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A glimpse in the dark? A first phylogenetic approach in a widespread freshwater snail from tropical Asia and northern Australia (Cerithioidea, Thiaridae)

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.


Introduction
Despite advances in the understanding of the family-level phylogeny of Cerithioidea Fleming, 1822, the taxonomical diversity in Thiaridae Gill, 1871Gill, (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, 2019dStrong and Lydeard 2019 and references therein) and recently the Neotropical Hemisinidae Fischer &Crosse, 1891 (Glaubrecht and, a more accurate circumscription of "core" Thiaridae began to emerge on the basis of molecular and/or morphological evidence (e.g., Glaubrecht 1993Glaubrecht , 1996Glaubrecht , 2011Holznagel 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 ecologi-cal 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(Glaubrecht , 1996Köhler and Glaubrecht 2001, 2003Glaubrecht 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 (1996Glaubrecht ( , 1999Glaubrecht ( , 2006Glaubrecht ( , 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. (2018aVeeravechsukij et al. ( , 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(Lesson -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(Schepman , 1915Rensch 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.
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. 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-bydot 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 Glob-al30-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 cal-Figure 1. Shells of "Thiara" aspera (Lesson, 1831 lipers (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.
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 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 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).
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) 1 st codon positions of cox1, 2) 2 nd codon positions of cox1, 3) 3 rd 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 PARTI-TIONFINDER 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 1 st and 2 nd codon positions of cox1 and the 16S sequences together in one partition (GTR + G model) and the 3 rd 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 (MC 3 ) 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 MC 3 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 PARTITION-FINDER. 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 GEN-BANK accession numbers are listed in Table 1.

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), Indo-nesia 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.

Systematic account
Thiaridae Gill, 1871Gill, (1823   . Results of the analysis of biometric data of "Thiara" aspera (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.  Iredale, 1943or 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.
? Melania rudis var. cylindrica Schepman, 1915: 27  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 (1874Brot ( -1879 and Glaubrecht and Podlacha (2010: 200), the name Melania aspera Lesson, 1830 has priority over the somewhat more fre-* a question mark indicates a tentative synonymisation quently used name Melania rudis (e.g., van Benthem Jutting 1937, 1956Subba 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 (1874Brot ( -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 (1874Brot ( -1879 Melania hybrida is based on a teratological specimen with an unusual aperture formation and is here tentatively synonymised with M. aspera.  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).
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.

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. "Thiara" aspera 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, 1956Subba 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 "Stenomelania" denisoniensis 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 "Stenomelania" denisoniensis. 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 ).
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.