color dark purple-red, glossy. Parietal callus thin, semitransparent. Umbilicus open, broad, deep. Holotype: height 28 mm, width 39 mm, 6 1/8 whorls (Fig. 2A–D). Reproductive system (Fig. 3): Spermathecal stalk as long as spermoviduct, swollen at basal half, slender till spermatheca, with a constriction dividing base into two expansions of equal length; basal expansion bacillary, with ten to twelve strong, corrugated folds along inside; distal expansion shuttle-shaped, with seven thin, strong zigzag folds inside. Spermatheca oval. Free oviduct long, about same length as basal expansion of spermathecal stalk. Proximal vagina muscular, smooth externally, gross, weaker than basal expansion of spermathecal stalk. Internal folds in proximal vagina continued with those of spermathecal stalk counting nine in number, strongly elevated, lamellate, smooth, gradually weakened towards atrium. Subsequent part of vagina gradually slender towards a sudden swell distally, with ten thin, wiggly folds inside. Distal 1/5 of vagina abruptly thickened, muscular, depressed, pale brown in color, with wide, flattened folds inside. Male genitalia twisted, wrapped by a thin, semi-translucent, muscular sheath. Flagellum moderate, robust, tapering, with

SATSUMA LONGKIAUWENSIS SP. NOV.

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FIGURE 2. Satsuma longkiauwensis sp. nov. A–D. shell of holotype (TMMT 0701). Scale bar = 10 mm. A. Apertural view; B. Lateral view; C. Basal view; D. Top view; E. Living specimen (paratype TMMT 0704). Photograph by ChungYi Hsiao.


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FIGURE 3. Reproductive system of Satsuma longkiauwensis sp. nov. (paratype TMMT 0705) A. Whole genitalia, vas deferens is cut to unfold the genitalia; B. Verge and penial caecum; C. Interior of genitalia. Scale bar for A and C = 10 mm, scale bar for B = 1 mm.

thickened base. Epiphallus as strong as base of flagellum with four to six unequally strong pilasters along inside, terminated with a shortly projecting, truncated oval, finely wrinkled verge entering penis. Penial caecum (appendix) not definite, with only an angled expansion surrounding verge. Pilasters around verge weak, wiggly, numbered 15–19. Proximal part of penis strong, thickened with 13 smooth, well-defined, strong pilasters. Subsequent part of penis slender with nine low, wiggly, corrugated pilasters inside. Distal part of penis



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extended again, about same length as distal swell of vagina, with weak, zigzag, corrugated pilasters internally. Atrium short, finely wrinkled inside. Radula and jaw: Radula ribbon length 7.6 mm, width 2.6 mm. Radula formula 56 + C + 56. Central tooth with blunt, wide, curved mesocone and two weak, symmetrical ectocones (Fig. 4A). Lateral teeth with narrower, outwardly directed mesocone and weak endocone and ectocone (Fig. 4B). Marginal teeth with more outwardly directed mesocone and prominently forked, pointed endocone and ectocone (Fig. 4C). Jaw dark brown, curved with high, wide vertical ribs, reduced on margins (Fig. 5).

FIGURE 4. Radula of Satsuma longkiauwensis sp. nov. (paratype, TMMT 0705). A. Central tooth; B. Lateral tooth (15th, left); C. Marginal tooth (45th, left). Scale bar = 20 µm.

FIGURE 5. Jaw of Satsuma longkiauwensis sp. nov. (paratype, TMMT 0705). Scale bar = 600 µm.

Distribution (Fig. 1): All specimens were collected in mountainous lowland between the river basin of the rivers Fengangxi and Sichungxi. Living individuals were collected only in Shouangliu, the type locality. Ecology: Live specimens were observed active in the field in September. They were nocturnal and herbivorous feeding on decaying fallen leaves and gramineous shoots (Fig. 2E). They were found on ground and crawling on rocks or lower tree trunks.


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Remarks: The reproductive system of Satsuma longkiauwensis sp. nov. differs from that of the largesized Satsuma species from Taiwan and Japan, e.g. S. bairdi (H. Adams, 1866), S. arisana (Kuroda, 1941) ssp., S. nux (Moellendorff, 1888), S. sphaeroconus (Pfeiffer, 1866) and S. mercatoria (Pfeiffer, 1845), in having a weak expansion instead of an elongated penial caecum externally and a hemispherical verge instead of elongated and thickened pilasters internally (Minato 1975; Wu & Chang 1975; Azuma & Azuma 1987; Chang 1994, 1997). The reproductive system of the new species is similar to that of S. succincta (H. Adams, 1866), Pancala bacca (Pfeiffer, 1866) and P. batanica pancala (Schmacker & Boettger, 1891) (Tabe 1937; Sinagawa 1979; Chang 1992; Hwang 1995), but differs from P. bacca and P. batanica pancala in having a larger shell, roundly angulated peripheries adjacent to the peristome, an angled and less swollen junction between epiphallus and penis, robust and non-twisted flagellum, a more dilated penis at the proximal part and a swollen distal vagina. Shell morphology of the new species differs from other camaenid species distributed in Hengchun Peninsula, as P. bacca, P. batanica pancala, S. pekanensis (Rolle, 1911) and S. contraria (Pilsbry & Hirase, 1909), in having a larger and/or dextral shell (Hsieh et al. 2006). Satsuma arisana tani (Kuroda, 1941), which is distributed in southeastern Taiwan, is also similar to the new species in shell dimensions (holotype: height 30.0 mm, width 41.0 mm) and shape. Unfortunately the type specimens were not traceable currently. However, it can be deduced from the original description and photographs of the holotype (Kuroda 1941) that the umbilicus of S. arisana tani is perfectly covered by the reflected superior columellar lip and completely closed. Among camaenid species from Taiwan, Satsuma succincta shows a high similarity in shell morphology and banding patterns. Nevertheless, this species is smaller (shell height 16.2–22.8 mm, shell width 23.0–31.2 mm, 6 syntypes, BMNH 1866.5.9.9) in shell dimensions than the new species. Both species are ground-living in the low land forests of south Taiwan, but not sympatric. The smoothly rounded junction between the outer and inferior lips, yellowish coloration of shell, the robust and smoothly curved flagellum, and the swollen distal vagina of S. longkiauwensis sp. nov. distinguish it further from S. succincta. Satsuma succinct and its allies show an angulated junction at peristome, reddish coloration of shell mostly, slender and wiggly flagellum and slender distal vagina (Chang 1989; Hwang 1995). All the 26 valid (sub-)species of Camaenidae from Taiwan can be separated using the key below. This identification key is based on shell characters of type specimens examined except for Satsuma arisana takahasii (Kuroda, 1941), S. arisana tani (Kuroda, 1941), S. nux paiwanis (Kuroda, 1941) and S. nux sericata (Kuroda, 1941). The type material of these four taxa could not be located. Anatomical characters are not used, as those are not available for all taxa.

Key to the camaenid species from Taiwan 1 2 3 4 5 6

shell surface hairy or tubercular................................................................................................................... 2 shell surface smooth to roughly striated ...................................................................................................... 3 inferior and outer lips armed with fold ........................................... Moellendorffia hiraseana Pilsbry, 1905 inferior and outer lips not armed with fold ................. Yakuchloritis hungerfordianus (Moellendorff, 1884) shell high conic; shell height > shell width ................................................................................................. 4 shell depressed conic; shell height < shell width........................................................................................ 7 shell color white or yellowish golden; with/without polymorphic bands ................................................... 5 shell color brown; with a peripheral band ..................................... Satsuma sphaeroconus (Pfeiffer, 1866) shell color yellowish golden ........................................ Satsuma albida mollicula (Pilsbry & Hirase, 1906) shell color white.......................................................................................................................................... 6 base flat; body whorl suddenly widened anterior to peristome in a dorsal view; shell thick ........................



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7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 -

........................................................................................ Satsuma albida insignis (Pilsbry & Hirase, 1906) base convex; body whorl regularly widened; shell thin ..........................Satsuma albida (H. Adams, 1870) shell color yellow......................................................................................................................................... 8 shell color light brown to dark brown........................................................................................................ 10 shell dextral...................................................................................................Pancala bacca (Pfeiffer, 1866) shell sinistral ............................................................................................................................................... 9 umbilicus open, ca. 1/6 of shell width; without band ......................... Satsuma formosensis (Pfeiffer, 1866) umbilicus half covered by reflexed columellar lip, < 1/10 of shell width; with polymorphic bands ........... .......................................................................... Pancala batanica pancala (Schmacker & Boettger, 1891) shell sinistral ............................................................................................................................................ 11 shell dextral .............................................................................................................................................. 12 spire lower; shell height/width ratio < 0.7; umbilicus open .... Satsuma contraria (Pilsbry & Hirase, 1909) spire higher; shell height/width ratio > 0.7; umbilicus half covered by reflexed columellar lip................... ................................................................................................................. Satsuma pekanensis (Rolle, 1911) shell width > 35 mm .................................................................................................................................. 13 shell width < 35 mm ................................................................................................................................. 17 shell large-sized, shell width > 50 mm; shell color dark brown ...Satsuma arisana arisana (Kuroda, 1941) shell medium-sized, shell width 35–45 mm; shell color yellowish brown................................................ 14 outer lip sinuous; inferior lip straight, with a weak fold.......................... Satsuma bairdi (H. Adams, 1866) outer lip smoothly curved; inferior lip smoothly curved, without fold ..................................................... 15 spire lower; shell height/width ratio = 0.67 .............................. Satsuma arisana takahasii (Kuroda, 1941) spire higher; shell height/width ratio = 0.7–0.74 ....................................................................................... 16 umbilicus totally covered by reflexed columellar lip ........................ Satsuma arisana tani (Kuroda, 1941) umbilicus open, not covered by reflexed columellar lip.......................... Satsuma longkiauwensis sp. nov. periphery carinate ...................................................................................................................................... 18 periphery round, at most roundly angulate ................................................................................................ 23 umbilicus not covered by reflexed columellar lip ...................................... Satsuma mellea (Pfeiffer, 1866) umbilicus partly covered by reflexed columellar lip ................................................................................. 19 periphery sharply carinate; surface immediately below periphery concave; inferior lip flat ........................ ........................................................................................................Satsuma friesiana (Moellendorff, 1884) periphery bluntly carinate; surface immediately below periphery convex; inferior lip smoothly curved 20 periphery angulate in front; whorls flat to weakly convex ........................................................................ 21 periphery round in front; whorls convex ................................................................................................... 22 whorls flat; base convex; axial striae dense........................................Satsuma succincta (H. Adams, 1866) whorls convex; base less convex; axial striae loose ......................... Satsuma nux paiwanis (Kuroda, 1941) spire higher; shell height/width ratio = 0.79; peristome very thick; whorls less convex .............................. ............................................................................................. Satsuma succincta rubrotincta (Kuroda, 1941) spire lower; shell height/width ratio = 0.69; peristome not particularly thickened; whorls convex ............. ............................................................................................. Satsuma succincta amblytropis (Pilsbry, 1901) umbilicus close; upper surface finely striated........................ Satsuma arisana takkiriensis (Kuroda, 1941) umbilicus partly covered; upper surface roughly rugous .......................................................................... 24 inferior lip curved; spire lower, shell height/width ratio = 0.66–0.77 . Satsuma litus (Chang & Tada, 2000) inferior lip flat; spire higher, shell height/width ratio > 0.79 .................................................................... 25 sculpture rough; surface of base granular; periphery roundly angulate.. Satsuma nux (Moellendorff, 1888) sculpture weak; surface of base finely striated; periphery round .... Satsuma nux sericata (Kuroda, 1941)


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Acknowledgements We gratefully acknowledge the assistance of Dr. Hsueh-Wen Chang and Hong-Da Zhu (National Sun Yat-Sen University), Dr. Sheng-Hai Wu (National Chung Hsing University), Dr. Hong-Chang Liu (Providence University), Chia-Wei Lin (National Taiwan Ocean University), Chuan-Chin Huang and Chih-Han Chang (both National Taiwan University) in the field works and Chung-Yi Hsiao for photographs. We thank the Institute of Ecology and Biological Resources (IEBR), Vietnam for their hospitality during our collecting visit in Vietnam. This study was partially supported by the National Science Council Grant, ROC (NSC-94-2313-B-00206, NSC-95-2313-B-002-035, NSC-95-2621-B-212-001).

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