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It is traditionally assumed that the relationship between wave steepness and wave age is independent of the wind wave growth state. In fact, the traditional relationship can not describe the whole course of wind wave growth. This paper assumes that the relationship between wave steepness and wave age changes with the variety of dimensionless fetch. Based on the relationship proposed by Hou and Wen (1990), a new relationship in the course of wind wave growth is revealed. Comparisons between the present study and other previous relationships show that this new relationship explains better the observations than the other existing relationships. In the case of small fetch, wave age value increases more quickly than other models while it is in opposition to that in the case of large fetch. The result in present paper can clearly reflect the whole course of wind wave growth, it is an improvement for traditional results.

SATSUMA HAGIOMONTIS SP. NOV. (FIGS 5, 6) Material examined Type specimen: Holotype: TMMT 0604 (from type locality, dry shell, tissue in alcohol). Five paratypes: all from type locality, TMMT 0615 (dry shell, tissue in alcohol, dissected); TMMT 0616 (dry shell, tissue in alcohol); BMNH 20060751, BMNH 20060752, ANSP 413674 (dry shell). Additional material: Museum of Zoology, National Taiwan University (NTUM): two dry shells, collected from Kuwarusu (= Taiwu, Pingtung County, southern Taiwan) in 1918. Type locality M. Beidawushan, Pingtung County, southern Taiwan (Fig. 1, Table 1). Diagnosis Shell and soft body white with grey pedal stripe; periphery keeled; shell height/diameter ratio close to 1.0; distal vagina short; penis corrugated on surface; penial caecum conical; genitalia with or without spinules. Etymology L. hagio: holy, L. montis: mountain. The type locality, Mt Beidawushan, is a holy mountain of two aboriginal tribes, the Paiwan and Rukai. Description Shell (Fig. 5): dextral, conical, thin, fragile, semitranslucent, medium sized. Apex obtuse. Whorls inflated. Peripheries bluntly angulated with keel extended to peristome. Base inflated. Pre-apertural constriction behind outer and inferior lips present. Shell surface smooth, glossy, with spiral striae. Shell colour white milky with thin periostracum. Aperture diagonal, ovate-lunate. Peristome thin, expanded, reflected at curved inferior lip. Superior columellar lip vertical, reflected, covering most of umbilicus. Inferior columellar lip oblique, continual to inferior lip. Band or stain: Polymorphism exists in this taxon. Some individuals exhibit a red-brown to black-purple stain around the umbilicus, columellar, columellar lip, inferior lip and apex, others have spiral colour band around the whorls or at the base. The outer lip exhibits no coloration. Band formula = 1 0 0 0 0 0 7 8; 1 0 0 0 5 0 7 0; 1 0 0 4 0 0 7 0; 1 0 0 4 0 0 7 8. Reproductive system (Fig. 6): Bursa stalk long, with apparently expanded and conical base, regularly tapering towards oval bursa copulatrix. Proximal vagina muscular, swollen, smooth externally, with 14–17 strong, smooth folds internally. Middle vagina less muscular, distinctly constrictive, twisted. Distal vagina short, one-quarter of vagina in length. Flagellum long, tapering, not swollen at base. Pilaster in epiphallus wide, weak, three in number. Penial caecum long, depressed-conical. Cecal pilaster two in number, moderately weak, not strongly corrugated on surfaces. Principal pilaster and proximal penis absent. Middle penis long, twisted, depressed, with seven to nine moderately strong, wiggly, corrugated pilasters inside. Distal penis short, constrictive, with three wide and low pilasters vanishing towards atrium internally. Of the two dissected specimens one (TMMT 0615) has many spinules on the surface of the pilasters in the middle penis. Distribution The species was found only in a narrow area of mid-altitude (1000–1400 m) forest near Mt Beidawushan, Pingtung County, southern Taiwan (Fig. 1, Table 1). Remarks These arboreal herbivores are nocturnal, perching under leaves more than 2 m above ground. Adults were found during summer (Fig. 5I, J). The species differs from S. albida in having a keeled periphery, lower spire with shell height/diameter ratio close to 1.0, short distal vagina; corrugated penis externally, conical penial caecum and spinules inside genitalia. The conical penial caecum and strongly corrugated penis are unique among white-shelled species in west Taiwan., Published as part of Wu, Shu-Ping, Hwang, Chung-Chi & Lin, Yao-Sung, 2008, Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan, pp. 437-493 in Zoological Journal of the Linnean Society 154 (3) on pages 447-449, DOI: 10.1111/j.1096-3642.2008.00415.x, http://zenodo.org/record/5446700

SATSUMA LUTEOLELLASP. NOV. (FIGS 29, 30) Material examined Type specimen: Holotype: TMMT 0611 (from type locality, dry shell, tissue in alcohol). Total 17 paratypes: seven paratypes from type locality: TMMT 0659 (dry shell, tissue in alcohol, dissected); TMMT 0640–0641, TMMT 0678 (dry shell, tissue in alcohol); BMNH 20060757, ANSP 413678, SMF 329387 (dry shell). Ten paratypes from Antong, Hualien: TMMT 0642 (juvenile, in alcohol); TMMT 0639, TMMT 0676– 0677 (dry shell, tissue in alcohol); NMNS 005405-2, BMNH 20060758–20060759, ANSP 413677 (N = 2), SMF 329386 (dry shell). Type locality Fuyuan, Hualien County, eastern Taiwan (Fig. 1, Table 1). Diagnosis Shell and soft body yellowish without pedal stripe; number of whorls more than six; shell highly conical, with shell height/diameter ratio = 1.5. Etymology L. luteola: yellow; L. -ella: diminutive suffix, referring to the small and yellowish shell. Description Shell (Fig. 29): Dextral, thin, conical, medium sized to large. Apex obtuse. Whorls inflated. Periphery bluntly angulated, arc-like. Base inflated. Pre-apertural constriction behind peristome absent. Shell colour yellowish white. Surface even, matt, with fine axial and spiral striae. Periostracum thin. Aperture diagonal, ovate, angulated curve between outer lip and inferior lip. Peristome thin. Columellar lip vertical, reflected covering umbilicus. Umbilicus mostly closed, crevice-like. Band or stain: Polymorphism exists in this taxon. Some individuals do not possess any colour band or stain, whilst some exhibit a red-brown umbilicus; however, the inferior lip and outer lip lack coloration. Others have a peripheral or sub-peripheral band around the whorls. Band formula = 0 0 0 0 0 0 0 0; 0 0 0 0 0 0 7 0; 1 0 0 4 0 0 7 0; 1 0 0 0 5 0 7 0; 1 (0 0 0) 5 0 7 0 (very rare). Reproductive system (Fig. 30): Bursa stalk tapering, not enlarged at base, shorter than spermoviduct. Bursa copulatrix oval to ellipsoid. Proximal vagina muscular, swollen, with 12–17 internal folds; middle vaginas of same width or more constricted, muscular; distal vagina medium in length, one-third length of vagina, same width as middle vagina, constricted towards atrium, with fine wrinkles inside. Flagellum conical, swollen at base with short and digitate tip. Penial caecum short, conical, blunt, furrowed externally; cecal pilasters two in number; remaining inner walls contain 12–14 weak ridges. Proximal penis elongated, muscular, twisted, unevenly furrowed externally, with strong folds internally; principal pilaster long, twice to threetimes length of penial caecum; middle penis constrictive, shortly elongated, with well-defined, strong and corrugated pilasters; distal penis of same strength as middle penis, containing strong, smooth pilasters inside. Spinules present in penial caecum, penis and vagina. Distribution The species was found in lowland broadleaf forest from Liyutan, Mataian, Fuyuan and Antong (Fig. 1, Table 1). It was thus considered to be distributed around the northern parts of the East Rift Valley, eastern Taiwan. Remarks Found from 2 to 3 m above the ground up to the canopy. Mature individuals are found in summer (Fig. 29M, N). Satsuma luteolella is similar to S. pilsbryi (described below) based on morphological characters. The former differs from the latter in having larger shell size, greater number of whorls, paler shell colour, slender base of bursa stalk, long principal pilaster, less expanded proximal penis and lack of a pre-apertural constriction. A dwarf form, which has similar shell dimensions (17.5 mm height, 13.0 mm diameter) to S. pilsbryi, was found in the lowland forest around Antong area, southern Hualien County, eastern Taiwan. They can be easily distinguished based on the characters listed above., Published as part of Wu, Shu-Ping, Hwang, Chung-Chi & Lin, Yao-Sung, 2008, Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan, pp. 437-493 in Zoological Journal of the Linnean Society 154 (3) on pages 475-477, DOI: 10.1111/j.1096-3642.2008.00415.x, http://zenodo.org/record/5446700

SATSUMA VIRIDIBASIS SP. NOV. (FIGS 35, 36) Satsuma albida (H. Adams). Chang et al., 1996: 25–30, fig. 2 (genitalia) [non albida] Material examined Type specimen: Holotype: TMMT 0613 (from type locality, dry shell, tissue in alcohol). Total five paratypes: four paratypes from type locality: TMMT 0663 (dry shell, tissue in alcohol, dissected); BMNH 20060756, ANSP 413676, SMF 329385 (dry shell). One paratype from Yangmingshan, Taipei: TMMT 0675 (dry shell, tissue in alcohol, dissected). Type locality Sikanshui, Xindian, Taipei County, northern Taiwan (Fig. 1, Table 1). Diagnosis Shell and soft body yellowish without pedal stripe; colour of base greenish; pre-apertural constriction present behind outer lip; number of whorls more than six; bursa stalk shorter than spermoviduct; principal pilaster short; proximal penis short, apparently expanded. Etymology L. viridis: green; L. basis: base, referring to the green body whorl. Description Shell (Fig. 35): Dextral, conical, thin, medium sized. Apex obtuse. Whorls inflated. Periphery bluntly angulated, arc-like. Base inflated. Pre-apertural constriction behind outer lip only. Periostracum thin. Shell colour light yellow above periphery, light yellowish green to cyan below periphery. Surface smooth, with fine axial and spiral striae. Aperture subvertical, ovate. Junction between outer lip and inferior lip curved. Peristome thin, expanded at inferior lip and outer lip. Columellar lip vertical, reflected. Umbilicus mostly covered by columellar lip, crevice-like. Band or stain: Band or stain is not present. Band formula = 0 0 0 0 0 0 0 0. Reproductive system (Fig. 36): The reproductive system is similar to that of S. vallis with following exceptions: (1) the flagellum is shorter than that of S. vallis, (2) the epiphallus is short and has three pilasters inside, and (3) spinules are present. Three individuals from the type locality were dissected. Distribution The species was found in lowland ( Remarks Found from 2 m to the canopy. Adult individuals were found in summer (Fig. 35E). The present species is morphologically similar to S. vallis but with larger shell dimensions, greenish shell base, more diagonal aperture, more expanded base, presence of genital spinules and short epiphallus. The specimens dissected and figured by Chang et al. (1996: fig. 2) are actually this present species (see remarks in S. albida). The species is similar to S. luteolella in shell dimensions, but S. viridibasis differs in having greenish shell base, pre-apertural constriction, shorter bursa stalk, short principal pilaster and more expanded proximal penis.

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). 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.

SATSUMA PILSBRYI SP. NOV. (FIGS 37, 38) Material examined Type specimen: Holotype: TMMT 0614 (from type locality, dry shell, tissue in alcohol, dissected). Nine paratypes: all from type locality, TMMT 0644, TMMT 0665–0670 (dry shell, tissue in alcohol); BMNH 20060766, ANSP 413684 (dry shell). Type locality Taiyuan, Taitung County, eastern Taiwan (Fig. 1, Table 1). Diagnosis Shell and soft body yellowish without pedal stripe; shell conical with small diameter; pre-apertural constriction present behind outer lip; base of bursa stalk conical; principal pilaster short; proximal penis short, swollen. Etymology This species is named after the American malacologist Henry A. Pilsbry (1862–1957) who described two species of this group. Description Shell (Fig. 37): Dextral, conical, thin, medium sized to small. Apex obtuse. Whorls inflated. Periphery bluntly angulated, arc-like. Base expanded. Preapertural constriction behind outer lip only. Shell colour light yellow. Periostracum thin. Surface evenly smooth, with fine axial and spiral striae. Aperture diagonal, ovate. Junction between outer lip and inferior lip an angulated curve. Peristome thin, expanded at inferior lip and outer lip. Inferior lip curved downward. Columellar lip vertical, reflected. Umbilicus mostly covered by columellar lip, crevice-like. Junction between columellar lip and inferior lip obtusely angulated. Band or stain: Polymorphism exists in this taxon. Some individuals lack colour band and stain, whilst some exhibit light red-brown apical spot, subperipheral band or umbilicus stain. Inferior and outer lips are colourless. Band formula = 0 0 0 0 0 0 0 0; 10005070. Reproductive system (Fig. 38): Bursa stalk short, fourfifths spermoviduct, tapering, swollen at base. Bursa copulatrix clavated. Proximal vagina muscular, swollen, smooth, with 14 pilasters internally; middle vagina short, constricted, thick-walled, with 17 weak and wiggly folds inside; distal vagina long, nearly one-third length of vagina, gradually constricted towards atrium, with low folds or fine wrinkles inside. Flagellum short and swollen at base, with apparently short and quickly tapering tip. Epiphallus short, with four pilasters inside. Penial caecum moderate, conical, blunt, smooth; cecal pilaster two in number; remaining inner walls of penial caecum contain several moderate ridges. Proximal penis short, swollen, smooth externally, with one deep groove on epiphallic side corresponding to principal pilaster internally; principal pilaster same length as penial caecum; middle penis constricted suddenly, regularly elongated, smooth externally, with seven to nine strong and smooth pilasters; distal penis short, only weak folds or fine wrinkles inside. Spinules present in vagina, penial caecum and proximal penis. Two individuals from the type locality were dissected. Distribution This species was found in lowland forest habitats around the Taiyuan valley, Taitung County, eastern Taiwan (Fig. 1, Table 1). Remarks The perching distance from ground is 2 m; also active on low shrubbery. Adults were found in summer (Fig. 37G). The shell morphology of this species is similar to that of S. vallis, but is differs in dimensions and shell colour. The genital morphology of this species is fundamentally similar to that of S. viridibasis and S. vallis with the following exceptions: (1) the proximal vagina is stronger, (2) the base of the bursa stalk is conical and (3) the penial caecum and proximal penis are strong, smooth and muscular. Satsuma pilsbryi is probably an intraspecific variation from southern populations of S. vallis and S. viridibasis in shell morphology. However, the well-defined geographical isolation and distinguishable reproductive system reveal it merits specific status.

SATSUMA KATIPOLENSISSP. NOV. (FIGS 27, 28) Material examined Type specimen: Holotype: TMMT 0607 (from type locality, dry shell, tissue in alcohol). Eight paratypes: all from type locality, TMMT 0679 (dry shell, tissue in alcohol, dissected); TMMT 0638 (dry shell, tissue in alcohol); NMNS 005405-5, BMNH 20060773– 20060774, ANSP 413688 (N = 2), SMF 329396 (dry shell). Type locality Zhiben, Taitung County, eastern Taiwan (Fig. 1, Table 1). Diagnosis Shell and soft body yellowish without pedal stripe; base of bursa stalk expanded; distal vagina short; principal pilaster short. Etymology From Katipol, the type locality name of the Puyuma aboriginal tribe, today known as Zhiben. Description Shell (Fig. 27): Dextral, extremely thin, fragile, translucent, medium to small. Apex obtuse. Whorls inflated. Periphery bluntly angulated to arc-like. Base expanded. Pre-apertural constriction behind peristome absent. Shell colour light yellow, turning pale white when periostracum peeled away. Surface smooth, with fine axial and spiral striae. Aperture diagonal, ovate-lunate to trapezoid. Junction between outer lip and inferior lip angulated obtusely. Peristome thin, expanded. Inferior lip curved downward. Columellar lip subvertical, slanting to aperture side, reflected covering umbilicus. Umbilicus mostly covered by columellar lip, crevice-like. Junction between columellar lip and inferior lip bluntly angulated. Band or stain: Polymorphism exists in this taxon. Band formula = 0 0 0 0 0 0 0 0; 0 0 0 0 0 0 7 0; 1 0 0 4 0 0 7 0 (Rare). Reproductive system (Fig. 28): Bursa stalk expanded at base, slender at apical half. Proximal vagina muscular, swollen, smooth, with 12–18 internal folds; middle vagina muscular, slender, constrictive, with fine, dense and wiggly folds internally; distal vagina swollen, short, one-fifth length of vagina. Flagellum long, conical, swollen at base, tapering at tip. Epiphallus contains four wide, low pilasters inside. Penial caecum depressed, conical, short; cecal pilaster two in number; remaining inner walls contain 17–25 weak ridges. Proximal penis not particular swollen, with a deep groove externally; principal pilaster medium in length, equal to cecal pilaster; middle penis long, muscular, unevenly furrowed, with five to six strong, wiggly and corrugated pilasters; distal penis smooth externally, with eight moderate and smooth internal folds that reduce gradually as fine wrinkles towards atrium. Spinules present in penis and vagina. Two individuals were dissected. Distribution This species is distributed in the low-altitude (Remarks Inhabiting from 2 to 5 m. Adults were found in autumn (Fig. 27G). This species is morphologically similar to S. huberi and can be distinguished by the following characteristics: (1) the shell is thinner, (2) the shell colour is never ivory-white, (3) the base of the bursa stalk is expanded, (4) the base of the flagellum is more swollen, (5) the vagina is shorter than the penis and (6) the penial caecum is short., Published as part of Wu, Shu-Ping, Hwang, Chung-Chi & Lin, Yao-Sung, 2008, Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan, pp. 437-493 in Zoological Journal of the Linnean Society 154 (3) on pages 473-475, DOI: 10.1111/j.1096-3642.2008.00415.x, http://zenodo.org/record/5446700

Wu, Shu-Ping, Hwang, Chung-Chi, Lin, Yao-Sung (2008): Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan. Zoological Journal of the Linnean Society 154 (3): 437-493, DOI: 10.1111/j.1096-3642.2008.00415.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00415.x

SATSUMA ALBIDA SPECIES COMPLEX Diagnosis: The colour of shell, periostracum and soft body is white to yellowish, never brownish as seen in other Satsuma. The shell is conical with shell height/ shell diameter ratios of 1.0–1.8. Description: Shell dextral, conical, thin to thick, small to medium size (see Table 2 for shell measurements). Shell white or yellowish with or without coloured bands and stains; shell of some species with greenish or golden tint. Bands highly polymorphic, pink, reddish purple or dark purple, variable in width and shape (Fig. 2). Whorls not obviously expanded. Periphery bluntly angulated to keeled. Base expanded or flattened. Surface glossy with curved striae and thin periostracum. Aperture diagonally tetragonal or ovate-lunate. Peristome thin, reflected. Pre-apertural constriction behind outer and inferior lips present on some species. Insertion of columellar lip dilated, not connected with insertion of outer lip. Callus between insertions thin, present only on some species. Umbilicus narrow, half to mostly covered by dilated columellar lip. Colour of soft body in accordance with shell, either white or yellowish. Longitudinal pedal stripe of grey colour present on ventral side of foot in white-coloured individuals. Yellow-shelled species do not exhibit such pedal stripe. The vagina and penis can be divided into three segments, namely proximal, middle and distal parts for convenience of description. Boundaries between parts can be recognized by apparent changes of the external and internal morphologies, such as the strength of parts and the number, shape and strength of internal folds. However, some species showed no apparent boundary between these parts. The verge around the epiphallic pore forms two to three strong pilasters (called caecal pilasters hereafter in this article) tapering to the apex of the penial caecum and usually fused as a single principal pilaster extending distally to the proximal penis. Bursa stalk long with 8–15 smooth and straight internal folds. Free oviduct short. Proximal vagina muscular, swollen, smooth with strong internal folds. Middle vagina constrictive with wiggly and dense internal folds in some species. Distal vagina and atrium villous, dull externally with fine corrugations internally. Flagellum slender or conical. Penis and epiphallus bundled by thin and semi-transparent penial muscle. Proximal penis absent, short or long, with variable morphology. Middle and distal penis thin, slender with strong pilasters to fine wrinkles internally. In some species, a few to numerous variably sized, tiny, flexible and nearly regular tetrahedral spinules were born on the surfaces of pilasters in part of the penial caecum, penis and vagina. A representative figure of the spinules is supplied only for Satsuma mollicula as the morphology of spinules is similar between species under the stereo microscope., Published as part of Wu, Shu-Ping, Hwang, Chung-Chi & Lin, Yao-Sung, 2008, Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan, pp. 437-493 in Zoological Journal of the Linnean Society 154 (3) on page 442, DOI: 10.1111/j.1096-3642.2008.00415.x, http://zenodo.org/record/5446700

Satsuma longkiauwensis sp. nov. (Figs. 2���5) Type material. Type locality: TAIWAN: Pingtung County, Shuangliu, 22 �� 13 ��� 7.3 ���N 120 �� 47 ��� 35.9 ���E, alt. 280 m; TMMT 0 701 (holotype): adult with decaying soft body in ethanol, collected by S.-P. Wu, 20 Oct. 2005; TMMT 0 705 (paratype): adult dissected for genitalia, radula and jaw, remaining soft parts in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0 704 (paratype): sub-adult dissected for radula and jaw, remaining soft parts in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0 706 (paratype): sub-adult soft body in ethanol, collected by S.-P. Wu, 13 Sept. 2005; TMMT 0 707 (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, 22 ��07��� 55.4 ���N 120 �� 47 ��� 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. 2 A���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 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 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. 4 A). Lateral teeth with narrower, outwardly directed mesocone and weak endocone and ectocone (Fig. 4 B). Marginal teeth with more outwardly directed mesocone and prominently forked, pointed endocone and ectocone (Fig. 4 C). 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 (15 th, left); C. Marginal tooth (45 th, left). Scale bar = 20 ��m. 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. 2 E). They were found on ground and crawling on rocks or lower tree trunks. 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., Published as part of Wu, Shu-Ping, Lin, Yao-Sung & Hwang, Chung-Chi, 2007, A new Satsuma species (Pulmonata: Camaenidae) endemic to Taiwan, pp. 59-68 in Zootaxa 1608 on pages 61-65, DOI: 10.5281/zenodo.178845, {"references":["Adams, H. (1866) Descriptions of fifteen new species land and freshwater shells from Formosa, collected by Robert Swinhoe, Esq., consul at Taiwan in that island. Proceedings of the Zoological Society, London, 1866, 316 - 319.","Kuroda, T. (1941) A catalogue of molluscan shells from Taiwan (Formosa), with description of new species. Memoirs of the Faculty of Science and Agriculture, Taihoku Imperial University, 22, 65 - 216.","Moellendorff, O. F. (1888) Mittheilungen aus dem Gebiete der Malakozoologie. Diagnoses specierum novarum sinensium. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 20, 38 - 44.","Pfeiffer, L. (1866) Descriptions of thirteen new species of land-shells from Formosa, in the collection of the late Hugh Cuming, collected by Mr. Robert Swinhoe, vice-consul of that island. Proceedings of the Zoological Society, London [1865], 828 - 831.","Pfeiffer, L. (1845) Diagnosen einiger neuer Heliceen. Zeitschrift fur Malakozoologie, 2, 152 - 158.","Minato, H. (1975) Systematic position of \" Coniglobus \" sphaeroconus (Pfeiffer, 1865) from Formosa (Pulmonata: Camaenidae). Venus, 34, 99 - 104.","Wu, W. K. & Chang, K. M. (1975) Anatomy of the land snail Ganesella (Coniglobus) nux (Moellendorff). Bulletin of Malacology, 2, 107 - 111.","Azuma, M. & Azuma, Y. (1987) Land molluscan fauna of Miyako Islands, the southwestern Okinawa, Japan, with descriptions of new taxa - I. Venus, 46, 159 - 165.","Chang, K. M. (1994) Anatomy of Satsuma bairdi (H. Adams, 1866) from Taiwan (Pulmonata). Venus, 53, 319 - 325.","Chang, K. M. (1997) Anatomy of Coniglobus arisanus (Kuroda) from Taiwan (Pulmonata: Camaenidae). Bulletin of Malacology, 21, 41 - 48.","Schmacker, B. & Boettger, O. (1891) Neue Materialien zur Charakteristik und geographischen Verbreitung chinesischer und japanischer Binnenmollusken II. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 23, 145 - 194.","Tabe, M. (1937) Morphological study of Coniglobus batanicus pancalus (Schmacker & Boettger) and C. b. boteltobagoensis Kuroda and on the systematic position of the genus Coniglobus. Zoological Magazine, 49, 12 - 20.","Sinagawa, K. (1979) Anatomy of Camaena cicatricosa and Pancala bacca and Pancala batanica pancala. Kainakama, 13, 4 - 7.","Chang, K. M. (1992) Dwarf type of Camaena batanica pancala (Schmacker and Boettger) from South Cape, Taiwan and its allied species (Pulmonata: Camaenidae). Bulletin of Malacology, 17, 11 - 20.","Hwang, C. C. (1995) Analysis of mitochondrial DNA phylogeny of Camaena (Pulmonata: Camaenidae). Unpublished Master thesis, National Sun Yat-sen University, Kaohsiung, Taiwan, 101 pp.","Rolle, H. (1911) Diagnosen neuer Arten. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 43, 30 - 32.","Pilsbry, H. A. & Hirase, Y. (1909) New land Mollusca of the Japanese empire. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 586 - 599, [1908].","Hsieh, B. C., Hwang, C. C. & Wu, S. P. (2006) Landsnails of Taiwan. Forestry Bureau Council of Agriculture, Executive Yuan, Taipei, Taiwan, 277 pp.","Chang, K. M. (1989) Anatomy of Coniglobus nux paiwanis (Kuroda) and Coniglobus pekanensis (Rolle) from south Taiwan (Pulmonata: Camaenidae). Bulletin of Malacology, 14, 1 - 8."]}

Satsuma A. Adams, 1868 Type species: Helix japonica Pfeiffer, 1847, by original designation, Published as part of Wu, Shu-Ping, Lin, Yao-Sung & Hwang, Chung-Chi, 2007, A new Satsuma species (Pulmonata: Camaenidae) endemic to Taiwan, pp. 59-68 in Zootaxa 1608 on page 61, DOI: 10.5281/zenodo.178845, {"references":["Adams, A. (1868) On the species of Helicidae found in Japan. Annals and Magazine of Natural History, ser. 4, 1, 459 - 472.","Pfeiffer, L. (1847) Diagnosen neuer Landschnecken. Zeitschrift fur Malakozoologie, 4, 145 - 151."]}