Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan Author Wu, Shu-Ping Author Hwang, Chung-Chi Author Lin, Yao-Sung text Zoological Journal of the Linnean Society 2008 2008-10-31 154 3 437 493 https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00415.x journal article 10.1111/j.1096-3642.2008.00415.x 0024-4082 5446700 AE7FA6D7-DB18-4868-9698-822F799FFCBD SATSUMA CAREOCAECUM SP. NOV. ( FIGS 15 , 16 ) Material examined Type specimen: Holotype : TMMT 0603 (from type locality, dry shell, tissue in alcohol). Seventeen paratypes : all from type locality, TMMT 0651 (dry shell, tissue in alcohol, dissected); TMMT 0622–0624 , TMMT 0646–0650 (dry shell, tissue in alcohol); BMNH 20060769–20060771 , ANSP 413686 ( N = 3), SMF 329393–329394 (dry shell). Figure 15. Satsuma careocaecum sp. nov. A–D, holotype (TMMT 0603, shell height = 19.1 mm). E, F, paratype (TMMT 0622, shell height = 17.8 mm). G, H, paratype (TMMT 0623, shell height = 17.9 mm). I, J, paratype (TMMT 0624, shell height = 17.7 mm). K, living specimen. Arrow indicates the pre-apertural constriction. Figure 16. Reproductive system of Satsuma careocaecum sp. nov. (paratype, TMMT 0651). A, whole genitalia; B, interior of genitalia. C, part of spermatophore showing its position in the female genitalia. D, cross-section of spermatophore at dash line. Scale bar for A–C = 5 mm, for D = 0.5 mm. See text for abbreviations. An abbreviation with a question mark indicates a tentative portion of genitalia. Type locality Guanziling, Tainan County, southern Taiwan ( Fig. 1 , Table 1 ) . Diagnosis Shell and soft body white with grey pedal stripe; base expanded; penial caecum absent. Etymology L. careo : lack; caeca : caecum, referring to the lack of a penial caecum. Description Shell ( Fig. 15 ): Dextral, conical, thin, rigid, medium sized. Apex obtuse. Whorls inflated. Periphery bluntly angulated to arc-like. Base expanded. Pre-apertural constriction behind outer and inferior lips present. Surface sheen with spiral striae. Shell colour white or white milky. Periostracum fine. Aperture diagonal, ovate-lunate. Peristome thin, expanded, reflected. Inferior lip curved downwards. Columellar lip oblique, reflected covering most of umbilicus. Junction of columellar lip and inferior lip roundly angulated. Band or stain: Polymorphism exists in this taxon. Most individuals exhibit a red-brown to black-purple stain around the umbilicus, columellar, columellar lip and apex; a sub-peripheral band is present in some specimens. The outer lip and inferior lip lack such coloration. Band formula = 0 0 0 0 0 0 0 0; 1 0 0 0 0 0 7 0; 1 0 0 0 5 0 0 0; 1 0 0 0 5 0 7 0. Reproductive system ( Fig. 16 ): Bursa stalk long, almost slender. Proximal and middle vagina equally thin without conspicuous boundary between them, smooth externally, bearing 8–12 smooth, irregular strength folds internally; distal vagina one-third length of vagina. Flagellum long, tapering, with slender tip. Epiphallus with four weak pilasters inside. Penial caecum absent. Segment corresponding to penial caecum with three weak pilasters inside; then continued by slender tubule to penis. No verge or apparent internal constriction observed. Penis weak, gradually becoming robust towards atrium; middle penis with five clear, smooth, thin pilasters; distal penis with five strong, irregular, wiggly pilasters. Spermatophore ( Fig. 16C ) found in bursa stalk of figured individual ( TMMT 0651); apical end partly digested in bursa copulatrix (not shown); tail tip remains in proximal vagina; cross-section at middle part ( Fig. 16D ) with three shallow valleculae and a projected fold. Six specimens were dissected. Distribution From Guanziling to Dongshan, northern Tainan County, southern Taiwan ( Fig. 1 , Table 1 ). Remarks The perching distance is typically from 3 m above ground to the canopy. Mature individuals often appear during winter ( Fig. 15K ). The absence of a penial caecum is the most distinct character among species of this species complex. Some Satsuma species , e.g. S. nux paiwanis ( Kuroda, 1941 ) , showed a reduced penial caecum ( Chang, 1989 ), whilst others, e.g. Pancala batanica pancala ( Schmacker & Boettger, 1891 ) and Pancala bacca ( Pfeiffer, 1866 [1865]), showed a lack of a penial caecum with an apparent verge at insertion of the epiphallus ( Chang, 1992 ; Hwang, 1995 ). A totally degenerated penial caecum as seen in S. careocaecum sp. nov. was not recorded in Satsuma . The species differs from S. lini in having a greater number of whorls, apical spot, sub-peripheral band and umbilicus spot; from S. hagiomontis sp. nov. and S. swinhoei sp. nov. in having bluntly angulated periphery; and from S. insignis in having thinner shell and regularly expanded body whorl.