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An impressive new camaenid, Entadella entadiformis gen. & sp. n. From Guangxi, China (Gastropoda: Pulmonata)

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An impressive, large bodied, new camaenid species, Entadella entadiformis Páll-Gergely & Hunyadi n. sp. is described from several localities in Guangxi Province (Southern China). The new genus, Entadella Páll-Gergely & Hunyadi n. gen. includes three species: E. athrix (Möllendorff, 1901), E. cavaleriei (Bavay, 1913), and E. entadiformis n. sp., and characterized by a large, nearly flat, brownish shell with rounded body whorl, smooth or finely tuberculated embryonic whorls and a very or moderately narrow umbilicus. The penial verge is small or middle-sized with lateral opening, the penial caecum is absent, the flagellum is short, and the vagina is very short. In order to provide basis for comparison of Entadella n. gen. with the type species of the genus Camaena, the reproductive anatomy, shell and radula of a Camaena cicatricosa specimen, collected from Guangxi, China, is also described.
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Journal of ConChology (2016), Vol.42, no.4 167
AN IMPRESSIVE NEW CAMAENID, ENTADELLA ENTADIFORMIS
GEN. & SP. N. FROM GUANGXI, CHINA (GASTROPODA:
PULMONATA)
Barna Páll- gergely1, andrás hunyadi2, Jamen uiriamu otani3 & takahiro asami1
1Department of Biology, Shinshu University, Matsumoto 390–8621, Japan
2Adria sétány 10G 2/5., Budapest 1148, Hungary, e- mail: hunand@freemail.hu
32- 2- 14 Shinmachi, Minakuchi, Koka, Shiga, 528–0038 Japan
Abstract An impressive, large bodied, new camaenid species, Entadella entadiformis Páll- Gergely & Hunyadi n. sp. is
described from several localities in Guangxi Province (Southern China). The new genus, Entadella Páll- Gergely & Hunyadi
n. gen. includes three species: E. athrix (Möllendorff, 1901), E. cavaleriei (Bavay, 1913), and E. entadiformis n. sp., and
characterized by a large, nearly flat, brownish shell with rounded body whorl, smooth or finely tuberculated embryonic whorls
and a very or moderately narrow umbilicus. The penial verge is small or middle- sized with lateral opening, the penial caecum
is absent, the flagellum is short, and the vagina is very short. In order to provide basis for comparison of Entadella n. gen.
with the type species of the genus Camaena, the reproductive anatomy, shell and radula of a Camaena cicatricosa specimen,
collected from Guangxi, China, is also described.
Key words Anatomy, taxonomy, land snail, new species, new genus
IntroductIon
Guangxi Province in southern China harbours
abundant limestone habitats suitable for species
rich land snail communities. In spite of this, the
terrestrial snail fauna of this province is little
known, mainly because malacologists working at
the end of the 19th and beginning of the 20th cen-
tury were rather focused on neighbouring areas
(Hunan, Hubei Provinces, Tonkin = Northern
Vietnam). The family Camaenidae Pilsbry, 1893
(including Bradybaenidae, see Wade et al., 2006,
2007; Gittenberger et al., 2012), which is a spe-
ciose group of large and often colourful land
snails inhabiting various habitats, is also nearly
unknown. Heude (1890) described only four
species of the Camaenidae from Guangxi (Si-
lin = Xilin county) and Yen (1939) reported only
five species from that province.
In the past 10 years or so, several new taxa were
described from Guangxi, mainly those belonging
to the families Clausiliidae (e.g. Nordsieck, 2005,
2012; Grego & Szekeres, 2011), Plectopylidae
(see Páll- Gergely & Hunyadi, 2013; Páll- Gergely
& Asami, 2014) and Hypselostomatidae (see
Jochum et al., 2014; Páll- Gergely et al., 2015). On
the other hand, few recent publications have
reported on the specious Camaenidae (e.g. Ding
et al. 2016). In this paper we describe a very large,
conspicuous species of the Camaenidae based on
shell and anatomical characters. This new spe-
cies cannot be included in any other camaenid
genera. Therefore we diagnose a new genus for
this new species and discuss its relationship with
other anatomically known camaenids. Two addi-
tional species, Chloritis athrix Möllendorff, 1901
and Helix (Chloritis) cavaleriei Bavay, 1913 are also
classified within the new genus.
MaterIal and Methods
Determination of the number of shell whorls (pre-
cision to 0.25 whorl) follows Kerney & Cameron
(1979: 13). Animals were killed with boiling water
and bodies were pulled out afterwards. Shells,
radulae and jaws were directly observed without
coating under a low vacuum SEM (Miniscope
TM- 1000, Hitachi High- Technologies, Tokyo).
Individual buccal masses were removed and
soaked in 2 M KOH solution overnight before
extracting the radula, which was preserved in
70% ethanol.
The holotype of Entadella entadiformis n. sp.
was collected when it was juvenile (approx. 3
whorls), but was raised in captivity by J.U. Otani.
Both of the anatomically examined Entadella
entadiformis n. sp. specimens were pulled out
of the shell after boiling, and the spermoviduct
and the bursa copulatrix were torn and remained
within the shell, and therefore their morphology
could not be examined. The ethanol- preserved
Contact author : pallgergely2@gmail.com
B PÁll-gergely et al
168
specimens were dissected under a Leica stereo-
microscope, equipped with a digital camera. In
the description of the reproductive system, we
used the terms “proximal” and “distal” relative
to the hepatopancreas.
The meaning of Chinese administrative units
are the following: Xian = district, Shi = town
region, Xiang and Zhen = community, Cun =
village, Zizhixian = autonomous region.
Abbreviations
HA: Collection András Hunyadi (Budapest,
Hungary)
HNHM: Hungarian Natural History Museum
(Budapest, Hungary)
MNHN: Muséum National d’Histoire Naturelle
(Paris, France)
OK: Collection Kenji Ohara (Nishinomiya, Japan)
PGB: Collection Barna Páll- Gergely (Moson-
magyaróvár, Hungary)
JUO: Collection Jamen Uiriamu Otani (Koka,
Japan)
systeMatIcs
Family Camaenidae Pilsbry, 1893
Genus Camaena Albers, 1850
Type species Helix cicatricosa (Müller, 1774).
Remarks A recent paper revised the sinistral
Camaena species of China including molecu-
lar phylogeny and genital anatomy (Ding et al.
2016). For comparative reasons we provide a
brief description of the protoconch and the geni-
talia of the type species.
Camaena cicatricosa (Müller, 1774)
Figs 1–2.
Specimen examined China, Guangxi, Liuzhou
Shi, Longtan Gongyuan, 200m a.s.l., 24°16.913'N,
109°24.568'E, leg. Nakahara, Y., Ohara, K., Okubo,
K. & Otani, J.U., 11.11.2004., HNHM 99698 (shell),
HNHM 99698a (body and prepared genitalia in
ethanol), HNHM 99698r (radula on double faced
adhesive tape).
Figure 1 Shell and protoconch of the anatomically examined Camaena cicatricosa (O.F. Müller, 1774) specimen
(HNHM 99698). D = 43.8mm.
entadella entadiformis gen. et sP. n. from China 169
Remarks on the shell Shell large, sinistral,
de pressed globular; protoconch large with rough,
irregular, but glossy surface (overall resembles to
a rumpled and smoothed metal foil).
Description of the genitalia (Figs 2A–E) One spec-
imen was anatomically examined. Atrium short;
penis relatively short, cylindrical, inner wall with
irregular, fine, transversal wrinkles; penial verge
opens terminally, extremely long, extends almost
to the distal end of the penis, cylindrical, becomes
slightly slimmer towards its end; epiphallus
approximately as long as penis, internally with
roughly four longitudinal folds; retractor mus-
cle short, thick, inserts approximately middle of
epiphallus, and attaches on diaphragm; there are
additional, weaker muscle fibres inserting on the
epiphallus; flagellum nearly as thick as epiphal-
lus, becomes slightly slimmer towards its end;
flagellum internally with three low and one high
Figure 2 Reproductive anatomy (A–E), jaw (F) and radula (G–I) of Camaena cicatricosa (O.F. Müller, 1774) (HNHM
99698). A: whole genitalia; B: inner structure of the penis; C: inner structure of the epiphallus (e) and the flagellum
(f); D: spermatophore; E: inner structure of the vagina; F: jaw; G: half of the radula; H: central and first lateral
teeth; I: marginal teeth.
B PÁll-gergely et al
170
longitudinal fold; vas deferens enters epiphallus
laterally; vagina approximately as long as penis,
internally with several irregular, serrated longi-
tudinal folds; bursa copulatrix with very long
stalk and a thickened, club- like sac; an ovoid
spermatophore with pointed end was found in
the bursa.
Description of the jaw and radula (Figs 2F–I) Jaw
odontognathous, with approximately 12 regular,
very strong ribs; radula elongated but not very
slender; central tooth present, unicuspid, slightly
smaller than first laterals; laterals and margin-
als arranged in transversal, more or less straight
rows; laterals 25, marginals at least 35, laterals
unicuspid, gradually becoming bi- and tricuspid;
marginals tricuspid, the endocone is the largest,
with rounded tip. The number of laterals and
marginals only estimated because the number
and shape of cusp changes gradually.
Remarks Solem (1992) gave a detailed descrip-
tion of the anatomy of Camaena cicatricosa col-
lected from Hong Kong. Our results largely
agree with that of Solem (1992), but some differ-
ences were also found. First, the penial verge in
our specimen is much longer than the specimen
from Hong Kong, in which this organ looks as if
retracted (Solem 1992, Fig. 2A). Second, the fla-
gellum is very long, vermiform in the specimen
examined by Solem (1992), but our specimen had
a relatively short, thick flagellum. The anatomi-
cal description of Hwang (2011), which was also
based on a specimen collected in Hong Kong,
agrees with that of Solem (1992). Our results
overall agree with the anatomy of C. cicatricosa
published by Ding et al. (2016) with the excep-
tion of sculpture of the inner penial wall and the
verge. Namely, the specimen we examined had
both surfaces rather finely reticulated, whereas
those of the specimen in Ding et al. (2016) was
finely longitudinally striated.
Camaenid spermatophores usually possess a
long tail. The spermatophore of the examined
specimen was ovoid with pointed end, and
looked complete, not decayed.
Entadella Páll- Gergely & Hunyadi n. gen.
Diagnosis Shell large, discoid, nearly flat, with
narrow or wide umbilicus, protoconch smooth or
covered with small tubercles to various degree.
Penis relatively short; penial verge small to
medium sized, with lateral opening; vagina very
short.
Type species Entadella entadiformis n. sp.
Species included E. athrix (Möllendorff, 1901), E.
cavaleriei (Bavay, 1913), Entadella entadiformis n. sp.
Comparisons Entadella n. gen. differs from
Camaena Albers, 1850 by the penial verge with
lateral opening, whereas that in Camaena it opens
terminally. Moreover, the vagina of Camaena is
much longer than that of Entadella n. gen. (for the
anatomy of C. cicatricosa and related species, see
Solem, 1992; Schileyko, 2003; Hwang, 2011; Ding
et al., 2016; and this study). Conchologically,
Camaena species have a higher spire and gener-
ally a rather globular shell very often with dark
bands. Moreover, the protoconch sculpture of
Camaena is irregular (resembles to a battered
metallic surface), whereas that of Entadella n. gen.
is smooth, or ornamented with small tubercles.
The radula and jaw morphology of Camaena and
Entadella n. gen. shows no notable differences.
The anatomy of Trichochloritis Pilsbry, 1891 (see
Schileyko, 2007), which is conchologically some-
what similar to the new genus, is very different
from that of Entadella n. gen. Most importantly
it lacks a flagellum, has a penial sheath, and the
retractor muscle inserts on the curvature of vas
deferens. Camaenella Pilsbry, 1893 has a nearly
globose shell and a very large, elongated penial
verge (Schileyko, 2003). Neocepolis Pilsbry, 1891
has a globose shell with an obtuse tooth on the
columella and with a strong fold on the parietal
wall (Sutcharit et al., 2007). Moreover, the vagina
of Neocepolis is much longer (Sutcharit et al.,
2007). Trachia Martens, 1860, which is widely dis-
tributed in Southeast Asia, also possesses a flat
shell with rather narrow umbilicus, but many
species have bands on the shell and they have
less whorls than Entadella n. gen. The main dif-
ference between the anatomies of Trachia and
Entadella n. gen. is that the former lacks a penial
verge and has a long vagina, whereas the new
genus has a small verge and possesses very short
vagina (Schileyko, 2003, 2013).
Moellendorffia Ancey, 1887 (and related genera:
Trichelix L. Pfeiffer, 1862 and Moellendorffiella
Pilsbry, 1905) is similar to Entadella n. gen. by the
granulated protoconch, the uniformly brown-
ish colour, the short flagellum and short penial
entadella entadiformis gen. et sP. n. from China 171
verge, which is nevertheless bilobed in Trichelix
and Moellendorffia (see Schileyko 2003). The shell
of Moellendorffia differs from that of Entadella n.
gen. by the prominently granulated and hirsute
teleoconch and the presence of teeth (lamellae) in
the aperture.
Derivation of name The shell of Entadella species
resemble to the seed of the tropical liana belong-
ing to the genus Entada (Fabaceae) in terms of
size, shape and colour. Entadella is the diminu-
tive form of Entada.
Remarks Based on shell characters, the genus
Entadella could be classified in the family
Helicarionidae and other related “limacoid”
groups as well. The absence of a caudal horn and
the odontognathous jaw clearly indicates that the
new genus is a member of Camaenidae.
Entadella athrix (Möllendorff, 1901)
Chloritis athrix Möllendorff, 1901: 73–74.
Chloritis (Trichochloritis) athrix, — Zilch, 1966: 300,
Plate 9, Fig. 22.
Diagnosis A medium sized, brownish or yel-
lowish, nearly flat camaenid species with very
narrow, but open umbilicus, semilunar aperture,
and regularly growing whorls.
Material examined Tonkin, Lao Kay, prob-
ably leg. Messager, NHMUK 1908.12.21.39–40;
Region de Muong- Hum, Tonkin, MNHN-
IM- 2012–27112/12; Tonkin, Muong Hum, coll.
Staadt, 1969, MNHN- IM- 2012–27113/4; Muong-
Hum, leg. Messager, MNHN- IM- 2012–27114/6;
Phong- Tho, leg. Messager, MNHN-
IM- 2012–27115/8; Phong- Tho, leg. Messager,
MNHN- IM- 2012–27116/3; Phong- Tho, leg.
Messager, MNHN- IM- 2012–27117/5; Muong Bo,
leg. Messager, MNHN- IM- 2012–27118/3 (albi-
nistic form); Muong Bo, leg. Messager, MNHN-
IM- 2012–27119/6; Muong Bo, leg. Messager,
MNHN- IM- 2012–27120/3; Tonkin, Région de Lao
Kay, leg. Messager, MNHN- IM- 2012–27121/3;
Tonkin, Lao Kay, leg. Messager, MNHN-
IM- 2012–27122/3; Tonkin, Lao Kay, leg. Messager,
MNHN- IM- 2012–27123/8; Tonkin, Région de Lao
Kay, leg. Messager, MNHN- IM- 2012–27124/5;
Tonkin, Région de Lao Kay, leg. Messager,
MNHN- IM- 2012–27125/50; Tonkin, Pakhé, leg.
Messager, MNHN- IM- 2012–27126/3; Pakhé, leg.
Messager, MNHN- IM- 2012–27127/9; Tonkin,
Pakhé, leg. Messager, MNHN- IM- 2012–27128/3
(albinistic form); Tonkin, Pakhé, leg. Messager,
MNHN- IM- 2012–27129/3; Tonkin, Pakhé, leg.
Messager, MNHN- IM- 2012–27130/2; Muong
Kong, leg. Messager, MNHN- IM- 2012–27131/2;
Muong Kong, leg. Messager, MNHN-
IM- 2012–27132/2; Muong Kong, leg. Messager,
MNHN- IM- 2012–27133/2 (one adult and one
juvenile shell); Muong Kong, leg. Messager,
MNHN- IM- 2012–27134/3; Ta Phin (N.
Vietnam), leg. Saurin, MNHN- IM- 2012–27135/1;
Tonkin, Bords de la Rein Nong (?), MNHN-
IM- 2012–27136/1; Tonkin, leg. Messager,
MNHN- IM- 2012–27137/2; Muong Bo, Dinh Lu,
leg. Messager, MNHN- IM- 2012–27138/3; Dinh
Lu, leg. Messager, MNHN- IM- 2012–27139/1;
Lang Léou, That Khé et Bac Kan, leg. Messager,
MNHN- IM- 2012–27140/2; Tonkin, leg. Messager,
MNHN- IM- 2012–27141/2; Tonkin, leg. Messager,
coll. Letellier 1949, MNHN- IM- 2012–27142/1.
Geographic range (Fig. 9) Entadella athrix is
known from a few localities in Northern Vietnam,
along the Chinese border.
Remarks The anatomy of this species is
unknown, but its shell is similar to that of
Entadella entadiformis n. sp. (for differences, see
under that species). Therefore, this species is
classified to Entadella n. gen.
Entadella cavaleriei (Bavay, 1913)
Fig. 3
Helix (Chloritis) Cavaleriei Bavay, 1913: 603–604,
Plate 21.
Diagnosis A large, brownish, nearly flat cama-
enid species with moderately wide umbilicus
and a semilunar aperture. Penis relatively short,
thickened, internally with parallel folds and a
medium sized, club- shaped verge with lateral
opening; vagina very short.
Material examined San Chouen, Chine
Meridionale, MNHN- IM- 2000–31769 (holotype);
Same data, MNHN IM- 2012–27177 specimen in
ethanol (anatomically examined).
Description of the genitalia (Figs 3B–D) One spec-
imen was anatomically examined. This specimen
was obviously part of the original series (col-
lector and collection site agrees), but was not
B PÁll-gergely et al
172
mentioned in the original description, therefore
it is not a paratype.
The whole body was very much fragile; only
the outer part of the body (including the buccal
mass, the sole and the posterior genitalia) could
be examined. Penis relatively short, thickened,
internally with dense, irregular, parallel folds;
penial verge blub- shaped, middle sized, with
lateral opening; epiphallus cylindrical, longer
than penis; epiphallus- flagellum region was
extremely fragile, it broke even due to the slight-
est touch with the forceps, therefore a retractor
muscle could not be observed; flagellum well-
developed, shorter than epiphallus, pointed,
rather slender; vas deferens cylindrical, rela-
tively thick; vagina short and thick, inner wall
Figure 3 A, Shell of the holotype of Entadella cavaleriei (Bavay, 1913) (MNHN- IM- 2000–31769). Reproductive
anatomy (B–D), jaw (E) and radula (F–H) of another specimen (MNHN IM- 2012–27177). B–C: penial verge from
two different views (white arrow indicates the laterally positioned opening); D: whole genitalia (note that a retrac-
tor muscle might be present, but could not examined due to the fragile condition of the specimen); E: jaw; F: half
of the radula; G: central and first lateral teeth; H: marginal teeth.
entadella entadiformis gen. et sP. n. from China 173
not examined; bursa copulatrix long, there is no
distinct sac- shaped bursa.
Description of the jaw and radula (Figs 3E–H) Jaw
odontognathous, with approximately twelve
very strong, irregular ribs; radula elongated but
not very slender; central tooth present, unicus-
pid, with slight indication of two additional side
cusps; central tooth smaller than the first later-
als; laterals and marginals are arranged in trans-
versal, more or less straight rows; laterals 14–15,
marginals at least 15 (the edge of the radula was
twirled, so the outermost marginals could not be
examined); laterals unicuspid, gradually becom-
ing bi- and tricuspid; marginals tricuspid. The
number of laterals and marginals only estimated
because the number and shape of cusps changes
gradually.
Geographic range This species is known from the
type locality only (“San Chouen in China meridi-
onali”), which could not be located on the map.
Remarks The shell of this species is similar to
that of Entadella entadiformis n. sp. in shell size
(D = 34.1, H = 15.1), shape and sculpture, espe-
cially the small tubercles near the suture of the
protoconch. Therefore, it is classified in the genus
Entadella.
Entadella entadiformis Páll- Gergely & Hunyadi
n. sp.
Figs 4–8.
Holotype China, Guangxi, Douanyaozu
Zizhixian, Baoan Xiang, Nongjiao, 607m,
24°05.580'N, 107°46.971'E, leg. Ishibe, T., Ohara,
K., Okubo, K. & Otani, J.U., 24.10.2011., HNHM
99699 (shell), HNHM 99699a (body and prepared
genitalia in ethanol), HNHM 99699r (radula on
double faced adhesive tape in ethanol).
Paratypes China, Guangxi, Douan Yaozu
Zizhixian, Baoan Xiang, Nongjiao, 607m,
24°05.580'N, 107°46.971'E, leg. Ishibe, T., Ohara,
K., Okubo, K. & Otani, J.U., 22.10.2011., JUO/1
paratype, OK/5 paratypes, PGB/1 paratype (ex
coll. K. Ohara); China, Guangxi, Bama Yaozu
Zizhixian, Jiaoyuetiankeng, 285m, 24°06.92707'N,
107°06.81754'E, leg. Nakahara, Y., Ohara, K.,
Okubo, K. & Otani, J.U., 15.07.2006., HNHM
99700 (1 paratype: shell, ethanol- preserved
body, prepared genitalia, radula on double faced
adhesive tape), JUO/1 paratype, OK/3 para-
types; China, Guangxi, Dahua Yaozu Zizhixian,
Qibainong Xiang, near bus stop at Nòngténg,
603m, 24°04.905'N, 107°41.100'E, leg. Ishibe, T.,
Ohara, K., Okubo, K. & Otani, J.U., 22.10.2011.,
JUO/2 paratypes; China, Guangxi, Heshan
Shi, Beisi Xiang, Baxianyan, 120m, 23°49.510'N,
108°55.942'E, leg. Nakahara, Y., Ohara, K.,
Okubo, K. & Otani, J.U., 14.11.2004., OK/1 para-
type; China, Guangxi, Laibin, leg. Takagi T.,
collection date unknown, OK/1 paratype (not
exact locality, not indicated on the map); China,
Guangxi, Laibin Shi, Wushan Xiang, eastern
end of Xiaopingyang Zhen, 130m, 23°24.351'N,
109°10.029'E, leg. Hunyadi, A., 7.10.2009, HA/1
paratype; China, Guangxi, Hechi Shi, Duan
Yaozu Zizhixian, Gaoling Cun, Dingfu Cun
W 2km, 320m, 24°03.197'N, 108°01.290'E, leg.
Hunyadi, A., 8.10.2009, HA/1 paratype; China,
Guangxi, Nanning Shi, Longan Xian, Longhushan
Senlin Gongyuan, Peri Hill, 140m, 22°57.485'N,
107°37.754'E, leg. Hunyadi, A., 11.10.2009, HA/5
paratypes; China, Guangxi, Hechi Shi, Bama
Xian, cliffs near Jiaole Cun, 590m, 24°7.045'N,
107°7.847'E, leg. Hunyadi, A. & Szekeres, M.,
10.09.2013, HA/12 adult paratypes + 7 juve-
nile paratypes; China, Guangxi, Dahua Yaozu
Zizhixian, prefectural border at Qibainong Xiang,
688m, 24°06.496'N, 107°41.466'E, leg. Ishibe, T.,
Ohara, K., Okubo, K. & Otani, J.U., 22.10.2011.,
JUO/2 paratypes.
Type locality China, Guangxi, Douan Yaozu
Zizhixian, Baoan Xiang, Nongjiao, 607m,
24°05.580'N, 107°46.971'E.
Measurements (in mm) shell diameter: 32.5–49.5,
shell height: 14.9–20.9 (n = 17, from different sam-
ples). See also Table 1.
Diagnosis A large, brownish, nearly flat cama-
enid species with very narrow, but open umbili-
cus, semilunar aperture and irregularly growing
whorls. Penis relatively short, thickened inter-
nally with parallel folds and a small, triangular,
pointed verge with lateral opening. Vagina very
short with longitudinal wrinkles internally.
Description of the shell (Figs 4C–D, 5) Shell large,
dextral, rather thin and light in face of its robust
appearance, light chocolate brown to dark orange
in colour; shell rather flat with slightly elevated
spire; shell outline rather oval from dorsal view
B PÁll-gergely et al
174
due to the shape of the body whorl which is
the widest on the opposite side of the shell
than the aperture; in one specimen from Laibin
the body whorl was the widest about a quarter
whorl behind the aperture; body whorl rounded,
somewhat shouldered by slightly depressed
from ventrolateral direction; whorls 6.25–7.25,
convex, separated by moderately deep suture;
protoconch 2.25–2.5 whorls, covered with rough,
elongated papillae which are arranged irregu-
larly on the first approximately 1.5 whorls but
gradually become regular and arranged in radial
rows; usually 8–10 tubercles stand in a single
row; occasionally the first 0.5–1 whorl is irregu-
larly wrinkled without tubercles; these tuber-
cles cover the whole protoconch in only some
populations (Qibainong Xiang and Nòngténg) in
others the protoconch is nearly entirely smooth,
only very few tubercles are visible near the
inner suture; teleoconch sculpture dominated
by radial wrinkles that are rather regular and
simple at the beginning of the teleoconch (first
Figure 4 Living specimens and shells of Entadella entadiformis Páll- Gergely & Hunyadi n. sp. A: Paratype from
Jiaoyuetiankeng in its original environment; B: living holotype (HNHM 99699); C: holotype (HNHM 99699); D:
paratype from Jiaoyuetiankeng (HNHM 99700).
entadella entadiformis gen. et sP. n. from China 175
0.5–2 whorls, depending on populations), but
become irregular and rugged gradually (there
are several small nodes at the back side); some
very fine, dense spiral lines are also discernible
between ribs; ventral side nearly smooth, only
very fine wrinkles and spiral lines are visible;
rough dorsal sculpture gradually changes on the
body whorl to smooth ventral surface; umbilicus
open but very narrow, its edge slightly covered
by peristome; aperture semilunar, elongated in
the direction of the umbilicus; slightly oblique
to the shell axis from lateral view; no inner ribs,
lamellae or teeth present; peristome lighter than
the shell, it is somewhat thickened and slightly
reflexed; callus absent, but there is a very fine cal-
cium layer on the parietal shell wall which results
in a very slightly lighter colour than the rest of
the shell.
Description of the body and the genitalia (Figs 4A–B,
6–7) Body dark bluish grey; caudal horn and
caudal foss absent; lateral zones of sole grey,
central zone light, but not delimited sharply;
extent of the white central zone variable in the
two examined specimen; in the holotype the
central zone only slightly wider than one of the
darker lateral zones, whereas in specimen from
Jiaoyuetiankeng the lateral grey zones are very
narrow; tentacles relatively short; no head wart
visible. Because of the absent apical whorls of the
bodies of the anatomically examined specimens,
the pallial complex could not be examined.
Figure 5 Protoconch of Entadella entadiformis Páll-
Gergely & Hunyadi n. sp. A–B: holotype (HNHM
99699); C–D: paratype from Jiaoyuetiankeng (HNHM
99700). Scales represent 1mm.
Figure 6 Genital anatomy of Entadella entadiformis Páll- Gergely & Hunyadi n. sp. A: holotype (HNHM 99699); B:
paratype from Jiaoyuetiankeng, (HNHM 99700).
B PÁll-gergely et al
176
Table 1 Measurements of Entadella entadiformis n. sp. from different localities.
Locality name Shell diameter (in mm) Shell height (in mm) no. of measured specimens
Nongjiao 38.6–49.5 16.9–20.4 8
Jiaoyuetiankeng 38.2–43.3 17.3–19.8 5
Qibainong Xiang 45.1–47.3 20.7–20.9 2
Nòngténg 39.6–39.8 18.6–18.7 2
Laibin 33.2 16.7 1
Baxianyan 32.5 14.9 1
Jiaole Cun 33.2–37.7 15.8–17.3 4
Xiaopingyang Zhen 33.5 17.6 1
Dingfu Cun 44.8 spire broken 1
Peri Hill 35.4–38.3 17.2–20.4 5
Figure 7 Inner structure of the reproductive organs of Entadella entadiformis Páll- Gergely & Hunyadi n. sp. Figs
A–D and G show the holotype (HNHM 99699), Figs E–F show the paratype from Jiaoyuetiankeng (HNHM 99700).
A: penis (p), epiphallus (e) and flagellum (f); B: flagellum; C–F: penial verge; G: vagina (v) and the stalk of the
bursa copulatrix (b). Figs C–D and E–F show the verge from two different views.
entadella entadiformis gen. et sP. n. from China 177
Two specimens were dissected belonging to dif-
ferent populations. Right ommatophoral retrac-
tor crosses the male and female genitalia; penis
relatively short, thickened, internally with irreg-
ular, parallel folds, some of which join together
to form a single fold; the folds run into a circular
wart- like structure which is situated at the proxi-
mal end of the penis, just under the penial verge;
penial verge rather triangular, pointed, with lat-
eral opening; epiphallus cylindrical, longer than
the penis, with a strong retractor muscle attached
around the middle; epiphallus internally with
4–5, strong, parallel longitudinal folds; flagellum
well- developed, slightly shorter than epiphallus,
with a pointed and curved apical part; the paral-
lel fold on the inner wall of epiphallus continues
inside flagellum; vas deferens cylindrical, rela-
tively thick; an additional retractor muscle runs
together with it; the two branches of the muscle
attach on the epiphallus and flagellum; vagina
short and thick, internally with parallel, irregu-
lar wrinkles and some fine transversal structure
between the folds; this inner structure continues
in the stalk of the bursa copulatrix, which is very
long.
Description of the jaw and radula (Fig. 8) Jaw
odontognathous, with approximately ten very
strong, irregular ribs. Radula elongated but not
very slender; central tooth present, unicuspid,
smaller than the first laterals; laterals and mar-
ginals are arranged in transversal, more or less
straight rows; laterals 19, marginals at least 16;
laterals unicuspid, gradually becoming bi- and
tricuspid; marginals tricuspid, but the outer-
most cusp can be also divided; the endocone
is the largest, it is rather pointed. The num-
ber of laterals and marginals only estimated
because the number and shape of cusps changes
gradually.
Derivation of name The name of the new species
refers to the similarity of the shell to the seeds of
Entada spp.
Geographic range (Fig. 9) This new species is
known from Guangxi Province, China.
Comparisons Entadella athrix is smaller and less
depressed (shell diameter: 24.6–30.2, shell height:
13.6–16.5), has fewer (5–6) whorls, and a yellow
corneous shell. Moreover, its aperture is less
oblique from lateral view, the growth rate of the
whorls is more regular, its dorsal sculpture much
weaker, and the protoconch is smooth, matt,
without obvious tubercles. Entadella cavaleriei has
much wider umbilicus, comparatively smaller
aperture, and slightly more angled body whorl.
The weak constriction of the last whorl and the
brown shell is superficially similar to Stegodera
angusticollis (Martens, 1875) and Traumatophora
triscalpta (Ancey, 1881). The former species is
sinistral, the last whorl is distorted and crowded
against the preceding whorls, and there is a lon-
gitudinal depression (sulcus) on the last whorl.
The latter species has three longitudinal lamel-
lae in the aperture, corresponding to longitudi-
nal depressions of the outer surface of the body
whorl.
The combination of the large, brownish, flat
shell and the narrow umbilicus distinguishes
Figure 8 Jaw (A–B) and radula (C–H) of Entadella
entadiformis Páll- Gergely & Hunyadi n. sp. Figs A, C,
E, G: holotype (HNHM 99699); Figs B, D, F, H: para-
type from Jiaoyuetiankeng (HNHM 99700). Figs C
and D show the half of the radula, Figs E and F show
the central and first lateral teeth, Figs G and H show
the marginals.
B PÁll-gergely et al
178
Entadella entadiformis n. sp. from all known cama-
enid snails from southern China and Northern
Vietnam.
Habitat Entadella entadiformis n. sp. inhabits
limestone rich primarily and secondary forests,
but does not seem to be associated with lime-
stone rocks strongly. It is relatively abundant in
its natural habitat.
acknowledgeMents
We are very grateful to Dominika Páll- Gergely for
inking the line drawings, Jonathan Ablett (NHM),
Virginie Héros and Philippe Maestrati (MNHN)
for providing access to their museum collec-
tions, and to Manuel Caballer Gutierrez (project
E- RECOLNAT: ANR- 11- INBS- 0004, MNHN) for
the images of Helix cavaleriei. The web article of
Hartmut Nordsieck about the Camaenidae of
Guangxi (www.hnords.de) was of a great help
for us. This study was supported by Grants- in-
Aid for Scientific Research (KAKENHI) from
Japan Society for the Promotion of Science to
T. Asami. Barna Páll- Gergely is an International
Research Fellow of the Japan Society for the
Promotion of Science. We are indebted to The
Biodiversity Heritage Library for the multitude
of rare literature made available to us (www.
biodiversitylibrary.org).
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