Aphally in the stylommatophoran land snail Phaedusa (Clausiliidae:
Phaedusinae) in Timor and its systematic implications
Frank Köhler and Guilherme Burg Mayer
Natural Sciences, Australian Museum, Sydney, Australia
ABSTRACT
The Clausiliidae (subfamily Phaedusinae) of the Lesser Sunda island of Timor are poorly studied.
To date, one species with a rather confusing taxonomic history, Phaedusa timorensis Nordsieck,
2007, has been recorded from West Timor. Based on the study of recently collected material, we
present the first record of clausiliids from Timor-Leste (East Timor). Two species, which live in
sympatry with each other, are recognised and described based on comparative shell
morphology and mitochondrial differentiation: Phaedusa ramelauensis n. sp. and
P. angustocostata n. sp. Both species are ovoviviparous, retaining shelled juveniles in their
oviducts. One population of P. ramelauensis consists of individuals that lack male organs
while having fully functional female organs. As with similar cases in other stylommatophoran
families, the aphally is hypothesised to be a byproduct of either within-species
polymorphism or hybridisation and to have no bearing on the taxonomy of these terrestrial
snails. Because aphally is the key characteristic of Renschiphaedusa Loosjes and Loosjes-van
Bemmel, 1973, this genus-level taxon is placed in the synonymy of Phaedusa H. and
A. Adams, 1855.
http://zoobank.org/urn:lsid:zoobank.org:pub:528EB46B-5E4F-49B6-AECE-4DFE20B4B4A0
ARTICLE HISTORY
Received 16 July 2015
Final version received 21
December 2015
KEYWORDS
Lesser Sunda Islands;
ovoviviparity;
Renschiphaedusa;
reproductive anatomy
Introduction
The Clausiliidae is a hyper-diverse family of terrestrial
stylommatophoran gastropods with an almost global
distribution and centres of diversity in western
Eurasia, eastern Asia and South America. Strangely,
Clausiliidae do not occur in North America or in Austra-
lia (Uit de Weerd and Gittenberger 2013). Members of
this family are characterised by their mostly sinistral,
turreted to fusiform shell and the possession of a
complex internal mechanism enabling them to close
off the aperture of their shell, the clausilial apparatus
(CA) (e.g., Nordsieck 2007). Clausiliid snails often
dwell on hard substrates, such as rock faces and tree
trunks, and are frequently found in mountainous
regions, in particular on limestone, where they tend
to have narrow ranges of distribution (Loosjes 1953).
They are simultaneous hermaphrodites and generally
thought to reproduce by outcrossing (Nordsieck 2007).
Clausiliid taxonomy relies heavily on shell charac-
ters, including features of the CA, and the group has
been subdivided into several subfamilies with strong
emphasis on these characters. However, a molecular
phylogenetic study has recently demonstrated that
most shell characters are homoplasious, including fea-
tures of the CA with the exception of the apertural for-
mation (Uit de Weerd and Gittenberger 2013). In this
study, seven highly supported clades have been recog-
nised, which are more or less geographically confined
and have been considered to represent subfamilies
(Uit de Weerd and Gittenberger 2013). One of these
clades comprises the Phaedusinae (including Serruli-
nae). This subfamily has a disjunct distribution in the
eastern Mediterranean to Caucasus region (‘Serrulinae’)
and southeastern and eastern Asia (‘Phaedusinae sensu
stricto’) (Uit de Weerd and Gittenberger 2013, fig. 4).
The Phaedusinae is the only subfamily represented in
the Indo-Australian Archipelago (IAA) with the south-
ernmost limits of its distribution being on the island
of Timor (Nordsieck 2007: map 1 on p. 19).
The Phaedusinae of the IAA were most recently sys-
tematically revised by Loosjes (1953, 1963) with sup-
plementary data provided subsequently by Loosjes
and Loosjes-van Bemmel (1973) and Nordsieck (2002,
2007). While this group is generally poorly documen-
ted, our knowledge of the Timorese species is particu-
larly sparse.
This poor knowledge is caused by a general lack of
study of the island’s malacofauna. Historically, the
works of von Martens (1865, 1867, 1892, 1897) and
Rensch (1931, 1935) on non-marine gastropods of
Timor are particularly relevant. However, these
studies are entirely based on samples from West
Timor (formerly a Dutch colony). The eastern part of
the island (then a Portuguese colony, later annexed
by Indonesia, and now an independent country,
Timor-Leste) had remained almost entirely unstudied.
© 2016 Crown Copyright in the Commonwealth of Australia 2016 Department of the Environment
CONTACT Frank Köhler frank.koehler@austmus.gov.au
MOLLUSCAN RESEARCH, 2016
http://dx.doi.org/10.1080/13235818.2016.1150771
Downloaded by [Australian Museum], [Mr Frank Köhler] at 17:15 08 August 2016