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Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan

November 2008
Zoological Journal of the Linnean Society 154(3):437 - 493

DOI:10.1111/j.1096-3642.2008.00415.x

Authors:

Shu-Ping Wu

University of Taipei, Taiwan

Chung-Chi Hwang

National University of Kaohsiung

The taxonomy of the endemic arboreal snail Satsuma albida species complex from Taiwan was unclear due to the animals' highly similar morphology, and their nocturnal and strict arboreal behaviour, leading to difficulties in collecting living specimens. This article is the first comprehensive comparative study on the systematics and taxonomy of this species complex using external morphology, anatomy of the reproductive system and molecular phylogeny. Consequently, two subspecies of S. albida are raised to species status, namely S. insignis and S. mollicula. Fourteen new species are also described. Fourteen of the 17 species showed polymorphism in banding pattern amongst populations and other species retained the whitish unity as seen in S. albida. Distributions of almost all taxa are geographically limited, with the exception of S. polymorpha sp. nov. The phylogeny of these species was reconstructed using 20 morphological characters and molecular data from the partial sequences of mtDNA CO1 and 16S rRNA genes, and the complete ITS2 sequence. The molecular phylogeny revealed three subclades (west, east and polymorpha clade) and revealed that these snails are monophyletic, originating from a ground-dwelling ancestor. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 437–493.

. Continued Locality Altitude WGS84 coordinate

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. Satsumaalbida species complex showing measurements of holotype/lectotype (italic)

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Satsuma albida. A–D, lectotype (BMNH 1878.1.28.229, shell height = 15 mm). E, living specimen. Arrow indicates the pre-apertural constriction.

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. GenBank accession numbers of taxa

…

+35

Reproductive system of Satsuma albida (TMMT P-0277, Dajin). A, whole genitalia; B, interior of genitalia. Scale bar = 5 mm. See text for abbreviations.

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Systematic revision of the arboreal snail Satsuma

albida species complex (Mollusca: Camaenidae) with

descriptions of 14 new species from Taiwan

SHU-PING WU1, CHUNG-CHI HWANG2* and YAO-SUNG LIN1,3

1Institute of Ecology and Evolutionary Biology, National Taiwan University, 1, Roosevelt Rd, Sec. 4,

Taipei, 10617, Taiwan

2Department of Bioresources, Da-Yeh University, No. 112 Shanjiao Rd, Dacun, Changhua, 51591,

Taiwan

3Department of Life Science, National Taiwan University, 1, Roosevelt Rd, Sec. 4, Taipei, 10617,

Taiwan

Received 21 February 2007; accepted for publication 26 September 2007

The taxonomy of the endemic arboreal snail Satsuma albida species complex from Taiwan was unclear due to the

animals’ highly similar morphology, and their nocturnal and strict arboreal behaviour, leading to difficulties in

collecting living specimens. This article is the ﬁrst comprehensive comparative study on the systematics and

taxonomy of this species complex using external morphology, anatomy of the reproductive system and molecular

phylogeny. Consequently, two subspecies of S. albida are raised to species status, namely S. insignis and S.

mollicula. Fourteen new species are also described. Fourteen of the 17 species showed polymorphism in banding

pattern amongst populations and other species retained the whitish unity as seen in S. albida. Distributions of

almost all taxa are geographically limited, with the exception of S. polymorpha sp. nov. The phylogeny of these

species was reconstructed using 20 morphological characters and molecular data from the partial sequences of

mtDNA CO1 and 16S rRNA genes, and the complete ITS2 sequence. The molecular phylogeny revealed three

subclades (west, east and polymorpha clade) and revealed that these snails are monophyletic, originating from a

2008, 154, 437–493.

ADDITIONAL KEYWORDS: anatomy – phylogeny – Pulmonate – Stylommatophora – taxonomy.

INTRODUCTION

The endemic camaenid arboreal snail Satsuma albida

(Adams, 1870) was ﬁrst collected in the south-west of

Taiwan by R. Swinhoe, a British consul. This whitish

conic snail was described as a new species by Adams

(1870), who provided a concise description and a

distinct hand drawing but without anatomical

description. Two subspecies, S. a. insignis (Pilsbry &

Hirase, 1906 [1905]) and S. a. mollicula (Pilsbry &

Hirase, 1909 [1908]), were described later based on

morphological characters. However, living specimens

are difficult to collect because of their arboreal and

nocturnal behaviour. Consequently, a clear taxonomy

of these highly similar animals has not until now been

provided.

The land snail genus Satsuma A. Adams, 1868 is a

varied group that is distributed throughout East Asia

including Taiwan and Japan (Kuroda, 1958; Wu &

Wu, 1998; Hsieh, Hwang & Wu, 2006). Taiwanese

Satsuma species exhibit strict terrestrial, arboreal or

semi-arboreal behaviour (Hsieh et al., 2006). Owing to

the whitish and highly conical shell, the strict arbo-

real snails have been suggested to be placed within

the genus Ganesella Blandford, 1863 (Pilsbry &

Hirase, 1906 [1905]) or the subgenus Luchuhadra

Kuroda & Habe, 1949 (Kuroda & Habe, 1949; Minato,

1976) together with some species distributed in the

*Corresponding author. E-mail: cchwang@mail.dyu.edu.tw

Zoological Journal of the Linnean Society, 2008, 154, 437–493. With 40 ﬁgures

Ryukyu Islands. Habe (1955) placed Luchuhadra as a

subgenus under Satsuma with deﬁnitive descriptions

of the reproductive system of S.(Luchuhadra)largilli-

erti (Pfeiffer, 1849), the type species of the subgenus.

Minato (1976) reported the reproductive system of S.

(Luchuhadra)amanoi (Kuroda, 1960), which accorded

with Habe’s (1955) viewpoint. However, the reproduc-

tive system of S. albida from northern Taiwan (Chang,

Chang & Hwang, 1996) and S. a. mollicula from

Kenting, southern Taiwan (Sinagawa, 1980), showed a

swelling on the posterior epiphallus and a small ﬂa-

gellum, suggesting that the Taiwanese taxa should be

removed from Luchuhadra (Chang et al., 1996). Fur-

thermore, in line with Pilsbry & Hirase’ (1906 [1905])

view, Chang et al. (1996) doubted the Taiwanese

species belonged to the genus Ganesella, distributed

from China to India, owing to the thin, highly pyrami-

dal and white-coloured shell and angulated periphery.

Nevertheless, their systematic status remains unclear.

The systematic and taxonomic revision of this

group can now be undertaken based on extensive

collection in Taiwan since 1995. Sufficient numbers of

living specimens were obtained for detailed examina-

tions. This article ﬁrst revises the taxonomy of the S.

albida species complex using morphological and ana-

tomical characteristics. The phylogeny of the species

complex is also reconstructed using morphological

and molecular characters to understand the relation-

ships amongst these highly similar species.

MATERIAL AND METHODS

SPECIMEN COLLECTION AND MORPHOLOGICAL DATA

Snails were collected from representative sites in

Taiwan (Fig. 1, Table 1). The collected snails were

anaesthetized using menthol in water, killed in hot

water and then ﬁxed and preserved in 70% ethanol.

The soft bodies of sexually mature specimens, e.g.

with fully expanded lips, were pulled out carefully

and subjected to dissection for examination of the

reproductive system. The dissections were performed

using a stereo microscope (Leica MZ7.5) with a

040 60 Km

120°E 121°E 122°E

25°N

24°N

23°N

22°N

Japan

Ryukyu Is.

Taiwa n

Philippines

China

Jiaoxi

Fushan

Renze

Mt Taipingshan

Shenmihu

Gekou

Liyutan

Mataian

Fuyuan

Ruisui

Antong

Taiyuan

Beinan

Zhiben

Taimali

Ludao Is.

Nanrenshan

Lujiaokeng

Mt Xiangshan

Sikanshui

Pinglin

Fuxing

Baling

Mingchi

Beipu

Youluo

Dananshi

Wula i

Guanziling

Dongshan

Nanhua

Shouka

Shuangliu

Mudan

Fenghuanggu

Shanlinxi

Baolai

Jiaxian

Tengzhi

Shanping

Liugui

Dajin

Qingshan

Mt Dahanshan

Mt Beidawu

Fengshan

Zhongpu

Figure 1. Sampling localities of all taxa in Taiwan. Bold line, Central Range; thin line, administrative boundary.

438 S.-P. WU ET AL.

Table 1. List of the sampling localities showing altitude (metres) and WGS84 coordinate

Locality Altitude WGS84 coordinate

Satsuma albida species complex from Taiwan

Wulai, Taipei 450 24°46′35.1″N, 121°30′11.6″E

Sikanshui, Taipei 280 24°54′03.9″N, 121°34′03.0″E

Pinglin, Taipei 360 24°56′28.7″N, 121°42′51.4″E

Lujiaokeng, Mt Yangmingshan, Taipei 370 25°11′28.4″N, 121°33′50.2″E

Mt Xiangshan, Taipei 120 25°01′38.1″N, 121°34′34.7″E

Jiaoxi, Yilan 30 24°49′44.1″N, 121°45′59.9″E

Fushan, Yilan 640 24°45′29.3″N, 121°35′34.2″E

Mt Taipingshan, Yilan 1600 24°29′43.6″N, 121°32′08.3″E

Renze, Yilan 600 24°32′36.9″N, 121°30′29.0″E

Shenmihu, Yilan 1300 24°22′43.5″N, 121°44′50.4″E

Baling, Taoyuan 690 24°40′43.3″N, 121°23′07.7″E

Mingchi, Taoyuan 1130 24°39′06.5″N, 121°28′18.3″E

Fuxing, Taoyuan 430 24°49′26.2″N, 121°21′19.4″E

Youluo, Hsinchu 430 24°41′07.5″N, 121°08′59.8″E

Beipu, Hsinchu 520 24°39′41.6″N, 121°04′28.9″E

Dananshi, Miaoli 540 24°35′48.7″N, 121°02′37.5″E

Shanlinxi, Nantou 1800 23°38′22.3″N, 120°47′31.7″E

Fenghuanggu, Lugu, Nantou 700 23°43′52.2″N, 120°47′22.2″E

Guanziling, Tainan 320 23°19′43.3″N, 120°30′59.5″E

Nanhua, Tainan 320 23°07′16.8″N, 120°34′07.6″E

Dongshan, Tainan 180 23°16′50.6″N, 120°25′50.7″E

Jiaxian, Kaohsiung 380 23°07′20.8″N, 120°36′59.4″E

Baolai, Kaohsiung 510 23°06′33.1″N, 120°42′10.9″E

Shanping, Kaohsiung 800 22°58′06.4″N, 120°41′10.8″E

Dajin, Kaohsiung 300 22°53′31.5″N, 120°38′50.1″E

Tengzhi, Kaohsiung 1360 23°03′24.9″N, 120°43′15.5″E

Liugui, Kaohsiung 350 22°59′24.3″N, 120°38′36.6″E

Gekou, Taroko National Park, Hualien 180 24°09′28.0″N, 121°37′20.1″E

Liyutan, Hualien 145 23°56′09.8″N, 121°30′42.8″E

Mataian, Hualien 180 23°40′22.0″N, 121°25′13.9″E

Ruisui, Hualien 1150 23°29′40.4″N, 121°22′33.3″E

Antong, Hualien 420 23°17′20.1″N, 121°20′42.7″E

Fuyuan, Hualien 200 23°35′15.4″N, 121°21′12.7″E

Taiyuan, Taitung 580 23°07′17.2″N, 121°16′46.0″E

Beinan, Taitung 1250 22°47′56.2″N, 121°02′13.6″E

Zhiben, Taitung 200 22°42′22.5″N, 121°02′07.0″E

Taimali, Taitung 900 22°36′36.8″N, 120°59′54.6″E

Mt Huoshaoshan, Ludao Is., Taitung 20 22°39′10.6″N, 121°29′03.4″E

Mt Beidawu, Taiwu, Pingtung 1400 23°09′29.4″N, 120°42′29.3″E

Mt Dahanshan, Pingtung 1200 22°24′39.1″N, 120°45′19.8″E

Qingshan, Sandimen, Pingtung 1000 22°48′54.0″N, 120°40′34.1″E

Shuangliu, Pingtung 280 22°13′07.3″N, 120°47′35.9″E

Shouka, Pingtung 450 22°14′35.6″N, 120°50′36.9″E

Mudan, Pingtung 225 22°07′55.4″N, 120°47′18.5″E

Nanrenshan, Kenting National Park, Pingtung 240 22°04′55.3″N, 120°51′08.3″E

Outgroup

Satsuma nux

Zhongpu, Chiayi, Taiwan 340 23°22′44.2″N, 120°32′58.2″E

Satsuma succincta

Fengshan, Kaohsiung, Taiwan 30 22°32′34.3″N, 120°23′29.7″E

Satsuma pekanensis

Beinan, Taitung, Taiwan 1250 22°47′56.2″N, 121°02′13.6″E

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 439

camera lucida attachment for drawing. Shell mea-

surements including shell height, shell diameter (the

greatest breadth) and number of whorls were made

following Kerney & Cameron (1979), using a digital

calliper to an accuracy of 0.1 mm. The type materials

of H. Adams and H. Pilsbry and additional materials

were inspected.

The major portion of the collection was selected as

type materials and general vouchers, and were

deposited in the National Taiwan Museum (TMMT),

the National Museum of Natural Science, Taiwan

(NMNS), the Natural History Museum, London

(BMNH), the Academy of Natural Sciences of Phila-

delphia (ANSP) and the Naturmuseum Senckenberg,

Frankfurt am Main (SMF). Specimens in the Nishi-

nomiya Shell Museum, Japan, and Museum of Zoology,

National Taiwan University, Taiwan (NTUM), were

also examined.

The following abbreviations for genitalia are used in

the ﬁgures: ag, albumen gland; at, atrium; bc, bursa

copulatrix; bs, bursa stalk; dp, distal penis; ep,

epiphallus; ﬂ, ﬂagellum; fo, free oviduct; hd, hermaph-

rodite duct; mp, middle penis; pc, penial caecum; pp,

proximal penis; prm, penial retractor muscle; pv, proxi-

mal vagina, mv, middle vagina, dv, distal vagina; so,

spermoviduct; vd, vas deferens.

The Satsuma albida species complex maintains a

polymorphic banding pattern of varied width, colora-

tion and shapes. For simpliﬁcation, the banding

pattern is coded using eight digits, modiﬁed from the

system used for other taxa (Jones, Leith & Rawlings,

1977; Lai, 1981). Besides the spiral bands on whorls,

additional purple–black stains on the umbilicus, the

lip or the apex are also coded. A number (1–8) indi-

cates that the band is present, whereas a ‘0’ means no

bands or stains are observed. A band formula may be

ranged from 00000000 for a shell of no band to

12345678 for a shell of eight bands and stains. Digits

in parentheses denote a smear displayed between and

connected with the bands or stains. The smears were

usually paler than bands and blurred the margins of

bands. A ‘0’ or zeros in parentheses means a zone of

smear without deﬁnite sharp band or stain.

Digit 1: an apical spot usually covering the embryonic

whorls only, sometimes extended to the anterior part

of the subsequent whorl.

Digit 2: a sub-sutural band that is close to or imme-

diately in contact with the suture.

Digit 3: a supra-peripheral band between the suture

and periphery.

Digit 4: a peripheral band covering the angulated

area of the periphery.

Digit 5: a sub-peripheral band, sometimes contacting

band 4.

Digit 6: a basal band positioned close to the

umbilicus.

Digit 7: an umbilicus spot covering the umbilicus and

columellar lip.

Digit 8: inferior lip stain covering the inferior lip and

area surrounding it.

Three stains or bands were not coded in this study

because of their ambiguity and rarity. (1) A stain on

the outer lip was also observed, but it was pale pink

and unclear. (2) A short and thin band along the

suture, comprising 1/4–1/8 of the body whorl anterior

to the upper insertion of the outer lip was seen

occasionally. The band is covered by or closely con-

tacted with the two bands surrounding the periphery

and is difficult to be coded. (3) The columella is

stained when the apex spot is present for the most

broken shells observed. In some cases, the columellar

stain is absent even when the apical spot is present.

Owing to difficulty in observing the columellar stain,

it is omitted herein.

In order to reconstruct a morphological phylogeny,

potentially phylogenetically informative characters

were selected. Ambiguous (e.g. number of internal

folds in genitalia and length of genital organ) and

highly polymorphic (e.g. banding pattern of shell)

characters were omitted. The selected characters

were solely used for species-level taxonomy. Charac-

Table 1. Continued

Locality Altitude WGS84 coordinate

Satsuma largillierti

Okinawa Island, Japan

Satsuma myomphala

Shimane, Honshu, Japan

Moellendorffia hiraseana 500 22°47′13.6″N, 121°03′0.1″E

Beinan, Taitung, Taiwan

Camaena longsonensis

Langson, N. Vietnam 1155 4°37′24.7″N, 108°36′51.7″E

440 S.-P. WU ET AL.

ter states for continuous variables were determined

by natural gaps of measurements between species.

Twenty morphological characters from shell, soft body

and genitalia were selected as follows:

1. Shell shape (shell height/shell diameter): 0, <0.8;

1, 0.8–1.1; 2, 1.1–1.3; 3, >1.3

2. Periostracum colour of shell spire: 0, brown; 1,

white; 2, yellowish; 3, golden

3. Periostracum colour of shell base: 0, brown; 1,

white; 2, yellowish; 3, golden; 4, greenish

4. Pre-apertural constriction behind inferior lip: 0,

absent; 1, present

5. Pre-apertural constriction behind outer lip: 0,

absent; 1, present

6. Aperture shape (aperture height/aperture diam-

eter): 0, <1; 1, >1

7. Peripheral keel: 0, absent; 1, present

8. Shell size (shell height x shell diameter in mm):

0, <400; 1, >400

9. Soft body colour: 0, dark grey; 1, white; 2,

yellowish

10. Dark pedal stripe: 0, absent; 1, present

11. Relative length of distal vagina to whole vagina:

0,0(=distal vagina absent); 1, 0–0.25; 2, 0.25–0.5

12. Bursa stalk base: 0, tapering smoothly; 1, swollen

or conic

13. Bursa stalk/spermoviduct length ratio: 0, >0.75

(equal length); 1, <0.75 (short bursa stalk)

14. Spinules: 0, absent; 1, present

15. Flagellum base (ratio of maximum ﬂagellum

width/ﬂagellum length): 0, slender (ratio <0.25);

1, swollen (ratio >0.25)

16. Penial caecum: 0, absent; 1, slender; 2, conic

17. Surface of penis near to epiphallus: 0, smooth; 1,

strongly corrugated

18. Deep groove on proximal penis: 0, absent; 1,

present

19. Principal pilaster: 0, absent/short; 1, medium (the

same length as penial caecum); 2, long (longer

than penial caecum)

20. Ratio of penial caecum/epiphallus between penial

retractor muscle and insertion to penial caecum:

0, <0.2; 1, 0.2–2.0; 2, >2.0

MOLECULAR DATA

Whole genomic DNA was extracted from pedal muscle

tissue of fresh, frozen or ethanol-preserved specimens

using the HotSHOT method (Truett et al., 2000).

Fragments of the partial cytochrome oxidase subunit

1 (CO1) and 16S rRNA gene were ampliﬁed using

primer pairs LCO1491/HCO2198 (Folmer et al., 1994)

and 16Sar/16Sbr (Palumbi, 1996). Sequences of the

complete ITS2 gene were ampliﬁed according to Wade

& Mordan (2000). Each 30-mL PCR mixture consisted

of 3 mL10¥reaction buffer containing 15 mMMgCl2,

0.2 mMdNTP, 0.1 units of Taq polymerase (Super-

Therm), 0.5 mM of each primer, 1 mL template DNA

and ddH2O. The conditions for thermal cycling, per-

formed on a Biometra Uno II, was 5 min at 95 °C for

pre-denaturing, 35 cycles of 1 min at 95 °C, 1 min at

50 °C and 1 min at 72 °C, and a ﬁnal extension at

72 °C for 7 min for both the CO1 and the 16S rRNA

genes. The amplicons were examined on a 2% agarose

gel for quality and fragment size, then puriﬁed using

Geneaid PCR Extraction Kit and sequenced on an

ABI 3730 automated sequencer. Chromatographs and

sequences were examined, initially compiled, aligned

and edited in BIOEDIT 7.0.1. (Hall, 1999) and

CLUSTAL X (Thompson et al., 1997).

PHYLOGENETIC ANALYSIS

Congruence between different gene sequences was

tested by performing the incongruence length test

(ILD) (Mickevich & Farris, 1981) with the homogene-

ity partition command of the program PAUP* 4.0b10

(Swofford, 2003). Invariable sites were removed

before applying the test (Cunningham, 1997). The

test was performed by heuristic search with TBR

branch-swapping and 1000 replicates. Gaps were

treated as missing. The best-ﬁtting model for the

aligned sequences of these genes was determined

using a hierarchical likelihood ratio test performed in

the program Modeltest 3.7 (Posada & Crandall, 1998).

The selected substitution model was then adopted in

the reconstruction of phylogeny based on the

neighbour-joining method (NJ) (Saitou & Nei, 1987).

The NJ analysis was conducted in PAUP using the

maximum-likelihood distance. Parsimony analysis

was conducted in PAUP by heuristic search under

TBR branch swapping, random taxa addition of ten

replicates and MulTrees option in effect. Gaps within

the alignment were treated as missing. Support for

nodes was evaluated by bootstrap analysis of 1000

replicates (Felsenstein, 1985). Satsuma (L.)largilli-

erti from Okinawa Island was included for compari-

son, as species of the subgenus Luchuhadra are

thought to be closely related to the S. albida species

complex. Representative species of Satsuma from

Taiwan and Japan were also included, i.e. S. pekan-

ensis (Rolle, 1911), S. succincta (Adams, 1866) and S.

myomphala (Martens, 1865) (Table 1). Camaena long-

sonensis (Morlet, 1891) collected from north Vietnam

and Moellendorffia hiraseana Pilsbry, 1905 were used

as outgroups.

The morphological dataset was analysed by parsi-

mony analysis using a branch-and-bound search with

default settings in PAUP. All characters were treated

as unordered and equally weighted. Multiple states in

the same species were interpreted as polymorphism.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 441

Agreement among equally parsimonious trees was

evaluated by the 50% majority rule consensus.

Support for nodes was tested by bootstrap analysis

(Felsenstein, 1985) with 1000 replicates. Because of

the incomplete anatomical data on most Satsuma

species, two species, S. nux and S. largillierti, were

included for comparisons. Character states for S. nux

were obtained from unpublished data of the present

authors. Characters, especially of the genitalia, of S.

largillierti were compiled from literature researches

(Kuroda & Habe, 1949; Habe, 1955; Azuma, 1995).

Due to the lack of anatomical information for C.

longsonensis, character states for the outgroup genus

Camaena were compiled from the literature, includ-

ing the anatomy of C. cicatricosa (Müller, 1774) and

C. xanthoderma (Moellendorff, 1882) (Pilsbry, 1895;

Sinagawa, 1979; Hwang, 1995). Unavailable

character states were treated as missing, which were

usually genital characters.

RESULTS

SYSTEMATIC DESCRIPTION

FAMILY CAMAENIDAE PILSBRY, 1895

GENUS SATSUMA A. ADAMS, 1868

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 measure-

ments). Shell white or yellowish with or without

coloured bands and stains; shell of some species

with greenish or golden tint. Bands highly polymor-

phic, 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 inser-

tions 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 here-

after in this article) tapering to the apex of the

penial caecum and usually fused as a single princi-

pal 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 vari-

ably sized, tiny, ﬂexible and nearly regular tetrahe-

dral spinules were born on the surfaces of pilasters in

part of the penial caecum, penis and vagina. A rep-

resentative ﬁgure of the spinules is supplied only for

Satsuma mollicula as the morphology of spinules is

similar between species under the stereo microscope.

SATSUMA ALBIDA (H. ADAMS, 1870) (FIGS 3, 4)

Helix (Satsuma)albida H. Adams, 1870: 378, pl. 27,

ﬁg. 9. (Nov., 1870)

Helix albida, Pfeiffer, 1876: 379–380; Moellendorff,

1884: 335–336; Tryon, 1887: 218, pl. 51, ﬁg. 53.

Helix (Satsuma)albida, Schmacker & Boettger, 1891:

159.

Ganesella albida, Pilsbry, 1895: 169; Pilsbry &

Hirase, 1906 [1905]: 736; Kuroda, 1941: 144; Kuroda,

1958: 145.

Satsuma albida, Minato, 1976: 84; Richardson, 1985:

268; Chang et al., 1996: 25–30 [non albida]; Hsieh

et al., 2006: 231.

Coniglobus (Luchuhadra)albida, Chang, 1984: 15, pl.

5, ﬁg. 28.

Satsuma albidum, Lai, 1990: 49. [incorrect gender

ending]

Satsuma (Coniglobus)albida, Ohara & Otani, 2002:

83.

Material examined

Type specimen: Only one specimen originally col-

lected by R. Swinhoe was found and is here desig

nated as the lectotype (BMNH 1878.1.28.229,

442 S.-P. WU ET AL.

Fig. 3A–D), which was part of H. Adams’ collection

purchased by the Natural History Museum, London.

The number of specimens was not mentioned in

Adams’ description, but a later mark on the case,

‘paratype’, indicated that there was probably more

than one specimen examined by H. Adams although

no specimen and designation of types can be traced.

Therefore, we prefer to designate the single speci-

men as lectotype.

Additional material: Nineteen specimens were

collected from Nanhua (N=1), Jiaxian (N=2), Baolai

(N=5), Shanping (N=1) and Dajin (N=10). Three

specimens from Dajin were deposited as vouchers,

TMMT P-0277 (dissected, in alcohol), BMNH

20060782 (shell) and ANSP 413693 (shell).

Type locality

Taiwan, Formosa. The locality ‘Taiwan’ indicated the

‘Taiwan Hsien (County), Taiwan Fu’ on south-west

Formosa at that time (Swinhoe, 1864), which is

equivalent approximately to part of Tainan County

and Kaohsiung County at present. The label accom-

panying with the specimen is marked as ‘Takow,

Formosa’ and is incongruent with H. Adams’ descrip-

tion. Takow (or Takao) located at Qijin, coastal part of

Kaohsiung City nowadays, is less likely to have this

species. We suggest that it is either mislabelled by

Table 2. Satsumaalbida species complex showing measurements of holotype/lectotype (italic)

Species NSH (mm) SD (mm) SH/SD Whorls

S. albida

15.0 14.0 1.07 5.8

15.1 (15.0–15.8) 13.7 (12.8–14.3) 1.11 (1.07–1.23) 5.5 (5.3–5.8)

S. hagiomontis

17.7 16.2 1.09 6.8

15.3 (14.0–17.7) 15.3 (14.0–16.2) 1.01 (0.99–1.09) 5.9 (5.1–6.8)

S. swinhoei

18.5 16.5 1.12 6.3

17.2 (16.3–18.5) 16.2 (14.6–17.0) 1.07 (1.01–1.18) 5.9 (5.8–6.3)

S. insignis

16.8 14.3 1.17 6.0

16.8 (16.1–20.2) 14.8 (12.5–15.6) 1.17 (1.10–1.35) 6.1 (6.0–6.8)

S. lini

17.6 14.9 1.19 5.8

17.3 (14.2–18.6) 14.8 (12.3–15.1) 1.17 (1.15–1.23) 5.8 (5.3–6.0)

S. phoenicis

19.4 16.0 1.21 6.0

18.1 (17.4–19.4) 15.5 (14.3–16.3) 1.19 (1.10–1.25) 5.9 (5.8–6.0)

S. careocaecum

19.1 14.8 1.29 6.6

17.8 (16.3–19.1) 14.6 (13.6–15.6) 1.23 (1.16–1.30) 6.1 (5.6–6.8)

S. polymorpha

14.2 13.9 1.03 5.5

13.9 (11.5–17.1) 13.4 (11.0–16.8) 1.06 (0.99–1.16) 5.0 (4.5–5.9)

S. mollicula

21.2 15.1 1.40 6.5

21.8 (20.4–23.5) 16.3 (15.1–16.7) 1.34 (1.28–1.44) 6.5 (6.5–6.9)

S. huberi

20.3 12.4 1.64 6.8

17.7 (16.5–20.7) 12.2 (11.5–14.5) 1.44 (1.39–1.64) 6.1 (5.8–6.8)

S. auratibasis

20.1 15.8 1.27 6.3

20.0 (17.5–22.2) 14.6 (13.0–15.8) 1.37 (1.27–1.54) 6.5 (5.9–7.0)

S. katipolensis

18.2 12.1 1.50 6.0

16.3 (15.9–18.9) 12.2 (12.0–13.2) 1.36 (1.24–1.50) 5.8 (5.5–6.0)

S. luteolella

23.5 15.3 1.54 6.6

21.0 (16.3–23.5) 13.9 (11.1–15.3) 1.50 (1.40–1.64) 6.5 (5.9–6.9)

S. kanoi

29.7 18.4 1.61 7.3

28.3 (25.9–29.7) 17.1 (15.9–18.6) 1.62 (1.51–1.81) 7.1 (6.9–7.8)

S. vallis

18.5 12.8 1.45 6.1

17.4 (16.0–18.5) 12.7 (12.4–13.2) 1.37 (1.24–1.45) 5.8 (5.5–6.1)

S. viridibasis

22.4 14.6 1.54 6.5

21.4 (20.8–23.3) 14.4 (13.9–15.8) 1.50 (1.44–1.55) 6.3 (6.0–6.5)

S. pilsbryi

17.9 12.3 1.46 5.8

16.9 (14.8–17.9) 12.1 (11.1–13.2) 1.40 (1.28–1.46) 5.9 (5.6–6.1)

SH, shell height; SD, shell diameter; SH/SD, shell height/shell diameter ratio; Whorls, number of whorls; N, sample size

of all type specimens for new species or type(s) and additional specimens measured in this study for the three already

described species, S. albida,S. insignis and S. mollicula; median (bold), minimum and maximum (parentheses).

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 443

1 0 0 0 5 0 7 8

0 0 (0 0) 0 0 0 0 1 (0 0 0) 5 0 7 0 1 0 (0 4) 5 0 7 8

1 0 0 4 0 0 7 0 1 0 0 4 0 0 7 8 1 0 3 0 5 0 7 0 0 0 0 0 0 0 0 0

0 0 0 0 0 6 0 0

0 (0 0) 0 (0) 0 0 0 0 0 0 0 0 0 7 0

1 0 0 0 0 0 7 0 1 0 0 0 0 6 7 0 1 0 0 0 0 6 7 8 1 0 0 0 0 0 7 8

1 0 3 0 5 0 7 8

1 0 0 0 5 6 7 8 1 0 0 0

(

5 6 7

)

8 1 0 0 4 0 6 7 8 0 0 0 0 0 6 7 0

Figure 2. Band formula and colour stain demonstrating polymorphism of banding patterns. Numbers 1–8 show the

position of the band. A ‘0’ means no bands or stains present. Digits in parentheses denoted a smear displayed between

and connected with the bands or stains. See text for deﬁnition of bands and stains in detail.

444 S.-P. WU ET AL.

H. Adams or was used as a convenient way to repre-

sent part of the ‘Taiwan Hsien’ area, like some of

H. Adams’ species (1866, 1870).

Diagnosis

Shell and soft body white with grey pedal stripe;

shell height equal to shell diameter; penis smooth-

surfaced, slender; penial caecum long, slender.

Etymology

L. albidus: white

Description

Shell (Fig. 3):dextral, conical, thin but rigid, medium

to small, higher than wide. Apex obtuse. Whorls

expanded; periphery bluntly angulated. Base inﬂated.

Pre-apertural constriction behind outer and inferior

lips present. Surface smooth, glossy with faint axial

and spiral striae. Shell white or white milky with ﬁne

periostracum. Aperture diagonal, ovate-lunate. Peris-

tome thin, expanded; inferior lip convex, reﬂected;

outer lip expanded at periphery but simple near to

insertion, straight in right-lateral view; superior end

of columellar lip vertical, distinctly dilated, covering

most of umbilicus; inferior end of columellar lip

oblique. Junction of columellar and inferior lips angu-

lated. Umbilicus narrow, covered by dilated columel-

lar lip, left only crevice.

Band or stain: Without any band or stain. Band

formula =00000000.

Reproductive system (Fig. 4):Bursa stalk swollen at

base, shorter than spermoviduct. Bursa copulatrix

oval or clavated. Proximal vagina muscular, gross and

smooth externally, bearing about 15 strong and

wiggly folds internally; middle vagina constrictive;

distal vagina long, one-third to half length of vagina,

ﬁnely wrinkled inside. Flagellum long, tapering, not

swollen at base. Epiphallus with three wide but weak

pilasters inside. Penial caecum long, slender, almost

equal in width until blunt tip. Cecal pilaster two in

number, weak, barely any more prominent than other

pilasters, which number about seven. Principal pilas-

ter absent; proximal penis not identiﬁed; middle penis

long, slender, muscular, approximately same strength

as penial caecum, with six to seven low, well-deﬁned,

wiggly, corrugated pilasters inside; distal penis

slender, same strength as middle penis, bearing weak

and smooth pilasters that reduce gradually as ﬁne

wrinkles towards atrium internally. Walls of penis

and penial caecum smooth, thin, semi-translucent;

internal pilasters visible from outside. Three indi-

viduals dissected, each from Dajin, Baolai and Shan-

ping. Specimen from Baolai with weakly swollen

middle penis. Specimen from Shanping with weakly

swollen distal penis.

Figure 3. Satsuma albida. A–D, lectotype (BMNH 1878.1.28.229, shell height =15 mm). E, living specimen. Arrow

indicates the pre-apertural constriction.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 445

Figure 4. Reproductive system of Satsuma albida (TMMT P-0277, Dajin). A, whole genitalia; B, interior of genitalia.

Scale bar =5 mm. See text for abbreviations.

446 S.-P. WU ET AL.

Distribution

This species was found in south-east Tainan and

northern Kaohsiung County including Nanhua,

Jiaxian, Baolai, Liugui, Shanping and Dajin (Fig. 1,

Table 1).

Remarks

These animals exhibit arboreal and herbivorous

behaviour, perching 2–4 m above the ground

(Fig. 3E).

Because it is hard to distinguish this species from

similar species without examination of specimens, the

reliability of identiﬁcation of most literature sources

(Pfeiffer, 1876; Moellendorff, 1884; Schmacker & Boet-

tger, 1891; Kuroda, 1941; Chang, 1984; Lai, 1990) was

left for further consideration. The specimens dissected

by Chang et al. (1996) were collected from Taipei

County, and have different morphologies on shell and

reproductive characters. Our dissected individual had

a shell height of 19.2 mm, diameter of 14 mm and

height/diameter ratio of 1.37. The illustrated reproduc-

tive system revealed a short ﬂagellum with a strongly

expanded base and digitate tip, a conical penial

caecum, a strongly swollen proximal vagina and an

expanded proximal penis, similar to Satsuma viridiba-

sis described below. The photographed shell in Chang

et al. (1996) has larger shell dimensions (height

17.7 mm, diameter 16 mm, height/diameter ratio 1.11)

than S. albida. This is, in fact, a shell of S. phoenicis

(described below) collected from the type locality of the

species by one of the authors C. C. Hwang. The authors

were not aware of the species diversity in this species

complex.

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).

Figure 5. Satsuma hagiomontis sp. nov. A–D, holotype (TMMT 0604, shell height =17.7 mm). E, F, paratype

(TMMT 0615, shell height =14 mm). G, H, paratype (TMMT 0616, shell height =14 mm). I, J, living specimens. Arrow

indicates the pre-apertural constriction.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 447

Figure 6. Reproductive system of Satsuma hagiomontis sp. nov. (paratype, TMMT 0615). A, whole genitalia;

B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

448 S.-P. WU ET AL.

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 aborigi-

nal tribes, the Paiwan and Rukai.

Description

Shell (Fig. 5):dextral, conical, thin, fragile, semi-

translucent, 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 =10000078;

10005070;10040070;10040078.

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. Flagel-

lum 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 Beidawus-

han, 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.

SATSUMA SWINHOEI SP.NOV.(FIGS 7, 8)

Material examined

Type specimen: Holotype: TMMT 0605 (from type

locality, dry shell, tissue in alcohol). Nine paratypes:

all from type locality, NMNS 005405-1 (dry shell,

tissue in alcohol, dissected); TMMT 0617–0620 (N=4,

dry shell, tissue in alcohol); BMNH 20060754, BMNH

20060755, ANSP 413675, SMF 329384 (dry shell,

tissue in alcohol).

Type locality

Qingshan, Pingtung County, southern Taiwan (Fig. 1,

Table 1).

Diagnosis

Shell and soft body white with grey pedal stripe;

periphery keeled; base of bursa stalk slender; distal

vagina short; penis surface smooth.

Etymology

In memory of the prominent naturalist, Mr Robert

Swinhoe (1836–1877) who discovered the ﬁrst S.

albida.

Description

Shell (Fig. 7):Dextral, conical, thin, rigid, medium

sized. Apex obtuse. Whorls expanded. Periphery

bluntly angulated, keeled. Base inﬂated. Pre-

apertural constriction behind outer and inferior lips

present. Surface smooth, glossy with spiral striae.

Shell colour milky white with ﬁne periostracum.

Aperture diagonal, ovate-lunate to tetragonal. Peris-

tome thin, expanded, reﬂected at curved inferior lip.

Superior columellar lip vertical, reﬂected, covering

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 449

umbilicus mostly. Inferior columellar lip oblique,

obtusely angulated at junction with inferior lip.

Band or stain: Polymorphism exists in this taxon.

Some individuals exhibit a light red-brown to purple

stain around and in the umbilicus. Band formula =

00000000;00040000;00000600;00

000670;10005078;10005678;1000

(567)8;10040678.

Reproductive system (Fig. 8):The genital morphology

of the present species is similar to that of S. hagi-

omontis sp. nov. but with following exceptions: (1)

the base of the bursa stalk is expanded only, not

conical; (2) the two cecal pilasters are weak, merely

stronger than the ridges in the penial caecum and

(3) the distal vagina is swollen and short, one-

seventh the length of the vagina. One individual

was dissected.

Figure 7. Satsuma swinhoei sp. nov. A–D, holotype (TMMT 0605, shell height =18.5 mm). E, F, paratype (BMNH

20060754, shell height =16.3 mm). G, H, paratype (TMMT 0617, shell height =17.2 mm). I, J, paratype (TMMT 0618,

shell height =17.3 mm). K, L, paratype (TMMT 0619, shell height =17.3 mm). M, N, paratype (NMNS 005405-1, shell

height =16.3 mm). O, P, paratype (TMMT 0620, shell height =18.2 mm). Q, R, living specimens. Arrow indicates the

pre-apertural constriction.

450 S.-P. WU ET AL.

Figure 8. Reproductive system of Satsuma swinhoei sp. nov. (paratype, NMNS 005405-1). A, whole genitalia;

B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 451

Distribution

The species was only found in a limited area of

mid-altitude (1000–1200 m) forest near Qingshan

Village, Pingtung County, southern Taiwan (Fig. 1,

Table 1).

Remarks

Inhabits from 2 m above the ground to the canopy.

Adults were found in summer and autumn (Fig. 7Q,

R).

The present species is close to S. albida genetically,

but the shell and genitalia morphologies resemble S.

hagiomontis sp. nov. (see Phylogeny). The species

differs from S. albida in having a keeled periphery,

short distal vagina and larger shell dimensions

(Table 2). Satsuma swinhoei sp. nov. differs from

S. hagiomontis sp. nov. in the opaque shell, straight

inferior columellar lip, tetragonal aperture, angulated

junction between the inferior lip and the columellar

lip, slender base of the bursa stalk and slender penial

caecum.

SATSUMA INSIGNIS (PILSBRY &HIRASE, 1906)

(FIGS 9, 10)

Ganesella albida insignis Pilsbry & Hirase, 1906

[1905]: 736. (15 Jan 1906)

Ganesella albida insignis, Kuroda, 1941: 144, No.

1062.

Coniglobus (Luchuhadra)albida insignis, Chang,

1984: 15.

Satsuma albida insignis, Richardson, 1985: 268;

Hsieh et al., 2006: 232.

Satsuma albidum insignis, Lai, 1990: 49. [incorrect

gender ending]

Material examined

Type specimen: Lectotype: ANSP 89989 (dry shell),

subsequently designated by Baker (1963). One para-

lectotype: ANSP 412186 (dry shell).

Additional material: Five specimens were collected

from Tengzhi, north-east Kaohsiung, southern Taiwan

and deposited as vouchers: TMMT P-0280-1 (dry shell,

tissue in alcohol, dissected); TMMT P-0278–P-0280

(dry shell, tissue in alcohol); BMNH 20060783, ANSP

413694 (dry shell).

Type locality

Hotawa, Taiwan (near Jiaxian, Kaohsiung County,

southern Taiwan at present).

Diagnosis

Shell and soft body white with grey pedal stripe; shell

thick; body whorl suddenly widened anterior to peris-

tome; aperture height smaller than width.

Etymology

L. insignis: distinguished.

Description

Shell (Fig. 9):Dextral, conical, thick, rigid, medium

sized. Apex obtuse. Whorls inﬂated. Periphery bluntly

angulated. Base ﬂat. Pre-apertural constriction

behind outer and inferior lips present. Shell colour

porcelain white. Surface smooth, with axial and spiral

striae. Body whorl suddenly widened anterior to

peristome. Aperture subvertical, trapeziform to

ovate-lunate. Peristome thin. Inferior lip expanded,

reﬂected; outer lip less expanded and reﬂected, in

right-lateral view curved forward; junction between

outer lip and inferior lip angulated acutely. Superior

columellar lip reﬂected. Umbilicus covered by col-

umellar lip mostly, crevice-like. Junction between ver-

tical columellar lip and curved inferior lip angulated.

Band or stain: Polymorphism exists in this taxon.

Most individuals exhibit a black-purple stain around

the columella, columellar lip and inferior lip. The outer

lip is paler in colour. Band formula =00000000;

00000678;10000678;10040078.

Reproductive system (Fig. 10):Bursa stalk long, regu-

larly tapering towards bursa copulatrix. Proximal

vagina moderately muscular, gross, smooth exter-

nally, with 13 wavy folds inside; middle vagina also

muscular, more constrictive than proximal part, with

12 moderate, smooth folds inside; distal vagina one-

third length of vagina. Flagellum long, tapering.

Epiphallus with three wide but weak pilasters inside.

Penial caecum short, tapering with a blunt tip; cecal

pilaster two in number, weak but still prominent

compared with ridges in penial caecum. Principal

pilaster and proximal penis absent; middle penis

moderately swollen, muscular, obviously striated

externally, with equally strong and elevated internal

pilasters six to eight in number; junction between the

middle and distal penis constrictive expeditiously;

distal penis swollen, muscular, with one strong, high

pilaster and four to seven wide but weak pilasters.

Three specimens were dissected.

Distribution

The animals were only found in the Tengzhi area,

north-east Kaohsiung, southern Taiwan (Fig. 1,

Table 1).

Remarks

Inhabits from above 5 m (Fig. 9Q, R). Life span is

probably more than 1 year. Mature individuals were

often observed during winter. The correct date of

publication was emended based on Clench & Turner

(1962). This species differs from S. albida in having a

452 S.-P. WU ET AL.

more ﬂattened base, more angulated periphery, more

protruding body whorl anterior to peristome, thicker

shell, larger shell size and greater number of whorls.

The protruding body whorl and thick shell can differ-

entiate this species from other white-shelled species

in west Taiwan.

SATSUMA LINI SP.NOV.(FIGS 11, 12)

Material examined

Type specimen: Holotype: TMMT 0602 (from type

locality, dry shell, tissue in alcohol, dissected). Three

paratypes: all from type locality, TMMT 0621 (imma-

ture, in alcohol); ANSP A21015 (shell with tissue in

alcohol); BMNH 20060753 (dry shell).

Type locality

Shanlinxi, Nantou County, central Taiwan (Fig. 1,

Table 1)

Diagnosis

Shell and soft body white with grey pedal stripe;

periphery angulated but not keeled; junction between

columellar and inferior lips angulated. Because of a

Figure 9. Satsuma insignis. A–D, lectotype (ANSP 89989, shell height =16.8 mm). E–H, paralectotype (ANSP 412186,

shell height =15.8 mm). I, J, (TMMT P-0278, shell height =16.5 mm). K, L, (TMMT P-0279, shell height =18 mm). M, N,

(BMNH 20060783, shell height =17 mm). O, P, (TMMT P-0280, shell height =20 mm). Q, living specimen. R, copulation.

Arrow indicates the pre-apertural constriction.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 453

Figure 10. Reproductive system of Satsuma insignis (TMMT P-0280-1, Tengzhi). A, whole genitalia; B, interior of

genitalia. Scale bar =5 mm. See text for abbreviations.

454 S.-P. WU ET AL.

lack of knowledge of its genitalia, distinguishing this

species from similar species based on shell character-

istics only is difficult.

Etymology

L. lini: the name is dedicated to the late Professor

Lin, Fei-Jan (1934–2004), a pioneering evolutionary

biologist from Taiwan.

Description

Shell (Fig. 11):Dextral, conical, medium sized. Apex

obtuse. Whorls inﬂated. Periphery bluntly angulated.

Base expanded. Pre-apertural constriction behind

outer and inferior lips present. Surface smooth, glossy

with spiral striae. Shell colour whitish. Periostracum

thin. Aperture diagonal, ovate-lunate, angulated

between outer lip and inferior lip. Peristome thin,

expanded, reﬂected at curved inferior lip. Superior

columellar lip reﬂected. Umbilicus covered by columel-

lar lip mostly, crevice-like. Columellar lip vertical to

subvertical, angulated at junction with inferior lip.

Band or stain: Polymorphism exists in this taxon. The

majority of individuals do not possess band or colour

stain, and display a whitish coloration. A rare form

exists with pink stain and smear from the third to the

fourth band position. Band formula =00000000;

00(00)0000(very rare).

Reproductive system (Fig. 12):No reproductively

mature specimens were collected. An immature geni-

talia is ﬁgured to demonstrate its basic conﬁguration.

Distribution

The species was found only in a narrow area of

mid-altitude forest (1600–1800 m) near Shanlinxi,

Nantou County, central Taiwan (Fig. 1, Table 1).

Remarks

Inhabits typically above 3 m. Adults were found

during winter (Fig. 11G). Only four specimens are

available; two of these have adult shell morphology

but genital development is incomplete. The other

individuals are sub-adults.

Figure 11. Satsuma lini sp. nov. A–D, holotype (TMMT 0602, shell height =17.6 mm). E, F, paratype (TMMT 0621,

shell height =17 mm). G, living specimen. Arrow indicates the pre-apertural constriction.

Figure 12. Reproductive system of Satsuma lini sp.

nov. (holotype, TMMT 0602), immature genitalia. Scale

bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 455

This species is morphologically similar to S. albida,

S. swinhoei sp. nov. and S. insignis, but its genetic

relationship is closer to S. insignis (see Phylogeny).

Compared with S. albida,S. lini has a larger, thinner

and more rigid shell, higher spire and more oblique

aperture. The species differs from S. swinhoei in

having a periphery that is not keeled, smaller shell

diameter and higher spire.

SATSUMA PHOENICIS SP.NOV.(FIGS 13, 14)

Satsuma albida (H. Adams), Chang et al., 1996:

25–30, ﬁg. 1 (shell). [non albida]

Material examined

Type specimen: Holotype: TMMT 0601 (from type

locality, dry shell, tissue in alcohol). Five paratypes:

all from type locality, NMNS 005405-6 (dry shell,

tissue in alcohol, dissected); TMMT 0645 (dry shell,

tissue in alcohol); BMNH 20060772, ANSP 413687,

SMF 329395 (dry shell).

Type locality

Fenghuanggu, Lugu, Nantou County, central Taiwan

(Fig. 1, Table 1).

Diagnosis

Shell and soft body white with grey pedal stripe;

proximal penis present, swollen, grooved; principal

pilaster present, bifurcate, not formed in single strong

pilaster.

Etymology

L. phoenicis: phoenix, translation of the type locality

‘phoenix valley’ (Fenghuanggu).

Description

Shell (Fig. 13):Dextral, conical, medium sized. Apex

obtuse. Whorls expanded. Periphery bluntly angu-

lated, extending to peristome. Base expanded. Pre-

apertural constriction behind outer and inferior lips

present. Surface glossy, with spiral striae. Shell

colour milky white, covered with ﬁne periostracum.

Aperture diagonal, ovate-lunate. Peristome thin,

expanded. Inferior lip reﬂected, curved downward.

Superior columellar lip vertical, reﬂected. Umbilicus

covered by columellar lip, crevice-like. Junction

between oblique inferior columellar lip and inferior

lip smoothly curved.

Band or stain: Band or stain is not present in this

species. Band formula =00000000.

Figure 13. Satsuma phoenicis sp. nov. A–D, holotype (TMMT 0601, shell height =19.4 mm). E, living specimen. Arrow

indicates the pre-apertural constriction.

456 S.-P. WU ET AL.

Figure 14. Reproductive system of Satsuma phoenicis sp. nov. (NMNS 005405-6). A, whole genitalia; B, interior of

genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 457

Reproductive system (Fig. 14):Bursa stalk long, taper-

ing with wiggly folds internally. Proximal vagina

muscular, furrowed externally; middle vagina muscu-

lar, constrictive, with weak folds inside; distal vagina

one-quarter length of vagina. Flagellum long, taper-

ing smoothly towards tip. Penial caecum long, taper-

ing with blunt tip; cecal pilasters two or three in

number, surrounding epiphallic pore. Principal pilas-

ter present, bifurcate, weak, not merged as single

strong pilaster. Proximal penis widened, muscular,

furrowed with a deep groove externally corresponding

to strong, elevated, corrugated internal pilasters;

middle penis short, suddenly constrictive from proxi-

mal penis, with weak and smooth pilasters inside;

distal penis moderately slender, with one to two weak

internal pilasters gradually weakened towards

atrium. Three specimens were dissected.

Distribution

Lugu area, Nantou County, central Taiwan (Fig. 1,

Table 1).

Remarks

Animals are arboreal, herbivores and nocturnal

perching under leaves 2–3 m above ground. Adults

were found during summer and autumn (Fig. 13E).

The swollen and grooved proximal penis and

bifurcate principal pilaster is unique among species

in west Taiwan. The shell of the present species

differs from that of S. albida in its larger size, and

being thinner, translucent and more rigidness;

differs from that of S. lini in larger measurements

(Table 2), more ﬂattened base and inferior lip, more

blunt periphery, more curved downward inferior lip

and, hence, smooth junction between the inferior lip

and the columellar lip. This species is morphologi-

cally similar to S. lini sp. nov., but the genetic

relationship was calculated to be closer to S. care-

ocaecum sp. nov. The photographed shell in Chang

et al. (1996, Fig. 1) belongs to this species (see

remarks in S. albida.)

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.

458 S.-P. WU ET AL.

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 spermato-

phore 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.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 459

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 =00000000;

10000070;10005000;10005070.

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 POLYMORPHA SP.NOV.(FIGS 17–20)

Material examined

Type specimen: Holotype: TMMT 0606 (from type

locality, dry shell, tissue in alcohol). Thirty-four

paratypes: 27 paratypes from type locality: TMMT

0680 (dry shell, tissue in alcohol, dissected); TMMT

0627–0630, TMMT 0632, TMMT 0673–0674 (dry

shell, tissue in alcohol); NMNS 005405-7–005405-11,

BMNH 20060776–20060781, ANSP 413689, ANSP

413690 (N=2), ANSP 413691 (N=2), ANSP 413692,

SMF 329397–329398 (dry shell); two paratypes from

Mataian, Hualien: TMMT 0631, TMMT 0671 (dry

shell, tissue in alcohol); one paratype from Ruisui,

Hualien: TMMT 0633 (dry shell, tissue in alcohol);

two paratypes from Fushan, Yilan: TMMT 0625 (dry

shell, tissue in alcohol), BMNH 20060775 (dry shell);

one paratype from Youluo, Hsinchu: TMMT 0626 (dry

shell, tissue in alcohol); one paratype from Shenmihu,

Yilan: TMMT 0672 (dry shell, tissue in alcohol,

dissected).

Type locality

Beinan, Taitung County, eastern Taiwan (Fig. 1,

Table 1).

Diagnosis

Shell and soft body white with grey pedal stripe; shell

height/diameter ratio close to 1.0; number of whorls

few, ﬁve in most individuals; proximal penis and

460 S.-P. WU ET AL.

principal pilaster distinctly short, weak, almost van-

ishing, absent in individuals from Shenmihu; internal

spinules absent in most individuals, present only in a

specimen from Shenmihu.

Etymology

Gr. polus: much; Gr. morph: shape, referring to the

high polymorphism of banding patterns.

Description

Shell (Figs 17, 18):Dextral, conical, thin, medium

sized. Apex obtuse. Whorls inﬂated. Periphery bluntly

angulated. Base expanded. Pre-apertural constriction

behind outer and inferior lips present. Surface even,

glossy, with faint spiral striae. Shell colour milky

white. Periostracum ﬁne, with polymorphic banding

patterns in most individuals. Aperture diagonal,

ovate-lunate to rectangular. Peristome thin, inﬂated,

Figure 17. Satsuma polymorpha sp. nov. A–D, holotype (TMMT 0606, shell height =14.2 mm). E, F, paratype

(TMMT 0625, shell height =16.7 mm). G, H, paratype (TMMT 0626, shell height =16 mm). I, J, paratype (TMMT 0627,

shell height =14.6 mm). K, L, paratype (TMMT 0628, shell height =14 mm). M, N, paratype (TMMT 0629, shell

height =14.5 mm). O, P, paratype (TMMT 0630, shell height =15.1 mm). Q, R, paratype (TMMT 0631, shell

height =13.8 mm). S, T, paratype (TMMT 0632, shell height =14.9 mm). Arrow indicates the pre-apertural constriction.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 461

Figure 18. Satsuma polymorpha sp. nov. A, B, paratype (TMMT 0633, shell height =17.1 mm). C–I, living specimens.

462 S.-P. WU ET AL.

Figure 19. Reproductive system of Satsuma polymorpha sp. nov. from Beinan, Taitung (paratype, TMMT 0680).

A, whole genitalia; B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 463

Figure 20. Reproductive system of Satsuma polymorpha sp. nov. from Shenmihu, Yilan (paratype, TMMT 0672).

A, whole genitalia; B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

464 S.-P. WU ET AL.

reﬂected at inferior lip. Superior columellar lip

slanted, reﬂected covering most of umbilicus. Junc-

tion between oblique columellar lip and inferior lip

angulated.

Band or stain: The taxon demonstrates the highest

polymorphism of its group. The banding patterns and

stains are varied among individuals. Band formula

=00000000;0(00)0(0)000;00000070;

10000070;10000078;10005078;10

(04)5078;10040070;10040078;103

05078;10305600;10305078.

Reproductive system (Figs 19, 20):Bursa stalk long,

slender, with expanded base. Proximal vagina barely

widened, furrowed externally, with 11–15 strong,

wiggly folds internally; middle vagina varied in

length, not as distinguishable as proximal vagina,

usually muscular sometimes with thick walls, con-

strictive, with ten weak, dense folds inside; distal

vagina long, two-ﬁfths to three-ﬁfths length of vagina.

Flagellum long, tapering, not swollen at base. Pilaster

in epiphallus three in number, wide, low. Penial

caecum long, robust, tapering, with blunt tip; cecal

pilaster two in number, weakly to strongly prominent.

Principal pilaster short, almost vanishing. Proximal

penis short, muscular, twisted, unevenly furrowed

externally, with principal pilaster and eight to ten

strong, corrugated pilasters; middle and distal penis

short, constrictive near atrium, with gradually van-

ishing pilasters inside. Spinules observed only in a

dissected individual from Shenmihu, Yilan (TMMT

0672, Fig. 20). Five individuals from the type locality,

Beinan (Fig. 19), two from Fushan, two from Shen-

mihu and one immature specimen from Ruisui were

dissected. The samples from Shenmihu are different

in having (1) an indistinct middle vagina where only

a short constriction was seen, (2) weak pilasters in

the penial caecum, (3) spinules and in lacking (4) a

proximal penis and principal pilaster (Fig. 20).

Distribution

This is the most widespread taxon of its group,

distributed in mid-altitude (800–1600 m) forest in

east Taiwan (from Shenmihu, Yilan County, to

Taimali, Taitung County) and in north Taiwan of

lower altitude (from Dananshi, Miaoli County, to

Fushan, Yilan County) (Fig. 1, Table 1).

Remarks

Perches 5 m above the ground; at times animals are

active in the tree canopy. Adults were found in

summer. Brood is frequently 20 (Fig. 18C–I).

The shells of northern populations (Miaoli,

Hsinchu, Taoyuan, Taipei and north Yilan County)

are solely white. Regardless of anatomical characters,

they share a common genetic structure with the poly-

morphic populations distributed in eastern Taiwan

(data not shown).

The species has a similar shell height/diameter

ratio to S. hagiomontis sp. nov., but the shell dimen-

sions are smaller than that of the latter species. Most

specimens of this species have a smaller shell and

lower number of whorls than other white-shelled

species in west Taiwan.

SATSUMA MOLLICULA (PILSBRY &HIRASE, 1909)

(FIGS 21, 22)

Ganesella albida mollicula Pilsbry & Hirase, 1909

[1908]: 593–594 (3 March 1909)

Ganesella albida mollicula, Kuroda, 1941: 145,

no. 1062; Sinagawa, 1980: 6–8, ﬁgs 4, 5.

Coniglobus (Luchuhadra)albida mollicula, Chang,

1984: 15.

Satsuma albida mollicula, Richardson, 1985: 268;

Hsieh et al., 2006: 232.

Luchuhadra albida, Chang, 1985: 7. [wrong

identiﬁcation]

Satsuma albidum molliculum, Lai, 1990: 49. [incor-

rect gender ending]

Material examined

Type specimen: Holotype: ANSP 95753 (dry shell),

ﬁxed by monotypy (Baker, 1963).

Additional material: Specimens were collected from

Nanrenshan, eastern Hengchun peninsula, Pingtung

County, southern Taiwan. Four specimens were

deposited as vouchers: TMMT P-0282 (dry shell,

tissue in alcohol, dissected); TMMT P-0281 (dry shell,

tissue in alcohol); BMNH 20060784, ANSP 413695

(dry shell).

Type locality

Toshun, South Cape of Formosa (south-east of Heng-

chun Peninsula, Pingtung County, southern Taiwan

at present).

Diagnosis

Shell colour honey yellowish to greenish gold; soft

body yellowish without pedal stripe; periphery keeled;

number of whorls more than six; base of bursa stalk

conic; bursa stalk shorter than spermoviduct; princi-

pal pilaster long.

Etymology

L. mollis: soft; -cula: diminutive suffix.

Description

Shell (Fig. 21):Dextral, conical, thin, fragile, semi-

translucent, medium sized. Apex obtuse. Whorls

inﬂated. Periphery angulated with keel. Base

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 465

inﬂated. Pre-apertural constriction behind peristome

absent. Shell colour honey yellowish to greenish gold,

turning pale white when periostracum peeled off.

Surface smooth, with ﬁne axial and spiral striae.

Aperture diagonal, ovate-lunate. Peristome thin. Infe-

rior lip and outer lip expanded. Junction between

outer lip and inferior lip curved. Superior columellar

lip slanted towards aperture, reﬂected covering most

of umbilicus. Umbilicus almost closed, only tiny

crevice-like behind columellar lip.

Band or stain: Most individuals have neither band

nor stain; rarely do individuals have red-brown

stain on the columellar and the columellar lip. The

inferior lip, outer lip, apex and whorls do not

have such coloration. Band formula =00000000;

00000070.

Reproductive system (Fig. 22):Bursa stalk short, half

length of spermoviduct, tapering, expanded at basal

part only. Bursa copulatrix clavated. Proximal

vagina muscular, swollen, smooth, with 10–13 inter-

nal folds; middle vaginas of equal strength, muscu-

lar; distal vagina short, one-third to one-quarter

length of vagina, gradually constrictive towards

atrium. Flagellum long, conically swollen at base,

with digitate tip. Pilaster in epiphallus wide, low,

three to four in number. Penial caecum long,

conical, with a blunt tip; cecal pilaster two in

number, strong; in one of the two dissected individu-

als, one cecal pilaster not merged into the principal

pilaster after insertion of epiphallus, simply reduced

as an independent and weak fold instead; remaining

inner walls in penial caecum with eight to ten weak

ridges. Proximal penis long, muscular, twisted,

strongly furrowed. Principal pilaster twice length of

cecal pilasters; middle penis depressed, bent, fur-

rowed externally, with ﬁve to nine strong internal

pilasters reduced gradually as ﬁne corrugations

towards atrium. Spinule present on internal sur-

faces of vagina, penial caecum and penis. Two indi-

viduals were dissected.

Distribution

In south Hengchun Peninsula, Pingtung County,

southern Taiwan (Fig. 1, Table 1).

Remarks

The perching distance from the ground is 2–5 m.

Mature individuals are often observed during

summer (Fig. 21E). The correct date of publication

was emendated based on Clench & Turner (1962).

This species is distinct from S. albida in having a

high spire, yellowish and thin shell, yellowish soft

body without grey pedal stripe, swollen ﬂagellum at

base, conical penial caecum, long principal pilaster,

Figure 21. Satsuma mollicula. A–D, holotype (ANSP 95753, shell height =21.2 mm). E, living specimen.

466 S.-P. WU ET AL.

Figure 22. Reproductive system of S. mollicula (TMMT P-0282). A, whole genitalia; B, interior of genitalia. Scale

bar =5 mm. C, spinules (scale bar =0.5 mm). See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 467

long and corrugated proximal penis, and distinct phy-

logenetic relationship (see Phylogeny), and hence it

is given full species status. The keeled periphery is

unique among the yellow-shelled species in east

Taiwan. The golden tint to the shell, long principal

pilaster, conical base of bursa stalk and short bursa

stalk distinguish this species from other species.

SATSUMA HUBERI SP.NOV.(FIGS 23, 24)

Material examined

Type specimen: Holotype: TMMT 0608 (from type

locality, dry shell, tissue in alcohol). Eleven

paratypes: all from type locality, TMMT 0654 (dry

shell, tissue in alcohol, dissected); TMMT 0634–0635,

TMMT 0652–0653 (dry shell, tissue in alcohol);

NMNS 005405-4, BMNH 20060764–20060765, ANSP

413683 (N=2), SMF 329391 (dry shell).

Type locality

Mt Dahanshan, Pingtung County, southern Taiwan

(Fig. 1, Table 1).

Diagnosis

Shell and soft body yellowish or white without pedal

stripe; shell thick, robust; distal vagina short; princi-

pal pilaster short.

Etymology

The name is dedicated to the late Father Franz Huber

(1913–1994), an Austrian priest who was also a biolo-

gist, in memory of his contribution to biological

science education in Taiwan.

Description

Shell (Fig. 23):Dextral, conical, hard, rigid, medium

sized. Apex obtuse. Whorls expanded. Periphery

curved, barely angulated. Base inﬂated. Pre-apertural

Figure 23. Satsuma huberi sp. nov. A–D, holotype (TMMT 0608, shell height =20.3 mm). E, F, paratype (TMMT 0634,

shell height =20.7 mm). G, H, paratype (TMMT 0635, shell height =19.3 mm). I–K, living specimens.

468 S.-P. WU ET AL.

Figure 24. Reproductive system of Satsuma huberi sp. nov. (paratype, TMMT 0654). A, whole genitalia; B, interior of

genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 469

constriction behind peristome absent. Shell colour

ivory-white to pale yellow. Surface even, with ﬁne

axial and spiral striae. Aperture subvertical, elliptical

to ovate. Junction between outer lip and inferior lip

curved. Peristome thin, expanded. Columellar lip ver-

tical, reﬂected, covering umbilicus. Umbilicus fully

closed.

Band or stain: Polymorphism exists in this taxon.

Most individuals are tinged with pale red-brown

spiral stain extending around the darkly stained

umbilicus. The apex has the same colour as the umbi-

licus. Such coloration is not observed at the inferior

lip and outer lip. Band formula =10000670;

1000(067)0;1(000)0070(very rare);100

40070(very rare).

Reproductive system (Fig. 24):Bursa stalk tapering,

expanded at base, shorter than spermoviduct. Bursa

copulatrix oval. Proximal vagina muscular, swollen,

smooth, with 13–14 internal folds; middle vaginas

constrictive, slender, muscular; distal vagina short,

one-ﬁfth length of vagina, swollen with one major

internal fold in addition to ﬁne corrugations. Flagel-

lum conical, swollen at base, with short and digitate

process. Penial caecum long and conical; cecal pilasters

two to three in number, strong; remaining inner walls

contained 15 weak ridges. Proximal penis long, mus-

cular, twisted, with unevenly ridged surface, internally

with seven strong and corrugated folds; principal

pilaster medium in length, equal to or barely longer

than cecal pilasters; middle penis obviously constric-

tive, smooth externally, with moderate and smooth

internal folds; distal penis of same width as middle

penis, with wide and low pilasters inside. Boundary

between middle penis and distal penis not clear.

Spinule present on pilasters of proximal and middle

penis, not seen in vagina. Three individuals were

dissected.

Distribution

This species was recorded only at Mt Dahanshan,

Pingtung County, southern Taiwan (Fig. 1, Table 1).

Remarks

The perching distance from ground is typically more

than 2 m. Adults can be found in winter and eventu-

ally breed in spring. Brood size is less than 30

(Fig. 23I–K).

The thick and robust shell is unique among the

Satsuma albida species complex. Some specimens of

S. insignis also have thick shells, but never as thick

and robust as shells in this species. The polymorphic

colour of shell, both white and yellow, is also unique.

SATSUMA AURATIBASIS SP.NOV.(FIGS 25, 26)

Material examined

Type specimen: Holotype: TMMT 0609 (from type

locality, dry shell, tissue in alcohol). Fifteen paratypes:

all from type locality, TMMT 0658 (dry shell, tissue

in alcohol, dissected); TMMT 0636–0637, 0655–0657

(dry shell, tissue in alcohol); NMNS 005405-3, BMNH

20060760–20060762, ANSP 413679–413681, SMF

329388–329389 (dry shell).

Type locality

Mudan, Pingtung County, southern Taiwan (Fig. 1,

Table 1).

Diagnosis

Spires and soft body yellowish without pedal stripe;

base and sometimes the entire last whorl pale gold;

number of whorls more than six; bursa stalk shorter

than spermoviduct; base of bursa stalk expanded;

principal pilaster long.

Etymology

L. aurat: golden; L. basis: base, referring to its pale

golden base.

Description

Shell (Fig. 25):Dextral, conical, thin, fragile, semi-

translucent, medium sized. Apex obtuse. Whorls

inﬂated to ﬂat. Periphery roundly curved. Base

inﬂated. Pre-apertural constriction behind peristome

absent. Shell colour on upper whorls light yellow; base

and sometimes the entire last whorl pale gold, turning

white when periostracum peeled away. Shell surface

smooth, with ﬁne axial and spiral striae. Aperture

diagonal, ovate-lunate to trapezoid. Inferior lip curved;

outer lip almost straight. Junction between outer lip

and inferior lip curved. Peristome thin, expanded.

Columellar lip vertical to subvertical, reﬂected cover-

ing umbilicus. Umbilicus mostly covered by columellar

lip, crevice-like. Junction between columellar lip and

inferior lip roundly angulated.

Band or stain: This taxon shows polymorphism. Some

individuals do not exhibit any colour band or stain

whilst others have a red-brown umbilicus and col-

umellar lip; otherwise, the apex, inferior lip and outer

lip lack such coloration. Some individuals have a

short and thin band immediately behind the insertion

of the outer lip. A rare colour pattern exists with

the peripheral band and the apical and umbilicus

spot. Band formula =00000000;10000070;

10040070(rare).

470 S.-P. WU ET AL.

Reproductive system (Fig. 26):The genital morphology

is highly similar to that of S. mollicula and can be

distinguished by (1) the longer bursa stalk of S.

auratibasis, which is three-ﬁfths the length of the

spermoviduct, than that of S. mollicula; and (2) the

longer epiphallus of S. auratibasis, which has a length

almost equivalent to the penis. The length of epiphal-

lus in S. mollicula is three-ﬁfths that of the penis.

Distribution

This species is distributed in lowland (<400 m) broa-

dleaf forest habitats around the north end of Hengchun

peninsula, Pingtung County, southern Taiwan, includ-

ing Mudan, Shuangliu and Shouka (Fig. 1, Table 1).

Remarks

The animals were found from 2 m above ground to the

tree canopy. Adults were found in spring and usually

perched higher than juveniles (Fig. 25I).

Satsuma auratibasis sp. nov. is similar to S.

mollicula based on the morphological characters

of shell and genitalia. Both were found distributed

on Hengchun peninsula, southern Taiwan, but

their distributions do not overlap. The former

species is distributed in the northern part of

the area, the latter in the south (Fig. 1).

Furthermore, molecular phylogeny indicates S.

auratibasis is close to S. huberi rather than S.

mollicula despite the morphological similarities.

Satsuma auratibasis differs from S. mollicula in

that its shell is larger, angulated at the periphery

that is never keeled, pale gold in colour, never

greenish gold as seen in S. mollicula. Moreover, the

golden tint is conﬁned to the base or the last whorl

only of S.auratibasis.S.auratibasis differs morpho-

logically from S. huberi in having gold-tinted shell

base, conical base of bursa stalk, long bursa stalk

and long principal pilaster.

Figure 25. Satsuma auratibasis sp. nov. A–D, holotype (TMMT 0609, shell height =20.1 mm). E, F, paratype (TMMT

0636, shell height =20.8 mm). G, H, paratype (TMMT 0637, shell height =18.4 mm). I, living specimen.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 471

Figure 26. Reproductive system of Satsuma auratibasis sp. nov. (paratype, TMMT 0658). A, whole genitalia;

B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

472 S.-P. WU ET AL.

SATSUMA KATIPOLENSIS SP.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 peris-

tome 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. Peris-

tome 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 =00000000;00000070;

10040070(Rare).

Reproductive system (Fig. 28):Bursa stalk expanded

at base, slender at apical half. Proximal vagina mus-

cular, 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

(<500 m) broadleaf forest habitats around Zhiben,

southern Taitung County (Fig. 1, Table 1).

Figure 27. Satsuma katipolensis sp. nov. A–D, holotype (TMMT 0607, shell height =18.2 mm). E, F, paratype

(TMMT 0638, shell height =16.1 mm). G, living specimen.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 473

Figure 28. Reproductive system of Satsuma katipolensis sp. nov. (paratype, TMMT 0679). A, whole genitalia;

B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

474 S.-P. WU ET AL.

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 character-

istics: (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.

SATSUMA LUTEOLELLA SP.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.

Figure 29. Satsuma luteolella sp. nov. A–D, holotype (TMMT 0611, shell height =23.5 mm). E, F, paratype (TMMT

0639, shell height =21.9 mm). G, H, paratype (TMMT 0640, shell height =20.5 mm). I, J, paratype (TMMT 0641, shell

height =21.7 mm). K, L, paratype (TMMT 0642, shell height =16.3 mm). M, N, living specimens.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 475

Figure 30. Reproductive system of Satsuma luteolella sp. nov. (paratype, TMMT 0659). A, whole genitalia; B, interior

of genitalia. Scale bar =5 mm. See text for abbreviations.

476 S.-P. WU ET AL.

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 diago-

nal, 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 =00000000;

00000070;10040070;10005070;

1(000)5070(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 exter-

nally; cecal pilasters two in number; remaining inner

walls contain 12–14 weak ridges. Proximal penis elon-

gated, muscular, twisted, unevenly furrowed exter-

nally, 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.

SATSUMA KANOI SP.NOV.(FIGS 31, 32)

Ganesella (albida var. ?) kasyotonis Kuroda & Kano in

Kuroda, 1941 (nomen nudum)

Ganesella kasyotonis, Kuroda, 1958 (nomen nudum)

Satsuma albida kasyotonis, Hsieh et al., 2006: 232.

Material examined

Type specimen: Holotype: TMMT 0610 (from type

locality, dry shell, tissue in alcohol). Seven paratypes:

all from type locality, TMMT 0662 (dry shell, tissue in

alcohol, dissected); TMMT 0660–0661 (dry shell,

tissue in alcohol); BMNH 20060767–20060768, ANSP

413685, SMF 329392 (dry shell).

Additional material: Nishinomiya Shell Museum,

Japan NCKG 3310: ﬁve dry, immature shells, col-

lected by T. Kano from Kasyoto Island (Lutao Island)

in 1937.

Type locality

Mt Huoshaoshan, Ludao Island, Taitung County,

eastern Taiwan (Fig. 1, Table 1)

Diagnosis

Shell and soft body yellowish without pedal stripe;

pre-apertural constriction present behind outer lip;

shell large; number of whorls seven.

Etymology

The name is dedicated to Mr Tadao Kano (1906–

1945), a Japanese explorer and naturalist; he devoted

his life to ﬁeld investigations and disappeared in the

rainforest of southern Asia. Mr Kano was the ﬁrst

collector of this species.

Description

Shell (Fig. 31):Dextral, conical, hard, rigid, medium

sized to large. Apex obtuse. Whorls inﬂated. Periph-

ery bluntly angulated to arc-like. Base inﬂated. Pre-

apertural constriction present behind outer lip only.

Shell colour faint yellow. Periostracum thin. Surface

smooth, with ﬁne axial and spiral striae. Aperture

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 477

diagonal, ovate. Junction between outer lip and infe-

rior lip angulated curved. Peristome thin, expanded

at inferior lip and outer lip. Inferior lip curved down-

ward. Columellar lip subvertical, reﬂected. Umbilicus

mostly covered by columellar lip, crevice-like. Junc-

tion between columellar lip and inferior lip obtusely

angulated.

Band or stain: Band or stain is not present. Band

formula =00000000.

Reproductive system (Fig. 32):Bursa stalk long, taper-

ing. Bursa copulatrix clavated. Proximal vagina

muscular, swollen, smooth, with 13–18 internal folds;

middle vaginas constricted, slender, muscular; distal

vagina short, one-twentieth length of vagina, as wide

as or wider than middle vagina, internally with wide

and low folds or ﬁnely corrugated only. Flagellum

short, swollen at base, with suddenly constricted, short

and digitate tip. Penial caecum moderately long,

conical; cecal pilasters two to three in number, strong,

not well merged into principal pilaster; remaining

inner walls contain 8–15 weak ridges. Proximal penis

long, muscular, twisted, unevenly furrowed externally,

with long principal pilaster lengthened twice that of

the cecal pilaster. Proximal penis with 15–22 weak

ridges; middle penis constricted, slender, with six

irregularly strong, corrugated and wiggly pilasters

inside; distal penis of same strength as middle penis,

with ﬁve to six strong and smooth internal pilasters.

Spinules present in penial caecum, proximal penis and

vagina. Two individuals were dissected.

Distribution

The species distributed only in Ludao Island, off

Taitung County, eastern Taiwan (Fig. 1, Table 1).

Remarks

Perching distance from the ground is 2 m to the

canopy. Animals were often found inhabiting banyan

trees (Fig. 31E). Satsuma kanoi has the largest shell

dimensions and number of whorls amongst this

group. Furthermore, it is conﬁned to Ludao Island

and is not sympatric with any species of this group.

This species is morphologically and phylogenetically

close to S. luteolella but differs from the latter in

having pre-apertural constriction, larger shell size,

greater number of whorls and short distal vagina.

SATSUMA VALLIS SP.NOV.(FIGS 33, 34)

Material examined

Type specimen: Holotype: TMMT 0612 (from type

locality, dry shell, tissue in alcohol). Five paratypes:

all from type locality, TMMT 0664 (dry shell, tissue in

Figure 31. Satsuma kanoi sp. nov. A–D, holotype (TMMT 0610, shell height =29.7 mm). E, living specimen. Arrow

indicates the pre-apertural constriction.

478 S.-P. WU ET AL.

Figure 32. Reproductive system of Satsuma kanoi sp. nov. (paratype, TMMT 0662). A, whole genitalia; B, interior of

genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 479

alcohol, dissected); TMMT 0643 (dry shell, tissue in

alcohol); BMNH 20060763, ANSP 413682, SMF

329390 (dry shell).

Type locality

Gekou, Taroko National Park, Hualien County,

eastern Taiwan (Fig. 1, Table 1).

Diagnosis

Shell and soft body yellowish without pedal stripe;

pre-apertural constriction present behind outer lip;

bursa stalk shorter than spermoviduct; spinule

absent; principal pilaster short; proximal penis short,

apparently expanded.

Etymology

L. vallis: a valley, after the type locality, ‘Taroko

valley’.

Description

Shell (Fig. 33):Dextral, conical, thin, semi-

translucent, medium sized to small. Apex obtuse.

Whorls expanded. Periphery bluntly angulated. Base

expanded. Pre-apertural constriction present only

behind outer lip. Shell colour pale yellow with

unevenly distributed white stripes. Periostracum

thin. Surface even, with ﬁne axial and spiral striae.

Aperture diagonal ovate. Junction between outer lip

and inferior lip roundly angulated. Peristome thin,

expanded at inferior lip and outer lip. Inferior

lip curved downward. Columellar lip subvertical,

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.

Although some individuals exhibit neither colour

banding nor staining, others possess a light brown

stain at the umbilicus and the apex. Band formula =

00000000;10000070(rare) (Fig. 2).

Reproductive system (Fig. 34):Bursa stalk short, two-

thirds length of spermoviduct, tapering, not enlarged

at base. Bursa copulatrix clavated. Proximal vagina

short, swollen, with 16–19 folds; middle vagina very

short, not particularly constricted, with 18–20 weak

and wiggly folds inside; distal vagina long, nearly half

length of vagina, same width as proximal vagina,

constricted towards atrium, with low folds or ﬁne

wrinkles inside. Flagellum short, conical, swollen at

base, with short, suddenly reduced and digitate tip.

Epiphallus long, with four pilasters inside. Penial

caecum moderate, conical, blunt, weakly furrowed on

surface; cecal pilaster two in number; remaining

inner walls of penial caecum contain six weak ridges.

Proximal penis short, expanded, weakly furrowed

externally, with one deep groove corresponding to

principal pilaster internally; principal pilaster same

length as penial caecum; middle penis constricted

suddenly, regularly elongated, smooth externally,

with six to eight strong pilasters; distal part short,

with only weak folds or ﬁne wrinkles inside. Spinules

absent. Two individuals from the type locality were

dissected.

Distribution

The species inhabits lowland (<200 m) forest around

the Taroko valley, Hualien County, eastern Taiwan.

The animals were usually found at the Gekou area

Figure 33. Satsuma vallis sp. nov. A–D, holotype (TMMT 0612, shell height =18.5 mm). E, F, paratype (TMMT 0643,

shell height =16 mm). G, living specimen. Arrow indicates the pre-apertural constriction.

480 S.-P. WU ET AL.

Figure 34. Reproductive system of Satsuma vallis sp. nov. (paratype, TMMT 0664). A, whole genitalia; B, interior of

genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 481

and Shenmigu valley of Taroko National Park (Fig. 1,

Table 1).

Remarks

The perching distance from the ground is typically

2–5 m. Active individuals were also observed on low

shrubs. Adults were found in summer (Fig. 33G).

Satsuma vallis is similar to S. katipolensis in shell

dimensions, but it differs from the latter in having

pre-apertural constriction, longer distal vagina,

slender base of bursa stalk, shorter bursa stalk, more

expanded proximal penis and lack of genital spinules.

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 angu-

lated, arc-like. Base inﬂated. Pre-apertural constric-

tion behind outer lip only. Periostracum thin. Shell

colour light yellow above periphery, light yellowish

green to cyan below periphery. Surface smooth, with

Figure 35. Satsuma viridibasis sp. nov. A–D, holotype (TMMT 0613, shell height =22.4 mm). E, living specimen.

Arrow indicates the pre-apertural constriction.

482 S.-P. WU ET AL.

Figure 36. Reproductive system of Satsuma viridibasis sp. nov. (paratype, TMMT 0675). A, whole genitalia;

B, interior of genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 483

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 =00000000.

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 (<500 m) forest

around the Taipei basin, northern Taiwan (Wulai,

Sikanshui, Pinglin, Mt Yangmingshan and Mt. Xiang-

shan) although not in the western part of the basin

(Fig. 1, Table 1).

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 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 con-

striction present behind outer lip; base of bursa stalk

conical; principal pilaster short; proximal penis short,

swollen.

Etymology

This species is named after the American malacolo-

gist 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. Pre-

apertural constriction behind outer lip only. Shell

colour light yellow. Periostracum thin. Surface evenly

Figure 37. Satsuma pilsbryi sp. nov. A–D, holotype (TMMT 0614, shell height =17.9 mm). E, F, paratype (TMMT

0644, shell height =17.6 mm). G, living specimen. Arrow indicates the pre-apertural constriction.

484 S.-P. WU ET AL.

Figure 38. Reproductive system of Satsuma pilsbryi sp. nov. (holotype, TMMT 0614). A, whole genitalia; B, interior

of genitalia. Scale bar =5 mm. See text for abbreviations.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 485

smooth, with ﬁne axial and spiral striae. Aperture

diagonal, ovate. Junction between outer lip and infe-

rior lip an angulated curve. Peristome thin, expanded

at inferior lip and outer lip. Inferior lip curved down-

ward. Columellar lip vertical, reﬂected. Umbilicus

mostly covered by columellar lip, crevice-like. Junc-

tion 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, sub-

peripheral band or umbilicus stain. Inferior and outer

lips are colourless. Band formula =00000000;

10005070.

Reproductive system (Fig. 38):Bursa stalk short, four-

ﬁfths 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 fun-

damentally 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.

PHYLOGENY

MOLECULAR DATA

A total of 484 (S. lini N =4, for each of the remaining

taxa N=30) individuals were successfully analysed in

their CO1 sequences. Sequences of the 16S rRNA gene

were analysed from 3–5 individuals from each of the

species. Sequences of the ITS2 genes were analysed

from one individual from each of the species. The ITS2

gene sequence of Camaena longsonensis was not avail-

able, which was difficult to amplify. Sequences were

veriﬁed by forward and reverse comparisons and were

deposited in GenBank (Table 3). Haplotypes of each

species were monophyletic with high bootstrap support

in a preliminary analysis using the neighbour-joining

method (data not shown). Moreover, no haplotypes

were shared between species. Therefore, only one

haplotype was selected as a representative for each of

the species and for the subsequent phylogenetic analy-

ses. Two segments of 21 and 49 bp in the 16S sequences

were excluded owing to highly variable and ambiguous

alignment. Atotal of 1506 bp of aligned sequences were

used in the following analyses containing 638 bp of

CO1, 362 bp of 16S rRNA and 506 bp of ITS2.

The ILD test for congruence between datasets was

marginally signiﬁcant (P=0.025). Cunningham (1997)

indicated that combining datasets with an ILD test P

value of greater than 0.01 retained or improved phy-

logenetic accuracy. Therefore, the sequences of the

three genes were combined for the subsequent analy-

ses. The combined dataset provided higher bootstrap

support and greater resolution than either dataset

alone. Of the characters, 609 bp (40%) were variable,

and 417 bp were parsimony informative. For the 23

species analysed, the heuristic searches resulted in

two trees of equal maximum parsimony (MP) with a

tree length of 1817 (CI =0.53, RI =0.59). All branches

were sufficiently (>50%) supported by bootstrap analy-

sis except for the (S. viridibasis,S. vallis,S. pilsbryi)

clade.

The general time-reversible model with a pro-

portion of invariable sites and a gamma-shaped

distribution of rates across sites (TvM +I+G) was

determined as the best-ﬁtting model for the combined

sequences of the three genes using the hierarchical

likelihood ratio test performed by the program Mod-

eltest 3.7 (Posada & Crandall, 1998). The model was

then adopted for the maximum-likelihood distance

486 S.-P. WU ET AL.

setting for NJ analysis. The topology of the NJ tree

was similar to the MP tree. Satsuma myomphala

from Honshu, Japan, diverged from species in

Ryukyu and Taiwan early. Satsuma largillierti from

Okinawa Island formed a separate clade from this

species complex and clustered with S. pekanensis and

S. succincta. The phylogeny of S. katipolensis was not

congruent in the MP and NJ trees. This species was

clustered with S. huberi and S. auratibasis in the NJ

tree, whereas it was closer to other species from

eastern Taiwan in the MP tree but with low bootstrap

value. The deep branches were not well resolved in

the NJ tree where levels of bootstrap support were

low. The 50% majority rule consensus tree (tree

length 1817, CI =0.53, RI =0.59) of the two MP trees

and the NJ tree was reconstructed by weighting each

of the MP trees as 0.5 and the NJ tree as 1 (Fig. 39).

MORPHOLOGICAL DATA

The parsimonious analysis on 20 morphological char-

acters (Table 4) resulted in 84 MP trees of 68 steps.

The 50% majority rule consensus (Fig. 40) produced a

tree of 70 steps (CI =0.59, RI =0.78, RC =0.46) with

high bootstrap support for most branchs. Bootstrap

analysis showed sufficient support at deep branches

and the clade (mollicula,auratibasis) only (Fig. 40).

Satsuma largillierti from Okinawa Island formed a

separate clade from this species complex and agreed

with molecular phylogeny data. However, it was not

clustered with S. nux. The 17 species of the S. albida

species complex from Taiwan formed a monophyly.

However, four species from west Taiwan, insignis,

lini,phoenicis and careocaecum, could not be sepa-

rated based on these characters. Satsuma polymor-

pha does not form an ancestral clade of species in

west Taiwan. The clustering of sister species in this

tree, e.g. (huberi,katipolensis) and (mollicula,aurati-

basis), agreed with the general impression but was

incongruent with molecular phylogeny data (Fig. 39).

The cladograms based on molecular and morphologi-

cal data (Figs 39, 40) presented three major clades

from Taiwanese species, as judged by the topology,

branch lengths and geographical distribution: an ‘east

clade’ (S. vallis,S. pilsbryi,S. viridibasis,S. kanoi,S.

luteolella,S. mollicula,S. katipolensis,S. huberi and

S. auratibasis), a ‘west clade’ (S. albida,S. swinhoei,S.

hagiomontis,S. insignis,S. lini,S. careocaecum and S.

Table 3. GenBank accession numbers of taxa

Species

GenBank accession number

CO1 16S rRNA ITS2

Satsuma albida EF057384 EF057354 EF204853

Satsuma insignis EF057382 EF057352 EF204863

Satsuma mollicula EF057374 EF057363 EF204859

Satsuma hagiomontis sp. nov. EF057386 EF057356 EF204865

Satsuma swinhoei sp. nov. EF057385 EF057355 EF204866

Satsuma lini sp. nov. EF057383 EF057353 EF204864

Satsuma phoenicis sp. nov. EF057381 EF057350 EF204852

Satsuma careocaecum sp. nov. EF057380 EF057351 EF204851

Satsuma polymorpha sp. nov. EF057378 EF057357 EF204862

Satsuma huberi sp. nov. EF057375 EF057364 EF204858

Satsuma auratibasis sp. nov. EF057376 EF057365 EF204860

Satsuma katipolensis sp. nov. EF057377 EF057366 EF204854

Satsuma luteolella sp. nov. EF057373 EF057362 EF204856

Satsuma kanoi sp. nov. EF057372 EF057361 EF204861

Satsuma viridibasis sp. nov. EF057371 EF057358 EF204850

Satsuma vallis sp. nov. EF057369 EF057360 EF204855

Satsuma pilsbryi sp. nov. EF057370 EF057359 EF204857

Satsuma nux EF057337 EF204778 EF204872

Satsuma arisana takkiriensis EF204828 EF204791 EF204885

Satsuma succincta EF204839 EF204802 EF204896

Satsuma pekanensis EF204833 EF204796 EF204890

Satsuma largillierti EF057387 EF057367 EF204904

Satsuma myomphala EU131663 EU131664 EU131665

Moellendorffia hiraseana EF204842 EF204805 EF204899

Camaena longsonensis EF057379 EF057368

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 487

phoenicis) and the ‘polymorpha clade’ (S. polymorpha).

High genetic diversity was observed amongst the dif-

ferent clades, with sufficient statistical support using

molecular data as shown by the branch length in

Figure 39. Nevertheless, genetic divergence among

each of the clades was different. Taxa of the east clade

were estimated to have lower genetic distances

between each other compared with species of the west

clade. The polymorpha clade is a sister taxon of the

west clade containing only one species, although the

morphological phylogeny (Fig. 40) did not agree with

the separate status of this clade. This species is unique

in having highly polymorphic banding patterns and

the most widespread geographical distribution in

eastern and northern Taiwan with low genetic diver-

gences (data not shown).

DISCUSSION

This study represents the ﬁrst comparative studies of

S. albida and its allied species using conchological,

anatomical and molecular characteristics. Taxonomi-

cal decisions, which were inconclusive in the past

based on morphological characters only (Kuroda,

1941; Kuroda, 1958; Sinagawa, 1980; Chang et al.,

1996), were made accordingly and included the erec-

tion of 14 new species, raising the taxonomic status of

two subspecies and the correction of previous identi-

ﬁcation. Raising two previous subspecies of S. albida,

S. insignis and S. mollicula, to full species was

suggested as they belonged to different clades

on the phylogenetic tree in addition to morphological

characters. Morphological evidence demonstrated

that the reproductive system of ‘S. albida’ ﬁgured by

Chang et al. (1996) was in fact that of S. viridibasis

sp. nov. based on similarities in their anatomical

morphology and close geographical distribution.

The general morphology of the reproductive system,

e.g. the presence of ﬂagellum and penial caecum, in the

S. albida species complex agrees with the known

structures of other Satsuma species (Kuroda & Habe,

1949; Habe, 1955; Minato, 1976; Chang, 1992).

However, an exception was observed in S. careocae-

cum, which showed a lack of the penial caecum. The

presence/absence of the penial caecum (=penial appen-

dix) was considered as a diagnostic character to

distinguish related camaenid (sub-)genera. That is,

Satsuma (including subgenus Satsuma s.s.,Luchu-

hadra and Coniglobus Pilsbry & Hirase, 1906 [1905])

presented a penial caecum of variable size, whereas

Pancala Kuroda & Habe, 1949 and Camaena Albers,

1850 and Camaena (Miyakoia) Minato, 1980 showed a

lack of penial caecum (Kuroda & Habe, 1949; Minato,

1980; Chang, 1992). Therefore, Pancala and Miyakoia

were assigned as subgenera of Camaena (Kuroda &

Habe, 1949; Minato, 1980). The species of Camaena

Table 4. Data matrix for phylogenetic analysis of Satsuma albida species complex. See text for character coding and

further explanation

11111 111112

1 2 3 456 78901234 567890

S. albida 2 1 1 111 00112100 010002

S. hagiomontis 1 1 1 110 10111100/1021001

S. swinhoei 1/21 1 110/110111000 010002

S. insignis 2 1 1 110 00112000 011001

S. lini 2 1 1 111 0011???? ??????

S. phoenicis 2 1 1 111 00112000 011112

S. careocaecum 2 1 1 110/100112000 000000

S. polymorpha 1 1 1 110/100112100/1011002

S. mollicula 3 3 3 001 10202111 121121

S. huberi 3 1/21/2001 00201001 121112

S. auratibasis 3 2 3 001 00202111 121121

S. katipolensis 3 2 2 001 00201101 121112

S. luteolella 3 2 2 001 00202001 121121

S. kanoi 3 2 2 011 01201001 121121

S. vallis 3 2 2 011 00202010 120111

S. viridibasis 3 2 4 011 00202011 120112

S. pilsbryi 3 2 2 011 00202101 120112

S. nux 0 0 0 000 01000110 120000

S. largillierti 1 2 2 000 0100?1? ? 0110? 2

Camaena 0 0 0 000 01000120 000000

488 S.-P. WU ET AL.

showed an elongated and conic penial verge at the

insertion of the epiphallus into the penis instead of a

protruding penial caecum (Pilsbry, 1895; Sinagawa,

1979; Hwang, 1995). The species of the genus Pancala,

Satsuma succincta,S. nux paiwanis and S. longkiau-

wensis Wu, Lin & Hwang, 2007 showed a weak and

hemispherical verge only, neither conical nor elongated

as a penial caecum (unpublished data).

The lack of a penial caecum observed in S. care-

ocaecum is unique not only in this species complex

but also in the genus Satsuma. No verge or similar

structure was observed in this species; instead,

there is only a constrictive tubule between the

epiphallus and the penis. With regard to develop-

ment, the lack of a penial caecum in S. careocaecum

may be considered as a pedomorphosis as all of the

subadult specimens dissected in the present study

showed the same absence (see the immature geni-

talia of a juvenile S. lini, Fig. 12). Anatomical evi-

dence showed that characters of the male genitalia

are better in distinguishing between species than

female gentialia, which agrees with Kameda, Kawa-

kita & Kato (2007). The characters of the ﬂagellum,

and portions at the insertion of the epiphallus,

which include the penial caecum, proximal penis,

internal pilasters and verge, are useful in taxonomy

at the speciﬁc level. Due to the lack of internal

morphology of genitalia for most Satsuma species,

the phylogenetic relationship based on morphologi-

cal characters presented in here is tentative. A com-

plete phylogenetic analysis is required to clarify

their evolutionary signiﬁcance.

Spinules of the genitalia were observed mainly in

species of the east clade, except S. vallis which does

not bear any spinules. The reverse was observed in

the other two clades, members of which usually do not

have spinules in the genitalia. Spinules were born in

genitalia of some individuals of S. hagiomontis,

S. swinhoei and S. polymorpha from Shenmihu. Occa-

sionally, female and male genitalia of the same indi-

0.1

S. lini

S. vallis

S. pilsbryi

S. viridibasis

S. kanoi

S. luteolella

S. mollicula

S. katipolensis

S. huberi

S. auratibasis

S. albida

S. swinhoei

S. hagiomontis

S. insignis

S. careocaecum

S. phoenicis

S. polymorpha

S. largillierti

C. longsonensis

S. succincta

S. pekanensis

S. myomphala

M. hiraseana

EAST

WEST

POLYMORPHA

87/59

68/*

*/53

93/82

94/69

71/68

100/99

100/100

94/100

90/97

64/70

99/94

59/*

81/*

*/*

90/76

90/75

98/99

86/85

93/100

Figure 39. The phylogenetic relationships of the Satsuma albida species complex from Taiwan based on the tree

reconstructed by the maximum-parsimonious method on a combined dataset of the partial CO1, 16S rRNA and complete

ITS2 genes. East, West and Polymorpha indicate the three major clades of the group in Taiwan. Numbers beside the nodes

refer to the percentage bootstrap support of 1000 replicates based on the maximum-parsimonious method (left) and the

neighbour-joining tree (right). An asterisk indicates bootstrap value of less than 50%.

SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 489

vidual do not have spinules simultaneously. Penial

armatures were observed in some Westraltrachia and

related genera in Australia (Solem, 1984), which

showed a hardened edge on the principal pilaster.

Hooked penial armatures were observed in several

species of the family Streptaxidae (Verdcourt, 2000).

However, a similar structure was not reported in

Satsuma and other East Asian camaenids partly

because of few reports on interior genitalia. The func-

tion and phylogenetic signiﬁcance of such penial

armature in the S. albida group remain unknown.

It was suggested to function partly as stimulator or

hold-fast surface during copulation (Gómez, 2001).

However, Solem (1984) doubted that the armatures

may be seasonally variable.

Three clades of distinct genetic characters were

found among the Taiwanese arboreal species (Fig. 39),

which agreed partly with the morphological phylog-

eny based on characters of the shell, soft body and

reproductive system (Fig. 40). The species of the east

clade uniformly have more conical and yellowish

shells, a yellowish soft body and stout male genitalia.

The species of the west and polymorpha clades have

less conical and white shells, white/grey soft body

with dark grey stripes on the foot and slender male

genitalia.

The nine species clustered in the east clade have

conspicuous morphological differences in the shell

and reproductive system. Furthermore, no common

haplotypes derived from mtDNA CO1 and 16S rRNA

gene sequences were shared between each of these

species. Therefore, no gene ﬂow is revealed between

the species and no sign of hybridization. However,

these species share short genetic distances (uncor-

rected proportional distance =0.006–0.052). Species

within this clade are considered to have arisen only

recently with prompt divergence that may have been

caused by geographical barriers and rapid speciation

via signiﬁcant morphological shifts to aspects of body

form and reproductive system in east Taiwan.

The west clade comprises seven species showing

high genetic divergence between the taxa (uncor-

rected proportional distance =0.032–0.15) although

their shell morphologies are similar to each other.

Long branch lengths on the phylogenetic tree imply

that these species may have diverged a long time ago.

S. lini

S. vallis

S. pilsbryi

S. viridibasis

S. kanoi

S. luteolella

S. mollicula

S. katipolensis

S. huberi

S. auratibasis

S. albida

S. swinhoei

S. hagiomontis

S. insignis

S. careocaecum

S. phoenicis

S. polymorpha

S. largillierti

Camaena

S. nux

100

92 88

Figure 40. Fifty per cent majority rule consensus tree of 84 maximum-parsimonious trees based on 20 morphological

characters of the Satsuma albida species complex from Taiwan. Tree length: 70, CI: 0.59, RI: 0.78, RC: 0.46. Numbers

above the branches are the percentage of the consensus tree, and those below are the bootstrap percentage of 1000

replicates. Bootstrap values of less than 50% were omitted.

490 S.-P. WU ET AL.

Furthermore, the seven species do not share haplo-

types from the molecular survey. All taxa have a

distinctive reproductive system that suggests repro-

ductive isolation is well established among species of

the west clade. Most species from this clade are

conﬁned to a narrow geographical range and altitude

limitation, and are dependent on primary broadleaf

forest. Thus, species formation of this early clade

arisen is considered to have resulted from dispersal or

vicariance events through a long evolutionary process

(Avise, 2000). The ﬁxed genetic structure and habitat

limitation is probably derived from habitat fractures

caused by the repetitive expansion and retreat of the

forests during glacial periods of the past million years

on Taiwan (Lin, 1970; Chai, 1972; Teng, 1990). The

similarity in shell morphology is probable due to

similar selective forces on ecology,

Satsuma polymorpha, despite being the most wide-

spread species in eastern and northern Taiwan and

having distinctly divergent banding pattern, does not

diverge into different species between populations.

Eastern populations of S. polymorpha are distributed

over a wide geographical range of mid-altitude forests

(south Yilan to Taitung) with high polymorphism

in banding pattern, whereas northern populations

(north Yilan and Taipei, Taoyuan, Hsinchu and

Miaoli) do not display such polymorphism, i.e. all

shells exhibit a whitish colour as seen in S. albida.

However, a similar genetic structure and haplotypes

are shared between populations (data not shown).

Reasons for the asymmetric evolution in the morpho-

logical and genetic variability of S. polymorpha from

the eastern and northern populations remain uncer-

tain. The molecular phylogenetic relationship sug-

gests that S. polymorpha arose at the same time as

the west clade, but no further divergence to species

level as seen in the west clade has taken place.

The Luchuhadra species in the Ryukyu Islands

formed a separate clade with other ground-dwelling

Satsuma in Taiwan showing that the major clades of

these arboreal species evolved independently after

the divergence from their common ancestor. This dis-

agrees with the suggestion of assigning the arboreal

S. albida group to Luchuhadra (Kuroda & Habe,

1949; Habe, 1955; Minato, 1976). Further phyloge-

netic investigations are required for a better resolu-

tion of the evolutionary history of the genus Satsuma,

including its nominal subgenera Coniglobus and

Luchuhadra, and its allies from East Asia. Therefore,

no generic determinations are proposed herein and

the genus Satsuma (including species in Satsuma s.s.,

Luchuhadra and Coniglobus) is suggested to be

applied tentatively to these arboreal snails.

Despite the controversies mentioned in the Intro-

duction on the systematics of these arboreal snails,

we consider that these three clades of Taiwanese

arboreal camaenids have arisen from their common

ancestor through the evolutionary process of specia-

tion in their areas of distribution. The pyramidal

shell, pale coloration, highly polymorphic banding

patterns and stains should be adaptive consequences

of the arboreal process from their dark-coloured,

depressed-conical and less polymorphic Satsuma-like

ground ancestor. Similar selective forces on arboreal

ecology during the adaptive evolution are considered

to have maintained the phenotypic coincidence of the

three groups. Morphological similarities of these arbo-

real camaenid snails illustrate apomorphy of these

taxa as compared with their ground-dwelling ances-

tor. Therefore, the three groups show similar colour

pattern and shell morphology but are located in dif-

ferent clades. The use of shell morphology and col-

oration may not be appropriate for taxonomy of this

group of animals. Anatomical study of the reproduc-

tive system and molecular analysis provide should be

used to clarify a group of organisms such as these.

ACKNOWLEDGEMENTS

We would like to thank Dr Hong-Chang Liu (Provi-

dence University) for samples from Okinawa Island,

the Institute of Ecology and Biological Resources

(IEBR) of Vietnam for samples from Vietnam, Dr

Si-Long Chen (Taipei Zoo) for some samples of S. kanoi

sp. nov., Chung-Yi Hsiao and Po-Feng Lee for photo-

graphs, and Hui-Ming Huang, Chi-Ying Kao, Jing-

Rong Lai, Chuan-Chin Huang, Chih-Han Chang, Wen-

Shin Lin, Chin-Feng Lin, Bin-Ling Shih, Chang-Yi

Tsai, Jing-Kuan Yang, Kuo-Chiang Liu and several

anonymous helpers for assistance in the ﬁeld. We

thank Dr Hsueh-Wen Chang (National Sun Yat-sen

University) for the loan of the microscope. We greatly

appreciate the friendly cooperation of the curators of

the following museums for help in examination and

deposition of material: Paul Callomon (ANSP), Fred

Naggs and the staff of Higher Invertebrates Section

(BMNH), Dr Ronald Janssen (SMF), Yoko Otani and

Kenji Ohara (Nishinomiya Shell Museum), Dr Jiann-

Neng Fang, Chun-Yan Sun (TMMT) and Kun-Hsuan

Lee (NMNS). Permissions for collection were kindly

offered by Kenting National Park, Taroko National

Park and Forestry Bureau (0961750119). This study

was supported by the National Science Council Grant,

ROC (NSC-94-2313-B-002-06, NSC-95-2313-B-002-

035, NSC-95-2621-B-212-001).

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SYSTEMATICS OF ARBOREAL SATSUMA IN TAIWAN 493

Convergent evolution of Amphidromus-like colourful arboreal snails and phylogenetic relationship of East Asian camaenids, with description of a new Aegistohadra species (Helicoidei: Camaenidae: Bradybaeninae)

Article

Full-text available

Apr 2022

East Asian terrestrial snails of the family Camaenidae Pilsbry, 1895a are diverse in terms of genus and species numbers, shell morphology and mode of living. This family also includes colourful conical arboreal snails that traditionally have been assigned to the genus AmphidromusAlbers, 1850. Yet, the present study shows that, despite their deceiving conchological similarity, some of these East Asian arboreal snails do not belong to the genus Amphidromus or the subfamily Camaeninae Pilsbry, 1895a. The presence of a dart complex comprising a mucous gland, a dart sac, an accessory sac and a proximal accessory sac, along with a pronounced penial caecum and molecular phylogenetic analyses revealed that former ‘Amphidromus’ dautzenbergi, ‘A.’ roemeri and ‘Camaena’ mirifica, and one additional new species belong to AegistohadraWu, 2004 (subfamily Bradybaeninae Pilsbry, 1934). Aegistohadra dautzenbergi, comb. nov. and Aegistohadra roemeri, comb. nov. are conical with colourful spiral bands, whereas Aegistohadra mirifica, comb. nov. and Aegistohadra zhangdanae, sp. nov. are heliciform to conical with colourful, variegated spiral and transverse banding patterns. DNA sequence analyses also revealed that each variety of Aegistohadra dautzenbergi could not be differentiated by mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) gene fragments. The phylogenetic position of Aegistohadra within the East Asian camaenids revealed that the similar appearance in shell morphology, microhabitat use and diet to arboreal snails in the genus Amphidromus is homoplastic. Moreover, the presence or absence of a dart complex is also homoplastic and is unsuitable for suprageneric classification. By contrast, the presence of a flagellum and a penial caecum is useful for the suprageneric classification.

Arboreal snail genus Amphidromus Albers, 1850 of Southeast Asia: Shell polymorphism of Amphidromus cruentatus (Morelet, 1875) revealed by phylogenetic and morphometric analyses

Article

Full-text available

Aug 2022
PLOS ONE

Species of colourful arboreal snails of the genus Amphidromus from Southeast Asia commonly exhibit high intraspecific variation in shell morphology. Although highly polymorphic Amphidromus specimens with different colouration have been collected at the same locality and were revealed to possess similar genital organs, there is yet no morphometric or DNA analyses of these different shell morphs. This study is the first to reveal that both striped and stripeless morphs of A. cruentatus from Laos and Vietnam belong to the same mitochondrial (COI and 16S rRNA) lineage. Although the shell colouration between the striped and stripeless morphs is markedly different, morphometric and shell outline-based analyses indicated an overall similarity in shell shape. We also revised the systematics of A. cruentatus, in which we treated similar related species, namely A. eudeli, A. fuscolabris, A. thakhekensis, A. gerberi bolovenensis, A. goldbergi, A. pengzhuoani, A. eichhorsti and A. pankowskiae as junior synonyms of A. cruentatus. Amphidromus daoae, A. anhdaoorum, A. stungtrengensis, A. yangbayensis and A. yenlinhae, which were formerly regarded as junior synonyms, are considered as species different from A. cruentatus based on shell morphology and morphometric analyses. Preliminary phylogenetic analyses also retrieved some Amphidromus species groups as distinct mitochondrial lineages.

Descriptions of four new dextral land snails of the genus Camaena (Gastropoda, Eupulmonata, Camaenidae) from south China

Article

Full-text available

Nov 2020

In this study, four new dextral camaenid from China are reported, based on shell morphology, reproductive system anatomy, and molecular phylogenetic analyses: Camaena funingensis Zhou, Wang & Lin, sp. nov. , Camaena gaolongensis Zhou, Wang & Lin, sp. nov. , Camaena maguanensis Zhou, Wang & Hu, sp. nov. , and Camaena yulinensis Zhou, Wang & Hu, sp. nov. Detailed descriptions of the morphological characteristics including shells and genitalia, DNA sequences, and living environments of the four new species are provided, with further comparisons with congeners.

Niche partitioning among three snail‐eating snakes revealed by dentition asymmetry and prey specialisation

Article

Full-text available

Feb 2021
J ANIM ECOL

The level of dentition asymmetry in snail‐eating snakes may reflect their prey choice and feeding efficiency on asymmetric land snails. The three species of Pareas snakes (Squamata: Pareidae) in Taiwan, which form partially sympatric distribution on the island, provide a potential case to test the hypothesis of niche partitioning and character displacement with regard to dentition asymmetry and specialisation in feeding behaviour. In this study, behavioural experiments confirmed that P. formosensis feeds exclusively on slugs, whereas P. atayal and P. komaii consumed both. However, P. atayal more efficiently preys on land snails than P. komaii, exhibiting a shorter handling time and fewer mandibular retractions. Micro‐CT and ancestral character reconstruction demonstrated the lowest asymmetry in P. formosensis (the slug specialist), the highest dentition asymmetry in P. atayal (the land snail specialist) and flexibility in P. komaii (the niche switcher): increased dentition asymmetry when sympatrically distributed with the slug eater (character displacement), and decreased asymmetry when living alone (ecological niche release). Ecological niche modelling showed that the distribution of P. formosensis is associated with the presence of slugs, while that of P. atayal could be explained by the land snails. Combining the results from morphology, phylogeny, behavioural experiments and ecological niche modelling, we showed that competition in the sympatric region might have facilitated character displacement among congeners, while the absence of competition in allopatric region has led to ecological niche release.

Shell chirality and phylogeography of land snail Satsuma batanica species group (Gastropoda: Stylommatophora: Camaenidae)

Article

Full-text available

Sep 2024
CONTRIB ZOOL

Chirality has been recognised as a potentially important mechanism in the speciation of land snails because it leads to reproductive isolation between individuals with different coiling directions. The Satsuma batanica species group comprises the sinistral S. batanica and the dextral S. bacca , despite the observed sympatric distributions of the two chiral morphs. This study examines the population structure, phylogeographic history and taxonomy of the species group by analyzing mitochondrial COI gene sequences. A total of 127 haplotypes were identified from 367 individuals collected from 39 locations. The phylogenetic trees, constructed using Bayesian inference and maximum likelihood methods, along with the median-joining network, identified 14 clades. The reconstructed ancestral states indicate that the most recent common ancestor of the S. batanica species group was sinistral. The frequent occurrence of the dextral morph, the shared haplotype, the sympatric distribution, and the co-occurrence of the two chiral morphs within the same phylogenetic clades recovered here do not support reproductive isolation and single-gene speciation based on chiral differences. All the names belonging to the S. batanica species group are considered synonyms because of their molecular and morphological similarities. This species is estimated to have originated approximately 615 thousand years ago from the northern margin of its current distribution. Two major genetic divergence stages occurred around 440 and 50–100 thousand years ago. The formation of ancient islands through orogenic activities is utilised to explain the phylogeographic history of the species.

Three new species of Camaena Albers, 1850 (Gastropoda, Stylommatophora, Camaenidae) from Guizhou and Guangxi, China

Article

Full-text available

Sep 2024

Three new species of camaenid land snail, Camaena qiannanensis Chen, Dai, Wu & Ouyang sp. n., Camaena feichenyui Chen, Dai, Wu & Ouyang sp. n. and Camaena qinghuai Chen, Dai, Wu & Ouyang sp. n. are described from Guizhou and Guangxi of southern China based on shell and genital morphology, and molecular phylogeny. The phylogenetic result shows that the genus Camaena is not a monophyletic group, with Camaenella located within it. The discovery enhances the species diversity of Camaena in Southern China and fills the gap between distribution areas of known species.

Three new species of Satsuma Adams, 1868 (Gastropoda, Camaenidae) from Taiwan

Article

Feb 2024

Revision of Burmochloritis Godwin-Austen, 1920 in Southeast Asia (Gastropoda: Stylommatophora: Camaenidae)

Article

Dec 2023

This paper revises the Southeast Asian Camaenidae that are characterized by a relatively large (>20 mm) shell with multiple reddish spiral bands on the body whorl and a finely pitted protoconch. The anatomy of 2 species is known, Burmochloritis kengtungensis Godwin-Austen, 1920 and Helix massiei L. Morlet, 1891, the latter from this study. Since both are similar to each other and differ from all other camaenids, we include both in the genus Burmochloritis Godwin-Austen, 1920. All additional species with such traits are known from empty shells only. They are provisionally also placed in Burmochloritis, although further information will be necessary to confirm their taxonomic position. Five new species are described from the collection of the Muséum national d’Histoire naturelle (Paris, France): Burmochloritis laotica Páll-Gergely & Neubert n. sp., B. muratovi Páll-Gergely n. sp., B. nordsiecki Páll-Gergely & Neubert n. sp., B. paya Páll-Gergely n. sp., and B. payella Páll-Gergely n. sp. Helix (Chloritis) marimberti Bavay & Dautzenberg, 1900 is a junior synonym of Burmochloritis luctativa (Mabille, 1889). Helix (Chloritis) marimberti var. carinata Bavay & Dautzenberg, 1909, which has been treated as a synonym of Helix marimberti, is here elevated to species level. Since it is a junior homonym of Helix carinata Link, 1807, the replacement name Burmochloritis laocaiensis Páll-Gergely, nom. nov., is proposed. With the current revision, Burmochloritis comprises 15 species and ranges through eastern Myanmar, northern Laos, and northern Vietnam, and there is a single species known from southern Laos. Chloritis anserina and C. theobaldi, which were formally included in Burmochloritis, are moved to the genus Bouchetcamaena Thach, 2018 because they lack multiple reddish bands.

Correction of two homonyms and an original spelling of Thach [2018]

Article

Jan 2019

Barna Páll-Gergely

Two substitution names are proposed for primary homonyms introduced for land snail of the family Camaenidae in Thach [2018].

MALACOLOGICA TAXONOMIC VANDALISM IN MALACOLOGY: COMMENTS ON MOLLUSCAN TAXA RECENTLY DESCRIBED BY N. N. THACH AND COLLEAGUES (2014-2019)

Article

Full-text available

May 2020

A New Satsuma Species (Pulmonata: Camaenidae) Endemic To Taiwan

Article

Full-text available

Oct 2007

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.

Phylogeography: The History and Formation of Species

Book

Jan 2000

JOHN C. AVISE

A new Satsuma species (Pulmonata: Camaenidae) endemic to Taiwan

Article

Oct 2007

CONFIDENCE LIMITS ON PHYLOGENIES: AN APPROACH USING THE BOOTSTRAP

Article

Jul 1985
EVOLUTION

Joseph Felsenstein

The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.

Type land snails in the Academy of Natural Sciences of Philadelphia part II. Land Pulmonata, exclusive of North America north of Mexico

Article