Print
A glimpse in the dark? A first phylogenetic approach in a widespread freshwater snail from tropical Asia and northern Australia (Cerithioidea, Thiaridae)
expand article infoDusit Boonmekam, Duangduen Krailas, France Gimnich§, Marco T. Neiber|, Matthias Glaubrecht|
‡ Silpakorn University, Nakhon Pathom, Thailand
§ Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany
| Universität Hamburg, Hamburg, Germany
Open Access

Abstract

Thiaridae are a speciose group of freshwater snails in tropical areas including a high number of described nominal taxa for which modern revisions are mostly lacking. Using an integrative approach, the systematic status of a group of thiarids from the Oriental region, including the nominal species Melania aspera and M. rudis, is reassessed on the basis of shell morphology and biometry, radula dentition patterns, and reproductive biology along with molecular genetic methods. Our results suggest that populations from the Oriental region cannot be distinguished on the basis of shell morphology, radula characters and their reproductive mode and are monophyletic based on mitochondrial sequences. Hence, M. rudis with M. aspera are regarded as belonging to the same species along with several other nominal taxa that were previously included in M. rudis. Moreover, populations from Thailand and Australia, from where the species was not previously recorded, could be shown to form a monophyletic group together with samples from Indonesia. However, a generic affiliation with Thiara, in which the investigated taxa were often included in the past, was not supported in our phylogenetic analyses, highlighting the need for a comprehensive revision of the genus-group systematics of Thiaridae as a whole.

Key Words

Cerithioidea, evolutionary systematics, Oriental region, Thailand

Introduction

Despite advances in the understanding of the family-level phylogeny of Cerithioidea Fleming, 1822, the taxonomical diversity in Thiaridae Gill, 1871 (1823) is still not well understood, and evolutionary systematic research in the sense of Glaubrecht (2010) in this particular family is still in its infancy. The Thiaridae, in earlier treatments subsumed under the name Melaniidae Children, 1823, have been used as a “rubbish bin” to accommodate all freshwater lineages belonging to the Cerithioidea. Only after the removal of the families Pachychilidae Fischer & Crosse, 1892, Melanopsidae Adams & Adams, 1854, Paludomidae Stoliczka, 1868, Pleuroceridae Fischer, 1885 (1863), and Semisulcospiridae Morrison, 1952) (Campbell 2019; Neiber and Glaubrecht 2019b, 2019c, 2019d; Strong and Lydeard 2019 and references therein) and recently the Neotropical Hemisinidae Fischer & Crosse, 1891 (Glaubrecht and Neiber 2019a), a more accurate circumscription of “core” Thiaridae began to emerge on the basis of molecular and/or morphological evidence (e.g., Glaubrecht 1993, 1996, 2011; Holznagel and Lydeard 2000; Lydeard et al. 2002; Glaubrecht et al. 2009; Strong 2011; Strong et al. 2011).

In addition to uncertainties in the delimitation of genera, research on thiarids is further complicated by the large disparity of shell characters among species, a large phenotypic plasticity within species and a high ecological adaptability that is, however, also known from other limnic Cerithioidea. This conchological variability has certainly led to an overestimation of the number of species in the past, as specifically shown for limnic lineages in the superfamily (Glaubrecht 1993, 1996; Köhler and Glaubrecht 2001, 2003, 2006; Glaubrecht and Köhler 2004; Glaubrecht et al. 2009), but may also cause problems in delimiting species resulting in an underestimation of the actual morphological disparity versus the taxonomical diversity, at least in some cases. These problems are exacerbated by the putatively widespread occurrence of parthenogenesis in different lineages of Thiaridae (Glaubrecht 1996) and the associated problems of what is actually meant by “species” in this case (e.g. Hausdorf 2011). Additionally, Thiaridae have also realised different life history strategies that were characterised by Glaubrecht (1996, 1999, 2006, 2011) by the duration of ontogenetic stages to remain within a specialised structure of the female, viz. the subhaemocoelic brood pouch. While in some thiarids only very early ontogenetic stages, i.e. embryos without shell, develop and are released as veligers (ovoviviparity), other thiarid species brood and even transform their subhaemocoelic brood pouch into a matrotrophic organ or “pseudoplacenta” that apparently nourishes the developing juveniles, as e.g. in the Southeast Asian thiarid Tarebia granifera (Lamarck, 1816) (euviviparity, see Glaubrecht 1996; Glaubrecht et al. 2009; Maaß and Glaubrecht 2012; Veeravechsukij et al. 2018b). Finally, some thiarids also have an extraordinarily high invasive potential, such as Melanoides tuberculata (Müller, 1774) and Tarebia granifera and today have an almost pantropical distribution (e.g., Brown 1994; Glaubrecht 1996).

To date, only few of the several dozen thiarid taxa have seen closer investigation. Glaubrecht et al. (2009) and Maaß and Glaubrecht (2012) surveyed the thiarid fauna of Australia. Dechruska et al. (2013) evaluated the status and identity of the nominal taxon Melania jugicostis Hanley & Theobald, 1876 from the Southeast Asian mainland, and Veeravechsukij et al. (2018a, 2018b) investigated the phylogeography and reproductive biology of T. granifera and its trematode parasites. However, many other named taxa have been rarely studied and, thus, remain enigmatic and even pure nomenclatorial “ghosts” with highly questionable status as evolutionary relevant entities, which hampers further insights into the systematics, biogeography, and evolution of these freshwater gastropods otherwise under scrutiny, e.g., in speciation and/or radiation studies.

Melania aspera Lesson, 1831, which was originally described from New Guinea (Lesson 1830–1831), is such an “enigmatic” taxon (Fig. 1), which Glaubrecht and Podlacha (2010) regarded as a possible senior synonym of the nominal species Melania rudis Lea & Lea, 1851. The latter taxon is usually regarded as belonging to Thiara Röding, 1798 and thought to be relatively widespread, being reported from several countries, occurring from India and Sri Lanka to Southeast Asia and the Indo-Australian archipelago (Schepman 1892, 1915; Rensch 1934; van Benthem Jutting 1937; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Patil and Talmale 2011, see also Fig. 2). However, actual distribution records are relatively scarce in the literature and the distinction from other nominal thiarid taxa remains uncertain so far.

Figure 1. 

Shells of “Thiaraaspera (Lesson, 1831). A. Holotype of Melania aspera Lesson, 1831, MNHN 21098, ‘La Nouvelle-Guinée’ [more specifically Manokwari on New Guinea Island, West Papua, Indonesia, see Glaubrecht and Podlacha 2010]; B. Syntype of Melania rudis Lea & Lea, 1851, USNM 119778, Amboyna; C. Syntype of Melania microstoma Lea & Lea, 1851, USNM 119722, mountain streams, isle of Negros, Philippines; D. ZMB 107002, Calcutta, India; E. ZMB 107003, Ceylon, Sri Lanka; F. ZMB 127534, Don Ko Canal, Nakhon Pathom, Thailand; G. ZMB 127535, Don Ko Canal, Nakhon Pathom, Thailand; H. ZMB 127535, Don Ko Canal, Nakhon Pathom, Thailand; I. ZMB 191279, Yehembang River, Bali, Indonesia; J. ZMB 191279, Yehembang River, Bali, Indonesia; K. ZMB 106472, Yehembang, Bali, Indonesia; L. East of Mendaya, stream southwest of Gumicik, Bali, Indonesia; M. ZMB 191278, stream at Tembeeha, road Tirobus-Kendari, Southwest Sulawesi, Indonesia; N. ZMB 107378, Banggai Islands, Peleng Island, West of Peninsula, Tataban river, Central Sulawesi, Indonesia; O. ZMB 107377, Banggai Islands, Peleng Island, West of Peninsula, Tataban river, Central Sulawesi, Indonesia.river; P, Q. ZMB 107617, Wabalarr, Roper River, Northern Territory, Australia; R. ZMB 106599, Berry Springs, Northern Territory, Australia. Scale bar: 1 cm.

Figure 2. 

Distribution and reproductive strategy of “Thiaraaspera (Lesson, 1831). Stars: type localities of a) Melania aspera Lesson, 1831, Monokwari, New Guinea, b) Melania rudis Lea & Lea, 1851, Amboyna and c) Melania microstoma Lea & Lea, 1851, mountain streams, isle of Negros, Philippines. Pie charts show the percentages of offspring in the brood pouch of female T. aspera in different size classes as defined in Glaubrecht et al. (2009), see inset. The numbers near the pie charts refer to the number of individuals examined per population. Filled circles: material preserved in ethanol; open circles: dry shells.

As a further contribution towards a better understanding of thiarid diversity, we here re-evaluate the identity of M. aspera and M. rudis on the basis of museum samples including available type material as well as material collected during ongoing surveys in Southeast Asia using shell morphology and biometry, radula dentition patterns, and reproductive biology along with molecular genetic methods. Nomenclatural issues and the synonymy of the genus Thiara are also discussed.

Material and methods

This study is mainly based on the examination of specimens in the collections of the Parasitology and Medical Malacology Research Unit, Department of Biology, Faculty of Science, Silpakorn University, Thailand and the Museum für Naturkunde, Berlin, Germany, and supplemented by material from other museums (see below). Additionally, new samples were collected using hand picking and scooping methods in Thailand and Australia. Specimens were fixed in 75–96% ethanol.

Collection acronyms

MNHN Muséum National d’Histoire Naturelle de Paris, France

SUT Silpakorn University, Nakhon Pathom, Thailand

USNM National Museum of Natural History, Washington, USA

ZMB Museum für Naturkunde, Berlin, Germany (formerly Zoologisches Museum Berlin)

Coordinates (WGS84) of localities were taken with a GPS device or determined as accurately as possible from a map. Sampling sites were then mapped on a dot-by-dot basis to a digitally reduced version of the drainage pattern map of the Indo-Australian region. This map was prepared using a relief map on the basis of the Global30-Arc-Second Elevation Data (GTOPO30) from the U.S. Geological Survey and a river map from the map server Aquarius Geomar; and then compiled using Adobe Photoshop CS3 and Adobe Illustrator. For the exact locality data, see the material examined section.

Shell characters

Specimens were photographed using a digital EOS 350D camera (Canon, Tokyo, Japan). Standard biometric parameters were taken from each shell using electronic callipers (accuracy 0.1 mm): shell height (H), shell width (W), aperture length (AL; measured from the upper apertural angle to the farthest point on the basal margin of the aperture), aperture width (AW; measured perpendicular to AL as the widest distance between outer apertural margin and outer margin of parietal callous), height of the body whorl (BW), and number of whorls (NW) as shown in Figure 3A. To reduce dimensionality a principal component analysis was conducted on log-transformed shell measurements using R 3.3.2 (R Core Team 2016). Only the minimal number of PCA axes that accounted for more than 95% of the cumulative variation were used for further testing.

Figure 3. 

Measured shell parameters. A: H – shell height; W – shell width; BW – body whorl height; AL – aperture length; AW – aperture width. B: he – height of embryonic shell; we – width of embryonic shell; de – maximum diameter at one whorl.

The Shapiro-Wilk test was conducted in R to test for normal distributions of PCA 1 and PCA 2 values, respectively, for the here proposed geographic subgroups, i.e., samples from 1) Thailand, 2) Indonesia, 3) India, and Sri Lanka, and 4) Australia. Since some of the Shapiro-Wilk tests were significant (p ≤ 0.05), the non-parametric Kruskal-Wallis rank sum test was conducted for PCA 1 and PCA 2 assuming the grouping of specimens according to geography followed by Dunn’s test (Bonferroni-corrected) as post-hoc test as implemented in the R package “dunn.test 1.3.5” (Dinno 2017) in case that the Kruskal-Wallis-rank-sum tests were significant.

Radula preparation

Shells of representative specimens were cracked with a small vice and removed from the soft body parts, which were afterwards examined and dissected with the aid of a Leica Wild MZ 9.5 stereo microscope (Leica Microsystems, Wetzlar, Germany). Radulae were extracted following the protocol of Holznagel (1998), fixed on aluminium stubs, and coated with platinum using a Polaron SC 7640 Sputter Coater (Quorum Technologies, East Grinstead, UK). Radulae were then viewed and photographed (oriented so that denticles on the teeth were well visible) with a scanning electron microscope (SEM) EVO LS10 (Zeiss, Oberkochen, Germany).

Content of brood pouch

The brood pouch was opened after removing the mantle and its content was counted under a Leica Wild MZ 9.5 stereo microscope. Both, shelled juveniles and embryos, were grouped into standard size classes as described in Glaubrecht et al. (2009). Embryos and juveniles from representative specimens were fixed on aluminium stubs, air-dried, coated with platinum using a Polaron SC 7640 Sputter Coater, and then viewed, photographed, and measured (Fig. 3B) with a EVO LS10 SEM. Parameters of the embryonic shell were measured from SEM images as shown in Figure 3B: diameter of first half whorl (de; measured as the maximal witdth of the shell after 0.75 turns of the suture line), width of first quarter whorl (he; measured parallel to de as the distance from the starting point of the suture to the point after 0.25 turns of the suture line), width of first half whorl (we; measured perpendicular to de as the distance from the starting point of the suture to the point after 0.5 turns of the suture line).

Molecular methods and phylogenetic analyses

Total genomic DNA was extracted from ethanol-preserved foot muscle tissue using a CTAB protocol as described by Winnepenninckx et al. (1993) from 31 thiarid specimens and Paludomus siamensis Blanford, 1903 as outgroup representing one of the cerithioidean families, which have been shown to be closely related to the Thiaridae (Wilson et al. 2004; Strong et al. 2011).

For phylogenetic analyses, fragments of the mitochondrial cytochrome c oxidase subunit 1 (cox1) gene and the 16 S rRNA (16S) gene were amplified by polymerase chain reaction (PCR) using the primer pairs LCO1490 (5’-GGT CAA CAA ATC ATA AAG ATA TTG G-3’; Folmer et al. 1994) plus HCO2198var (5’-TAW ACT TCT GGG TGK CCA AAR AAT-3’; Rintelen et al. 2004) and 16S_F_Thia2 (5’-CTT YCG CAC TGA TGA TAG CTA G-3’; Neiber and Glaubrecht 2019a, see also Gimnich 2015) plus H3059var (5’-CCG GTY TGA ACT CAG ATC ATG T-3’; Wilson et al. 2004), respectively. Amplifications were conducted in 25 µl volumes containing 50–100 ng DNA, 1× PCR buffer, 200 mM of each dNTP, 0.5 mM of each primer and 1 U of Taq polymerase. After an initial denaturation step of 3 min at 94 °C, 35 cycles of 30 s at 94 °C, 60 s at 45–62°C and 60–120 s at 72°C were performed, followed by a final extension step of 5 min at 72°C. PCR products were purified using a NucleoSpin Extract II Kit (Macherey–Nagel, Bethlehem, PA, USA). Both strands of the amplified gene fragments were cycle-sequenced using the primers employed in PCR with the Big Dye Terminator chemistry version 1.1 (Applied Biosystems, Inc., Waltham, MA, USA). Sequences were visualised on an ABI 3130xl or ABI 3730xl Genetic Analyzer (Applied Biosystems, Inc.).

Forward and reverse sequence reads were assembled with CODONCODE ALIGNER v. 3.7.1 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye. For information on vouchers, see Table 1. The protein coding cox1 sequences were aligned with MUSCLE (Edgar 2004) as implemented in MEGA 6 (Tamura et al. 2013) under default settings. The 16S sequences of were aligned with MAFFT (Katoh and Standley 2013) using the Q-INS-i iterative refinement algorithm and otherwise default settings, because this algorithm has been described to perform better for the alignment of sequence data sets that may contain deletions and insertions than alternative multiple sequence alignment methods (Golubchik et al. 2007).

Museum registration numbers, GenBank accession numbers and locality data for the specimens used in the molecular phylogenetic analyses. Abbreviations for countries: AUS – Australia, IDN – Indonesia, IND – India, THA – Thailand.

Taxon Museum number Extraction number Country Latitude Longitude GenBank accession number
cox1 16 S rRNA gene
Thiaraaspera SUT 0311020 11449 THA 13°38'08"N, 100°05'03"E MK879291 MK879427
SUT 0312070 11446 THA 13°48'08"N, 100°02'06"E MK879292 MK879428
SUT 0311044 9603 THA 13°38'08"N, 100°05'03"E MK879290
ZMB 191268 2200 IDN 03°39'28"S, 122°13'52"E MK879296 MK879434
ZMB 191488 4558 IDN 08°38'39"S, 115°16'38"E MK879297 MK879435
ZMB 107377 6494 IDN 01°32'18"S, 122°51'28"E MK879293 MK879429
ZMB 107378 6495 IDN 01°32'18"S, 122°51'28"E MK879294 MK879430
ZMB 107617 7586 AUS 14°56'02"S, 133°10'26"E MK879295 MK879433
ZMB 107617 8743 AUS 14°56'02"S, 133°10'26"E MK879431
ZMB 107617 8744 AUS 14°56'02"S, 133°10'26"E MK879432
Stenomelaniadenisoniensis ZMB 106682 7599 AUS 14°55'47"S, 133°08'44"E MK879288 MK879425
ZMB 106632 7602 AUS 15°00'42"S, 133°14'25"E MK879287 MK879424
Thiara amarula ZMB 191489 2886 IDN 01°26'43"S, 127°29'01"E MK879289 a MK879426 a
ZMB 107472 6496 IDN 03°35'28"S, 128°08'42"E MK094074 MK098355
Thiara winteri ZMB 106554 1043 IDN 08°23'38"S, 114°45'04"E MK879301 MK879439
ZMB 190261 1055 IDN 02°35'34"S, 120°54'10"E MK879302 MK879440
Thiara cf. winteri ZMB 106472 1001 IDN 08°23'38"S, 114°45'04"E MK879298 MK879436
ZMB 191279 2232 IDN 08°23'36"S, 114°45'04"E MK879299 MK879437
ZMB 191279 4559 IDN 08°23'36"S, 114°45'04"E MK879300 MK879438
Mieniplotia scabra ZMB 107382 6514 IDN 00°48'33"N, 127°17'40"E MK879279 MK879416
ZMB 107564 7340 AUS 14°55'38"S, 133°07'06"E MK879280 MK879417
ZMB 127495 9574 THA 07°55'15"N, 099°15'47"E MK879285 MK879422
SUT 0312060 9578 THA 12°51'15"N, 099°59'49"E MK879278 MK879415
SUT 0311024 9580 THA 14°54'04"N, 100°03'48"E MK879276 MK879413
SUT 0311040 9582 THA 13°25'07"N, 099°57'18"E MK879277 MK879414
ZMB 127470 9589 THA 08°27'09"N, 098°28'01"E MK879284 MK879421
ZMB 127468 9599 THA 12°56'54"N, 099°28'52"E MK879283 MK879420
ZMB 107962 9779 THA 16°37'38"N, 100°56'43"E MK879282 MK879419
ZMB 107869 9781 THA 08°38'18"N, 099°44'59"E MK879281 MK879418
SUT 0311009 9787 THA 16°11'33"N, 099°15'51"E MK879275 MK879412
Melanoides tuberculata ZMB 200313 7530 IND 11°34'45"N, 076°34'55"E MK879274 MK879411
Paludomus siamensis ZMB 107721 7334 THA 14°26'15"N, 098°51'11"E MK879286 MK879423

Maximum likelihood (ML), Bayesian Inference (BI), and maximum parsimony (MP) approaches were used to reconstruct the phylogenetic relationships. The sequence data set was initially divided into four partitions for the nucleotide model-based ML and BI approaches: 1) 1st codon positions of cox1, 2) 2nd codon positions of cox1, 3) 3rd codon positions of the cox1, and 4) the 16S. To select an appropriate partitioning scheme and evolutionary models the sequence data set was analysed with PARTITIONFINDER v. 1.1.1 (Lanfear et al. 2012) conducting an exhaustive search and allowing for separate estimation of branch lengths for each partition using the Bayesian information criterion as recommended by Luo et al. (2010). Models to choose from were restricted to those available in MRBAYES v. 3.2.6 (Ronquist et al. 2012) as well as in GARLI v. 2.1 (Zwickl 2006). As best-fit partitioning scheme, the PARTITIONFINDER analysis suggested to combine the 1st and 2nd codon positions of cox1 and the 16S sequences together in one partition (GTR + G model) and the 3rd codon positions of cox1 in a second partition (HKY + G model).

The BI analysis was performed using MRBAYES v. 3.2.6. Metropolis-coupled Monte Carlo Markov chain (MC3) searches in MRBAYES were run with four chains in two separate runs for 50,000,000 generations with default priors, trees and parameters sampled every 1000 generations under default heating using the best-fit model as suggested by PARTITIONFINDER. Diagnostic tools in MRBAYES, including Estimated Sample Size (ESS) values > 200, were used to ensure that the MC3 searches had reached stationarity and convergence. The first 5,000,000 generations of each run were discarded as burn-in.

Heuristic ML analysis was performed with GARLI using the best-fit models as suggested by PARTITIONFINDER. Support values were computed by bootstrapping (BS) with 1,000 replicates.

Heuristic MP searches were carried out with PAUP v. 4.0b10 (Swofford 2002) using 100 random-addition-sequence replicates and TBR branch swapping. Support values were computed by bootstrapping with 1,000 replications.

Alternative phylogenetic hypotheses were tested using the approximately unbiased (AU) test (Shimodeira 2002) as implemented in the program CONSEL (Shimodeira and Hasegawa 2001). Information on vouchers and GENBANK accession numbers are listed in Table 1.

Results

Biometric analyses

The first two principal components (PCA 1 and PCA 2) account for > 95% of the cumulative variation in shell parameters. The plot of PCA 1 vs PCA 2 (Fig. 4A) shows that the clusters of specimens that were grouped according to geographic origin widely overlap. Especially the clusters of specimens from Thailand and Indonesia (corresponding to mitochondrial Clades A and B, Fig. 4) and the clusters of specimens from Australia (corresponding to mitochondrial Clade C, Fig. 4) also widely overlap. The Kruskal–Wallis rank sum tests were significant for PCA 1 (p < 5.0 × 10−6) and PCA 2 (p < 2.0 × 10−16), i.e., at least one group stochastically dominates one other group in each of the tests. Dunn’s test for PCA 1 found significant differences between the groups including samples from Indonesia and Australia (p < 0.0001) as well as between the groups including samples from Australia and Thailand (p < 0.0075), respectively, but not for pairwise comparisons of the other groups (Fig. 4B). Dunn’s test for PCA 2 found significant differences between the following groups: Indonesia vs Australia (p < 0.0031), Indonesia vs Thailand (p < 0.0001), Australia vs Thailand (p < 0.0007), Australia vs India/Sri Lanka (p < 0.0004), and Thailand vs India/Sri Lanka (p < 0.0001), but not for Indonesia vs India/Sri Lanka (Fig. 4C). However, both for PCA 1 and PCA 2 the comparison of ranges shows that the ranges of all pairs of geographic groups overlap and therefore do not allow a diagnostic separation of these groups on the basis of the biometric data. The included type specimens of the nominal taxa M. rudis and M. microstoma fall within the convex hull spanned by specimens sampled from Thailand, Indonesia, Australia, India, and Sri Lanka in the PCA 1 vs PCA 2 plot; only the holotype of the nominal taxon M. aspera lies outside this area (Fig. 4A), although closely resembling the examined syntypes of M. rudis and M. microstoma with respect to shell sculpture and overall shape.

Figure 4. 

Results of the analysis of biometric data of “Thiaraaspera (Lesson, 1831) specimens from Australia (yellow), Indonesia (green), Thailand (red) and India/Sri Lanka (blue) and type material of Melania aspera Lesson, 1831 (holotype, triangle), Melania rudis Lea & Lea, 1851 (syntype, square) and Melania microstoma Lea & Lea, 1851 (syntype, diamond). A. Scatter plot of the first two axes of the principal component analysis (PCA) of biometric data. Coloured lines indicate the outline of the convex hull for each geographic group; B, C. Boxplots of PCA 1 (B) and PCA 2 (C); bars above the box plots indicate significant differences of groups resulting from testing with Dunn’s test.

Figure 5. 

Bayesian 50% majority-rule consensus tree based on partial sequences mitochondrial cytochrome c oxidase subunit 1 (cox1) and 16S rRNA (16S) genes. Support values at nodes refer to Bayesian posterior probabilities (left), Maximum Likelihood (middle) and Maximum Parsimony (right) bootstrap values. AUS: Australia, IDN: Indonesia, THA: Thailand. Numbers at tips refer to DNA vouchers in the collection of the ZMB, see also Table 1.

Phylogenetic analyses

A clade including Thiara amarula (Linnaeus, 1758) (the type species of Thiara Röding, 1798), T. winteri (Busch, 1842) in Philippi (1842–1844), T. cf. winteri from Bali, and the specimens identified as Thiara aspera from Thailand, Indonesia, and Australia as well as “Stenomelaniadenisoniensis (Brot, 1877) in Brot (1874–1879) was recovered in all three analyses (BI: 1.00, BS (ML): 92, BS (MP): 96). However, Thiara is paraphyletic with respect to “S.denisoniensis. Thiara amarula grouped together with T. winteri and T. cf. winteri from Bali in a clade (BI: 1.00, BS (ML): 97, BS (MP): 93). A sister group relationship of T. amarula and T. winteri was recovered in the BI and ML analyses (BI: 0.99, BS (ML): 83) but not in the MP analysis (BS (MP): < 50). Within this clade, the clades including specimens of T. amarula, T. winteri, and T. cf. winteri from Bali, respectively, were supported (BI: 1.00, BS (ML): 96–100, BS (MP): 100). The clade containing T. amarula, T. winteri, and T. cf. winteri was recovered as the sister group of a clade containing “S.denisoniensis and specimens from Thailand, Australia, and Indonesia (including also a single specimen from Bali) assigned to the T. aspera on basis of conchological similarity with rather high support (BI: 1.00, BS (ML): 92, BS (MP): 96). Within this clade, “S.denisoniensis was recovered as the sister group (BI: 1.00, BS (ML): 94, BS (MP): 90) of T. aspera, which in turn formed a rather well-supported clade (BI: 1.00, BS (ML): 90, BS (MP): 79). Thiara aspera specimens from Australia (Clade C) formed a maximally supported clade. The T. aspera specimens from Thailand grouped together with a single individual from Bali in a well-supported clade (Clade A; BI: 1.00, BS (ML): 92, BS (MP): 100), which was sister to another supported clade (Clade B; BI: 1.00, BS (ML): 77, BS (MP): 100) that included T. aspera specimens from Sulawesi.

The included specimens of Mieniplotia scabra (Müller, 1774) formed the sister clade of a specimen of Melanoides tuberculata from India in the BI analysis, albeit without support. To test alternative phylogenetic hypotheses, we conducted four AU tests: 1) the monophyly of M. scabra (p = 0.252) and 2) the monophyly of T. winteri plus the T. cf. winteri specimens from Bali (p = 0.156) could not be rejected, whereas 3) the monophyly of T. aspera, T. winteri and T. cf. winteri (p < 0.001) and 4) the monophyly of Thiara excl. “S.denisoniensis (p = 0.033) but including the T. aspera specimens was rejected at a confidence level of α = 0.05.

Systematic account

Thiaridae Gill, 1871 (1823)

Thiara Röding, 1798

Vesica Humphrey, 1797: 58 [unavailable, published in a work rejected for nomenclatural purposes, see International Commission on Zoological Nomenclature 1912: 116–117; among the mentioned species is Vesica thiara Humphrey, 1797 (unavailable) = Helix amarula Linnaeus, 1758].

Thiara Röding, 1798: 109 [type species: Helix amarula Linnaeus, 1758, by subsequent designation of Herrmannsen 1849 in Herrmannsen 1847–1849: 576].

Melania Lamarck, 1799: 75 [type species: Helix amarula Linnaeus, 1758, by monotypy].

Melanigenus Renier, 1807: pl. 8 [unavailable, published in a work rejected for nomenclatural purposes, see International Commission on Zoological Nomenclature 1956: 290].

Melas Montfort, 1810: 322–324 [unjustified emendation of Melania Lamarck, 1799].

Melanidia Rafinesque, 1815: 144 [unjustified emendation of Melania Lamarck, 1799].

Melanea – Sowerby 1818 in Sowerby 1818–1822: 33 [incorrect subsequent spelling of Melania Lamarck, 1799].

? Spirilla Gray, 1824: 254 [unavailable, published in synonymy; mentioned as Spirilla spinosa (quoting a label or note attributed to G. Humphrey as “Spirilla spinosa, freshwater spiral spined shell, from Admirality Island, New Guinea”) under Melania setosa Swainson, 1824 (= Thiara cancellata Röding, 1798, see Swainson 1824: 13–15 and Wilkins 1957: 167–169) and as being conspecific with the nomenclaturally unavailable Buccinum aculeatum Lister, 1692: pl. 1055, fig. 8. Mentioned as a synonym by Férussac 1824: 318, Gray 1825: 524, Oken 1833: 133, Gray 1847: 152, Wilkins 1957: 167 as well as by Agassiz 1842: 84, Agassiz 1847: 348 and Herrmannsen 1848 in Herrmannsen 1847–1849: 491 in nomenclators, with the name attributed to Humphrey 1797 (where it could not be found). Used by Favre 1869: 79 (attributing the name to G. Humphrey but without reference to the work of Gray 1824) for a subgenus of Fusus Bruguière 1789 in Bruguière 1789–1792 and in a very different meaning from that of Gray 1824 and therefore not regarded here as having been made available from that work].

Spirella Oken 1833: 61 [incorrect subsequent spelling of the unavailable Spirilla Gray, 1824].

Melacantha Swainson, 1840: 341 [type species: Helix amarula Linnaeus, 1758 by subsequent designation of Herrmannsen 1849 in Herrmannsen 1847–1849: 26].

Thaira Gray 1840: 148 [incorrect subsequent spelling of Thiara Röding, 1798].

Amarula Sowerby, 1842: 61 [type species: Helix amarula Linnaeus, 1758, by monotypy].

Melanium – Busch 1842 in Philippi 1842–1845: 4 [incorrect subsequent spelling of Melania Lamarck, 1799].

Tiara Gray 1847: 152 [incorrect subsequent spelling of Thiara Röding, 1798].

Thaera Agassiz 1847: 367 [unavailable, emendation for Thaira as used by Gray 1840: 148 proposed in synonymy in a nomenclator].

Lithoparches Gistel, 1848: ix [nom. nov. pro Melania Lamarck, 1799; type species: Helix amarula Linnaeus, 1758, by typification of the replaced name].

Hydrognoma Gistel, 1848: 169 [nom. nov. pro Melania Lamarck, 1799, type species: Helix amarula Linnaeus, 1758, by typefication of the replaced name].

Tiaropsis Brot, 1871: 298 [non Agassiz 1849: 289–298; type species: Melania winteri Busch, 1842 in Philippi 1842–1844: Melania, 1, pl. 1 figs 1, 2 by subsequent designation of Brot 1874 in Brot 1874–1879: 7].

Cerithomelania Moore, 1899: 233–234 [type species: Helix amarula Linnaeus, 1758 by original designation].

? Ripalania Iredale, 1943: 209 [type species: Melania queenslandica Smith, 1882 by monotypy].

? Setaeara Morrison, 1952: 8 [type species: Thiara cancellata Röding, 1798 by original designation].

Remarks

Many names have been proposed for the group of Thiaridae that is currently regarded as representing Thiara Röding, 1798. Several of these names are objective junior synonyms of Thiara having the same type species (Helix amarula Linnaeus, 1758), and several others are nomenclaturally unavailable. A few, like Ripalania Iredale, 1943 or Setaeara Morrison, 1952, may actually be synonyms of Thiara. However, those hypotheses should be further tested using molecular genetic approaches. Therefore, these nominal genera were only tentatively included in the synonymy of Thiara.

“Thiara” aspera (Lesson, 1831)

Figs 1, 6, 7

Melania aspera Lesson, 1831 in Lesson (1830–1831: 357–358) [type locality: “La Nouvelle-Guinée” (= New Guinea), restricted to Manokwari by Glaubrecht and Podlacha (2010)].

Melania rudis Lea & Lea, 1851: 186 [type locality: ‘Amboyna’ (= Ambon)].

Melania microstoma Lea & Lea, 1851: 186 [type locality: mountain streams, isle of Negros, Philippines].

? Melania armillata Lea & Lea, 1851: 195–196 [type locality: India].

? Melania broti Reeve, 1859 in Reeve (1859–1861: pl. 22 fig. 160) [type locality: Ceylon (= Sri Lanka)].

? Melania hybrida Reeve, 1859 in Reeve (1859–1861: pl. 13 fig. 163) [type locality: not given].

? Melania chocolatum Brot, 1860: 256–257, pl. 16, fig 2 [type locality: “Ceylon” (= Sri Lanka)].

? Melania (Tiaropsis) rudis var. spinosa Brot, 1877 in Brot (1874–1879: 306) [type locality: not given, see also Brot (1868: 33, pl. 1, fig. 7)].

? Melania (Tiaropsis) drilliiformis Martens, 1897: 305 [nomen nudum].

? Melania fortitudinis Fulton, 1904: 51–52, pl. 4, fig. 3 [type locality: “Soekaboemi, Java” (= Sukabumi, Java)].

? Melania rudis var. cylindrica Schepman, 1915: 27 [type locality: West Ceram, Kairatu (= West Seram Island, Kairatu)].

Diagnosis

Thiarid with a turreted, subcylindrical to elongate-ovoid, strongly ornamented high-spired shell with usually rather flattened whorls and a narrowly pyriform aperture that at most reaches half the total shell height, but usually less. Ornamentation of the shell consisting of sinuous axial ribs that usually reach to the base of the body whorl and spiral chords that form nodes where they intersect the ribs; spiral chords usually present on the entire whorl but strongest at the base of body whorl.

Remarks

The examined type specimens of M. aspera, M. rudis, and M. microstoma correspond well to each other in overall shell shape and sculpture and are here regarded as conspecific because of this. As already noted by Brot (1874–1879: 307) and Glaubrecht and Podlacha (2010: 200), the name Melania aspera Lesson, 1830 has priority over the somewhat more frequently used name Melania rudis (e.g., van Benthem Jutting 1937, 1956; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Patil and Talmale 2011 as T. rudis). The holotype of Melania aspera is unusual in possessing a very small aperture in relation to overall shell height, possibly explaining its isolated position in the PCA 1 vs PCA 2 scatter plot (Fig. 4A). The nominal taxa M. armillata, M. broti, and M. chocolatum described from India or Sri Lanka were regarded by Brot (1874–1879) as closely related to M. rudis and are here tentatively synonymised with M. aspera, largely following the views of Rensch (1934) and van Benthem Jutting (1937, 1956) who synonymised these taxa with M. rudis. According to Brot (1874–1879: 307–308) Melania hybrida is based on a teratological specimen with an unusual aperture formation and is here tentatively synonymised with M. aspera. The nominal taxon Melania (Tiaropsis) rudis var. spinosa Brot, 1877 is an individual variation of M. aspera with somewhat longer shoulder spines. The original figures and descriptions of Melania fortitudinis and M. (Tiaropsis) rudis var. cylindrica from Java and West Seram Island also correspond well with the holotype of M. aspera and are herein treated as synonyms of the former.

Type material examined

Holotype of Melania aspera Lesson, 1831, MNHN 21098, “La Nouvelle-Guinée”; syntype of Melania rudis Lea & Lea, 1851, USNM 119778, “Amboyna”; syntype of Melania microstoma Lea & Lea, 1851, USNM 119722, ‘mountain streams, isle of Negros, Philippines’.

Additional material examined

(w: ethanol preserved samples). India: Kolkata, ZMB 107002. Sri Lanka: Colombo, ZMB 107003. Thailand: Samut Sakhon Province, Klong Don Ko, SUT 0311020, ZMB 127535, SUT 0311044, SUT 0311053, ZMB 127534, w; Nakhon Pathom province, Pond in Silpakorn University campus, SUT 0312069, SUT 0312070 = ZMB 127536, w. Indonesia: Bali: South Bali, Yehembang River, ZMB 191279, ZMB 191279a, w, South Bali, at Yehembang, ZMB 106472, w; east of Mendaya, stream southwest of Gumicik, ZMB 191488; Sulawesi: South Sulawesi, Kalena catchment, Angkona river, ZMB 192751, w; southeast Sulawesi, Pohara river, at Pohara, road Kendair to Kolaka, ZMB 191261, w; southeast Sulawesi, Simbune river, 1 km northeast of Raterate, road Kendari to Kolaka, ZMB 191262, ZMB 191262a, w; southeast Sulawesi, stream at Tembeeha, road Tirobus to Kendari, ZMB 191278, w; central Sulawesi, Banggai Islands, Peleng Island, West Peninsula, Tataban river, ZMB 107378, w, ZMB 107377,w. Australia: Northern Territory: Berry Springs, ZMB 106704, w, ZMB 106599a, w, ZMB 127616, w; Wabalaar, Roper River, ZMB 107617, w, ZMB 107614, w, ZMB 127645, w; Salt creek, ZMB 127619, w, ZMB 127636, w, ZMB 127637, w; Roper Bar, ZMB 127620, w; Queensland: O’Shanassy, ZMB 107280.

Shell

Turreted, subcylindrical to elongate-ovoid, corneous to dark brown, with up to nine whorls (the early whorls usually eroded) (Fig. 1; for juvenile shells, see Fig. 6). Whorls rather flat to convex, separated by a slightly impressed to distinctly impressed, undulating suture. Whorls slightly constricted below the suture, ornamented with sinuous ribs and spiral chords that usually form nodules at their intersections. Radial sculpture usually strongest on the upper half of the whorls, with the nodules at the shoulder of the whorls usually largest, sometimes forming spines. Towards the lower part of the body whorl the spiral sculpture often becomes the dominant sculptural element, forming distinct parallel chords. Aperture pyriform, angled in its upper part and rather narrow, wider at the base and appearing truncated in frontal view. Columella thickened, almost straight to curved, abruptly terminating basally. Shell size H = 7.6–48.0 mm, W = 3.1–22.0 mm (Table 2).

Figure 6. 

Juvenile and embryonic shells of “Thiaraaspera (Lesson, 1831), SUT 0311020, Samut Sakhon Province, Klong Don Ko. A. Lateral view; B. Apical whorls, lateral, C. Apical view. D. Details of the protoconch. Scale bars: 450 µm (A); 250 µm (B); 200 µm (C); 100 µm (D).

Shell parameters of “Thiaraaspera (Lesson, 1831) specimens from Thailand, Indonesia and Australia, with min./max. values, mean, standard deviation (SD), and number of whorls.

Voucher Country, region n Measurements (mm) NW
H W AL AW BW
USNM 119778 Indonesia, Ambon Island 1 23.7 9.7 9.8 5.1 15.5 4
USNM 119722 Philippines, Negros Island 1 20.3 7.7 7.1 3.3 12.3 6
MNHN 21098 Indonesia, West Papua 1 25.0 7.8 6.9 3.5 12.6 7
GSUBg 14265 Indonesia, Java 1 48.0 22.0 22.0 10.0 28.1 7
ZMB 107002 India, Calcutta 1 17.4 7.5 5.7 2.5 12.0 3
ZMB 107003 Sri Lanka, Colombo 5 Range 13.3–16.6 5.3–6.6 4.3–5.3 2.3–2.5 9.0–11.3 4–5
Mean 14.3 5.9 4.8 2.4 9.8
SD 1.2 0.4 0.3 0.1 0.8
SUT 0311053 Thailand, Samut Sakhon 30 Range 7.6–12.8 3.1–5.5 2.8–6.2 1.6–3.4 4.4–7.8 4–7
Mean 9.5 3.9 4.0 2.3 5.7
SD 1.1 0.6 0.6 0.4 0.9
SUT 0311020 Thailand, Samut Sakhon 52 Range 14.1–24.3 7.1–11.1 7.0–11.1 3.2–5.4 9.7–16.2 6–7
Mean 17.7 8.7 8.7 4.2 12.0
SD 2.5 1.0 1.0 0.5 1.5
SUT 0311044 Thailand, Samut Sakhon 1 22.9 10.9 10.5 4.8 15.2 6
SUT 0312070 Thailand, Nakhon Pathom 12 Range 14.4–19.8 5.3–7.9 5.7–6.8 2.4–4.2 7.6–12.0 5–8
Mean 17.1 6.8 7.0 3.4 9.9
SD 1.5 0.7 0.9 0.5 1.2
ZMB 191488 Indonesia, Bali 2 Range 19.8–22.1 8.3–9.4 8.0–8.8 4.1–4.7 12.8–14.5 5
Mean 20.9 8.9 8.4 4.4 13.7
SD 1.1 0.6 0.4 0.3 0.9
ZMB 191278 Indonesia, Sulawesi 19 Range 14.3–18.2 6.2–7.8 6.1–8.9 3.1–3.9 9.3–12.3 4–5
Mean 16.1 6.7 7.2 3.6 10.4
SD 1.0 0.3 0.6 0.2 0.7
ZMB 107377 Indonesia, Sulawesi 10 Range 13.6–20.3 5.5–7.5 4.5–7.3 2.7–4.0 7.7–11.7 4–5
Mean 16.8 6.5 6.2 3.4 9.6
SD 2.1 0.7 0.8 0.4 1.3
ZMB 107378 Indonesia, Sulawesi 19 Range 12.9–19.6 5.2–7.1 5.1–7.8 2.6–3.9 7.7–11.2 4–5
Mean 16.9 6.3 6.3 3.2 9.9
SD 1.5 0.4 0.7 0.4 0.8
ZMB 191279 Indonesia, Bali 17 Range 16.8–27.0 7.0–9.9 7.9–12.0 3.3–5.1 11.0–17.2 4–6
Mean 22.7 8.6 9.9 4.3 14.5
SD 3.7 1.0 1.4 0.6 2.2
ZMB 106472 Indonesia, Bali 16 Range 17.9–22.8 6.4–9.3 7.0–10.6 3.1–5.0 11.4–16.2 4–7
Mean 20.1 7.5 8.5 3.8 12.7
SD 1.4 0.6 0.9 0.4 1.2
ZMB 127538 Indonesia, Bali 20 Range 18.4–30.4 8.6–13.7 9.3–14.9 4.4–7.4 12.8–20.7 4–6
Mean 24.8 11.0 11.8 5.6 16.8
SD 3.0 1.2 1.4 0.7 1.9
ZMB 191268 Indonesia, Sulawesi 5 Range 29.3–41.5 11.5–16.0 12.1–18.7 5.7–8.4 12.3–26.8 4–6
Mean 35.3 13.5 15.3 6.9 20.9
SD 4.9 2.0 2.5 1.1 5.3

Operculum

The operculum is typical for thiarids, oval and paucispiral, light to dark brown, and with the nucleus being excentric in the lower left corner.

Juvenile shell

The shells of the juveniles in the brood pouch had up to five whorls, with a maximum height of about 2.5 mm. The protoconch is smooth, with the radial and spiral sculpture developing on the first teleoconch whorls (Fig. 6). For measurements of the embryonic shell, see Table 3.

Measurements of parameters of the juvenile protoconch of “Thiaraaspera (Lesson, 1831) of specimens obtained from the brood pouch.

Voucher Country, region n Measurements (μm)
he we de
ZMB 127534 Thailand, Samut Sakhon 3 Range 48.0–63.2 96.0–120.0 312.7–395.7
Mean 54.0 106.7 352.8
ZMB 127535 Thailand, Samut Sakhon 2 Range 56.3–71.4 107.0–114.3 354.3–366.2
Mean 63.9 110.7 352.8
ZMB 191278 Indonesia, Sulawesi 2 Range 34.0–72.4 76.0–91.1 202.0–252.4
Mean 53.2 84.6 227.2
ZMB 191488 Indonesia, Bali 1 83.3 95.2 259.5

Radula

Taenioglossate (Fig. 7), resembling other thiarids. As in all thiarids the central tooth or rachidian is significantly wider than tall; all specimens have a central cusp flanked by three to six triangular denticles on both sides, resulting in up to 12 denticles and a typically 4–5/1/4–5 pattern at the upper cutting edge (Fig. 7A, C, E, Table 4). The laterals are equipped with three to six smaller denticles on the inner side, and three to six denticles outside from the large main cusps (Fig. 7A, C, E, Table 4). The marginal teeth are moderately long, spoon-shaped, with a varying number of 6–10 denticles (Fig. 7B, D, F, Table 4).

Figure 7. 

Radulae of “Thiaraaspera (Lesson, 1831) from Thailand. A, B. SUT 0312070, Nakhon Pathom province, pond at Silpakorn University campus; A. Central and lateral teeth; B. Marginal teeth; C, D. SUT 0311020, Samut Sakhon province, Klong Don Ko; C. Central and lateral teeth; D. Marginal teeth. E, F: SUT 0311053, Samut Sakhon Province, Klong Don Ko; E. Central and lateral teeth; F. Marginal teeth. Scale bars: 35 µm (A, E); 5 µm (B, F); 25 µm (C); 10 µm (D).

Variation of cusps on the radula teeth of “Thiaraaspera (Lesson, 1831) specimens.

Voucher Country, region n Marginal teeth Lateral teeth (left) Lateral teeth (right) Rachidian
SUT 0311053 Thailand, Samut Sakhon 4 6–8 3-1-3 3-1-3 4–5-1-4–5
SUT 0311020 Thailand, Samut Sakhon 4 6–8 3-1-3 3-1-3 4–5-1-4–5
SUT 0312070 Thailand, Nakhon Pathom 2 7–8 5-1-5 4-1-4 4-1-4
SUT 0312069 Thailand, Nakhon Pathom 2 9–10 3-1-3 3-1-3 3-1-3
ZMB 191278 Indonesia, Sulawesi 2 7–8 3-1-3 3-1-3 4-1-4
ZMB 191488 Indonesia, Bali 1 10 6-1-6 6-1-6 6-1-5
ZMB 191279 Indonesia, Bali 3 6–7 4-1-4 4-1-4 4–5-1-4–5
ZMB 106472 Indonesia, Bali 2 6–7 3-1-3 3-1-3 4-1-4

Reproductive strategy

The results of the analysis of brood pouch content are summarised in Figure 2. Juveniles of up to 2 mm (rarely also larger) were found in the populations from Thailand, Indonesia (Bali and Sulawesi) and Australia (Northern Territory) suggesting an euviviparous reproductive strategy for “T.aspera, i.e, the taxon was found to give birth to crawling and shelled juveniles in accordance with the definitions in Glaubrecht et al. (2009). In a few populations in Thailand and on Bali, gravid females with only early embryos, i.e., veliger larvae in the brood pouch were found.

Distribution

Thiaraaspera as here understood is a widespread species, with records from Sri Lanka and India (Subba Rao 1989), Myanmar and Cambodia (van Benthem Jutting 1956), Indonesia (Rensch 1934; van Benthem Jutting 1956), and the Philippines (Lea and Lea 1851; van Benthem Jutting 1956). As our results indicate, the taxon is also present in Thailand and northern Australia, from where it was not previously reported (Fig. 2).

Discussion

The results of our phylogenetic analyses show that the nominal taxon Melania winteri Busch, 1842 is closely related to Thiara amarula and can be classified with the same genus. However, the nominal species Melania aspera Lesson, 1830 (= Melania rudis Lea & Lea, 1851), which has often been classified as a member of Thiara (e.g., van Benthem Jutting 1937, 1956; Subba Rao 1989; Ramakrishna and Dey 2007; Budha 2010; Patil and Talmale 2011 under the name T. rudis) cannot be included within that genus on the basis of our data without broadening the concept of Thiara to an extent that it encompasses almost the entire conchological diversity of Thiaridae because “Stenomelaniadenisoniensis Brot, 1877, which is conchologically similar to Stenomelania Fischer, 1885 or Melanoides Olivier, 1804, clusters within Thiara s. lat. in our phylogenetic analyses and an approximately unbiased test rejected the monophyly of T. amarula, T. winteri, T. cf. winteri, and “T”. aspera, i.e., excluding “Stenomelaniadenisoniensis. Pending a phylogenetic analysis of the entire family, we here retain the species in Thiara, but indicate the tentative placement by quotation marks.

Our phylogenetic analyses further show that “T.aspera exhibits little genetic variation throughout the Indo-Malayan Archipelago and the Southeast Asian mainland, although populations vary considerably with regard to shell shape, and especially sculpture, confirming previous surveys on thiarid species, which showed also an extraordinary plasticity of the shell (Glaubrecht et al. 2009). We here report the presence of “T.aspera in Thailand for the first time, albeit in anthropogenic habitats. Previous surveys of the Thai freshwater snail fauna, e.g., by Brandt (1974) did not record the species. Thus, as his years-long surveys were exhaustive it is safe to assume that the species is probably introduced but additional surveys should be carried out to clarify whether the taxon also occurs in natural habitats in this country and was only overlooked in the past.

We also report “T.aspera here for Australia for the first time, where the taxon was found in natural habitats in the Northern Territory and in north-western Queensland. The populations from Australia were found to be somewhat differentiated genetically from the remaining specimens of “T.aspera from Thailand and Indonesia included in the phylogenetic analyses and also slightly differ conchologically, i.e., the spiral sculpture almost disappears on the upper half of the teleoconch whorls. Further analysis should therefore confirm whether these differences are constant and would allow a taxonomic separation of the Australian populations.

Unfortunately, no samples could be included in the phylogenetic analyses from either India or Sri Lanka. However, as the examined material closely resembles the holotype of Melania aspera in shell characters (although this specimen is exceptional because of its very small aperture in relation to total shell height which may explain its isolate position in Fig. 4A), the populations from these two countries are here regarded as belonging to the species. Therefore, “T.aspera has to be considered as a widespread species, ranging from India and Sri Lanka across the Southeast Asian mainland and islands into Australia (Fig. 2).

At present, our data on “T.aspera do not allow to assess whether the observed differences of juvenile stages in the brood pouch of the female indicate differences in the reproductive strategy, or rather individual or seasonal variations. The close phylogenetic relationships among these populations (Fig. 5), however, let the latter two explanations appear more likely in our opinion. Therefore, we consider “T.aspera a euviviparous species, although it has to be stated that repeated periodic sampling would be necessary to resolve this issue conclusively.

Conclusions

These results highlight the need for a comprehensive revision of the genus-group systematics of Thiaridae as a whole. However, mitochondrial DNA markers are fraud with difficulties in some freshwater cerithioideans (Köhler and Deein 2010; Whelan and Strong 2015; Köhler 2016) and probably also in Thiaridae. Likewise, there appears to be a confusing variability in shell and reproductive features in thiarids, which is in stark contrast to a conserved radular morphology as compared to some other cerithioidean families (Glaubrecht 1996). A stable system of the family, which ought to include the type species of all named genus-group taxa, can be expected to emerge only after phylogenetic analyses based on suitable molecular markers and/or detailed morphological data become available. A stable system of the family then could serve as a basis for a better understanding of the evolutionary systematics and phylogeography of the group.

Acknowledgements

We thank Thomas von Rintelen and Christine Zorn for access to the collection housed at the Museum für Naturkunde, Berlin. We are indebted to the German Academic Exchange Service DAAD for grants in support of research in Thailand and the Deutsche Forschungsgemeinschaft DFG for a grant (DFG GL 297/19-1) making research in Australia possible. We are also indebted to the support fund of the Faculty of Science, Silpakorn University, Thailand and the Thailand Research Fund (The Royal Golden Jubilee Ph.D. Programme PHD/0195/2551) for funding. We thank Vince Kessner (Adelaide River) and Richard Willan (Darwin) for helping with the fieldwork and handling of collections in Australia. We also thank Frank Köhler (Sydney) and Zoltán Fehér (Budapest) for constructive comments on a draft version of the manuscript.

References

  • Adams H, Adams A (1854) The genera of Recent Mollusca; arranged according to their organization. London, van Voorst. Vol. 1, Part 10, Van Voorst, London, 289–320. [pls 37–40]
  • Agassiz L (1842) Nomina systematica generum molluscorum, tam viventium quam fossilium, secundum ordinem alphabeticum disposita, adjectis auctoribus, libris in quibus reperiuntur, anno editionis, etymologia et familiis ad quas pertinent. In: Agassiz L (Ed.) Nomenclator zoologicus, continens nomina systematica generum animalium tam viventium quam fossilium. Secundum ordinem alphabeticum disposita, adjectis auctoribus, libris, in quibus reperiuntur, anno editionis, etymologia et familis, ad quas pertinent, in singulis classibus. Jent & Gassmann, Soloduri, 1–98. https://doi.org/10.5962/bhl.title.12420
  • Agassiz L (1847) Nomenclatoris zoologici index universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tam viventium quam fossilium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nec non variis adnotationibus et emendationibus. Jent & Gassmann, Soloduri, 393 pp. https://doi.org/10.5962/bhl.title.1819
  • Agassiz L (1849) On the naked-eyed Medusae of the shores of Massachusetts in their perfect state of development. Contributions to the natural history of the Acalephae of North America Part 1. Memoirs of the American Academy of Arts and Sciences (New Series) 4: 221–312. [pls 1–8] https://doi.org/10.2307/25058163
  • Brandt RAM (1974) The non-marine aquatic Mollusca of Thailand. Archiv für Molluskenkunde 105: 1–423.
  • Brot A (1860) Description de nouvelles espèces de Mélanies. Revue et Magazin de Zoologie pure et appliquée 12: 254–267. [pls 16–17]
  • Brot A (1868) Matériaux pour servir à l’étude de la famille des mélaniens. Additions et corrections au catalogue systématique des espèces qui composent la famille des mélaniens. Georg, Genève, 64 pp. [3 pls] https://doi.org/10.5962/bhl.title.11249
  • Brot A (1871) Catalogue of the Recent species of the family Melanidæ. American Journal of Conchology 6: 271–325.
  • Brot A (1874–1879) Die Melaniaceen (Melanidae) in Abbildungen nach der Natur mit Beschreibungen. Systematisches Conchylien-Cabinet von Martini und Chemnitz 1(24): 1–488. [pls 1–49] https://doi.org/10.5962/bhl.title.124284
  • Brown DS (1994) Freshwater Snails of Africa and their Medical Importance (2nd edn). Taylor & Francis, London & Bristol, 609 pp.
  • Campbell DC (2019) Semisulcospiridae Morrison, 1952. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the World – A Distribution Atlas. Johns Hopkins University Press: Baltimore, 81–85.
  • Children JG (1823) Lamarck’s genera of shells. Quaterly Journal of Science, Literature and the Arts 15: 216–258.
  • Dechruska W, Krailas D, Glaubrecht M (2013) Evaluating the status and identity of “Melaniajugicostis Hanley & Theobald, 1876 – an enigmatic thiarid gastropod in Thailand (Caenogastropoda, Cerithioidea). Zoosystematics and Evolution 89: 293–310. https://doi.org/10.1002/zoos.201300015
  • Favre E (1869) Description des mollusques fossiles de la Craie des environs de Lemberg en Galicie. Georg, Genève & Bale, 187 pp. [13 pls]
  • Férussac AEJPJF d’Audebard de (1824) Sur la structure de la Melania setosa, par M. J. E. Gray. (Zoolog. Journ., no. 2, 1824, p. 253, pl. 8, f. 6 à 8.). Bulletin des Sciences naturelles et de Géologie 3: 318–319.
  • Fischer P (1885) Manuel de Conchyliologie et de Paléontologie Conchyliologique. Fascicule 8. Savy, Paris, 689–784.
  • Fischer P, Crosse H (1891) Etudes sur les mollusques terrestres et fluviatile du Mexique et du Guatemala. Mission scientifique au Mexique et dans l’Amérique Centrale. Recherches zoologiques. Partie 7. Volume 2. Livraison 12. Imprimerie nationale, Paris, 33–39. [pls 49–52]
  • Fischer P, Crosse H (1892) Etudes sur les mollusques terrestres et fluviatile du Mexique et du Guatemala. Mission scientifique au Mexique et dans l’Amérique Centrale. Recherches zoologiques. Partie 7. Volume 2. Livraison 13. Imprimerie nationale, Paris, 40–49. [pls 53–54]
  • Fleming J (1822) The Philosophy of Zoology, a general view of the structure, functions and classification of animals. Volume 2, Constable & Co., Edinburgh, 618 pp. https://doi.org/10.5962/bhl.title.24597
  • Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.
  • Fulton H (1904) On some species of Melania and Jullienia from Yunnan and Java. The Journal of Malacology 11: 51–52. [pl. 4]
  • Gill T (1871) Arrangement of the families of mollusks. Smithsonian Miscellaneous Collections 227: 1–49.
  • Gimnich F (2015) Molecular approaches to the assessment of biodiversity in limnic gastropods (Cerithioidea, Thiaridae) with perspectives on a Gondwanian origin. Dissertation, Humboldt Universität zu Berlin, Berlin.
  • Gistel J (1848) Naturgeschichte des Thierreichs für höhere Schulen. Hoffmann, Stuttgart, 216 pp. [32 pls]
  • Glaubrecht M (1993) Mapping the diversity: geographical distribution of the freshwater snail Melanopsis (Gastropoda: Cerithioidea: Melanopsidae) with focus on its systematics in the Mediterranean Basin. Mitteilungen aus dem Hamburger Zoologischen Museum und Institut 90: 41–97.
  • Glaubrecht M (1996) Evolutionsökologie und Systematik am Beispiel von Süß- und Brackwasserschnecken (Mollusca: Caenogastropoda: Cerithioidea): Ontogenese-Strategien, paläontologische Befunde und Historische Zoogeographie. Backhuys Publisher, Leiden, 499 pp. [25 pls]
  • Glaubrecht M (1999) Systematics and the evolution of vivparity in tropical freshwater gastropods (Cerithioidea: Thiaridae sensu lato) – an overview. Courier Forschungsinstitut Senckenberg 215: 91–96.
  • Glaubrecht M (2006) Independent evolution of reproductive modes in viviparous freshwater Cerithioidea (Gastropoda, Sorbeoconcha) – a brief review. Basteria 69(Supplement 3): 28–32.
  • Glaubrecht M (2009) On “Darwinian mysteries” or molluscs as models in evolutionary biology: from local speciation to global radiation. American Malacological Bulletin 27: 3–23. https://doi.org/10.4003/006.027.0202
  • Glaubrecht M (2010) Evolutionssystematik limnischer Gastropoden. Habilitationsschrift, Mathematisch-Naturwissenschaftliche Fakultät, Humboldt-Universität zu Berlin, Berlin.
  • Glaubrecht M (2011) Towards solving Darwin’s “mystery”: speciation and radiation in lacustrine and riverine freshwater gastropods. American Malacological Bulletin 29: 187–216. https://doi.org/10.4003/006.029.0211
  • Glaubrecht M, Brinkmann N, Pöppe J (2009) Diversity and disparity ‘down under’: Systematics, biogeography and reproductive modes of the ‘marsupial’ freshwater Thiaridae (Caenogastropoda, Cerithioidea) in Australia. Zoosystematics and Evolution 85: 199–275. https://doi.org/10.1002/zoos.200900004
  • Glaubrecht M, Köhler F (2004) Radiating in a river: systematics, molecular genetics and morphological differentiation of viviparous freshwater gastropods endemic to the Kaek River, central Thailand (Cerithioidea, Pachychilidae). Biological Journal of the Linnean Society 82: 275–311. https://doi.org/10.1111/j.1095-8312.2004.00361.x
  • Glaubrecht M, Neiber MT (2019a) Hemisinidae Fischer & Crosse, 1891. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the world – A Distribution Atlas. Johns Hopkins University Press, Baltimore, 51–55.
  • Glaubrecht M, Neiber MT (2019b) Thiaridae Gill, 1873 (1823). In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the world – A Distribution Atlas. Johns Hopkins University Press, Baltimore, 86–89.
  • Glaubrecht M, Podlacha K (2010) Freshwater gastropods from early voyages into the Indo-West Pacific: The ‘melaniids’ (Cerithioidea, Thiaridae) from the French ‘La Coquille’ circumnavigation, 1822–1825. Zoosystematics and Evolution 86: 185–211. https://doi.org/10.1002/zoos.201000002
  • Golubchik T, Wise MJ, Easteal S, Jermiin LS (2007) Mind the gaps: evidence of bias in estimates of multiple sequence alignments. Molecular Biology and Evolution 24: 2433–2442. https://doi.org/10.1093/molbev/msm176
  • Gray JE (1824) On the structure of Melania setosa. The Zoological Journal 1: 253–255. [pl. 8]
  • Gray JE (1825) Reply to Mr. Swainson on Neritina Corona and Melania setosa. The Zoological Journal 1: 523–526.
  • Gray JE (1840) Eastern zoological gallery. Synopsis of the Contents of the British Museum 42: 89–152.
  • Gray JE (1847) A list of the genera of recent Mollusca, their synonyma and types. Proceedings of the Zoological Society of London 15: 129–219.
  • Hanley SCT, Theobald W (1876) Conchologia Indica. Illustrations of the Land and Freshwater Shells of British India. L. Reeve & Co., London, 65 pp. [160 pls] https://doi.org/10.5962/bhl.title.96210
  • Herrmannsen AN (1847–1849) Indicis generum malacozoorum primordia. Nomina subgenerum, generum, familiarum, tribuum, ordinum, classium; adjectis auctoribus, temporibus, locis systematicis atque literariis, etymis, synonymis. Praetermittuntur Cirripedia, Tunicata et Rhizopoda. Vol. II. Fischer, Cassellis, 717 pp. https://doi.org/10.5962/bhl.title.10147
  • Holznagel WE (1998) A nondestructive method for cleaning gastropod radulae from frozen, alcohol-fixed, or dried material. American Malacological Bulletin 14: 181–183.
  • Holznagel WE, Lydeard C (2000) A molecular phylogeny of North American Pleuroceridae (Gastropoda: Cerithioidea) based on mitochondrial 16S rDNA sequences. Journal of Molluscan Studies 66: 233–257. https://doi.org/10.1093/mollus/66.2.233
  • Humphrey G (1797) Museum Calonnianum. Specification of the various Articles which compose the magnificent Museum of Natural History collected by M. de Calonne in France, and lately his Property: consisting of an Assemblage of the most beautiful and rare Subjects in Entomology, Conchology, Ornithology, Mineralogy, &c. [Humphrey], London, 84 pp.
  • International Commission on Zoological Nomenclature (1912) Opinions rendered by the International Commission on Zoological Nomenclature. Opinions 38 to 51. Smithsonian Institution Washington Publications 2060: 89–117.
  • International Commission on Zoological Nomenclature (1956) Opinion 427. Opinions and Declarations rendered by the International Commission on Zoological Nomenclature 14: 281–310.
  • Iredale T (1943) A basic list of the fresh water Mollusca of Australia. The Australian Zoologist 10: 188–230.
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 32: 772–780. https://doi.org/10.1093/molbev/mst010
  • Köhler F (2016) Rampant taxonomic incongruence in a mitochondrial phylogeny of Semisulcospira freshwater snails from Japan (Cerithioidea: Semisulcospiridae). Journal of Molluscan Studies 82: 268–281. https://doi.org/10.1093/mollus/eyv057
  • Köhler F, Deein G (2010) Hybridisation as potential source of incongruence in the morphological and mitochondrial diversity of a Thai freshwater gastropod (Pachychilidae, Brotia H. Adams, 1866). Zoosystematics and Evolution 86: 301–314. https://doi.org/10.1002/zoos.201000013
  • Köhler F, Glaubrecht M (2001) Toward a systematic revision of the Southeast Asian freshwater gastropod Brotia H. Adams, 1866 (Cerithioidea: Pachychilidae): an account of species from around the South China Sea. Journal of Molluscan Studies 67: 281–318. https://doi.org/10.1093/mollus/67.3.281
  • Köhler F, Glaubrecht M (2003) Morphology, reproductive biology and molecular genetics of ovoviviparous freshwater gastropods (Cerithioidea: Pachychilidae) from the Philippines, with description of a new genus Jagora. Zoologica Scripta 32: 35–59. https://doi.org/10.1046/j.1463-6409.2003.00100.x
  • Köhler F, Glaubrecht M (2006) A systematic revision of the Southeast Asian freshwater gastropod Brotia (Cerithioidea: Pachychilidae). Malacologia 48: 159–251.
  • Lamarck JBPA de Monet de (1799) Prodrome d’une nouvelle classification des coquilles, comprenant une rédaction appropriée des caractères génériques, et l’établissement d’un grand nombre de genres nouveaux. Mémoires de la Société d’Histoire Naturelle de Paris 1: 63–91. [1 table]
  • Lamarck JBPA de Monet de (1816) Tableau encyclopédique et méthodique des trois règnes de la nature. Vingt-troisième partie. Mollusques et polypes divers. Agasse, Paris, 391–488. [pls 1–98]
  • Lanfear R, Calcott B, Ho SYW, Guindon S (2012) PARTITIONFINDER: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29: 1695–1701. https://doi.org/10.1093/molbev/mss020
  • Lea I, Lea HC (1851) Description of a new genus of the family Melaniana, and of many new species of the genus Melania, chiefly collected by Hugh Cuming, Esq., during his zoological voyage in the East, and now first described. Proceedings of the Zoological Society of London 18: 179–197.
  • Lesson RP (1830–1831) Zoologie du voyage autour du monde, exécuté par ordre du roi, sur la corvette de sa majesté, La Coquille, pendant les années 1822, 1823, 1824 et 1825. Zoologie. Tome second. 1re partie. Arthus Bertrand, Paris, 471 pp.
  • Linnaeus C (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Salvius, Holmiæ, 824 pp. https://doi.org/10.5962/bhl.title.542
  • Lister M (1692) Appendix ad Historiæ Conchÿliorum librum. IV de buccinitis, ÿsue lapidibus, qui buccina omnigena ualdè referant. Authoris, Londini, 16 pp.
  • Luo A, Qiao H, Zhang Y, Shi W, Ho SYW, Xu W, Zhang A, Zhu C (2010) Performance of criteria for selecting evolutionary models in phylogenetics: a comprehensive study based on simulated datasets. BMC Evolutionary Biology 10: 242. https://doi.org/10.1186/1471-2148-10-242
  • Lydeard C, Holznagel WE, Glaubrecht M, Ponder WF (2002) Molecular phylogeny of a circum-global, diverse gastropod superfamily (Cerithioidea: Mollusca: Caenogastropoda): pushing the deepest phylogenetic limits of mitochondrial LSU rDNA sequences. Molecular Phylogenetics and Evolution 22: 399–406. https://doi.org/10.1006/mpev.2001.1072
  • Maaß N, Glaubrecht M (2012) Comparing the reproductive biology of three “marsupial”, eu-viviparous gastropods (Cerithioidea, Thiaridae) from drainages of Australia´s monsoonal north. Zoosystematics and Evolution 88: 293–315. https://doi.org/10.1002/zoos.201200023
  • Martens E von (1897) Süss- und Brackwasser Mollusken des Indischen Archipels. In: Weber M (Ed.) Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien. Vierter Band. Erstes Heft. Brill, Leiden, 1–331. [pls 1–12] https://doi.org/10.5962/bhl.title.46994
  • Montfort PD de (1810) Conchyliologie systématique, et classification méthodique des coquilles; offrant leurs figures, leur arrangement générique, leurs descriptions caractéristiques, leurs noms; ainsi que leur synonymie en plusieurs langues. Ouvrage destiné à faciliter l’étude des coquilles, ainsi que leur disposition dans les cabinets d’histoire naturelle. Coquilles univalves, non cloisonnées. Tome second. Schœll, Paris.
  • Moore JES (1899) On the divergent forms at present incorporated in the family Melaniidae. Proceedings of the Malacological Society of London 3: 230–234.
  • Morrison JPE (1952) World relations of the melanians. The American Malacological Union News Bulletin and Annual Reports 6: 6–9.
  • Müller OF (1774) Vermium terrestrium et fluviatilium, seu animalium infusoriorum, helminthicorum, et testaceorum, non marinorum, succincta historia. Volumen alterum. Heineck et Faber, Havniæ, Lipsiæ, 214 pp. https://doi.org/10.5962/bhl.title.12733
  • Neiber MT, Glaubrecht M (2019a) Unparalleled disjunction or unexpected relationships? Molecular phylogeny and biogeography of Melanopsidae (Caenogastropoda: Cerithioidea), with the description of a new family and a new genus from the ancient continent Zealandia. Cladisics. https://doi.org/10.1111/cla.12361
  • Neiber MT, Glaubrecht M (2019b) Melanopsidae H. Adams and A. Adams, 1854. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the world – An Distribution Atlas. Johns Hopkins University Press, Baltimore, 56–61.
  • Neiber MT, Glaubrecht M (2019c) Pachychilidae Fischer and Crosse, 1892. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the World – A Distribution Atlas. Johns Hopkins University Press, Baltimore, 62–67.
  • Neiber MT, Glaubrecht M (2019d) Paludomidae Stoliczka, 1868. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the World – A Distribution Atlas. Johns Hopkins University Press, Baltimore, 68–73.
  • Oken L (1833) Voyage autour du monde: executé par ordre du Roi sur la corvette: La Coquille pendant les années 1822–1825, publié par L. I. Duperrey, Capitain de frégate. Zoologie par Lesson. Bertrand, Paris, T. I. 26; 4. 360; II. 30; Atlas in Fol. Isis 17: 25–156.
  • Olivier GA (1804) Voyage dans l’Empire Othoman, l’Égypte et la Perse, fait par ordre du gouvernement, pendant les six premières années de la République. Tome second. Agasse, Paris, 466 pp. [pls 18–33]
  • Patil SG, Talmale SS (2011) Land and freshwater Mollusca. In: Venkatamaran K (Ed.) Fauna of Madhya Pradesh (including Chhattisgarh). State Fauna Series 15 (Part 3). Zoological Survey of India, Kolkata, 1–30.
  • Philippi RA (1842–1844) Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien, unter Mithülfe mehrerer deutscher Conchyliologen. Erster Band. Fischer, Cassel, 204 pp. [48 pls] https://doi.org/10.5962/bhl.title.10589
  • R Core Team (2016) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. https://www.R-project.org/
  • Ramakrishna, Dey A (2007) Handbook on Indian freshwater Mollusca. Zoological Survey of India, Kolkata, 399 pp.
  • Reeve LA (1859–1861) Monograph of the genus Melania. In: Reeve LA (Ed.) Conchologia Iconica; or, illustrations of the shells of molluscous animals. Vol. XII. Reeve, London. [pls 1–59]
  • Renier SA (1807) Tavole per servire alla classificazione e conoscenza degli animali. Tipografia del Seminario, Padova, 1 pp. [8 pls]
  • Rensch B (1934) Die Molluskenfauna der Kleinen Sunda-Inseln Bali, Lombok, Sumbawa, Flores und Sumba III. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 65: 389–422.
  • Rintelen T von, Wilson AB, Meyer A, Glaubrecht M (2004) Escalation and trophic specialization drive adaptive radiation of freshwater gastropods in ancient lakes on Sulawesi, Indonesia. Proceedings of the Royal Society of London B 271: 1541–1549. https://doi.org/10.1098/rspb.2004.2842
  • Röding PF (1798) Museum Boltenianum sive catalogus cimeliorum e tribus regnis naturæ quæ olim collegerat Joa. Fried Bolten, M. D. p. d. per XL. annos proto physicus Hamburgensis. Pars secunda continens conchylia sive testacea univalvia, bivalvia & multivalvia. Trapp, Hamburgi, 199 pp.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MRBAYES 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
  • Schepman MM (1892) Land- and freshwater shells collected by Dr. H. ten Kate in Soemba, Timor and other East-Indian Islands. Notes from the Leyden Museum 14: 145–160. [pl. 6]
  • Sowerby J (1818–1822) The Mineral Conchology of Great Britain; or coloured figures and descriptions of those remains of testaceous Animals or shells, which have been preserved at various times and depths in the earth. Vol. III. Arding & Merrett, London, 194 pp. [102 pls]
  • Strong EE, Colgan DJ, Healy JM, Lydeard C, Ponder WF, Glaubrecht M (2011) Phylogeny of the gastropod superfamily Cerithioidea using morphology and molecules. Zoological Journal of the Linnean Society 162: 43–89. https://doi.org/10.1111/j.1096-3642.2010.00670.x
  • Strong EE, Lydeard C (2019) Pleuroceridae P. Fischer, 1885. In: Lydeard C, Cummings K (Eds) Freshwater Mollusks of the world – A Distribution Atlas. Johns Hopkins University Press, Baltimore, 74–80.
  • Subba Rao NV (1989) Handbook freshwater Mollusca of India. Zoological Survey of India, Calcutta, 289 pp.
  • Swainson W (1824) Description of two new and remarkable fresh water shells: Melania setosa and Unio gigas. Quarterly Journal of Science, Literature and the Arts 17: 13–17.
  • Swainson W (1840) A Treatise on Malacology: or the natural classification of shells and shell-fish. Longman, Orme, Brown, Green & Longmans, London, 417 pp. https://doi.org/10.5962/bhl.title.8027
  • Swofford DL (2002) PAUP*. Phylogenetic Analysis Using Parsimony and other methods. Version 4.0b10. Sinauer Associates, Sunderland.
  • Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution 30: 2725–2729. https://doi.org/10.1093/molbev/mst197
  • van Benthem Jutting T (1937) Non marine mollusca from fossil horizons in Java with special reference to the Trinil fauna. Zoologische Mededelingen 20: 83–180. [pls 4–12]
  • van Benthem Jutting WSS (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian Archipelago. Treubia 23: 259–493.
  • Veeravechsukij N, Namchote S, Neiber MT, Glaubrecht M, Krailas D (2018a) Exploring the evolutionary potential of parasites: larval stages of pathogen digenic trematodes in their thiarid snail host Tarebia granifera in Thailand. Zoosystematics and Evolution 94: 425–460. https://doi.org/10.3897/zse.94.28793
  • Veeravechsukij N, Krailas D, Namchote S, Wiggering B, Neiber MT, Glaubrecht M (2018b) Molecular phylogeography and reproductive biology of the freshwater snail Tarebia granifera in Thailand and Timor (Cerithioidea, Thiaridae): morphological disparity versus genetic diversity. Zoosystematics and Evolution 94: 461–493. https://doi.org/10.3897/zse.94.28981
  • Whelan NV, Strong EE (2015) Morphology, molecules and taxonomy: extreme incongruence in pleurocerids (Gastropoda, Cerithioidea, Pleuroceridae). Zoologica Scripta 45: 62–87. https://doi.org/10.1111/zsc.12139
  • Wilkins GL (1957) The Cracherode shell collection. Bulletin of the British Museum (Natural History). Historical Series 1: 123–184. [pls 20–25]
  • Wilson AB, Glaubrecht M, Meyer A (2004) Ancient lakes as evolutionary reservoirs: evidence from the thalassoid gastropods of Lake Tanganyika. Proceedings of the Royal Society of London B: Biological Sciences 271: 529–536. https://doi.org/10.1098/rspb.2003.2624
  • Zwickl DJ (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. PhD Thesis, University of Texas at Austin, Austin.

1 * a question mark indicates a tentative synonymisation
2 * a question mark indicates a tentative synonymisation