Australian Museum, Sydney NSW, Australia
Molecular phylogeny and taxonomic revision of the genera Baudinella Thiele,
1931, Retroterra Solem, 1985 and Molema K€ ohler, 2011 endemic to the coastal
Kimberley, Western Australia (Gastropoda, Camaenidae)
FRANCESCO CRISCIONE and FRANK K
€
OHLER
Abstract
Baudinella Thiele, 1931, Retroterra Solem, 1985 and Molema K€ ohler, 2011 are three genera of camaenid land snail endemic to coastal regions of the
Western Australian Kimberley. Each of these genera has fairly distinct shells, but all exhibit a rather similar configuration of the reproductive system,
which is characterized by lack of a penial sheath and presence of an elongated and coiled bursa copulatrix. By combining comparative morphology of
shell and penial anatomy with analyses of the mitochondrial DNA fragments 16S rRNA (16S) and cytochrome c oxidase subunit 1 (COI), the phyloge-
netic relationships amongst representative species of these genera are addressed to test the monophyly of taxa and to identify new species. Our results
show that all three genera are members of a single camaenid radiation. Five new species are described: two new species of Baudinella, B. magna n.
sp. from the Institute and Montesquieu Archipelagos and B. margaritata n. sp. from Pitta Gorge in the Prince Regent Reserve, one new species of
Molema, Molema tenuicostata n. sp., from near Talbot Bay, and two new species of Retroterra, R. dichroma, and R. nana, from the Prince Regent
Reserve.
Key words: Stylommatophora – Helicoidea – mitochondrial DNA – genetic distances – new species
Introduction
For its remoteness and inaccessibility, the Western Australian
Kimberley has remained the most poorly known region in Aus-
tralia in terms of its biodiversity. However, several large surveys
conducted in the past four decades have improved the documen-
tation of biotic patterns throughout the Kimberley, which has
since emerged as one of Australia’s biodiversity hot spots
(McKenzie 1991; Gibson and McKenzie 2012).
With respect to camaenid land snails, the Kimberley is argu-
ably the major hot spot in Australia, for no other Australian
region of comparable size harbours more species. The inventory
of Kimberley land snails, including camaenids, has made signifi-
cant leaps forward during recent years. Collectively, the most
significant contributions to the current knowledge of the Kimber-
ley Camaenidae are the works of Alan Solem, who systemati-
cally surveyed and completely revised the entire fauna. In all,
Solem (1981a,b, 1984, 1985, 1988) newly described more than
100 camaenid species. At the time, Solem’s works on the Kim-
berley fauna quadrupled the number of known species highlight-
ing the Camaenidae as the taxonomically and ecologically most
diverse land snail group in the Kimberley. Ongoing revisions
have doubled species numbers again. Most significantly, K€ ohler
(2011a) newly described 83 species from islands off the Kimber-
ley coast. In addition, a series of taxonomic revisions contained
descriptions of many new species in several genera (K€ ohler
2010a,b, 2011b,c; Criscione et al. 2012; K€ ohler and Johnson
2012; Criscione and K€ ohler 2013a,b; K€ ohler and Criscione
2013a) raising the number of known species to 250.
Systematic studies have demonstrated that land snails are not
equally distributed over the land surface but that rainfall patterns,
soil type and topography govern the distribution of vegetation
types and, hence, patterns of species richness of local land snail
communities throughout the Kimberley (Solem and McKenzie
1991; Gibson and K€ ohler 2012; K€ ohler et al. 2012). Accord-
ingly, the richest land snail communities are found in vine
thicket patches in the high-precipitation zone of the north-wes-
tern coastal region between the Admiralty Gulf to the north and
the Camden Sound to the south (Solem and McKenzie 1991).
Fifteen camaenid genera are represented in this part of the
Kimberley, several of which are exclusively found here, such as
Youwanjela K€ ohler and Shea, 2012 (K€ ohler and Shea 2012).
Amongst these endemic camaenids of the high rainfall zone, the
taxonomy of three presumably closely related genera is revised,
Baudinella Thiele, 1931, Retroterra Solem, 1985, and Molema
K€ ohler, 2011. Their close relationships are underpinned by sev-
eral shared features of the shell and genital anatomy (Solem
1985; K€ ohler 2011a).
Baudinella comprises seven presently recognized species, most
of which are island endemics. Two species are found on the
mainland: B. baudinensis (Smith, 1893) occurs on Baudin Island
and Cape Voltaire, and B. regia Solem, 1985 in the Prince
Regent Nature Reserve (Fig. 1). Solem (1991) also listed two
undescribed taxa from coastal localities in the York Sound and
Prince Frederick Harbour. Baudinella species have repeatedly
been found in sympatry with species of Setobaudinia Iredale,
1933 having very similar, small and flat shells with reflected
apertural lips and widely open umbilici (K€ ohler 2011a; Criscione
and K€ ohler 2013c). Despite these similarities, the two genera
clearly differ in their genital anatomy and are phylogenetically
distinct (Solem 1985; Criscione and K€ ohler 2013c).
Retroterra is restricted to the wettest part of the Kimberley,
the Prince Regent Reserve (Solem 1985, 1991), including two
nearby islands, Unwins and Bongaree Islands, and Hanover Bay
(K€ ohler 2011a; Fig. 2). Retroterra species differ from other cam-
aenids in having a rather large, flat shell with extremely wide
umbilicus and regular radial sculpture. Typical features of the
genital anatomy include a well-developed penial verge and long,
convoluted bursa copulatrix (Solem 1985).
The only species of Molema, M. stankowskii K€ ohler, 2011, is
only known from north-west of Molema Island, Talbot Bay. The
species has a discoidal shell with a relatively large aperture whose
lip is expanded and reflected bearing no indentations. A large
stimulatory pilaster is a distinctive penial feature (K€ ohler 2011a).
Corresponding author: Frank K€ ohler (frank.koehler@austmus.gov.au)
Contributing authors: Francesco Criscione (francesco.criscione@austmus.
gov.au)
J Zoolog Syst Evol Res (2014) 52(4), 273--284
Accepted on 16 February 2014
© 2014 Blackwell Verlag GmbH J Zoolog Syst Evol Res doi: 10.1111/jzs.12065