Valiguna flava (Heynemann, 1885) from Indonesia and Malaysia: Redescription and comparison with Valiguna siamensis (Martens, 1867) (Gastropoda : Soleolifera : Veronicellidae)

2008·4.5 MB·English

by  Suzete Rodrigues Gomes

THE VELIGER
TheVeliger 50(3):163-170 (October 1, 2008) © CMS, Inc., 2007
Valigunaflava (Heynemann, 1885) from Indonesia and Malaysia:
Redescription and Comparison with Valiguna siamensis (Martens, 1867)
(Gastropoda: Soleolifera: Veronicellidae)
SUZETE RODRIGUES GOMES,1 JULIANE BENTES PICANCO,2
MENNO SCHILTHUIZEN3 and JOSE WILLIBALDO THOME4
1Departamento de Zoologia, Instituto de Biociencias, Universidade Federal do Rio Grande do Sul. Av. Bento
Goncalves, 9500, Cep 91.501-970, Porto Alegre, Rio Grande do Sul, Brazil
(e-mail: [email protected])
2Faculdade de Biociencias, Pontificia Universidade Catolica do Rio Grande do Sul. Av. Ipiranga, 6681,
Cep 90.619-900, Porto Alegre, Rio Grande do Sul, Brazil
(e-mail: jbp_yahoo.com.br)
3Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Locked Bag 2073, 88999 Kota
Kinabalu, Malaysia and National Museum ofNatural History 'Naturalis', P.O. Box 9517, 2300 RA Leiden,
the Netherlands
(e-mail: [email protected])
4Escritorio de Malacologia e de Biofilosofia, Praca Dom Feliciano, 39, sala 1303 (Ed. Guanabara - Centro),
90020-160, Porto Alegre, Rio Grande do Sul, Brazil
Abstract. We redescribe and figure Valiguna flava, an almost unknown Southeast-Asian land slug. Detailed
morphology, radula, jaw and living specimens of Valigunaflava were studied for the first time, based on material
recently collected in Borneo and on the holotype. VI. flava is also compared with Valiguna siamensis, the only other
species ofthe genus, which is for the first time recorded in China.
Key Words: Morphology, anatomy, land slug, Borneo, Australasian region.
INTRODUCTION he considered a new species very close to Vaginula
schneideri (VI. isseli Hoffmann, 1941), he reconsidered
Veronicellidae Gray, 1840 includes a large number of Valiguna, this time as a genus. Hoffmann (1941)
species ofland slugs totally without shell (Thomeet al., included in Valiguna, VI. schneideri and VI. isseli,
2006). Currently, 27 generic taxa are recognized for the species in which the vas deferens does not open
family, which are distributed widely in the humid terminally (acrocaulis form, such as species of Sarasi-
tropicsand subtropics (Hoffmann, 1925; Forcart, 1953; nula) norbasally in the penis(pleurocaulisform, such as
Thome, 1975; Gomes & Thome, 2004). For the species ofSemperula), having an intermediateform, the
Oriental and Australian regions, Gomes & Thome acropleurocaulis orpseudopleurocaulis form.
(2004) recognized six different genera (total of 13 Recently, Gomes & Thome (2004) examined the
species): Filicaulis Simroth, 1913, Laevicaulis Simroth, holotype of Vaginulaflava Heynemann, 1885 (consid-
1913, Sarasinula Grimpe & Hoffmann, 1924, Semper- ered by Hoffmann, 1925 a synonym of Semperula
ula Grimpe & Hoffmann, 1924, Valiguna Grimpe & maculata) and also the original and subsequent
Hoffmann, 1925 and Veronicella Blainville, 1817. descriptions of VI. schneideri and VI. isseli (Simroth,
Originally, the taxon Valiguna was proposed by 1895; Grimpe & Hoffmann, 1925a, b; Hoffmann, 1925,
Grimpe & Hoffmann (1925a) as a subgenus of 1941). They concluded that these three species are
Semperula to include Vaginula schneideri Simroth, synonyms and that Valiguna flava is the valid name
1895 from Eastern Sumatra (Indrapura, Tandjong since it has priority. Gomes & Thome (2004) also
Kuba). However, in the same year that Grimpe & proposed the inclusion ofSemperula siamensis into the
Hoffmann (1925a) proposed the taxon Valiguna, they genus Valiguna because this species also has an
rejected it and considered Vaginula schneideri a acropleurocaulis penis. According to Gomes & Thome
subspecies of Semperula siamensis (Martens, 1867) (2004), VI. flava has records from Nias, Borneo and
(Hoffmann, 1925; Grimpe & Hoffmann, 1925b). Only Indrapura (Indonesia), and Valiguna siamensis from
later, when Hoffmann (1941) found specimens which Galle (Sri Lanka) and Petshaburi (Thailand). Both
Page 164 The Veliger, Vol. 50, No. 3
species were insufficiently described in the original
description. Vaginulus reticulatus Westerlund, 1883,
which is a synonym of VI. siamensis according to
Gomes & Thome (2004), was redescribed by Thome
(1984), although VI. flava has not been studied since
then.
Ourprimary purpose is, forthefirst time, to describe
and figure in detail the morphology, radula, jaw, and
living specimens of Valigunaflava. The study is based
on samples recently collected from Borneo and on the
holotype. The species is also compared with Valiguna
siamensis, the only other species ofthe genus, based on
material from China and on the paratypes of Vaginulus
reticulatus (synonym of VI. siamensis).
MATERIAL AND METHODS
Six living and adult specimens of VI.flava, which were
collected in Borneo and deposited in the Museum
BORNEENSIS of Universiti Malaysia Sabah, were
analyzed. The holotype and paratype of this species,
deposited in the Natural History Museum (BMNH-
1880.10.6), London, England were also studied.
VI. Jlava was also compared with four specimens of
VI. siamensis deposited in the Museum ofthe Institute
of Zoology, Chinese Academy of Sciences, Beijing
(China) (lots ZMIZ01091, ZMIZ01092), from Yunnan
and Guangxi, China. The type-material of Vaginulus
Figures 1-2. External characteristics of Valiguna flava. 1,
reticulatus Westerlund, 1883 (13 specimens) (synonym dorsal position of a living specimen (BOR/MOL 3411); 2,
of VI. siamensis according to Gomes & Thome, 2004), ventral position ofa fixed specimen (BOR/MOL 3439).
deposited at Swedish Museum of Natural History
SMNH
(Stockholm, Sweden) was also examined (lots Valiguna isseli Hoffmann, 1941:234.
SMNH
6427, 3753).
The specimens ofboth species were dissected undera
stereomicroscope for studying the internal structures.
Diagnostic Features
Drawings were done using a camera lucida. Two jaws
and two radulae (from each species) were extracted Themain diagnostic structure inVeronicellidae isthe
under the stereomicroscope and later analyzed using a penis. The penis of Valigunaflava has a glans and a
scanning electron microscope (SEM Philips XL 30). base completely distinct from each other. The base is a
The terminology used and the morphological and cylindrical structure, without peculiarities in outer
anatomical characteristics described and illustrated are surface. The glans starts from a flap that surrounds
those usually considered diagnostic in the Veronicelli- the distal extremity of the penis base. First, it is
dae (Hoffmann, 1925; Coifmann, 1935; Forcart, 1953; cylindrical, but then itcurves, forminga peak, inwhose
Barker, 2001; Thome et al., 2002; Gomes & Thome, extremity the vas deferens opens. In the dorsum ofthe
2001, 2004). curvature (penis apex) is a cylindrical structure covered
by dentate and serrated formations (Figures 9-12).
Valiguna flava (Heynemann, 1885)
(Figures 1-12) External Characteristics
Vaginulaflava Heynemann, 1885:10-11. The specimens are relatively large and they have an
Vaginula schneideri Simroth, 1895:7-8. oval body (Figures 1-2). The notum (dorsal region) is
Semperula (Valiguna) schneideri; Grimpe & Hoffmann smooth only with some widely spaced granules. There
1925a: 391-392. are some scattered blackish spots and also a narrow
Semperula siamensis schneideri; Hoffmann, 1925: 181- light line on the dorsum (Figure 1). This last one is in
182; Grimpe & Hoffmann, 1925b:18-19, 31-33. the middle of the notum and is not always clearly
Valiguna schneideri; Hoffmann, 1941:236. visible. The notum ground coloration ranges from pale
S. R. Gomes et al., 2007 Page 165
Figures 3-8. Radula andjaw of Valigunaflava (BOR/MOL 3409). 3, entirejaw; 4, lateral (L), central (C) and lateral (L) teeth,
respectively; 5. lateral view ofa central tooth; 6. medium part ofthe right halfofthe radula; 7, lateral view ofa lateral tooth; 8,
lateral teeth in the edges ofthe radula.
yellowish brown to dark reddish brown. The hypono- body length (45-60), body width (22-30), sole width
tum is also pigmented from pale yellowish brown to (2.5-4.8) and hyponotum width (5.2-10.5).
dark reddish brown, depending of the notum colora-
tion. However, it isalwaysmuchlighterthanthenotum Digestive System
and has a homogeneous coloration, without spots or
lines or only with few tiny blackish spots (Figure 2). Themouth isfollowedby a buccal bulb (= pharynx),
The soleislight beigeandverynarrow, havinglessthan where the radula and thejaw lie. There are two salivary
half of the hyponotum width. The female pore is glands with very slender, ramified and delicate acini
situated at ca. 45% ofthe length ofthe body measured connected to the buccal bulb. The buccal bulb is
from the front, and it is far from the pedal groove by connected to the esophagus, which is followed by the
ca. 2/5 the width of the hyponotum. In all specimens gastric crop. The latter is barely delimited from the
thefemaleporeis surrounded by a slender line ofblack esophagus, both having almost the same diameter. The
pigmentation (Figure 2). gastric crop leads into a stomach, which is long (twice
Holotype measurements (mm): body length (70.00), longer than wide). Two lobules of the digestive gland
body width (29.00), sole width (5.44) and hyponotum open into the stomach, one anterior and another one
width (8.11). Measurements (mm) (6 other specimens): posterior. The anterior lobe does not totally cover the
Page 166 The Veliger, Vol. 50, No. 3
Figures 9-12. Reproductive system in Valigunaflava (BOR/MOL 3409). 9, complete reproductive system; 10-12, three different
positions of the penis (cb, bursa copulatrix; db, duct ofthe bursa copulatrix; dd, distal posterior deferens; df, vas deferens; dp,
proximal posterior deferens; fc, fertilization complex; fg, female genital pore; gb, albumen gland; gh, hermaphroditic gland; gn,
glans; jd, canalis junctor; It, long tubules, md, middle deferens; ol, spermioduct; ov, oviduct; pk, peak, in whose extremity the
deferens opens; pi, penial gland papilla; pr, prostate; re, rectum; rp, penis retractormuscle; st, short tubules; sv, seminal vesicle; tf,
structure covered by dentate and serrated formations; vg, penis base. Scale bar: 1 mm.
S. R. Gomes et al., 2007 Page 167
anterior intestinal loop. The intestine starts from the bursa copulatrix. The canalis junctor (Barker, 2001;
stomach, followsto theanteriorregionwhere itforms a Gomes & Thome, 2004) penetrates in the duct of the
loop (the anterior intestinal loop) and then continues bursa copulatrix, near to the halfofthe total length of
back to the posterior region. Near the height of the theduct. The oviductiswidernearto the female genital
female pore, the intestine penetrates in the body wall, pore, involving the duct of the bursa copulatrix
where the rectum begins. The rectum continues to the (Figure 9).
end ofthe body where it opens via the anus. The latter The penis has a glans and a base well differentiated.
is located centrally in the body and is totally hidden The base of the penis is somewhat cylindrical and
over the sole. The anal opening is represented by a smooth. The glans starts from an surrounding structure
fissure. Neither a circular opening (with a defined (as a flap) at the distal extremity ofthe penis base; it is
border) nor an opercular membrane is found. The anal initially cylindrical and smooth, but quickly tapers and
opening floor is formed by well developed folds that curves forming a peak, in whose extremity the vas
sometimes seem papillae. The nephridium does not deferens opens. In the dorsum of the curvature is a
have an external aperture. It is probably connected to cylindrical structure covered by dentate and serrated
the rectum within the body wall. formations, which is in the distal extremity ofthe penis
The jaws are well arquated and narrow. They are (Figures 9-12).
formed by an average of24 wide and superposed ribs. The penial gland is small and has a length similar
The three ribs of the middle are somewhat higher and to the length of the penis. It is formed by a papilla
less distinct from the others (Figure 3). and by about 15 short tubules (they exceed a little the
Theradula has 93-95 teethpertransverserow, which height of the pericardium). The papilla is relatively
is formed by one symmetrical central or rhachis tooth long, with around half of the tubules' length. The
(Figures 4-5), flanked on both sides by 46—47 lateral tubules are not differentiated in groups according to
teeth (Figures 6-7). The dental formula is C/ length, although subtle differences in tubule lengths is
1+L46—47/2. The lateral teeth are smaller towards the observed.
edges ofthe radula (Figure 8).
Distribution: Islands of Sumatra and Borneo (Fig.
Pedal and Pallial Nerves, Pedal Aortic System 18).
One pair ofpallial and one ofpedal nerves run from Natural history: They were found on the forest floor, at
the central nervous system towards the posterior night and early in the morning, in onecasemore or less
extremity of the body cavity (Barker, 2001). They are clustered around a rotten log.
parallel and together from the central nervous system
until neartheheight ofthe bursacopulatrix. After, they Type-material: Holotype. INDONESIA, Borneo: P. E.
are parallel but separated from each other and run like Gerrard col. (BMNH 1880.10.6.4). Other materials.
that until the end ofthe body cavity. The pedal aortic MALAYWSIA, Sabah: Danum Valley Field Centre, ca.
artery begins at the level ofthe central nervous system 60 km of Lahad Datu, primary dipterocarp forest,
m
and runs between the nerves (centrally) until around ca. 300 asl. (04°58'00"N 117 48'00"E), 2 specimens,
the level ofthe bursa copulatrix. 2003.80.W, P. Craze & M. Schilthuizen col. (BORV
MOL 3409); Danum Valley Borneo Rainforest Lodge,
Pedal Gland 1 specimen, 2000.84.W, C. Rutjes col. (BOR/MOL
3410); Kota Kinabalu Distr., Poring Hot Springs,
m
The pedal gland, located on the anteriorextremity of 660 asl. (06°02'87"N 116°42'16"E), 1 specimen,
the sole under the head, is short and straight. It is 2001.59.W, M. Schilthuizen & P. Koomen col. (BOR/
broad in its proximal portion (in the aperture), MOL 3411); Interior Prov., Tenom (behind Perkasa
m
narrowing in the middle and with the posterior Hotel), 327-400 asl. (05°07'17"N 115°56'40"E), 1
extremity rounded and somewhat broadened (produc- specimen, 2003.89.W, M. Schilthuizen col. (BOR/MOL
ing a goblet-shape). It has two different areas: one 3438); Kinabalu National Park (Poring Hot Springs),
external lighter and one internal yellowish. The 500 m asl., 1 specimen, 2003/03/24 (2003.26.W), M.
posterior extremity of the gland receives a wide pedal Schilthuizen col. (BOR/MOL 3439).
gland artery (Coifmann, 1935). The lot deposited at Natural History Museum in
London (England) (BMNH 1884.1.10.1), which is
Reproductive System identified as a paratype of VI. flava, is not VI. flava.
In thisspecimen (adult), thathas not been dissected yet,
The bursa copulatrix (= spermatheca or gametolitic the penis (the main diagnostic characteristic) is
gland) is almost circular, but somewhat concave in the completely different. It is probably an unknown species
middle. It has a cylindrical duct a little longer than the ofVeronicellidae.
Page 168 The Veliger, Vol. 50, No. 3
Figures 13-17. Reproductivesystemin Valigwtasiamensis(ZMIZ01092). 13,completereproductivesystem; 14,detailofthebursa
copulatrix; 15-17, three different positions ofthe penis (cb, bursa copulatrix; db. duct ofthe bursa copulatrix; dd, distal posterior
deferens; df, vasdeferens; dp, proximal posteriordeferens; fc, fertilizationcomplex; fg, female genital pore; gb, albumen gland; gh,
hermaphroditic gland; gn, glans; he, honeycomb aspect structure; jd, canalis junctor; It, long tubules, md, middle deferens; ol,
spermioduct;ov, oviduct;pk, peak, inwhoseextremitythedeferensopens;pi,penialglandpapilla; pr,prostate; re,rectum;rp,penis
retractor muscle; st, short tubules; sv, seminal vesicle; vg, penis base). Scale bar: 1 mm.
S. R. Gomes et al., 2007 Page 169
• Valigunaflava a
^
"T" Valigunasiamensis
_|
I
Figure 18. Map showing the distribution of Valigunaflavaand Valigunasiamensis, considering bibliography records and the lots
recently collected in the north ofBorneo and China.
DISCUSSION on the reproductive system (as copulatrix bursa region
andpenial gland) arevery similarbetween both species.
TVIh.efylavsahaarned sVeIv.ersailamemnosripshoalroegiccleaalrlychcalroascetersipsetciicess., The body form differs between VI. flava and VI.
siamensis. The first one has an oval body while the
although important differences are also found between
other is longer and narrower, although in both species
them.
The penis is the structure that most notably thhyeposonloetwuimd.thUnisfosrmtaulnlaetretlhya,nthhaelfcothleorwaitditonh ocfanthneortigbhet
discriminates VI. flava from VI. siamensis and even compared since the specimens of VI. siamensis from
from the other species of the family. The dentate and
China were all completely discolored. The specimens of
serrated formation found in the distal extremity ofthe Vs. reticulatus examined by Thome (1984). and again
penis of VI. flava is very characteristic and it is found by us, were also discolored. Externally, VI. flava can be
only in this species ofVeronicellidae. In VI. siamensis, identified mainly by presence ofwide blackish spots in
the vas deferens also opens in a lateral peak and a well the notum, although there is some variation in the
hdaesvelaopfeodrmbaatsieonislfiokuenda."Bhuot,neiynctohmisb"spelcoiceastetdheinpentihse intVeIn.sitsyiaomfentshies,notpruemvigorusoluyndrceodleosrcartiiboend(FbiygurTeho1)m.e
distal extremity (Figures 13-17). It is a complex (1984) when he studied Va. reticulatus from Galle
structure, which is practically absent in juvenile (synonyms), is recorded for the first time to China
specimens. The bursa copulatrix region has also some (Yunnan and Guangxi) (Figure 18).
differences. In VI. siamensis, the bursa copulatrix duct
is extended in the medium region, assuming a domed Acknowledgements. Weare grateful to Centro de Microscopia
aspect. In VI. siamensis the bursa duct is fairly longer e Microanalises (CEMM) ofPontificia Universidade Catolica
than the copulatrix bursa, different from VI. flava. In do Rio Grande do Sul for the SEM photomicrographs; to
this species, the bursais small and assumesa formfrom Peter Koomen forthe photo ofthe living specimen; Suzete R.
spherical to elliptical. In both species, the canalis Gomes, Juliane Picanco and Jose W. Thome gratefully
jtuhnectbourrspaeniettsrelafte(sasininthmeabnuyrsaotchoeprulsapterciixesduocft,Venrootnii-n zaDecenksnegonrwvaltoeelfdvugilemlysenaftcionkanCnoicewinlatleidfsgiucepospeofriTtneacnfncroiolamlogs"iucCpoop.no"sretMlehfnornoNmoactSihceohinRlatolhyuadile-
cellidae). Netherlands Embassy, Kuala Lumpur.
The other internal characteristics are very similar
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