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Land snails and slugs

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 ZOOLOGY

Land Snails and Slugs

Menno Schilthuizen

A
fter arthropods, the Mollusca are the most species-rich animal phylum
on land, where they come in three forms, all belonging to the class of
Gastropoda: shelled snails, shell-less slugs, and an intermediate form
called semislugs, which have a small, externally visible shell that is, however,
not large enough to contain the body.
Despite their diversity, the Bornean land gastropods are poorly investigated.
Although many species were described from collections made in the 19th
and early 20th century, these descriptions were often not unambiguous and
many genera are therefore in urgent need of revision. In the 1980s and 1990s,
malacologist Jaap Vermeulen published a series of important monographs in
the journal Basteria covering the microsnail family Diplommatinidae and
several other groups that show high endemism on limestone outcrops. Other
than that, however, until recently, very few comprehensive treatments have
appeared.
More recently, attempts have been underway to provide a complete
inventory (estimated richness: 350 species) of the terrestrial malacofauna of
the state of Sabah, which so far has resulted in a paper (Vermeulen et al., 2015;

Gunung Penrissen: The Roof of Western Borneo

ZooKeys, 531: 1–139) providing diagnoses of 79 species (of which 48 new
to science) and a yet unpublished manuscript for a field guide by Vermeulen,
Liew & Schilthuizen. Moreover, a cybertaxonomic website aiming to provide
updated access to all species from Borneo has been set up by Thor-Seng Liew
of Universiti Malaysia Sabah (borneanlandsnails.myspecies.info). A better
overview of Borneo’s estimated 1,500 species is urgently needed, because
of the increasingly important role snails are beginning to play in ecological
studies on the island.
Outside of limestone outcrops, very few surveys of the malacofauna of
the interior of Sarawak have been carried out. Only an inventory of Lanjak
Entimau by Huong & Liew (2011) is available. In this chapter, I describe a
(very incomplete) attempt at a first inventory of the malacofauna of Penrissen.
Despite the wetness of the habitat, snails are not abundantly present in tropical
rainforests on acid soils. To obtain a first impression of the Penrissen snail
fauna, one has to check every leaf and rock, and go through large amounts of
leaf litter and soil. Even such a thorough search will only yield small numbers
of specimens.

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 LAND SNAILS AND SLUGS

At Penrissen, we did precisely that. On April 5th and 6th, 2016, both
by day and at night, we walked trails and other areas (incl. the golf course)
around the Borneo Highlands Resort. In addition, I reviewed photographic
records taken on March 18th, 2016. During these walks, the vegetation, the
surfaces of tree trunks and branches as well as rocks, leaf litter and deadwood
were checked for the presence of gastropods and/or their shells. In addition,
three quantities of ca. five litres each of leaf litter and soil were collected from
the base of rock faces and trees. This material was first coarsely sieved with a
10 mm mesh width beetle sieve in the field. Then enriched by flotation and the
flotsam was dried and sieved in a sieve cascade and sorted for shells under a
dissection microscope. Live specimens were photographed.
In total, this procedure yielded 90 individuals were found (including
both dead and alive ones), belonging to at least 25 species. The checklist
provides their names as well as the locations and numbers. Below, I give some
additional information on the species found.
The Bornean land snails consist of two large and unrelated groups,
superficially similar, but in body structure very different. First, the
“prosobranchs”, which have gills, separate sexes, a shell piece that closes off
the aperture (“operculum”), and eyes at the base of the tentacles. Second,
the “pulmonates”, which have lungs, are hermaphrodites (they are male
and female at the same time), don’t have an operculum, and carry their
eyes at the tips of their tentacles. Prosobranchs are much more vulnerable
to habitat disturbance than pulmonates. We found two prosobranch families,
Cyclophoridae and Diplommatinidae. We recorded four species of the former
Gunung Penrissen: The Roof of Western Borneo

family: (1) Alycaeus globosus Adams 1870 (Fig. 1), recognizable by its small

Fig. 1. (left). Alycaeus globosus, a colourful representative of so-called prosobranch

snails, which carry their eyes at the base of the tentacles, and are able to close off the
shell with an “operculum”, a small door-like shell-piece. Fig. 2 (right). A juvenile
Cyclophorus, presumably C. perdix borneensis.
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 LAND SNAILS AND SLUGS

number of somewhat inflated whorls and widely-spaced, fine radial ribs. This
species is known from limestone and montane forest throughout Sarawak
and Sabah. (2) A juvenile of a Cyclophorus species (Fig. 2). The image does
not provide enough features to reliably identify the species, but based on the
coloration and the sculpture, as well as the location, it seems probable that
this is C. perdix borneensis (Metcalfe, 1851). (3) A species of Ditropopsis
was found that is similar to D. koperbergi (Zilch, 1955), a species so far only
recorded from Kalimantan and Sabah, except for the following differences:
on the last whorl there are two spiral ribs instead of just one; also, the shell
is relatively tall conical, more like D. trachychilus than D. koperbergi. For
the time being, the specimen is best referred to as D. cf. koperbergi, although
it might, in fact, represent a new species. (4) Japonia barbata (Pfeiffer,
1855) (Fig. 3) carries numerous spiral ribs of which three are spectacularly
adorned with long periostracal hairs (hence the name barbata, which means
“bearded”).
We found two species of the microsnail family Diplommatinidae,
which includes many species endemic to limestone hills, but also more
widespread taxa that are not confined to karst. One of these was recorded here:
Diplommatina whiteheadi Smith 1898. This species was found in Sarawak
once before, in Lanjak Entimau, close to the present locality. The second
diplommatinid, D. busanensis Godwin Austen 1889 (Fig. 4) was found on
dead leaves on the ground. It is a well-known species from the limestone hills

Gunung Penrissen: The Roof of Western Borneo

Fig. 3 (left). Japonia barbata, with its curious “beard” of bristles on the periostracum,
the protein layer covering the shell. Fig. 4 (right). Diplommatina busanensis, two
individuals of this tiny, anticlockwise-coiled snail foraging on a dead leaf.

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 LAND SNAILS AND SLUGS

south of Kuching, but this is the first record from higher elevation and away
from limestone.
All the following families belong to the pulmonates. First, of the
Valloniidae, a single empty shell of Ptychopatula orcella (Stoliczka 1873)
was found in leaf litter along the Kalimantan trail. This is a widespread
species throughout Southeast Asia. Similarly, the only subulinid found was
the common and widespread (and possibly introduced) Allopeas gracile
(Hutton 1834), found on the golf course.
The large-bodied and interesting family Dyakiidae was represented by
two species. First, the beautiful Dyakia regalis (Benson 1850) (Figs. 5-7).
Interestingly, the population is variable in both colour and coiling direction. We
found three sinistral (anticlockwise-coiled) adults of the dark form, one dextral
Gunung Penrissen: The Roof of Western Borneo

Fig. 5 (left). Dyakia regalis, a clockwise coiled specimen of this variable species. Fig.
88 6 (top right). Dyakia regalis, an anticlockwise-coiled specimen. Fig. 7 (bottom right).
Dyakia regalis, anticlockwise-coiled, pale form.
 LAND SNAILS AND SLUGS

(clockwise coiled) juvenile of the dark form, one sinistral juvenile of the dark
form, and two sinistral adults of the pale form. In shell shape and sculpture,
the pale and dark forms appear identical, and so do the dextral and sinistral
forms, and it all appears to represent variation within the species. Among
the specimens figured on Bornean Terrestrial Snails (borneanlandsnails.
myspecies.info) there are also multiple colour forms, and Benson himself, in
the original description, mentions two varieties, a uniform dark and a striped
one. Such intraspecific colour variation is commonly reported in land snails.
Variation in coiling direction, however, is rare, because normally, dextral and
sinistral individuals cannot mate. The second dyakiid was a juvenile empty
shell, very similar to Everettia cf. planispira Liew, Schilthuizen & Vermeulen
2009.
The Chronidae, consisting of small snails, difficult to identify, were
represented by at least three species of the genus Kaliella. The snails of this
genus are live-bearing, meaning that the eggs hatch into baby snails while
still in the mother’s body. Indeed, in some of the
specimens from Penrissen, such offspring
were present inside the collected live
specimens. We found the following
species. Kaliella calculosa (Gould
1852), a widespread species (it
occurs throughout Southeast Asia
and the Pacific). Kaliella sp. “A”,
an unidentified species from leaf

Gunung Penrissen: The Roof of Western Borneo

litter at the foot of large boulders
on the golf course. The combination
of shell characters of this species
could not be reliably matched to any
known Borneo species. Finally, Kaliella
microconus (Mousson 1865) (Fig. 8) was
found alive on vegetation. This species Fig. 8. Kaliella microconus, a tiny
has been recorded throughout Borneo as (2 mm tall) snail from the family
well as in other parts of Southeast Asia Chronidae.
and the Pacific.
The largest family in Penrissen was the Ariophantidae, represented
by seven species. The largest of these was (1) Hemiplecta buettikoferi
Schepman 1896 (Fig. 9). The specimens are externally very similar to the H.
humphreysiana commonly encountered in the lowlands of Sabah. However,
I dissected one adult and found that the genitalia are different from those
described for H. humphreysiana by Godwin Austen (1901; as cited on Bornean

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 LAND SNAILS AND SLUGS

Terrestrial Snails), but more resemble the description given by Schepman

(1896) for H. buettikoferi from Kapit. (2) Macrochlamys tersa (Issel 1874)
(Fig. 10) was found on one of the large boulders on the golf course. M.
tersa is widespread and common throughout Sabah and Sarawak, and often
encountered in secondary vegetation. One broken shell, characteristic of (3)
Rahula raricostulata (Smith 1894) was recorded from leaf litter along the
Kalimantan trail. The species was described from southern Sarawak. (Fig.
11). (4) A species that, in shell shape and sculpture, strongly resembles M.
physotrochus (a recently described species from Sabah and Sarawak). The shell
Gunung Penrissen: The Roof of Western Borneo

Fig. 9 (left). Hemiplecta buettikoferi, a common and large (shell ca. 4 cm across) snail
from primary forest. Fig. 10 (top left). Macrochlamys tersa, a small (shell diameter ca.
4 mm) and common snail throughout Borneo. Like all ariophantids, it has a “horn” at
the trailing tip of the foot. Fig. 11 (bottom left). Microcystina cf. physotrochus, a tiny,
dark species of the ariophantid genus Microcystina in wet weather on a twig of moss.
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 LAND SNAILS AND SLUGS

is, however, very dark, as is the body of the animal. The live snail therefore,
appears nearly black, which, in M. physotrochus from lowland in Sabah, is not
the case to such a degree. It is also remarkable that the protoconch in all living
(sub)adults was strongly eroded (see Fig. 11), suggesting the animal may be
affected by the acidity of the environment. (5) We found a Vitrinula species
that I tentatively place in Vitrinula thisbe (Smith, 1895) (Fig. 12), originally
described from Mount Rabong. The animal has two finger-like shell lobes, the
right longer than the left, which are used to sweep over the surface of the shell
when it is crawling. The shell is transparent and almost completely smooth.
The Ariophantidae also contain the only two semislugs found. The most
common of these (Fig. 13) was a slender animal, pale yellowish grey, with
indistinct dark grey markings on the mantle lobes and the sides of the foot.

Gunung Penrissen: The Roof of Western Borneo

Fig. 12 (top). Vitrinula cf. thisbe. The species of the genus Vitrinula have (as their
name belies) a glass-like shell. This species also carries a brown spiral band just
beneath the shell’s edge. Fig. 13 (bottom). Damayantia simrothi is a semislug: a slug
with a small, externally visible shell, too small for the animal to withdraw into.
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 LAND SNAILS AND SLUGS

Because of its long, laterally compressed foot, it is probably a member of the

genus Damayantia, most probably (6) D. simrothi Collinge 1903, described
from Kuching. The species was found along all trails, often in relatively large
numbers (in various sizes, presumably including many juveniles) on leaves,
especially at night. We collected ten individuals. We could have collected more,
were it not for the annoying habit of these and other related ariophantids, that
Godwin Austen (1903) describes as follows: “As is well known by those who
have collected these exotic slugs, and noticed by Semper and others, some
species fling the tail right and left with great force and rapidity when they are
touched and will thus throw themselves off the hand”. The other semislug
was (7) a Microparmarion species (Fig. 14), photographed by Pui Yong Min.
I am unable to assign this specimen to a described species; externally, it most
resembles an unidentified semislug that we found in Sabah, Upper Padas (Fig.
7 in Schilthuizen & Liew, 2008. Basteria, 72: 287-306).
The only certainly introduced species, was the bradybaenid Bradybaena
similaris Férussac, 1821, also known as “Asian tramp snail” (Fig. 16).
It was very common on the signboards and lamp-posts of the golf course.
This is an introduced circumtropical species, common in disturbed habitats
throughout Southeast Asia. Its native range is possibly mainland east Asia.
The Bradybaenidae are often united with the Camaenidae, the latter family
represented at Penrissen only by Ganesella acris (Benson, 1859) (Fig. 15).
This is a widespread species throughout south-east Asia, also known from
Sabah and Peninsular Malaysia.
The largest native snail of Borneo, Bertia brookei (Adams & Reeve 1848)
Gunung Penrissen: The Roof of Western Borneo

was the only representative found of the Trochomorphidae: one juvenile on

the Batu Pangah trail, feeding on mushrooms, and one subadult empty shell
in the same area.

Fig. 14 (left). Microparmarion sp. A yet unidentified species of semislug. Fig. 15

(right).Ganesella acris, like many Camaenidae, has a pattern of tiny granules on its
92 shell.
 LAND SNAILS AND SLUGS

Figure 16. Bradybaena similaris, the “Asian tramp snail”, a common introduced
species throughout Asia.

Finally, two species of the minuscule Charopidae were found: First,

Charopa sp. “argos”. This minute (1.05 mm diameter) species, characterized
by the white translucent shell, widely spaced radial ribs, and open umbilicus
(the “hole” at the bottom of the shell encircled by the whorls), is endemic to
Borneo, and common throughout Sabah, Brunei, Sarawak, and Kalimantan.
However, it remains nameless until it is included in a taxonomic revision
(Vermeulen, Liew & Schilthuizen, in prep.) of the entire genus Charopa from

Gunung Penrissen: The Roof of Western Borneo

Borneo. Second, a Philalanka species that somewhat resembles Philalanka
rugulosa Vermeulen, Liew & Schilthuizen 2015, an endemic species from
Mount Kinabalu. The Penrissen material is smaller (shell width less than 2
mm), and the edge of the umbilicus is more sharply defined, due to an internal
columellar cord.
In the streams crossing the golf course, we also found large numbers of
two species of freshwater snail: one belonging to the Lymnaeidae and one to
the Thiaridae. Since the focus of this inventory was land snails and semislugs,
we did not identify this material any further.
Overall, this brief inventory of the malacofauna of Penrissen offers
interesting prospects for further study. The variability in colour and coiling
direction of Dyakia regalis is a welcome addition to the limited number of
species of land snails that show such variability. This will permit future research
on the evolution of these character systems. The Penrissen malacofauna also
offers a good representation of families that are in urgent need of taxonomic
revision, especially Ariophantidae. We recommend further malacological
studies be undertaken at this interesting location.
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 CHECKLIST OF SNAILS AND SLUGS

Classification largely follows the website Bornean Terrestrial Molluscs, http://

borneanlandsnails.myspecies.info. The abbreviations refer to the localities:
KT, Kalimantan Trail; PT, Batu Pangah Trail; GC, Golf Course. The numbers
refer to the individuals found, with “L”, live and “S”, empty shells.

Family Scientific Name Locality

Cyclophoridae Alycaeus globosus Adams, 1870 KT; 3L
Cyclophorus perdix borneensis PT; 1L
Ditropopsis cf. koperbergi (Zilch, 1955) PT; 1L
Japonia barbata (Pfeiffer, 1855) KT: 2L
Diplommatinidae Diplommatina whiteheadi Smith, 1898 KT; 5S
Diplommatina busanensis Godwin Austen,
KT; 7L
1889
Valloniidae Ptychopatula orcella (Stoliczka, 1873) KT; 1S
Subulinidae Allopeas gracile (Hutton, 1834) GC; 5S
Trochomorphidae Bertia brookei (Adams & Reeve, 1848) PT; 1L+1S
Dyakiidae Dyakia regalis (Benson, 1850) PT; 7L
Everettia cf. planispira Liew, Schilthuizen
KT; 1S
& Vermeulen, 2009
Chronidae Kaliella calculosa (Gould, 1852) PT; 1S
Gunung Penrissen: The Roof of Western Borneo

Kaliella sp. A GC; 2S

Kaliella microconus (Mousson, 1865) KT; 3L
Ariophantidae Hemiplecta buettikoferi Schepman, 1896 PT; 6L+1S
Macrochlamys tersa (Issel, 1874) GC; 1L
Rahula raricostulata (Smith, 1894) KT; 1S
Microcystina cf. physotrochus Vermeulen,
PT; 5L+1S
Liew & Schilthuizen, 2015
Vitrinula cf. thisbe (Smith, 1895) PT; 2L
PT, KT;
Damayantia simrothi Collinge, 1903
10L
Microparmarion sp. PT; 1L
Bradybaenidae Bradybaena similaris Férussac, 1821 GC; 3L
Camaenidae Ganesella acris (Benson, 1859) PT, KT; 6L
Charopidae Charopa sp. “argos” KT; 1S
Philalanka cf. rugulosa Vermeulen, Liew
94 & Schilthuizen, 2015
KT; 6S

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