Phylogenetic Position of African Punctoid Snails (Stylommatophora, Punctoidea, Trachycystinae)
Museum of New Zealand Te Papa Tongarewa, 169 Tory Street, Wellington 6011, New Zealand
Taxonomy 2022, 2(2), 227-235; https://doi.org/10.3390/taxonomy2020017
Submission received: 27 April 2022
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Revised: 31 May 2022
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Accepted: 31 May 2022
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Published: 2 June 2022
Abstract
:The punctoid land snail family Charopidae, as currently defined, is a paraphyletic assemblage of taxa with Gondwanan distribution. It is represented in Africa largely by the pinwheels (genus Trachycystis and allies) and afrodontas (genus Afrodonta and allies), as well as a few additional genera, such as Reticulapex, Pilula, and Helenoconcha. Herein, a Bayesian inference phylogenetic analysis (using four molecular markers) is conducted to test whether these taxa belong to the Charopidae and, if so, what their position is in the phylogenetic tree. It is concluded that Reticulapex and Pilula do not belong to the Punctoidea and are thus transferred to the Acavidae and Helicarionoidea, respectively. The pinwheels and afrodontas form a monophyletic group, the most basal branch of “Charopidae”, here classified as the subfamily Trachycystinae. It possibly represents an old southern African lineage potentially dating back to the split of Gondwana, while the remaining “Charopidae” and Punctidae can be found in Zealandia and Australia, and later, in the Americas and Europe. If further studies support the present findings, the elevation of Trachycystinae to the family level might be warranted. Finally, Flammoconchinae is also identified as a New Zealand subfamily of “Charopidae”.
1. Introduction
The land snail family Charopidae, as currently defined, has a Gondwanan distribution [1], being represented in Africa largely by two groups: the pinwheels and the afrodontas [2]. The pinwheel snails have discoid or low conical (helicoid) shells that lack apertural dentition and have been historically classified in the genus Trachycystis Pilsbry, 1893 (type species: Helix bisculpta Benson, 1851). Presently, the several previously accepted subgenera of Trachycystis are considered full genera [3,4]. The afrodontas, on the other hand, have smaller discoid shells bearing apertural dentition and have been historically classified in the genus Afrodonta Melvill & Ponsonby, 1908 (type species: Afrodonta bilamellaris Melvill & Ponsonby, 1908, by subsequent designation of [5]). Like Trachycystis above, Afrodonta has recently been split into several genera [6].
Both pinwheels and afrodontas were originally assigned to the family Endodontidae (e.g., [7,8,9,10,11]), but were later transferred to the Charopidae when that taxon was elevated to family level by Solem [11,12]. Both the Charopidae and Endodontidae belong to the superfamily Punctoidea, alongside the families Punctidae and Cystopeltidae [1]. Solem [12] was mainly interested in the Pacific Punctoidea and proposed—based solely on penial anatomy—that both Afrodonta and Trachycystis were closely related to Graeffedon Solem, 1983, from Samoa and Tonga (currently in Charopinae). Schileyko [13] established the family Trachycystidae to allocate Trachycystis, but subsequently [14] returned that genus to the Endodontidae within the subfamily Trachycystinae (Afrodonta was kept in subfamily Endodontinae).
Most subsequent authors (e.g., [15,16,17]), however, followed Solem’s classification. Hence, Trachycystinae has typically been considered synonymous with Charopinae [16,17], and sometimes includes taxa from places other than southern Africa, such as Saint Helena and South America (e.g., [4,12]).
In the molecular phylogenetic analysis of Punctoidea conducted by [1], there was a single African representative, belonging to the genus Chalcocystis Watson, 1934. Those authors showed that Chalcocystis did not belong to the Charopinae; rather, it was the most basal branch of the clade comprised of a paraphyletic Charopidae and the family Punctidae. They remarked that the subfamily Trachycystinae could have some biological reality, provided that the group was circumscribed solely to African taxa. Therefore, herein, that assertion is tested by including newly sequenced Trachycystinae species in the molecular framework of [1].
2. Material and Methods
2.1. Taxon Sampling
The present species selection contains 13 species (from nine genera) representing not only pinwheels and afrodontas, but also another two genera that have been included in the Trachycystinae [3,4,18]: Reticulapex Emberton & Pearce, 2000 and Pilula E. von Martens, 1898. Unfortunately, no suitable specimens of Helenoconcha Pilsbry, 1892 could be procured; this genus is endemic to Saint Helena and it is potentially related to Trachycystis senso latu, according to [12].
Furthermore, sequences from four genera of New Zealand Charopidae that are considered basal within the family (F. Brook, pers. comm. 2021) were also included: Cavellia Iredale, 1915, Flammoconcha Dell, 1952, Pseudallodiscus Climo, 1971, and Therasiella Powell, 1948. This was conducted to further put to test whether Trachycystinae is the most basal branch of Charopidae, as suggested by [1].
The full set of taxa used here can be seen in Table 1. Samples were obtained from the following collections: NMSA = Kwa-Zulu Natal Museum (Pietermaritzburg, South Africa); OZD = Department of Zoology, University of Otago (Dunedin, New Zealand); and UF = Florida Museum of Natural History (Gainesville, USA). DNA sequences of other Punctoidea were obtained from NCBI GenBank (from the study of Salvador et al., 2020; Table 2). The family-level classification of Punctoidea used here follows Salvador et al. (2020).
Based on the observed morphological features of the shell and soft body, it was suspected that Reticulapex and Pilula were not members of the Punctoidea. They seem to more closely resemble, respectively, members of the Acavidae and Helicarionoidea/Trochomorphoidea. DNA sequences of species belonging to those groups were obtained from GenBank (Table 2) and included in the analysis. The classification of Acavidae, Helicarionoidea, and Trochomorphoidea used here follows [18].
The outgroup for the phylogenetic analysis (see below) was composed of species from basal lineages within the Stylommatophora: Achatinidae, Rhytididae, and Streptaxidae. DNA sequences of members of those families were likewise obtained from GenBank (Table 2).
2.2. DNA Extraction, Amplification and Sequencing
The specimens used herein were fixed and preserved in ethanol (70% to 98%). Larger specimens had a small section of their foot clipped for DNA extraction, while very minute specimens were used completely. DNA extraction followed the standard protocol of the QIAGEN DNEasy® Blood & Tissue Kit, with a repetition of the final step to increase the yield.
The same markers used by [1] were adopted herein: (1) the barcoding fragment of the mitochondrial COI gene (primers LCO/HCO of [19]), which was circa 650 bp long; (2) the mitochondrial 16S rRNA gene (primers 16SarL/16SbrH of [20]), circa 450 bp; and (3) a continuous fragment of nuclear DNA, amplified in two fragments, encompassing the 3′ end of the 5.8S rRNA gene, the ITS2 region, and the 5′ end of the 28S rRNA gene (primers LSU-1/LSU-3 and LSU-2/LSU-5 of [21,22]), circa 1300 bp in total.
The settings of the PCR amplification were as follows. COI and 16S: initial denaturation at 96 °C (3 min); 35 cycles of denaturation at 95 °C (30 s), annealing at either 48 °C (COI) or 50 °C (16S) (1 min), and extension at 72 °C (2 min); and final extension at 72 °C (5 min). ITS2+28S: initial denaturation at 95 °C (3 min); 40 cycles of denaturation at 95 °C (30 s), annealing at either 50 °C (ITS2 section) or 45 °C (28S section) (1 min), and extension at 72 °C (5 min for ITS2 section or 2 min for 28S section); and final extension at 72 °C (4 min). Small variations in the protocol were pursued regarding the annealing temperatures and number of cycle steps for repeats of the samples that failed to amplify in the first round.
PCR success was assessed via agarose gel electrophoresis. The PCR products were cleaned with ExoSAP-IT™ (Affymetrix Inc., Santa Clara, CA, USA) following the manufacturer’s protocol. Samples were sent out to Massey Genome Service (Massey University, Palmerston North, New Zealand) to be Sanger sequenced.
2.3. Sequence Assembly and Alignment
Sequences were quality proofed and assembled in Geneious Prime (v.2020.2.2, Biomatters Ltd., Auckland, New Zealand), and the consensus was uploaded to GenBank. See Table 1 for the registration numbers.
Alignment of the consensus sequences was likewise conducted in Geneious Prime using the MAFFT plugin (v.7.450; [23,24]) with the default settings. The resulting alignment of each marker (COI, 16S, and ITS+28S) was visually proofed for inconsistencies. The alignment was then run through Gblocks [25], using the least restrictive settings, to eliminate poorly aligned or data-deficient positions that could interfere with the analyses. The resulting alignments were then concatenated for a single phylogenetic analysis.
2.4. Phylogenetic Analysis
Phylogenetic analyses were performed via Bayesian inference using MrBayes (v.3.2.7 [26]) via the CIPRES Science Gateway (v. 3.3 [27]). Two concurrent analyses were run, each with 4 Markov chains of 60 million generations (the first 20% discarded as ‘burn-in’), the default priors, nst = 6, rates = invgamma, temperature parameter = 0.1, sampling every 1000 generations, and with the substitution model parameters unlinked across the markers (COI, 16S, and ITS+28S). MCMC convergence was assessed by examining the standard deviation of split frequencies (~0.01) and the potential scale reduction factor (PSRF~1.0), as well as trace plots in Geneious [28].
3. Results
Including the outgroup, 64 terminal taxa were used in the analysis (Table 1 and Table 2). After the exclusion of positions with Gblocks (see above), the resulting COI sequences were 653 bp long, 16S were 405 bp long, and ITS2+28S were 1055 bp long. Thus, the concatenated sequences used for the analyses were 2011 bp long.
In a first trial, the African species Trachycystis tollini and Xerocystis capensis (Table 1) formed a sister pair with very long branches inside the clade formed by all other Trachycystinae (Figure 1). This was deemed an artefact due to the short and fragmentary sequences of both species (Table 1), and they were thus removed from further analyses, as they were likely confounding the relationships within the Trachycystinae. Their exclusion did not change the topology of the tree outside of the Trachycystinae.
As hypothesized based on the overall morphology, Reticulapex and Pilula are not members of the Trachycystinae, or even the Punctoidea. Reticulapex belongs to the family Acavidae, with good support (posterior probability or PP = 0.94; Figure 1). The position of Pilula is not so clear; however, it seems more closely related to the superfamily Helicarionoidea (PP = 0.86), recovered as paraphyletic here (Figure 1).
The arrangement of the Punctoidea is similar to that of [1], with Endodontidae being the first branch, followed by the Cystopeltidae (including Australian and South American branches), and a paraphyletic Charopidae that includes the Punctidae. Nevertheless, the addition of African species and potentially basal New Zealand Charopidae to the dataset has brought more resolution to the “Charopidae”.
Trachycystinae was recovered as a monophyletic group with some support (PP = 0.90; Figure 1) and the most basal branch of the “Charopidae” and Punctidae clade, though with low support (PP = 0.79; Figure 1). A well-supported group formed by the New Zealand Flammoconcha and Therasiella (both represented by their type species) was recovered as basal to all other “Charopidae” and Punctidae (PP = 1.0; Figure 1).
4. Discussion
The present results support Trachycystinae as a valid clade at the subfamily level, restricted to African taxa of afrodontas and pinwheels. Other African taxa tested (Reticulapex and Pilula) that had been previously assigned to the Punctoidea [3,4,18] are excluded from this superfamily and transferred to the Acavidae and Helicarionoidea, respectively.
Trachycystinae is a monophyletic group, the most basal one in the family “Charopidae” (Figure 1). It possibly represents an old southern African lineage potentially dating back to the split of Gondwana, while the Cystopeltidae were distributed in Australia and South America, and the remaining “Charopidae” in Zealandia and Australia (and later, in the Americas, and eventually Europe with the Punctidae) [1]. The restricted distribution of the Trachycystinae in southern Africa is a common pattern shared by other relict family and subfamily-level taxa of land snails [29]. There is no fossil record available of Trachycystinae dating from the Cretaceous or Paleogene, when the split might have occurred [29,30]. To my knowledge, the oldest fossils currently known are three species of Trachycystis from the Early Miocene in Kenya [31], which imply a different and possibly broader geographic distribution of the subfamily in the past. Further fossil records belong to the extant species Xerocystis capensis and to “Trachycystis” sp. from the Pliocene of South Africa [32,33].
If further studies support the present findings, the elevation of the Trachycystinae to the family level (as sister to the “Charopidae” and Punctidae) might be warranted, similar to the status of the recently redefined Cystopeltidae [1].
It was not possible, however, to obtain a clear internal structure of the Trachycystinae (Figure 1). It is uncertain at this point whether this is due to missing sequences of some markers for a few species or to a lack of further species. Nevertheless, the present phylogeny hinted that the genus-level classification of some species (in the Chalcocystis and Trachycystis), as well as the validity of some genera, might warrant further attention in future studies.
Finally, even though it was not part of the objectives of this study, the arrangement of the present phylogeny (Figure 1) indicates that Flammoconchinae is a valid taxon in the subfamily level within the “Charopidae”. Excluding Trachycystinae, this newly defined Flammoconchinae is basal to all other “Charopidae” and Punctidae (Figure 1). It is endemic to New Zealand and contains the genera Flammoconcha and Therasiella, both of which have similar shells to some species of African pinwheels. The phylogenetic position of the genus Calymna F.W. Hutton, 1883, often classified in this subfamily, requires further investigation. No fossil record of this subfamily is known [1].
5. Conclusions
Based on the present results, the following revised taxonomic classification, listing only the genera that were present in the analysis, is proposed:
| Superfamily Rhytidoidea Pilsbry, 1893 |
| Family Acavidae Pilsbry, 1895 |
| Genus Reticulapex Emberton & Pearce, 2000 |
| Superfamily Punctoidea Morse, 1864 |
| Family Endodontidae Pilsbry, 1895 |
| Family Cystopeltidae Cockerell, 1891 |
| Family Punctidae Morse, 1864 |
| Family “Charopidae” Hutton, 1884 |
| Subfamily Charopinae Hutton, 1884 |
| Subfamily Flammoconchinae Schileyko, 2001 |
| Genus Flammoconcha Dell, 1952 |
| Genus Therasiella Powell, 1948 |
| Subfamily Trachycystinae Schileyko, 1986 |
| Genus Afrodonta Melvill & Ponsonby, 1908 |
| Genus Chalcocystis H. Watson, 1934 |
| Genus Chilocystis H. Watson, 1934 |
| Genus Phortion Preston, 1910 |
| Genus Psichion Gude, 1911 |
| Genus Trachycystis Pilsbry, 1893 |
| Genus Xerocystis H. Watson, 1934 |
| Superfamily Helicarionoidea Bourguignat, 1877 |
| Helicarionoidea incertae sedis |
| Genus Pilula E. von Martens, 1898 |
Funding
This research received no external funding.
Institutional Review Board Statement
Not applicable.
Informed Consent Statement
Not applicable.
Data Availability Statement
All data are present in this article. Accession numbers of genetic sequences hosted on GenBank are provided.
Acknowledgments
I am grateful to Igor Muratov (NMSA), John Slapcinsky (UF), and Martyn Kennedy (OZD) for granting access to the specimens and samples under their care; to Fred Brook for the discussion about which basal New Zealand charopid species to include in the analysis; to Ton de Winter for the discussion and comments on an earlier version of this manuscript; and to two anonymous reviewers for their helpful comments.
Conflicts of Interest
The author declares no conflict of interest.
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Figure 1.
Bayesian inference phylogenetic tree based on CO1, 16S, and ITS2+28S with a focus on African taxa. Posterior probabilities are shown on nodes. Scale bar = substitutions per site.
Figure 1.
Bayesian inference phylogenetic tree based on CO1, 16S, and ITS2+28S with a focus on African taxa. Posterior probabilities are shown on nodes. Scale bar = substitutions per site.
Table 1.
List of species used in this study, with GenBank accession numbers for each marker, and voucher data (registration number and locality of provenance). * Fragmentary sequences, excluded from final analysis.
Table 1.
List of species used in this study, with GenBank accession numbers for each marker, and voucher data (registration number and locality of provenance). * Fragmentary sequences, excluded from final analysis.
| Species | COI | 16S | ITS2+28S | Voucher | Provenance |
|---|---|---|---|---|---|
| African Charopidae | |||||
| Afrodonta bilamellaris Melvill & Ponsonby, 1908 | ON365470 | ON374086 | ON376035 | NMSA-Mol 0W9618 | South Africa, KwaZulu-Natal, Port Shepstone, Four Man’s Hill |
| Chalcocystis viridula (Connolly, 1939) | — | ON374088 | ON376037 | NMSA-Mol 0W9407 | South Africa |
| Chilocystis calorama (Melvill & Ponsonby, 1899) | ON365472 | ON374089 | ON376038 | NMSA-Mol 0W9636 | South Africa, KwaZulu-Natal, Port Shepstone, Simuma Hill |
| Chilocystis scolopendra (Melvill & Ponsonby, 1903) | ON365473 | ON374090 | ON376039 | NMSA-Mol 0W7837 | South Africa, KwaZulu-Natal, Port Shepstone, Hlokohloko Valley |
| Phortion oconnori (Preston, 1912) | ON365475 | ON374092 | ON376041 | NMSA-Mol 0W8889 | South Africa, Western Cape, Bainskloof Pass |
| Psichion inclara (Morelet, 1889) | ON365478 | ON374095 | ON376043 | NMSA-Mol 0W9163 | South Africa, Eastern Cape, Woody Cape Nature Reserve |
| Trachycystis bathycoele (Melvill & Ponsonby, 1892) | ON365481 | — | — | NMSA-Mol 0P0178 | South Africa, Limpopo, Entabeni Forest |
| Trachycystis glanvilliana (Ancey, 1890) | ON365482 | ON374097 | ON376047 | NMSA-Mol 0W7149 | South Africa, KwaZulu-Natal, Karkloof Falls |
| Trachycystis tollini (Benson, 1856) | — | — | ON376048 * | NMSA-Mol 0W9407 | South Africa, Western Cape, Platbos Forest Reserve |
| Xerocystis capensis (L. Pfeiffer, 1841) | — | — | ON376049 * | NMSA-Mol 0W8893 | South Africa, Western Cape, De Hoop Nature Reserve |
| African taxa, not Charopidae | |||||
| Pilula cordemoyi (G. Nevill, 1870) | ON365476 | ON374093 | — | UF 415477 | Réunion Island, Basse Valley |
| Reticulapex michellae K.C. Emberton et al., 2010 | ON365479 | — | ON376044 | UF 421124 | Madagascar, Andriantantely Massif |
| Reticulapex sp. | — | — | ON376045 | UF 421119 | Madagascar, Andriantantely Massif |
| New Zealand Charopidae | |||||
| Cavellia buccinella (Reeve, 1852) | ON365471 | ON374087 | ON376036 | OZD Cabuc-1 | New Zealand, Northland, Kaipara |
| Flammoconcha cumberi (Powell, 1941) | ON365474 | ON374091 | ON376040 | OZD FB29 | New Zealand, West Coast, Grange Ridge |
| Pseudallodiscus ponderi Climo, 1971 | ON365477 | ON374094 | ON376042 | OZD Ppond-1 | New Zealand, Auckland, Pahiatua Hill Scenic Reserve |
| Therasiella celinde (Gray, 1850) | ON365480 | ON374096 | ON376046 | OZD FB116 | New Zealand, Waikato, Pokeno |
Table 2.
List of species used in the phylogenetic analysis for which data were retrieved from GenBank, with accession numbers for each marker, and provenance data of the voucher specimen.
Table 2.
List of species used in the phylogenetic analysis for which data were retrieved from GenBank, with accession numbers for each marker, and provenance data of the voucher specimen.
| Species | COI | 16S | ITS2+28S | Provenance of Voucher |
|---|---|---|---|---|
| Charopidae | ||||
| Allodiscus dimorphus (Reeve, 1852) | MN792581 | MN756708 | MN782439 | New Zealand, Auckland, Waitakere Ranges, Titirangi, Atkinson Track |
| Alsolemia longstaffae (Suter, 1913) | MN792582 | MN756709 | MN759313 | New Zealand, Southland, Colac Bay |
| Chalcocystis aenea (F. Krauss, 1848) | MN792590 | MN756717 | MN782447 | South Africa, KwaZulu-Natal, Hluhluwe |
| Charopa coma (Gray, 1843) | MN792591 | MN756718 | MN782448 | New Zealand, Auckland, Waitakere Ranges, Titirangi, Paturoa Stream |
| Fectola infecta (Reeve, 1852) | MN792600 | MN756727 | MN782457 | New Zealand, Waikato, Coromandel Peninsula, Port Charles |
| Flammulina zebra (Le Guillou, 1842) | MN792601 | MN756728 | MN782458 | New Zealand, Tasman, Lake Daniells |
| Mitodon wairarapa (Suter, 1890) | MN792607 | MN756732 | MN782464 | New Zealand, Southland, Stewart Island, Mason Bay, Gutter |
| Mocella eta (Pfeiffer, 1853) | MN792608 | MN756733 | MN782465 | New Zealand, Northland, Umuheke Bay |
| Neophenacohelix giveni (Cumber, 1961) | MN792609 | MN756743 | MN782466 | New Zealand, Northland, Whangarei, Coronation Reserve |
| Otoconcha dimidiata (L. Pfeiffer, 1853) | MN792614 | MN756738 | MN782471 | New Zealand, Northland, Whangarei, Bream Head |
| Phacussa helmsi (Hutton, 1882) | MN792618 | MN756742 | MN782475 | New Zealand, West Coast, Greymouth, Point Elizabeth |
| Phenacohelix pilula (Reeve, 1852) | MN792619 | MN756744 | MN782476 | New Zealand, Northland, Whangaruru North Head |
| Radioconus amoenus (Thiele, 1927) | MN792623 | MN756749 | MN782481 | Brazil, Santa Catarina, Florianópolis, Gruta do Triângulo |
| Radiodiscus sp. | MN792625 | MN756751 | MN782483 | Brazil, Bahia, Ilhéus |
| Radiodiscus sp. | MN792626 | MN756752 | MN782484 | Chile, Chiloé, Chiloé National Park, Chepu |
| Ranfurlya constanceae Suter, 1903 | MN792627 | MN756753 | MN782485 | New Zealand, Auckland Islands, Adams Island |
| Sinployea atiensis (Pease, 1870) | MN792628 | MN756754 | MN782486 | Cook Islands, Rarotonga, Tupapa Valley |
| Stenacapha hamiltoni (Cox, 1868) | MN792629 | MN756755 | MN782487 | Australia, Tasmania, Central Plateau, Viormy |
| Suteria ide (Gray, 1850) | MN792630 | MN756756 | MN782488 | New Zealand, Manawatu-Wanganui, Bushy Park |
| Therasia thaisa Hutton, 1883 | MN792631 | MN756757 | MN782489 | New Zealand, Southland, Clifden, Clifden Limestone Cave System |
| Cystopeltidae | ||||
| Cystopelta bicolor Petterd & Hedley, 1909 | MN792592 | MN756719 | MN782449 | Australia, Tasmania, Bronte Park |
| Diemenoropa kingstonensis (Legrand, 1871) | MN792616 | MN756740 | MN782473 | Australia, Tasmania, Skullbone Plains, Kenneth Lagoon |
| Lilloiconcha cf. gordurasensis (Thiele, 1927) | MN792604 | MN756731 | MN782461 | Brazil, Alagoas, Pedra Talhada Biological Reserve |
| Lilloiconcha gordurasensis (Thiele, 1927) | MN792605 | — | MN782462 | Brazil, São Paulo, São Paulo, Burle Marx Park |
| Lilloiconcha superba (Thiele, 1927) | MN792606 | — | MN782463 | Brazil, Alagoas, Pedra Talhada Biological Reserve |
| Scelidoropa officeri (Legrand, 1871) | MN792617 | MN756741 | MN782474 | Australia, Tasmania, Flinders Island, Brougham Sugarloaf |
| Zilchogyra sp. | MN792632 | — | MN782490 | Brazil, São Paulo, Cotia, Morro Grande Reserve |
| Endodontidae | ||||
| Libera fratercula (Pease, 1867) | MN792603 | MN756730 | MN782460 | Cook Islands, Rarotonga, Tupapa |
| Punctidae | ||||
| Laoma leimonias (Gray, 1850) | MN792602 | MN756729 | MN782459 | New Zealand, Northland, Kaihu, Maropiu Road |
| Paralaoma servilis (Shuttleworth, 1852) | MN792615 | MN756739 | MN782472 | New Zealand, Southland, Colac Bay |
| Phrixgnathus celia Hutton, 1883 | MN792620 | MN756745 | MN782477 | New Zealand, Southland, Stewart Island, Mason Bay |
| Punctum californicum Pilsbry, 1898 | MN792621 | MN756746 | MN782478 | USA, California, San Francisco, Presidio, Lincoln Boulevard |
| Punctum pygmaeum (Draparnaud, 1801) | MN812719 | MN756747 | MN782479 | UK, Monmouthshire, Monmouth, Pentwyn Farm |
| Punctum randolphii (Dall, 1895) | MN792622 | MN756748 | MN782480 | Canada, British Columbia, Pemberton, Riverside Trail, Lillooet River |
| Helicarionoidea | ||||
| Antiquarion evelynensis Hyman & Köhler, 2020 | MN654044 | MN654081 | — | Australia, Queensland, Milllaa Millaa |
| Antiquarion ravenshoe Hyman & Köhler, 2020 | MN654053 | MN654089 | — | Australia, Queensland, Tully |
| Mysticarion insuetus Iredale, 1941 | KY662466 | KY662376 | — | Australia, New South Wales, Newcastle |
| Nitor subrugatus (Reeve, 1852) | MH248130 | MH255873 | — | Australia, New South Wales, Tooloom National Park |
| Sarika resplendens (Philippi, 1847) | MT364982 | MT365763 | MT365707 | Thailand |
| Stanisicarion freycineti (Férussac, 1821) | MN073628 | MN073747 | — | Australia, Queensland, Eastern Escarpment Conservation Area |
| Rhytidoidea | ||||
| Acavus phoenix (L. Pfeiffer, 1854) | — | — | AY014083 | Unknown |
| Ampelita lamarei (L. Pfeiffer, 1853) | — | — | KP230504 | Madagascar, Msoala Peninsula |
| Embertoniphanta goudotiana (Férussac, 1839) | — | — | KP230517 | Madagascar, Tsingy Beanka, Belitsaka |
| Rhytida greenwoodi (Gray, 1850) | KT970868 | KT970900 | KP230525 | New Zealand, Waikato, Raglan |
| Trochomorphoidea | ||||
| Asperitas trochus (O. F. Müller, 1774) | MT654630 | MT651546 | MT651601 | Indonesia |
| Dyakia hugonis (L. Pfeiffer, 1864) | MT803064 | MT741748 | MT741915 | Malaysia, Sabah, Segaliud Lokan Forest Reserve |
| Rhinocochlis nasuta (Metcalfe, 1852) | MT803094 | MT741771 | MT741938 | Malaysia, Sarawak, Sibu, Lanjak Entimau |
| Outgroup | ||||
| Gulella caryatis (Melvill & Ponsonby, 1898) | HQ328133 | HQ328323 | GQ330510 | Namibia |
| Subulina octona (Bruguière, 1789) | JX988066 | JX988353 | MF444887 | Palau |
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MDPI and ACS Style
Salvador, R.B. Phylogenetic Position of African Punctoid Snails (Stylommatophora, Punctoidea, Trachycystinae). Taxonomy 2022, 2, 227-235. https://doi.org/10.3390/taxonomy2020017
AMA Style
Salvador RB. Phylogenetic Position of African Punctoid Snails (Stylommatophora, Punctoidea, Trachycystinae). Taxonomy. 2022; 2(2):227-235. https://doi.org/10.3390/taxonomy2020017
Chicago/Turabian StyleSalvador, Rodrigo Brincalepe. 2022. "Phylogenetic Position of African Punctoid Snails (Stylommatophora, Punctoidea, Trachycystinae)" Taxonomy 2, no. 2: 227-235. https://doi.org/10.3390/taxonomy2020017
