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of the
FLORIDA STATE MUSEUM
Biological Sciences
VOLUME 29 1983 NUMBER 3
NON-MARINE MOLLUSKS OF BORNEO
II PULMONATA: PUPILLIDAE, CLAUSILIIDAE
III PROSOBRANCHIA: HYDROCENIDAE, HELICINIDAE
FRED G. THOMPSON
AND
S. PETER DANCE
UNIVERSITY OF FLORIDA
GAINESVILLE
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NON-MARINE MOLLUSKS OF BORNEO
II PULMONATA: PUPILLIDAE, CLAUSILIIDAE
III PROSOBRANCHIA: HYDROCENIDAE, HELICINIDAE
FRED G. THOMPSON AND S. PETER DANCE1
ABSTRACT: The Bornean land snails of the families Pupillidae, Clausiliidae, Hydrocenidae, and
Helicinidae are reviewed based on collections from 38 localities in Sarawak and Sabah and on
previous records from the island. The following species are recorded: PUPILLIDAE-
Pupisoma orcula (Benson), Costigo puluiusculum (Issel) new combination, Costigo molecul-
ina Benthem-Jutting, Nesopupa moreleti (Brown), N. malayana Issel; Boysidia (Dasypupa)
salpinx new subgenus and species, B. (D.) paini new species. The following Malayan species
are referred to the subgenus Dasypupa: B. (D.) terae (Tomlin), B. (D.) megaphona (Benthem-
Jutting), B. (D.) elephas (Benthem-Jutting), and B. (D.) perigyra (Benthem-Jutting). CLAU-
SILIIDAE: Phaedusa borneensis (Pfr.), P. filicostata stenotrema new species. HYDROCE-
NIDAE: Georissa bangueyensis Smith, G. borneensis Smith, G. everetti Smith, G. gomanto-
nensis Smith, G. hosei Godwin-Austen, G. kobelti Gredler, G. niahensis Godwin-Austen, G.
saulae (Benthem-Jutting) new combination, G. scalinella (Benthem-Jutting) new combina-
tion, G. similis Smith, G. williamsi Godwin-Austen, G. hadra new species, G. pyrrhoderma
new species, and G. xesta new species. Names placed in synonymy include G. flavescens
Smith (= G. gomantonensis) and G. hungerfordi Godwin-Austen (= G. williamsi). No HELI-
CINIDAE were collected. Species previously recorded from the island are listed.
RESUMEN: Los caracoles terrestres de Borneo, pertenecientes a las families Pupillidae,
Clausiliidae, Hydrocenidae y Helicinidae fueron revisados en base a colecciones de 38
localidades en Sarawak y Sabah, asi como a registros previous de la isla. So registraron las
siguientes species: PUPILLIDAE: Pupisoma orcula (Benson), Costigo puluiusculum (Issel)
nueva combinaci6n, Costigo moleculina Benthem-Jutting, Nesopupa moreleti (Brown), N.
malayana Issel; Boysidia (Dasypupa) salpinx nuevo subgenero y especie, B. (D.) paini nueva
especie. Las siguientes species malayas se refieren al subgenero Dasypupa: B. (D.) terae
(Tomlin),B. (D.) megaphona (Benthem-Jutting), B. (D.) elephas (Benthem-Jutting) yB. (D.)
perigyra (Benthem-Jutting). CLAUSILIIDAE: Phaedusa borneensis (Pfr.), P. filicostata
stenotrema nueva especie. HYDROCENIDAE: Georissa bangueyensis Smith, G. borneensis
Smith, G. everetti Smith, G. gomantonensis Smith, G. hosei Godwin-Austen, G. kobelti
Gredler, G. niahensis Godwin-Austen, G. saulae (Benthem-Jutting) nueva combinaci6n, G.
scalinella (Benthem-Jutting) nueva combinaci6n, G. similis Smith, G. williamsi Godwin-
Austen, G. hadra nueva especie, G. pyrrhoderma nueva especie y G. xesta nueva especie.
Nombres ubicados en sinonimia incluyen G. flavescens Smith (=G. gomantonensis) y G.
hungerfordi Godwin-Austen (= G. williamsi). No se colectaron especimenes de HELI-
CINIDAE. Se mencionan species previamente registradas en la isla.
TABLE OF CONTENTS
Introduction ................................................. ............. 102
Acknowledgements ................. .............. .... ..................102
II Pulmonata
Pupillidae ..................... .................................... ..........103
Clausilliidae ........... ...........................................109
'Dr. Thompson is Curator in Malacology at the Florida State Museum, University of Florida, Gainesville, FL32611 U.S.A. Dr.
Dance is at South Bank House, Broad Street, Hay-on-Wye (via Hereford) HR3 5DB, United Kingdom.
THOMPSON, F. G. and S. P. DANCE. 1983. Non-Marine Mollusks of Borneo. II Pulmonata:
Pupillidae, Clausiliidae. III Prosobranchia: Hydrocenidae, Helicinidae. Bull. Florida State
Mus., Biol. Sci. 29(3):101-152.
BULLETIN FLORIDA STATE MUSEUM
III Prosobranchia, Archeogastropoda
Hydrocepidae ................................................................ 112
Helicinidae ................................................................ 128
Literature Cited ................................................................. 129
INTRODUCTION
In 1970 the first of a projected series of papers was published on
mollusks of Borneo (Dance 1970), based upon collections made by Dr. G.
E. Wilford and his associates of the Geological Survey, Malaysia. It dealt
with the Streptaxidae. This paper, comprising parts II and III of the series,
covers two families of Pulmonata as well as the Archeogastropoda.
The land snail fauna of Borneo is poorly known, even though more than
350 species have been reported from the island (Martens 1908, Solem 1964).
The majority of the species recorded are endemic and have been collected
in Sarawak, Brunei, and Sabah, an area comprising about a third of the
island. The southern portion, Kalimantan, is virtually unknown malacolog-
ically, except for a few species collected randomly at coastal localities by
travelers. Many species have been recorded from the island only once, and
their original descriptions frequently lack important diagnostic details.
Few distributional data have been available, and many species have been
reported merely from "Borneo."
The collections on which the present studies are based originated from
the same places in northern Borneo from which most species have been
recorded previously (Fig. 1). These collections are rich in small or minute
taxa that usually frequent limestone, and most are abundantly represented
(Dance 1970:149-150). Larger species, such as the Cyclophoridae, Camae-
nidae, and Bradybaenidae, are poorly represented, and our material of
these does not constitute a major addition to the known fauna. Almost all of
our material consists of dried shells. No live specimens were preserved for
anatomical studies.
As was anticipated, our material contains many new taxa. On the other
hand many taxa previously recorded from Borneo are missing from our
material. From this we estimate that less than 25% of the Bornean fauna is
known. Our accounts of particular families list all species recorded from
Borneo and satellite islands to facilitate future investigations by other
workers.
ACKNOWLEDGEMENTS
We are grateful to the following people who have helped us in various ways. Dr. Alan
Solem (Field Museum of Natural History), Dr. Ronald Janssen (Senckenbergischen Naturfor-
schenden Gesellschaft) and Dr. Fred Naggs (British Museum, Natural History) loaned speci-
mens essential to this report. Mr. Terrence E. Crowley compared specimens from our
material with types in the British Museum (Natural History). Dr. F. E. Loosjes, Wageningen,
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 103
Netherlands, compared relevant specimens of Clausiliidae. Kurt Auffenberg, Florida State
Museum, helped prepare illustrations and performed numerous other minor but essential
tasks. The excellent drawings of Boysidia were made by Wendy Zomlefer, Staff Illustrator,
Florida State Museum. Micrographs of hydrocenid shells were made by Kurt Auffenberg with
an HITACHI 5-415A scanning electron microscope in the Department of Zoology, University
of Florida.
II PULMONATA
PUPILLIDAE
The classification used in this paper follows Pilsbry (1927:vii-xii) and
Baker (in Pilsbry 1935:191-209). Thiele (1931:503-514) disregarded name
priorities and used VERTIGINIDAE Stimpson 1851 to replace PUPILLI-
DAE Turton 1831. Zilch (1959:146-174), who rearranged the pupillid gen-
era among five families on the basis of shell characteristics, employed a
different nomenclature for family and subfamily units.
NESOPUPINAE STEENBERG 1925
Pupisoma orcula (BENSON)
This widely distributed species has not been reported from Borneo
previously. It was found at the following stations. SABAH: Kinabatangan
Valley, Batu Tumanggong Besar, 1 mi downstream from Sakau,
05034'N, 118020'E, (UF 35936). SARAWAK: First Div., Bau area, Gunong
Kapor (UF 35933); First Div. S of Serian, Gunong Selabor, 0055'N,
110025'E, (UF 35939,35931); Fourth Div. SW of Marudi, Beluru area, Bukit
Vrong, 0400(YN, 114012'E, (UF 35932); Fourth Div., Lower Tutoh Valley,
nr. Gunong Mulu; Melinau Paku Cave, 04003'N, 114048'E, (UF 35935);
Fourth Div., Lower Tutoh Valley, nr. Gunong Mulu, small hill towards
headwaters of Melinau Paku Riv., 04003'N, 114048'E, (UF 35934); Fourth
Div., Lower Tutoh Valley, nr. Gunong Mulu, junct. Melinau Riv. and
Melinau Paku Riv. (UF 35937); Fourth Div., small hill N of lower Melinau
Riv. 04003'N, 114048'E, (UF 35938); Fourth Div., Tatau Valley, Bukit
Sarang, 02045'N, 113002'E, (UF 35929); Fifth Div., Gunong Budah, Meda-
lam Valley, trib. to Limbang Riv., 04008'N, 115005'E, (UF 35930).
Costigo pulvisculum (Issel)
Helix pulvisculum Issel 1874; Ann. Mus. Civ. Stor. Nat. Genova 6:406; pl. 5,
figs. 24-27.-Tryon 1887; Man. Conch 3:191; pl. 42, figs. 55-57.
Pupisoma pulvisculum(Issel), Smith 1898; Proc. Malac. Soc. London 3:
29.-Pilsbry 1920; Man. Conch. 26:30-31.
TYPE LocALITY.-Sarawak.
DISTRIBUTION.-Now recorded from the following localities. SABAH:
southeast portion, 15 mi NNW Kalabakan, head of Cowie Harbour (UF
35940); Kinabatangan Valley ca 1 mi downstream fr. Laab
BULLETIN FLORIDA STATE MUSEUM
0527'N, 11759'E, (UF 35941). SARAWAK: Fifth Div., Limbang Valley, ca
2 mi NE junct. Medalam Riv. and Limbang Riv. 04020'N, 11458'E, (UF
35942). Smith (1898:29) also recorded the species from Lombok.
REMARKS.-Pilsbry (1920:30-31) suggested that this species is synonym-
ous with P. orcula (Benson), but we do not agree. The specimens before us
comply well with Issel's description and figures. They are about half the
size of P. orcula, they lack the spiral sculpture typical of Pupisoma, and
they have a granulose-striate surface, as in Costigo. Consequently, we
consider pulvisculum to be an edentate species of Costigo.
Costigo moleculina benthem-Jutting
Costigo moleculina Benthem-Jutting 1940; Treubia 17:331; figs. 1-2.-1952;
Treubia 21:352-353; fig. 31.
TYPE LOCALITY.-West Java, forest between Tjisolok (Pelabuanratu)
and the hot springs (Tjipanes), some miles inland.
DIsTRBUTION.-Previously known only from the type locality. We now
add the following station in Borneo. SABAH: Kinabatangan Valley, Batu
Tumanggong Besar, 1 mi downstream from Sukau, 05034'N, 118020'E (UF
35943).
REMARKS.-Apart from their smaller size, our two specimens agree well
with the original description. Costigo gracilis (Haas) from Luzon is similar
in appearance and intermediate in length, though more slender (Haas
1937:9-10, figs. 21-23). Both species differ from other Costigo species by
having only two teeth within the aperture, a columellar, and a parietal
lamella. Additional material may show that the two forms are conspecific.
Measurements in mm for our specimens and the types of the two nominate
forms are:
length width
Costigo gracilis, type 2.0 1.25
Costigo moleculina, type 2.2 1.4
Costigo moleculina, Sabah 1.83 1.27
Costigo moleculina, Sabah 1.77 1.24
The following Nesopupinae also have been recorded from Borneo.
Nesopupa moreleti (Brown)
Pupa moreleti Brown 1870; J. Conchyl. 18:393.-Pfeiffer 1877; Monogr.
Helic. Viv. 8:391.-Smith; Ann. Mag. Nat. Hist. (6), 13:458.
Nesopupa moreleti (Brown) Pilsbry 1920; Man. Conch. 25:339-340; pl. 32,
figs. 1-3, 6.
TYPE LocALITY.-Labuan Island, off the north coast of Borneo.
DIsTRBUTION.-Pilsbry (1920:339-340) recorded the species from the
Philippine Islands.
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 105
Nesopupa malayana (Issel)
Vertigo malayana Issel 1874; Ann. Mus. Civ. Stor. Nat. Genova 6:416; pl. 5,
figs. 30-32.
Nesopupa malayana (Issel) Pilsbry 1920; Man. Conch. 25:342-343; pl. 32,
figs. 14-16.
TYPE LOCALITY.-"Borneo."
DIsTRIBUTION.-Pilsbry (1920:343) recorded the species from the Phil-
ippine Islands.
GASTROCOPTINAE PILSBRY 1917
Boysidia and Hypselostoma are closely related genera that differ only
in the position of the aperture and in sculpture. Boysidia has an adnate
peristome and lacks spiral striations. Hypselostoma has a solute peristome
and has spiral striations in addition to axial sculpture. When Pilsbry (1917)
reviewed Boysidia and Hypselostoma, most known species were assign-
able to either genus with little difficulty. The generic affinities of a few
were less clear. Since then the number of known taxa has more than
doubled (Rensch 1932, 1934; Haas 1937; Tomlin 1939; Benthem-Jutting
1949, 1960, 1961, 1962; this paper). Species were assigned to either genus
because of the position of the aperture. No attention was given to other
characteristics of the shell, nor was any allowance made for the possibility
that a solute aperture may have evolved more than once from the typical
condition in Boysidia. Consequently both genera contain a heterogeneous
assortment of species. Neither is comprehensible in its present composi-
tion, and further generic subdivision is necessary. Three species occur in
Borneo which exemplify this problem.
Boysidia Ancey 1981
The following three Bornean species are placed in a new subgenus
because they cannot be assigned to Boysidia s. s. nor to Hypselostoma
Benson 1856. The reasons for this are as follows. Characteristics of the
sculpture are similar to Boysidia, the advanced apertures of two are typical
of Hypselostoma, and characteristics of the aperture barrier are a typical
for either genus, but apparently derive from the generalized pattern that
occurs in Boysidia. The new subgenus also accommodates some Malayan
species.
Dasypupa NEW SUBGENUS
TYPE SPECIES: Boysidia (Dasypupa) salpinx new species.
A subgenus of Boysidia in which the columellar lamella is tubercular
and deeply recessed within the aperture (Figs. 2, 4), not blade-like as in
Boysidia Ancey or Paraboysidia Pilsbry. The parietal-angular lamellae are
BULLETIN FLORIDA STATE MUSEUM
concrescent into a short, flexed, bilobed blade (Fig. 5). The shell is conical,
spire depressed-conical to elongate-conical, with minute rough periostra-
cal fimbriations parallel to the lines of growth (Figs. 2-6). Spiral striations
are absent in contrast to Hypselostoma, in which some species of Dasypupa
have been placed. Occasional raised spiral threads may be present due to
fusion of short segments of granular sculpture. The peristome may be
adnate to or solute from the previous whorl.
ETYMOLOGY.-Dasypupa, from the classical Greek dasys meaning
shaggy, plus pupa, and alludes to the rough microsculpture of the shell.
Boysidia (Dasypupa) salpinx NEW SPECIES
SHELL (Figs. 2-6, 7-8).-Small-sized, about 1.8-2.1 mm high and 2.2-2.6
mm wide. Spire depressed-conical. Shell about 0.70-0.89 times as high as
wide. Opaque; reddish-brown with a dull sheen. Umbilicus open, about
1/4 width of shell, showing previous whorl (Fig. 3). About 4 whorls.
Embryonic whorls 1.5, protruding, rounded, sculptured with a dense mesh
of fine granules. Succeeding whorls angular at periphery, sculptured with
irregularly spaced growth striations that are interspersed with finer, rough
periostracal threads (Fig. 6). Last whorl bluntly angular and flattened
above and below periphery; weakly furrowed above and below angle
along last quarter whorl. Last half whorl with a furrow along umbilical
wall. Aperture projecting forward for a short distance beyond previous
whorl, tilted upward slightly due to ascent of last quarter whorl (Fig. 8).
Aperture about as wide as high; nearly quadrangular, with a rounded base.
Peristome broadly expanded. Interior of aperture with 4-6 teeth. Parietal-
angular lamella fused and extending forward nearly to edge of aperture.
Palatal plica, basal plica, and columellar lamella short, compressed-
tubercular and deeply immersed in aperture (Fig. 4). Occasionally a small
tubercular supracolumellar and subcolumellar lamella may also be present.
Outer surface weakly indented over palatal plica.
Measurements in mm of holotype and three paratypes (UF 35946)
follow.
length width aper. h. aper. w. whorls
Holotype 2.02 2.26 1.24 1.27 4.0
Paratype 1.74 2.51 1.24 1.30 3.9
Paratype 1.86 2.17 1.15 2.21 4.0
Paratype 2.11 2.60 1.21 1.27 4.1
Measurements for three specimens from UF 35945 are:
length width aper. h. aper. w. whorls
2.11 2.33 1.24 1.30 4.0
2.57 2.79 1.30 1.33 4.4
2.70 2.82 1.55 1.47 4.5
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 107
TYPE LOCALITY.-Gunong Subis, a limestone massif about 40 mi SW of
Mira, Niah area, Fourth Div., Sarawak, Borneo, 030 51'N, 113045'E. HOLO-
TYPE: UF 35944; collected in 1962 by G.E. Wilford. Paratypes: UF 36946
(26 specimens), UF 36281 (2 SEM spec.), S. Dance coll. (8 spec.); British
Museum (Natural History) (2); Sarawak Museum, Kuching (2); same data
as holotype.
DISTRIBUTION.-Known from northwestern Borneo from the type local-
ity and two other stations widely separated from it. SARAWAK: First Div.,
Bau area, entrance of main cave at Gunong Kapor, 01017'N, 11004'E (UF
35945); First Div., Gunong Selabor, an isolated limestone mountain S of
Serian, 00055'N, 11025'E (UF 35947, 35948).
REMARKs.-This snail occurs at three widely separated localities, yet we
can find no significant differences that will distinguish any of the popula-
tions. Specimens from Gunong Kapor (UF 35945) tend to be larger and
more frequently have a subcolumellar lamella, but other characteristics of
the shell are typical for the species.
ETYMOLOGY.-Salpinx, from the classical Greek meaning a trumpet, in
allusion to the forward projecting, flared aperture.
Boysidia (Dasypupa) paini NEW SPECIES
SHELL (Figs. 9, 10).-Medium-small, 2.67-2.95 mm long; broadly coni-
cal, major width 0.83-1.01 times length of shell; minor width 0.73-0.84 times
length of shell. Umbilicus narrow, about 1/6-1/9 times width of shell; base
of last whorl angulate around umbilicus. Color reddish-brown, lusterless,
opaque. Whorls 4.0-4.4. Embryonic whorls 1.5, protruding and rounded;
with very fine granular sculpture. Succeeding whorls angular along
periphery and with a deeply impressed suture; flattened above and below
peripheral angle. Last quarter whorl ascending slightly; with a shallow
impressed zone above and below peripheral angle, weakly constructed
behind aperture. Whorls sculptured with growth striations that are
interpersed with coarse, elongate, periostracal granules aligned parallel to
growth striations. Aperture slightly oblique, rhomboid, higher than wide,
about 0.52-0.60 times height of shell; interior hyaline reddish-brown. Peris-
tome broadly expanded, tightly appressed against previous whorl (Fig.
10). Interior of aperture with four deeply immersed teeth. Parietal-angular
lamellae fused and bilobed; angular portion low and flexed to the right.
Palatal plica, basal plica, and columellar lamella small, nearly equal in size,
compressed-tubercular. Outer surface weakly indented over palatal plica.
Measurements in mm of three specimens are:
maj. min.
length width width aper. h. aper. w. whorls
Holotype 2.95 2.64 1.92 1.64 1.49 4.4
Paratype 2.91 2.95 2.23 1.71 1.61 4.2
Paratype 2.67 2.20 1.95 1.46 1.24 4.0
BULLETIN FLORIDA STATE MUSEUM
Specimens from Gunong Budah (UF 35950) differ by being slightly
narrower and having a narrower umbilicus about 1/9 the width of the shell.
Other aspects of the shell are typical.
TYPE LOCALITY.-Foot of cliff about 400 yards north of the main
entrance to Deer Cave, Melinau Paku Valley, Fourth Div., Sarawak, Bor-
neo, 04005'N, 114053'E. HOLOTYPE: UF 35949; collected by J. R. D. Wall
in 1964. PARATYPES: UF 35951 (6); S. P. Dance Collection (3).
DISTRIBUTION.-Known in Borneo from one other station near the type
locality. SARAWAK: Fifth Division, SW flank Gunong Budah, Medalam
Valley, a tributary of the Limbang River; 04008'N, 115005'E (UF 35950).
REMARKS:-Boysidia (D.) pain is distinguished from other Dasypupa
by its broadly conical shape and adnate aperture. It is similar in appearance
to B. salpinx, but the latter species has a solute aperture and a wider
umbilicus. It is less similar in shape to B. (D.) procera, and differs in other
respects as discussed under that species.
ETYMOLOGY.-Boysidia (Dasypupa) paini is named for Mr. Tom Pain of
London, England, well known as the author of many papers on tropical
snails.
Boysidia (Dasypupa) procera NEW SPECIES
SHELL (Figs. 11, 12).-Medium-sized, about 3.1-3.4 mm long; elongate-
conical, major width 0.69-0.74 times height of shell, minor width 0.59-0.60
times height of shell; sides of spire straight. Shell lusterless, opaque,
reddish-brown, with 5.0-5.1 whorls; Embryonic whorls 1.5, rounded, pro-
truding. Succeeding whorls with a deeply impressed suture, angulate at the
periphery, nearly flattened above periphery. Last whorl slightly con-
stricted behind peristome, with a low cord-like peripheral keel and a
shallow, impressed groove above keel. Base of last whorl with an angulate
rim around umbilicus, which is about 1/5 width of shell. Postembryonic
whorls sculptured with oblique, irregular growth striations and rough
granular periostracal growth threads. Aperture free from and slightly
advanced beyond previous whorl, subcircular, nearly as high as wide,
about 0.39-0.45 times length of shell. Peristome broadly expanded; outer lip
and columellar lip oblique and nearly parallel. Interior of aperture with 4-5
deeply immersed teeth; parietal-angular lamella bilobed with angular
segment lower, flexed to the right and advancing nearly to edge of aper-
ture; palatal plica, basal plica, and columellar lamella subequal, small and
compressed-tubercular; a very small supracolumellar lamella may also be
present. Outer surface weakly indented over palatal plica.
Measurements in mm of two complete specimens are:
maj. min.
length width width aper. h. aper. w. whorls
Holotype 3.41 2.51 2.02 1.55 1.58 5.1
Paratype 3.10 2.14 1.86 1.33 1.33 5.0
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 109
TYPE LOCALITY.-Southwest flank of Gunong Budah, Medalam Valley,
a tributary of the Limbang River, Fifth Division, Sarawak, Borneo 04008'N,
115005'E. HOLOTYPE: UF 38022; collected by G. E. Wilford in 1966.
PARATYPES: UF 38023 (1 complete specimen and two fragments); same
data as for the holotype.
DISTRIBUTION.-Known only from the type locality.
REMARKS.-This species is distinguished by its elongate shell, solute
aperture, nearly round peristome and wide umbilicus. It is most similar to
B. (D.) paini, which occurs at the same locality, but the latter species has a
shorter, broader shell, fewer whorls, an adnate peristome, a narrower
umbilicus, and a rectangular-shaped peristome. The aperture also is rela-
tively larger, being more than half the length of the shell.
ETYMOLOGY.-The specific epithet procera is from the Latin procerus
meaning drawn out or elongate, in allusion to the relatively elongate shell
compared to other Bornean species.
The following Malayan species are referred to the subgenus Dasypupa.
Boysidia (Dasypupa) terae (Tomlin)
Hypselostoma terae Tomlin 1939; J. Conch. 21:146; pl. 12, Fig. 2.-
Benthem-Jutting 1949; Bull. Raffles Mus. (19):56.-1949; Bull. Raffles
Mus. (21): 21; figs. 10.
TYPE LOCALITY.-Bukit Chintamani, Penang.
Boysidia (Dasypupa) megaphona (Benthem-Jutting)
Hypselostoma megaphona Benthem-Jutting 1949; Bull. Raffles. Mus.
(21):21-23; figs. 11.
TYPE LOCALITY.-Buki Charas, near Kuantan, Penang.
Boysidia (Dasypupa) elephas (Benthem-Jutting)
Hypselostoma elaphas Benthem-Jutting 1949; Bull. Raffles Mus. (21):23-24;
figs. 12.
TYPE LOCALITY.-Bukit Tenggek, Penang.
Boysidia (Dasypupa) perigyra (Benthem-Jutting)
Hypselostoma perigyra Benthem-Jutting 1949; Bull. Raffles Mus. (21):25-
26; figs. 14.
TYPE LOCALITY.-Bukit Tabun, Kanching, Selangor.
CLAUSILIIDAE
Loosjes (1953) gave a thorough review of the CLAUSILIIDAE of the
Indo-Australian region. He recorded five species as occurring in Borneo:
BULLETIN FLORIDA STATE MUSEUM
Phaedusa borneensis (Pfeiffer) (pp. 28-31)
Phaedusa dorsoplicata Loosjes (pp. 31-33)
Phaedusa filicostata filialis (Martens) (pp. 50-53)
Pseudonenia scalariformis Loosjes (pp. 78-79)
Paraphaedusa schwaneri (Martens) (pp. 144-147)
We have two species in our material from northern Borneo. One is well
known. The other represents a new subspecies.
Phaedusa borneensis (Pfeiffer)
(Figures 13-15, 19)
Three lots of this species come from within the range recorded by
Loosjes. SARAWAK: First Div., south of Serian, Semabang entrance to
Lobang Batu Cave, western side of Gunong Selabor, 00055'N, 110025'E
(UF 35928); First Div., Bau Area, Gunong Kapor, entrance to main cave,
01017'N, 110004'E (UF 35924); Fourth Div., Niah area, Gunong Subis, ca 40
mi SW of Miri, 03051'N, 113045'E (UF 35927).
Phaedusa filicostata stenotrema NEW SUBSPECIES
SHELL (Figs. 16-18,20-21, 25).-Terrete-fusiform, sides of spire slightly
convex, nearly straight-sided, medium-sized, about 17-21 mm long, about
4.28-5.26 times as long as wide. Lower whorls narrow, last three whorls
comprising 0.62-0.67 length of shell. Brown in color, barely transparent,
shiny; interior of aperture light brown; peristome white. Whorls 9.5-11.2,
weakly arched with a moderately impressed suture. A barely visible sub-
sutural spiral cord occurs along lower whorls. Embryonic shell with 3
whorls, sculptured with fine thread-riblets that weakly crenulate the
suture. Following whorls sculptured with similar oblique thread-riblets
that are frequently poorly defined. Riblets becoming stronger and more
widely spaced on last whorl. Penultimate whorl with 10-15 thread-
riblets/mm; back of last whorl with 8-11 riblets/mm. Aperture sub-
elliptical, narrow, about 0.68-0.75 times as wide as high; free from and
projecting forward from previous whorl (Figs. 24, 25). Plane of aperture
nearly parallel to axis of shell. Peristome broadly expanded, narrowest
above sinulus, widest along basal lip. Edge of outer lip straight-sided or
weakly concave. Sinulus slightly deeper than wide, pointed at apex. Super-
ior lamella thick, oblique, slightly converging toward outer lip, short,
confined to parietal edge of aperture, widely separated and offset to the
center from spiral lamella. Inferior lamella extending into aperture for one
whorl, converging toward, but separated from upper end of spiral lamella;
inferior lamella gradually decreasing in height posteriorly and anteriorly
where it flexes to descend partially down columellar side of aperture.
Columellar lamella lower and shorter than inferior lamella, not visible from
aperture. Spiral lamella about as high as inferior lamella; oblique, sloping
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 111
toward principal plica; spiral lamella beginning on parietal wall 1/4 whorl
behind aperture and continuing for 3/4 whorl to ventral side. Principal
plica extending from middle of right side to a little to left of dorsal side.
Palatal barrier consisting of 4-7 palatal plicae confined to left side in a
crescent that may be on a lunella-like callus (Figs. 24, 25). Upper palatal
plica about 1/3 as long as principal plica and diverging from it. Lowest
plica short, and oblique. Other plicae short and tubercular or absent.
Clausilium plate 2.1 mm long, 1.3 mm wide; sides diverging to widest point
about 1/3 of distance from end, weakly pointed and folded (Figs. 22,23).
Pedicel long and slender, lying at about 700 to plate (clausilium from
specimen 19.5 mm long).
Measurement in mm of three specimens are:
length width aper. h. aper. w. whorls
Holotype 20.5 3.9 4.6 3.2 11.2
Paratype 17.1 4.0 4.4 3.3 9.5
Paratype 19.1 3.9 4.6 3.4 10.8
TYPE LOCALITY.-Melinau Limestone, near Gunong Mulu, lower Tutoh
Valley, Fourth Division, Sarawak, Borneo (04003'N, 114048'E). HOLO-
TYPE: UF 36426, collected by G. E. Wilford in 1961. PARATYPES: UF
35926 (4 complete specimens, 5 fragments), Rijksmuseum van Natuurlijke
Historic 55518 (3 specimens); S. P. Dance (1 specimen); same data as
holotype.
DISTRIBUTION.--Known only from the type locality.
REMARKS.-Phaedusa filicostata (Stoliczka) includes six other subspe-
cies (Loosjes 1953: 38-53). Five are confined to the Malayan Peninsula. One,
P. f. filialis (von Martens) is found in southeastern Kalimantan, Borneo. P. f.
stenotrema is most similar to P. f. tenuicosta (Nevill), a subspecies widely
distributed in the Malayan Peninsula. It differs from tenuicosta by its
usually smaller size, relatively more obese shell with a lower L/W ratio,
lower whorl count and narrower aperture in which the outer lip is straight-
sided or slightly concave in outline. In addition the axial riblets of steno-
trema are nearly straight in contrast to the wavy riblets of tenuicosta.
Loosjes gave measurements for eleven population samples of tenuicosta.
His sample of smallest specimens from Kelantan are as narrow as steno-
trema, but they are longer and have a higher whorl count. P. f. filialis has
little similarity to P. f. stenotrema. It is a much larger subspecies with flat
whorls and heavy palatal plicae (Loosjes 1953:50-53). P. f. stenotrema is
sympatric with P. borneensis (Pfr). The latter is distinguishable from
stenotrema by its swollen neck and its more rugose sculpture (Figs. 13-15,
19).
ETYMOLOGY.-stenotrema, from the classical Greek stenos meaning
narrow, and trema meaning hole, alluding to the narrow aperture charac-
teristic of the new subspecies.
BULLETIN FLORIDA STATE MUSEUM
III PROSOBRANCHIA, ARCHAEOGASTROPODA
Two families are represented in the Bornean fauna, the HYDROCE-
NIDAE and the HELICINIDAE. Their classification within the ARCHEO-
GASTROPODA follows Thompson (1980). As neither family has been
reviewed for many years, complete synonymies are given for the Bornean
species.
HYDROCENIDAE
INTRODUCTION
Hydrocenids are an important component of the land snail fauna of
Borneo. More species are found here than are known from any other area of
comparable size. Seven genus-group taxon names have been proposed for
the Hydrocenidae: Hydrocena Pfeiffer 1847, Georissa Blanford 1864,
Chondrella Pease 1871, Georissops Pilsbry and Hirase 1908, Omphalorissa
Iredale 1933, Monterissa Iredale 1944, Petrorissa Kuroda and Habe, in
Habe 1949. No review of the family has been made since 1876 (Pfeiffer),
and there is no general agreement as to the status of the generic names. For
purposes of this paper we follow the traditional use of Georissa for south-
east Asiatic and Pacific species, with Chondrella and Georissops as subgen-
era. Petrorissa is a synonym of Georissa s.s. All of the Bornean species
belong in Georissa s.s. because of their opercular structure. The shell
sculpture varies greatly between the Bornean species and nearly encom-
passes the range of variation known for the family. The protoconch charac-
teristically is elongate-ovate in shape and is set off from the postembryonic
shell as a small dome-like cap. It is minutely sculptured with a dense mesh
of fine reticulating raised threads (Figs. 26-31). No characteristics of spe-
cific or group significance in this sculpture were noticed among the Bor-
nean species. Resorption of the internal whorl septa occurs. In all Bornean
species the septum lies between 1.0-1.5 whorls behind the aperture (Figs.
32-38). The remnant of the the axis is a short tube-like cavity, which
apparently accommodates a caecum on the ventral side of the stomach. In
Hydrocena s.s. the septum extends into the shell for only a half whorl and
the axial tube is reduced to a triangular pit.
Opercula are available for all of the species studied. No conspicuous
characteristics of specific distinction are apparent. The opercula are con-
centric, calcareous, and broadly ovate. The outer surface is smooth with a
few fine concentric striations. The inner surface has a peg originating from
the lower left quadrant. The peg is moderately stocky and projects inward
at an angle of about 250 to the face of the operculum. Each operculum is
smaller than the aperture and is retractable within the shell for about a
quarter whorl. The opercula of various species are illustrated (Figs. 39-42).
Thirteen species have been described from Borneo; eight are repres-
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 113
ented in the material at hand. Most of the earlier descriptions are brief,
some are in Latin, and their accompanying illustrations are sketchy and
uninformative. It was necessary in many cases to compare our material
with type specimens in order to establish correct identifications, for this
could not be determined from the literature. We find it essential to redes-
cribe the species represented in our material. A list of previously named
species recorded from Borneo and satellite islands is given below along
with their current status.
Previous authors
Georissa bangueyensis Smith 1895
Georissa borneensis Smith 1895
Georissa everetti Smith 1895
Georissa flavescens Smith 1895
Georissa gomantonensis Smith 1893
Georissa hosei Godwin-Austen 1889
Georissa hungerfordi
Godwin-Austen 1889
Georissa kobelti Gredler 1902
Georissa niahensis Godwin-Austen 1889
Hydrocena saulae Benthem-Jutting
1966
Hydrocena scalinella Benthem-Jutting
1966
Georissa similis Smith 1893
Georissa williamsi Godwin-Austen 1889
This report
valid species (?)
valid species
valid species
syn. of G. gomantonensis Smith
valid species
valid species
syn. of G. williamsi G.-A.
valid species
valid species (?)
Georissa saulae (Benthem-
Jutting), valid species
Georissa scalinella (Benthem-
Jutting), valid species
valid species
valid species
The following new taxa are added to this list
Georissa hadra new species
Georissa pyrrhoderma new species
Georissa xesta new species
No doubt additional species remain to be discovered in northern Bor-
neo. Apparently some species are restricted to single isolated limestone
hills, and large areas of suitable habitat have not yet been explored for
mollusks. As Kalimantan is virtually terra incognita for mollusks our con-
cepts of Bornean hydrocenids will need to be revised eventually.
SPECIES RELATIONSHIPS
The Bornean Georissa are divisible into four species groups which are
rather well defined by shell characteristics.
1.) The hosei group: Medium- to large-sized species (2-4 mm) with
slender apical whorls separated by a very deep suture, and with a
peripheral spiral row of obliquely compressed nodes (other spiral
rows of nodes may be present). G. hadra; G. hosei, G. kobelti, G.
BULLETIN FLORIDA STATE MUSEUM
scalinella, G. saulae, and perhaps G. niahensis. Related species
occur in the Philippine Islands. G. monterosatiana from Malaya
may also belong here.
2.) The borneensis group: Medium-sized (about 2 mm), broadly coni-
cal species with reddish shells. Sculpture consisting of growth
striations and/or spiral threads (a spiral row of fluted nodes may
be present on the shoulder). G. borneensis and G. pyrrhoderma.
Related species are not known from other areas.
3.) The everetti group: Medium-sized (about 2 mm), broadly conical
species, citron yellow in color, sculptured with spiral or obliquely
spiral threads and growth striations or threads. G. everetti and G.
gomantonensis. Similar species are not known from adjacent
islands or the mainland.
4.) The williamsi group: Small- to medium-sized species (1.5-3.0 mm)
with a narrowly conical shell, light brown in color, sculptured with
spiral threads. Growth striations and riblets may also be present. G.
williamsi, G. xesta, G. similis, and G. bangueyensis. Similar species
occur on other islands in Indonesia, the Philippine Islands, and
Southeast Asia.
It is not realistic to deduce zoogeographic patterns of relationships at
present. Hydrocenids are small species seldom collected, and the mollus-
can fauna of vast expanses of Southeast Asia remains unknown. Any faunal
comparisons made at present would be highly tenuous.
ECOLOGICAL DEPLOYMENT
Species of Georissa live on limestone or in close association with cal-
careous terrain. Saul (1966:7) reported that the Bornean Hydrocena(=Geo-
rissa) scalinella (Benthem-Jutting) and H. (=G.) saulae (Benthem-Jutting)
occur on moist limestone among mosses and other plants in cool, shaded
places. Kurt Auffenberg informs me that on Luzon he found G. rufescens
Moellendorff only on damp limestone in cool, shaded habitats. Some
species are adapted to a wider array of microhabitats. Berry (1965)
reported that the Malayan G. monterosatiana Godwin-Austen and Nevill is
found on mossy limestone, bare rocks, soil, and tree trunks near limestone
hills. Berry (1961) stated that G. monterosatiana feeds on algae, fragments
of moss, and lichens. None of the species reported in this paper was
collected alive, and no data are available on their ecological deployment.
Judging from the abundance and distribution of the material at hand, some
species, such as G. hosei, G. gomantonensis, G. williamsi, and G. similis, are
generalized in their deployment, similar to G. monterosatiana. Apparently,
other species such as G. hadra, G. pyrrhoderma, G. scalinella, and G.
saulae, are more restricted, for they are known only from single samples
from localities where several collections were made.
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 115
BORNEAN SPECIES ERRONEOUSLY DESCRIBED As HYDROCENIDAE
cornea Pfeiffer, Hydrocena; Proc. Zool. Soc. London 1854:308. (Type
locality: Borneo and Bashee Island). = A synonym of Assiminea subcor-
nea Neville 1885; ASSIMINEIDAE (Abbott 1958:267).
glabrata Pfeiffer, Hydrocena; Proc. Zool. Soc. London 1854:308. (Type
locality: Island of Borneo). = Omphalotropis glabrata (Pfeiffer), ASSI-
MINEIDAE (Abbott 1958:268)
THE BORNEAN SPECIES
Georissa hadra, NEW SPECIES
SHELL (Figs. 43-46).--Relatively large, about 3-4 mm long. Spire pink,
grading to yellowish-orange below. Elongate-conical with an attenuate
apex due to the deeply impressed suture; spire straight-sided. Shell about
0.66-0.75 times as wide as high. Shell wall very thick (Fig. 32) with 3.5-4.1
whorls in larger specimens (3.6 in holotype). Whorls flattened above
periphery and narrowly shouldered along suture. Protoconch subcircular,
superficially smooth. Subsequent whorls with numerous fine spiral lira-
tions and 1-3 rows of enlarged and obliquely flattened spiral nodes (Fig. 45,
46). Nodes most distinct on spire; usually reduced in size or obsolete on
body-whorl; occasionally another spiral row of nodes may be present along
outer edge of shoulder near suture. Spiral sculpture incised by rather
coarse, oblique, irregular growth striations. Spiral sculpture very weak on
base of shell. Umbilical shield indented, bounded by a narrow crest that is
continuous with outer edge of columellar lip. Columellar callus wide, with
a concave face; edge of callus slightly bulging below middle into aperture
opening. Aperture about 0.37-0.45 times length of shell; subovate in shape.
Axis of aperture at about 28-330 to axis of shell; plane of aperture in lateral
profile 64-670 to shell axis. Outer lip thickened internally, nearly blunt-
edged, straight in lateral profile.
Measurements based on fifteen specimens selected to demonstrate
variation are as follow holotypee in parentheses): length, 3.16-4.03 mm
(3.66); width, 2.36-2.67 mm (2.36); aperture height, 1.36-1.71 mm (1.36).
TYPE LocALITY.-Butik Besungai, a small limestone hill 0.5 miles
southwest of Batu Gading, and about 4 miles northeast of Long Lama,
Baram Valley, Fourth Division, Sarawak (03052'N, 114025'E). HOLO-
TYPE: UF 36107; collected 16 June 1962 by D. J. Stevens. PARATYPES:
UF 36108 (100), UF 36109 (4 SEM mounts), S.P. Dance Coll. (46); SMF
255738/5; BMNH. (5); Sarawak Museum (3); same data as holotype.
DISTRIBUTION.-Known only from the type locality.
REMARKS.-This snail belongs to a complex of Bornean species that also
includes G. hosei Godwin-Austen, G. saulae (Benthem-Jutting), and G.
scalinella (Benthem-Jutting). The group also includes G. rufescens Moel-
lendorff from Luzon, Philippine Islands, and G. styloptyche Moellendorff
BULLETIN FLORIDA STATE MUSEUM
from Coron Island, Calamianes Archipelago. The group is characterized
by having spiral series of enlarged nodes and a deeply channelled suture
producing nearly scalariform whorls on the spire. G. hadra differs from all
other described species of this group by its large size, larger whorl count,
thick shell, and pink spire.
ETYMOLOGY.-The specific name, hadra, is from the classical Greek,
hadros, meaning well developed, stout, and alludes to the characteristically
thick shell.
Georissa hosei GODWIN-AUSTEN
Georissa hosei Godwin-Austen 1889; Proc. Zool. Soc. London: 353; pl. 39,
Fig. 11.-Smith 1893; Linn. Soc. Zool. 24:351; pl. 25, Fig. 27.
SHELL (Figs. 47-49a).-Small, 1.9-2.6 mm long. Adults elongate-conical
with an attenuated spire and a very deep suture. Shell about 0.69-0.81 times
as wide as high (smaller specimens proportionately wider). Whorls 2.8-3.3.
Protoconch bulbous, superficially smooth. Subsequent whorls flattened
above, and with strong spiral sculpture dominated by an enlarged suprape-
ripheral series of crispate nodes and usually by a similar, but smaller
subperipheral series; supraperipheral series bordered above by 5-8 fine
spiral lirae broken into short segments by axial threads (Fig. 49a); there are
4-6 similar spiral lirae between supraperipheral and subperipheral series;
base with 3-8 similar lirae that diminish near umbilical callus. Aperture
ovate, about 0.41-0.51 times height of shell, slightly higher than wide;
parietal wall lying at about 450 to shell axis; umbilical callus strongly
indented; outer lip inserting on base of penultimate whorl far in front of
parietal edge. Aperture very oblique in lateral profile, lying at about 59-640
to axis of shell.
Measurements for three population samples are given in Table 1. UF
35919 contains the largest specimens seen. Measurements and proportions
given above are based on these.
TABLE 1.-Georissa hosei: measurements of specimens from three population samples
selected to show variation. Two sets are selected from UF 35919. One consisting
of large specimens. The other of medium-sized specimens. Sets from UF 35920
and UF 35915 are the largest specimens in the samples.
UF 35919 UF 35919 UF 35920 UF35915
Number 8 6 5 5
Length 1.92-2.57 1.52-1.86 1.49-1.77 1.36-1.86
Width 1.55-1.86 1.27-1.46 1.24-1.40 1.27-1.55
Aper. h. 0.84-1.09 0.71-0.87 0.74-0.87 0.74-0.84
Whorls 2.8-3.1 2.4-2.7 2.4-2.7 2.3-2.7
Wid./len. 0.69-0.81 0.78-0.88 0.79-0.85 0.83-0.93
Wid./len. 0.41-0.51 0.44-0.50 0.47-0.56 0.45-0.55
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 117
OPERCULUM (Fig. 39).--Concentric; calcareous; broadly ovate. Outer
surface smooth with a few fine incremental striations. Inner surface with
peg originating from outer left cones; peg moderately stocky, projecting
baso-laterally at an angle of about 250 to axis, and inward at about 270 to
face of operculum; peg with a low knob on outer edge.
TYPE LOCALITY.-"Borneo." Lectotype by present designation: British
Museum (Natural History) 89.12.7.72; collected by C. Hose, 1889; one of
two syntypes.
DISTRBUTION.-Widely distributed in Sarawak, Borneo. Known from
the following localities SARAWAK: Fourth Div., Baram Valley, Bukit
Besungai, ca 4 mi NE Long Lama (03o52' N, 114025'E) (UF 35916, 35918);
Fourth Div., Tatau Valley, Bukit Sarang (02045'N, 113002'E) (UF 35914,
35915, 35921); Fourth Div., Gunong Subis, Niah area, ca 40 mi SW Miri
(03051'N, 113045'E) (UF 35917); Fourth Div., on Baram Riv., Batu Gading,
25 mi above Marudi (03053'N, 11430'E) (UF 35919); First Div., Gunong
Selabor, S of Serian, Semabang entrance to Lobang Batu Cave (0055'N,
11025'E) (UF 35920); Fourth Div., Batu Gading area, mouth of Labang
Tukeng Cave, nr Kejin Trib. of Baram River, ca 4 mi SE Long Lama
0350'N, 114028'E) (UF 35913).
Smith (1893:351) recorded the species from Jambusan (Sarawak), a
place we have not been able to locate. Presumably it comes from within the
range given above.
REMARKS.-This species is variable in the length of the spire and in the
intensity of the sculpture. Some population samples, consisting of less
mature shells, have a proportionately shorter spire, and thus appear pro-
portionately stocky. Occasionally the sculpture is less pronounced, but in
all specimens examined the supraperipheral series of crispate nodes is
present.
Four other Bornean species have similar sculpture: Georissa scalinella
Benthem-Jutting, G. saulae Benthem-Jutting, G. kobelti Gredler, and G.
hadra Thompson and Dance. They differ from G. hosei in color, size, and
details of sculpture, as discussed under those species.
Georissa kobelti GREDLER
Georissa kobelti Gredler 1902; Nachr. Deut. Malak. Gesellschaft 34:61.-
Zilch 1973; Arch. Molluskenkunde 103:265, Taf. 12, Fig. 11.
SHELL (Figs. 50-52).- Moderately small, about 2-3 mm long. Elongate-
conical, about 0.66-0.79 times as wide as high. Color uniform light yellow-
brown. Spire straight-sided with a deeply impressed suture. Whorls 2.7-3.9,
uniformly rounded, regularly increasing in-size. Protoconch ovate, bul-
bous, minutely granulose. First postembryonic whorl with about 10-14
fine, wavy spiral threads that are crossed by finer, irregularly spaced
growth striations. On lower whorls some threads become enlarged and
BULLETIN FLORIDA STATE MUSEUM
rugose, bearing numerous oblique scale-like nodes (Fig. 52); enlarged
spiral threads usually interspersed with smaller threads; penultimate whorl
with about 8-12 enlarged threads above insertion of lip; occasional speci-
mens with a conspicuously enlarged supraperipheral row of nodes (Fig.
52). Last whorl in large specimens with reduced sculpture. Spiral sculpture
rather uniform over surface of whorls except on base around umbilical area
where spiral cords become obsolete. Umbilical callus indented, without
conspicuous bounding ridge. Aperture subovate, about 0.37-0.48 times
height of shell; parietal-columellar margin straight-edged or weakly con-
vex. Axis of aperture at 24-360 to axis of shell; plane of aperture at 30-410 to
axis of shell. Peristome uniformly rounded to columella; not thickened
internally; straight-edged in lateral profile.
Measurements in mm of five specimens selected to show variation are
as follows.
length width aperture whorls
SMF 215893/4 3.00 1.98 1.12 3.8
SMF 215893/4 2.33 1.74 0.99 3.2
UF 36171 2.39 1.71 0.96 3.1
UF 36171 2.11 1.61 0.90 2.9
UF 36171 2.08 1.64 0.90 2.8
TYPE LOCALITY.-Niah, Baram (Sarawak, Borneo). LECTOTYPE:
Senckenbergische Naturforschende Gesellschaft 215893a (Zilch 1973:265).
DISTRIBUTION.-Known only from a single locality in the Niah Hills in
northern Borneo. We have examined the following specimens: SARAWAK:
Fourth Div., Beluru Area, SW of Marudi, eastern part of Bukit Vrong
(04000'N, 114012'E) (UF 36170,4 spec.). In addition we have examined two
syntypes of G. kobelti (SMF 215893/4).
REMARKs.-This species is closely related to G. hosei Godwin-Austen. It
differs from G. hosei, as well as others in the hosei group by its scale-like
nodes arranged on spiral threads. In addition, none of the spiral threads are
conspicuously enlarged on the lower whorls.
Georissa saulae (BENTHEM-JUTTING)
Hydrocena saulae Benthem-Jutting 1966; J. Conch. 26:40-41; fig. 2.
TYPE LOCALITY.-Laying Cave, Crocker Range, Keningau, Sabah. Holo-
type in the Zoological Museum, University of Amsterdam.
This species is known only from the type locality. It and the closely
related G. scalinella (Benthem-Jutting) occur far to the east of other Bor-
nean species of thehosei group. G. saulae and G. scalinella are very closely
related. G. saulae differs from G. scalinella by having a more elevated spire
and less scalariform whorls. The two may be only subspecifically distinct.
Additional material from the Crocker Range may clarify their relationship.
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 119
A specimen of G. saulae is illustrated (Fig. 53, 54) so that its sculptural
characteristics can be compared. The surface is covered with closely
spaced, oblique, lamelliform axial threads that are more crowded and
elevated than in related species, except G. scalinella. Superimposed on the
axial threads are a few spiral lirations. The peripheral liration has enlarged
flattened nodes and usually there is a subperipheral series also. Measure-
ments of five specimens, UF 25057 (2), UF 36110 (1), and UF 36111 (2), are
as follow: length of shell, 1.71-1.88mm; width, 1.27-1.38mm; aperture
height, 0.67-0.35; 3.3-3.5 whorls; width/length, 0.72-0.78; aperture height/
shell length, 0.38-0.40.
Shell lengths are comparable to measurements given by Benthem-
Jutting; the widths are not. Benthem-Jutting measured the greatest width
of specimens whereas our measurements of width are made perpendicular
to the axis of the shell.
Georissa scalinella (BENTHEM-JUTTING)
Hydrocena scalinella Benthem-Jutting 1966; J. Conch. 26:39-40, Fig. 1.
TYPE LOCALITY.-Lahad Datu Caves on Teck Guan Estate, Sabah.
Known only from the type locality. Holotype in the Zoological Museum,
University of Amsterdam.
Georissa niahensis GODWIN-AUSTEN
Georissa niahensis Godwin-Austen 1889; Proc. Zool. Soc. London 1889:
373-4; pl. 39, Fig.8.
TYPE LOCALITY.-Niah Hills (Sarawak), Borneo. Lectotype by present
designation: British Museum (Natural History) 89.12.7.69; collected by A.
Everett, one of two syntypes. The lectotype is the specimen figured by
Godwin-Austen.
REMARKS.-Our sample contains no specimens identifiable with this
species. We have examined the lectotype and the smaller syntype. Mea-
surements of the lectotype are: length, 3.22 mm; width, 2.11 mm; aperture
height, 1.36 mm; 4.2 whorls; width/length, 0.66; aperture/length, 0.42 (See
Table 2 to compare with G. williamsi). The smaller syntype is immature.
Godwin-Austen gives slightly larger measurements for the lectotype. He
also states that the syntypes were ruddy ochre in color. Currently they are
faded nearly white.
G. niahensis is similar in sculpture to G. williamsi but is much larger. G.
niahensis also shows similarities to the hosei group in the depth of the suture
and the relatively rapidly expanding whorls, but it lacks the node-like
sculpture found among species of that group. Additional collections from
the Niah Hills are needed to clarify the status of this species.
BULLETIN FLORIDA STATE MUSEUM
TABLE 2.-Georissa williamsi Godwin-Austen and G. xesta, new species. Measurements in
mm of large specimens selected to show maximum variation. Smaller speci-
mens of G. williamsi from UF 35908 completely encompass the variation
tabulated for UF 35912 and UF 35911. Holotype of G. xesta in parentheses.
williamsi xesta
UF 35908 UF 35912 UF 35911 UF 35968-9
Number 9 8 10 7
Length 1.83-2.29 1.30-1.73 1.46-1.73 1.58-2.05 (1.83)
Width 1.40-1.55 1.02-1.24 1.09-1.20 1.15-1.33 (1.21)
Aper. h. 0.71-0.81 0.59-0.71 0.59-0.71 0.65-0.71 (0.68)
Whorls 3.4-4.0 3.3-3.8 3.0-3.4 3.2-3.8 (3.7)
Wid./len. 0.65-0.76 0.65-0.79 0.68-0.77 0.65-0.73 (0.66)
Aper./len. 0.33-0.41 0.35-0.43 0.39-0.45 0.35-0.41 (0.37)
Ceorissa everetti SMITH
Georissa everetti Smith 1895; Proc. Zool. Soc. London 1895; 125; pl. 4, Fig.
15.
SHELL (Figs. 55-57).-Small, about 1.95-2.20 mm long. Ovate-conical in
shape, about 0.69-0.77 times as wide as high. Spire convex in outline. Light
yellow; occasionally with an orange-tinted spire. Whorls 3.3-3.6. Proto-
conch bulbous, globose, superficially smooth. Subsequent whorls uni-
formly rounded with a deeply impressed suture; sculptured with oblique
incremental thread-riblets that are decussated by oblique spiral threads
(Fig. 57). Incremental riblets most distinct on spire, becoming obsolete on
last half of body whorl. Spiral threads usually strongest on body whorl.
Continuous and uniform from suture to base. Umbilical area indented and
bordered by a columellar lip. Aperture subovate in shape, about 0.41-0.48
times length of shell. Axis of aperture at about 53-590 to shell axis. Plane of
aperture in lateral profile lying about 33-360 to shell axis. Columellar callus
rounded in front, concave in outline along aperture opening. Basal lip
conspicuously thickened internally. Outer lip slightly sinuous in lateral
profile.
Measurements based on ten specimens are given (UF 36157). No popu-
lation differences are apparent between the two samples examined.
Length, 1.95-2.20 mm; width, 1.46-1.55 mm; aperture height, 0.87-1.00 mm.
TYPE LOCALITY.- "Rumbang, W. Sarawak." We have not been able to
find this place on any maps. Presumably it is in the vicinity of Kuching.
Holotype: British Museum (Natural History) 93.6.7.69.
DISTRIBUTION.-Widely distributed in Sarawak, Borneo, but apparently
sparse in occurrence. We have examined specimens from the following
localities. SARAWAK: Fourth Div., Niah Area, Gunong Subis, a limestone
massif about 40 mi SW of Miri (03051'N, 113045'E) (UF 36156); First Div.,
Wah Mountain, immediately S of Bra-Ang Wah Village, 23 mi S Kuching
(01013'N, 11016'E) (UF 36157).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 121
REMARKS.-This species was founded on a single specimen, and has not
been reported since its original discovery. The holotype was compared
with our specimens, and they agree well in all important characteristics. G.
everetti is unique within the genus because of the decussate sculpture
produced by oblique spiral threads and the oblique axial riblets. The
thickened basal lip is also unusual.
Georissa gomantonensis SMITH
Georissa gomantonensis Smith 1893; J. Linn. Soc. Zool. 24:351; pl. 25, fig.
25.
Georissa flavescens Smith 1895; Proc. Zool. Soc. London 1895:126; pl. 4, fig.
17.
SHELL (Figs. 58-60).-Small, about 1.88-2.14 mm long; broadly ovate-
conical, about 0.76-0.85 times as wide as high; sides of spire weakly convex
due to slightly increased rate of descent of last whorl. Shell citron yellow in
color. Whorls 3.0-3.2. Protoconch globose and superficially smooth. Sub-
sequent whorls inflated and evenly rounded with a deeply impressed
suture, and with a narrow, shallow channel on shoulder bordering suture.
Channel bordered laterally by single row of oblique, compressed, small
nodes (Fig. 58). Whorls sculptured below nodes with spiral threads parallel
with suture and about as wide as their interspaces. Spiral threads strongest
above periphery, weakly developed or obsolete on base; 17-21 spiral
threads on penultimate whorl between upper suture and insertion of lip.
Umbilical callus weakly indented, bounded laterally by a narrow crest.
Aperture subovate, height about 0.44-0.48 times height of shell; parietal-
columellar margin weakly concave in outline. Axis of aperture at 32-360 to
shell axis; plane of aperture in lateral profile 28-340 to shell axis. Outer lip
very slightly sinuous in lateral profile, thin. Basal lip slightly thickened
within.
Measurement of eight specimens (UF 36172) selected to show maxi-
mum variation: length of shell, 1.86-2.14 mm; width of shell, 1.46-1.74 mm;
aperture height, 0.90-0.96 mm.
TYPE LOCALITIES.-Georissa gomantonensis: Gomanton, N. Borneo
(=Sabah). Holotype: British Museum (Natural History) 92.7.20.39. Georissa
flavescens: Gomanton, N.E. Borneo (=Sabah). Lectotype by present
designation: British Museum (Natural History) 93.6.8.11, one of 3 syntypes.
DISTnIBUTION.-Known from scattered localities in eastern Sabah, Bor-
neo. We have examined specimens from the following places. SABAH:
Kinabatangan Valley, limestone scarp 3 mi NNW Laab, 7 mi E Lamag
(0532'N, 117059'E) (UF 36172, 36173 SEM); Kinabatangan Valley, north-
ern flank Bukit Gomantong, 22 mi S of Sandakan (0503(YN, 11810'E) (UF
36174); Kinabatangan Valley, NE end Bukit Kuntos nr Mile 6 on track from
Suanlamba to Gomantong Caves, 22 mi S of Sandakan (05031'N, 118"17'E)
BULLETIN FLORIDA STATE MUSEUM
(UF 36176); Kinabatangan Valley, Batu Tumanggong Besar, 1 mi down-
stream from Sukau (05034'N, 118020'E) (UF 36176); Tabin Valley, trib.
lower Segama Riv., limestone ridge W of lower Tabin River (05024'N,
118048'E) (UF 36177); head of Cowie Harbour, limestone outcrop ca 15 mi
NNW Kalabakan (0428'N, 11728'E) (UF 36178).
REMARKS.-This species is similar in size, shape, color, and sculpture to
G. everetti Smith. It differs from the latter species by its broader shape, its
shoulder channel bordering the suture and its sculpture. G. gomantonensis
is 0.76-0.85 times as wide as high; the subsutural channel is bordered by a
row of small, obliquely compressed nodes and the spiral threads run
parallel to the suture. G. everetti is 0.69-0.77 times as high, lacks a channel
on the shoulder of the whorls, lacks a spiral row of compressed nodes, the
spiral threads are oblique to the suture and are crossed by oblique axial
threads. Other Bornean species of similar shape are G. borneensis Godwin-
Austen, G. bangueyensis Smith, and G. pyrrhoderma Thompson and
Dance. These species differ so much in color and sculpture, however, that
no close relationship to G. everetti and G. gomatonensis should be
assumed.
Georissa borneensis SMITH
Figures 61-62
Georissa borneensis Smith 1895; Proc. Zool. Soc. London 1895:126; pl. 4,
Fig. 18.
TYPE LocALITY.-Gomanton, N.E. Borneo (=Sabah). Lectotype by
present designation: British Museum (Natural History) 94.7.20.61, one of
six syntypes; collected by A. Everett. Known only from the type locality.
REMARKS.-The original description incorrectly defines and illustrates
this species. None of the six syntypes has the dimensions given by Smith,
and the original figure shows an ovate-conical shell in contrast to the
turbinate-conical shell of the species. Smith stated that the outside of the
shell is reddish and the interior is bright red. The syntypes now are faded
nearly white. In addition to the color of the aperture the species is charac-
terized as follows. It has strong growth striations that become almost
rib-like in some instances; spiral sculpture is absent. The peristome is
internally thickened. The shell is turbinate-conical in shape with the
periphery lying well below the middle of the body whorl, and the suture is
channelled with a narrow shoulder (Fig. 62). Measurements in mm of the
type specimens are as follows (lectotype in parenthesis): length, 2.20-2.42
(2.39); width, 1.78-1.83 (1.77); aperture height, 0.93-0.96 (0.96); whorls,
3.3-3.5 (3.4); width/length, 0.73-0.80 (0.74); aperture/length, 0.39-0.40
(0.40).
Georissa borneensis differs from other Bornean species by its thickened
peristome, sculpture, shape, and suture. We associate it with G. pyrrho-
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 123
derma because of similarities in color. Otherwise it is intermediate in aspect
between G. pyrrhoderma and the G. williamsi species group.
Georissa pyrrhoderma, NEW SPECIES
SHELL (Figs. 63-65).-Small, adults about 1.9-2.1 mm long. Orange-red
in color; interior hyaline. Broadly conical, about 0.79-0.86 times as high as
wide. Spire very slightly convex in outline. Whorls of spire relatively small
and with a deeply impressed suture. Whorls 2.8-3.1. Protoconch nearly
circular and bulbous. Subsequent whorls strongly inflated and uniformly
rounded; sculptured with relatively weak spiral threads and numerous
close growth striations; about 18-21 spiral threads on penultimate whorl
above insertion of lip; spiral threads usually absent on base of shell. Occa-
sional specimens with a supraperipheral spiral row of small compressed
nodes (Figs. 64,65), which when present are predominant on the middle
whorl. Umbilical callus slightly concave, very wide compared with other
Bornean species; not bounded laterally by a narrow crest. Aperture sub-
ovate in shape; relatively large, height about 0.48-0.54 times length of shell.
Parietal-columellar margin straight. Axis of aperture at 28-340 to shell axis.
Plane of aperture at 26-340 to shell axis. Peristome uniformly thin, straight in
lateral profile.
Measurements based on sixteen specimens selected to show maximum
variation are given holotypee in parentheses). Length of shell, 1.92-2.23 mm
(2.23); width of shell,. 1.64-1.80 mm (1.80); height of aperture, 0.99-1.12
mm (1.12).
Most of the shells comprising the type series are weathered and
encrusted and few are unblemished. The holotype is a fresh specimen but
has a small puncture in the penultimate whorl behind the insertion of the
lip.
TYPE LOCALITY.-Borneo, Sarawak, First Division, western side of
Gunong Selabor, Semabang entrance to Lobang Batu Cave. Gunong Sela-
bor is an isolated limestone mountain south of Serian, in southwestern
Sarawak (00055' N, 110025'E). HOLOTYPE: UF 36183 collected in 1962 by
G.E. Wilford. PARATYPES: UF 36184 (100), UF 36185 (4 SEM specimens)
SMF 255739/5; S.P. Dance Coll. (53); BMNH (5); Sarawak Museum (3).
DIsTRIuTIoN.-Known only from the type locality.
REMARKS.-G. pyrrhoderma belongs to a group of Bornean species that
also includes G. borneensis Smith. The group consists of species having
broadly conical, reddish shells. G. pyrrhoderma is distinguished from G.
borneensis by its sculpture of conspicuous spiral threads. Other aspects of
the shell cannot be adequately compared because of insufficient material
of G. borneensis. The two species are geographically remote from each
other on Borneo and on Banggi Island.
ETYMOLOGY.-The name pyrrhoderma is from the Greek, pyrrhos,
BULLETIN FLORIDA STATE MUSEUM
meaning red or flame-colored, and derma, meaning skin or hide, in allusion
to the color of the shell.
Georissa williamsi GODWIN-AUSTEN
Georissa williamsi Godwin-Austen 1889; Proc. Zool. Soc. London 1889:
353; pl. 39, Fig. 10.
Georissa hungerfordi Godwin-Austen 1889; Proc. Zool. Soc. London 1889:
354; pl. 39, Fig. 9.
Georissa lowi Smith 1893; J. Linn. Soc. Zool. 24:351.
Georissa javana Moellendorff 1897; Nachr. Blatt. Deut. Malak. Gesell.
29:97.-Paravincini 1935; Arch. Moll. 67:179.-Laidlaw 1935; Bull. Raffles
Mus. 10:99, Figs. 2-6, Zilch 1973; Arch. Moll. 103:265; pl. 12, fig. 8.
Hydrocena (Georissa) javana (Moellendorff) Benthem-Jutting 1940; Treu-
bia, 17:332.-Benthem-Jutting 1948; Treubia 19:544-545, Figs. 5 (shell), 6
(operculum):1959; Beaufortia:59.
Georissa javana intermediaMoellendorff 1897; Nachr. Blatt. Deut. Malak.
Gesell. 29:97.-Zilch 1973; Arch. Moll. 103:265; pl. 12, Fig. 9.
SHELL (Figs. 66-68).-Minute-sized, reaching a length of 2.3 mm.
Yellowish-brown to rust colored. Elongate-conical, about 0.65-0.76 times
as high as wide in large specimens; smaller specimens relatively broader
(Table 2). Spire slightly convex in outline. Columella narrow; internal
partition extending to 1.0 whorls behind aperture. Whorls 3.5-4.0 in large
specimens. Protoconch bulbous, superficially smooth. Subsequent whorls
sculptured with rather heavy spiral cords that parallel suture. Shells over 2
mm long with 7-8 cords on penultimate whorl and 12-16 cords on last
whorl. Cords strongest on shoulder of whorl, weakest on base. Interspaces
with occasional weak growth striations. Sculpture variable; some speci-
mens nearly smooth on last half of body whorl. Umbilical callus small;
strongly indented. Aperture broadly ovate, 0.33-0.41 times length of shell.
Plane of aperture in lateral profile at about 30-340 to axis of shell. Axis of
aperture at 27-450 to axis of shell. Columellar callus wide; rounded. Outer
lip slightly sinuous in lateral profile; insertion not noticeably advanced
forward as in G. hosei group.
Measurements of large specimens from three population samples are in
Table 2.
TYPE LOCALITIES.-Georissa williamsi: Borneo. Holotype: British Mu-
seum (Natural History) 89.12.7.71; collected by C. Hose. Georissa hunger-
fordi: Borneo. Lectotype by present designation: British Museum (Natural
History) 91.3.17.864; collected by Sir Hugh Low; one of two syntypes.
Georissa lowi: Borneo. Holotype same as G. hungerfordi because Smith
proposed G. lowi as a substitute name. He assumed that G. hungerfordi
G.-A. 1889 was preoccupied by G. hungerfordiana Moellendorff 1885
which is not so because the two are not primary homonyms. Georissa
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 125
javana: Gunung Cede, Java, 7500 ft alt. Lectotype: Senckenbergische
Naturforschende Gesellschaft SMF 227955 (Zilch 1973:265; Taf. 12, fig. 8).
Georissa javana intermedia: Gunung Cede, Java, 5000 ft alt. Lectotype:
Senckenbergische Naturforschende Gesellschaft SMF 227966 (Zilch
1973:265; Taf. 12, Fig. 9).
DISTRIBUTION.-Widely distributed on Java (Benthem-Jutting 1948:
544), Sumatra (Benthem-Jutting 1959:59), Christmas Island (Laidlaw 1935)
and Borneo. Known from the following localities in Borneo. SARAWAK:
First Division, SE end of Bau Limestone outcrop (01003'N, 11027'E) (UF
35908); Fourth Division, Tatau Valley, Bukit Sarang (02"45'N, 113002'E)
(UF 35906, 35907, 35910); 25 mi S Kuching, 1 mi NE Pankalan Ampat, hill
0.5 mi E Temerang (01o11'N, 11012'E) (UF 35905); Beluru Area, SW of
Marudi, Bukit Vrong (0400'N, 11412'E) (UF 35911, 35912); Beluru Area,
SW of Marudi, nr. Bukit Kudi (0400(YN, 114012'E) (UF 35909).
REMARKS.-Georissa williamsi belongs to a species group that also
includes G. xesta Thompson and Dance, G. bangueyensis Smith, G. niahen-
sis Godwin-Austen, and G. similis Smith. Similar species are recorded from
nearby islands. The group is characterized by its elongate-conical shape,
small size, and spiral cords. G. niahensis is known only from two speci-
mens, and it is difficult to discuss its characteristics other than to say it is a
much larger, stockier species, reaching a length of 3.6 mm. G. similis is
similar in size and appearance to G. williamsi, except that it has radial ribs
in addition to spiral cords, and is more slender. Other differences are
discussed under G. similis.
G. williamsi occurs in numerous isolated colonies, and some are distinc-
tive in the size attained by adults and in the intensity of the sculpture,
particularly on the later whorls. We can find no differences that will
separate any of the named forms listed in the synonymy. We suspect that
species named from other places will also be proved synonymous with
williamsi when they become better known.
Georissa xesta, NEW SPECIES
SHELL (Figs. 69-70).-Small, adults about 1.6-2.0 mm long; elongate-
conical, about 0.65-0.73 times as wide as high. Spire straight-sided. Shell
thin. Periostracum yellowish to rust-colored. Whorls 3.2-3.8 in adult shells.
Protoconch bulbous, superficially smooth. Subsequent whorls strongly
arched with a deeply impressed suture; sculptured with very low, weak,
spiral threads, which may be absent in some specimens; spiral threads,
when present, equally developed as far as umbilical callus, crossed by
irregularly spaced growth striations. Umbilical region conspicuously
indented. Columellar callus thick and rounded. Aperture subcircular,
weakly concave across parietal margin; aperture about 0.35-0.41 times
length of shell. Axis of aperture at about 27-340 to shell axis, plane of
BULLETIN FLORIDA STATE MUSEUM
aperture at about 16-220 to shell axis. Outer lip weakly sinuous in lateral
profile.
Measurements of the holotype and six paratypes are given in Table 2.
TYPE LOCALITY.-A small limestone ridge quarried for rock 5 mi W
Kudat, Sabah, Borneo (06057'N, 1160 48'E). HOLOTYPE: UF 35968; col-
lected in 1966 by G.E. Wilford. PARATYPES: UF 35969 (63), UF 35970 (5),
S.P. Dance Coll. (10), SMF 255740/6, BMNH (5) same data as the holotype.
DISTRInUTIN.-Known only from the type locality.
REMARKS.-This subspecies differs from G. williamsi in the intensity of
its spiral sculpture. Upon superficial examination G. xesta appears to be
smooth, but spiral sculpture is apparent when viewed with strong reflected
light. Its relationship to G. bangueyensis Smith is discussed under that
species.
ETYMOLOGY.-The specific epithet xesta is derived from the Greek
xestos, meaning smooth or shaven, alluding to its weak sculpture.
Georissa bangueyensis SMITH
Georissa bangueyensis Smith 1895; Proc. Zool. Soc. London 1895:126; pl. 4,
fig. 16.
TYPE LOCALITY.-Banguey Island (=Banggi Island), Sabah, Borneo.
Lectotype by present designation: British Museum (Natural History)
93.6.7.9, one of six cotypes; collected by A. Everett. Known only from the
type locality.
REMARKS.-The status of this species is uncertain. It is similar to G.
xesta, but it is stockier and has a proportionally larger aperture. Measure-
ments for the type series are as follow (lectotype in parenthesis): length,
1.62-1.81 mm (1.70); width, 1.21-1.30 mm (1.21); aperture height, 0.72-0.75
mm (0.75); whorls 3.2-3.5 (3.4); width/length, 0.69-0.75 (0.71); aperture/
length, 0.42-0.44 (0.44) (see Table 2 for comparison with G. xesta). The
sculpture consists of barely evident growth striations and a few weak spiral
striations. Smith (1895:126) stated that the species is reddish in color. The
type specimens are now faded to nearly white. The strong umbilical callus
mentioned by Smith does not differ significantly from that of related
species. Additional specimens from Banggi Island may show that G. ban-
gueyensis and G. xesta are conspecific, but we do not feel warranted in
making this assumption at present.
Georissa similis SMITH
Georissa similis Smith 1893; J. Linnean Soc. Zool. 24:351; pl. 25, Fig. 26.
Hydrocena (Georissa) gomantonensis (Smith), Solem 1964; Sabah Soc. J.
2:8; Fig. I, 1.
SHELL (Figs. 73-75).-Medium-sized, about 2.5-2.9 mm long; elongate-
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 127
conical, about 0.62-0.70 times as high as wide; spire straight-sided, or very
weakly convex in profile. Color light brown, glossy. Columella large,
tubular; internal partition extending to 1.5 whorls behind aperture (Fig.
38). Shell thin, translucent. Large specimens with about 3.0-3.5 whorls (3.3
in holotype). Protoconch bulbous, ovate in shape from above (Fig. 31).
Subsequent whorls strongly arched with a very deeply impressed suture;
sculptured with fine spiral threads and stronger, weakly sigmoid, retrac-
tively slanting riblets that are almost as wide as their interspaces. Riblets
continuous but diminished onto the base. Spiral threads and riblets give a
matted appearance to surface of shell. Umbilical area indented; bounded
laterally by a thin crest continuing from outer edge of columellar lip. Callus
wide, rounded, bulging below middle; weakly granular. Aperture semi-
lunar in shape, about 0.35-0.41 times height of shell; axis of aperture at
about 28-310 to axis of shell; plane of aperture in lateral profile at about
22-30 to shell axis.
Measurements in mm of eleven large specimens (UF 36094) selected to
show maximum variation are as follow: length 2.51-2.91; width, 1.74-1.86;
aperture height, 1.00-1.12.
TYPE LOCALITY.-Georissa similis: Gomanton Hill, North Borneo
(=Sabah). Lectotype by present designation: British Museum (Natural
History) 92.7.23.51, one of two syntypes. Georissa gomantonensis: Goman-
ton, North Borneo (=Sabah). Holotype: British Museum (Natural History)
92.7.20.39.
DISTRIBUTION.-Known only from extreme northeastern Borneo from
limestone outcrops in the Kinabatangan and Segama Valley and the
Gomantong Hills. We have examined specimens from the following locali-
ties. SABAH: Kinabatangan Valley, Bukit Kolop, 4 mi WSW Latangan
(05020'N, 117032'E) (UF 36099); Kinabatangan Valley, Batu Tumanggong
Besar, 1 mi downstream from Sukau (05034'N, 11820'E) (UF 36096);
limestone outcrop along Kinabatangan River 1 mi downstream from Laab
(05027'N, 11759'E) (UF 36094, 36100); Kinabatangan Valley, limestone
scarp, 3 mi NNW Laab, 7 mi. E. Lamag (05032'N, 11757'E) (UF 36168);
limestone hill nr. Mile 2, on track from Suanlamba to Gomantong Caves,
18 mi S Sandakan (05036'N, 118017'E) (UF 36097); northern flank Bukit
Gomantong, 22 mi S Sandakan (05030'N, 11810'E) (UF 36169); limestone
ridge in Tabin Valley, a tributary of lower Segama River (05022'N,
118054'E) (UF 36098); ridge W of Tabin River, tributary to lower Segama
River (05024'N, 118048'E) (UF 36095).
REMARKS.-Georissa similis is similar in size and appearance to G.
williamsi and G. xesta. It differs from G. williamsi and G. xesta by its more
slender shape, fewer whorls, and its axial riblets. The obesity and number
of whorls are variable among the three taxa and overlap occurs between
them in these characters. G. similis is readily and consistently distinguisha-
ble from the others by its sculpture.
BULLETIN FLORIDA STATE MUSEUM
The type specimens of G. similis are immature and do not show the axial
sculpture as well as do larger, more developed specimens.
HELICINIDAE
No helicinids are present in the collection. The following species are
recorded from Borneo.
Geophorus agglutinans eos WAGNER
Geophorus agglutinans eos Wagner 1911; Martini-Chemnitz Syst. Conchyl.
Cab., n.f., abt. 18 (2):143; Taf. 31, figs. 1-4.
TYPE LOCALITY.-Gunung Sekerat, on the east, Kutei Prov., Kalimantan,
Indonesian Borneo. Known only from the type locality.
Sulfurina martensi (ISSEL)
Helicina martensi Issell 1874; Ann. Mus. Civ. Stor. Nat. Genova 6:444-445;
pl. 2, figs. 23-25.
Helicina citrina var. minor Martens 1873; Malak. Blatt. 20:161.
Sulfurina martensi (Issel) Wagner 1905; Helicinenstudien, in: Denk. Akad.
Wien 77:375; taf. 2, figs. 8 a-c.-Wagner 1911; Martini-Chemnitz Syst.
Conch. Cab., n.f., abt. 18 (2):24; Taf. 3, figs. 22-24.
TYPE LOCALITY.-Borneo.
DIsTIBUTION.-Labuan Is., Palawan Is., Borneo (Wagner 1911:24).
Aphanoconia usukanensis(GODWIN-AUSTEN)
Helicina usukanensis Godwin-Austen 1889; Proc. Zool. Soc. London
1889:352; pl. 39, Fig. 7.-Smith 1895; Proc. Zool. Soc. London 1895:125.
TYPE LOCALITY.--Usukan Island, Borneo.
DISTRIBUTION.-Known from Mengalun Island, Usukan Island, Banggi
Island, Mantangak Island, Palawan, Sibutu Island, Bilatan Island, Sulu
Islands (Smith 1895:125).
Aphanoconia borneensis(MARTENS)
Helicina borneensis Martens 1864; Monatsb. Berlin Akad.: 120.-Pfeiffer
1865; Monogr. Pneum. Viv. 3:238.-Martens 1868; Ostasien Zool. 2:171.-
Issel 1874; An. Mus. Civ. Stor. Nat. Genova 6:81- Sowerby 1866; Thes.
Conch. 3:286; pl. 270, Figs. 194, 196, 211, 212.
Aphanoconia borneensis, Wagner 1905; Helicinenstudien, in: Denk. Akad.
Wien 77:47; Taf. 5; Figs. 19a-c.-Wagner 1911; Martini-Chemnitz. Syst.
Conch. Cab., n.f., abt. 18(2):175; pl.35, Figs. 1-4.
TYPE LOCALITY.-Singdawang, Kalimantan, Indonesian Borneo. Known
only from the type locality.
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 129
Aphanoconia trichroa WAGNER
Aphanoconia trichroa Wagner 1905; Helicinenstudien, in: Denk. Akad.
Wien 77: 44; Taf. 5, figs. 15-16.--Wagner 1911; Martini-Chemnitz Syst.
Conch. Cab., n.f., abt. 18(2):173; Taf. 35, figs. 10-13.
TYPE LOCALITY.-Banguei (=Banggi Island), Sabah, Borneo.
DISTmBUTION.- Known only from Banggi Island and Sibatu Island off
the northeast coast of Borneo (Wagner 1911:173).
LITERATURE CITED
Abbott, T. 1958. The gastropod genus Assiminea in the Philippines. Proc. Acad. Nat. Sci.
Philadelphia 110:213-278.
Baker, H.B. 1935. Review of the anatomy of Pupillidae and related groups. In H.A. Pilsbry,
Manual of Conchology (Ser. 2) 28:191-226.
Benthem-Jutting, W.S.S., van. 1948. Systematic studies on the non-marine Mollusca of the
Indo-Australian Archipelago. Pt.2. Javanese operculate land snails of the families
Hydrocenidae, Helicinidae, Cyclophoridae, Pupinidae and Cochlostomatidae. Treubia,
91:540-603.
S1949. The Malayan species of Boysidia, Paraboysidia, Hypselostoma and
Gyliotrachela (Gastropoda, Pulmonata, Vertiginidae), with a catalogue of all the species
hitherto described. Bull. Raffles Mus. 21:5-47.
1952. Systematic studies on the non-marine Mollusca of the Indo-Australian
Archipelago, Pt. 3. Javanese pulmonate land snails of the families Ellobiidae to Limacidae.
Treubia 21:291-435.
S1959. Catalogue of the non-marine Mollusca of Sumatra and of its Satellite
Islands. Beaufortia 83:41-191.
1 960. Some notes on land and freshwater Mollusca of Malaya. Basteria
24:10-20.
S1961. Additional new species and new localities of the Family Vertiginidae
and the Genera Oophana and Opisthostoma from Malaya. Bull. Raffles Mus. 26:34-48.
S1962. Coquilles terrestres nouvelles de quelques collins calcaires du Cam-
bodje et du Sud Vietnam,. J. Conchyl. 102:3-15.
Berry, A.G. 1961. The habitats of some minute cyclophorids, hydrocenids and vertiginids on a
Malayan limestone hill. Bull. Singapore Nat. Mus. 30:101-105.
S1965: Reproduction and breeding fluctuations in Hydrocena monterosati-
ana, a Malayan limestone archeogastropod. Proc. Zool. Soc. London 144:219-227.
Dance, S.P. 1970. Non-Marine Molluscs of Borneo, I. Streptaxacea: Steptaxidae. J. Conch.
27:149-162.
Gredler, P.V. 1902. Zur Conchylien-fauna von Borneo and Celebes. Deut. Malak. Gesell.
34:53-63.
Haas, F. 1937. Neue and Kritische Pupilliden. Arch. fur. Moll. 69:1-12.
Habe, T. 1949. The study of land snails ( in Japanese). Kosei-sha, Japan.
Issel, A. 1874. Molluschi Borneensi, illustrazione delle specie terrestri e d'acqua dolce, raccolte
nell' Isola Borneo, dai Signori G. Doria e O. Beccari. Ann. Mus. Civico di Storia Nat.
Genova 6:366-478.
Laidlaw, F.F. 1935. Notes on a collection of terrestrial Mollusca from Christmas Island. Bull.
Raffles. Mus. 10:95-100.
Loosjes, F.E. 1953. Monograph of the Indo-Australian Clausiliidae (Gastropoda, Pulmonata,
Clausiliidae, Phaedusinae). Beaufortia (31):1-226.
Martens, E., von. 1908. Beschreibung einiger in ostlichen Borneo von Dr. Martin Schmidt
BULLETIN FLORIDA STATE MUSEUM
gesammelten Land und Susswasser-Conchylien. Zool. Mus. Mitteilungen Berlin 4:249-292;
pls. 5-6.
Pilsbry, H.A. 1916-1918. Manual of Conchology, (Ser. 2) 24:i-xii, 1-380.
S1918-1920. Manual of Conchology, (Ser. 2) 25:i-ix, 1-401.
S1927-1935. Manual of Conchology, (Ser. 2) 28:i-xii, 1-226.
Rensch, 1934. Zur Landschneckenfauna von Timor. Sitzungsb. Gesellschaft Naturforschende
Fr. Berlin. 1934:311-336.
S1931. Zur Kenntnis der Molluskenfauna Timors. Zoologische Jahrbucher,
Abt. Syst. Okologie Geogr. 30:429-455.
Saul, M. 1966. Shell collecting in the limestone caves of Borneo. New York Shell Club Notes
(125):5-9.
Smith, E.A. 1893. Descriptions of new species of land-shells from Borneo. Proc. Linnean Soc.
Zool. London. 24:341-352.
S1895. On a collection of land-shells from Sarawak, British North Borneo,
Palawan, and other neighboring Islands. Proc. Zool. Soc. London 1895:97-127.
S1898. A list of the land-shells of the island of Lombok with descriptions of
new species. Proc. Malac. Soc. London 3:26-32.
Solem, A. 1964. A collection of non-marine mollusks from Sabah. Sabah Soc. J. 2:1-40.
Thiele, J. 1931. Handbuch der Systematischen Weichtierkunde, I. i-vi, 1-778. Berlin.
Thompson, F.G. 1980. Proserpinoid land snails and their relationships within the Archeogas-
tropoda. Malacologia 20:1-33.
Tomlin, J.R., Le B. 1939. New Land Shells from the Malay Peninsula. J. Conch. 21:146-147.
Zilch, A. 1959. Gastropoda, Euthyneura. Handbuch der Palaozoologie 6(2):i-xii, 1-834.
S1973. Die Typen und Typoide des Natur-Museums Senckenberg, 52. Arch.
Moll. 103:263-272.
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 131
i-Q
SABAH .
0 50 I1O Ila 2 km
'2*
Figure 1. Map of Borneo showing stations from which material for this study was collected.
BULLETIN FLORIDA STATE MUSEUM
F1
Figures 2-6. Boysidia (Dasypupa) salpinx, n.sp., paratypes (UF 36281) Fig. 2 (X18), Fig. 3
(X22), Fig. 4 (X44), Fig. 5 (X44), Fig. 6 (X60).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
A_
7 8
Figures 7-8. Boysidia (Dasypupa) salpinx, n.sp. Holotype (UF 35944).
Figures 9-10. Boysidia (Dasypupa) paini, n.sp. Holotype (UF 35949).
134 BULLETIN FLORIDA STATE MUSEUM
11 12
Figures 11-12. Boysidia (Dasypupa) procera, n.sp. Holotype (UF 38022).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 135
0
0 0 0
Figures 13-18. Bornean clausiliids. Figs. 13-15.-Phaedusa borneensis (Pfr.) (UF 35928). Figs.
16-18.-Phaedusa filicostata stenotrema, n.ssp. Holotype (UF 36426).
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Figures 19-21. Bornean clausiliids. Fig. 19.-Phaedusa borneensis (Pfr.) (UF 35928). Figs.
20-21.-Phaedusa filicostata stenotrema, n.ssp. Holotype (UF 36426).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS 137
22
24 25
23
Figures 22-25. Phaedusa filicostata stenotrema n.ssp. Figs. 22, 23. Clausilium. lateral and
frontal views. Figs. 24, 25. Paratypes (UF 35926); location of plicae within last whorl. Scale
equals 5 mm.
BULLETIN FLORIDA STATE MUSEUM
Figures 26-31. Protoconch and embryonic sculpture of Bornean Georissa. Fig. 26.-G. hosei
G.-A. (UF 35919) (X92). Fig. 27.-G. hadra, n.sp. (UF 36109) (X92). Fig. 28.-G. kobelti Gr. (UF
36179) (X92). Fig. 29.-G. sauli (Ben.-Jutt.) (UF 36111) (X92). Fig. 30.-G. pyrrhoderma, n.sp.
(UF 36185) (X92). Fig. 31.-G. similis Smith (UF 36100) (X122).
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1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
0
0
Figures 32-38. Axial tubes in Bornean Georissa. Fig. 32.-G. hadra n.sp. (UF 36109) (X17). Fig.
33.-G. hosei G.-A. (UF 35919) (X21). Fig. 34.-G. pyrrhoderma n.sp. (UF 36185) (X21). Fig.
35.-G. everetti Smith (UF 36158) (X21). Fig. 36.-G. gomantonensis Smith (UF 36173) (X21).
Fig. 37.-G. similis Smith G.-A. (UF36100) (X28). Fig. 38.-G. williamsi G.-A. (UF 35912) (X28).
BULLETIN FLORIDA STATE MUSEUM
@
H
0
0
Figures 39-42. Inner view of opercula of some Bornean Georissa. Fig. 39.-G. hosei G.-A. (UF
35919) (X72). Fig. 40-.G. xesta, n.sp. (UF 35970) (X72). Fig. 41.-G. pyrrhoderma, n.sp. (UF
36185) (X72). Fig. 42.-G. similis Smith (UF 36100) (X108).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
Figures43-46. Georissa hadra, n.sp. Paratypes (UF36109). Fig. 43 (X18). Fig. 44 (X18). Fig.45
(X30). Fig. 46 (X44).
BULLETIN FLORIDA STATE MUSEUM
Figures 47-49a. Geoissa hosei odwin-Austen (UF 35919). Figs. 47-49 (X27). Fig. 49a (X70).
Figures 47-49a. Georissa hosei Godwin-Austen (UF 35919). Figs. 47-49 (X27). Fig. 49a (X70).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
Figures 50-52. Georissa kobelti Gredler (UF 36179). Figs. 50-51 (X32). Fig. 52 (X82).
119r
~4?61-~ I*Q-
BULLETIN FLORIDA STATE MUSEUM
Figures 53-54. Georissa sauli (Benthem-Jutting) (UF 36110). Fig. 53 (X26). Fig. 54 (X68).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
Figures 55-57. Georissa everetti Smith (UF 36158). Figs. 55-56 (X29). Fig. 57 (X77).
It
ik, %A~o k
BULLETIN FLORIDA STATE MUSEUM
Figures 58-60. Georissa gomantonensis Smith (UF 36173). (X28).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
61
62
Figures 61-62. Georissa borneensis Smith. Lectotype (BMNH 94.7.20.61). Scale equals 2 mm.
1 I I
BULLETIN FLORIDA STATE MUSEUM
Figures 63-65. Georissa pyrrhoderma, n.sp. Paratypes (UF 36185). Figs. 63,64 (X29). Fig. 65
(X38).
Vol. 29, No. 3
1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
Figures 66-68. Georissa williamsi Godwin-Austen (UF 35912) (X41).
BULLETIN FLORIDA STATE MUSEUM
Figures. 69-70. Georissa xesta, n.sp. Paratypes (UF 35970) (X33).
Vol. 29, No. 3
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1983 THOMPSON AND DANCE: BORNEAN NON-MARINE MOLLUSKS
72
Figures. 71-72. Georissa banguenyensis Smith. Lectotype (BMNH 93.6.7.9). Scale equals 1
mm.
BULLETIN FLORIDA STATE MUSEUM
0
Figures 73-75. Georissa similis Smith. (UF 3100) (X47).
Figures 73-75. Georissa similis Smith. (UF 36100) (X47).
0
Vol. 29, No. 3
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