2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.981.5358353583MonographArchitaenioglossaGastropodaBiodiversity & ConservationBiogeographyPalaeontologySystematicsCenozoicAsiaGenus-level revision of the Alycaeidae (Gastropoda, Cyclophoroidea), with an annotated species cataloguePáll-GergelyBarna1pallgergely2@gmail.comhttps://orcid.org/0000-0002-6167-7221SajanSheikh23https://orcid.org/0000-0002-2785-6824TripathyBasudev2MengKaibaryer4AsamiTakahiro5https://orcid.org/0000-0001-5706-0272AblettJonathan D.6https://orcid.org/0000-0002-7277-1934Plant Protection Institute, Centre for Agricultural Research, Herman Ottó út 15, Budapest, H-1022, HungaryCentre for Agricultural ResearchBudapestHungaryZoological Survey of India, Prani Vigyan Bhawan, M Block, New Alipore, Kolkata 700053, West Bengal, IndiaZoological Survey of IndiaKolkataIndiaWildlife Institute of India, Chandrabani, Dehradun 248 002, Uttarakhand, IndiaWildlife Institute of IndiaDehradunIndiaNational Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences, Beijing, ChinaInstitute of Zoology, Chinese Academy of SciencesBeijingChinaDepartment of Biology, Shinshu University, Matsumoto 390-8621, JapanShinshu UniversityMatsumotoJapanMollusca Section, Invertebrates Division, Department of Life Sciences, The Natural History Museums, London SW7 5BD, United KingdomNatural History MuseumLondonUnited Kingdom
Corresponding author: Barna Páll-Gergely (pallgergely2@gmail.com)
Academic editor: Frank Köhler
20202910202098112207C44C797-C412-5A71-BAE0-32A55E6FA5DC5194AAC8-6B8A-473F-8A41-470A60182A0B42484222304202015082020Barna Páll-Gergely, Sheikh Sajan, Basudev Tripathy, Kaibaryer Meng, Takahiro Asami, Jonathan D. AblettThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.http://zoobank.org/5194AAC8-6B8A-473F-8A41-470A60182A0B
412 species-group names (including 11 replacement names), and 14 genus-group names of the Alycaeidae have been introduced to date. Type materials of 85% (336) of the known species and subspecies were examined, a further 5% (19) of the taxa were studied using available non-type material, and for another 6% (22) the original descriptions were sufficiently detailed to evaluate their taxonomic status. Only 3% of the taxa (12) could not be examined. Special attention was paid to the sculpture of the embryonic whorls and the sutural tube-microtunnel system in order to provide a novel classification for this group.
In this study 363 taxa (320 species or 43 subspecies) are accepted within the family Alycaeidae. Of these, 22 have been described by the lead author and his coauthors in previous publications. In addition, there are 18 species that were formerly classified in Cycloryx and now belong to Pincerna due to its synonymy with Cycloryx. Among the remaining 323 species, 209 (65%) are transferred here to another genus, whilst 114 (35%) have remained in their original genus.
Seven genera are accepted. While some questions (e.g., the distinction between Pincerna and Alycaeus) remained unanswered, this revision made three main achievements: (1) The Dicharax species were identified based on the absence of spiral striation on the entire shell; (2) the Metalycaeus species were identified based on the spiral striation of the protoconch; (3) and Stomacosmethis was separated from Alycaeus based on the extremely short sutural tube.
Five nominal species are being synonymised with other species, and eight species are now treated as subspecies. The following replacement names are proposed: Dioryxurnulaniosiensis Páll-Gergely, nom. nov. for Alycaeusurnulavar.daflaensis Godwin-Austen, 1914; Dioryxurnularotundus Páll-Gergely, nom. nov. for Alycaeusurnulavar.globosus Godwin-Austen, 1914; Pincernacrenilabrisjuttingae Páll-Gergely, nom. nov. for Alycaeuscrenilabrislaevis van Benthem Jutting, 1959; Pincernacrenilabriskorintjiensis Páll-Gergely, nom. nov. for Alycaeuscrenilabrislatecostatus van Benthem Jutting, 1959; Dicharaxconicusjatingaensis Páll-Gergely, nom. nov. for Alycaeusconicusvar.nanus Godwin-Austen, 1914; Metalycaeusgodwinausteni Páll-Gergely, nom. nov. for Alycaeusneglectus Godwin-Austen, 1914; and finally Metalycaeussuhajdai Páll-Gergely, nom. nov. for Alycaeusvarius Godwin-Austen, 1914.
land snailmuseum collectionssystematicstaxonomySynthesis of Systematic Resources226506501100000780European Commissionhttp://doi.org/10.13039/501100000780Citation
Páll-Gergely B, Sajan S, Tripathy B, Meng K, Asami T, Ablett JD (2020) Genus-level revision of the Alycaeidae (Gastropoda, Cyclophoroidea), with an annotated species catalogue. ZooKeys 981: 1–220. https://doi.org/10.3897/zookeys.981.53583
Introduction
The Alycaeidae are operculate land snails in the superfamily Cyclophoroidea. Approximately 360 Asian species and subspecies have been described so far, and classified into 14 genera or subgenera. Alycaeids inhabit a vast area that stretches from the Western Ghats (India) through the Himalaya to Japan in the east, the Chinese Gansu and Shaanxi provinces in the north and Indonesia to the south (Godwin-Austen 1882–1920; Gude 1921; van Benthem Jutting 1948, 1959; Minato 1988; Gittenberger et al. 2017; Aravind & Páll-Gergely 2019). The alycaeid shell is characterised by a tube, which is closed at its outer end, and opens into the inside of the shell just behind the operculum. This tube is in contact with numerous, extremely narrow tunnels, which are formed by the outermost shell layer (Páll-Gergely et al. 2016).
Some terrestrial caenogastropod genera lacking such a sutural tube have provisionally been assigned to the Alycaeidae. Laotia Saurin, 1953, which includes two species, has been included either in the Diplommatinidae, because of its similarity with Helicomorpha, or in the Alycaeidae, because of its resemblance to the alycaeid Chamalycaeus (Saurin 1953; Páll-Gergely 2014). The latest publication on Laotia placed this genus in Alycaeidae (Do et al. 2015). The Madagascan endemic Boucardicus Fischer-Piette & Bedoucha, 1965 (Madecataulus Fischer-Piette & Bedoucha, 1965 is a synonym, see Emberton and Pearce 1999) has also been placed in Alycaeidae due to a similar shell shape and radula (Emberton 2002; Egorov 2019). Following recent extensive surveys, there are now approximately 200 accepted Boucardicus species (Fischer-Piette et al. 1993; Emberton and Pearce 1999; Emberton 2002; Emberton et al. 2010; Balashov and Griffiths 2015).
Our study covers the systematics of the Alycaeidae sensu stricto, a group that is characterised by the possession of an external tube that runs along the suture (see above). Although anatomical and radular characters are known for some species (Godwin-Austen 1882–1920; Tielecke 1940; Venmans 1956; Emberton and Pearce 1999; Emberton 2001), those can only be used as supplementary information to hypothesise about the relatedness of the genera, and cannot be used for the appropriate generic placement of species at the current time. Thus, the classification presented here is primarily based on morphological characters of the shell.
The current generic subdivision of the Alycaeidae was established over a century ago and no genus-level revision has been proposed since the publication of Kobelt’s (1902) monograph. Arguably, most authors did not examine the type species of genera, especially those of Alycaeus, Chamalycaeus and Dicharax, when attributing new alycaeid species to any of these genera. Moreover, some allegedly diagnostic characters of the genera and subgenera may not reflect evolutionary relationships because these character states have probably evolved in convergence. For example, the low spire was regarded as the key trait of Chamalycaeus (Kobelt 1902). However, most Chamalycaeus, Metalycaeus and Dicharax species are low-spired, and even Alycaeusjousseaumei has a depressed shell, while, there are Dicharax and Metalycaeus species that are high spired (Páll-Gergely et al. 2017). Similarly, Dicharax was defined on the basis of a swelling behind the peristome (Kobelt 1902). This trait, however, occurs in several species of the genera Metalycaeus and Dicharax, and the strength of the swelling is very variable across Dicharax and Metalycaeus species (Páll-Gergely et al. 2017). Lastly, the outer surface traits of the operculum, which defined the genera Pincerna, Stomacosmethis and Metalycaeus, are also variable within species, and, on the other hand, show similar morphology between species not closely related.
The aim of this study is to provide an updated generic classification of the Alycaeidae based on the principles of our former paper (Páll-Gergely et al. 2017) focusing on two key traits largely neglected in previous publications, because these are presumed to be more useful to distinguish natural groups. Firstly, the sculpture on the outer shell surface and secondly, the length and sculpture of the area where the sutural tube is situated and the surface is differently ribbed from the other whorl range. Microtunnels functioning as complex gas exchange device are present with the sutural tube, which could provide useful traits for alycaeid systematics, at least in some of the groups (Páll-Gergely et al. 2016).
Taxonomic history of Alycaeidae
Pfeiffer (1858) divided the fourteen species of the genus Alycaeus (equivalent to present-day Alycaeidae) known at the time into two informal groups, namely a species with subturbinate shells (“Subturbinati”) and those with depressed shells (“Depressi”). Benson (1859) named three sections within Alycaeus as follows. (1) Alycaeus: “the last whorl constricted somewhat remotely from the aperture, tumid on both sides of the constriction”; (2) Charax: “constriction broad, contiguous to the aperture, and divided more or less remotely from it, across the whorl, by a ridge which is hollow internally”; (3) Dioryx: “constriction narrow, and immediately behind the aperture; the sutural tube arising proportionally nearer to the peristome than in Alycaeus and Charax”. In Benson’s (1859) system, all the three groups were further sub-divided into unnamed subgroups on the basis of shell shape (Alycaeus and Dioryx) and the morphology of the swelling between the constriction and the aperture (Charax). Pfeiffer (1876) introduced the name Orthalycaeus as a subgenus of Alycaeus and divided it into four subgroups. The name was established without description, but contained 26 species, which made it available. Pfeiffer seemingly intended this name to be used as what we would call today a nominotypical subgenus. He did not select a type species, which was subsequently done by Kobelt (1879: 191), who selected Alycaeusgibbus as the type species of Orthalycaeus. Because this species is also the type species of Alycaeus (also by subsequent designation), these two genus names are objective synonyms. Kobelt and Möllendorff (1897) recognised two genera within the family Alycaeidae: Dioryx and Alycaeus (with the subgenera Orthalycaeus, Chamalycaeus, Charax). Kobelt’s (1902) monograph was based on the same system as the one by Kobelt and Möllendorff (1897), but he treated the Alycaeinae as a subfamily of the Cyclophoridae. Kobelt (1902) recognised Dioryx as a distinct genus and subdivided Alycaeus into four ‘sections’: Alycaeus, Chamalycaeus Kobelt & Möllendorff, 1897 (incorrect attribution of authors, see under Chamalycaeus), Dicharax Kobelt & Möllendorff, 1900 (replacement name for Charax Benson, 1859, non Charax Scopoli, 1777 [Pisces]) and Metalycaeus Pilsbry, 1900. Metalycaeus was described for two Japanese species (A.melanopoma and A.hirasei). Later, A.tsushimanus was also included in Metalycaeus (Pilsbry & Y. Hirase, 1909a). Metalycaeus has only been reported from Japan, and was diagnosed on the basis of a thickened ring on the outer side of the operculum. In later publications it was not accepted as a distinct taxon, but treated as a junior synonym of Chamalycaeus (see Minato 1988). Preston (1907) described the subgenus Pincerna for Alycaeus (Pincerna) liratula Preston, 1907. According to Preston Pincerna has an alycaeiform shell, which is higher than wide, and the operculum with a “circular cup” on its outer surface. The subgenera Cycloryx and Raptomphalus were described by Godwin-Austen (1914). The former is characterised by the ovate-conoid shell and an extremely short tube, which is often pear or club-shaped, whereas the latter has a conspicuous keel on the umbilical margin. Stomacosmethis Bollinger, 1918 was defined on the basis of a pipe, tongue or cup-shaped structure on the outer side of the operculum and included two species from southern Celebes and eastern Borneo. Three genus-group taxa (Sigmacharax Kuroda, 1943, Cipangocharax Kuroda, 1943, Awalycaeus Kuroda, 1951) were described from Japan. These have been used either as genera or as subgenera of Chamalycaeus and can be distinguished from each other as follows: Sigmacharax has a peculiar, sigmoid last whorl with an ovate aperture having an interior ridge, Cipangocharax has a thick, “shelly” operculum with closely imbedded spiral cuticular lamellae on its outer surface, and Awalycaeus has a very short distance between the starting point of the tube and the peristome, its operculum is situated at the aperture, not deeper as in other groups.
In our own works we have defined Metalycaeus by the presence of a spirally striated protoconch, and several species from China, Vietnam, Laos, and Japan have been placed in this genus (Páll-Gergely and Asami 2017; Páll-Gergely et al. 2017). We also found that Chamalycaeus possesses a protoconch without spiral striae, and a teleoconch with spiral striation. Consequently, most species previously classified as Chamalycaeus have been transferred to Metalycaeus or Dicharax. The latter is characterised by the absence of spiral striation on the entire shell (Páll-Gergely et al. 2017). We further synonymised Cycloryx with Pincerna, which is accepted as a distinct genus (Páll-Gergely 2017).
Materials and methods
Specimens (shells and radulae) were examined using a low vacuum SEM (Miniscope TM-1000, Hitachi High-Technologies, Tokyo) directly without coating. The ethanol-preserved specimens were dissected under a Zeiss stereomicroscope, and photographs were taken using a Keyence LHX5000 digital microscope.
Photographs of shells were taken using various photographic equipment in our laboratories and in museum collections. Photographs of types deposited in the IZCAS, ZSI, SMF are published here. In cases of the other museums the photographs of types are mostly available online, or they will be published by us in separate papers.
Locality data cited as verbatim from the specimen labels, and no English translations are provided in most cases.
Differences in size are indicated in the generic diagnosis using the following terms: very small (smaller than 3 mm), small (3–4 mm), medium-sized (4–6 mm), large (6–8 mm), very large (larger than 8 mm). We distinguish three regions of the teleoconch, following Páll-Gergely et al. (2017): Region 1 (R1) ranging from the beginning of the teleoconch to the beginning of the differently ribbed region where the sutural tube lies, Region 2 (R2) extending from the end of R1 to the constriction (i.e., the length of R2 usually corresponds with the length of the sutural tube, see Páll-Gergely et al. 2016; Páll-Gergely and Asami 2017), and Region 3 (R3) ranging from the constriction up to the peristome.
In order to maintain consistency with the editorial conventions of MolluscaBase (2020), initials of first names of authors are indicated in all cases where a given author shares the same family name with another malacologist (i.e., Y. Hirase, L. Pfeiffer).
Alycaeusgibbosulus Stoliczka, 1872: ALY03, Malaysia, Malay Peninsula, Baling, after a large bridge of the Baling River, 05°40.950'N, 100°54.883'E, 100 m, leg. Fatley, R., Juhász, A., Majoros, G., Motochin, R., Páll-Gergely, B., 22.07.2016, HNHM 104424.
Stomacosmethisdohrni (O. Boettger, 1893): Indonesia, Kalimantan Selatan, Beramban, leg. Yansen Chen, Apr 2012, HNHM 104426.
Stomacosmethisbalingensis (Tomlin, 1948): Malaysia, Malay Peninsula, Baling, after a large bridge of the Baling River, 05°40.950'N, 100°54.883'E, 100 m, leg. Fatley R., Juhász A., Majoros G., Motochin R., Páll-Gergely B., 22.07.2016., HNHM 104425.
Alycaeusgibbosulus Stoliczka, 1872: Malaysia, Penang, Penang National Park, Around Monkey beach, 5°28.457'N, 100°11.165'E, 81 m a.s.l. (ALY30 in molecular study), leg. Hirano, T., 21.07.2016, HNHM 104857.
Stomacosmethisbalingensis (Tomlin, 1948): Malaysia, Malay Peninsula, Baling, after a large bridge of the Baling River 05°40.950'N, 100°54.883'E, leg. Fatley, R., Harl, J., Juhász, A., Majoros, G., Motochin, R., Páll-Gergely, B., 22.07.2016. (2016.07.22A, specimen1), HNHM 104861.
Stomacosmethisperakensis (Crosse, 1879): 2016.07.22A, Malaysia, Malay Peninsula, Baling, after a large bridge of the Baling River, 05°40.950'N, 100°54.883'E, leg. Fatley, R., Harl, J., Juhász, A., Majoros, G., Motochin, R., Páll-Gergely, B., 22.07.2016, HNHM 104430.
Abbreviations
AMNHAmerican Museum of Natural History (New York, USA);
ANSPAcademy of Natural Sciences (Philadelphia, USA);
D shell diameter;
HA Collection András Hunyadi (Budapest, Hungary);
HBUMM Mollusc collection of the Museum of Hebei University (Baoding, China);
HNHMHungarian Natural History Musem (Budapest, Hungary);
IZCASNational Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences (Beijing, China);
MCZMuseum of Comparative Zoology (Massachusetts, USA);
MNHNMuséum National d’Histoire Naturelle (Paris, France);
NHMWMuseum of Natural History of Vienna (Vienna, Austria);
NZSI National Zoological Collection of the Zoological Survey of India (when cited specimens deposited in the ZSI);
NSMTNational Museum of Nature and Science, Tsukuba, Japan;
PGB Collection Barna Páll-Gergely (Mosonmagyaróvár, Hungary);
RBINSRoyal Belgian Institute of Natural Sciences (Brussels, Belgium);
RMNHNational Museum of Natural History Naturalis (Leiden, The Netherlands);
SMFSenckenberg Forschungsinstitut und Naturmuseum (Frankfurt am Main, Germany);
UFFlorida Museum of Natural History, University of Florida (USA);
UMMZUniversity of Michigan, Museum of Zoology (Ann Arbor, USA);
UMZC University Museum of Zoology (Cambridge, United Kingdom);
USNMSmithsonian National Museum of Natural History (Washington, USA);
ZMBMuseum für Naturkunde (Berlin, Germany);
ZSIZoological Survey of India (Kolkata, India).
Description and assessment of morphological charactersShell morphologyProtoconch sculpture (Fig. 1)
The protoconch is spirally striated in Metalycaeus (Fig. 1E, F) (rarely unstriated, see M.laevis), and “smooth” (glossy, finely granular or very finely pitted) in all the other genera (Fig. 1A–D). Two species (Alycaeusconformis and A.gibbosulus) exhibit oblique striae on the protoconch (Fig. 2C; Foon & Liew, 2017). They are placed in Alycaeus because the protoconch of A.rolfbrandti (a species otherwise similar to the type species, A.eydouxi) is finely scaly/tuberculated in oblique lines at the end of the protoconch (Fig. 2B). This sculpture is seemingly an intermediate character state between the smooth (Fig. 2A) and obliquely striated types (Fig. 2C).
Among the genera with usually depressed shell shape (Dicharax, Chamalycaeus, Metalycaeus), the protoconch is also low (depressed) in the majority of Dicharax species. It is, however, often more elevated in the other two genera than what we would expect from the overall low spire. This was not the case for every single species, but generally Chamalycaeus and Metalycaeus possess more elevated protoconch than Dicharax. When the protoconch sculpture is not clearly visible due to corrosion, general differences in the protoconch shape between Dicharax and Chamalycaeus/Metalycaeus may help in generic classification. In the genera with generally higher spire (Alycaeus, Dioryx, Pincerna, Stomacosmethis) the protoconch is as elevated as what we would expect from the high spire.
R1 sculpture
Spiral striation is absent only in Dicharax. All the other genera possess some spiral striae of varying development. These spiral striae consist of microscopic elevated ridges arranged in clearly visible spiral lines. Some spiral striae visible in a few Dicharax species (e.g., D.candrakirana, D.depressus), however, is appear to be a part of the inner shell layers and may not be homologous with those of Chamalycaeus and Metalycaeus. The strength of radial ribbing is also informative (usually strong in Chamalycaeus and Pincerna, weak in Alycaeus and Stomacosmethis). Dioryx has overall weak sculpture, whereas it is highly variable in Metalycaeus.
Length of R2
Stomacosmethis is characterised by very short R1 (with a short, tumid, sometimes pear-shaped tube), whereas Alycaeus possesses very long R1 (ca. 0.5 whorl). Most Pincerna species, especially the ones classified in Cycloryx previously, possess a short tube, but some species have a longer tube than usual for that genus. Distinction between longer-tubed Pincerna and Alycaeus is the most problematic part of the current classification. The remaining genera (Chamalycaeus, Dicharax, Dioryx, Metalycaeus) exhibit high variability in terms of the tube length.
Sculpture of R2
Highly variable within each genus with the exception of Dioryx, which has no elevated R2 ribs. Typically, Chamalycaeus and Metalycaeus species possess widely spaced, sharp ribs. However, Metalycaeusvinctus has widely spaced, sharp ribs, but its putative sister species M.minatoi has a smooth outer surface in R2 without any elevated ribs. This seems to indicate that this trait may differ substantially even between closely related species. Similar examples are found among some Himalayan Metalycaeus species, and also among the Chinese Dicharaxmoellendorffi vs. other Dicharax species. Typical Dicharax species possess R2 ribs that curve towards the aperture.
Development of R3
In most alycaeid genera, except for Dioryx, R3 is strongly developed. The commonly weak shell sculpture may suggest the monophyly of Dioryx. The R3 area is occasionally reduced in other genera also, such as in Chamalycaeusmicroconus, C.mixtus, Dicharaxakioi, and Alycaeusconformis, which are classified in their respective genera based on other characters.
Operculum
The inner side is with or without central nipple. When present, its extent and height may vary between or within species (Páll-Gergely et al. 2017). The outer surface is usually smooth, but can have a closely coiled spiral lamella (in Dicharax and Metalycaeus, see Páll-Gergely et al. 2017), which may result in a circular ring (Dicharaxbison, Metalycaeusnipponensis). It is unknown whether the pipe, tongue or cup-shaped structure in some Pincerna and Stomacosmethis species is homologous with the similarly circular structure of Dicharax and Metalycaeus. Metalycaeus, Pincerna and Stomacosmethis were originally defined on the basis of opercular characters. The outer surface of the operculum can also be finely granulated and flaky with short calcareous spikes or scaffold-like calcareous deposits. These traits are generally (but not always) consistent within each species, making them useful for species recognition (Foon and Liew 2017; Páll-Gergely et al. 2017). Although in some cases opercular characters may suggest relatedness, it does not appear to be useful for subdividing the Alycaeidae into genera. Thus, we do not use opercular traits in our system.
Anatomy
Females of seven species belonging to seven genera were examined. See corresponding locality data under Materials and methods.
Alycaeuseydouxi Venmans, 1956: ovarium elongated, spindle-shaped, bursa copulatrix curved, relatively slender, opens near centre of ovarium, strongly extends beyond ovarium posteriorly, receptaculum seminis small, rounded (Fig. 3B, C).
Alycaeusgibbosulus Stoliczka, 1872: ovarium wide with pointed anterior and blunt posterior end, bursa copulatrix large, extends beyond ovarium posteriorly, opens at middle part of ovarium near its base, bursa has a thickened posterior portion; receptaculum seminis small, oval (Fig. 3D, E).
Female genital anatomy of AlycaeidaeA positioning of females during anatomical examination B, CAlycaeuseydouxi Venmans, 1956 (NHMUK 20160702, V142, specimen5) D, EAlycaeusgibbosulus Stoliczka, 1872 (ALY03) F, GDioryxmessageri (Bavay & Dautzenberg, 1900), NHMUK 20140343 H, IDicharaxtokunoshimanusprincipalis (Pilsbry & Y. Hirase, 1909) HNHM 104428 (2016.10.01B, spec2) J, KMetalycaeusminatoi Páll-Gergely, 2017 HNHM 104427 (2015.12.14A, female28) L, MStomacosmethisdohrni (O. Boettger, 1893) HNHM 104426 N, OStomacosmethisbalingensis (Tomlin, 1948), HNHM 104425 (2016.07.22A, sp2). Abbreviations: B: bursa copulatrix; H: head; O: ovarium; OV: oviduct; P: trematode parasite (found in the pericardium); PC: pericardium; PO: position of the operculum (operculum removed); R: rectum; RS: receptaculum seminis; S: sole. Upper images of each pair (B, D, F, H, J, L, N) shows the genitalia before removing the rectum (with grey shading). Scale bars: 1 mm. Photograph and drawings: Barna Páll-Gergely.
https://binary.pensoft.net/fig/468648
Dioryxmessageri (Bavay & Dautzenberg, 1900): ovarium oval, anterior end pointed, posterior end blunt, bursa copulatrix relatively small, oval, strongly extends beyond ovarium posteriorly, its stalk slender, opens posterior to centre of ovarium, receptaculum seminis elongate (Fig. 3F, G).
Dicharaxtokunoshimanusprincipalis (Pilsbry & Y. Hirase, 1909): Ovarium elongated, pointed posteriorly, bursa copulatrix relatively slender with blunt bursa, does not extend beyond ovarium, receptaculum seminis strongly elongated (Fig. 3H, I).
Metalycaeusminatoi Páll-Gergely, 2017: ovarium slender, pointed anteriorly and rounded posteriorly, bursa copulatrix, rounded, does not extend beyond ovarium, opens near opening of ovarium, receptaculum seminis small, rounded (Fig. 3J, K).
Stomacosmethisdohrni (O. Boettger, 1893): shape of ovarium could not be examined due to its decayed condition, but it is probably oval, bursa copulatrix large, elongate, strongly extends beyond ovarium posteriorly, opens at centre of ovarium, receptaculum seminis small, oval (Fig. 3L, M).
Stomacosmethisbalingensis (Tomlin, 1948): ovarium peanut-shaped, bursa copulatrix was damaged, its posterior part could not be dissected out, opens at anterior part of ovarium, near ovarium opening, receptaculum seminis elongated, a complicated spermoviduct was found in bursa copulatrix: its head is pointed drop-shaped, both ends of the head connected to a slender stalk that forms a flattened loop, the entire length of the stalk is continuous, forming a ring (Fig. 3N, O).
Our knowledge of genital anatomy of terrestrial operculate snails is far more limited than that of pulmonates, probably because dissection of the soft body is more difficult. Firstly, the reproductive organs are not so clearly separated as in pulmonates, but are attached to neighbouring tissues and organs. Secondly, tissues of ethanol-preserved animals are far more fragile. Therefore, it is much more difficult to see the boundaries and junctions of certain organs. For the current study much of the ethanol-preserved material was not in a suitable condition for reproductive anatomy. More than half of the available material was not used for this reason.
So far, the reproductive anatomy of the Alycaeidae is little known. Tielecke (1940) published a few notes without figures on two alycaeid taxa. Although we have dissected a few female specimens, our observations of reproductive anatomy are insufficient to feed into our classification between genera. Considerable differences could be observed in the relative size of the bursa copulatrix, thickness and origin of the bursa’s stalk, shape of the bursa and receptaculum seminis. The taxonomic value of these traits must be clarified by further observations. Nevertheless, the bursa copulatrix originates from the lateral side of the ovarium, which may probably be a synapomorphic character of the Alycaeidae. In contrast, the bursa starts from the terminal (distal) end of the ovarium in all the anatomically examined specimens of Cyclophoridae (Tielecke 1940).
Radula
Radulae of nine species belonging to five genera were examined: Alycaeuseydouxi Venmans, 1956 (Fig. 4A), Alycaeusgibbosulus Stoliczka, 1872 (Fig. 4B), Dioryxmessageri (Bavay & Dautzenberg, 1900) (Fig. 4C), Chamalycaeus sp. (Fig. 4D, Suppl. material 1: Fig. S1), Chamalycaeuseveretti (Godwin-Austen, 1889) (Fig. 4E), Dicharaxitonis (Kuroda, 1943) (Fig. 4F), Dicharax(?)okinawaensis (Uozumi, Yamamoto & Habe, 1979) (Fig. 4G), Stomacosmethisbalingensis (Tomlin, 1948) (Fig. 4H), Stomacosmethisperakensis (Crosse, 1879) (Fig. 4I). See corresponding locality data under Materials and methods. For descriptive note on the radular traits, see Table 1 and Fig. 4.
Radula traits of Alycaeidae. For the species examined by Venmans (1956) see Radula under “Description and assessment of morphological characters” (page 14).
Taxon
Morphology of central tooth
Reference
Alycaeusconformis
see remarks
Venmans 1956
Alycaeuseydouxi
see remarks
Venmans 1956
Alycaeuseydouxi
5 cusps, broad, central cusp blunt
this study
Alycaeusgibbosulus
5 cusps, broad, central cusp blunt
this study
Chamalycaeuseveretti
5 cusps, broad, central cusp pointed
this study
Dicharaxalticola
5 cusps, broad, central cusp pointed
Páll-Gergely et al. 2017
Dicharaxananensis
5 cusps, broad, central cusp pointed
Yano et al. 2013
Dicharaxbicrenatus
5 cusps, broad, central cusp pointed
Godwin-Austen 1884 (1882–1920)
Dicharaxcristatus
5 cusps, broad, central cusp pointed
Páll-Gergely et al. 2017
Dicharaxdepressus
5 cusps, broad, central cusp pointed
Páll-Gergely et al. 2017
Dicharaxfimbriatus
5 cusps, broad, central cusp pointed
Páll-Gergely et al. 2017
Dicharaximmaculatus
5 cusps, broad, central cusp pointed
Páll-Gergely et al. 2017
Dicharaxitonis
5 cusps, broad, central cusp pointed
this study
Dicharaxlongituba
5 cusps, broad, central cusp pointed
Benthem-Jutting 1948
Dicharaxokinawaensis
5 cusps, broad, central cusp pointed
this study
Dicharaxplanorbulus
5 cusps, broad, central cusp pointed
Heude 1882–1890
Pincernamaolanensis
7 cusps, broad, central cusp pointed
Luo et al. 2009
Dioryxmessageri
7 cusps, broad, central cusp blunt
this study
Dioryxsetchuanensis
5 cusps, broad, central cusp blunt
Heude 1882–1890
Metalycaeusminatoi
5 cusps, broad, central cusp pointed
Páll-Gergely & Asami, 2017
Metalycaeusvinctus
5 cusps, broad, central cusp pointed
Páll-Gergely & Asami, 2017
Metalycaeuszayuensis
5 cusps, broad, central cusp pointed
Zhang et al. 2008
Pincernathieroti
see remarks
Venmans 1956
Pincernayanseni
5 cusps, broad, central cusp pointed
Páll-Gergely 2017
Stomacosmethisbalingensis
3 cusps, elongated, central cusp pointed
this study
Stomacosmethishochstetteri (synonym of jagori)
3 cusps, elongated, central cusp pointed
Bollinger 1918
Stomacosmethisjagori
3 cusps, elongated, central cusp pointed
Thiele 1929
Stomacosmethisjagori
1 cusp, elongated, central cusp pointed
Sarasin & Sarasin, 1899
Stomacosmethiskapayanensisselangoriensis
see remarks
Venmans 1956
Stomacosmethiskuekenthali
1 cusp, elongated, central cusp pointed
Sarasin & Sarasin, 1899
Stomacosmethisperakensis
3 cusps elongated, central cusp pointed
this study
Stomacosmethisporcilliferus
3 cusps elongated, central cusp pointed
Bollinger 1918
Stomacosmethissarasinorum
5 cusps, elongated, central cusp pointed
Bollinger 1918
Dicharax(?)panshiensis
6 cusps ?, broad, central cusp pointed
Chen 1989
Venmans (1956) described the radular morphology of four alycaeid species. The structure of the radula in Alycaeuseydouxi and S.kapayanensis (specimens were collected as Batu Caves, which is the type locality of S.kapayanensisselangoriensis, see Foon and Liew 2017) are especially interesting. Venmans (1956) published drawings of the radula of a specimen of Alycaeuseydouxi which had an elongated, spatula-like central tooth without any side cusps. Our observations of the same species from the same locality were, however, strikingly different. The specimens we examined had a blunt central tooth with a blunt, central cusp, and four pointed side-cusps. In order to confirm the identification of Venmans, we examined his specimens. We confirmed that they were indeed A.eydouxi shells. Furthermore, so far, the only Stomacosmethis radula with a blunt central tooth with cusps has been that of S.kapayanensis figured by Venmans (1956). It is highly likely that Venmans mixed the radular drawings or radulae of those two species. Therefore, in our analysis of alycaeid radulae we ignore the data of Venmans (1956).
The radula morphology of 28 species are known from the available literature and this study (excluding the results of Venmans 1956). In every species the radular teeth are arranged in v-shaped rows, each transverse row with seven teeth (2-1-1-1-2). The central tooth is strongly constricted in its middle part. The lateral and two marginal teeth have slighter median constriction of the plates and are seemingly longer and slenderer than the central tooth, except for Stomacosmethis, where they are of comparable width to the central teeth. The lateral and marginal teeth are similar in terms of shape of the cusps to the central teeth in all examined specimens.
There are generally two types of central teeth. One type has one round tooth with 5–7 cusps, and the other is elongated with 1–5 (usually 1–3) cusps. The former type is common among all the genera (Alycaeus, Chamalycaeus, Dicharax, Dioryx, Metalycaeus, Pincerna), whereas the latter, elongated type has only been observed in Stomacosmethis species. The placement of S.balingensis in Stomacosmethis is mainly based on its radular morphology, which is similar to that of sympatric S.perakensis. The first type of central teeth can be further subdivided into groups with a blunt (Alycaeus, Dioryx) or pointed (Chamalycaeus, Dicharax, Metalycaeus, Pincerna) central cusp. However, a Chamalycaeus species we examined had a blunt central cusp. The type with a pointed central cusp is a probably plesiomorphic character, which is visible in many terrestrial caenogastropods (e.g., Cyclophorus, Cyclotus, Japonia, see Egorov 2009). Together with conchological, anatomical and molecular phylogenetic information the radular traits may provide insights about relationships of alycaeid genera (see Concluding remarks).
Genus-level diversity
Of the 14 nominal genus-group taxa that have been described (Table 2), we accept seven. The classification proposed in this study is based on unique character states (Dicharax, Dioryx, Metalycaeus) and on unique combinations of character states (other genera) (Table 3). The number of accepted species-level taxa is: Alycaeus: 7, Chamalycaeus: 26, Dicharax: 164, Dioryx: 30, Metalycaeus: 61, Pincerna: 37, Stomacosmethis: 37.
Genus-group taxa of the Alycaeidae. Abbreviations: M: monotypy, OD: original designation, SD: subsequent designation. Valid genera are marked with an asterisk.
shell very small to medium sized, usually depressed, R2 of variable length
Pincerna
Dicharax
smooth, usually low
variable
1–11
5–7 cusps, broad, central cusp pointed
protoconch + teleoconch without spiral striae
Dioryx
smooth
variable
3.5–9
5–7 cusps, broad, central cusp blunt
shell globular or high-spired, sculpture reduced, R3 absent
Metalycaeus
spirally striated, elevated
variable
3–10
5 cusps, broad, central cusp pointed
protoconch spirally striated
Pincerna
smooth
very short to short
2.5–6
5 cusps, broad, central cusp pointed
shell very small to medium sized, high spired, R2 short
Alycaeus, Chamalycaeus, Stomacosmethis
Stomacosmethis
smooth
very short
3–13
elongated, usually with 1 central cusp only, or central cusps with 1–2 small cusp at each side
tiangular, colourful shell, R2 very short
Pincerna
Besides taxonomic problems at the species level (see under Annotated list of species group taxa), some aspects of grouping species into genera turned out to be especially challenging. As a result, the generic boundaries are not completely clear. This may be due to repeated evolution of morphological traits and the presence of the large numbers of species in the genera. In the absence of phylogenetic analyses, the current classification is tentative. We anticipate that some adjustments are inevitable as our understanding of the evolutionary history of this family improves. Some remarks about the species boundaries (numbers correspond with Fig. 5):
(1) Metalycaeus is characterised by the presence of spirally striated protoconch. However, in rare cases the striation is strongly reduced (see M.laevis).
(2) Dicharax is characterised by the absence of spiral striation on both the protoconch and teleoconch. However, striation is present in a few Dicharax species (see remarks under D.candrakirana, and D.depressus). Striae are not elevated threads but are probably a part of the inner shell layers; this structure would not be homologous with the striae present in the other genera. Dicharax is a diverse genus containing 166 species and subspecies. However, meaningful subdivision is not possible at present.
(3) Dioryx and Alycaeus are clearly separated based on the presence of R3, globular shape and weak sculpture of Dioryx. Although the R3 in Alycaeusconformis renders its shell somewhat similar to the shell of Dioryx species, we consider Dioryx to be a recognisable group, which would be monophyletic.
(4) Distinction between Alycaeus and Pincerna is probably the most problematic issue in the system presented here. The genus Cycloryx (treated as a synonym of Pincerna) originally contained species with a very small shell, which has elevated spire usually with strong ribs and very short tube, and taxa from the Himalaya region and northern Myanmar. However, similar species are disjunctly found from Sumatra (Pincernayanseni), northern Vietnam (A.costulosus) and Borneo (A.globosus). For example, the shell of P.yanseni is hardly distinguishable from that of Hymalayan taxa. The shells of Pincernaliratula and P.thieroti in the Malay Peninsula may be larger than typical Cycloryx, and their sutural tubes may be longer than the extremely short tube of Cycloryx. These two species, however, seem to belong to the same group as Cycloryx according to their strong ribbing, generally short tube and high spire. Furthermore, A.vanbuensis and A.costulosus both of which occur in northern Vietnam, are only distinguishable in tube length. The former is similar to the type species of Pincerna, and the latter to Himalayan species of Cycloryx. Recognition of Pincerna and Cycloryx as different genera requires to classify A.vanbuensis and A.costulosus into those genera on the basis of the tube length, which would not be acceptable. For these reasons, Cycloryx has been treated as a synonym of Pincerna (see Páll-Gergely 2017). Alycaeusmouhoti has a smoother and larger shell with the simple (not double) peristome than otherwise very similar A.vanbuensis and is also similar to Alycaeuseydouxi (type species of Alycaeus) possessing commonly large and similarly shaped shell with a rather long tube. Alycaeusmouhoti has shell characteristics that connect Alycaeus and Pincerna. Thus, a morphological continuum is present from A.eydouxi, A.mouhoti and A.vanbuensis to A.costulosus, which looks like a typical Cycloryx species. Generic subdivision on this basis would inflate the genus Alycaeus enormously, and A.eydouxi was separated from Pincerna and Stomacosmethis in our molecular phylogeny. Therefore, in this revision by a conservative approach, we included only those species in Alycaeus that are similar to the type species in terms of the very large shell, extremely long sutural tube (and R2) and strongly inflated body whorl.
(5) Chamalycaeus species have a depressed shell with reticulated sculpture and a sutural tube and R2 which vary in length. Pincerna species have higher spire, sculpture dominated by radial ribs and sometimes a short tutural tube. Pincernacrenilabris seemingly connect the two genera by its rather globular shell and medium-sized tube.
(6) The genus Stomacosmethis is characterised by colorful triangular shell mostly with flat whorls and a very short tube. Pincerna species from the distributional range of Stomacosmethis differ by having round whorls and a short tube. All Stomacosmethis species that were examined for radular traits possess unique, simple teeth (Fig. 4H, I). However, a similar type of radula has been found in A.balingensis, which could be classified as Pincerna based on the shell shape (round whorls). For this reason, A.balingensis is here moved to Stomacosmethis. This example suggests that Pincerna and Stomacosmethis might not be mutually monophyletic. Further examinations are necessary to verify the generic position of Pincerna species.
(7) One of the most important results of this study is discovery of distinct differences between Alycaeus and Stomacosmethis, which form two groups without taxa that exhibit overlapping traits of morphology.
Relationships between alycaeid genera. Solid lines indicate partly unclear generic borders. See explanation in the text.
https://binary.pensoft.net/fig/468650Annotated list of species-group taxa
In this study we list 412 species-group names including eleven replacement names (seven of them proposed in this study) and five nomina nuda (Table 4). Types of 336 species and subspecies (85%) were examined, and of 19 taxa non-type specimens were examined (5%), whilst we relied on the sufficiently detailed original descriptions of the 22 taxa (6%). For 17 taxa (4%) no material was available to be examined in this study.
List of all alycaeid species-group names. Subgenera are treated at the same level as genera in the “previous classification”. Previous classification does not include our papers (Páll-Gergely 2017, Páll-Gergely & Asami 2017, Páll-Gergely et al. 2017, Páll-Gergely & Auffenberg 2019). Abbreviations for species examined: DOD: detailed original description was sufficient for generic placement; NE: not examined; NT: non-type material; T: types.
Taxon
Previous classification
This study
Remarks
Rank (species or subspecies)
Specimens examined
abdoui
Dicharax
Dicharax
sp
T
abei
Awalycaeus
Dicharax
sp
NT
aborensis
Chamalycaeus
Metalycaeus
sp
T
akhaensis
Raptomphalus
Dicharax
sp
T
akioi
Cipangocharax
Dicharax
sp
T
akiratadai
Awalycaeus
Dicharax
sp
T
akyabensis
Alycaeus
Dicharax
ssp
T
alticola Foon & Liew, 2017
Alycaeus
Stomacosmethis
ssp
DOD
alticola Páll-Gergely & Hunyadi, 2017
Dicharax
Dicharax
sp
T
altispirus
Alycaeus
Stomacosmethis
sp
T
amphora
Dioryx
Dioryx
sp
T
ananensis
Cipangocharax
Dicharax
sp
DOD
anapetes
nomen nudum
anceyi
Alycaeus
Pincerna
sp
T
andamaniae
Chamalycaeus
Chamalycaeus
sp
T
anghamiensis
Dioryx
Dioryx
ssp
T
anonymus
Alycaeus
Dicharax
sp
T
anthostoma
Dicharax
Dicharax
sp
T
armillatus
Dicharax
Chamalycaeus
sp
T
asaluensis
Dicharax
Dicharax
sp
T
ataranensis
Dicharax
Dicharax
sp
T
avae
Dicharax
Dicharax
sp
T
awaensis
Chamalycaeus
Metalycaeus
sp
T
awalycaeoides
Metalycaeus
Metalycaeus
sp
T
bacca
Dioryx
Dioryx
sp
T
balingensis
Alycaeus
Stomacosmethis
sp
T
barowliensis
Alycaeus
Dicharax
sp
T
bawai
Dicharax
Dicharax
sp
T
beddomei
Alycaeus
Metalycaeus
sp
T
bembex
Cycloryx
Pincerna
sp
T
bhutanensis
Chamalycaeus
Metalycaeus
sp
T
bicrenatus
Dicharax
Dicharax
sp
T
biexcisus
Cipangocharax
Dicharax
sp
T
bifrons
Dicharax
Dicharax
sp
T
birugosus
Dicharax
Dicharax
sp
T
bison
Dicharax
Dicharax
sp
T
blanfordi
Alycaeus
Dicharax
sp
T
brahma
Chamalycaeus
Metalycaeus
sp
T
broti
nomen nudum
burrailensis
Cycloryx
Pincerna
sp
T
burroiensis
Cycloryx
Dicharax
sp
T
burtii
Alycaeus
Metalycaeus
sp
T
busbyi
Chamalycaeus
Chamalycaeus
sp
T
calopoma
Alycaeus
Stomacosmethis
sp
DOD: detailed original description was sufficient for generic placement; NE
canaliculatus
Chamalycaeus
Chamalycaeus
sp
T
canaliculus
Chamalycaeus
Dicharax
junior synonym of birugosus
sp
T
candrakirana
Dicharax
Dicharax
sp
DOD
carinatus
Alycaeus
Stomacosmethis
sp
NT
cariniger
Dioryx
Dioryx
sp
T
caroli
Chamalycaeus
Metalycaeus
sp
T
caudapiscis
Dicharax
Dicharax
sp
T
celebensis
Chamalycaeus
Chamalycaeus
sp
T
chanjukensis
Alycaeus
Metalycaeus
junior synonym of brahma
sp
T
chaperi
Alycaeus
Alycaeus
junior synonym of gibbosulus
sp
DOD: detailed original description was sufficient for generic placement; NE
charasensis
Alycaeus
Stomacosmethis
ssp
DOD
chennelli
Dicharax
Dicharax
sp
T
christae
Alycaeus
Stomacosmethis
sp
T
clementsi
Alycaeus
Stomacosmethis
ssp
DOD
cochinensis
Dioryx
Dioryx
sp
T
commutatus
Raptomphalus
Metalycaeus
junior synonym of brahma
sp
T
compactus
Dioryx
Dioryx
sp
T
compressicosta
Dicharax
Metalycaeus
junior synonym of heudei
syn
T
conformis
Alycaeus
Alycaeus
sp
T
congener
Alycaeus
Stomacosmethis
sp
T
conicus
Alycaeus
Dicharax
sp
T
constrictus
Cycloryx
Pincerna
sp
T
costacrassa
Alycaeus
Stomacosmethis
ssp
DOD
costata
Cycloryx
Pincerna
sp
T
costulosa
Alycaeus
Pincerna
sp
T
crassicollis
Chamalycaeus
Dicharax
sp
DOD: detailed original description was sufficient for generic placement; NE
crassus
Dicharax
Dicharax
ssp
T
crenatus
Dicharax
Dicharax
sp
T
crenilabris
Alycaeus
Pincerna
sp
T
crenulatus
Dicharax
Metalycaeus
sp
T
crispatus
Dicharax
Dicharax
sp
T
cristatus
Dicharax
Dicharax
sp
T
cucullatus
Dicharax
Dicharax
sp
T
cyclophoroides
Chamalycaeus
Dicharax
sp
T
cyphogyrus
Chamalycaeus
Metalycaeus
sp
T
daflaensis Godwin-Austen, 1876
Dicharax
Dicharax
sp
T
daflaensis Godwin-Austen, 1914
Dioryx
Dioryx
replaced by Dioryxurnulaniosiensis nom. nov.
ssp
T
dalingensis
Dicharax
Dicharax
sp
T
damsangensis
Dicharax
Dicharax
sp
T
dautzenbergi
Dioryx
Dioryx
replacement name for major Bavay & Dautzenberg, 1900
sp
T
davisi
Chamalycaeus
Dicharax
sp
T
degenerans
Alycaeus
Stomacosmethis
ssp
T
depressus
Alycaeus
Dicharax
sp
T
diagonius
Dicharax
Dicharax
sp
T
difficilis
Cycloryx
Pincerna
sp
T
digitatus
Dicharax
Dicharax
sp
T
dihingensis
Cycloryx
Pincerna
ssp
T
dikrangensis
Alycaeus
Metalycaeus
sp
T
diminutus
Chamalycaeus
Dicharax
sp
T
diplochilus
Chamalycaeus
Dicharax
sp
T
distinctus
Chamalycaeus
Metalycaeus
sp
T
distortus
Dioryx
Dioryx
sp
T
ditaceus
Sigmacharax
Dicharax
ssp
T
diyungensis
Dicharax
Dicharax
ssp
T
dohertyi
Dicharax
Dicharax
sp
DOD: detailed original description was sufficient for generic placement; NE
dohrni
Alycaeus
Stomacosmethis
sp
T
dolichodeiros
Chamalycaeus
Dicharax
sp
DOD: detailed original description was sufficient for generic placement; NE
dolomiticus
Alycaeus
Metalycaeus
junior synonym of rathouisianus
syn
T
dongiensis
Dioryx
Dioryx
sp
T
draco
Dicharax
Dicharax
sp
T
duoculmen
Raptomphalus
Dicharax
sp
T
duorugosus
Dicharax
Dicharax
sp
T
duplicatus
Chamalycaeus
Dicharax
sp
T
edei
Chamalycaeus
Dicharax
sp
T
elegans
Cycloryx
Pincerna
sp
T
elevatus
Alycaeus
Dicharax
sp
T
ellipticus
Dicharax
Dicharax
sp
T
everetti
Dicharax
Chamalycaeus
sp
T
excisus
Chamalycaeus
Chamalycaeus
sp
T
expanstoma
Chamalycaeus
Dicharax
sp
T
expansus Foon & Liew, 2017
Alycaeus
Stomacosmethis
ssp
DOD
expansus Heude, 1890
Alycaeus
Metalycaeus
junior synonym of muciferus
syn
T
expatriatus
Chamalycaeus
Dicharax
sp
T
eydouxi
Alycaeus
Alycaeus
replacement name for C.gibbum Draparnaud, 1805
sp
NT
fargesianus
Chamalycaeus
Dicharax
sp
T
feddenianus
Dioryx
Dioryx
sp
NT
fimbriatus
Dicharax
Dicharax
sp
T
footei
Dicharax
Dicharax
sp
T
fractus
Dicharax
Metalycaeus
junior synonym of heudei
syn
T
fraterculus
Dicharax
Dicharax
sp
T
fruhstorferi
Chamalycaeus
Chamalycaeus
sp
T
fultoni
Alycaeus
Stomacosmethis
sp
T
galbanus
Alycaeus
Stomacosmethis
sp
T
gemma
Dicharax
Dicharax
sp
T
gemmula
Dicharax
Dicharax
sp
T
generosus
Cycloryx
Dicharax
sp
T
gibbosulus
Alycaeus
Alycaeus
sp
T
gibbus
Alycaeus
Alycaeus
replaced by eydouxi
sp
NT
glaber
Dicharax
Dicharax
sp
T
globosus Godwin-Austen, 1914
Dioryx
Dioryx
replaced by rotundus nom. nov.
ssp
T
globosus H. Adams, 1870
Alycaeus
Pincerna
sp
T
globuloides
Dioryx
Dioryx
sp
T
globulosus
Dioryx
Dioryx
sp
T
globulus
Dicharax
Dicharax
sp
T
godwinausteni nom. nov.
Alycaeus
Metalycaeus
nom. nov. pro A.neglectus Godwin-Austen, 1914
sp
T
granum
Cycloryx
Pincerna
sp
T
graphiaria
Cycloryx
Pincerna
sp
T
graphica
Cycloryx
Pincerna
sp
T
habiangensis
Dicharax
Dicharax
sp
T
harimensis
Chamalycaeus
Dicharax
junior synonym of japonicus
syn
T
hebes
Dicharax
Dicharax
sp
NT
helicodes
Alycaeus
Metalycaeus
junior synonym of muciferus
syn
T
heudei
Dicharax
Metalycaeus
sp
T
hirasei
Chamalycaeus
Metalycaeus
sp
T
hochstetteri
Alycaeus
Stomacosmethis
junior synonym of jagori
syn
DOD: detailed original description was sufficient for generic placement; NE
hosei
Alycaeus
Stomacosmethis
sp
NT
huberi
Alycaeus
Stomacosmethis
junior synonym of somnueki
syn
T
humilis
Dicharax
Dicharax
sp
NT
hungerfordianus
Chamalycaeus
Metalycaeus
sp
T
ibex
Metalycaeus
Metalycaeus
sp
T
ikanensis
Alycaeus
Stomacosmethis
ssp
DOD
imitator
Dicharax
Dicharax
sp
T
immaculatus
Dicharax
Dicharax
sp
T
inflatus Godwin-Austen, 1874
Chamalycaeus
Metalycaeus
sp
T
inflatus Möllendorff, 1886
Chamalycaeus
Dicharax
replaced by moellendorffi
sp
T
ingrami
Chamalycaeus
Dicharax
sp
T
itonis
Sigmacharax
Dicharax
sp
NT
jagori
Alycaeus
Stomacosmethis
sp
T
japonicus
Chamalycaeus
Dicharax
sp
NT
jatingaensis nom. nov.
Alycaeus
Dicharax
nom. nov. pro A.nanus Godwin-Austen, 1914
ssp
T
jaintiacus
Dicharax
Dicharax
sp
T
jousseaumei
Chamalycaeus
Alycaeus
sp
T
juttingae nom. nov.
Alycaeus
Pincerna
nom. nov. pro laevis van Benthem Jutting, 1959
ssp
DOD: detailed original description was sufficient for generic placement; NE
kamakiaensis
Alycaeus
Metalycaeus
sp
T
kapayanensis
Alycaeus
Stomacosmethis
sp
T
kelantanensis
Alycaeus
Stomacosmethis
sp
T
kengtungensis
Raptomphalus
Metalycaeus
sp
T
kessneri
Chamalycaeus
Chamalycaeus
sp
T
kezamaensis
Dicharax
Dicharax
sp
T
khasiacus
Dicharax
Dicharax
sp
T
khunhoensis
Cycloryx
Pincerna
sp
T
kinabaluana
Alycaeus
Pincerna
ssp
T
kiuchii
Cipangocharax
Dicharax
sp
T
kobeltianus
Dioryx
Dioryx
sp
T
korintjiensis nom. nov.
Alycaeus
Pincerna
nom. nov. pro latecostata van Benthem Jutting, 1959
ssp
DOD: detailed original description was sufficient for generic placement; NE
koshuensis
Chamalycaeus
Dicharax
ssp
NT
kuekenthali
Chamalycaeus
Stomacosmethis
sp
T
kurauensis
Alycaeus
Stomacosmethis
ssp
DOD
kurodai
Metalycaeus
Dicharax
junior synonym of spiracellum
syn
T
kurodatokubeii
Chamalycaeus
Dicharax
sp
T
kurzianus
Dicharax
Dicharax
sp
T
labrirubidum
Dioryx
Dioryx
sp
T
laevicervix
Chamalycaeus
Metalycaeus
ssp
T
laevis Pilsbry & Y. Hirase, 1909
Chamalycaeus
Metalycaeus
sp
T
laevis van Benthem Jutting, 1959
Alycaeus
Pincerna
replaced by juttingae nom. nov.
ssp
DOD: detailed original description was sufficient for generic placement; NE
lahupaensis
Raptomphalus
Dicharax
sp
T
laosensis
Metalycaeus
Metalycaeus
sp
T
Pincernalatecostata van Benthem Jutting, 1959
Alycaeus
Pincerna
replaced by korintjiensis nom. nov.
ssp
DOD: detailed original description was sufficient for generic placement; NE
latecostatus Möllendorff, 1882
Chamalycaeus
Metalycaeus
sp
T
latestriata
nomen nudum
lectus
Dicharax
Dicharax
sp
T
lenticulus
Dicharax
Dicharax
sp
T
levis
Alycaeus
Dicharax
sp
T
libonensis
Chamalycaeus
Metalycaeus
sp
T
liratula
Pincerna
Pincerna
sp
T
logtakensis
Alycaeus
Dicharax
sp
T
lohitensis
Alycaeus
Metalycaeus
sp
T
longituba
Alycaeus
Dicharax
sp
NT
luyorensis
Raptomphalus
Metalycaeus
sp
T
macgregori
Chamalycaeus
Metalycaeus
sp
T
magnificus
Raptomphalus
Metalycaeus
sp
T
magnus
Alycaeus
Dicharax
sp
T
major
Dioryx
Dioryx
replaced by dautzenbergi
sp
T
major (granum var.)
Cycloryx
Pincerna
senior synonym of A.mangutensis Godwin-Austen, 1914
sp
T
makarsae
Dicharax
Dicharax
ssp
T
mangutensis
Cycloryx
Pincerna
junior synonym of A.granumvar.major Godwin-Austen, 1893
sp
T
maolanensis
Dioryx
Pincerna
sp
T
maosmaiensis
Alycaeus
Dicharax
sp
T
margarita
Cycloryx
Pincerna
sp
T
matchacheepiorum
Alycaeus
Stomacosmethis
sp
DOD
mediocris
Alycaeus
Dicharax
ssp
T
melanopoma
Metalycaeus
Metalycaeus
junior synonym of nipponensis
syn
T
menglunensis
Dioryx
Dioryx
sp
T
messageri
Dioryx
Dioryx
sp
T
microconus
Chamalycaeus
Chamalycaeus
sp
T
microcostatus
Dicharax
Dicharax
sp
T
microdiscus
Chamalycaeus
Dicharax
sp
T
micropolitus
Dicharax
Dicharax
sp
T
microstoma
Alycaeus
Alycaeus
junior synonyom of sculptilis
syn
DOD: detailed original description was sufficient for generic placement; NE
minatoi
Metalycaeus
Metalycaeus
sp
T
minimus
Dicharax
Dicharax
ssp
T
minor (birugosus var.)
Dicharax
Dicharax
junior synonym of birugosus
syn
T
minor (constrictus var.)
Cycloryx
Pincerna
junior synonym of constrictus
syn
DOD: detailed original description was sufficient for generic placement; NE
minor (graphicus var.)
Cycloryx
Pincerna
junior synonym of graphicus
syn
DOD: detailed original description was sufficient for generic placement; NE
minor (jagori var.)
Alycaeus
Stomacosmethis
junior synonym of jagori
syn
DOD: detailed original description was sufficient for generic placement; NE
minor (paviei var.)
Alycaeus
Metalycaeus
junior synonym of heudei
syn
T
minor (pilula var.)
Dioryx
Dioryx
junior synonym of pilula
syn
DOD: detailed original description was sufficient for generic placement; NE
minor (vestitus var.)
Alycaeus
Dicharax
junior synonym of vestitus
syn
DOD: detailed original description was sufficient for generic placement; NE
mixtus
Chamalycaeus
Chamalycaeus
sp
T
miyazakii
Chamalycaeus
Dicharax
sp
NT
moellendorffi
Chamalycaeus
Dicharax
replacement name for inflatus Möllendorff, 1886
sp
T
monadicus
Dioryx
Dioryx
sp
T
montanus
Chamalycaeus
Dicharax
sp
T
mouhoti
Alycaeus
Pincerna
sp
T
muciferus
Chamalycaeus
Metalycaeus
sp
T
multicostulata
Cycloryx
Pincerna
sp
T
multidentatus
junior synonym of fimbriatus
syn
T
multirugosus
Dicharax
Dicharax
sp
T
muluana
Alycaeus
Pincerna
ssp
T
mundulus
Alycaeus
Metalycaeus
sp
T
muroharai
Chamalycaeus
Dicharax
ssp
T
muspratti
Raptomphalus
Dicharax
sp
T
mutatus
Dicharax
Dicharax
sp
T
nagaensis
Chamalycaeus
Dicharax
sp
T
nakashimai
Sigmacharax
Dicharax
sp
T
nanus Godwin-Austen, 1914
Alycaeus
Dicharax
replaced by jatingaensis nom. nov.
ssp
T
nanus Möllendorff, 1886
Chamalycaeus
Dicharax
junior synonym of diminutus
syn
T
nattoungensis
Alycaeus
Dicharax
ssp
T
neglectus Godwin-Austen, 1914
Alycaeus
Metalycaeus
replaced by godwinausteni nom. nov.
sp
T
neglectus Heude, 1885
Chamalycaeus
Metalycaeus
junior synonym of rathouisianus
syn
T
nicobaricus
Alycaeus
Alycaeus
junior synonym of reinhardti
syn
DOD: detailed original description was sufficient for generic placement; NE
niosiensis nom. nov.
Dioryx
Dioryx
nom. nov. pro Alycaeusdaflaensis Godwin-Austen, 1914
ssp
T
nipponensis
Chamalycaeus
Metalycaeus
sp
T
nishii
Chamalycaeus
Dicharax
sp
T
nitidus
Chamalycaeus
Dicharax
sp
T
nongtungensis
Dicharax
Dicharax
sp
T
notatus
Dicharax
Dicharax
sp
T
notus
Dicharax
Dicharax
sp
T
nowgongensis
Alycaeus
Dicharax
sp
T
oakesi
Raptomphalus
Metalycaeus
sp
T
obscurus
Dicharax
Dicharax
sp
T
ochraceus
Dicharax
Dicharax
sp
T
oglei
Alycaeus
Chamalycaeus
sp
T
oharai
Metalycaeus
Metalycaeus
sp
T
okamurai
Cipangocharax
Dicharax
sp
NT
okinawaensis
Chamalycaeus
Dicharax
sp
NT
okuboi
Metalycaeus
Metalycaeus
sp
T
oligopleuris
Chamalycaeus
Dicharax
sp
T
omissus
Chamalycaeus
Dicharax
sp
T
oshimanus
Chamalycaeus
Dicharax
sp
T
otiphorus
Cycloryx
Pincerna
sp
NT
pachitaensis
Dicharax
Dicharax
sp
T
panggianus
Alycaeus
Metalycaeus
sp
T
panshiensis
Chamalycaeus
Dicharax
sp
DOD: detailed original description was sufficient for generic placement; NE
parvulus
Chamalycaeus
Dicharax
sp
T
paucicostata
Cycloryx
Pincerna
sp
T
paviei
Alycaeus
Metalycaeus
junior synonym of heudei
syn
T
peilei
Dicharax
Dicharax
sp
T
pentagonus
Alycaeus
Dicharax
junior synonym of anthostoma
syn
T
perakensis
Alycaeus
Stomacosmethis
sp
T
perplexus
Alycaeus
Chamalycaeus
sp
T
physis
Alycaeus
Metalycaeus
sp
NT
pilsbryi
Chamalycaeus
Chamalycaeus
synonym of japonicus
sp
T
pilula
Dioryx
Dioryx
sp
NT
pingoungensis
Dioryx
Dioryx
sp
T
pisum
Dioryx
Dioryx
ssp
T
placenovitas
Cipangocharax
Dicharax
sp
T
planorbulus
Chamalycaeus
Dicharax
sp
T
plectocheilus
Dicharax
Dicharax
sp
T
plicilabris
Chamalycaeus
Dicharax
sp
T
pocsi
Dioryx
Dioryx
sp
T
politus
Alycaeus
Dicharax
sp
T
polygonoma
Dicharax
Metalycaeus
sp
T
porcilliferus
Alycaeus
Stomacosmethis
sp
T
praetextus
Alycaeus
Stomacosmethis
sp
T
pratatensis
Alycaeus
Dicharax
sp
T
principalis
Chamalycaeus
Dicharax
ssp
T
prosectus
Dicharax
Metalycaeus
sp
T
purus
Chamalycaeus
Dicharax
sp
T
pusillus
Alycaeus
Dicharax
sp
T
pygmaea
Alycaeus
Pincerna
ssp
T
pyramidalis
Alycaeus
Alycaeus
sp
T
quadrasi
Chamalycaeus
Metalycaeus
sp
T
rabongensis
Alycaeus
Pincerna
ssp
T
rarus
Chamalycaeus
Chamalycaeus
sp
T
rathouisianus
Chamalycaeus
Metalycaeus
sp
T
rechilaensis
Dicharax
Dicharax
sp
T
regalis
Alycaeus
Stomacosmethis
sp
DOD
reinhardti Mörch, 1872
Alycaeus
Chamalycaeus
sp
T
reinhardti Pilsbry, 1900
Alycaeus
Dicharax
replaced by pilsbryiKobelt 1902
sp
T
requiescens
Alycaeus
Dioryx
sp
T
reticulatus
Alycaeus
Chamalycaeus
sp
T
richthofeni
Dicharax
Chamalycaeus
sp
T
rimatus
Alycaeus
Stomacosmethis
sp
T
robustus
Dicharax
Dicharax
sp
T
roebeleni
Alycaeus
Stomacosmethis
sp
T
rolfbrandti
Alycaeus
Alycaeus
sp
T
rosea
Dioryx
Dioryx
sp
T
rotundatus
Alycaeus
Metalycaeus
sp
T
rotundus
Dioryx
Dioryx
nom. nov. pro globosus Godwin-Austen, 1914
ssp
T
rubinus
Alycaeus
Metalycaeus
sp
T
rugosus
Dicharax
Metalycaeus
sp
T
ruyangensis
Dioryx
Dioryx
sp
DOD
rywukensis
Dicharax
Dicharax
ssp
T
sabangensis
Alycaeus
Chamalycaeus
ssp
T
sadoensis
Chamalycaeus
Dicharax
ssp
T
sadongensis
Alycaeus
Stomacosmethis
sp
T
sandowayensis
Chamalycaeus
Dicharax
sp
T
sarasinorum
Stomacosmethis
Stomacosmethis
sp
T
satsumanus
Chamalycaeus
Metalycaeus
sp
T
scepticus
nomen nudum
sculptilis
Chamalycaeus
Chamalycaeus
sp
T
sculpturus
Alycaeus
Dicharax
sp
T
selangoriensis
Alycaeus
Stomacosmethis
ssp
DOD
semperi
Metalycaeus
Metalycaeus
sp
T
senyumensis
Alycaeus
Stomacosmethis
sp
DOD
serratus
Alycaeus
Dicharax
sp
T
setchuanensis
Dioryx
Dioryx
sp
T
shiibaensis
Chamalycaeus
Dicharax
sp
T
shiosakimasahiroi
Awalycaeus
Dicharax
sp
DOD
shiotai
Sigmacharax
Dicharax
ssp
T
sibbumensis
Alycaeus
Metalycaeus
sp
T
simplicilabris
Dicharax
Dicharax
junior synonym of cristatus
syn
T
sinensis
Chamalycaeus
Metalycaeus
sp
T
smithi
Chamalycaeus
Dicharax
junior synonym of cristatus
syn
T
solidus
Dicharax
Dicharax
ssp
T
somnueki
Alycaeus
Stomacosmethis
sp
DOD
somwangi
Alycaeus
Alycaeus
sp
DOD
sonlaensis
Alycaeus
Dicharax
sp
DOD
specus
Alycaeus
Chamalycaeus
sp
T
spiracellum
Dicharax
Dicharax
sp
T
spratti
Cycloryx
Stomacosmethis
sp
T
stoliczkii
Chamalycaeus
Dicharax
sp
T
strangulatus
Dicharax
Dicharax
sp
NT
strigatus
Chamalycaeus
Dicharax
sp
DOD: detailed original description was sufficient for generic placement; NE
stuparum
Dicharax
Dicharax
sp
T
stylifer
Dicharax
Metalycaeus
sp
T
subculmen
Dicharax
Dicharax
sp
T
subdigitata
Dicharax
Dicharax
ssp
T
subfossilis
Chamalycaeus
Chamalycaeus
sp
T
subhumilis
Dicharax
Dicharax
sp
T
subinflatus
Chamalycaeus
Metalycaeus
sp
T
sublimus
Chamalycaeus
Chamalycaeus
ssp
T
succineus
Dicharax
Dicharax
sp
T
suhajdai nom. nov.
Dioryx
Metalycaeus
nom. nov. pro Alycaeusvarius Godwin-Austen, 1914
sp
T
sumatranus
Alycaeus
Chamalycaeus
sp
T
summus
Cycloryx
Pincerna
sp
T
swinhoei
Dioryx
Dioryx
sp
T
sylheticus
Alycaeus
Dicharax
sp
T
tadai
Chamalycaeus
Dicharax
sp
NT
takahashii
Chamalycaeus
Dicharax
sp
T
tanegashimae
Chamalycaeus
Dicharax
sp
T
tanghali
Alycaeus
Chamalycaeus
sp
T
tangmaiensis
Chamalycaeus
Dicharax
sp
T
tangmaiensis
Dioryx
Dioryx
sp
T
tenellus
Cycloryx
Pincerna
sp
T
teriaensis
Dicharax
Metalycaeus
sp
T
theobaldi
Dicharax
Dicharax
sp
T
thieroti
Alycaeus
Pincerna
sp
T
thompsoni
Cycloryx
Pincerna
sp
T
tokunoshimanus
Chamalycaeus
Dicharax
sp
T
tomotrema
Chamalycaeus
Metalycaeus
sp
T
toruputuensis
Dicharax
Metalycaeus
sp
T
trigonostoma
nomen nudum
troglodytes
Chamalycaeus
Chamalycaeus
sp
T
tsushimanus
Chamalycaeus
Dicharax
sp
T
umbonalis
Chamalycaeus
Dicharax
sp
T
urceolus
Dioryx
Dioryx
sp
T
urnula
Dioryx
Dioryx
sp
T
vallis
Pincerna
Pincerna
sp
DOD
vanbuensis
Dioryx
Pincerna
sp
T
variabilis
Cycloryx
Pincerna
ssp
T
varius Godwin-Austen, 1914
Dioryx
Metalycaeus
replaced by suhajdai nom. nov.
sp
T
varius Pilsbry & Y. Hirase 1905
Chamalycaeus
Metalycaeus
sp
T
vesica
Alycaeus
Metalycaeus
sp
T
vestitus
Alycaeus
Dicharax
sp
T
vinctus
Chamalycaeus
Metalycaeus
sp
T
virgogravida
Alycaeus
Stomacosmethis
ssp
DOD
vulcani
Alycaeus
Chamalycaeus
sp
T
wilhelminae
Alycaeus
Stomacosmethis
sp
DOD
woodthorpi
Dicharax
Dicharax
sp
T
yamneyensis
Alycaeus
Metalycaeus
sp
T
yanoshigehumii
Chamalycaeus
Dicharax
sp
T
yanoshokoae
Awalycaeus
Dicharax
sp
DOD
yanseni
Pincerna
Pincerna
sp
T
yetayensis
Alycaeus
Metalycaeus
ssp
T
zayuensis
Chamalycaeus
Metalycaeus
sp
see remarks
zhuangiyucuii
Alycaeus
Metalycaeus
junior synonym of heudei
syn
T
Of the 395 taxa, 32 are considered synonyms, although no recent revision has been undertaken for some more specific geographic areas, such as the Himalaya region. Consequently, 362 species-group taxa (320 species and 43 subspecies) of the Alycaeidae are currently accepted. Twenty-two were described by us in previous publications, and there are 18 species, that were formerly classified in Cycloryx that now belong to Pincerna due to its synonymy with Cycloryx. Of the 323 remaining species (excluding our taxa and Cycloryx), 209 (65%) are here classified in a new genus, whilst 114 (35%) remain in their previously classified genus. Most of these changes in generic placement resulted from two reasons. Firstly, morphological traits for generic definitions in the preceding studies were not able to classify the currently recognised taxa in morphologically distinct groups, and thus, probably the genera did not reflect evolutionary relationships. Secondly, the type species of each genus were not examined adequately before assigning a new species in these genera.
Alycaeids possess complex shell morphology compared to many other land snail groups, and exhibit a considerable magnitude of variation between populations, which could be interpreted as intraspecific or interspecific variation. Thus, a lumping approach would recognize much fewer taxa than a splitting approach. We employed the former approach, which would be most appropriate when examining widespread and variable taxa such as Dicharaxcristatus, D.fimbriatus or Metalycaeusmuciferus (see Páll-Gergely et al. 2017). This would be most practical for systematic handling of enormous morphological variability among the currently available specimens in the Alycaeidae. The ‘splitting’ approach to these groups would have resulted in recognition of twice as many or even more species. For example, Foon and Liew (2017) have described several species from Peninsular Malaysia based on small quantitative differences in shell sculpture and size, which would be appropriate for subspecific distinction. Although the present study did not include revision of northeastern Indian Alycaeidae, we found that some of Godwin-Austen’s species exhibit rather minor differences (M.brahma and its two new synonyms, and D.birugosus, and its new synonym, D.canaliculus are examples). Accordingly, geographic variation of the currently recognized alycaeid species diversity largely stem from difference between the splitting and lumping approaches employed by the authors.
Superfamily Cyclophoroidea Gray, 1847
Cyclophoridae Gray, 1847: 181.
AnimaliaGastropodaCyclophoridaeBFA3C5CC-80A1-546E-8834-C0CCAC32234BAlycaeidaeW. T. Blanford, 1864AlycaeinaeW.T. Blanford, 1864: 465.Alycaeinae– Godwin-Austen, 1886: 186. (subfamily of Cyclophoridae); Bouchet and Rocroi 2005: 23, 248; Bouchet et al. 2017: 28, 340. (subfamily of Cyclophoridae)Alycaeidae– Kobelt & Möllendorff, 1897: 146; Egorov 2013: 33.Diagnosis.
Shell with a complex gas exchange system consisting of a common external sutural tube and several extremely narrow, perpendicularly running microtunnels, formed by the outermost shell layer (Páll-Gergely et al. 2016). Bursa copulatrix connecting to lateral side of ovarium.
Remarks.
In the past this group has been treated as a subfamily of the Cyclophoridae, and as a family of its own right. The complex gas exchange system, combined with the unique position of the bursa copulatrix (both are important synapomorphic characters), seems to justify the distinction of this group as an independent family.
(Fig. 6A), SD Nevill (1878: 290). Cyclostomagibbum Eydoux, 1838, is a junior homonym of Cyclostomagibbum Draparnaud, 1805. Thus, Venmans (1856) proposed Alycaeuseydouxi Venmans, 1956 as a replacement name.
Gray (1850) originally included two species within Alycaeus (A.gibbus Eydoux, 1838 and A.strangulatus L. Pfeiffer, 1846) without selecting either of them to be the type species.
Type species of alycaeid genus-group taxa AAlycaeuseydouxi Venmans, 1956 (SMF 109290; type species of Alycaeus) BChamalycaeusfruhstorferi (Möllendorff, 1897), lectotype (SMF 109481; type species of Chamalycaeus). Photographs: Barna Páll-Gergely (A) and Frank Walther (B).
https://binary.pensoft.net/fig/468651Diagnosis.
Shell very large (D: 8–15 mm), triangular, with body whorl being dominant due to very long R2; protoconch smooth, obliquely striated, or transitional character state of the two; R1 usually finely reticulated, due to fine radial ribs and fine spiral striation; R2 long or very long (usually almost reaching 0.5 whorl), smooth or with lamella-like, straight, dense ribs; umbilicus narrow. Operculum thin or relatively thickened (can have both calcareous and proteinaceous layers, Foon and Liew 2017), without elevated outer structure (although scaffold-like calcareous structure and appressed radially spiral lamellae can be present, see Foon and Liew 2017). Central tooth with five cusps, broad, central cusp blunt.
Differential diagnosis.
The sculpture of Alycaeus and Chamalycaeus (smooth protoconch, spirally striated, weakly ribbed teleoconch) are identical, although Chamalycaeus tend to have stronger ribs. The distinction is based on the narrow (Alycaeus) and wide (Chamalycaeus) umbilicus. Furthermore, Alycaeus shells are larger, more colourful (reddish or yellowish) and the very long (i.e., ca. 0.5 whorl-long) R2 in Chamalycaeus is very rare.
Typical Pincerna has a relatively short tube and a strongly ribbed teleoconch, whereas typical Alycaeus possesses a long tube and its teleoconch is weakly ribbed. Some species (P.anceyi, P.mouhoti) form connections between the two genera. However, we prefer to maintain the distinction between Pincerna and Alycaeus due to the many species characteristic to both respective genera.
Alycaeus is easily distinguished differs from Stomacosmethis which has a yellowish-orange, triangular shell, and a very short tube.
Distribution.
This genus is known from northern Laos and northern Vietnam until the southern end of the Malay Peninsula (Fig. 7).
Remarks.
Regarding the authorsip of Alycaeus (i.e., Baird vs. Gray), we follow Petit (2012: 24–25).
Distribution of Alycaeus Gray, 1850 (dark shaded area) and Dioryx Benson, 1859 (light shaded area).
https://binary.pensoft.net/fig/468652AnimaliaGastropodaCyclophoridae837DB68C-0DD0-5883-9C14-30B400006183AlycaeusconformisFulton, 1902Fig. 2CAlycaeusconformisFulton, 1902: 68–69.Alycaeusconformis– Venmans 1956: 81–82, figs 1, 2 (radula, see Results on radula); Páll-Gergely et al. 2016: fig. 2.; Foon and Liew 2017: 30–33, figs 7E, 14, 31D.Type locality.
“Perak”.
Material examined.
Perak, NHMUK 1902.5.28.22-23 (2 syntypes); Thailand, Phuket Island, Khao Phra Thaeo Non-hunting Area, Bangpae waterfall, 8°2'6.09"N, 98°23'12.68"E, leg. B. Páll-Gergely & G. Majoros, July 2010, HNHM 99714 (6 shells examined by Páll-Gergely et al. 2016); NHMW 111541 (10 shells, ex NHMW 36649).
Remarks.
Protoconch with oblique ribs; R1 densely, finely, regularly ribbed with some very weak spiral striation; R2 very long, with dark and light stripes, the lighter being slightly narrower and more elevated from the surface.
Alycaeusconformis and A.gibbosulus have a characteristic, oblique striation on the protoconch (Fig. 2). However, the protoconch of A.rolfbrandti is also strongly sculptured (mamillated), and at the end of the protoconch oblique striae can be seen. Therefore, there is a continuous transition from the smooth Alycaeus-type protoconch sculpture to that of A.conformis and A.gibbosulus. Due to the similarity in protoconch sculpture and the geographic proximity (they are also found in mixed museum samples), A.conformis and A.gibbosulus are presumably closely related.
AnimaliaGastropodaCyclophoridae3C5BDE98-12C5-5B70-BE2E-A6701ED8F079AlycaeuseydouxiVenmans, 1956Figs 2A, 6A, 8CyclostomagibbumEydoux, 1838: 6, pl. 117, fig 1. (non Cyclostomagibbum Draparnaud, 1805)Alycaeusgibbus– Reeve 1878: pl. 1, species 3.Alycaeus (Alycaeus) gibbus– Kobelt 1902: 344–345.Alycaeus (Orthalycaeus) gibbus– Godwin-Austen 1914: 427, pl. 156, figs 5, 5a.AlycaeuseydouxiVenmans, 1956: 87, figs 6, 7 (radula). (nom. nov. pro Cyclostomagibbum Eydoux, 1838, non Cyclostomagibbum Draparnaud, 1805)Alycaeuseydouxi– Egorov 2013: fig. 58a; Páll-Gergely et al. 2017: 9–10, fig. 3A.Type locality.
“que dans les grottes formées dans l’intérieur des montagnes de marbre qui s’élèvent au milieu de la plaine oú est bâtie la ville de Turanne, en Cochinchine”.
Protoconch matte, without spiral lines; R1 fine, dense, rather regular ribs with weak spiral striation; R2 long, with dense, lamellae-like, elevated ribs, which are most elevated closer to the suture. Below the ribs, the microtunnels are visible as narrow light bands between the darker, thicker stripes (visible where there are weathered areas of the shell).
Habe (1965) reported “Dioryx gibbus (Reeve)” from “Kao Phlong, north or Sara Buri, Central Thailand”, without publishing a picture. This almost certainly refers to a different species.
Living specimens of Alycaeuseydouxi Venmans, 1856. Thủy Sơn (Water Mountain), Ngũ Hành Sơn (Five Elements or Marble Mountains), Da Nang. 16°0.254’N, 108°15.756’E. Photograph: Junn Kitt Foon.
Penang, coll. Dr. Stoliczka, NZSI M.24998 (1 syntype); Perak, Kwala Kangsar ex coll. Grübauer, NHMW 36649 (7 shells, other 10 shells are A.conformis: NHMW 111541).
Remarks.
Shell sculpture as in A.conformis. The types of A.chaperi were not examined by us. We follow Möllendorff (1886, 1891) and Foon and Liew (2017) in treating it as a synonym of A.gibbosulus.
Habe’s (1965) record of this species from “Khao Chong, Trang Province, peninsular Thailand” almost certainly refers to Alycaeusgibbosulus.
Spire low, but upper whorls (not only the protoconch) elevated; Protoconch glossy, R1 with very weak, irregular growth lines and even weaker, fine spiral striation; R2 very long, with wider darker stripes and lighter, narrower channels between, the channels are somewhat elevated from the surface.
AnimaliaGastropodaCyclophoridae652C7B44-50B6-553A-A4BE-0E24E621F6ECAlycaeuspyramidalisBenson, 1856AlycaeuspyramidalisBenson, 1856: 225.Alycaeuspyramidalis– Reeve 1878: pl. 1, species 6; Godwin-Austen 1914: 427, pl. 156, figs 6, 6a; Gude 1921: 216.Alycaeus (Alycaeus) pyramidalis– Kobelt 1902: 348–349.Type locality.
“ad collem Therabuin, vallis Tenasserim”.
Material examined.
Therabuin of Therapen (?) hill in Tenasserim, NHMUK 1888.12.4.937–938 (2 possible syntypes); No locality data, UMZC I.102830 (2 possible syntypes).
Remarks.
The two syntypes in the NHM and one of the shells from Cambridge were weathered. The third shell from Cambridge is in a good state, and its sculpture could be examined. Protoconch without particular sculpture, rather matte; R1 with low, irregular growth ridges; R2 relatively short, but much longer than typical in Stomacosmethis, the surface is irregularly wrinkled, and possibly ribbed near the suture.
Habe’s (1965) record of this species from “Khao Chong, Trang Province, peninsular Thailand” refers to Alycaeusgibbosulus.
Laos, Kalkberge ca. 20 km östl. Takek, leg. Brandt 08.09.1963, SMF 262541 (1 shell; labelled as the holotype of “Alycaeus carinatus Brandt”, but not mentioned by Maassen 2006); locality data as above, SMF 262541 (5 shells, labelled as paratypes of “Alycaeus carinatus Brandt”, but not mentioned in Maassen 2006); South-Central Laos, Khammouan Province, ca. 9 km NE of Thakhek (Muang Khammouan), NW exposition cliff, limestone, clay, black soil in limestone pockets, on and under rocks in dry secondary forest on and under, alt. 190 m, 17°26.757'N, 104°52.937'E, leg. Abdou, A. & Muratov, I.V., 27.11.2007., MNHN-IM-2012-27321 (19 complete shells + some shell fragments).
Remarks.
Protoconch irregularly ribbed, squamous, the last ca. 0.25 whorl with oblique ribs similar to those of A.conformis and A.gibbosulus; R1 with regular, fine, low ribs without spiral striae; R2 long with dense, lamella-like ribs (very similar to those of A.eydouxi).
The shells in the Senckenberg Museum are part of the original series of the species collected by Brandt, but since Maassen did not state that he examined them, they are not part of the type series.
“Thailand, Lub Lae Cave, an isolated limestone hill in Chonburi Province at 13°07'16"N, 101°36'05"E”.
Remarks.
We were unable to examine shells of Alycaeussomwangi, but the original description provides enough information to allow for generic placement. Protoconch without spiral striae, R2 very long, with regular, low ribs.
Alycaeus (Chamalycaeus) fruhstorferi (Fig. 6B) by monotypy, see also Remarks.
Diagnosis.
Shell very small to medium sized (D: 2–5 mm), usually flattened, discoid or low triangular, protoconch smooth (or very finely pitted), elevated even if the spire is low; R1 usually roughly reticulated due to spiral striation and radial ribs (sometimes prominent); R2 from short to very long, with widely spaced, sharp, elevated ribs; R3 normally developed. Operculum usually thin, without notable outer structures. Radula is known for a single species (central tooth with five cusps, broad, central cusp pointed).
Differential diagnosis.
See under Alycaeus and Table 3. Metalycaeus species are identical, with the exception of the spirally striated protoconch.
Distribution.
Chamalycaeus is distributed from the southeastern Himalaya Region, the Malay Peninsula, Sumatra, Java, Borneo, Sulawesi, and the Philippine Palawan Island (Fig. 9).
Distribution of Chamalycaeus Möllendorff, 1897 (dark shaded area) and Metalycaeus Pilsbry, 1900 (light shaded area).
https://binary.pensoft.net/fig/468654Remarks.
Kobelt and Möllendorff (1897) listed the species of “Pneumonopoma”, which included all members of the genus Alycaeus. They introduced “Subgenus Chamalycaeus n.” (Kobelt and Möllendorff 1897: 148), indicating it as a new subgenus, and included 44 species within their new group. The subgenus Chamalycaeus, however, was previously mentioned in another paper in the same volume of the Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, in which Möllendorff (1897b: 93) described Alycaeus (Chamalycaeus) fruhstorferi. Möllendorff’s (1897b) paper was published in July-August, whereas that of Kobelt and Möllendorff (1897) was published in September-October. Accordingly, the genus Chamalycaeus was described by Möllendorff (1897) and its type species is Alycaeus (Chamalycaeus) fruhstorferi by monotypy. On the other hand, Alycaeus (Chamalycaeus) fruhstorferi was not listed in Kobelt’s and Möllendorff’s (1897) paper, which indicates that they aimed to describe A.fruhstorferi after their revision of Alycaeus. Moreover, Kobelt’s (1902) monograph referred to Chamalycaeus as it was introduced by Kobelt and Möllendorff (1897). Almost all subsequent treatments erroneously attributed the name Chamalycaeus to Kobelt & Möllendorff, 1897 (Kobelt 1902; Gude 1921; Yen 1939; Zilch 1957; Azuma 1980; Minato 1982b, 1987a, 2005; Minato and Yano 1988; Egorov 2005; Zhang et al. 2008) and referred to Alycaeusandamaniae Benson, 1861 as the type species as subsequent designated by Gude (1921). The ICZN Code 70.2 states the following: “If it is found that an earlier type species fixation has been overlooked, the overlooked fixation is to be accepted and any later fixations are invalid. If this is considered to cause instability or confusion the case is to be referred to the Commission for a ruling”. Therefore, we must examine whether the correction of the type fixation would cause instability. In our view, confusion or instability would be caused only if the majority of authors who have described species within Chamalycaeus were unaware of the shell morphology of Alycaeusandamaniae (incorrectly selected as the type species for Chamalycaeus). No detailed description of Alycaeusandamaniae has ever been published, and our revision suggests that most authors who described Chamalycaeus species did not examine samples of Alycaeusandamaniae. Thus, we find no reason to present this issue to the Commission. Instead, we follow Egorov (2013) in accepting Möllendorff (1897b) as the author of Chamalycaeus. Thus, in accordance with Art. 70.2 of the Code we clarify that Alycaeus (Chamalycaeus) fruhstorferi is the type species of Chamalycaeus Möllendorff, 1897 by monotypy.
AnimaliaGastropodaCyclophoridaeFEFD001D-63D2-5CCA-A735-31508704D77AChamalycaeusandamaniae(Benson, 1861)AlycaeusandamaniaeBenson, 1861: 28–29.Alycaeusandamaniae– Reeve 1878: pl. 2, species 10; Godwin-Austen, 1914: 430–431.Alycaeus (Chamalycaeus) andamaniae– Kobelt 1902: 352–353; Gude 1921: 223–224.Chamalycaeus (Chamalycaeus) andamaniae– Ramakrishna et al. 2010: 52.Type locality.
“ad portum Blair Insulæ Andamanicæ”.
Material examined.
Andaman Islands, UMZC I.103175 (holotype [single specimen mentioned in the original description], photographs examined); Camorta, leg. De Roepstorff, NHMUK 1903.7.1.2708 (1 specimen).
Remarks.
Protoconch elevated, finely granulated, no signs of spiral lines; R1 with equally strong spiral lines and irregular ribs; R2 short, with sharp, widely spaced, lamella-like ribs.
AnimaliaGastropodaCyclophoridae8F791FD6-A80D-53C1-8E27-3EDF3A867692Chamalycaeus(?)armillatus(Benson, 1856)AlycaeusarmillatusBenson, 1856: 227.Alycaeusarmillatus– Reeve 1878: pl. 5, species 38; Godwin-Austen 1914: 406, pl. 151, figs 3, 3a.Alycaeus (Dicharax) armillatus– Kobelt 1902: 365; Gude 1921: 236–237.Type locality.
“ad Thyet-Mio cum præcedente (= A. sculptilis)”.
Material examined.
UMZC 102995 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, no spiral striation visible.
The specimen largely matches Benson’s original description, and therefore we consider it to be the holotype. The photographs of that specimen show some signs of spiral striation. However, those striae may be part of the lower shell layers, and not raised threads as in other Chamalycaeus species. Consequently, the spiral striae on the holotype of A.armillatus may not be homologous with the ones in Chamalycaeus species; we would need fresh shells to confirm this. For the time being, we refer to this species as Chamalycaeus(?)armillatus.
The shells labelled as A.armillatus in the NHM (Thayet-myo, Pegu, coll. Blanford, NHMUK 1906.4.4.71, 6 shells) belong to another (probably undescribed) Chamalycaeus species based on the shorter R3, the shallower constriction between R2 and R3, and the smaller distance between the inner and outer peristomes.
Protoconch elevated, with very finely pitted surface, no signs of spiral striae; R1 with irregular ribs and spiral striae of the same strength; R2 short, with regular, straight, sharp ribs.
Golf von Siam: Koh-Samui, coll. Möllendorff, SMF 109468 (lectotype, designated by Zilch 1957); Same data, SMF 109469 (4 paralectotypes).
Remarks.
Protoconch low, rather matte, very finely granulated, without spiral lines; R1 densely, rather regularly ribbed, the ribs are quite sharp, there is a hardly visible spiral striation between each of the ribs; R2 short, with ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridaeDCBDDFC2-A13C-5F3A-BFA4-D8D8C2DDCBA5Chamalycaeuscelebensis(E. von Martens, 1891)Fig. 10AAlycaeuscelebensisE. von Martens, In: Weber, 1891: 217–218.Alycaeus (Chamalycaeus) celebensis– Kobelt 1902: 354.Type locality.
“Celebes: Luwu”.
Material examined.
Luwu, Celebes, M. Weber, ZMB/MOLL 44738 (photographs of a shell [possible syntype] were examined).
Remarks.
Protoconch elevated, no spiral lines visible, although the suture is filled with dirt and the photographs are not of high quality. R1 with strong widely spaced, ribs with a fine spiral striation; R2 with denser, straighter ribs, although this part of the shell was somewhat corroded. This species is placed in Chamalycaeus due to the colourless shell and biogeographic location. A closer examination of the protoconch would be important to rule out its affinity with Metalycaeus, although the occurrence of that genus in Celebes would be surprising.
Shells of Chamalycaeus Möllendorff, 1897 species AChamalycaeuscelebensis (E. von Martens, 1891), possible syntype (ZMB/MOLL 44738) BChamalycaeuskessneri Vermeulen, 1996, paratype (SMF 311351). Photographs: Barna Páll-Gergely (B) and Christine Zorn (A).
https://binary.pensoft.net/fig/468655AnimaliaGastropodaCyclophoridae5908F974-537A-55AF-AF81-A4B9820C501DChamalycaeuseveretti(Godwin-Austen, 1889)AlycaeuseverettiGodwin-Austen, 1889: 347, pl. 37, figs 5, 5a.Alycaeus n. sp. – Aldrich 1889: 25, pl. 3, figs 2, 2a, 2b (later mentioned A.broti, but this name was not made available).Alycaeuseveretti– E. A. Smith 1895: 116.Alycaeus (Dicharax) everetti– Kobelt 1902: 369.Chamalycaeus (Dicharax) everetti– Zilch 1957: 145.Type locality.
“Niah Hills”
Material examined.
Niah Hills, Borneo, NHMUK 1889.12.7.33 (holotype [single specimen mentioned in the original description]).
Remarks.
Aldrich (1889) did not give a name for the “Alycaeus sp.”, species he figured and described, because he thought it might be A.spiracellum, which he was unable examine for comparison. He mentioned that “if new, I propose the name Alycaeus broti for it”. This action does not make the name available, because under ICZN Art. 11.5 “To be available, a name must be used as valid for a taxon when proposed”, which was not the case for A.broti; therefore, the name Alycaeusbroti is not available. Smith (1895) mentioned that he compared A.everetti specimens with his “A. broti”, and they were identical.
Protoconch elevated, no spiral lines visible; R1 with very fine, irregular ribs and spiral lines; R2 short, with sharp, straight, widely spaced ribs.
“Insel Bongao zwischen Sulu und Borneo” (from the title).
Material examined.
Sulu-Inseln, Insel Bongao (Tawi-Tawi-Gr.), leg. Möllendorff 1890, coll. O. Boettger, SMF 109479 (holotype [single adult specimen mentioned in the original description]); Same data, SMF 109480 (4 paratypes).
Remarks.
Protoconch elevated without spiral lines; R1 with weak, widely spaced, irregular ribs and somewhat stronger spiral striation; R2 relatively short, with widely spaced, elevated, sharp ribs.
Java, leg. Fruhstorfer, coll. Möllendorff, SMF 109481 (lectotype, designated by Zilch 1957); Same data, SMF 109482 (5 paralectotypes); Mons Gede, 4000’, W. Java, leg. Fruhstorfer, Aug. 1892, E.R. Sykes colln. 1954, NHMUK 20150361 (4 specimens).
Remarks.
Protoconch elevated, no spiral lines visible; R1 rather regularly ribbed with sharp ribs, and with somewhat weaker spiral striation; R2 relatively long, with widely spaced, sharp ribs.
Tengasa Monkey Temple, Nusa Penida, Indonesia, 8°45'S, 115°31'E, leg. A.J. Witten, 1993, NHMUK 20000248 (paratype); Indonesia, Nusa Penida, Tengasa Monkey Temple, 8°45'S, 115°31'E, Secondary forest, leg. A.J. Whitten, 1993, ex coll. J.J. Vermeulen 4080, SMF 311351 (1 paratype). Indonesia, South Kalimantan, Nateh, leg. Yansen Chen, April 2012 (6 shells).
Remarks.
The examined paratype was badly weathered, only the elevated protoconch with some spiral lines on R1 and the short tube were visible. Based on these, C.kessneri remains classified in the genus Chamalycaeus.
The shells from Nateh were considerably smaller than typical C.kessneri, but agreed with that species in terms of shell shape, the short tube, and the spiral striation. Although these six shells were also weathered, one of them was in a relatively good condition. None of the shells showed signs of spiral striation on the protoconch, therefore the placement of this species in Chamalycaeus seems to be justified.
Malakka: Bukit Pondong (Perak), SMF 109493 (lectotype, designated by Zilch 1957); Same data, SMF 109494 (2 paralectotypes).
Diagnosis.
Protoconch rather low, without obvious spiral lines, the granules following a near spiralling arrangement, but not at all similar to the multiple, narrow spiral striae typical to most Metalycaeus species; R1 with rather regular ribs and strong spiral lines; R2 extremely short, with only ca. five ribs which are blunt (probably bent?). Operculum unknown.
Shells of Chamalycaeus Möllendorff, 1897 species AChamalycaeusmicroconus (Möllendorff, 1887), lectotype (SMF 109493) BC.mixtus Zilch, 1957, holotype (SMF 109510). All images: Barna Páll-Gergely, courtesy Ronald Janssen.
Malakka: Bukit Pondong (Perak), SMF 109510 (holotype); Same data, SMF 109511 (4 paratypes).
Remarks.
Protoconch as in C.microconus; R1 with rather irregular, widely spaced, low ribs, with somewhat stronger spiral striae; R2 extremely short, consists of ca. eight ribs which are bent in the direction of their anterior neighbours.
AnimaliaGastropodaCyclophoridaeF37C80C4-C0C1-5004-B37F-4C24AE888E1EChamalycaeusoglei(Godwin-Austen, 1914)AlycaeusogleiGodwin-Austen, 1914: 362, pl. 148, fig. 2.Alycaeusoglei– Gude 1921: 213.Alycaeus (Alycaeus) oglei– Ramakrishna et al. 2010: 49.Type locality.
“Sadia”; “Dihing, 500 ft”.
Material examined.
Noa Dihing, 500 f. (2 shells in the vial) & Sadia, 350 f. (1 shell in the vial), leg. M. Ogle, NHMUK 1903.07.01.2491 (syntypes). The box labelled A.oglei contained two glass vials.
Remarks.
Protoconch elevated, no spiral lines visible; R1 with rather irregular, low ribs, and weaker spiral striation; R2 very long with widely spaced, sharp ribs.
AnimaliaGastropodaCyclophoridae15F2F26B-5C08-5934-88BE-596C8A22DCD8Chamalycaeusperplexus(Godwin-Austen, 1914)AlycaeusperplexusGodwin-Austen, 1914: 380, pl. 155, fig. 11.Alycaeusperplexus– Gude 1921: 214.Alycaeus (Alycaeus) perplexus– Ramakrishna et al. 2010: 50.Type locality.
“Khasi Hills”.
Material examined.
Khasi Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2756. (3 syntypes).
Remarks.
Protoconch elevated, no spiral lines visible; R1 irregularly, weakly wrinkled and as strongly spirally striated; R2 moderately long, with wider darker and narrower lighter stripes alternating, the overall surface is nearly smooth, rather irregularly wavy.
“Philippine Islands, Palawan Prov., 50 km SW of Quezon, along trail from Ransang to Tau’t Batu Caves, 90–390 m a.s.l., 8°53'N, 117°35'E”.
Material examined.
Only the holotype (UF 525657) is known.
Remarks.
R1 rather strongly and irregularly ribbed with weaker spiral striation; R2 + R3 short, less than 90° combined; R2 shorter than R3; ribs on R2 lamella-like; spiral striation also visible on R2; R3 with spiral striation and weaker ribs than those on R1.
The placement of this species into the genus Chamalycaeus is based on biogeographic information alone, since the protoconch, which is necessary for generic allocation, is absent in the only available shell (Páll-Gergely and Auffenberg 2019).
AnimaliaGastropodaCyclophoridae4DD86E08-CAF5-50E8-ADCD-4932F38E7564Chamalycaeusreinhardti(Mörch, 1872)Alycaeus (Charax) reinhardi(sic) Mörch, 1872a: 22.Alycaeus (Charax) reinhardi(sic) – Mörch 1872b: 315.AlycaeusnicobaricusReeve, 1878: pl. 4, species 29.Alycaeusreinhardti– Godwin-Austen 1895: 455; Godwin-Austen 1914: 431; Gude 1921: 216–217.Alycaeus (Alycaeus) reinhardti– Kobelt 1902: 349; Subba Rao and Mitra 1991: 26, pl. 3, fig. 4.; Ramakrishna et al. 2010: 50.Type locality.
“Bords de la rivière Galathea, sur la terre, sous les feuilles mortes” and “Kar Nicobar”.
Material examined.
Great Nicobar, coll. Godwin-Austen, NHMUK 1903.7.1.2711.
Remarks.
Spire elevated, shell slightly wider than high; protoconch elevated, no spiral lines visible; R1 with irregular ribs, and somewhat weaker spiral lines; R2 relatively long, with widely spaced, regular, sharp ribs.
The original spelling of the species was reinhardi, which was corrected to reinhardti by Kobelt (1902). This was a justified emendation under the Article 32.5. of the ICZN Code, because it was obvious that Mörch (1872a) named the new species after the collector Reinhardt.
Godwin-Austen (1895) mentioned that the type of this species is from Great Nicobar Island, and the form from Camorta is named f. minor by Mörch. However, we have not found the publication in which Mörch introduced that name.
Reeve’s A.nicobaricus was not examined by us, but it was considered to be a junior synonym of A.reinhardti by Gude (1921).
W-Java, Djampang, 2000’, leg. H. Fruhstorfer, 1895, coll. O. Boettger, SMF 57196 (syntype, labelled as holotype, photographs examined).
Remarks.
The original description does not mention the number of examined specimens. Thus, we consider the specimen labelled holotype (SMF 57196) as a syntype.
Spire quite elevated, shell approximately as high as it is wide; protoconch very finely granulated, no spiral lines visible; R1 with strong, rather irregular ribs and somewhat weaker spiral striation; R2 short, with widely spaced, sharp ribs.
AnimaliaGastropodaCyclophoridae0A37AD22-1337-51B9-A4AB-96DD473FB75CChamalycaeusrichthofeni(W. T. Blanford, 1863)AlycaeusrichthofeniW.T. Blanford, 1863: 324.Alycaeusrichthofeni– Reeve 1878: pl. 3, species 23; Godwin-Austen 1914: 428, pl. 151, fig. 9.Alycaeus (Dicharax) richthofeni– Kobelt 1902: 376; Gude 1921: 268.Type locality.
“Molmain”.
Material examined.
Tenasserim, Moulmein, NHMUK 1906.5.5.24 (holotype [single specimen mentioned in the original description]).
Remarks.
The holotype is strongly weathered. Protoconch strongly elevated, no spiral lines visible; R1 rather regularly ribbed with some weak signs of spiral striation; R2 long, with widely spaced ribs, which were probably sharp in the fresh shell.
AnimaliaGastropodaCyclophoridae45FAE379-94D6-5EEA-84EF-2AB1E9150911Chamalycaeussculptilis(Benson, 1856)AlycaeussculptilisBenson, 1856: 226–227.Alycaeussculptilis– Reeve 1878: pl. 4, species 32, figs a, b; Godwin-Austen 1914: 398, 412, pl. 139, figs 7, 7a; pl. 155, fig. 8.AlycaeusmargaritaTheobald in Hanley & Theobald, 1874: pl. 97, fig. 7 (renamed A.microstoma by Reeve 1878)AlycaeusmicrostomaReeve, 1878: pl. 4, species 28.Alycaeus (Chamalycaeus) sculptilis– Kobelt 1902: 362; Gude 1921: 233.Chamalycaeus (Chamalycaeus) sculptilis– Ramakrishna et al. 2010: 55.Type locality.
“ad Thyet-Mio prope fluvium Irawadi, non procul a finibus provinciæ Burmanicæ Britannicæ”.
Material examined.
Bens. col., Thyet Myo”, UMZC I.102845 (3 shells, type status uncertain); Pegu, Thayet-myo, “typical”, “aperture figured”, NHMUK 1906.4.4.70.
Remarks.
Protoconch elevated, no spiral lines visible; R1 with rather regular, widely spaced ribs, and somewhat weaker spiral lines; R2 long, with widely spaced ribs; there is a lamella on each rib which is slightly bent in the direction of the anterior neighbour.
Reeve (1878) named the shell figured by Hanley and Theobald (1874) on pl. 97, fig. 7, as Alycaeusmicrostoma, and published drawings (pl. 4, species 28). The type specimens were not examined by us, but that species was considered to be a synonym of Alycaeussculptilis Benson, 1856 by Godwin-Austen (1914).
The syntype is weathered and was glued to a piece of black paper by its R2 area, therefore limited information could be gained during its examination. The shell is depressed and conical; protoconch strongly weathered, but there were no signs of spiral striation near the suture; R1 regularly and strongly ribbed with very weak spiral striation; length of R2 could not be fully seen, but has low, dense riblets and fine spiral lines. We received photographs and good quality drawings of newly collected shells from Thor-Seng Liew and Jaap Vermeulen (pers. comm. August 2019), and those confirmed that this species is a Chamalycaeus due to the colourless shell, long R2, and relatively strong ribs.
AnimaliaGastropodaCyclophoridae134C8378-B11B-5AA8-A89C-764CD42CEE3EChamalycaeussubfossilis(P. Sarasin & F. Sarasin, 1899)AlycaeussubfossilisP. Sarasin & F. Sarasin, 1899: 63–64, pl. 4, figs 46, 46a, pl. 5, fig. 66, pl. 8, fig. 91.Alycaeus (Chamalycaeus) subfossilis– Kobelt 1902: 363–364.Chamalycaeussubfossilis– Páll-Gergely and Auffenberg 2019: 378, fig. 2A.Type locality.
Protoconch elevated, finely pitted, no spiral lines visible; R1 with dense riblets and some weak spiral striation; R2 short, with somewhat elevated ribs that are similar to those on R1.
Lothar Forcart selected a specimen (NHMB 2265a) and labelled it as the lectotype, but never published this action (Ambros Hänggi, pers. comm. 2020 June). We previously referred to that specimen as a lectotype (Páll-Gergely and Auffenberg 2019); however, that was not a valid lectotype selection. Thus, here we designate the same specimen selected by Locard (Páll-Gergely and Auffenberg 2019: fig. 2a) as the lectotype.
AnimaliaGastropodaCyclophoridae681691B1-A8D6-5F2B-83DF-FEB6B0878447Chamalycaeussumatranus(E. von Martens, 1900)Fig. 12AAlycaeus (Orthalycaeus) sumatranusE. von Martens, 1900: 6–7.Alycaeus (Alycaeus) sumatranus– Kobelt 1902: 351.Alycaeussumatranus– van Benthem Jutting 1959: 77–78, fig. 5.Type locality.
The original description does not mention the number of examined specimens. Thus, we consider the specimen labelled holotype (ZMB/MOLL, 51748) as a syntype.
Protoconch elevated, no spiral striation visible; R1 with rather irregular, low ribs and spiral striation roughly of the same strength; R2 long, with widely spaced, sharp ribs.
“Munipur. Exact locality not recorded; somewhere on the northern side of the valley”.
Material examined.
Munipur, figured by Godwin-Austen, NHMUK 1903.7.1.2671 (6 syntypes).
Remarks.
Protoconch elevated, but no spiral lines are visible; R1 with widely spaced, regular ribs and fine spiral striation; R2 long, with widely spaced, regular, sharp ribs.
“Mittel-Sumatra: Höhle von Pauh bei Fort de Kock”.
Material examined.
Mittel-Sumatra: Höhle von Pauh bei Fort de Kock, leg. Thienemann, 13.03.29., ZMB/MOLL 76101 (1 syntype; photographs examined).
Remarks.
Protoconch low, no spiral lines visible; R1 with widely spaced, sharp regular ribs and very weak spiral striation; R2 very short, with ribs which are similar to those on R1.
AnimaliaGastropodaCyclophoridaeFA3E1E03-B3E6-5F0A-A6CE-EE805F22C1ACChamalycaeusvulcani(W. T. Blanford, 1863)AlycaeusvulcaniW.T. Blanford, 1863: 323.Alycaeusvulcani– Reeve 1878: pl. 2, species 17; Godwin-Austen 1914: 413–414, pl. 151, figs 5, 5a; Gude 1921: 221–222.Alycaeus (Alycaeus) vulcani– Kobelt 1902: 352.Type locality.
“on the upper portion of the isolated peak of Puppa, an extinct volcano lying ca. 40 miles E. S. E. of the town of Pu-gán in the territories of the king of Ava”.
Material examined.
Ava, Burma, MCZ 135705 (1 shell, labelled as syntype); Puppadoung, ex coll. Theobald, NHMUK 1888.12.4.939–942 (4 shells, possible syntypes); Puppa, Ava, Burma, coll. H.F. Blanford, ex coll. auctoris, NHMUK (8 shells, possible syntypes); Puppa Hill, Ava, leg. Blanford, Crosse coll. 1899, Sykes coll. 1954, NHMUK (2 shells, possible syntypes).
Remarks.
Protoconch rather elevated, without spiral striation; R1 with elevated, regular, sharp ribs and without spiral striation; R2 long with widely spaced, sharp ribs.
The absence of spiral striation on the entire shell is unusual for Chamalycaeus, and characteristic for Dicharax. However, the general shell shape, the strong, equidistant ribs, and the elevated protoconch suggests that this species belongs to Chamalycaeus.
AnimaliaGastropodaCyclophoridae92ADF073-0314-52A2-B1D9-3D1072C3B8D2DicharaxKobelt & Möllendorff, 1900CharaxBenson, 1859: 177.DicharaxKobelt & Möllendorff, 1900: 186 (new replacement name for Charax Benson, 1859, non Charax Scopoli, 1777 [Pisces]).Chamalycaeus (Dicharax)– Thiele 1929: 108; Wenz, 1938: 478; Egorov 2013: 37.Chamalycaeus (Sigmacharax)Kuroda, 1943: 8.Chamalycaeus (Cipangocharax)Kuroda, 1943: 11.Chamalycaeus (Awalycaeus)Kuroda, 1951: 73–74.Chamalycaeus (Awalycaeus)– Egorov 2013: 35–36.Chamalycaeus (Cipangocharax)– Egorov 2013: 36.Chamalycaeus (Sigmacharax)– Egorov 2013: 37–38.Dicharax– Páll-Gergely et al. 2017: 10; Páll-Gergely and Asami 2017: 14 (Awalycaeus, Cipangocharax and Sigmacharax are synonyms).Type species.
Type species of alycaeid genus-group taxa ADicharaxhebes (Benson, 1857) (SMF 109244; type species of Dicharax) BD.(?)abei (Kuroda, 1951) (NSMT 50125; type species of Awalycaeus) CD.(?)biexcisus (Pilsbry, 1902) (NSMT 263; type species of Cipangocharax) DD.(?)itonis (Kuroda, 1943) (NSMT 78866; type species of Sigmacharax). Close-up images of the aperture are not to scale. All photographs: Barna Páll-Gergely.
https://binary.pensoft.net/fig/468658Diagnosis.
Shell very small to very large (D: 1–11 mm), in most cases the spire low (dorsal side flattened), spire rarely elevated (shell globular); protoconch low in nearly all species, smooth or finely pitted, not spirally striated; R1 usually glossy, sometimes ribbed (ribs can vary from weak to strong), but spiral lines almost always absent; R2 of variable length, typically with prominent ribs which are bent in an anterior direction, but many species have smooth R2 or straight ribs; R3 well developed, often with blunt or sharp swelling, in some taxa reduced (mostly ‘Awalycaeus’). Operculum thin or with various outer funnel-like structure resulting from modifications of the multispiral lamina. Central tooth typical for the family: 5–7 cusps, broad, central cusp pointed.
Differential diagnosis.
This genus can be recognised by the absence of spiral striation on the entire shell (protoconch and teleoconch). Very few species with spiral striation are classified in this genus.
Distribution.
Dicharax inhabits a large geographic area from the southeastern Himalayan region to Japan, and through the Malay Peninsula to the southern arc of the Malay Archipelago up to Sumatra and Java. There are also isolated occurrences in the Western Ghats of India and in the southwestern Himalaya (see Fig. 14).
Distribution of Dicharax Kobelt & Möllendorff, 1900.
https://binary.pensoft.net/fig/468659Remarks.
Cipangocharax, introduced as the subgenus of Chamalycaeus, was described for a single species, Alycaeusbiexcisus. The diagnosis of Cipangocharax was in fact the abbreviated description of Alycaeusbiexcisus. Kuroda (1943) indicated some features in italics, emphasising the importance of these characters to distinguish Cipangocharax from other members of Chamalycaeus. These characters were the extraordinary thickness of the operculum, and the closely coiled outer belt on the outer surface of the operculum. The Japanese Chamalycaeus species described since Kuroda’s (1943) paper showed that there are transitional character states between the thick and belted operculum of A.biexcisus and the thin and unbelted opercula of most Japanese Chamalycaeus species (e.g., Minato 1993). For example, the operculum of Cipangocharaxkiuchii is relatively slim, whereas that of “Chamalycaeus” miyazakii is exceptionally thickened. Consequently, the thickness of the operculum is not a distinguishing feature between Cipangocharax and other Japanese species assigned to Chamalycaeus. The outer opercular belt is missing in C.placenovitas (a species being otherwise very similar to A.biexcisus), therefore this character is also not stable within the genus. Moreover, the outer belt is known to be present and absent within the same species, or even population (see under Chamalycaeusnipponensis and Dicharaxsimplicilabris, see Páll-Gergely et al. 2017). The other distinctive character mentioned by Kuroda (1943) is the sinuated columellar margin. This region is not sinuated either in C.placenovitas, or in C.okamurai. Therefore, this character is also not stable within the genus. Moreover, Japanese Chamalycaeus species with unstriated protoconchs show an extraordinary diversity in terms of the formation of the aperture (C.expanstoma, C.okamurai, C.yanoshigehumii), indicating that the morphological variation is very high between species. Consequently, among the Japanese species with unstriated protoconch, it would not be legitimate to classify certain species into separate (sub)genera from the others. Furthermore, the species classified into the genus Sigmacharax also do not differ considerably from the rest of Japanese species with a smooth protoconch. Therefore, based on the absence of the spiral striation on the entire shell, these species are classified in the genus Dicharax. The overlapping ribs near the tube (Fig. 15) may a synapomorphic character of Japanese and Korean Dicharax, but this character was also found in the Chinese species D.alticola (Páll-Gergely et al. 2017), which is, due to the geographic distance, probably only distantly related. The morphological variation within the genus Dicharax (especially in northeastern India and in the Malay Archipelago) is so large, that at the current time we do not find it meaningful to separate the Japanese and Korean species into a separate subgenus within Dicharax.
R2 ribs of Japanese Dicharax Kobelt & Möllendorff, 1900 species A, BDicharax(?)abei (Kuroda, 1951), NSMT 50125 C, DDicharax (?) biexcisus (Pilsbry, 1902), NSMT 263 E, FDicharax(?)itonis (Kuroda, 1943), NSMT 78866. All images: Barna Páll-Gergely.
https://binary.pensoft.net/fig/468660
Awalycaeus is a peculiar group of alycaeids due to the reduced (short, un-swollen) R3. However, in Awalycaeusyanoshokoae there is a moderately developed R3, which can be interpreted as an intermediate form between Awalycaeus and the rest of Japanese alycaeids which have a smooth protoconch. Given that the other shell characters (absence of spiral striation, merged R2 ribs) are similar to the other Japanese species, we also treat Awalycaeus as a synonym of Dicharax.
Such ‘over spitting’ of generic taxa inhabiting Japan has also been documented in the pulmonate family Clausiliidae, which is a character-rich family such as the Alycaeidae (Páll-Gergely et al. 2019). Nordsieck (1998) stated that the Japanese clausiliid genera and subgenera correspond only to subgenera and species groups of Western Palaearctic clausiliids. This claim was confirmed by recent molecular phylogeny (Motochin et al. 2017).
For the sake of simplicity, this genus is divided into three sections: typical (with curved R2 ribs), atypical (without the typical R2 sculpture), and those species from Japanese and Korean localities (including species formerly classified into Awalycaeus, Cipangocharax, and Sigmacharax).
Protoconch slightly elevated, no spiral lines visible; R1 with rather regular, moderately elevated ribs without spiral lines inbetween; R2 with ribs being lamella-like.
Patung, Hubei: China, coll. Boettger ex coll. Möllendorff, SMF 39225 (lectotype of A.anthostoma, designated by Yen 1939); same data, SMF 39226 (12 paralectotypes of anthostoma); China, Hupé, Coll. Möllendorff, ex Oberwimmer, ex David D. Thaanum Jan. 1947, MCZ 180902 (3 paralectotypes of anthostoma).
Remarks.
In our earlier paper (Páll-Gergely et al. 2017) we overlooked that Gredler (1891) already synonymised A.pentagonus with A.anthostoma.
Protoconch low, glossy, without spiral lines; R1 rather regularly ribbed, ribs low, no spiral striation visible; R2 relatively long, with ribs curved towards the aperture, forming a smooth surface.
“Neuglo,” “Phulong” and “Dihung River, N. Cachar, north of Asalu”.
Material examined.
Dihung, N. Cachar, coll. Godwin-Austen, NHMUK 1903.7.1.2636. (2 syntypes); Asalu, North Cachar, NHMUK 1903.7.1.2761 (probably figured sample, with images on the sides of the box). See under D.crispatus.
Remarks.
Syntypes: protoconch low, spiral lines not visible; R1 irregularly ribbed, without spiral lines; R2 moderately long, with lamella-like, straight ribs.
Other sample.
The specimens are conspicuously variable in term of shell size (smallest: D = 2.7 mm, H = 2.1 mm; largest shell: D = 4.0 mm, H = 2.9 mm), the sculpture of R1 (nearly smooth to strongly, regularly ribbed) and the sculpture of R2. Despite the large variability, we consider all shells to belong to the same species since the variation is continuous between the extreme morphological forms. Protoconch low without spiral striation. R2 of some specimens typical Dicharax-like (ribs are curved towards the aperture), whereas those of other specimens are more lamella-like and less curved. Note that the shells with straight, lamella-like ribs on R2 are not weathered, which demonstrates that the two types of ribbing are a part of the intraspecific variation.
AnimaliaGastropodaCyclophoridaeE849669B-A1B7-57A5-A2D0-5BE06CA92FE8Dicharaxavae(W. T. Blanford, 1863)AlycaeusavaeW. T. Blanford, 1863: 323–324.Alycaeusavae– Reeve 1878: pl. 3, species 20; Godwin-Austen 1914: 406–407, pl. 151, fig. 6.Alycaeus (Dicharax) avae– Kobelt 1902: 365; Gude 1921: 238.Type locality.
“The hills east of Mandalay and Ava”.
Material examined.
Shan Hills, E of Ava, Burma, coll. Blanford, NHMUK 1906.4.4.61 (6 syntypes).
Remarks.
Protoconch low without spiral lines; R1 irregularly, densely ribbed, no spiral lines visible; R2 relatively short; ribs lamella-like, curved towards the aperture, but they are not in contact.
AnimaliaGastropodaCyclophoridaeFACAD3D0-4574-5406-920B-F715297DAE01DicharaxbisonPáll-Gergely & Hunyadi, 2017DicharaxbisonPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 32–34, figs 21A, 22, 23.Type locality.
Holotype (HNHM 99703) and several paratypes, see the original description for further details.
Remarks.
Protoconch low, rather matte; R1 regularly, densely ribbed, ribs low, without spiral lines; ribs becoming slightly more widely spaced towards end of R1; R2 very densely ribbed, ribs curved towards the aperture; for more details see the original description.
“Thailand, Chiang Rai Province, approx. 9 km south-southwest from Mae Sai, Wat Tham Pla, 400 m a.s.l., 20°19.723'N, 99°51.817'E”.
Material examined.
HNHM 100177 (holotype).
Remarks.
Protoconch glossy; R1 also glossy, with very fine, irregular growth lines; R2 with dense, curved ribs (ca. 46 altogether), for more details see the original description.
AnimaliaGastropodaCyclophoridae32CA960F-0FFB-556E-861E-BA5AA58CF0C7Dicharaxchennelli(Godwin-Austen, 1886)AlycaeuschennelliGodwin-Austen, 1886: 192–193, pl. 48, fig. 2.Alycaeuschennelliand chennelli var. – Godwin-Austen 1914: 387.Alycaeus (Dicharax) chennelli– Kobelt 1902: 366; Gude 1921: 240–241.Chamalycaeus (Dicharax) chennelli– Ramakrishna et al. 2010: 57.Type locality.
Piknui, Naga Hills, leg. A. Chennell, NHMUK 1903.7.1.2612 (8 syntypes); Lhota Naga Hills, leg. Chennell, NHMUK 1903.7.1.2613 (10 shells = “chennelli var.”).
Remarks.
Protoconch rather glossy, low, no spiral lines visible; R1 glossy, with widely spaced sharp ribs which are present only near the suture, and without spiral lines; R2 with curved ribs, typical to Dicharax.
The only stable character which distinguishes D.chennelli from D.diagonius is the presence of a lower apertural bay in the former, whilst it is absent in the latter.
AnimaliaGastropodaCyclophoridaeB471B14C-295D-590A-9905-BD8287837E42Dicharaxconicus(Godwin-Austen, 1871)AlycaeusconicusGodwin-Austen, 1871: 87–88, pl. 3, fig. 1.Alycaeusconicus– Reeve 1878: pl. 1, species 9; Godwin-Austen 1914: 387–388, pl. 143, figs 4, 4a, 4b; Gude 1921: 208.Alycaeus (Alycaeus) conicus– Kobelt 1902: 342; Ramakrishna et al. 2010: 47.Type locality.
“Was abundant on the Limestone Hill east of Kopili river, North Cachar District, and was occasionally also found in other places, but rare”.
Material examined.
Samiamri, E of the Kopili R., leg. Godwin-Austen, NHMUK 1903.7.1.2674 (12 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 is similar to protoconch by being moderately glossy and sculptureless; R2 short, with regular ribs curved towards the aperture, forming a relatively wide, flat area, when viewed from above.
Jatinga valley, N. Cachar, NHMUK 1903.7.1.2675 (12 syntypes); Khasi Hills, NHMUK 1903.7.1.2565 (1 shell, included with type lot, but not mentioned in the original description and not considered as part of the type series).
Etymology.
The replacement name (jatingaensis) refers to the type locality (Jatinga Valley).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 also glossy without spiral lines; R2 relatively short, with regular ribs; each rib has a lamella-like horizontal projection towards their anterior neighbours.
Alycaeusconicusvar.nanus Godwin-Austen, 1914 is a primary homonym of Alycaeusnanus Möllendorff, 1886 (treated as a synonym of A.diminutus). Both taxa have been used as valid with this combination after 1899, thus, a replacement name is given to the junior homonym.
AnimaliaGastropodaCyclophoridae50659405-971A-5059-BF1A-099F3CB67458Dicharaxcrenatus(Godwin-Austen, 1871)AlycaeuscrenatusGodwin-Austen, 1871: 90–91, pl. 3, fig. 5.Alycaeuscrenatus– Reeve 1878: pl. 1, species 1, figs a, b; Godwin-Austen 1914: 388–389, pl. 143, figs 8, 8a, 8b.Alycaeus (Dicharax) crenatus– Kobelt 1902: 366; Gude 1921: 241.Chamalycaeus (Dicharax) crenatus– Ramakrishna et al. 2010: 57.Type locality.
“On Burrail Range, N. Cachar, at ca. 5000 feet”.
Material examined.
Mokarsa, Khasi Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2642 (2 syntypes). Note that the type locality does not match with the locality of the type sample, but the type locality was clarified in Godwin-Austen 1914 in 1897–1914: 389, and the drawing in the original description is identical with the two syntypes.
Remarks.
Protoconch rather glossy, low, without spiral lines; R1 rather regularly, finely ribbed without spiral striation; R2 relatively short, with regular ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridaeDF8EDCE3-CB6C-50AE-AECA-C5FE3A42FB01Dicharaxcristatus(Möllendorff, 1886)AlycaeuscristatusMöllendorff, 1886: 168, pl. 5, fig. 6.Alycaeus (Charax) cristatus– Kobelt and Möllendorff 1897: 150.Alycaeus (Dicharax) cristatus– Kobelt 1902: 367.Alycaeus (Chamalycaeus) smithiFulton, 1907: 157, pl. 10, fig. 5.Alycaeus (Charax) fimbriatusvar.simplicilabrisBavay & Dautzenberg, 1912: 53–54, pl. 6, fig. 18.Chamalycaeus (Dicharax) cristatus– Zilch 1957: 146, pl. 6, fig. 23.Dicharaxcristatus– Páll-Gergely et al. 2017: 34–43, figs 12C, D, 13C, 24–27, 28A–D, 29C, D, 30 (smithi Fulton, 1907 and simplicilabris Bavay & Dautzenberg, 1912 are synonyms).Type locality.
“in provinciae sinensis Hunan parte meridionali”
Material examined.
Süd-Hunan: China, coll. Möllendorff 1886, SMF 39231 (lectotype, designated by Yen 1939); Same data, SMF 39232 (11 paralectotypes); Same data, SMF 39233 (2 paralectotypes); for types of the synonymised names see Páll-Gergely et al. (2017).
Remarks.
Protoconch low, glossy, without spiral lines; R1 with regular, low ribs and without spiral striation; R2 relatively long, with ribs curved towards the aperture and reaching each other.
AnimaliaGastropodaCyclophoridae6D3E8243-24A5-5F13-B1B4-21DE74A7CA51Dicharaxcucullatus(Theobald, 1870)AlycaeuscucullatusTheobald, 1870: 396–397, pl. 18, fig. 2.Alycaeuscucullatus– Reeve 1878: pl. 2, species 12; Godwin-Austen 1914, Vol. II: 407, pl. 155, fig. 5.Alycaeus (Dicharax) cucullatus– Kobelt 1902: 367–368; Gude 1921: 244–245.Type locality.
Protoconch low, rather matte, no spiral lines visible; R1 regularly, strongly ribbed without spiral striation; R2 Relatively long, with all ribs curved towards the aperture, and they are almost in contact.
Damsang, W. Bhutan, leg. Robert, NHMUK 1903.7.1.2677 (12 syntypes).
Remarks.
Protoconch moderately glossy, low, no spiral lines visible; R1 with regular ribs and without spiral striae; R2 is moderately long, the upper part of the ribs are horizontal (in cross-sectional view the ribs are T-shaped); in most cases the ribs do not reach each other.
Protoconch low, glossy, very finely granulated, no spiral lines visible; R1 finely, regularly ribbed without spiral lines; R2 long, with strong signs of Byne’s disease; the ribs are curved towards the aperture and reach each other (typical Dicharax structure), forming a glossy, nearly smooth surface.
Pac-Kha, leg. Messager, MNHN-IM-2000-27165 (1 syntype); for additional specimens see Páll-Gergely et al. (2017).
Remarks.
Protoconch low; R1 glossy with regular, dense ribs, which gradually transform to an irregularly ribbed section having widely spaced ribs at end of R1; ribs low and blunt on whole shell; R2 very densely ribbed, ribs curved towards aperture.
Shells of a single sample had some spiral striation, which is highly unusual in this genus (Páll-Gergely et al. 2017)
AnimaliaGastropodaCyclophoridae0C950916-5FA8-5ADD-9946-B6DDDE4066DFDicharaxdiagonius(Godwin-Austen, 1871)AlycaeusdiagoniusGodwin-Austen, 1871: 88–89, pl. 3, fig. 2.Alycaeusdiagonus[sic] – Reeve 1878: pl. 1, species 2.Alycaeus (Dicharax) diagonius– Kobelt 1902: 368–369; Gude 1921: 247–248.Alycaeusdiagonius– Godwin-Austen 1914: 389–390, pl. 143, figs 5, 5a, 5b.Chamalycaeus (Dicharax) diagonius– Ramakrishna et al. 2010: 58.Type locality.
“The Diyung valley, north of Asálú, in Cachar District”.
Material examined.
Diyung valley, N. of Asalu, N. Cachar, coll. Godwin-Austen, NHMUK 1903.7.1.2678 (10 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 also without spiral lines, its sculpture is similar to that of the protoconch; R2 short, with regular ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridae54D2665E-8C43-5B06-B93C-B2C8AB393EFCDicharaxdigitatus(H. F. Blanford, 1871)AlycaeusdigitatusH. F. Blanford, 1871: 41–42, pl. 2, fig. 4.Alycaeus (Dicharax) digitatus– Kobelt 1902: 369; Gude 1921: 248.Alycaeusdigitatus– Godwin-Austen 1914: 339–340, pl. 134, figs 5, 5a.Chamalycaeus (Dicharax) digitatus– Ramakrishna et al. 2010: 59.Type locality.
“apud Darjeeling in vallo Rungno fluminis Himalayæ Sikkimensis”.
Material examined.
Rechila Pk., Sikkim, leg. W. Robert, NHMUK 1903.7.1.1253 (1 shell, probably not syntype, but figured by Godwin-Austen 1914).
Remarks.
Protoconch low, without spiral lines; R1 with very fine ribs, no spiral lines visible; R2 moderately long, ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridae4F07CCC8-E90E-5E84-AF00-66847D2B7059Dicharaxdiminutus(Heude, 1885)Alycoeus[sic] diminutus Heude, 1885: 96, pl. 24, figs 5, 5a.Alycaeusdiminutus– Heude 1886: 210.Alycaeus (Orthalycaeus) diminutus– Möllendorff 1886: 170.Alycaeus (Orthalycaeus) nanusMöllendorff 1886: 170, pl. 5, fig. 8.Alycaeus (Chamalycaeus) diminutus– Kobelt and Möllendorff 1897: 148; Kobelt 1902: 354.Alycaeusnanus– Tarruella and Domènech 2011: 72.Dicharaxdiminutus– Páll-Gergely et al. 2017: 50–52, figs 33A–C (nanus Möllendorff, 1886 is a synonym).Type locality.
“in ditione Tchen-k’eou”.
Material examined.
Hunan, China, coll. Möllendorff ex coll. Heude, SMF 39255 (1 syntype of A.diminutus [“minutus” on the label]); for additional specimens see Páll-Gergely et al. (2017).
Remarks.
Protoconch low, glossy, without spiral lines, R1 with regular, dense, low ribs, no spiral striation visible; R2 short, with dense ribs curved towards the aperture.
Malakka: Bukit Pondong (Perak), coll. Möllendorff, SMF 109476 (lectotype, designated by Zilch 1957); Same data, SMF 109477 (5 paralectotypes); Perak, leg. Hungerford, NHMUK 1891.3.17.779–782 (4 possible syntypes, these are labelled as types, but this is questionable).
Remarks.
Protoconch low, rather matte, without spiral lines; R1 with similar sculpture to that of the protoconch; R2 very short, with ca. 20 regular ribs, ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridaeEBEDD32F-8C21-5C33-B9A3-1BF9B1CD2E35DicharaxdracoPáll-Gergely & Hunyadi, 2017DicharaxdracoPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 52–54, fig. 34A.Type locality.
Holotype (HNHM 99705) and a few paratypes (see the original description).
Remarks.
Protoconch low, rather glossy; R1 with low, regular ribs, ribbing weaker at beginning of R1, but stronger at end of R1; R2 with ribs curved towards aperture.
Cheng-Kou County, Chong-qing, China, HMT-218a, deposited in IZCAS (syntype: labelled as lectotype, but probably there was no valid lectotype designation). No type specimens deposited in American museums were reported by Johnson (1973).
Remarks.
This species could be examined for the first time, since it was not examined in our previous paper (Páll-Gergely et al. 2017). Protoconch low, smooth; R1 without spiral striation, its beginning is densely, finely ribbed, which gradually changes to a more widely-spaced, strongly ribbed surface. R2 and R3 are of comparable length, R2 ribs dense, low, blunt, not elevated. The most similar species is D.fargesianus, which has denser ribs on R1 and R3, and more marked swelling on R3.
The examined specimen has a shorter R3 than the one illustrated by Heude (1890) (see also Páll-Gergely et al. 2017: fig. 69I). This raises some doubts about the identity of this type.
https://binary.pensoft.net/fig/468661AnimaliaGastropodaCyclophoridae7ABF13F3-94F8-57A7-819D-C4804156A485Dicharaxfimbriatus(Bavay & Dautzenberg, 1912)Alycaeus (Charax) fimbriatusBavay & Dautzenberg, 1912: 52–53, pl. 6, figs 14–17.ChamalycaeusplicilabrismultidentatusYen, 1939: 29, pl. 2, fig. 33.Dicharaxfimbriatus– Páll-Gergely et al. 2017: 54–61, figs 13E, 35–37, 38A–D, 39 (multidentatus Yen, 1939 is a synonym); Inkhavilay et al. 2019: 14, fig. 5B.Type locality.
“Pac-Kha”.
Material examined.
Pac-Kha, leg. Messager, MNHN-IM-2000-27166 (1 syntype); for additional specimens see Páll-Gergely et al. (2017).
Remarks.
Protoconch low, rather matte; R1 rather regularly ribbed with blunt, but strong ribs; rib density decreases towards the end of region; R2 extremely densely ribbed; ribs curved towards aperture, forming a nearly smooth surface.
Haut Tonkin, leg. Messager, MNHN-IM-2000-27168 (1 syntype), for additional specimens see Páll-Gergely et al. (2017).
Remarks.
Protoconch low, glossy; R1 coarsely, rather irregularly ribbed, ribs weaker on edge of body whorl; R2 very finely and densely ribbed, ribs low, curved towards the aperture.
Khasi Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2566 (2 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 glossy, no ribs or spiral striation visible; R2 very short, with only ca. 14 ribs; ribs curved towards the aperture, which do not reach each other.
“Phunggum, a Naga village at head of the Lanier valley, at 5,000 feet”.
Material examined.
Phunggum, Lahupa Naga Hills, Munipur, NHMUK 1903.7.1.2486 (13 syntypes).
Remarks.
Protoconch low, matte, no spiral lines visible; R1 with irregular, rough wrinkles, especially near the suture, but no spiral lines are visible; R2 relatively long, with regular ribs curved towards the aperture; the ribs are bent, nearly reach each other forming a nearly smooth surface.
Habiang Garo, W. Khasi, leg. Godwin-Austen, NHMUK 1903.7.1.2649 (1 syntype).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 glossy, without ribs and spiral lines; R2 very short, only ca. 13 ribs are present; the ribs are slightly curved towards the aperture at their tops, and do not reach each other (typical Dicharax).
AnimaliaGastropodaCyclophoridae8E82FB8F-578B-55F0-9F76-B6A3209407BCDicharaxhebes(Benson, 1857)AlycaeushebesBenson, 1857: 204–205.Alycaeushebes– Reeve 1878: pl. 6, species 52; Godwin-Austen 1886: 191, pl. 43, figs 1, 1a–c.Alycaeus (Dicharax) hebes– Kobelt 1902: 371; Gude 1921: 255.Alycaeushebes– Godwin-Austen 1914: 374–375, pl. 145, figs 5, 5a, 5b.Chamalycaeus (Dicharax) hebes– Ramakrishna et al. 2010: 61.Dicharaxhebes– Páll-Gergely et al. 2017: 10, fig. 5.Type locality.
“ad Teria Ghát”.
Material examined.
Khasi Hills, Teria Ghat, leg. Godwin-Austen, NHMUK 1903.7.1.2658 (17 specimens); Vorder-Indien, Khasi Berge, coll. Möllendorff, SMF 109244; NHMUK 1888.12.4.908-910, leg. Theobald, possible syntypes. The shell, which was believed to be a possible type specimen (No locality, UMZC I.102635), belongs to another Dicharax species.
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 with low, irregular growth ridges, but otherwise glossy without spiral lines; R2 moderately long, with regular ribs nearly reaching each other; the ribs near the beginning of R2 are bent in an anterior direction, the ribs near the end of R2 are bent in a posterior direction, and the ribs in the middle section of R2 are T-shaped in cross sectional view.
AnimaliaGastropodaCyclophoridaeE823CE0E-73E8-5CAA-A07D-E44A7BB1F1EADicharaxhumilis(W. T. Blanford, 1862)AlycaeushumilisW. T. Blanford, 1862: 136–137.Alycaeushumilis– Reeve 1878: pl. 5, species 40; Godwin-Austen 1914: 408–409, pl. 151, fig. 8.Alycaeus (Dicharax) humilis– Kobelt 1902: 372; Gude 1921: 255–256.Type locality.
“ad Akouktoung, ad ripas fluminis Irawaddi, in provincia Burmana Pegu”.
Material examined.
River Bank, Myanoung, Pegu, NHMUK 1906.4.4.69 (1 shell); Pegu, coll. C. Bosch ex coll. H. Rolle, SMF 192340 (4 shells).
Remarks.
The only available specimen housed in the NHM was weathered; Protoconch low, with any recognisable sculpture; R1 with irregular, fine ribbing which turns into a widely spaced, strongly ribbed area at the end of the region, no spiral lines visible; R2 relatively short, weathered. SMF sample: protoconch low, rather glossy; R1 also glossy, with widely spaced, strongly ribs near the end of the region, no spiral lines visible; R2 relatively short, ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridae2C1CA3A4-190C-550B-A34C-5F89DBDCB46DDicharaximitatorPáll-Gergely & Hunyadi, 2017DicharaximitatorPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 64–65, figs 34B, 38E, F, 42A, B.Type locality.
“China, Guangxi, Bose Shi, Leye Xian, Moli Cun, cliffs S of the village on the left side of the Buliu River, 540 m, 24°39.436'N, 106°43.245'E”.
Material examined.
Holotype (HNHM 99706) and a few paratypes, see Páll-Gergely et al. (2017).
Remarks.
Protoconch without any recognisable sculpture, although it was weathered in examined shells; R1 smooth, glossy, with sharp, widely spaced, regular ribs near suture and inside umbilicus; R2 finely, densely ribbed, ribs are curved towards aperture at end of R2, but in curved in posterior direction at beginning of R2.
Nongjinghi, Jiantia Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2686 (14 syntypes).
Remarks.
Protoconch low, without spiral lines; R1 smooth except for some rough wrinkles near the suture, no spiral striation visible; R2 moderately long, with regular ribs, which are curved towards the aperture.
Protoconch matte, R1 smooth, no spiral lines visible (although the entire shell is somewhat weathered); R2 of normal length, the ribs are overall low, they are slightly curved towards the aperture.
Kezama, Naga Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2556 (1 syntype).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 with strong ribs and without spiral striation; R2 moderately long, ribs curved towards the aperture (typical Dicharax structure).
Protoconch low, glossy, no spiral lines visible; R1 glossy, with some spiral lines, which are, however, not present on the surface but are found on parts of the inner layers of the shell and visible through the semi-transparent upper layer (thus, not homologous with the spiral striation of other genera); R2 short, with lamella-like, sharp ribs, which are slightly curved towards the aperture; there is quite large gap between the ribs.
AnimaliaGastropodaCyclophoridae7B13D96B-6D2A-59A1-931B-4564FEFF7C30Dicharaxlongituba(E. von Martens, 1864)AlycaeuslongitubaE. von Martens, 1864: 1 20.Alycaeuslongituba– E. von Martens 1867: 151, pl. 4, fig. 8.Alycaeus (Dicharax) longituba– Kobelt 1902: 373.Chamalycaeuslongituba– van Benthem Jutting 1948: 573–575, fig. 29.Type locality.
“Sumatra bei Kepahiang”. Later (Martens 1867) more precisely: “Sumatra, am Ostabhang der mittleren Bergkette bei Kepahiang”.
Material examined.
Mt Gede, West Java, 4000 ft., H. Fruhstorfer, 1898, E. R. Sykes Collection, Acc. no. 1825, NHMUK 20150127 (3 shells).
Remarks.
Protoconch low, no spiral lines visible; R1 rather regularly, finely ribbed without spiral lines; R2 very long, with regular ribs, which are curved towards the aperture, and reach each other (typical Dicharax).
AnimaliaGastropodaCyclophoridae1EE219F4-6179-52FB-B82C-C4E51AC5310EDicharaxmaosmaiensis(Godwin-Austen, 1922)AlycaeusmaosmaiensisGodwin-Austen, 1922: 365, text figs.Type locality.
“Khasi Hills, near Cherrapunji, at the mouth of the Maosmai cave”.
Material examined.
Maosmai, nr Cherrapoonjee, Khasi, NHMUK 20191067 (1 syntype separated in a vial with pink wool + 4 additional syntypes).
Remarks.
Protoconch low, rather matte, no spiral lines visible; R1 with very widely spaced wrinkles without spiral striation; R2 moderately long, the ribs are bent and do not reach each other (typical Dicharax structure).
AnimaliaGastropodaCyclophoridaeEF211232-F41C-5A09-80D0-BB7EAC0F8B55DicharaxmicrocostatusPáll-Gergely, 2017DicharaxmicrocostatusPáll-Gergely in Páll-Gergely et al., 2017: 66, fig. 34C.Type locality.
“China, Sichuan, Taian Zhen, Qingchenghoushan, Dujiangyan Shi, Cuiyinghu to upper station of Jinli cable station, 1273 m, 30°56.27110'N, 103°28.75198'E”.
Material examined.
Holotype (HNHM 99708) and a few paratypes, see the original description.
Remarks.
Protoconch low, we only had weathered material available to study and therefore the sculpture could not be examined; R1 regularly, finely ribbed; R2 very densely ribbed, ribs curved towards aperture.
Malakka, Bukit Pondong (Perak), coll. Möllendorff, SMF 109496 (lectotype, designated by Zilch 1957); Same data, SMF 109497 (3 paralectotypes); Perak, leg. Hungerford, NHMUK 1891.3.17.794–796 (3 possible paralectotypes).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 irregularly, finely ribbed without spiral lines; R2 with ribs curved towards the aperture (typical Dicharax).
AnimaliaGastropodaCyclophoridae2D0C614C-AA34-55F2-B643-7759A79F83D1DicharaxmicropolitusPáll-Gergely & Hunyadi, 2017DicharaxmicropolitusPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 66–68, figs 34D, 42C, D, 43.Type locality.
“China, Sichuan, Taian Zhen, Qingchenghoushan, Dujiangyan Shi, Cuiyinghu to upper station of Jinli cable station, 1273 m, 30°56.27110'N, 103°28.75198'E”.
Material examined.
Holotype (HNHM 99709) and a few paratypes, see the original description.
Remarks.
Protoconch low, glossy; R1 almost smooth, with only very inconspicuous, irregular growth lines; R2 very densely ribbed, ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridaeD4B249CC-5EF9-51F1-BA17-4CBD52DF76B4Dicharaxnitidus(W. T. Blanford, 1862)AlycaeusnitidusW. T. Blanford, 1862: 141.Alycaeusnitidus– Reeve 1878: pl 3, species 25; Godwin-Austen 1914: 421–422, pl. 151, figs 4, 4a.Alycaeus (Chamalycaeus) nitidus– Kobelt 1902: 360; Gude 1921: 230–231.Chamalycaeus (Chamalycaeus) nitidus– Ramakrishna et al. 2010: 54.Type locality.
“prope Tongoop in Arakan”.
Material examined.
Manya Khyoung, Arakan, coll. Blanford, NHMUK 1906.4.4.54 (3 possible syntypes).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 glossy, without spiral lines; R2 short, with a few ribs; each rib lamella-like ribs, which is slightly curved towards the aperture.
AnimaliaGastropodaCyclophoridaeD108A9B7-ED2F-598A-AB8A-56A089BA6418Dicharaxnotatus(Godwin-Austen, 1876)AlycaeusnotatusGodwin-Austen, 1876: 176, pl. 7, figs 9, 9a, 9b.Alycaeusnotatus– Godwin-Austen 1886: 191–192, pl. 43, figs 2, 2a–c; Godwin-Austen 1914: 358–359, pl. 145, figs 8, 8a.Alycaeus (Dicharax) notatus– Kobelt 1902: 374; Gude 1921: 262.Chamalycaeus (Dicharax) notatus– Ramakrishna et al. 2010: 64; Tripathy et al. 2018: 789.Type locality.
“On the slopes of Torúpútú Peak at 3000 feet”.
Material examined.
Toruputu Peak, Dafla Hills, 3000, NHMUK 1903.7.1.2672 (4 syntypes); Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2544 (2 syntypes). Both samples are in the same box, but in different vials.
Remarks.
Protoconch low without spiral lines; R1 irregularly, strongly ribbed without spiral striation; R2 long, with dense ribs which are curved towards the aperture, but do not usually reach each other.
Fort Stedman, Burma, coll. Woodthorpe, NHMUK 1903.7.1.3065 (15 syntypes).
Remarks.
Protoconch low, without spiral lines; R1 rather regularly, weakly ribbed without spiral striae; R2 relatively short, with ribs curved towards the aperture that reach each other.
Koliaghur nr. Tezpur, Assam, NHMUK 1903.7.1.2682 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, without spiral lines; R1 nearly smooth, there are some rough wrinkles near the suture; R2 short, with ribs, which are curved towards the aperture that reach each other.
Malakka: Perak, coll. Möllendorff, SMF 109226 (lectotype, designated by Zilch 1957); Same data, SMF 109227 (2 paralectotypes).
Remarks.
Protoconch low, without spiral striae; R1 with widely spaced, strong ribs, which are the most prominent near the suture and become lower away from it; R2 very short, consists of ca. 15 ribs, which are curved towards the aperture.
Malakka: Bukit Pondong (Perak), coll. Möllendorff, SMF 109507 (lectotype, designated by Zilch 1957); Same data, SMF 109508 (4 paralectotypes).
Remarks.
Protoconch low, without spiral striae; R1 finely, regularly ribbed, without spiral striae; R2 extremely short, with ca. six ribs, which are curved towards the aperture.
Protoconch low, rather matte; R1 regularly, densely ribbed, ribs low; ribs becoming slightly more widely spaced towards end of R1; R2 very densely ribbed, ribs curved towards the aperture.
AnimaliaGastropodaCyclophoridaeB547A7EE-155F-5485-89A2-487B8AF83FFADicharaxplicilabris(Möllendorff, 1886)AlycaeusplicilabrisMöllendorff, 1886: 167, pl. 5, fig. 5.Alycaeus (Chamalycaeus) plicilabris– Kobelt and Möllendorff 1897: 149; Kobelt 1902: 361.Chamalycaeusplicilabrisplicilabris– Yen 1939: 29, pl. 2, fig. 32.Chamalycaeus (Chamalycaeus) plicilabrisplicilabris– Zilch 1957: 145, pl. 5, fig. 14.Dicharaxplicilabris– Páll-Gergely et al. 2017: 68–70, fig. 33D.Type locality.
“in provincia sinensi Hunan”.
Material examined.
China, Prov. Hunan, coll. O. Boettger ex coll. Möllendorff, SMF 39229 (lectotype, designated by Yen 1939); same data, SMF 39229 (4 paralectotypes).
Remarks.
Protoconch low, rather glossy; R1 regularly, densely ribbed; R2 with ribs curving towards the aperture.
AnimaliaGastropodaCyclophoridae2F27C70E-FFA0-5F20-9133-2F653082C751Dicharaxpolitus(W. T. Blanford, 1865)AlycaeuspolitusW. T. Blanford, 1865: 83–84.Alycaeuspolitus– Reeve 1878: pl. 5, species 39; Godwin-Austen 1914: 422, pl. 139, figs 5, 5a; Gude, 1921: 214–215.Alycaeus (Alycaeus) politus– Kobelt 1902: 348.Type locality.
Protoconch low, glossy, no spiral lines visible; R1 glossy, without notable sculpture; R2 short, with regular ribs, which are curved towards the aperture.
HNHM 99704 (holotype) and a few paratypes, see original description.
Remarks.
Protoconch, normally elevated (not higher or lower than what would be expected from the overall shell shape), it is matte, without any notable sculpture; R1 regularly ribbed; in fresh shells ribs sharp and strongly curved towards aperture; R2 very densely ribbed, ribs with T-shaped cross sectional view.
“Thailand, Chiang Rai Province, Doi Tung, 50 m before Wat Phra That Doi Tung, around the car park, 1350 m a.s.l., 20°19.540'N, 99°49.987'E”.
Material examined.
HNHM 100178 (holotype).
Remarks.
Protoconch glossy; R1 also glossy, with irregular growth lines; R2 bears dense, curved ribs (ca. 46–48 in total), for more details see the original description.
S. Sylhet Hills, leg. W. Channel, NHMUK 1903.7.1.55 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, no spiral lines visible; R1 with widely spaced ribs with the very slight indication of spiral lines; R2 relatively short, with some blunt (weathered), regular ribs, which have lamella-like horizontal projections reaching to the neighbouring ribs (typical Dicharax).
“Tongmai Town, (30°01'N, 95°E), Bomi County, Tibet Autonomous Region, China”.
Material examined.
CASIZ TM 0010054 (holotype) deposited in IZCAS: Tong-Mai Town, Bo-Mi County, Tibet Autonomous Region, China, leg. Chen De-niu, 1980.6.20; CASIZ TM 0010056 (paratype): same as holotype.
Remarks.
Protoconch low, without notable sculpture; R1 rather dense, low ribs, no spiral striae visible; R2 + R3 90° combined; R2 with low ribs, ribs similar to those on R1; R3 with a prominent, blunt swelling.
The shell of Dicharaxtangmaiensis is similar to some other northeastern Indian Dicharax species with fringed peristome (e.g., D.cucullatus). Future investigation should reveal whether this species is really distinct from other Himalayan species, since no comparisons were made in the original description.
https://binary.pensoft.net/fig/468662AnimaliaGastropodaCyclophoridae838693A7-69A7-5590-825F-BEEB773C3E30Dicharaxtheobaldi(W. T. Blanford, 1862)AlycaeusTheobaldiW. T. Blanford, 1862: 142–143.Alycaeustheobaldi– Reeve 1878: pl. 5, species 44; Godwin-Austen 1914: 359–360, pl. 149, figs 3, 3a, 3b; Godwin-Austen 1914: 382–383, pl. 145, figs 4, 4a.Alycaeus (Dicharax) theobaldi– Kobelt 1902: 377–378; Gude 1921: 272–273.Chamalycaeus (Dicharax) theobaldi– Ramakrishna et al. 2010: 67.Type locality.
“in montibus Khasi”.
Material examined.
Khasi Hills, coll. W. T. Blanford, NHMUK 1906.4.4.60 (2 possible syntypes).
Remarks.
Protoconch low, lacks any signs of spiral striation; R1 with widely spaced ribs but no spiral lines; R2 short, with ribs which are curved towards the aperture.
Protoconch somewhat elevated but lacks any signs of spiral striation; R1 with widely spaced ribs but no spiral lines; R2 short, with ribs which are curved towards the aperture.
AnimaliaGastropodaCyclophoridae4ADADD11-73EC-5E23-8B45-1DDB3B0E9E21Dicharaxvestitus(W. T. Blanford, 1862)AlycaeusvestitusW. T. Blanford, 1862: 138–139.Alycaeusvestitusvar.minorW. T. Blanford, 1862: 138.Alycaeusvestitus– Reeve 1878: pl. 1, species 3; Godwin-Austen 1914: 424–425, pl. 139, figs 2, 2a; Gude 1921: 220–221.Alycaeus (Alycaeus) vestitus– Kobelt 1902: 352.Dicharaxvestitus– Páll-Gergely et al. 2017: 71.Type locality.
“in montibus Arakanensibus”.
Material examined.
Moditoung, NHMUK 1906.4.4.53 (holotype [single specimen of both the nominotypical form and var. minor were mentioned in the original description]), and two additional non-type specimens in the same lot from Alori Khyoung and Mamya Khyoung.
Remarks.
All three specimens are strongly weathered; therefore, their sculpture could not be fully distinguished. Protoconch low, without recognisable sculpture; R1 seemingly smooth; R2 with dense ribs, which were all broken.
Protoconch low, matte; R1 with very low, but rather regular ribbing near the suture (ribs nearly absent at the edge of the body whorl); R2 of normal length, the ribs are curved towards the aperture, nearly reaching each other.
Atypical or questionable Dicharax speciesAnimaliaGastropodaCyclophoridae91825F23-4F44-50BC-9341-01B2BD3E0E36Dicharax(?)abdouiPáll-Gergely, 2017DicharaxabdouiPáll-Gergely in Páll-Gergely et al., 2017: 14, fig. 6.Dicharaxabdoui– Inkhavilay et al. 2019: 14, fig. 4F.Type locality.
“Laos, Khammouane Province, approx. 9 km NE of Thakhek (Muang Khammouan), 190 m, 17°26.757'N, 104°52.937'E, on and under rocks in dry secondary forest on and under NW exposed cliffs”.
Material examined.
MNHN IM-2012-27329 (holotype) and 2 paratypes (MNHN-IM-2012-27328).
Remarks.
Protoconch low, nearly smooth, with extremely fine pits arranged in spiral rows (not homologous with the spiral striation of Metalycaeus species); R1 nearly smooth, with low, widely spaced ribs near suture and in umbilicus; R2 very short, with low, dense regular ribs (ca. 20).
“Barowli Gorge, Durrang District, Assam, foot of the Akha Hills”.
Material examined.
Akha Hills, Barowli River, coll. Godwin-Austen, NHMUK 1903.7.1.2683 (holotype [single specimen mentioned in the original description]).
Remarks.
The entire shell is quite weathered, but the following observations could be made: protoconch low, without spiral striae; R1 glossy, with widely spaced, strong ribs (present only near the suture) and without spiral lines; R2 short, with dense, low ribs; R2 of fresh shells is probably smooth.
AnimaliaGastropodaCyclophoridae0F5F409C-311A-5AF2-ABEB-2D0726781C72Dicharax(?)alticolaPáll-Gergely & Hunyadi, 2017DicharaxalticolaPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 14–20, figs 8A, 9A–D, 10A, B, 11, 12A, B, 13A, B.Type locality.
Holotype (HNHM 99702) and several paratypes (see Páll-Gergely et al. 2017).
Remarks.
Protoconch low, seemingly smooth but rather matte; R1 somewhat regularly ribbed; rib density of R2 higher than that of R1, ribs on R2 low, not curved, rather sharp, connected to each other near tube (similar to Japanese “Awalycaeus” and “Cipangocharax” species); for more details see the original description.
Ataran, Burma, ex Dr. F. Stoliczka, NZSI M.8073 (holotype [single specimen mentioned in the original description]).
Remarks.
The holotype is in a strongly corroded state; therefore, the sculpture could not be examined in detail. This species is putatively placed in Dicharax due to the overall smooth shell and the fringed aperture.
Dicharaxataranensis (Godwin-Austen, 1914), holotype (NZSI M.8073). All images: Sheikh Sajan.
https://binary.pensoft.net/fig/468663AnimaliaGastropodaCyclophoridae4D0B0942-6276-5A10-BF00-6269FCB9DF99Dicharax(?)barowliensis(Godwin-Austen, 1914)AlycaeusbarowliensisGodwin-Austen, 1914: 352, pl. 141, fig. 4.Alycaeusbarowliensis– Gude 1921: 205.Alycaeus (Alycaeus) burowliensis[sic] – Ramakrishna et al. 2010: 46.Type locality.
“Barowli River, Akha Hills, Durrang, Assam”.
Material examined.
Barowli R. Durrang, Assam, coll. Godwin-Austen, NHMUK 1903.7.1.2723 (holotype [single specimen mentioned in the original description]).
Remarks.
Only the holotype is known. The outermost shell layer is entirely weathered and the sculpture is not visible. The protoconch is seemingly low. Based on this character, A.barowliensis is tentatively classified in the genus Dicharax.
“India, Karnataka State, Chamarajanagar District, Malai Mahadeshwara Hills, 1010 m a.s.l., 12.04911°N, 77.56369°E, from the base of a big tree, next to the road near the temple (the habitat has lots of lianas and stones with a good amount of litter in dry deciduous forest)”.
Material examined.
ZSI/WGRC/9865 (holotype), for other examined shells see the original description.
Remarks.
Protoconch somewhat elevated, rather glossy without notable sculpture; first whorl of R1 irregularly, finely ribbed, with ribs becoming stronger, rarer and more regular towards end of R1; R2 with 24–28 elevated, blunt, regular ribs; for description of cross-sectional view see original description.
AnimaliaGastropodaCyclophoridaeD126CCD0-A814-5759-A5F2-E6D06D481E3DDicharax(?)bicrenatus(Godwin-Austen, 1874)AlycaeusbicrenatusGodwin-Austen, 1874: 148, pl. 3, fig. 5.Alycaeus (Dicharax) bicrenatus– Kobelt 1902: 365; Gude 1921: 238–239.Alycaeusbicrenatus– Godwin-Austen 1884: pl. 51, fig. 4; Godwin-Austen 1914: 386–387, pl. 144, figs 5, 5a, 5b.Chamalycaeus (Dicharax) bicrenatus– Ramakrishna et al. 2010: 56.Type locality.
“Kopamedza Peak Naga Hill, 8–9,000 feet, in forest”.
Material examined.
Kopamedza, Naga Hills, NHMUK 1903.7.1.2490 (7 syntypes in two vials).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 very finely, regularly ribbed without spiral lines; R2 moderately long, with regular ribs, which are curved towards the aperture, however the space between the ribs is much larger than in typical Dicharax.
AnimaliaGastropodaCyclophoridae46A3DFAD-1027-591F-89DB-2D09989F269CDicharax(?)bifrons(Theobald, 1870)AlycaeusbifronsTheobald, 1870: 396, pl. 18, fig. 1.Alycaeusbifrons– Reeve 1878: pl. 6, species 48; Godwin-Austen 1914: 407, pl. 139, figs 3, 3a.Alycaeus (Dicharax) bifrons– Kobelt 1902: 365–366; Gude 1921: 239.Type locality.
Protoconch low, rather matte, no spiral lines visible; R1 irregularly wrinkled near the suture, this sculpture becomes stronger anteriorly, and near the end of R1 there are widely spaced, strong ribs, which extend not only to the suture area but to the edge of the body whorl; no signs of spiral striae visible on R1; R2 moderately long, with widely spaced, lamella-like, straight, rather low ribs.
Shells of Dicharax(?)birugosus (Godwin-Austen, 1893) AAlycaeuscanaliculus Godwin-Austen, 1914, (NHMUK 1903.7.1.2764) BA.birugosus Godwin-Austen, 1893, syntype (NHMUK 1903.7.1.2628). Photographs: Kevin Webb (NHM).
https://binary.pensoft.net/fig/468664Remarks.
Alycaeusbirugosus and A.canaliculus are practically identical and both of them inhabit the Khasi Hills. Thus, the latter is moved to the synonymy of the former.
Protoconch low, rather glossy, without spiral lines; R1 without spiral lines; R2 short, with blunt, straight ribs. Specimens labelled as “birugosus var.” are smooth on R1, whereas typical shells are more strongly sculptured.
Comments relating to “var. minor”: protoconch low, rather glossy, without spiral lines; R1 glossy, without spiral lines; R2 short, with regular, blunt, not bent ribs.
Chwegalé, Arakan Hills, NHMUK 1906.4.4.177 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, no spiral lines visible; R1 without spiral lines; R2 long, with widely spaced but blunt ribs, which are curved towards the aperture (especially near the tube, far from the tube the ribs are straighter); curved ribs are situated far apart from each other.
The single shell is strongly weathered. Protoconch low, its sculpture is not visible; there are no signs of spiral lines on R1; R2 with dense ribs, their fine structure is not visible. Based on the low protoconch this species is putatively classified in Dicharax.
“Indonesia, East Java: Malang, Sempu Island, limestone rocks in lowland rainforest at entrance of Kelabang Cave, 44 m a.s.l., 8°26'58"S 112°41'28"E”.
Material examined.
Photographs of the holotype (MZB 19025) were examined.
Remarks.
Protoconch low without spiral striae; R1–R3 smooth but spirally striated on the umbilical side. This spiral striation is assumed not to be homologous with that of Metalycaeus species (i.e., it is probably part of the lower shell layers, not elevated from the shell surface), and similar to the structure observed in some D.depressus shells (see Páll-Gergely et al. 2017). R2 smooth from above, with ca. seven narrow lines, no elevated ribs discernible.
No specimens were examined. The general shape and the sculpture of the species is similar to Dicharaxlongituba according to the original description. Therefore, Alycaeuscrassicollis is tentatively classified in Dicharax.
AnimaliaGastropodaCyclophoridaeF39A5CAB-EAD3-59E0-A7BB-9CABD571E526Dicharax(?)crispatus(Godwin-Austen, 1871)AlycaeuscrispatusGodwin-Austen, 1871: 91–92, pl. 4, fig. 1.Alycaeuscrispatus– Godwin-Austen 1875: 8, pl. 4, fig. 3.Alycaeus (Dicharax) crispatus– Kobelt 1902: 367; Gude 1921: 242–243.Alycaeuscrispatus– Godwin-Austen 1914: 371–372, pl. 145, figs 1, 1a, 1b; Godwin-Austen 1914: 389.Chamalycaeus (Dicharax) crispatus– Ramakrishna et al. 2010: 58; Tripathy et al. 2018: 789.Type locality.
“Khasia, Jiantia and N. Cachar Hills”.
Material examined.
Shibak, Habiang Garo Hills, NHMUK 1903.7.1.2635 (5 syntypes); Same container (probably same locality), NHMUK 1903.7.1.2759 (11 syntypes).
Godwin-Austen (1914: 372) explained that the Alycaeuscrispatus variety from north Cachar in his previous paper (Godwin-Austen 1871: 93) was renamed A.asaluensis. The originally figured sample (Godwin-Austen 1871: pl. 4, fig. 1) is from Shibak, Gabir valley (Godwin-Austen, 1914: 372).
Remarks.
Protoconch moderately elevated, matte, no spiral lines visible; R1 rather regularly ribbed, also without spiral lines; R2 relatively short, with regular, widely spaced, sharp ribs.
The placement of the species in the genus Dicharax is based on the absence of spiral striation on the entire shell; however, the sharp R2 ribs are characteristic of the genus Chamalycaeus. The shape of protoconch shows some variation within species. Namely, typical crispatus and typical cristatusminimus shells have only slightly elevated protoconchs, whereas it is characteristically Chamalycaeus-like (strongly elevated) in D.crispatusmakarsae specimens.
Habiang Garo Hills, W. Khasi, leg. Godwin-Austen, NHMUK 1906.4.4.176 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch rather low, R1 with strong, widely spaced ribs which are most prominent near the suture and disappear on the edge of the body whorl; R2 of normal length, ribs blunt, and at the anterior end of the region ribs curved towards aperture.
Rywuk, Garo Hills, South base of, coll. Godwin-Austen, NHMUK 1903.7.1.2637 (2 syntypes).
Remarks.
Protoconch moderately elevated, smooth; R1 with strong, widely spaced ribs without spiral striation; R2 of normal length and ribs curved towards aperture.
AnimaliaGastropodaCyclophoridae9A13BCB8-4A4E-55BF-9F1B-02B1EF28E127Dicharax(?)daflaensis(Godwin-Austen, 1876)AlycaeusdaflaensisGodwin-Austen, 1876: 176–177, pl. 7, figs 12, 12a, 12b.Alycaeus (Dicharax) daflaensis– Kobelt 1902: 368; Gude 1921: 245–246.Alycaeusdaflaensis– Godwin-Austen 1914: 354–355, pl. 145, figs 11, 11a, 11b.Chamalycaeus (Dicharax) daflaensis– Ramakrishna et al. 2010: 58; Tripathy et al. 2018: 789.Type locality.
“Torúpútú Peak, 7000 feet”.
Material examined.
Toruputu Peak, Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2497 (lectotype: here designated, and 6 paralectotypes). The type sample contained two vials, one with two and the other with four shells. The one with four shells contained three larger shells and one which was conspicuously smaller. That smaller shell differs from the larger ones in terms of other shell characters, such as the spire height (lower than the others), the sculpture of R1 (smoother than the others), the strength of the swelling on R3 (less elevated than that of the others), and the lobes of the peristome (less conspicuous than those of the others). Therefore, one of the larger shells is selected here as lectotype to avoid further confusion.
Remarks.
Protoconch low, rather matte, no spiral lines visible; R1 irregularly, finely, densely ribbed, some spiral lines visible but these are probably part of the layer below the outermost one; R2 relatively short, smooth, only lighter, narrow and slightly thicker, darker stripes alternating.
We could not find the type specimens in the NHM. According to the original description they are in Aldrich’s collection, which is housed in the Michigan Museum (Dance 1986). We contacted the Dr. Taehwan Lee (Michigan Museum) who reported that the type sample of A.dohertyi could not be found in the UMMZ. We classify this species in Dicharax because the original description did not mention spiral striation, which rules out Metalycaeus, and mentions that it has a rather long tube, which rules out Cycloryx (= Pincerna). The crenulated peristome is characteristic for many Dicharax species.
No type specimen housed in American museums were reported by Johnson (1973). The non-type specimen figured by Yen (1939: pl. 2, fig. 37), is similar to “Chamalycaeus” helicodes (= synonym of Metalycaeusmuciferus, see Páll-Gergely et al. 2017). This species was putatively classified into the genus Dicharax by Páll-Gergely et al. (2017).
Tsanspu Valley, leg. Oakes, NHMUK 1903.7.1.3582 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, no spiral lines visible; R1 glossy, with rough wrinkles near the suture and without any spiral lines; R2 very short, with alternating thicker/darker and narrow/lighter stripes; overall surface of R2 smooth.
Protoconch low, no spiral lines visible; R1 glossy, without any notable sculpture; R2 very short, with alternating thicker darker, and narrower lighter stripes; overall surface of the region smooth.
Naraindhur, Cachar, leg. F. Ede, NHMUK 1903.7.1.1665 (8 syntypes in 2 vials).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 without spiral striation; R2 very long, ribs very slender, relatively sharp, straight; at the edge of the body whorl space between ribs is ca. 3–4 × larger than the ribs themselves.
AnimaliaGastropodaCyclophoridae97BEF8A6-C8CB-5502-8BA9-EB5E699B0658Dicharax(?)ellipticusPáll-Gergely, 2017DicharaxellipticusPáll-Gergely in Páll-Gergely et al. 2017: 23–25, figs 15A, B, 16.Type locality.
“Vietnam, Quang Ninh Province, Ha Long Bay area, Tien Ong Cave on Hang Trai Island, collected inside the cave, 20°48.96'N, 107°07.33'E”.
Material examined.
RMNH 5004014 (holotype) and some other paratypes, see the original description.
Remarks.
Protoconch low, smooth, glossy; R1 also smooth, glossy, R2 also entirely smooth, with alternating narrow light (= microtunnels) and thick dark (= area between the microtunnels) transverse stripes.
AnimaliaGastropodaCyclophoridaeAD88A9AC-16F4-597E-B2B1-4930F3936EBCDicharax(?)expatriatus(W. T. Blanford & H. F. Blanford, 1860)AlycaeusExpatriatusW. T. Blanford & H. F. Blanford, 1860: 123–124.Alycaeusexpatriatus– Reeve 1878: pl. 5, species 45; Godwin-Austen 1914: 433–434.Alycaeus (Dicharax) expatriatus– Kobelt 1902: 369–370; Gude 1921: 250–251.Chamalycaeus (Dicharax) expatriatus– Ramakrishna et al. 2010: 59.Chamalycaeusexpatriatus– Raheem et al. 2014: 46, fig. 25C.Dicharax(?)expatriatus – Aravind & Páll-Gergely 2018: 59, fig 1B.Type locality.
“Haud raro ad Neddoowuttom ghat, ad latus septentrionale montium “Nilgiri” Indiæ australis et circa 3000–4000 ped. alt.”.
Material examined.
Neddiwuttom, Nilgiris, NHMUK 1906.4.4.58 (lectotype, hereby designated and 5 paralectotypes).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 rather glossy, with low, irregular growth ridges and without spiral striae; R2 short with low, widely spaced ribs curved towards aperture.
The lectotype designation of Aravind and Páll-Gergely (2018) was invalid because no explicit statement was made to designate the lectotype. Thus, we here designate that shell to be the lectotype, which was named as such and figured by Aravind & Páll-Gergely (2018: fig. 1B).
China, Tchen-k’eou, MCZ 167229 (4 syntypes). According to Johnson (1973) paratypes are also present in the USNM (inv. number: 472341). These were not examined by us.
Remarks.
Protoconch low, rather matte; R1 regularly, densely ribbed; R2 extremely densely ribbed, ribs low, not typical to Dicharax.
AnimaliaGastropodaCyclophoridaeDFB497E9-31EB-51F4-83F9-6C46EAF949F8Dicharax(?)footei(W. T. Blanford & H. F. Blanford, 1861)AlycaeusFooteiW. T. Blanford & H. F. Blanford, 1861: 348, pl. 1, fig. 3.Alycaeusfootei– Reeve 1878: pl. 4, species 35; Godwin-Austen 1914: 432–433Alycaeus (Dicharax) footei– Kobelt 1902: 370; Gude 1921: 251–252.Chamalycaeus (Dicharax) footei– Ramakrishna et al. 2010: 60.Chamalycaeusfootei– Raheem et al. 2014: 46, Fig. 25D.Dicharax(?)footei – Aravind & Páll-Gergely 2018: 61, figs 1C, D.Type locality.
“Habitat in montibus Kolamulliis dictis”.
Material examined.
Kolamalai Hills, nr. Trichinopoly, coll. W.T. Blanford, NHMUK 1906.4.4.57 (lectotype, hereby designated); for additional examined specimens see Aravind & Páll-Gergely (2018).
Remarks.
Protoconch low, without spiral structure; R1 smooth, glossy, without spiral striation; R2 long, with straight, low ribs; in some specimens, especially near the beginning of the tube, the ribs look as if they have been “pushed” in an anterior direction (ribs similar to typical Dicharax, but lower).
The lectotype designation of Aravind and Páll-Gergely (2018) was invalid, because no explicit statement was made to designate the lectotype. Thus, we here designate that shell to be the lectotype, which was named as such, and figured by Aravind & Páll-Gergely (2018: fig. 1D).
AnimaliaGastropodaCyclophoridaeE2195F0A-241E-5025-884C-6935264FABF3Dicharax(?)gemma(Godwin-Austen, 1914)AlycaeusgemmaGodwin-Austen, 1914: 355–356, pl. 149, figs 6, 6a.Alycaeus (Dicharax) gemma– Gude 1921: 252.Chamalycaeus (Dicharax) gemma– Ramakrishna et al. 2010: 60; Tripathy et al. 2018: 789.Type locality.
“No .7 Camp, Dikrang Valley, Dafla Hills”.
Material examined.
No. 7. camp, Dikrang Valley, Dafla, NHMUK 1903.7.1.2601 (3 syntypes in two vials).
Remarks.
All three available shells were weathered, but the following observations could be made: protoconch low, without spiral lines; R1 with rough wrinkles near the suture, but the region is without spiral lines; R2 very short, the fine structure of the ribs could not be observed.
AnimaliaGastropodaCyclophoridaeC894D558-72AD-544E-8432-BA2045ED30EDDicharax(?)gemmula (Benson, 1859)AlycaeusgemmulaBenson, 1859: 179–180.Alycaeusgemmula– Reeve 1878: pl. 5, species 37; Godwin-Austen 1886: 190, pl. 48, figs 4, 4a–c.Alycaeus (Dicharax) gemmula– Kobelt 1902: 370–371; Gude 1921: 252–253.Alycaeus (Charax) gemmula– Godwin-Austen 1914: 340.Chamalycaeus (Dicharax) gemmula– Ramakrishna et al. 2010: 60; Tripathy et al. 2018: 789.Type locality.
“in valle Rungun”.
Material examined.
Darjling, coll. Blanford, NHMUK 1906.4.4.55 (1 specimen, labelled as “typical”, holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 also glossy, R2 moderately long, with thicker darker and very narrow, lighter stripes alternating; overall R2 surface nearly smooth.
AnimaliaGastropodaCyclophoridaeF2786BFE-431E-5837-AAB6-514876F80DF4Dicharax(?)glaber(W. T. Blanford, 1865)AlycaeusglaberW. T. Blanford, 1865: 84.Alycaeusglaber– Reeve 1878: pl. 4, species 31; Godwin-Austen 1914: 418–419, pl. 151, fig. 1.Alycaeus (Dicharax) glaber– Kobelt 1902: 371; Gude 1921: 253.Type locality.
Holotype (HNHM 99707) and a few paratypes, see the original description.
Remarks.
Protoconch low, glossy; R1 almost completely smooth, with only very inconspicuous, irregular growth lines; R2 extremely densely ribbed, with low, blunt ribs; for cross-sectional view of R2 see original description.
AnimaliaGastropodaCyclophoridae9C897CA8-52F2-54A0-AE55-22255518AAF3Dicharax(?)ingrami(W. T. Blanford, 1862)AlycaeusingramiW. T. Blanford, 1862: 135–136.Alycaeusingrami– Reeve 1878: pl. 6, species 54; Godwin-Austen 1886: 193–194, pl. 44, figs 1, 1a–c; Godwin-Austen 1914: 421.Alycaeus (Chamalycaeus) ingrami– Kobelt 1902: 357; Gude 1921: 228–229.Type locality.
Protoconch low, matte, without spiral striation; R1 with regular, strong, rather dense ribs, near the suture the ribs are sharp, elevated; R2 long, with rather dense, sharp ribs; at the anterior end of R2 the ribs are slightly bent in anterior direction.
AnimaliaGastropodaCyclophoridae2F504B6F-054F-5037-810D-21A4E6D6A887Dicharax(?)khasiacus(Godwin-Austen, 1871)AlycaeusKhasiacusGodwin-Austen, 1871: 90, pl. 3, fig. 4.Alycaeuskhasiacus– Reeve 1878: pl. 1, species 8; Godwin-Austen 1914: 356, 376–377, 393, pl. 143, figs 7, 7a, 7b.Alycaeus (Dicharax) khasiacus– Kobelt 1902: 372–373; Gude 1921: 257–258.Alycaeuskhasiacusvar. – Godwin-Austen 1914: 356.Chamalycaeus (Dicharax) khasiacus– Ramakrishna et al. 2010: 62.Type locality.
“On the highest parts of the Khasi and Jiantia Hills”.
Material examined.
Lailangkote, Khasi Hills, NHMUK 1903.7.1.2650 (29 syntypes).
Remarks.
Protoconch low, glossy, no spiral lines visible; R1 with similar sculpture to that of the protoconch; R2 short, with alternating thicker/darker and narrow/lighter stripes, resulting in a nearly smooth surface.
AnimaliaGastropodaCyclophoridaeAACAE32D-DC0F-5612-AE7A-0BD50D83C627Dicharax(?)kurzianus(Theobald & Stoliczka, 1872)Fig. 20AlycaeuskurzianusTheobald & Stoliczka, 1872: 330, pl. 11, fig. 2.Alycaeuskurzianus– Reeve 1878: pl. 3, species 22; Godwin-Austen 1914: 409–410, pl. 151, figs 7, 7a.Alycaeus (Dicharax) kurzianus– Kobelt 1902: 373; Gude 1921: 258–259.Type locality.
“Nattoung in provincia Barmana, Prome dicta”.
Material examined.
Nr. Prome, Pegu, leg. F. Stoliczka, NHMUK 1903.7.1.2700 (3 probable syntypes [labelled as “kurtzianus”; the locality do not match with the original description, but the specimens are identical to the figured one]); Nattoung, W. Prome, coll. Theobald, NZSI M.24974 (1 syntype), 25021 (6 syntypes).
Remarks.
Protoconch low, glossy, without spiral lines; R1 also glossy, without any notable sculpture; R2 moderately long, with lamella-like ribs, which are straight in the first half of the region, but are gradually more curved towards the aperture in the second half of R2.
Dicharax(?)kurzianus (Theobald & Stoliczka, 1872) Nattoung, W. Prome, coll. Theobald, syntype (NZSI M.24974). All images: Sheikh Sajan.
https://binary.pensoft.net/fig/468665AnimaliaGastropodaCyclophoridaeEA66F64F-21A0-5C67-961D-879605E5E556Dicharax(?)lectus(Godwin-Austen, 1914)Fig. 21AlycaeuslectusGodwin-Austen, 1914: 340, pl. 136, figs 5, 5a, b.Alycaeus (Dicharax) lectus– Gude 1921: 259.Chamalycaeus (Dicharax) lectus– Ramakrishna et al. 2010: 62; Tripathy et al. 2018: 789.Type locality.
“in valle Rungun”.
Material examined.
Near Chaukkalan, Darjeeling, coll. Dr. F. Stoliczka, NZSI M.8074 (holotype [single specimen mentioned in the original description]).
Remarks.
The holotype was strongly weathered, but no spiral striation could be found on the relatively intact protoconch and R1 surface; the protoconch was also low, therefore this species is classified in Dicharax. The R2 region was so weathered that its original structure could not be seen.
Dicharax(?)lenticulus (Godwin-Austen, 1874), holotype (NZSI M.8075). All images: S. Sajan.
https://binary.pensoft.net/fig/468667AnimaliaGastropodaCyclophoridae5ABFCD6F-1BAF-586B-A26D-A151E708DC3EDicharax(?)levis(Godwin-Austen, 1914)AlycaeuslevisGodwin-Austen 1914: 394, pl. 138, figs 3, 3a.Alycaeuslevis– Gude 1921: 209–210.Alycaeus (Alycaeus) levis– Ramakrishna et al. 2010: 48.Type locality.
“Gaziphimi, Lahupa Naga Hills, Munipur”.
Material examined.
Munipur, coll. Godwin-Austen, NHMUK 1903.7.1.2631 (holotype [single specimen mentioned in the original description]).
Remarks.
The entire shell was strongly weathered; the fine sculpture could not be fully examined. Protoconch low, no spiral lines visible; R1 without any recognisable sculpture; R2 moderately long, ribs dense and their fine structure could not be examined. The species is placed in the genus Dicharax on the basis of its low protoconch.
AnimaliaGastropodaCyclophoridae83FA6444-7421-559F-9623-96ABF3E44197Dicharax(?)logtakensis(Godwin-Austen, 1914)AlycaeuslogtakensisGodwin-Austen, 1914: 394–395, pl. 155, fig. 6.Alycaeuslogtakensis– Gude 1921: 209–210.Alycaeus (Alycaeus) logtakensis– Ramakrishna et al. 2010: 48.Type locality.
“Logtak Lake, Munipur (...) on a low hill near the northern shore”.
Material examined.
On low hill. Logtak Lake, Munipur, coll. Godwin-Austen, “figured”, NHMUK 1903.7.1.2639 (1 syntype).
Remarks.
Protoconch elevated, but no spiral lines were visible; R1 with widely spaced ribs without spiral striae; R2 relatively long with widely spaced, sharp ribs.
Naga Hills, fr. Beddome, NHMUK 1903.7.1.1480 (1 syntype).
Remarks.
Protoconch low, weathered, no spiral lines visible; R1 also without spiral lines; R2 very long (similar to Chamalycaeusheudei), some of the ribs have a horizontal projection in an anterior direction; near the suture these projections sometimes reach the neighbouring ribs; the entire shell is somewhat weathered, therefore it is not possible to decide if all ribs had these horizontal projections or only some of them.
AnimaliaGastropodaCyclophoridae4D04347F-2E33-507D-BA1E-B6C1C75366D2Dicharax(?)moellendorffi(Kobelt & Möllendorff, 1887)AlycaeusinflatusMöllendorff, 1886: 168, pl. 5, fig. 7.Alycaeus (Chamalycaeus) moellendorffiKobelt & Möllendorff, 1897: 149; Kobelt 1902: 359.Chamalycaeusmoellendorffi– Yen 1939: 30, pl. 2, fig. 35.Chamalycaeus (Chamalycaeus) moellendorffi– Zilch 1957: 144, pl. 5, fig. 12.Dicharax(?)moellendorffi – Páll-Gergely et al. 2017: 27–30, fig. 18.Type locality.
“Dau-dshou provinciae sinensis Hunan”.
Material examined.
China: Dau-dshou (Hunan), coll. Möllendorff 1886, coll. Boettger, SMF 39236 (lectotype, designated by Yen 1939); same data, SMF 39237 (2 paralectotypes); same data, SMF 39238 (8 paralectotypes).
Remarks.
Protoconch low, glossy, without spiral striation; R1 with dense, rather regular, low ribs and without spiral striation; R2 relatively long, with darker wider and lighter narrower stripes alternating, the overall surface of R2 is smooth.
Sikkim, at 11,000 ft, coll. Dr. Stoliczka, NZSI M.8082 (syntype, labelled as holotype).
Remarks.
The original description does not mention the number of available specimens. Thus, the shell labelled as holotype is considered to be a syntype.
Protoconch low, without spiral striation, R1 with low, rather regular ribbing; surface of R2 wavy, ribs are blunt and only slightly elevated from the surface.
“Hills at head of the Lanier River, Naga Hills, ca. 5–6,000 feet”.
Material examined.
Head of Lanier R.N.E. Munipur, coll. Godwin-Austen, NHMUK 1903.7.1.2485 (2 syntypes).
Remarks.
Protoconch low, rather glossy, no spiral lines visible; R1 smooth, without spiral striation, only some irregular, rough wrinkles visible near the suture; R2 moderately long, nearly smooth with alternating darker and lighter stripes.
E. Naga, coll. R.H. Beddome, NHMUK 1912.4.16.273 (16 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 with very widely spaced regular ribs and without spiral striation; R2 long, some ribs slightly curved towards the aperture, but mostly straight, lamella-like.
AnimaliaGastropodaCyclophoridae3A94D8CB-DD0E-511D-B0CB-AD59D566DA91Dicharax(?)mutatus(Godwin-Austen, 1876)AlycaeusmutatusGodwin-Austen, 1876: 177–178, pl. 7, figs 11, 11a.Alycaeus (Dicharax) mutatus– Kobelt 1902: 373–374; Gude 1921: 260–261.Alycaeusmutatus– Godwin-Austen 1914: 357, pl. 145, figs 9, 9a.Chamalycaeus (Dicharax) mutatus– Ramakrishna et al. 2010: 63; Tripathy et al. 2018: 789.Type locality.
“On Torúpútú, Tánir, and Shengorh Peaks, at 6–7000 feet elevation, in the dead leaves and moss about the roots of the forest”.
Material examined.
Toruputu Peak, Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2495 (9 syntypes).
Remarks.
Protoconch low, finely granulated, no spiral lines visible; R1 rather regularly, finely ribbed without spiral lines; R2 relatively short, with narrow, greyish, and somewhat more thickened yellowish corneous alternating stripes; entire R2 surface almost smooth, but narrow greyish stripes slightly elevated from the surface.
AnimaliaGastropodaCyclophoridae2C390697-188D-57CA-BD21-709D8EB99753Dicharax(?)nagaensis(Godwin-Austen, 1871)Alycaeusingramivar.nagaensisGodwin-Austen, 1871: 92, pl. 5, fig. 2.Alycaeusnagaensis– Godwin-Austen 1884: pl. 51, figs 3, 7; Godwin-Austen 1886: 195, pl. 44, figs 3, 3a–c; Godwin-Austen 1914: 396–397, pl. 143, figs 2, 2a, 2b.Alycaeus (Chamalycaeus) nagaensis– Kobelt 1902: 359; Gude 1921: 230.Chamalycaeus (Chamalycaeus) nagaensis– Ramakrishna et al. 2010: 54.Type locality.
“Neighbourhood of Asálú, rather local in its distribution, but abundant”.
Material examined.
Asalu, N. Cachar, NHMUK 1903.7.1.2615 (7 syntypes in 2 vials).
Remarks.
Protoconch rather low without any signs of spiral striae; R1 regularly ribbed without spiral striation; R2 long with widely spaced, sharp ribs (typical for Metalycaeus).
Nattoung Hills, Mendon District, Pegu, Burma, coll. Theobald, NZSI M.8036 (holotype, [single specimen mentioned in the original description]).
Remarks.
Protoconch low, without spiral striation; beginning of R1 nearly smooth, its end with rather widely spaced, wide, but low ribs; R2 long, with rather sharp, straight ribs.
Protoconch slightly elevated, but no spiral lines visible; R1 also without spiral lines; R2 moderately long with relatively dense but spaced, sharp ribs. The slightly elevated protoconch and the morphology of the ribs is similar to that of Chamalycaeus, but they are closer to each other than usually occurs in that genus.
Protoconch low, no spiral lines visible; R1 also without spiral striation; R2 short, with dense ribs; the fine morphology of the ribs could not be examined due to corrosion.
“Pachita village (Camp no. 7 of the Expeditionary Force, 1874–75), Dafla Hills, Assam”.
Material examined.
Pachita village, Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2614 (3 syntypes).
Remarks.
Protoconch low, without spiral striae; R1 also without spiral striation; R2 short, with very low ribs; this region is characterised by alternating thicker/dark, and narrower/white stripes.
AnimaliaGastropodaCyclophoridaeD3B436FA-2774-58BA-B830-5E0743865EE5Dicharax(?)panshiensis(Chen, 1989)ChamalycaeuspanshiensisChen, 1989: 157.“Chamalycaeus” panshiensis – Páll-Gergely et al. 2017: 107. Type locality.
“Hongn-an forestry centre, Panshi County (43°15'N, 126°10'E), Jilin Province, China”.
Material examined.
Type specimen presumably lost.
Remarks.
The type specimens could not be found in Beijing during a recent search, and they are thus considered lost. The original description is not sufficient for correct generic placement. A Dicharax species have been collected near the type locality of C.panshiensis (Guoyi Zhang pers. comm.) reminiscent of Korean and Japanese Dicharax species, which suggests that C.panshiensis belongs to the genus Dicharax. However, since no specimens are available and the original description is also useless, collection of topotypic specimens would be necessary.
Naga Hills, coll. Preston, NHMUK 1915.1.4.1281 (1 syntype).
Remarks.
The syntype is strongly weathered, the original sculpture could not be examined. Alycaeuspeilei is placed in the genus Dicharax on the basis of the seemingly low protoconch.
AnimaliaGastropodaCyclophoridaeF9F34B4C-702C-5ADD-8050-D79D8B738105Dicharax(?)plectocheilus(Benson, 1859)AlycaeusplectocheilusBenson, 1859: 180.Alycaeusplectocheilus– Reeve 1878: pl. 2, species 14; Godwin-Austen 1914: 342–343, pl. 134, figs 4, 4a–c.Alycaeus (Dicharax) plectochilus[sic] – Kobelt 1902: 375.Alycaeusplectocheilus, large var. – Godwin-Austen 1914: 342–343, pl. 133, figs 3, 3a–c.Alycaeus (Dicharax) plectochilus[sic] – Gude 1921: 264–265.Chamalycaeus (Dicharax) plectochilus– Ramakrishna et al. 2010: 65.Dicharaxplectocheilus– Páll-Gergely et al. 2017: 54.Type locality.
“in valle Rungun”.
Material examined.
Darjiling, Rungun Valley, coll. Blanford, NHMUK 1906.4.4.184 (2 syntypes); Vorder Indien, Rungun Valley, Darjiling, leg. Hungerford 1889, leg. O. Boettger, SMF 109254 (2 shells); Damsang, Daling District, NHMUK 1903.7.1.1256 (25 shells of “large var.”).
Remarks.
Protoconch low, rather glossy, no spiral lines visible; R1 glossy, very finely ribbed without spiral lines; R2 moderately long, it forms a nearly smooth area with alternating thicker/darker and narrower/lighter stripes.
AnimaliaGastropodaCyclophoridae12AF3161-0162-5C2A-9D46-E9AFD384243FDicharax(?)pusillus(Godwin-Austen, 1871)AlycaeuspusillusGodwin-Austen, 1871: 89–90, pl. 3, fig. 3.Alycaeuspusillus– Reeve 1878: pl. 1, species 7; Godwin-Austen 1914: 379–380, 398, pl. 143, figs 6, 6a, 6b; Gude 1921: 215–216.Alycaeus (Alycaeus) pusillus– Kobelt 1902: 348; Ramakrishna et al. 2010: 50.Type locality.
“near Jawai”, “on the banks of the Kopili river on the road from Jawai to Asálú, viâ Súfai”.
Material examined.
Jawai, Jaintia, NHMUK 1903.7.1.2688 (5 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 glossy, without notable sculpture; R2 short, with alternating thicker/darker and narrower/lighter stripes; the overall surface of R2 is seemingly smooth.
“Rechila Peak, Daling District, on Sikhim-Bhutan Frontier, 10,300 feet”.
Material examined.
Rechila Pk., Sikkim, leg. N. Robert, NHMUK 1903.7.1.1252 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch low, without spiral striae; R1 also without spiral striae and with low, dense, regular ribs; R2 with low, regular ribs; the entire R2 surface is wavy.
Mai-i, Sandoway Dist., Arakan, leg. Stoliczka, NHMUK 1903.7.1.2558 (holotype [single specimen mentioned in the original description]).
Remarks.
Protoconch moderately elevated, no signs of spiral striae; R1 regularly ribbed, without spiral striae; R2 very short, ribs lamella-like, straight, seemingly do not differ from the ribs of R1.
The specimen in lot 1903.7.1.2666 is weathered, so the original sculpture could not be fully examined, but a second type lot of A.sculpturus contained four shells in relatively good condition (NHMUK 1903.7.1.2667).
Protoconch moderately elevated, no spiral lines visible; R1 with widely spaced, strong ribs and without spiral lines; R2 of normal length, with widely spaced, sharp ribs.
Laisen Valley, Jiantia Hills, NHMUK 1903.7.1.2487 (1 syntype).
Remarks.
The syntype was strongly corroded. Protoconch low without spiral lines; R1 smooth with rough wrinkles near the suture; R2 moderately long, the fine structure of the ribs could not be examined because the ribs have been damaged.
“The earliest Miocene (Aquitanian, 23–21 Ma) Hang Mon Formation at Hang Mon in Northern Vietnam”.
Remarks.
This species fits the range of morphological variation of Dicharaxfimbriatus, which is an extant species from the same geographic area. However, we find it more useful to keep this taxon as a separate species due to its age. The strikingly similar appearance of an early Miocene species (Aquitanian, 23–21 Ma) to species living today indicates the high level of morphological stability of Dicharax.
AnimaliaGastropodaCyclophoridaeF6A26778-EEE1-51C5-BA78-8AEE3EE8617DDicharax(?)stoliczkii(Godwin-Austen, 1874)AlycaeusStoliczkiiGodwin-Austen, 1874: 147, pl. 3, fig. 3.Alycaeusstoliczkii– Reeve 1878: pl. 6, species 53; Godwin-Austen 1914: 399–400, pl. 144, figs 3, 3a, 3b.Alycaeus (Chamalycaeus) stoliczkai[sic] – Kobelt 1902: 363; Gude 1921: 233–234.Chamalycaeus (Chamalycaeus) stoliczkai[sic] – Ramakrishna et al. 2010: 55.Type locality.
“Angaoluo Peak, Nágá Hills at 7,000 feet”; “further to the east at Kezakenomih, and at the head of the Lanier River at ca. 5,000 feet where the specimens are much larger”.
Material examined.
Naga Hills, NHMUK 1903.7.1.2622 (3 syntypes).
Remarks.
Protoconch low, without spiral structure; R1 rather regularly ribbed without spiral structure, R2 long, with sharp, straight, low ribs. The sharp ribs are characteristic for the genus Dicharax, but in case of A.stoliczkii the ribs are much lower than in any Chamalycaeus species. Therefore, based on the low protoconch without spiral lines, we classify this species in the genus Dicharax.
AnimaliaGastropodaCyclophoridaeD234939D-4459-5FFB-B52C-21E5213B63FEDicharax(?)strangulatus(L. Pfeiffer, 1846)CyclostomastrangulatumL. Pfeiffer, 1846: 86.Alycaeusstrangulatus– Reeve 1878: pl. 6, species 47; Godwin-Austen 1914: 337, pl. 136, figs 1, 1a.Alycaeus (Dicharax) strangulatus– Kobelt 1902: 376; Gude 1921: 269.Chamalycaeus (Dicharax) strangulatus– Ramakrishna et al. 2010: 66.Dicharaxstrangulatus– Sajan et al. 2020: 523, Figs 1A, B, 2A–L.Type locality.
“Bengalia”.
Material examined.
“possible syntype” NHMUK 1856.9.15.18 (1 shell); Mussoorie, N.W Himalaya,”figd”, NHMUK 1903.7.1.2501 (at least 5 shells); NHMUK 1928.7.28.85–104 (from general collection). See also newly collected specimens examined in Sajan et al. (2020).
Remarks.
Protoconch low, without spiral striae; R1 with irregular, low ribs and without spiral striation; R2 short, with alternating wider/darker and narrower/lighter stripes; entire surface nearly smooth.
The syntypes should be in the Indian Museum (see Nevill 1878), but they have not been located there. This species is tentatively classified in the genus Dicharax based on the flat shell and low protoconch, and its resemblance to Dicharax(?)khasiacus (see the original description).
Naga Hills, leg. Doherty, NHMUK 1903.7.1.2687 (3 syntypes).
Remarks.
Protoconch low, no spiral lines visible; R1 glossy, without spiral striae; R2 short; ribs slightly bent, although the exact fine sculpture could not be determined due to corrosion of the shell.
Dicharaxsubhumilis (Möllendorff, 1897), lectotype (SMF 109224). Photographs: Barna Páll-Gergely, courtesy Ronald Janssen.
https://binary.pensoft.net/fig/468670AnimaliaGastropodaCyclophoridae110EB0D8-B041-5244-8D56-11509E37CF40Dicharax(?)succineus(W. T. Blanford, 1862)AlycaeussuccineusW. T. Blanford, 1862: 139–140.Alycaeussuccineus– Reeve 1878: pl. 2, species 16; Godwin-Austen 1914: 424, pl. 151, fig. 2.Alycaeus (Dicharax) succineus– Kobelt 1902: 377; Gude 1921: 271–272.Type locality.
“in montibus Arakanensibus”.
Material examined.
Tangoop Pass, Arakan Hills, NHMUK 1906.404.52 (1 syntype).
Remarks.
Protoconch rather low, without spiral striae; R1 regularly ribbed without spiral striation; R2 long with widely spaced, sharp ribs. Alycaeussuccineus is classified in Dicharax on the basis of the low protoconch and the absence of spiral striation, but the elevated, sharp R2 ribs are characteristic of the genera Metalycaeus and Chamalycaeus.
Protoconch low, no spiral lines visible; R1 with rather regular ribs and without spiral striation; R2 long, with widely spaced, ribs lamella-like, straight, but relatively low (typical Chamalycaeus character).
Fort Stedman, Burma, coll. Woodthorpe, NHMUK 1903.7.1.3064 (22 syntypes in two vials); Burma, E. R. Sykes Collection, Acc. no. 1825, NHMUK 20150126 (2 shells).
Remarks.
Protoconch glossy, low, no spiral lines visible; R1 with rather irregular, dense, low ribs without spiral striation; R2 with alternating thicker/darker and narrow/lighter stripes, the overall surface is smooth.
Japanese and Korean DicharaxAnimaliaGastropodaCyclophoridae1DE1573B-6048-5196-9808-89D63B1075EFDicharax(?)abei(Kuroda, 1951)Fig. 13BAwalycaeusabeiKuroda, 1951: 73–74, text figs 1–3.Awalycaeusabei– Kuroda and Abe 1980: 20; Minato 1982a: 121–123, fig. 4; Azuma 1982: 13, pl. 4, fig. 44; Hanshin Shell Club 1986: pl. 2, figs 7–9; Minato 1988: 17; Yano et al. 2016: 57, fig. 4C-a, b.Type locality.
“徳島縣(阿讃境に近く)城壬山々頂附近” (Tokushima, [near the Sakai boundary], near the top of Mt. Jiyauwau).
Holotype is deposited in the NC-H006 (Kazunori Hasegawa, pers. comm. 2015), and was not examined by us.
Protoconch low, rather matte, very finely granulated; R2 with 30–32 low, blunt, irregular ribs, which are usually in contact with each as they near the tube; at the edge of the body whorl they are distinct, the gap between them being slightly smaller than the width of a rib; no spiral lines visible on R1.
Protoconch low, very finely granulated, moderately matte; R2 with ca. 26 blunt, relatively low ribs; near the suture most ribs form connections (projections) with their neighbours or are completely merged; at the edge of the body whorl the space between the ribs is slightly less than the width of the ribs; no spiral lines visible on R1.
Protoconch low, very finely granulated, only slightly glossy; R2 with ca. 23 relatively low riblets occasionally fused to each other near the tube; space between the riblets at the edge of the body whorl is roughly as wide as the ribs; no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridae747D6B03-F703-558C-9235-BAB2E865E8C8Dicharax(?)ananensis(Yano, Tada & Matsuda, 2013)CipangocharaxananensisYano, Tada & Matsuda, 2013: 29–37, figs 1–4.Type locality.
“Wakasugidani, Suii-cho, Anan city, Tokushima Prefecture, Shikoku, Japan”.
Remarks.
No specimens were examined by us, but the original description provides enough information of the key characters: protoconch low, glossy; R1 with rather dense, strong ribs; R2 short, with very dense, low ribs; R3 longer than R2, without swelling.
Protoconch low, finely granulated, rather matte; R2 with ca. 26 low, blunt, irregular ribs, which are connected to each other near the tube, gradually becoming free towards the edge of the body whorl (here the distance between neighbouring ribs is very small, much less than the width of a rib); no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridae1DB7B20D-F763-5FC7-96D5-DD1B4CB87A76Dicharax(?)cyclophoroides(Pilsbry & Y. Hirase, 1909)AlycaeuscyclophoroidesPilsbry & Y. Hirase, 1909b: 9, pl. 5, figs 1, 2.Alycaeuscyclophoroides– Pilsbry 1926: 454.Chamalycaeus (Metalycaeus) cyclophoroides– Kuroda 1936: 170.Type locality.
Protoconch low, finely granulated, rather matte; R2 with ca. 28 low, blunt riblets, which form connections with each other near the tube; they are distinct at the edge of the body whorl, the space between them is slightly smaller than the width of a rib; no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridae452F5130-914B-5170-8A2F-305C565D7A4BDicharax(?)cyclophoroideskoshuensis(Kuroda, 1936)Chamalycaeus (Metalycaeus) cyclophoroideskoshuensisKuroda, 1936: 174.Chamalycaeuscyclophoroideskoshuensis– Hanshin Shell Club 1986: pl. 2, figs 3, 4.Type locality.
According to Kwon and Habe (1979)Chamalycaeuskoshuensis Kuroda, 1936 is a synonym (“local form”) of Alycaeuscyclophoroides. The holotype is deposited in the NC (NC-H203) (not examined by us).
Protoconch low or slightly elevated; R2 with ca. 26, relatively elevated and distinct ribs which are not in contact even at near the tube; at the edge of the body whorl the distance between the ribs is more than twice as wide as the width of a rib; there are no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridae6B395E6D-2EA1-5547-85EB-D54BA6E3064DDicharax(?)expanstoma(Minato, 1982)ChamalycaeusexpanstomaMinato, 1982b: 126–127, pl. 1, figs 1–4.Chamalycaeusexpanstoma– Minato 1988: 15, pl. 3, figs 1, 2.Type locality.
Protoconch low, very finely granulated, moderately glossy, no spiral lines visible; R2 with ca. 14–20 (14 Sakurai coll., 20 on the holotype) low, blunt ribs; ribbing resembles irregular wrinkled surface; near the tube some ribs reach the neighbouring ones, and partly fuse with them; at the edge of the body whorl the ribs are not in contact, the spaces between ribs almost reaches the width of a single rib; no spiral lines on R1.
Ura-Hikimi-Kyo, Hikimi-mach, Mino-gun, Shimane Pref., NSMT-Mo 78866 (5 shells). The holotype is deposited in NC-H004 (Kazunori Hasegawa, pers. comm.) (not examined by us).
Remarks.
Protoconch low, very finely granulated, moderately glossy; R2 with ca. 24 ribs; they are low and the entire ribbing is not clear; the ribs are not distinct entities, their boundaries are obscure since their edges merge; no spiral lines visible on R1.
Protoconch low, very finely granulated, not glossy; however, the protoconch of the holotype was quite weathered; ca. 16 ribs on R2 (individual ribs are wider than in the nominotypical subspecies); R2 ribbing similar as in the nominotypical subspecies; no spiral lines visible on R1.
ANSP 78777a (lectotype of A.harimensis, designated by Baker 1964, photographs examined); ANSP 78817a (lectotype of A.reinhardti, designated by Baker 1964, photographs examined); 紀伊周参見 (Kii Susami), Toru Inaba collection, NSMT-Mo 66474 (5 shells) (“japonicus harimensis”).
Remarks.
Protoconch low, glossy; R2 with 18–22 low, blunt ribs connected to each other near the tube, but free at the edge of the body whorl (here the distance between two ribs is ca. as wide as the width of a single rib); no spiral lines visible on the teleoconch.
AnimaliaGastropodaCyclophoridae358BBE74-2906-5097-96DD-CCEC427394B1Dicharax(?)japonicussadoensis(Pilsbry & Y. Hirase, 1903)Alycaeusharimensisvar.sadoensisPilsbry & Y. Hirase, 1903: 128–130.Chamalycaeusjaponicussadoensis– Minato 1988: 14.Type locality.
“Aikawa, Sado”.
Material examined.
ANSP 83895a (lectotype, designated by Baker 1964, photographs examined); 佐渡長江川左岸北斜面 標高200 m ナラ トチ クリ 雑木林 1 個体 1982-11 矢田政治 (Sado, Nagae-gawa, sa-gan, kita-shamen, hyoko 200 m, nara, tochi, kuri zoki-bayashi, 1 kotai, 1982-11, Yada Masaji = Sado, Nagae river, left bank, north slope, altitude 200 m, mixed forest of oak [nara], Japanese horse-chestnut [tochi], and Japanese chestnut [kuri], 1 specimen, Nov. 1982, Masaji Yada), NSMT-Mo 60464 (1 shell).
Remarks.
The shell we examined was weathered, only the shape of protoconch (low) could be seen and its sculpture was not visible; R2 with ca. 22 low, blunt, irregular ribs which are joined to their neighbours at least near the tube; no spiral lines visible on the shell.
Protoconch similar to that of biexcisus; R2 with 16, widely spaced, low, blunt ribs; spaces as wide as or slightly wider (at the edge of the body whorl) than individual ribs; in the case of the three shells in the Sakurai collection some of the ribs on R2 are joined to the neighbouring ones near the tube; no spiral lines visible on R1. Operculum relatively slim with low outer belt and without nipple (see also original description).
NSMT-Mo 64217 (holotype); 岐阜縣1揖斐郡池田町 (Gifu-ken, Ibi-gun, Ikeda-cho = Gifu Prefecture, Ibi County, Ikeda Town), coll. Sakurai, NSMT-Mo 79094 (2 shells).
Remarks.
The protoconch of the holotype probably has a growth disorder, and its sculpture could not be fully examined. It was low, very finely granulated, and the last 0.25 of the whorl was wrinkled. The specimens in the Sakurai collection have normally developed protoconchs, which were low and finely granulated, without spiral lines; R2 has ca. 30–32 low, blunt ribs; most ribs are completely merged with the neighbouring ribs nearer the tube; ribs free at edge of body whorl; space between ribs ca. 2 × as wide as an individual rib; no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridaeBAC4E2AA-72FB-5315-9EAE-E6CDD233AC04Dicharax(?)miyazakii(Takahashi & Habe, 1976)ChamalycaeusmiyazakiiTakahashi & Habe, 1976: 27–28, text fig. 1.Chamalycaeusmiyazakii– Minato 1988: 15, pl. 3, figs 7, 8.Type locality.
福岡縣1粕屋郡古賀町薬王寺 (Fukuoka-ken, Kasuya-gun, Koga-machi, Yakuoji = Fukuoka Prefecture, Kasuya County, Koga Town, Yakuoji [=temple], coll. Sakurai, NSMT-Mo 79095 (13 shells).
Remarks.
Protoconch low, very finely granulated, but rather glossy; R2 with ca. 20 low, blunt riblets; they are very low near the tube (it is difficult to decide whether they overlap or not); at the edge of the body whorl the space between the ribs is roughly as wide as a single rib; no spiral lines visible on R1; operculum relatively thick, at least as thick as that of C.kiuchii.
Protoconch low, very finely granulated, moderately glossy (the protoconch of the holotype was partly weathered); R2 with ca. 20 low ribs, clearly separated at edge of body whorl; generally ribs more distinct than in S.itonisitonis; here space between ribs is as wide as individual ribs; no visible space between the ribs closer to the tube, except for those situated close to the end of the tube; no spiral lines visible on R1.
Protoconch low, relatively glossy, finely granulated with some fine radial growth lines; R2 with ca. 26 low riblets, more distinct at edge of body whorl, but overall very obscure; no spiral lines visible on R1.
Protoconch low, moderately glossy, very finely granulated, with fine wrinkles on the last 0.25 of whorl; R2 with ca. 28 low, hardly separable, blunt ribs; close to the tube boundary between ribs hardly visible, they possibly overlap; at the edge of the body whorl there is some distance between ribs, smaller than width of an individual rib; no spiral lines visible on R1.
Protoconch low, relatively glossy, extremely finely granulated; R2 with ca. 24 low, blunt, irregular ribs, which usually overlap with the neighbouring rings nearer the tube; distance between ribs at edge of body whorl ca. half rib width; no spiral lines visible on R1.
This species has a lip that is strongly extended in the direction of the umbilicus. Also, the thickened operculum is very peculiar with a compressed hourglass shape.
This species was originally described as a fossil species. However, a living individual was collected on Mount Nekumachiji (26°41'01.18"N, 128°08'17.85"E), Okinawa Island (Hiroshi Fukuda, pers. comm. 2016).
We have not found the holotype in the NSMT-Mo (inventory number 57766 according to the original description). We place the species in this genus because the original description does not mention spiral lines.
AnimaliaGastropodaCyclophoridaeB3C1D4BD-F4DA-5F08-AE85-5D2D5A1E15F9Dicharax(?)oshimanus(Pilsbry & Y. Hirase, 1904)AlycaeusoshimanusPilsbry & Y. Hirase, 1904b: 7–8.Chamalycaeusoshimanus– Azuma 1982: 9, pl. 3, fig. 26; Minato 1988: 13.Type locality.
Protoconch low, finely granulated, moderately glossy; R2 with ca. 32 low, blunt ribs; in one of the specimens the ribs are free from each other even near the tube; distance between ribs at edge of body whorl larger than a rib width; in the other, weathered specimens (Sakurai coll.) the regular ribs stand close to each other, similarly to that of C.laevis; no spiral lines on the teleoconch.
Protoconch low, smooth, but the last 0.5 whorl of the protoconch was slightly weathered; ca. 24 ribs on R2 , well separated and wide near beginning of tube, gradually becoming narrower, and dense towards end of tube; near tube ribs situated very close to each other, some of them overlap with their neighbours; ribs clearly separated at edge of body whorl, but spaces still much narrower than a rib width; no spiral lines visible on R1.
Operculum with a big central nipple on its inner side; however, the outer belt was absent. The columellar margin of the aperture is not sinuated. The aperture forms a connection between the Cipangocharax-type and normal-type aperture.
AnimaliaGastropodaCyclophoridaeD3169C07-8B0E-5436-8D79-0EBCA7E0A9EFDicharax(?)purus(Pilsbry & Y. Hirase, 1904)AlycaeuspurusPilsbry & Y. Hirase, 1904c: 617.Chamalycaeuspurus– Minato 1988: 12.Type locality.
“Tokunoshima, Ōsumi”.
Material examined.
ANSP 87683 (1 syntype, photographs examined).
Remarks.
Protoconch low, moderately glossy; R1 with low riblets, R2 with denser, blunt ribs, which are connected to each other near the tube.
Protoconch low, very finely granulated, moderately glossy, no wrinkles present; R2 with ca. 22 low, blunt, but well separated ribs; near tube some ribs merge with neighbouring ribs, but distance between ribs at edge of body whorl ca. as wide as a single rib (or slightly wider); no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridaeE1A51944-62A1-54C9-8183-22C520CFD4F4Dicharax(?)shiosakimasahiroi(Yano, Matsuda & Nishi, 2016)AwalycaeusshiosakimasahiroiYano, Matsuda & Nishi in Yano et al. 2016: 55–57, figs 2A–G, 3C, D, 4B-a–B-d.Type locality.
No specimens were examined by us, but the photographs of the original description provide sufficient details of the fine sculpture. Protoconch low, rather glossy; R1 with widely spaced, elevated, sharp ribs; R2 with much denser, blunter ribs.
AnimaliaGastropodaCyclophoridae58E0D59A-9287-5A08-A1BD-EDD87A6F5353Dicharax(?)spiracellum(A. Adams & Reeve, 1850)CyclostomaspiracellumA. Adams & Reeve, 1850: 56, pl. 14, fig. 1.Alycaeusspiracellum– E. von Martens 1867: 150–151; Reeve 1878: pl. 4, species 33; Godwin-Austen 1889: 346, pl. 37, fig. 6.Alycaeus (Dicharax) spiracellum– Kobelt 1902: 376.AlycaeuskurodaiPilsbry & Y. Hirase 1908: 60, pl. 4, figs 1–4.Alycaeuskurodae(sic) – Pilsbry 1926: 454.Chamalycaeus (Metalycaeus) kurodai– Kwon and Habe 1979: 26.DicharaxspiracellumPáll-Gergely 2019: 192, fig 1A–D. (kurodai Pilsbry & Y. Hirase, 1908 is a synonym)Type locality.
“Borneo, under decayed vegetable matter in the forests” (spiracellum), “Cheju (Quelpart) Island” (kurodai). Páll-Gergely (2019) emended the type locality of C.spiracellum to “probably Cheju Island, South Korea”. See Remarks.
Material examined.
Borneo, ‘Samarang pl. 14. F. 1’ (in pencil), coll. Mrs Lombe Taylor, NHMUK 1874.12.11.233 (possible syntype); Borneo, NHMUK 1889.12.7.27 (1 shell); 済州島 (Seishūtō = Jeju Island), coll. Hirase (#275) NSMT-Mo 7593 (4 shells, labelled as A.kurodai); ANSP 95742 (1 syntype of A.kurodai) (photographs examined).
Remarks.
protoconch low, rather matte, with irregular fine wrinkles; R2 with ca. 24 low ribs; boundaries between ribs hardly visible; ribs situated very close to each other near tube; distance between ribs at edge of body whorl roughly a rib width; no spiral lines visible on R1.
Dicharaxspiracellum (A. Adams & Reeve, 1850) was described from Borneo, based on shells from the expedition of HMS Samarang (1843–1846). Alycaeuskurodai Pilsbry & Y. Hirase, 1908, described from the Korean Cheju Island, is conchologically identical to Dicharaxspiracellum and is a junior synonym of the former species (Páll-Gergely 2019). The type specimens of Dicharaxspiracellum were probably obtained when the Samarang visited Cheju Island, and the specimens were most likely mislabelled (Páll-Gergely 2019). Therefore, the type locality of D.spiracellum was emended to “probably Cheju Island, South Korea” (Páll-Gergely 2019).
AnimaliaGastropodaCyclophoridaeA97FFB05-4517-5452-A3D4-8C682F92CAD6Dicharax(?)spiracellumduplicatus(Kuroda & Miyanaga, 1943)ChamalycaeuskurodaiduplicatusKuroda & Miyanaga, 1943: 130, 136, text fig. 4.Chamalycaeus (Metalycaeus) kurodai– Kwon and Habe 1979: 26 (treats duplicatus as a synonym = “local form”).Dicharaxspiracellumduplicatus– Páll-Gergely 2019: 192.Type locality.
“Reisui, Zenra-Nandō, southern coast of Tyōsen”.
Remarks.
No specimens of this taxon were examined by us. The holotype is deposited in the Nishinomiya Museum (NC-H204; Kazunori Hasegawa, pers. comm. 2015). The drawing in the original description shows a shell with two parallel R3 swellings, which justifies the distinction of that form at least at subspecies level.
AnimaliaGastropodaCyclophoridae3192F2BF-C4F3-579C-B002-DD6DED91C3DBDicharax(?)tadai(Kuroda & Kawamoto, 1956)ChamalycaeustadaiKuroda & Kawamoto in Kawamoto & Tanabe, 1956: 13, 86, figs 11, 12.Chamalycaeustadai– Azuma 1982: 11–12, pl. 4, fig. 39; Hanshin Shell Club 1986: pl. 2, figs 1, 2; Minato 1988: 15.Type locality.
“萩市見島, 日崎” (Hisaki, Mishima, Hagishi [Hagi City in Yamaguchi Prefecture]).
Protoconch low, very finely granulated; moderately glossy; R2 with 18–24 irregular, low, blunt ribs, connected to each other near tube and sometimes further away from tube (near edge of body whorl) also; distance between ribs at edge of body whorl usually ca. a rib width; no spiral lines visible on R1.
The authors (Kuroda and Kawamoto 1956) wrote the following: “Having lamellar plate in the form of spiral plates on the outer surface is a feature of the Metalycaeus group” (translated by J.U. Otani).
Protoconch low, finely granulated, moderately glossy; R2 with 32–36 low, blunt, well separated ribs; nearer the tube there is some distance between most of the ribs, but some of them join with their neighbours; distance between ribs at edge of body whorl 1–2 × as large as a rib width; no spiral lines visible on R1.
“Limestone outcrops of the Koumori-do Cave, Honjo-Kazato, Saeki City, Oita Prefecture”.
Material examined.
NSMT-Mo 77464 (holotype).
Remarks.
Protoconch low, very finely granulated, moderately glossy; with fine wrinkles on the last 0.25 of whorl; R2 with ca. 42 low, blunt ribs, which are situated very close to each other near the tube; near the middle of the tube the ribs reach each other and form connections with their neighbours; space between ribs at edge of body whorl roughly twice as large as a rib width; no spiral lines visible on R1.
Protoconch low, finely granulated, rather matte; R2 with ca. 20 low, blunt, regular ribs which are some distance from each other even near the tube; distance between ribs at edge of body whorl is larger than a rib width; no spiral lines visible on the teleoconch.
Minato (1988) treated Chamalycaeustanegashimae as a subspecies of Chamalycaeussatsumanus. According to our classification, Chamalycaeussatsumanus and Sigmacharaxtanegashimae belong to different genera (Dicharax and Metalycaeus, respectively).
AnimaliaGastropodaCyclophoridae22EE3E22-32C8-5484-A789-058A5C323194Dicharax(?)tokunoshimanus(Pilsbry & Y. Hirase, 1904)AlycaeustokunoshimanusPilsbry & Y. Hirase, 1904c: 617–618.Chamalycaeustokunoshimanus– Azuma 1982: 9, pl. 3, fig. 27.Chamalycaeustokunoshimanustokunoshimanus– Minato 1988: 13.Type locality.
“Tokunoshima, Ōsumi”.
Material examined.
ANSP 87505 (lectotype, designated by Baker 1964, photographs examined).
Remarks.
Protoconch low, finely granulated; R1 with low, irregular wrinkles; R2 with very low, regularly spaced ribs, which are in contact with each other near the tube.
AnimaliaGastropodaCyclophoridaeFCD81336-F51D-57BA-84CD-7A9BBBA7736CDicharax(?)tokunoshimanusprincipalis(Pilsbry & Y. Hirase, 1909)AlycaeustokunoshimanusprincipalisPilsbry & Y. Hirase, 1909a: 587.Chamalycaeustokunoshimanus– Azuma 1982: 9–10, pl. 3, fig. 28.Chamalycaeustokunoshimanusprincipalis– Minato 1988: 13.Type locality.
Protoconch low and finely granulated, the last 0.25 of the protoconch whorl is finely wrinkled; R2 with ca. 34–40 low, blunt ribs; they are situated close to each other near tube, but probably do not form connections with each other; distance between ribs at the edge of body whorl ca. as wide as a rib width; R1 with extremely fine spiral lines between ribs (probably not homologous with the spiral striations in Metalycaeus species).
AnimaliaGastropodaCyclophoridaeB2B16438-A7CF-5DBB-92E1-32FFAA0FEE4ADicharax(?)tokunoshimanusmediocris(Pilsbry & Y. Hirase, 1909)AlycaeustokunoshimanusmediocrisPilsbry & Y. Hirase, 1909a: 587.Type locality.
Protoconch relatively low, very finely granulated, moderately glossy; R2 with ca. 22 low, blunt, irregular ribs which are connected to each other near the tube but are free further away from there; distance between ribs at edge of body whorl less than the a rib width; no spiral lines are visible on the teleoconch.
Protoconch low, very finely granulated, moderately glossy, but the protoconch of the holotype was slightly weathered; however, those in the Sakurai collection were not; R2 with ca. 25 low, blunt, but well separated ribs; even near the tube the ribs seem to be free from each other; distance between ribs at edge of body whorl ca. as wide as a rib, or slightly wider; no spiral lines visible on R1.
AnimaliaGastropodaCyclophoridae1761A3EA-F62C-5B07-AB4E-C14BC73E2D6ADicharax(?)yanoshokoae(Yano & Matsuda, 2016)Awalycaeussp. – Kawase et al. 2012: 85–88, fig. 5a.AwalycaeusyanoshokoaeYano & Matsuda in Yano et al. 2016: 53–55, figs 1A–I, 3A, B, 4A-a–d.Type locality.
“extended region of limestone area of Saruda-do Cave, Okina, Hidaka-mura, Takaoka-gun, Kochi Prefecture”.
Remarks.
No specimens were examined by us, but the photographs of the original description provide sufficient details of the fine sculpture. Protoconch low, rather glossy; R1 with relatively dense, strong ribs; R2 with low, very dense ribbing.
Shell small to very large (3.5–9 mm), globose, sometimes the body whorl is angled or keeled; protoconch smooth, not spirally striated; R1 usually very finely reticulated due to fine radial ribs and fine spiral striation, or smooth; R2 short to long, usually without ribs (superficially smooth, with alternating lighter and darker stripes); R3 practically absent (the inner opening of the sutural tube is situated close to the aperture). Operculum thin, usually proteinaceous (“horny”), in some species can be thicker, calcified. Central tooth with 5–7 cusps, broad, central cusp blunt.
Differential diagnosis.
The combination of the globular shell, the reduced sculpture (only very fine spiral and radial lines are present), and the absence of the R2 region characteristic for the vast majority of alycaeid species make Dioryx species distinguishable from other alycaeid genera.
Distribution.
Dioryx is distributed from the southeastern Himalayan region to Taiwan in the east, and down to the northern part of the Malay Peninsula and southern Vietnam in the south (Fig. 7).
Remarks.
Dioryx is primarily classified based on the reduced R3. This character state is found in other genera of the Alycaeidae as well, such as in Alycaeusconformis, Chamalycaeusmicroconus, and in Awalycaeus (treated as a synonym of Dicharax). Even presuming that similar morphology has appeared multiple times in the Alycaeidae, we can reasonably assume the monophyly of Dioryx due to the uniform, simple sculpture across all species, and the generally globose shell.
AnimaliaGastropodaCyclophoridae5BAC7BEB-F533-5022-9640-AA7077F092F2Dioryxamphora(Benson, 1856)Fig. 26AlycaeusAmphoraBenson, 1856: 226.Alycaeusamphora– Reeve 1878: pl. 2, species 15.Dioryxamphora– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 336–337; Gude 1921: 198–199, fig. 31.Alycaeus (Dioryx) amphora– Godwin-Austen 1914: 429–430, pl. 153, figs 11, 11a, 11b.Type locality.
“ad Moulmein, et in valle Tenasserim”.
Material examined.
UMZC I.102850 (1 syntype); NHMUK 1906.5.5.23 (figured by Godwin-Austen; this sample contains 2 Dioryxamphora shells, and a third shell of another Dioryx species).
Remarks.
Protoconch matte, without spiral striae; R2 with regular spiral striation, and weaker, irregular radial lines; the overall sculpture is weak; R2 very long, reaches 150°, its surface smooth, with alternating narrow light and slightly thicker dark stripes.
The examined shell was weathered. Protoconch without spiral striae; R1 with very fine spiral striation and weaker, irregular growth lines; R2 short, smooth, alternating narrow/lighter and thicker/darker stripes.
AnimaliaGastropodaCyclophoridae9AC91EAE-4AC4-5552-836E-66255EBC87C8DioryxcarinigerMöllendorff, 1897DioryxcarinigerMöllendorff, 1897a: 41.Dioryxcariniger– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 337; Zilch 1957: 141, pl. 5, fig. 1; Páll-Gergely et al. 2017: 10; Inkhavilay et al. 2019: 15, fig. 5E, F.Type locality.
“Prope oppidum Luang-Prabangin regione Laos”.
Material examined.
Hinter Indien: Lakon (Laos-Gebiet), coll. Möllendorff, SMF 109263 (lectotype, designated by Zilch 1957); Same data, SMF 109264 (paralectotype).
Remarks.
Protoconch very finely tuberculated; R1 without radial lines, only very weak growth ridges may be visible, but the region bears visible, closely spaced spiral striae; R2 relatively long, with alternating darker and lighter stipes, lighter ones only slightly narrower than dark ones.
AnimaliaGastropodaCyclophoridaeA60241DD-878B-54DF-BBE7-1C4301833693Dioryxcochinensis(Godwin-Austen, 1914)Alycaeus (Dioryx) cochinensisGodwin-Austen, 1914: 428–429, pl. 156, figs 7, 7a.Dioryxcochinensis– Páll-Gergely et al. 2017: 10.Type locality.
“Cochin China”.
Material examined.
Cochin China, NHMUK 20170011 (holotype [single specimen mentioned in the original description], measurements of the specimen: D = 6.1, H = 7.0). See also Remarks.
Remarks.
Protoconch without spiral striae; R1 with very fine spiral striation and weaker, irregular, wavy growth lines; R2 very short, smooth, narrow light and thicker dark stripes alternating.
According to the original description of A.cochinensis, only a single shell was found “stuck on the same slab with Alycaeus gibbus” and originated from the collection of Hugh Cuming. We were unable to locate the holotype in the type collection of the NHM, but found a shell in the general collection labelled “Dioryx cochinensis G.-A. ms., Cochin China (Cuming)”, with a black glue mark on its body whorl indicating that it was previously mounted as indicated in the original description. This specimen is identical with the figures of Godwin-Austen (1914) (characteristic shell shape from apertural view: pl. 156, fig. 7; and the very short tube: pl. 156, fig. 7a). Therefore, we identify this specimen as the holotype of Dioryxcochinensis. The measurements provided by Godwin-Austen (1914) are the following: “major diam: 5.5; alt. axis 4.75 mm”. Our measurements of the specimen are, however, somewhat larger, namely D: 6.1 mm, H: 7 mm. This is probably due to the inaccuracy of the original measurements (as usual in Godwin-Austen’s specimens), and we do not question the status of the specimen.
Five shells in the Senckenberg Museum (Tonkin, Bac-Kan, Cho-Ra, leg. Messager, coll. Rolle ex coll. Bosch, SMF 192287) were labelled as syntypes, although they clearly belong to a different species than the single syntype in MNHN Paris, which agrees with the figures in Bavay and Dautzenberg (1900b). This single shell is identical to Dioryxdistortus, but smaller. A comprehensive revision would probably reveal that the two names are synonyms.
The following remarks are based on the syntype from Paris: protoconch matte; R1 with irregular, very weak, widely spaced ribs and dense spiral striae of the same strength, although due to small space between spiral striae, they are much more conspicuous than the rare radial lines; R2 long, with very thin lighter and darker thicker stripes, the overall surface is smooth, glossy.
AnimaliaGastropodaCyclophoridae72F6B891-D3DD-503B-86D7-C528A8CEB50FDioryxdautzenbergiPáll-Gergely, 2017Alycaeus (Dioryx) majorBavay & Dautzenberg, 1900a: 118–119.Alycaeus (Dioryx) major– Bavay and Dautzenberg 1900b: 452, pl. 11, figs 4–6.Dioryxmajor– Kobelt 1902: 338–339.Alycaeusmajor– Rees 1964: 63, pl. 5, fig. 26.DioryxdautzenbergiPáll-Gergely in Páll-Gergely et al., 2017: 10, fig. 4A. (replacement name for Alycaeus (Dioryx) major Bavay & Dautzenberg, 1900, non Alycaeus (Dioryx) granumvar.major Godwin-Austen, 1893)Type locality.
Protoconch glossy; R1 with very weak growth lines and no spiral striae; R2 relatively short, smooth, very narrow lighter and thicker, darker stripes alternating.
AnimaliaGastropodaCyclophoridae1D39AED0-21A5-57C4-AF0F-BDD4CEAD189FDioryxdistortus(Haines, 1855)CyclostomadistortumHaines, 1855: 158, pl. 5, figs 5–8.Alycaeusdistortus– Reeve 1878: pl. 3, species 24.Dioryxdistortus– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 337.Type locality.
“Siam”.
Material examined.
Siam: coll. Haines, AMNH 62467 (1 syntype).
Remarks.
Protoconch matte, without spiral striation; the overall sculpture very weak; first whorl of R1 with spiral striation and widely spaced, irregular radial lines; later spiral lines disappear and radial lines become regular and dense; R2 very long, reaches ca. 150°, it is entirely smooth, only slimmer light and thicker darker stripes alternate.
The holotype was weathered, but the protoconch seemed to be without sculpture. R2 very long, smooth.
AnimaliaGastropodaCyclophoridae97C6A266-CE70-57B7-80F7-BAC8D06114F8Dioryxfeddenianus(Theobald, 1870)AlycaeusFeddenianusTheobald, 1870: 397, pl. 18, fig. 4.Alycaeusfeddenianus– Reeve 1878: pl. 2, species 18.Dioryxfeddenianus– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 338; Gude 1921: 199, fig. 32.Alycaeus (Dioryx) feddenianusGodwin-Austen 1914: 415, pl. 153, fig. 12.Type locality.
“Shan States”.
Material examined.
Upper Salwin, NHMUK 1888.12.4.927–930 (4 shells).
Remarks.
Protoconch matte, without spiral striation; R1 with widely spaced spiral striae, and weaker, much denser radial lines; R2 exceeds 90°, it is smooth, with slimmer lighter and thicker darker stripes alternating.
AnimaliaGastropodaCyclophoridaeF0C2A791-CE29-5752-B0F7-8E6375CD400FDioryxglobuloidesZilch, 1957AlycaeusglobulusMöllendorff, 1885: 162.Dioryxglobulus– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 338; Yen 1939: 28, pl. 2, fig. 28.DioryxglobuloidesZilch, 1957: 141, pl. 5, fig. 2 (replacement name for globulus Möllendorff, 1885, non Godwin-Austen, 1874).Dioryxglobuloides– Páll-Gergely et al. 2017: 10, fig. 4D.Type locality.
“in regione Badung provinciae sinensis Hubei”.
Material examined.
Patung, Hupei: China, coll. Möllendorff, SMF 39221 (lectotype, designated by Zilch 1957); Same data, SMF 39222 (10 paralectotypes).
Remarks.
Protoconch rather matte, extremely finely granulated; R1 with irregular, fine, low wrinkles; R2 short, with darker, thick and lighter, narrow stripes alternating.
AnimaliaGastropodaCyclophoridae58EE2BD6-AAC6-5793-BB4E-E5219E870B54Dioryxglobulosus(Godwin-Austen, 1914)Alycaeus (Dioryx) globulosusGodwin-Austen, 1914: 368–369, pl. 157, figs 1, 1a.Dioryxglobulosus– Gude 1921: 200; Ramakrishna et al. 2010: 74; Tripathy et al. 2018: 789.Type locality.
“Luyor, Tsanspu Valley”.
Material examined.
Luyor, Abor, 7,200 f., leg. Oakes, NHMUK 1903.7.1.3528 (7 syntypes in two different vials).
Remarks.
Protoconch glossy; R1 with irregular, weak ribs and very weak spiral striae; R2 relatively short, with very narrow lighter and darker thicker stripes; overall surface smooth.
AnimaliaGastropodaCyclophoridaeDD4F5E74-F287-52EA-BD05-7FA19F690F72Dioryxkobeltianus(Möllendorff, 1875)Alycaeus Kobeltianus Möllendorff, 1875: 121–122. Alycaeus (Dioryx) kobeltianus– Schmacker and Boettger 1890: 121, pl. 2, fig. 3.Dioryxkobeltianus– Kobelt and Möllendorff 1897: 146; Kobelt 1902: 338; Yen 1939: 29, pl. 2, fig. 30; Zilch 1957: 141, pl. 5, fig. 3; Varga 1972: 136, figs 17, 18; Páll-Gergely et al. 2017: 10, fig. 4E.Type locality.
“Berge bei Kiukiang”.
Material examined.
Kiukiang, China, coll. Möllendorff 1874, SMF 39298 (syntype, labelled as holotype [the number of available shells was not stated in the original description]).
Remarks.
The original description does not mention the number of examined specimens. Thus, we consider the specimen labelled as holotype (SMF 39298) syntype.
Protoconch very finely granulated; R1 with fine, rather irregular growth lines and very fine spiral lines; R1 relatively short, smooth, with alternating dark, wider and light, slimmer stripes.
Khargan, Attaran valley, coll. W. Theobald, NZSI M.8056 (syntype, labelled as holotype [the original description is based on multiple specimens]).
Remarks.
Protoconch smooth, without spiral striation, R1 with irregular, fine wrinkles; R2 ca. 0.25 whorl long, with irregularly arranged (probably teratological condition?) lighter, narrow, and thicker darker stripes.
“Menglun, Mengla County (21°09'N, 101°02'E), Yunnan Province, China”.
Material examined.
Meng-Lun town, Meng-La County, Xi-Shuang-Ban-Na Dai Autonomous Prefecture, China, leg. Chen De-Niu, 1994.5.3, TM 046653 (holotype, deposited in IZCAS); TM 086956 (paratype) same as holotype.
Remarks.
Protoconch finely granulate, matte; R1 consisting with fine, irregular growth lines, and dense, fine, regular spiral striation; R2 long, reaches 180°, glossy, with lighter, slimmer, and darker, thicker stripes interchanging, and with fine, obscure spiral sculpture; constriction deep and relatively long.
Protoconch rather glossy, R1 with widely spaced, low, irregular ribs and dense, rather prominent spiral striae; R2 of ca. 90°, with alternating thinner/lighter and thicker/darker stripes.
protoconch finely granulate; R1 with irregular, fine growth lines (no spiral striation visible); R2 smooth, with alternating white/narrow and darker/thicker stripes.
AnimaliaGastropodaCyclophoridae7920030B-00AF-5A93-A7CD-55A4BA02E7D2Dioryxpilula(Gould, 1859)AlycaeuspilulaGould, 1859: 424–425.Dioryxpilulaand pilulavar.minor – Kobelt and Möllendorff 1897: 147.Alycaeus (Dioryx) pilula– Bavay and Dautzenberg 1900b: 456–457.Dioryxpilula– Kobelt 1902: 339; Yen 1939: 28, pl. 2, fig. 29; Zilch 1957: 141; Varga 1972: 136, figs 19, 20; Páll-Gergely et al. 2017: 10.Alcaeus[sic] pilulaJohnson 1964: 127.Type locality.
“Hong Kong, China”.
Material examined.
Hong Kong, ex coll. Möllendorff, ex coll. Mus. Heude, MCZ 167223 (1 shell).
Remarks.
Protoconch extremely finely granulated, rather matte; R1 with irregular growth ridges and very fine spiral striation; R2 relatively short, with extremely fine alternating wider/darker and slimmer/lighter stripes.
Johnson (1964) did not find the type lot in the Smithsonian Museum. The sample we examined was collected at the type locality.
Protoconch matte, without notable sculpture; R1 smooth, with very low, irregular growth lines; R2 ca. 0.25 of whorl long, smooth, with alternating thinner/lighter and thicker/darker stripes.
Shell shape typical Dioryx; apex relatively elevated, matte; R1 with irregular, widely spaced, low ribs and very slightly stronger, dense spiral striation, which is the strongest on the first whorl of R1; R2 long, with dense alternating lighter/narrow and darker/thicker stripes; overall surface smooth.
Protoconch rather matte, R1 consists with very weak, dense, inconspicuous growth lines, extremely weak spiral striation; R2 smooth, with alternating narrow/lighter and thicker/darker stripes.
This species differs from D.messageri in stable characters and they are found in the RBINS in multiple mixed samples. Therefore, we handle D.rosea as a separate species. A redescription will be presented in a separate paper.
“Ruyang Town, Ruyuan County (24°7'N, 113°2"E), Guangdong Province, China”.
Remarks.
The holotype (IZCAS TM 132794) and a paratype (IZCAS TM 132795–132805) were photographed by one of us (Meng Kaibaryer). The shell height was 4.7 and 4.6 mm respectively, whereas it is given as 7.1 mm in the original description. Moreover, the examined specimens were much more corpulent than the one imaged in Hu et al. (2004), indicating that they belong to another species. Consequently, the types of this species are considered lost.
Shells of Dioryx Benson, 1859 species ADioryxruyangensis Hu, Yin, & Chen, 2004, holotype (IZCAS TM 132794) B label of the holotype C figures from the original description of Dioryxruyangensis Hu, Yin, & Chen, 2004. Photographs: Kaibaryer Meng (A, B).
https://binary.pensoft.net/fig/468674AnimaliaGastropodaCyclophoridae4D86D6CC-8224-5BFD-A154-F402DBEC1D99Dioryxsetchuanensis(Heude, 1885)Alycoeus[sic] setchuanensis Heude, 1885: 97, pl. 24, figs 6, 6a–c.Dioryxsetchuanensis– Kobelt and Möllendorff 1897: 147; Kobelt 1902: 340; Páll-Gergely et al. 2017: 10.Type locality.
Protoconch matte; R1 with inconspicuous, irregular growth lines and very fine, dense spiral striation (strength of spiral striation is variable); R2 smooth, with alternating lighter/slimmer and darker, slightly thicker stripes.
According to Johnson (1973) syntypes are also present in the USNM (inventory number: 472337).
AnimaliaGastropodaCyclophoridaeFF2351DC-EF98-5864-8740-FBF96D0D8D48Dioryxswinhoei(H. Adams, 1866)Alycaeus (Dioryx) swinhoeiH. Adams, 1866: 318, pl. 33, figs 11, 11a.Alycaeusswinhoei– Reeve 1878: pl. 3, species 21.Dioryxswinhoei– Kobelt and Möllendorff 1897: 147; Kobelt 1902: 340; Hsieh et al. 2006: 88 + figures; Hwang 2014: 8, fig. 1F.Type locality.
“Takow, Formosa”.
Material examined.
Formosa, NHMUK 1866.5.9.8 (lectotype, designated by Hwang 2014); Same data, NHMUK 1866.5.9.9 (1 paralectotype); Formosa: Sammaipo, leg. Hirase, 1906, 1378a (probably locality code of Hirase), coll. Jetschin ex coll. K.L. Pfeiffer, SMF 109780 (2 shells).
Remarks.
Both the lectotype and the paralectotype were strongly weathered. Protoconch without notable sculpture; R1 with some radial sculpture; R2 short, with alternating narrow/lighter and thicker/darker stripes, overall surface smooth. The specimens deposited in the Senckenberg Museum were not weathered, and the sculpture could be examined in more detail: protoconch rather glossy, R1 with very fine, irregular radial lines and weak spiral striation.
“Tongmai Town, (30°01'N, 95°E), Bomi County, Tibet Autonomous Region, China”.
Material examined.
Tong-Mai Town, Bo-Mi County, Tibet Autonomous Region, China, leg. Chen De-Niu & Gao Jia-Xiang, 1980.6.20, CASIZ TM 008902 (holotype, deposited in IZCAS); same data as holotype, leg. Gao Jia-Xiang, CASIZ TM 008903–008913 (paratypes).
Remarks.
Protoconch without notable sculpture; R1 with fine, irregular growth lines (some of them are almost represented as widely-spaced ribs) and very fine spiral striation; R2 ca. 0.25 whorl long, with alternating darker/wider and very lighter/narrow stripes.
Protoconch very finely granulated, R1 with hardly visible, oblique ribs, R2 normal length, smooth with alternating darker/wider and lighter/narrower stripes.
Japvo Peak, Naga, 10,000 f., leg. Godwin-Austen, NHMUK 1903.7.1.2530 (4 syntypes, one of the separated by pink wool).
Remarks.
Protoconch smooth, R1 with irregular, very fine radial lines; R2 moderately long (ca. 90°), seemingly smooth, with alternating lighter/slimmer and darker/thicker stripes.
The replacement name niosiensis refers to the type locality (Niosi Ridge).
Remarks.
Alycaeus (Dioryx) urnuladaflaensis Godwin-Austen, 1914 is a junior primary homonym of Alycaeusdaflaensis Godwin-Austen, 1876. Therefore, a replacement name is proposed here.
AnimaliaGastropodaCyclophoridae1AEAE428-1569-55D2-8CC5-DCC342632DF3DioryxurnularotundusPáll-Gergelynom. nov.Alycaeus (Dioryx) urnulavar.globosusGodwin-Austen, 1914: 363, pl. 153, fig. 8 (non Alycaeusglobosus H. Adams 1870).Dioryxurnulavar.globosa– Gude 1921: 202.Type locality.
“Brahmakund, E. Assam”.
Material examined.
Brahmakund, E. Assam, leg. M. Ogle, NHMUK 1903.7.1.2532 (17 syntypes).
Etymology.
The replacement name rotundus (Latin for spherical, globular, round, circular) refers to the shell shape of this subspecies.
Remarks.
Protoconch matte; R1 with irregular, low riblets and somewhat weaker spiral striation; R2 ca. 90° in length, smooth, with alternating lighter/slimmer and darker/thicker stripes.
Alycaeus (Dioryx) urnulavar.globosus Godwin-Austen, 1914 is a junior primary homonym of Alycaeusglobosus H. Adams 1870. Here we propose rotundus as a replacement name for the junior homonym.
The whole shell is nearly smooth; protoconch matte; R1 with irregular, very fine growth lines; R2 moderately long (ca. 90°), it has alternating slimmer/lighter and thicker/darker stripes.
Alycaeus (Metalycaeus) melanopoma Pilsbry, 1900 (Fig. 31A) (junior synonym of Chamalycaeusnipponensis [Reinhardt, 1877], see Minato 1988), SD Thiele (1929: 108).
Diagnosis.
Shell small to very large (D: 3–10 mm), spire usually low, protoconch usually elevated, spirally striated (key character); spiral striation of protoconch very rarely absent (see under M.laevis); R1 usually reticulated with sometimes prominent radial ribs; spiral lines always present on the teleoconch; R2 rarely short, usually long to very long, sometimes entirely smooth, but typically with widely spaced, straight, sharp ribs; R3 usually well-developed, although can be reduced. Operculum thin or relatively thickened, sometimes with funnel-shaped outer surface caused by the modification of the multispiral outer laminae. Radula usual for the family (central tooth with five cusps, broad, central cusp pointed).
Differential diagnosis.
Metalycaeus is recognised based on the spirally striated protoconch, which distinguishes Metalycaeus from species assigned to all other alycaeid genera.
Distribution.
Metalycaeus is widely distributed from the southeastern Himalaya to the south of Honshu Island (Japan) and the northern islands of the Philippines, but it does not extend any further south than northern Laos and Vietnam (Fig. 9).
The Metalycaeus records in Páll-Gergely and Auffenberg (2019) from Borneo (Chamalycaeuseveretti) and Sumatra (Chamalycaeussumatranus) later proved to be Chamalycaeus species.
Remarks.
The type species of Raptomphalus (Metalycaeusmagnificus) (Fig. 31B) and M.oakesi have a prominently keeled umbilicus which serve as the distinctive character for the genus Raptomphalus. A less prominently keeled umbilicus is observable in other species, such as Chamalycaeusvulcani and Metalycaeusbrahma, which can be interpreted as intermediate forms between the non-keeled umbilicus of alycaeids and the keeled umbilicus of Raptomphalus. Moreover, a variety of Metalycaeusbrahma has much less prominent umbilical keel than typical species in the genus. Therefore, the keeled umbilicus cannot serve as a distinctive character between genera. Consequently, Raptomphalus is a synonym of Metalycaeus.
For sake of simplicity, this genus is divided here into typical and atypical (or questionable) species. The first includes those having sharp, widely spaced R2 ribs, whereas the second includes species with R2 sculpture other than typical, including those which could not be examined.
Type species of alycaeid genus-group taxa AAlycaeusmelanopoma Pilsbry, 1900 (synonym of Chamalycaeusnipponensis (Reinhardt, 1877)), lectotype (ANSP 78815; type species of Metalycaeus); BMetalycaeusmagnificus (Godwin-Austen, 1914), syntype (NHMUK 1903.7.1.3115; type species of Raptomphalus). Photographs: downloaded from the website of ANSP (A), Harold Taylor (B).
https://binary.pensoft.net/fig/468676Typical MetalycaeusAnimaliaGastropodaCyclophoridae7E1D7FAD-3595-50D1-9383-AAB355F8129CMetalycaeusbeddomei(Godwin-Austen, 1914)AlycaeusbeddomeiGodwin-Austen, 1914: 386, pl. 149, figs 5, 5a.Alycaeusbeddomei– Gude 1921: 205–206; Ramakrishna and Mitra 2002: 21.Alycaeus (Alycaeus) beddomei– Ramakrishna et al. 2010: 46.Type locality.
“Naga Hills”.
Material examined.
Naga Hills, coll. Beddome, NHMUK 1912.4.16.294 (5 syntypes).
“Brahmakund” (brahma); “Chanjuk La, Tsanspu Valley, 4300 ft., Lat. 29°25', Long. 95°20'” (chanjukensis); “Bhutan” (commutatus).
Material examined.
Brahmakund, E. Assam, leg. M. Ogle, NHMUK 1903.7.1.2610 (16 syntypes of A.brahma); Dihung valley, Singpho Hills, leg. M. Ogle, NHMUK 1903.7.1.2611 (1 specimen of “Metalycaeus brahma (Godwin-Austen, 1886) var.”); Chanjuk La, Tsanspu valley, Lat. 29°25', Long. 95°20’, coll. Oakes, NHMUK 1903.7.1.3583 (3 syntypes of A.chanjukensis); Bhutan, ex Beddome coll., NHMUK 1912.4.16.293 (2 syntypes of A.commutatus).
Remarks.
Protoconch elevated, spirally striated; R1 spirally striated; R2 very long, ribs on R2 sharp and widely spaced.
The variety “M. brahma var.” has a much less prominent keel inside the umbilicus, which supports our view regarding the invalid status of Raptomphalus.
Alycaeuscommutatus and Alycaeuschanjukensis do not differ from M.brahma in any important shell characters. The shell and aperture shapes, sculpture, and ratios of shell regions (R1 and R2) are identical. Thus, those two species are considered as synonyms of M.brahma.
Shells of Metalycaeusbrahma (Godwin-Austen, 1886) A syntype of A.brahma (NHMUK 1903.7.1.2610) B syntype of A.commutatus Godwin-Austen, 1914 (NHMUK 1912.4.16.293) C syntype of A.chanjukensis Godwin-Austen, 1914 (NHMUK 1903.7.1.3583). All photographs: Kevin Webb (NHM).
https://binary.pensoft.net/fig/468677AnimaliaGastropodaCyclophoridae0387AB5D-E5C8-5DE5-B7F7-5B81338DBED8Metalycaeusburtii(Godwin-Austen, 1874)AlycaeusBurtiiGodwin-Austen, 1874: 149–150, pl. 3, fig. 9.Alycaeusburtii– Reeve 1878: pl. 3, species 27; Godwin-Austen 1914: 352–353, pl. 144, figs 8, 8a.Alycaeus (Alycaeus) burti[sic] – Kobelt 1902: 342; Ramakrishna et al. 2010: 47; Tripathy et al. 2018: 789.Alycaeusburti[sic] – Gude 1921: 206–207.Type locality.
“Foot of the Bhutan Himalaya at the debouchement of the Barowli River, Assam”.
Material examined.
Barowli Gorge, Tezpur Dist., Assam, NHMUK 1903.7.1.2492 (holotype [single specimen mentioned in the original description]).
Philippines (N-Luzon), ‘ad Digallorin vicum insulæ Luzon’ (Digollorin Bay: 16°50'N, 122°26'E), coll. C. Semper, 18 August 1860, SMF 158415 (2 syntypes); Luzon, Philippines, V W. MacAndrew Collection, Acc. no. 1563, NHMUK 20150120 (2 shells).
Remarks.
Protoconch elevated, spirally striated; R1 with irregular ribs and spiral striae of ca. the same strength, or slightly even stronger than the ribs; R2 relatively short, with alternating darker/wider and slimmer/lighter stripes; its surface is nearly smooth.
AnimaliaGastropodaCyclophoridaeAF0EA8B1-D3BF-5159-8266-741D517167AEMetalycaeuscrenulatus(Benson, 1859)AlycaeuscrenulatusBenson, 1859: 180–181.Alycaeuscrenulatus– Reeve 1878: pl. 5, species 43; Godwin-Austen 1871: 90, pl. 3, fig. 4; Godwin-Austen 1914: 337–338, pl. 133, figs 1, 1a–c.Alycaeus (Dicharax) crenulatus– Kobelt 1902: 367; Gude 1921: 242.Chamalycaeus (Dicharax) crenulatus– Ramakrishna et al. 2010: 57; Tripathy et al. 2018: 789.Type locality.
“in valle Rungun”.
Material examined.
Damsang, W. Bhutan, leg. Robert, NHMUK 1903.7.1.1254 (7 shells).
Remarks.
Protoconch elevated, spirally striated; R1 with rather regular ribs and weaker spiral striae; R2 short with widely spaced, high, sharp ribs.
“In insula Catanduanes leg. J. Quadras, prope vicum Caramuan insulae Luzon”.
Material examined.
Philippinen: Caramuan (Luzon, Camarines), coll. Möllendorff, SMF 109473 (lectotype, designated by Zilch 1957); Philippinen: Catanduanes, SMF 109475 (8 paralectotypes); Catanduanes, Philippines, ‘S&F Ad. 31/10/07, da Costa Collection’, V. W. MacAndrew Collection, Acc. no. 1563, NHMUK 20150119 (3 shells).
Remarks.
Protoconch elevated, spirally striated; R1 with dense, weak ribs and spiral striae of the same strength; R2 moderately long, with relatively dense and low ribs.
Toruputu Peak, Dafla Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2494 (2 syntypes).
Etymology.
We dedicate this species to H. H. Godwin-Austen (1834–1923), who described the majority of the alycaeid species in the Himalaya region.
Remarks.
Protoconch elevated, spirally striated; R1 rather glossy, but very fine reticulated (ribs and spiral striation or roughly equal strength); R2 short, with widely spaced, sharp ribs.
Alycaeusneglectus Heude, 1885 belongs to the genus Metalycaeus due to the spirally striated protoconch. This is the case even if it is treated here as a junior synonym of Metalycaeusrathouisianus (Heude, 1882). Thus, A.neglectus Godwin-Austen, 1914 is a primary as well as a secondary junior homonym of Alycaeusneglectus Heude, 1885 and thus, a replacement name (godwinausteni) must be given.
Formosa, leg. Hungerford, NHMUK 1891.3.17.790–791 (probable syntypes, see Hwang 2014).
Remarks.
Protoconch elevated, spirally striated; R1 very finely, irregularly ribbed, and striated with equally strong striae; R2 very short, consists of less than 10, sharp, widely spaced, rather irregular ribs which are slightly bent in the direction of their anterior neighbours.
AnimaliaGastropodaCyclophoridaeB6715413-F59A-5617-B2F2-31066BE12613Metalycaeusinflatus(Godwin-Austen, 1874)AlycaeusinflatusGodwin-Austen 1874: 146, pl. 3, fig. 1.Alycaeus (Chamalycaeus) inflatus– Kobelt 1902: 356–357; Gude 1921: 227–228.Alycaeusinflatus– Godwin-Austen 1914: 392–393, pl. 144, figs 1b–d.Alycaeusinflatusvar. – Godwin-Austen 1914: 394, pl. 144, figs 1, 1a.Chamalycaeus (Chamalycaeus) inflatus– Ramakrishna et al. 2010: 53.Type locality.
“Naga Hills under Japvo Peak”; “Yémi, Phúnngum, and Gaziphimi at the head of the Lanier River”.
Material examined.
Japvo Peak, 10000, Naga Hills, NHMUK 1903.7.1.2536 (3 syntypes in 2 vials).
Remarks.
Protoconch elevated, spirally striated; R1 with very weak, irregular ribs, and approximately as strong spiral lines; R2 relatively long, with sharp, straight, widely spaced ribs.
AnimaliaGastropodaCyclophoridae728E54D6-63F5-571B-8D18-BD929AFC6F77Metalycaeuskamakiaensis(Godwin-Austen, 1914)AlycaeuskamakiaensisGodwin-Austen, 1914: 375–376, pl. 141, fig. 8.Alycaeuskamakiaensis– Gude 1921: 209.Alycaeus (Alycaeus) kamakiaensis– Ramakrishna et al. 2010: 48.Type locality.
“Kamakia Temple Hill near Gowhatty, Assam”.
Material examined.
Kamakia Hill, Gowhatty, Assam, NHMUK 1903.7.1.2705 (1 syntype).
Remarks.
Protoconch elevated, spirally striated; R1 spirally striated; R3 moderately long, ribs are sharp and widely spaced.
Kengtung, Siam Frontier, leg. Woodthorpe, NHMUK 1903.7.1.3037 (holotype [single specimen mentioned in the original description]).
Remarks.
The whole shell is weathered; therefore, the sculpture could only be examined in very small regions of the shell. Protoconch elevated, some slight signs of spiral striation are visible; R1 regularly ribbed, spiral lines are not visible; R2 very long, the ribs are very widely spaced and sharp.
AnimaliaGastropodaCyclophoridae14C9D465-7B10-54FE-A918-6BDB4784328AMetalycaeuslaosensisPáll-Gergely, 2017MetalycaeuslaosensisPáll-Gergely in Páll-Gergely et al., 2017: 86–87, fig. 47C.Metalycaeuslaosensis– Inkhavilay et al. 2019: 16, fig. 6D.Type locality.
“Northern Laos, Phongsaly Province, old forest near stream approx. 1 km SW of a stream and Nam Ou (river) confluence, 493 m, 21°44.663'N, 102°10.999'E”.
Material examined.
Holotype (MNHN IM-2012-27172) and paratypes, see Páll-Gergely et al. (2017).
Remarks.
Protoconch elevated, spirally striated, R1 with irregular, low, widely spaced ribs and fine spiral striation; R2 short, regularly, densely ribbed, ribs rather straight in posterior part of region, ribs gradually curved towards aperture.
AnimaliaGastropodaCyclophoridae01B270F7-E744-5173-A787-DCE350500C1BMetalycaeuslatecostatus(Möllendorff, 1882)Alycaeuslatecostatus– Möllendorff, 1882: 346, pl. 10, fig. 7.Alycaeus (Orthalycaeus) latecostatus– Möllendorff 1886: 171.Alycaeus (Chamalycaeus) latecostatus– Kobelt and Möllendorff 1897: 148; Kobelt 1902: 358.Chamalycaeuslatecostatus– Yen 1939: 29–30, pl. 2, fig. 34.Chamalycaeus (Chamalycaeus) latecostatus– Zilch 1957: 143, pl. 5, fig. 11.Metalycaeuslatecostatus– Páll-Gergely et al. 2017: 88, fig. 47B.Type locality.
“Bei Lien-tschou auf Tropfsteinfelsen”.
Material examined.
China: Lo-fou-schan (Kwangtung), leg. Möllendorff, coll. Kobelt, SMF 39215 (lectotype, designated by Yen 1939); Same data, SMF 39299 (2 paralectotypes); Same data, SMF 39216 (8 shells).
Remarks.
Protoconch elevated, spirally striated; R1 with very strong ribs with very weak spiral lines between them; R2 short, with widely spaced, sharp ribs, which are usually slightly bent in the direction of their anterior neighbours.
AnimaliaGastropodaCyclophoridaeD5FE8F67-D3A0-52B2-B859-7C746CEF5A9CMetalycaeuslohitensis(Godwin-Austen, 1914)AlycaeuslohitensisGodwin-Austen, 1914: 362–363, pl. 137, figs 1, 1a.Alycaeuslohitensis– Gude 1921: 210.Alycaeus (Alycaeus) lohitensis– Ramakrishna et al. 2010: 48; Tripathy et al. 2018: 789.Type locality.
“Brahmakund, Eastern Assam”.
Material examined.
Brahmakund, Eastern Assam, leg. Ogle, NHMUK 1903.7.1.2493 (19 syntypes); Brahmakund, E. R. Sykes Collection, Acc. no. 1825, NHMUK 20150125 (1 shell) (matches the description and the figure in the orig. description).
Remarks.
Protoconch elevated, spirally striated; R1 with rather regular, widely spaced ribs with somewhat weaker spiral striae between the ribs; R2 long, with sharp, widely spaced ribs.
Both shells were somewhat weathered. Protoconch elevated, spirally striated; R1 spirally striated; R2 relatively long, with dense, low ribs; they might be sharper in fresh shells.
AnimaliaGastropodaCyclophoridaeB776AC86-9F45-5EF7-B427-95E26219E1EFMetalycaeusmacgregori(Godwin-Austen, 1914)AlycaeusmacgregoriGodwin-Austen, 1914: 356–357, pl. 141, figs 2, 2a, 2b.Alycaeus (Chamalycaeus) macgregori– Gude 1921: 229.Chamalycaeus (Chamalycaeus) macgregori– Ramakrishna et al. 2010: 54; Tripathy et al. 2018: 789.Type locality.
“Dafla Hills”.
Material examined.
Dafla Hills, NHMUK 1903.7.1.2521 (holotype [fixed by original designation]); Shengorh, Dafla Hills, coll. Godwin-Austen, NZSI M.8058 (2 shells, labelled as co-type = paratypes).
Remarks.
Protoconch elevated, spirally striated; R1 finely ribbed and spirally striated; R2 relatively short with widely spaced ribs, but the fine structure of the ribs was not visible due to corrosion.
AnimaliaGastropodaCyclophoridae6D49DDFD-5AFB-5E42-97DB-32A82DBDCC8DMetalycaeusmuciferus(Heude, 1885)Fig. 33Alycoeus[sic] muciferus Heude, 1885: 96, pl. 24, figs 1, 1a.Alycaeus (Orthalycaeus) muciferus– Möllendorff 1886: 171.AlycaeushelicodesGredler, 1888: 365.AlycaeusexpansusHeude, 1890: 129, pl. 38, fig. 2. syn. nov.Alycaeus (Chamalycaeus) muciferus– Kobelt and Möllendorff 1897: 149; Kobelt 1902: 359.Metalycaeusmuciferus– Páll-Gergely et al. 2017: 88–93, figs 58, 59, 60, 61 (helicodes Gredler is a synonym).“Chamalycaeus” expansus – Páll-Gergely et al. 2017: 105, fig. 69L. Type locality.
China, Tchen-k’eou, MCZ 167230 (7 syntypes of A.muciferus); Tchen-k’eou, China, Pe-shang, Hunan, leg. Fuchs, coll. Gredler, SMF 192225 (2 paralectotypes of A.helicodes, lectotype in the Franziskanergymnasium Bozen [Bolzano, Italy], designated by Zilch 1974); HMT-215a, syntype: labelled as lectotype of A.expansus deposited in IZCAS: Tchen-k’eou (Cheng-kou County, Chong-qing, China) (Fig. 33).
Remarks.
Protoconch conspicuously elevated from dorsal surface, its end with inconspicuous spiral striations; R1 with irregular, widely spaced ribs and fine spiral sculpture; R2 with dense, sharp, elevated ribs.
No type specimens of Alycaeusexpansus were reported in American museums (Johnson 1973). We were not able to examine the types deposited in the Beijing Museum before the revision of Chinese Alycaeidae (Páll-Gergely et al. 2017), but could do so now. The general shell shape, colour, sculpture, and the lengths and proportions of R2 and R3 agree with those of A.muciferus, which was described from the same type locality. Consequently, we move Alycaeusexpansus to the synonymy of A.muciferus.
https://binary.pensoft.net/fig/468678AnimaliaGastropodaCyclophoridae1446E8DD-A9F0-5CFC-88C5-3C5D2CA3A25DMetalycaeusmundulus(Godwin-Austen, 1914)AlycaeusmundulusGodwin-Austen, 1914: 357–358, pl. 149, fig. 8.Alycaeusmundulus– Gude 1921: 212.Alycaeus (Alycaeus) mundulus– Ramakrishna et al. 2010: 49; Tripathy et al. 2018: 789.Type locality.
“Torúpútú, Dafla Hills”.
Material examined.
Toruputu, Dafla Hills, leg. Godwin-Austen, NHMUK 1906.1.1.955 (holotype [single specimen mentioned in the original description]).
Remarks.
The entire shell was weathered, but the following information could be obtained: protoconch strongly elevated, some spiral striation visible near suture (all other parts are strongly weathered); R1 with regular, strong ribs and spiral striae; R2 moderately long, rib structure could not be observed.
Hakone Mts, Japan, leg. Schmacker, ANSP 78815 (lectotype of A.melanopoma, designated by Baker 1964, photographs examined); Japan, Yedo, Doenitz l., coll. Reinhardt, SMF 185407 (labelled as ‘typus’); 東京都 (Tōkyō-to), 多摩市 (Tama-shi), 蓮光寺聖跡記念館 境内 (Renkō-ji Seiseki Kinenkan keidai) [Tama City, Tokyo Metropolitan Prefecture], NSMT-Mo 79102 (91 shells).
Remarks.
Protoconch elevated, its last 0.5 whorl with spiral lines; ribbing on R2 is quite variable; there are 14–22, usually elevated and distinct ribs. In some specimens the ribs are relatively irregular, and they seemingly join to each other near the tube. This is, however, rather due to the fact that the shells are weathered (despite the fact that they were collected alive), because the elevated ribs are destroyed and appear lower. R1 with clearly visible, fine spiral striation.
In the examined sample, 91 specimens had an operculum, and of those 53 of them had the outer black circle present. In case of 27 specimens the outer circle was entirely missing, and in 11 cases only small fragments of the rings remained.
“Chanjuk La, in Tsanspu Valley 4300 ft., Lat. 29°25´, Long. 95°20´”.
Material examined.
Abor Hills, Lat. 29°25, Long. 95°20, NHMUK 1903.7.1.3578 (5 syntypes).
Remarks.
In the original description the subgeneric name Raptomphalus is not used. However, the species is introduced after Alycaeus (Raptomphalus) magnificus, and the characteristically keeled umbilicus is similar in both species.
All four shells were strongly weathered, but the characteristic elevated protoconch was recognisable. Moreover, some indication of spiral striation was visible on both the protoconch and R1, therefore the classification of M.oakesi in Metalycaeus is justified. R2 long, with widely spaced, sharp ribs.
AnimaliaGastropodaCyclophoridae552F466C-74F3-5A1E-9C96-2CEBD35EC09EMetalycaeusoharaiPáll-Gergely & Hunyadi, 2017MetalycaeusoharaiPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 93–96, figs 48C–F, 62A, B, 63C, D.Type locality.
Holotype (HNHM 99712) and many other samples, see Páll-Gergely et al. (2017).
Remarks.
Protoconch elevated, finely spirally striated; R1 with sharp ribs and clearly visible, fine spiral lines resulting in a reticulated surface; R2 ribs ca. 2–3 × more densely arranged as in R1, ribs very sharp and high.
AnimaliaGastropodaCyclophoridaeC4CA630A-81B5-5379-8556-8E0322A1F9A6Metalycaeusphysis(Benson, 1859)AlycaeusphysisBenson, 1859: 179.Alycaeusphysis– Reeve 1878: pl. 6, species 51; Godwin-Austen 1914: 342, pl. 134, figs 1, 1a.Alycaeus (Chamalycaeus) physis– Kobelt 1902: 361; Gude 1921: 231–232, fig. 34.Alycaeus (Orthalycaeus) physis– Rees 1964: 64, pl. 5, fig. 27.Chamalycaeus (Chamalycaeus) physis– Ramakrishna et al. 2010: 55.Type locality.
“in valle Rungit (alt. 2000 ped.), prope Darjiling”.
Spire low, but protoconch elevated, spirally striated; R1 rather irregularly ribbed, with weak spiral lines; R2 long, with waved surface; there are light and dark stripes alternating.
AnimaliaGastropodaCyclophoridae9A78308B-B92B-50E9-9651-996E8F367495Metalycaeuspolygonoma(W. T. Blanford, 1862)AlycaeuspolygonomaW. T. Blanford, 1862: 140–141.Alycaeuspolygonus[sic] – Reeve 1878: pl. 2, species 11.Alycaeus (Dicharax) polygonoma– Kobelt 1902: 375; Gude 1921: 265–266.Alycaeuspolygonoma– Godwin-Austen 1914: 423, pl. 141, fig. 5.Type locality.
“in montibus Arakanensibus”.
Material examined.
Tongoop Pass, Arakan Hills, NHMUK 1906.4.4.51 (2 syntypes in different vials).
Remarks.
Protoconch elevated, spirally striated; R1 with irregular, fine ribs and spiral lines of approximately the same strength; R2 long, with sharp, widely spaced ribs.
AnimaliaGastropodaCyclophoridaeE175F738-BB97-5EE1-AEE3-2C0CB372FBA5Metalycaeusprosectus(Benson, 1857)AlycaeusprosectusBenson, 1857: 203–204.Alycaeusprosectus– Reeve 1878: pl. 6, species 49; Godwin-Austen 1914: 380–381, pl. 143, figs 1, 1a, 1b.Alycaeus (Dicharax) prosectus– Kobelt 1902: 375–376; Gude 1921: 266–267.Chamalycaeus (Dicharax) prosectus– Ramakrishna et al. 2010: 65.Type locality.
Protoconch elevated, spirally striated; R1 with irregular, low ribs and spiral striae of the same strength; R2 relatively long, with sharp, relatively widely spaced, straight ribs.
AnimaliaGastropodaCyclophoridaeC118395E-6835-5CCB-A5C7-B124C1EF07A1Metalycaeusquadrasi(Möllendorff, 1895)AlycaeusquadrasiMöllendorff in Quadras & Möllendorff, 1895: 83.Alycaeus (Chamalycaeus) quadrasi– Kobelt 1902: 362.Chamalycaeus (Chamalycaeus) quadrasi– Zilch 1957: 145, pl. 6, fig. 18.Metalycaeusquadrasi– Páll-Gergely and Auffenberg 2019: 385, fig. 8B.Type locality.
“prope vicum Buguey provinciae Cagayan”.
Material examined.
Philippinen, Buguey (Luzon, Cagayan), leg. Möllendorff, SMF 109515 (lectotype, designated by Zilch 1957); Same data, SMF 109516 (5 paralectotypes).
Remarks.
Protoconch elevated, spirally striated; R1 with extremely dense, fine ribs and spiral lines of approximately of the same strength; R2 relatively short, with widely spaced, sharp ribs.
“E collibus juxta civitatem Song-kiang (松江), provinciæ Kiang-sou, ad montes districtus Tong-lieou, sed non ubique” (Alycaeusrathouisianus); “ad rupestres colles civitatis Kien-té, provinciæ Ngan-houé legeram” (Alycaeusneglectus); “Ad calcarios lapides juxtà civitatem Tchen-kiang (Kiang-sou)”.
Material examined.
China, Tsing-p’on, MCZ 167143 (23 syntypes of A.rathouisianus); Sung-kiang: China, coll. Möllendorff ex coll. Heude, SMF 39243 (5 syntypes of A.rathouisianus); same data, 39303 (3 syntypes of A.rathouisianus); China, Tchein-kiang (Kiang-sou), MCZ 167209 (29 syntypes of A.dolomiticus); HMT-221a (labelled as lectotype of Alycaeusdolomiticus, Fig. 34) deposited in IZCAS: locality information were deleted; HMT-221 (labelled as paralectotype of Alycaeusdolomiticus), same data with HMT-221; HMT-220a (1 syntype, labelled as lectotype of Alycaeusneglectus, Fig. 35), deposited in IZCAS: Tong-lieu (Dong-Liu town, Dong-Zhi County, Chi-Zhou City, An-Hui Province, China); Same data, HMT-220 (1 syntype, labelled as paralectotype of Alycaeusneglectus).
Remarks.
Alycaeusdolomiticus Heude, 1890 is a synonym of Metalycaeusrathouisianus (see Páll-Gergely et al. 2017).
According to Johnson (1973), paratypes of A.neglectus are present in the MCZ (inv. number: 167222) and the USNM (inventory number: 472336). In the revision of Chinese Alycaeidae (Páll-Gergely et al 2017) we examined the former sample, but the locality (“China, Tong-lieou” in Anhui Province) did not match the type locality given in the original description (“Kien-té” in Zhejiang Province). Consequently, we did not consider them as type specimens; moreover, on a hand-written label of the MCZ 167222 sample the following text was written: “not types, only holotype in Heude’s coll.”. We have now been able to examine the specimens in the Beijing museum, and found that the specimens labelled as types also have the same type locality (“Tong-lieu”). The Beijing specimens were also labelled as lectotype (HMT-221a), and paralectotypes (HMT-221). It appears that although the locality on the type specimen labels and that in the original description do not match, the shells labelled as types are indeed types of A.neglectus. Since we did not find traces of the lectotype designation, we consider all type specimens of A.neglectus syntypes.
Since we did not consider the specimens in the MCZ types in Páll-Gergely et al. (2017), we had to rely on the original description regarding the identity of A.neglectus, and based on the low protoconch in the figure of the original description, we tentatively classified it in Dicharax. However, this opinion must now be revised: the shells labelled as types in the Beijing Museum and in the MCZ are identical, and they are also identical to typical Alycaeusrathouisianus shells. Thus, A.neglectus is considered a synonym of A.rathouisianus.
All three shells were strongly weathered. Protoconch moderately elevated, with slight indication of spiral striae; R1 with regular, widely spaced ribs and very slight indication of spiral striae; R2 relatively short, with widely spaced, sharp ribs.
AnimaliaGastropodaCyclophoridae4B25B40B-E359-5AE7-A94E-2B14A24A0C74Metalycaeussatsumanusawaensis(Pilsbry & Y. Hirase, 1904)AlycaeusawaensisPilsbry & Y. Hirase, 1904a: 117.Chamalycaeusawaensis– Azuma 1982: 11, pl. 3, fig. 34.Chamalycaeussatsumanusawaensis– Minato 1988: 14.Metalycaeussatsumanusawaensis– Páll-Gergely and Asami 2017: 4.Type locality.
Protoconch moderately elevated, very finely granulated, the last 0.5 whorl with fine spiral striae; R2 with 18–20 ribs; the ribs are widely spaced even near the tube; at the edge of the body whorl the gap between the ribs is ca. as wide as a rib itself; the first 0.75–1 whorl of R1 has clearly visible spiral striae.
AnimaliaGastropodaCyclophoridae51FE7038-B691-5E5F-91F7-8BA11FBC7B61Metalycaeussatsumanuslaevicervix(Pilsbry & Y. Hirase, 1904)AlycaeuslaevicervixPilsbry & Y. Hirase, 1904c: 618.Chamalycaeussatsumanuslaevicervix– Minato 1988: 14.Metalycaeussatsumanuslaevicervix– Páll-Gergely and Asami 2017: 4.Type locality.
“Kuchinoerabushima, Ōsumi”.
Material examined.
Kuchinoerabushima, leg. Hirase, 1904, ANSP 87699 (1 syntype, photographs examined); label on the right: 大隅永良部嶋 (Osumi, Erabu-jima = Osumi, Erabu Island), label on the left: 大隅永良部島 is corrected by insertion as 大隅口之永良部島 (Osumi, Kuchinoerabu-jima = Osumi, Kuchinoerabu Island), Hirase coll., NSMT-Mo 2053 (3 shells).
Remarks.
Protoconch moderately elevated, rather roughly granulated, the granules are arranged into spiral lines on the last 0.25–0.5 of whorl; R2 with ca. 15 ribs.
AnimaliaGastropodaCyclophoridae54FA0DC3-6D30-555F-9F67-91FD28EC6D17Metalycaeussatsumanus(Pilsbry, 1902)AlycaeussatsumanaPilsbry, 1902b: 548–549.Chamalycaeussatsumanus– Azuma 1982: 10, pl. 3, fig. 31.Chamalycaeussatsumanussatsumanus– Minato 1988: 14.Metalycaeussatsumanus– Páll-Gergely et al. 2017: 103, 104; Páll-Gergely and Asami 2017: 4.Type locality.
Protoconch elevated, with spiral lines on its last whorl; R2 with ca. 22 widely spaced, low, blunt, regular ribs; the distance between the ribs at the edge of the body whorl is at least 2–3 × wider than the width of a rib; R1 with conspicuous spiral striation.
Alycaeusawaensis Pilsbry & Y. Hirase, 1904 and Alycaeuslaevicervix Pilsbry & Y. Hirase, 1904 are classified as a subspecies of Chamalycaeussatsumanus by Minato (1988), which we also follow here. Sigmacharaxtanegashimae was additionally classified under Chamalycaeussatsumanus in the same study, but these species belong to different genera according to our revision.
Protoconch elevated, finely spirally striated, consisting of ca. 1.5 whorls; R1 consisting of 1.75–2 whorls, very densely, finely ribbed and finely spirally striated, resulting in a fine, dense reticulated surface; R2 + R3 short, ca. 70–80° combined; R2 slightly shorter than R3; ribs on R2 more widely spaced and more elevated than those on R1, although still low; spiral striation also visible on R2 (Páll-Gergely and Auffenberg 2019).
AnimaliaGastropodaCyclophoridae99C4C0A4-C258-59FE-A6E9-823694F0FDEEMetalycaeussinensis(Heude, 1882)AlycaeussinensisHeude, 1882: 7, pl. 12, figs 13, 13a.Alycaeussinensis– Möllendorff 1882: 345.Alycaeus (Chamalycaeus) sinensis– Kobelt and Möllendorff 1897: 149; Kobelt 1902: 363.Metalycaeussinensis– Páll-Gergely et al. 2017: 103–104, fig 55.Type locality.
“Ad radices saxorum inter folia decidua in districtu Tong-lieou (東流), provinciæ Ngan-houé”.
Material examined.
China, Tong-lieou, MCZ 167216 (lectotype, designated by Páll-Gergely et al. 2017).
Remarks.
Protoconch relatively low, with very fine matte surface and barely visible spiral lines; R1 with rather regular ribs and much weaker spiral lines between; ribs become more widely spaced towards the end of R1; R2 with relatively low, widely spaced ribs, however all available shells were weathered.
AnimaliaGastropodaCyclophoridaeBBC2ED4F-F68C-5205-9CD0-3591F1F917F6Metalycaeusstylifer(Benson, 1857)AlycaeusstyliferBenson, 1857: 204.Alycaeusstylifer– Reeve 1878: pl. 6, species 46; Godwin-Austen 1914: 344, pl. 133, figs 2–2a–c.Alycaeus (Dicharax) stylifer– Kobelt 1902: 376–377; Gude 1921: 269–270, fig. 35.Chamalycaeus (Dicharax) stylifer– Ramakrishna et al. 2010: 66.Type locality.
Gazifhimi, N.E. Manipur, leg. Godwin-Austen, NHMUK 1903.7.1.2489 (7 syntypes, one specimen indicated with pink wool, other shells in another glass vial).
Remarks.
Protoconch elevated, spirally striated; R1 with weak, irregular ribs and spiral striae of similar strength; R2 relatively long, with sharp, widely spaced ribs.
AnimaliaGastropodaCyclophoridaeF90B9DF2-44C8-50F2-810C-BD059E8FC4F8MetalycaeussuhajdaiPáll-Gergelynom. nov.Alycaeus (Dioryx) variusGodwin-Austen, 1914: 402, pl. 157, figs 7, 7a. (non Alycaeusvarius Pilsbry & Y. Hirase, 1905)Dioryxvarius– Gude 1921: 204; Ramakrishna et al. 2010: 75.Type locality.
“Lhota Naga Hills”.
Material examined.
Lhota Naga, coll. Godwin-Austen, NHMUK 1903.7.1.2574 (1 syntype).
Etymology.
The specific name is dedicated to Szilárd Suhajda, the first Hungarian mountaineer to reach the peak of K2 (formerly known as Mount Godwin-Austen). Godwin-Austen was the first to fix the height and position of that mountain.
Remarks.
Metalycaeusvarius (Godwin-Austen, 1914) and Metalycaeusvarius (Pilsbry & Y. Hirase, 1905) are primary (and also secondary) homonyms. Therefore, the newer taxon needs a replacement name.
The syntype was strongly weathered, but the following observations could be made: protoconch moderately elevated, and although weathered, clearly spirally striated; R1 with strong, rather irregular ribs and weaker spiral striation; R2 very long, with elevated, sharp ribs near the tube (in other places the ribs are not visible due to corrosion).
The constriction is longer than in usual Dioryx, and the protoconch is spirally striated. Therefore, this species is transferred from Dioryx to Metalycaeus.
Philippinen: Montalban bei Manila, leg. Möllendorff, SMF 109520 (lectotype, designated by Zilch 1957); Same data, SMF 109521 (2 paralectotypes).
Remarks.
Protoconch elevated, spirally striated; R1 very densely ribbed and spirally striated of approximately the same strength; R2 relatively short, with widely spaced, sharp ribs.
AnimaliaGastropodaCyclophoridae9C3717D1-3C0C-5A88-892C-9FA7C507001EMetalycaeusyamneyensis(Godwin-Austen, 1914)Alycaeusyamnayensis(in the headings) and A.yamneyensis (in the text) Godwin-Austen, 1914: 368, pl. 156, fig. 2.Alycaeusyamneyensis– Gude 1921: 222–223.Alycaeus (Alycaeus) yamneyensis– Ramakrishna et al. 2010: 51; Tripathy et al. 2018: 789.Type locality.
“Yamne Valley, Abor Hills”.
Material examined.
Yamney valley, Abor Hills, NHMUK 1903.7.1.3114 (3 shells, one of them separated with pink wool).
Remarks.
Protoconch elevated, spirally striated; R1 rather regularly ribbed and some spiral striation visible; R2 extremely long, with widely spaced, sharp ribs.
This species was spelled both “yamnayensis” and “yamneyensis” in the original description. The former is considered an incorrect original spelling, because the name is derived from the Yamne Valley.
AnimaliaGastropodaCyclophoridaeF7B65B3F-CB66-5D60-A3DD-D743BD598363Metalycaeusvarius(Pilsbry & Y. Hirase, 1905)AlycaeusvariusPilsbry & Y. Hirase, 1905: 729.Chamalycaeusvarius– Hsieh et al. 2006: 87 + figures.Type locality.
ANSP 83291 (lectotype, designated by Baker 1964, photographs examined); for other examined specimens see Páll-Gergely and Asami (2017).
Remarks.
Protoconch elevated, consists of 1.5 whorls, spirally striated except for the first 0.5 whorl which is smooth; R2 with ca. 18 ribs, which are relatively far from each other even close to the suture; the ribs are low, but all available shells were somewhat weathered.
“Tashan Town, Zayu County (28°04'N ,97°E), Tibet Autonomous Region, China”.
Material examined.
Probably lost, see remarks.
Remarks.
The specimens labelled as the types (Tashan Town, Zayu County, Nyingchi City, Tibet Autonomous Region, China, leg. Chen De-Niu, 1980.7.13, IZCAS TM_094538, holotype; IZCAS TM_094540, paratype) of C.zayuensis represent a species different from the one characterised in the original description. Namely, the real C.zayuensis has a smooth R2 with dense stripes, and finely ribbed R1, whereas the ones we examined have elevated, sharp, widely spaced R1 and R2 ribs. Furthermore, the ones in the Beijing Museum have sharp swelling on R3 with an incision at the middle, whereas the ones in the original description possess a relatively low, simple R3 swelling.
Chamalycaeuszayuensis was previously moved to the genus Metalycaeus based on the spiral striation (Zhouxing and Wechuan 2015; Páll-Gergely et al. 2017).
https://binary.pensoft.net/fig/468681Atypical (or questionable) MetalycaeusAnimaliaGastropodaCyclophoridae6A7BC84A-C7FE-5080-B5F8-26FF1DA9B1C7Metalycaeus(?)aborensis(Godwin-Austen, 1914)AlycaeusaborensisGodwin-Austen, 1914: 364, pl. 149, fig. 9.Alycaeus (Chamalycaeus) aborensis– Gude 1921: 223.Chamalycaeus (Chamalycaeus) aborensis– Ramakrishna et al. 2010: 52; Tripathy et al. 2018: 789.Type locality.
“Bapu Peak, Abor Hills”.
Material examined.
Abor Hills, leg. Oakes, NHMUK 1903.7.1.3.102 (5 syntypes, one of them separated with pink wool).
Remarks.
Protoconch relatively low and spirally striated; R1 with rather regular, strong ribs and weaker spiral striation; R2 very long, there are no visible ribs which would elevate from the surface of the shell, alternate thicker/darker and slimmer/lighter lines, where the lighter ones represent microtunnels.
AnimaliaGastropodaCyclophoridae08F99FC3-E976-5216-B918-440EAAD25946Metalycaeus(?)awalycaeoidesPáll-Gergely & Hunyadi, 2017Metalycaeus(?)awalycaeoides Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 74, figs 47A, 48A, B, 49A, B.Type locality.
Protoconch elevated and some remains of spiral striation visible near suture; R1 irregularly ribbed and weakly spirally striated; R2 relatively short, R2 ribs strongly developed, curved towards aperture, nearly reaching their neighbours.
AnimaliaGastropodaCyclophoridae3B45BD64-FBF0-583C-AD95-2106D5385B9AMetalycaeus(?)dikrangensis(Godwin-Austen, 1914)AlycaeusdikrangensisGodwin-Austen, 1914: 355, pl. 148, figs 6, 6a.Alycaeusdikrangensis– Gude 1921: 209.Alycaeus (Alycaeus) dikrangensis– Ramakrishna et al. 2010: 48.Type locality.
“Toruputu Peak, Dafla Hills”.
Material examined.
Toruputu Peak, Dafla Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2533 (13 syntypes in 3 different vials).
Remarks.
Protoconch elevated, spirally striated; R1 regularly ribbed, spirally striated; R2 very long, alternate thicker/darker and slimmer/lighter stripes, the ribs are only very slightly elevated from the surface.
AnimaliaGastropodaCyclophoridae1B0D8BE8-8C20-5642-ABEB-8E98F709C0CBMetalycaeus(?)ibexPáll-Gergely & Hunyadi, 2017Metalycaeus(?)ibexPáll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 84, fig. 55E.Type locality.
Sculpture of protoconch could not be examined due to strong corrosion; R1 rather regularly ribbed, with some additional, incomplete riblets between main ribs; spiral structure visible on some, not strongly weathered parts of R1; R2 very densely ribbed.
AnimaliaGastropodaCyclophoridae56944E6D-C895-5471-A03B-9F2ED27D9F75Metalycaeus(?)laevis(Pilsbry & Y. Hirase, 1909)AlycaeuslaevisPilsbry & Y. Hirase, 1909a: 588.Chamalycaeuslaevis– Minato 1988: 15, pl. 2, figs 6–8.Metalycaeuslaevis– Páll-Gergely and Asami 2017: 4.Type locality.
The NSMT sample we examined contained six shells with clearly visible protoconch sculpture. Four shells had spiral lines on the protoconch but in the case of two shells no spiral striae were visible. In all cases the protoconch was elevated. R2 with ca. 36 ribs, which are only just elevated from the surface of the shell. They are seemingly fully attached to their neighbours. There are no spiral lines visible on R1. The spiral striae stop at the protoconch-teleoconch border.
AnimaliaGastropodaCyclophoridaeFF1A1C75-992C-5D3C-90AA-833E156CFD5AMetalycaeus(?)libonensis(Chen, Li & Luo, 2003)Fig. 37ChamalycaeuslibonensisChen et al. 2003: 619–620, figs 1–3.Metalycaeus(?)libonensis – Páll-Gergely et al. 2017: 105.Type locality.
“Feng Dong, Libo County (25°4'N, 107°8'E), Guizhou Province, China”.
Protoconch relatively elevated, some spiral striation is visible on the photographs; R1 with relatively widely-spaced ribs and fine spiral striation; R2 + R3 ca 90° combined R2 slightly shorter than R3; R2 ribs sharp but relatively low, similar to those on R1; R3 with a low but relatively sharp swelling.
Yamney Valley, Abor Hills, NHMUK 1903.7.1.3115 (1 syntype).
Remarks.
The entire shell weathered, but the following observations could be made. Protoconch rather elevated, some spiral striae are recognisable near the suture; R1 without visible spiral lines; R2 short, very densely ribbed, but the fine morphology of the ribs could not be seen.
Holotype (NSMT-Mo 78937) and paratypes, see Páll-Gergely and Asami (2017).
Remarks.
protoconch elevated, spiral lines mostly visible on last 0.25 whorl; R1 with rather irregular, widely spaced ribs and fine spiral lines; R2 short, very densely ribbed, alternating very narrower/lighter and wider/dark bands.
AnimaliaGastropodaCyclophoridaeE4B5E75A-E7D5-566D-BD39-287865FE8EE2Metalycaeus(?)okuboiPáll-Gergely & Hunyadi, 2017Metalycaeus(?)okuboi Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 97–98, figs 53A, B, 62C, D, 64, 65C–F.Type locality.
Holotype (HNHM 99713) and several other samples, see Páll-Gergely et al. (2017).
Remarks.
Protoconch moderately elevated, spiral lines visible on the last whorl of protoconch (although there was no clear distinction between protoconch and teleoconch); R1 irregularly ribbed, ribs low but relatively sharp at the end of R1; there are signs of very weak spiral lines between ribs on R1; R2 ribs not erected but situated horizontally. For more details see the original description.
AnimaliaGastropodaCyclophoridaeDCF05423-28DF-58EB-81AA-C73041CB6401Metalycaeus(?)panggianus(Godwin-Austen, 1914)AlycaeuspanggianaGodwin-Austen, 1914: 367, pl. 156, figs 3, 3a.Alycaeuspanggianus– Gude 1921: 213–214.Alycaeus (Alycaeus) panggianus– Ramakrishna et al. 2010: 50; Tripathy et al. 2018: 789.Type locality.
“Sibbum, Abor Hills”.
Material examined.
Sibbum, Abor Hills, coll. Oakes, NHMUK 1903.7.1.3143 (holotype [single specimen mentioned in the original description]).
Remarks.
In the original description the subgeneric name Raptomphalus is not noted. However, the species is introduced after Alycaeus (Raptomphalus) magnificus, and the characteristically keeled umbilicus is similar to this species.
Protoconch elevated, spirally striated; R1 regularly ribbed, with recognisable spiral striae; R2 moderately long, with dense ribs, but their fine structure could not be examined due to the weathered condition of the holotype.
Protoconch elevated, very finely spirally striated; R1 irregularly wrinkled, glossy, without spiral lines; R2 moderately long, with low, blunt, simple ribs.
AnimaliaGastropodaCyclophoridaeF72FEB00-AFFB-5950-A5C4-D92E19595427Metalycaeus(?)sibbumensis(Godwin-Austen, 1914)AlycaeussibbumensisGodwin-Austen, 1914: 367, pl. 156, figs 4, 4a.Alycaeussibbumensis– Gude 1921: 219.Alycaeus (Alycaeus) sibbumensis– Ramakrishna et al. 2010: 51; Tripathy et al. 2018: 789.Type locality.
“Sibbum, Abor Hills”.
Material examined.
Sibbum, Abor Hills, leg. Oakes, NHMUK 1903.7.1.3142 (2 syntypes).
Remarks.
Protoconch elevated, spirally striated; R1 irregularly ribbed, with some signs of spiral striation; R2 very long, with alternating darker and lighter stripes forming a nearly smooth surface; the darker stripes probably represent the very low ribs.
AnimaliaGastropodaCyclophoridae8FCEFA02-9963-57ED-A2DC-F3FC58F5E96CMetalycaeus(?)toruputuensis(Godwin-Austen, 1914)Fig. 38AlycaeusTheobaldi– Godwin-Austen, 1876: 175–176, pl. 7, fig. 10. (operculum).Alycaeustheobaldi– Nevill 1878: 290.AlycaeustoruputuensisGodwin-Austen 1914: 359–360, pl. 149, figs 3, 3a, b, pl. 145, fig. 10 (operculum).Alycaeus (Dicharax) toruputuensis– Gude 1921: 274.Chamalycaeus (Dicharax) toruputuensis– Ramakrishna et al. 2010: 68.Type locality.
“on the slopes of Torúpútú peak in the Dafla Hills” (in Godwin-Austen 1914).
Material examined.
Slope of Toruputu peak, Dafla Hills, Assam, coll. Godwin-Austen, NZSI M.8037 (holotype, “the specimen figured is the type shell”, “type in Indian Museum, Calcutta”: Godwin-Austen 1914: 360); Toruputu Peak, Dafla Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2496 (6 paratypes in two different vials, one shell is indicated with pink wool).
Remarks.
Protoconch elevated, spirally striated; R1 with rather regular, strong ribs and much weaker spiral lines; R2 smooth, somewhat wavy, with thick darker and slimmer lighter stripes alternating.
Pincernaliratula Preston, 1907 (Fig. 39B) (introduced as a subgenus of Alycaeus, but apparently used on genus level), by monotypy.
Diagnosis.
Shell very small to medium sized (D: 2.5–6 mm), globose triangular, usually higher than wide; protoconch smooth, without spiral striation; R1 usually with strong, widely-spaced ribs and weak spiral striation; R2 smooth to prominently ribbed (in those cases not different from R1), very short, pyriform (in typical Cycloryx species) or somewhat longer in Pincerna-like shells; umbilicus often closed by the reflected columellar extension in many species. Operculum thin, in some species with a low circular structure, or calcareous ridges radiating outwards of the nucleus on the outer surface. Radula of a single species is known, see Table 1 (central tooth with 5 cusps, broad, central cusp pointed).
Type species of alycaeid genus-group taxa APincernaconstricta (Benson, 1851), syntype (NHMUK 1906.4.4.41; type species of Cycloryx) BPincernaliratula (Preston, 1907), syntype (NHMUK 1907.5.20.191; type species of Pincerna) CStomacosmethissarasinorum (Bollinger, 1918), lectotype (NHMB 2411.a; type species of Stomacosmethis). Photographs: Harold Taylor (A, B); Eduard Stoeckli (C).
The distribution of Pincerna seemingly consists of two disjunct geographic areas, namely the southeastern Himalaya to northern Vietnam and northern Laos, and Sumatra, the southern part of the Malay Peninsula, and northern Borneo (Fig. 40).
Remarks.
The relationship between Alycaeus, Pincerna, and Cycloryx is the most problematical point of the current revision. The genus Cycloryx Godwin-Austen, 1914 (type species: Cyclostomaconstrictum Benson, 1851, OD) was erected as a subgenus of Alycaeus Gray, 1850, and was described on the basis of the ovately conoid shell shape, the regular ribbing on the upper whorls, and the extremely short, often clubbed or pear-shaped sutural tube (Godwin-Austen 1914). Godwin-Austen (1914) only included species from northeastern India and Burma (Rakhin = Arakan, and the Shan States) in this genus. However, the diagnosis of Cycloryx matches several species extralimital to the distributional range as defined by Godwin-Austen, namely those from northern Vietnam, Borneo, China’s Guizhou Province, the Malay Peninsula, and Sumatra. The most striking example is the Sumatran Pincernayanseni and the northern Vietnamese Alycaeuscostulosus, which both look so similar to Indian and Burmese species that it was challenging to find any differences between those two and the Indian and Burmese taxa. One of those “extralimital Cycloryx” is Alycaeusliratulus Preston, 1907, known from the Malay Peninsula and Sumatra, which has been placed in its own subgenus, Pincerna Preston, 1907. Originally, Pincerna was diagnosed on the basis of a “circular cup” on the outer surface of the operculum. The outer surface of operculum, however, has limited taxonomic value at the generic level in the Alycaeidae, especially since the outer rings have evolved in multiple alycaeid genera (Dicharax, Metalycaeus, Stomacosmethis; see this study and Páll-Gergely et al. 2017). Moreover, the most similar species to P.liratula, P.thieroti, lacks the circular cup on the outer surface of the operculum. Consequently, no important conchological characters distinguish Cycloryx and Pincerna, and thus, they have been synonymised (Páll-Gergely 2017). See also under Genus-level diversity (p. 13).
Pincerna is globular and sparsely ribbed, Stomacosmethis is triangular and densely ribbed. Stomacosmethisbalingensis is globular, but densely ribbed, forming a connection between the two genera, but the radula morphology unambiguously indicates its position within Stomacosmethis. We retain the two genera separate due to the unique radular morphology of all known Stomacosmethis species. Furthermore, there are many species with typical features of both respective genera. See also under Alycaeus (p. 17).
Protoconch very finely granulated, matte; R1 nearly smooth, with irregular, very fine growth lines, and some fine ribbing near suture, spiral striation not visible; R2 short, darker thicker and somewhat narrower light stripes, but overall surface smooth.
The shell shape is similar to species from the genus Pincerna, but there are no strong ribs in the upper whorls of the shell, the tube is elongated, and not extremely short and piriform. The tube is shorter than usual for the genus Alycaeus, but the shell shape agrees with the other species classified within the genus.
AnimaliaGastropodaCyclophoridaeECC0F7F6-6E2C-583C-AA49-C9227C8AE12FPincernabembex(Benson, 1859)AlycaeusbembexBenson, 1859: 178–179.Alycaeusbembex– Reeve 1878: pl. 5, species 42; Godwin-Austen 1884: pl. 51, fig. 5.Alycaeus (Alycaeus) bembex– Kobelt 1902: 342.Alycaeus (Cycloryx) bembex– Godwin-Austen 1914: 346–347, pl. 147, figs 1, 1a; Gude 1921: 275–276.Cycloryxbembex– Ramakrishna et al. 2010: 70.Type locality.
“in valle Rungun”.
Material examined.
Darjiling, NHMUK 1906.4.4.44 (6 possible syntypes, one of them separated in a different vial and marked with a “T”).
Remarks.
Protoconch rather matte without any notable sculpture; R1 with widely spaced ribs and several, much finer radial lines between the larger ribs; R2 extremely short, with ca. 12 very narrow light stripes, and much thicker darker stripes.
Protoconch matte, without spiral striation; R1 with regular, rather dense, sharp, elevated ribs and occasionally extremely fine spiral striation; R2 extremely short, with four or five ribs only, sculpture of R2 does not differ from that of R1.
AnimaliaGastropodaCyclophoridae5F1BBAE8-5014-516E-B964-CE64BC3E39B3Pincernaconstricta(Benson, 1851)Fig. 39ACyclostomaconstrictumBenson, 1851: 184–195.Alycaeusconstrictusvar.minorBenson, 1859: 181.Alycaeusconstrictus– Reeve 1878: pl. 5, species 41.Alycaeus (Alycaeus) constrictus– Kobelt 1902: 343.Alycaeus (Cycloryx) constrictus– Godwin-Austen 1914: 347–348, pl. 147, figs 4, 4a; Godwin-Austen 1914: 348–349, pl. 154, figs 1, 1a; Gude 1921: 277–278.Alycaeus (Cycloryx) constrictus, var. minor – Godwin-Austen 1914: 348.Cycloryxconstrictus– Ramakrishna et al. 2010: 70.Pincernaconstricta– Páll-Gergely 2017: 218, fig. 1A.Type locality.
Protoconch matte, without spiral lines; R1 with very sharp, widely spaced ribs, and extremely fine spiral lines; R2 very short, with alternating darker/wider and much narrower/lighter stripes (altogether ca. 12), overall surface smooth.
Toruputu Peak, Dafla Hills, Dikrang valley, Dafla, leg. Godwin-Austen, NHMUK 1903.7.1.2596 (18 syntypes).
Remarks.
Protoconch matte, no spiral lines visible; R1 with regular, widely spaced, sharp ribs and extremely fine spiral striation; R2 very short, with ca. five ribs which are similar to those of in R2 but denser.
Tonkin, Phong Tho, leg. Messager, MNHN-IM-2000-31786 (1 syntype); Tonkin, Phong Tho, probably leg. Messager, coll. Staadt, 1969, MNHN-2012-27043 (4 shells, 2 of them typical costulosus, 2 other shells small vanbuensis).
Remarks.
Protoconch matte, no spiral lines visible; R1 strongly, regularly ribbed with hardly visible, extremely fine spiral striation; R2 extremely short, with ca. seven narrow, light stripes, the overall surface is smooth.
Some Indian species originally included in Cycloryx (e.g., P.graphicavar.dihingensis, P.thompsoni, P.burrailensis) are so similar to P.costulosa that it was challenging to find any notable differences. Pincernacostulosa must evidently be classified in Pincerna in the face of the large geographic gap between it and the Indian/Burmese species.
W-Java, Djampang, 2000’, leg. H. Fruhstorfer, 1893, coll. O. Boettger, SMF 57197 (syntype, labelled as holotype [number of available shells was not mentioned in the original description]); Java, Djampong (?), Res.: Preauger-Regentschappen, leg. ??? bauer (label not readable), coll. Oberwimmer, NHMW 111538 (3 shells).
Remarks.
The spire is high, but the protoconch is relatively low, it is rather matte, without spiral lines; R1 with some spiral lines, especially at the end of the region; There are widely spaced, strong ribs on R1 which are bunt at the beginning, but sharp at the end of the region; R2 short, with elevated, sharp, widely spaced ribs with some weak spiral striation between the ribs.
This species fits in the genus Chamalycaeus as well as into Pincerna. It is tentatively placed here due to the elevated spire. Further studies should focus on its systematic relationships.
“Kajo Aro Estate, Mt. Korintji, among fallen leaves, 1450 m alt.”.
Etymology.
The replacement name korintjiensis refers to the type locality (Mt. Korintji).
Remarks.
The holotype and the paratype were not examined by us, but according to the original description the differences between this and the nominotypical subspecies are only minor (more distantly ribbed than the nominotypical subspecies).
Alycaeuscrenilabrislatecostatus van Benthem Jutting, 1959 is a junior primary homonym of Alycaeuslatecostatus Möllendorff, 1882 (treated as Metalycaeus here). Thus, a replacement name (korintjiensis) is proposed to replace the junior homonym.
This subspecies is dedicated to and named after Woutera Sophie Suzanna van Benthem Jutting (1899–1991), who described this taxon under the name Alycaeuscrenilabrislaevis.
Remarks.
The holotype and the three paratypes were not examined by us, but according to the original description, this and the nominotypical subspecies differ only in the sculpture of R3 and the formation of the peristome.
Alycaeuscrenilabrislaevis van Benthem Jutting, 1959 is a primary homonym of Alycaeuslaevis Pilsbry & Y. Hirase, 1909, therefore a replacement name (juttingae) is proposed here.
Protoconch matte, no spiral lines visible; R1 with widely spaced, strong, sharp, regular ribs and much weaker riblets between the main ribs; R2 extremely short, the few (ca. five) ribs are somewhat elevated from the surface.
Shengorh Pk, Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2594 (1 syntype).
Remarks.
Protoconch matte, without spiral striation; R1 with relatively dense, regular, sharp ribs and extremely fine, pitted spiral striation between the ribs; R2 with ca. eight ribs, which are similar to those of R1.
AnimaliaGastropodaCyclophoridae9D1E07B3-918A-54E8-9763-3FA38974C42APincernaglobosa(H. Adams, 1871)AlycaeusglobosusH. Adams, 1871: 794.Alycaeusglobosus– Godwin-Austen 1889: 346–347, pl. 37, figs 3, 3a; E. A. Smith 1895: 116.Alycaeus (Alycaeus) globosus– Kobelt 1902: 345.Type locality.
“Busan, near Sarawak, Borneo”.
Material examined.
Busan Hills, Borneo, “typical” NHMUK 1889.12.7.30–31 (4 shells, possible syntypes); Borneo, ex coll. Fulton, London, NHMW 21003 (2 shells).
Remarks.
Protoconch relatively low in face of the elevated spire, glossy, R1 regularly ribbed with widely spaced, sharp ribs and extremely fine spiral striation; R2 extremely short, with alternating light/very narrow and darker/wider stripes; there are five or six light stripes, and they are very slightly elevated from the surface.
AnimaliaGastropodaCyclophoridae844DC18F-FAFB-58A7-A8C2-8E13994B68E7Pincernaglobosakinabaluana(E. A. Smith, 1895)Alycaeusglobosusvar.kina-baluanaE. A. Smith, 1895: 116.Type locality.
“Kina-Balu, N. Borneo”.
Material examined.
Kina Balu, NHMUK 1893.6.7.104–107 (4 shells, possible syntypes).
Remarks.
Similar to the nominotypical form but has a wider peristome and the end of R1 is almost smooth.
AnimaliaGastropodaCyclophoridae3944BE93-2A95-5733-903E-ABDCDE4C7C9APincernaglobosamuluana(E. A. Smith, 1895)Alycaeusglobosusvar.muluanaE. A. Smith, 1895: 116.Type locality.
“Mulu Mountain”.
Material examined.
Mulu Mt., N. Borneo, leg. A. Everett, NHMUK 1892.2.5.11–14 (4 syntypes).
Remarks.
As in the nominotypical form in the mentioned traits.
AnimaliaGastropodaCyclophoridae6C2A4F01-A8B7-502B-AF66-9789CB4632A6Pincernaglobosapygmaea(E. A. Smith, 1895)Alycaeusglobosusvar.pygmaeaE. A. Smith, 1895: 116.Type locality.
“Mulu Mountain”.
Material examined.
Mulu Mt., N. Borneo, NHMUK 1892.7.20.33 and 1893.6.8.10 (2 syntypes).
Remarks.
Protoconch matte, R1 with strong, regular, widely spaced, sharp ribs without spiral striations; R2 short with ribs that are similar to the ones on R1 in morphology and density.
Smith (1895) advised considering all forms of Alycaeusglobosus under one species because there is “such a general resemblance throughout the series”. Although this is true, the size difference between pygmaea and muluana are extreme. If they were truly collected from the same site, then they should be treated as distinct species. However, the accuracy of the localities is unknown.
AnimaliaGastropodaCyclophoridae431353A1-8CF7-5DDF-8637-2CE194BEAFACPincernaglobosarabongensis(E. A. Smith, 1895)Alycaeusglobosusvar.rabongensisE. A. Smith, 1895: 116.Type locality.
Protoconch without spiral striae; R1 with regular, elevated, sharp ribs and weak spiral striae; R2 very short, with 6–8 darker/thicker stripes and lighter/narrower ones which are relatively well elevated from the surface.
Margarita, near Sadia, Assam, coll. Godwin-Austen, NHMUK 1903.7.1.2511 (1 syntype).
Remarks.
Protoconch matte, R1 with widely spaced, regular, sharp ribs and extremely fine spiral striation; R2 very short, with ca. 3 ribs, which are denser than those on R1.
Protoconch matte, no spiral lines visible; R1 with extremely widely spaced, sharp ribs and extremely fine spiral striation; R2 very short, with few ribs similar to those on R1 in morphology, but denser.
AnimaliaGastropodaCyclophoridae2309DE69-13B4-5BAC-886D-89D120A96B77Pincernagraphica(W. T. Blanford, 1862)AlycaeusgraphicusW. T. Blanford, 1862: 137–138.Alycaeusgraphicus– Reeve 1878: pl. 4, species 34.Alycaeusgraphicusvar. – Theobald 1870: 398, pl. 18, fig. 3.Alycaeus (Dioryx) graphicusvar.minorGodwin-Austen, 1874: 149, pl. 3, fig. 4.Alycaeus (Alycaeus) graphicus– Kobelt 1902: 345.Alycaeus (Cycloryx) graphicus– Godwin-Austen 1914: 361, 419–420, pl. 146, figs 1, 1a, 1b; Gude 1921: 280–281.Alycaeus (Cycloryx) graphicusvar.minor– Godwin-Austen 1914: 403, pl. 144, figs 9, 9a.Cycloryxgraphicus– Ramakrishna et al. 2010: 72.Type locality.
“in montibus Arakanensibus provinciam Burmanam Pegu ad Arakan secernentibus”.
Protoconch glossy, no spiral lines visible; R2 with widely spaced, regular, sharp ribs and without spiral striation; R2 extremely short, consisting of six or seven ribs that are similar to the ones on R1, but are denser.
Protoconch matte, no spiral lines visible; R1 with regular, widely spaced, sharp ribs and extremely fine spiral striation; R2 extremely short, with ca. five narrow, light stripes, which are somewhat elevated from the surface.
Protoconch matte, without spiral striation; R1 with rather low, regular, widely spaced ribs without spiral striation; R2 very short, with ca. six ribs which are similar to those on R1 in morphology but are more densely situated.
Khunho Peak, Naga Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2520 (1 syntype).
Remarks.
Protoconch matte, without spiral striation; R1 with widely spaced, sharp, regular ribs that have extremely fine spiral striation; R2 very short, with ca. five ribs similar to those of R1 in density and height.
Ke-lan-tan, Malay Penins., ex coll. Preston, 1909, ANSP 99391 (1 syntype, photographs examined); Ke-lan-tan, Purchased of Mr. H.B. Preston, NHMUK 1907.5.20.191–192 (2 syntypes).
Remarks.
Protoconch relatively low, no spiral lines visible; R1 with widely spaced, relatively low ribs, with finer spiral striation; R2 moderately short, with strong, irregular ribs and weak spiral striation; outer surface of the operculum rather glossy, multispiral, with an elevated ring on the edge of the nucleus.
Mangat valley, Jiantia Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2518 (8 syntypes).
Remarks.
Protoconch matte, R1 with regular, widely spaced, rather low ribs; no spiral striation observed; R2 very short, with ca. four ribs which are similar to the ones on R1 in terms of density and morphology.
This species was described as A.mangutensis by Godwin-Austen in 1914, and that name was later used as a valid taxon name (Gude 1921; Ramakrishna et al. 2010). However, the name Alycaeus (Dioryx) granumvar.major Godwin-Austen, 1893 for the same taxon was validly introduced, and is available. Thus, the species must be called Pincernamajor (Godwin-Austen, 1893), and the name Alycaeusmangutensis Godwin-Austen, 1914 is a junior objective synonym.
“Maolan Town, Libo County (25°03'N, 108°00'E), Guizhou Province, China”.
Material examined.
Mao-Lan Town, Libo County, Qian-Nan Prefecture, Guizhou Province, China, leg. Luo Tai-Chang, 2001.7.9., IZCAS TM 047081 (holotype).
Remarks.
Protoconch without spiral striae; R1 with strong, widely-spaced ribs and very fine spiral striation; R2 + R3 nearly 0.5 whorl together, approximately the same length; R2 long, without elevated ribs.
Pincerna(?)maolanensis (Luo, Zhang & Zhuo, 2009), holotype (IZCAS TM 047081). White line indicates the length of the sutural tube. Photographs: Kaibaryer Meng.
https://binary.pensoft.net/fig/468686AnimaliaGastropodaCyclophoridaeDDD27FF0-497F-555F-9E8A-46E388CC0551Pincernamargarita(Theobald, 1874)AlycaeusmargaritaTheobald in Hanley & Theobald, 1874: 39, pl. 95, fig. 10.Alycaeusmargaritus– Reeve 1878: pl. 1, species 5.Alycaeus (Alycaeus) margarita– Kobelt 1902: 347.Alycaeus (Cycloryx) margarita– Godwin-Austen 1914: 416.Alycaeusmargarita– Gude 1921: 211–212.Type locality.
Protoconch matte, without spiral striation; R1 with strongly elevated and sharp, regular, widely spaced ribs; R2 extremely short, with ca. five ribs which are denser than those on R1, but similar in terms of morphology.
AnimaliaGastropodaCyclophoridae3287A303-236A-5033-8A02-81A3F9A9BDA2Pincernamouhoti(L. Pfeiffer, 1862)AlycaeusmouhotiL. Pfeiffer, 1862: 275, pl. 36, figs 1, 2.Alycaeusmouhoti– Reeve 1878: pl. 3, species 19; Páll-Gergely et al. 2017: 10; Inkhavilay et al. 2019: 12, figs 4A, B.Alycaeus (Alycaeus) mouhoti– Kobelt 1902: 347.Type locality.
Apex matte, no spiral lines visible; R1 with rather regular, low ribs and extremely fine spiral striation (mostly visible near the suture); R2 moderately long, smooth, narrow lighter and thicker darker stripes alternating.
The tube length is variable within this species (based on Chinese and Vietnamese samples, B. Páll-Gergely, unpublished information); in some specimens its length can reach to a 0.5 whorl, whereas in others it is shorter than a 0.25 whorl.
“Head of the Lanier River, Lahupa Naga Hills, N.E. Manipur”.
Material examined.
Munipur, head of Lanier Rr, leg. Godwin-Austen, NHMUK 1903.7.1.2557 (1 syntype).
Remarks.
Protoconch glossy; R1 with dense, relatively low, but sharp ribs and extremely fine spiral striation; R2 very short, with ca. five ribs similar to those of R1 in terms of morphology and density.
AnimaliaGastropodaCyclophoridae94DBC630-AAED-5885-9222-14D8CD76961CPincernaotiphorus(Benson, 1859)AlycaeusotiphorusBenson, 1859: 178–179.Alycaeusotiphorus– Reeve 1878: pl. 4, species 30.Alycaeus (Alycaeus) otiphorus– Kobelt 1902: 347.Alycaeus (Cycloryx) otiphorus– Godwin-Austen 1914: 349, pl. 147, figs 2, 2a, 2b; Gude 1921: 283.Cycloryxotiphorus– Ramakrishna et al. 2010: 73.Type locality.
“ad Pankabari (1000 ped. alt.) et in valle Rungun (4000 ped.) prope Darjiling Himalayanum”.
Material examined.
No locality data, UMZC I.102555 (1 syntype); Darjiling, “compared with typical sp in Museum Cambridge”, NHMUK 1903.7.1.2565 (1 shell).
Remarks.
The whole shell was weathered, but the smooth protoconch and the remains of regular ribs on R1 are visible; R2 very short, probably smooth.
AnimaliaGastropodaCyclophoridae050910C7-B59C-5346-8283-741D5BD063A8Pincernapaucicostata(Godwin-Austen, 1914)AlycaeuspaucicostatusGodwin-Austen, 1914: 361, pl. 147, figs 5, 5a.Alycaeus (Cycloryx) paucicostatus– Gude 1921: 284.Cycloryxpaucicostatus– Ramakrishna et al. 2010: 73.Type locality.
“Torúpútú Peak, Dafla Hills”.
Material examined.
Toruputu Pk, Dafla Hills, NHMUK 1903.7.1.2595 (2 syntypes).
Remarks.
Protoconch glossy, without spiral lines; R1 with very sharp, elevated ribs and extremely fine, but well visible spiral striation; the density of ribs decreases in an anterior direction; R2 very short, with ca. five ribs, similar to those of the end of R1 but are slightly denser.
Rechila Pk, W. Bhutan, leg. Godwin-Austen, NHMUK 1903.7.1.2573 (12 syntypes).
Remarks.
Protoconch matte, without spiral striation; R1 with dense, weak, irregular ribs and only slightly weaker spiral striation; R2 very short, with ca. five narrow, light stripes, the surface of R2 nearly smooth.
Shan Hills, on same slide as N. 22, leg. Fedden, NHMUK 1906.5.5.87 (holotype [single specimen mentioned in the original description]).
Remarks.
The holotype was strongly weathered. Protoconch without any visible sculpture, R1 and R2 with widely spaced ribs; between the main elevated ribs there are some additional lower ribs.
Mont Lano, Perak, coll. de Morgan, MNHN-IM-2000-31799 (1 syntype); West Malaysia: Selangor: Serendah Forest Reserve, Bukit Takun (400 m), alive on low shrubs, leg. Sow-Yan Chan, Hiong Eng Ng & Leo Nguang 27.10.1996, ex coll. Chan, 1997, SMF 311321 (5 shells; similar to the ones in the original description).
Remarks.
Protoconch very finely granulated, rather glossy; R1 with rather irregular, low, widely spaced ribs and very prominent spiral sculpture, which is visible on the whole shell except for the protoconch; R2 very short, consists of darker and thicker, and narrow and light stripes, which are very slightly elevated from the surface.
Entire shell somewhat weathered; protoconch matte; R1 regularly, densely ribbed, spiral striation not visible, but additional, lower radial lines present between the ribs; R2 very short, ca. five ribs, similar to those on R1 in density.
This species may be conspecific with P.graphicusvar.dihingensis because there is only a slight difference in the density of ribbing (coarser and more oblique in thompsoni). Pincernathompsoni has a completely closed umbilicus, but due to the lack of large shell material we are unable to determine whether this represents individual variability or a stable character. Alternatively, the closed umbilicus may be a feature of more mature (more developed) specimens.
AnimaliaGastropodaCyclophoridaeD8B3E62C-E378-51A2-975A-425355781F39Pincerna(?)vallisZ.-Y. Chen & M. Wu, 2020PincernavallisZ.-Y. Chen & M. Wu, 2020: 42, figs 1, 4A, B, 5A.Type locality.
“China, Hubei, Wufeng Tujiazu Autonomous County, Chaibuxi National Forest Park, 30.216N, 110.199E, 1220 m a.s.l.”.
Material examined.
Photographs of the holotype (HBUMM 10017-spec. 1) and the paratype (HBUMM 10017-spec. 2), were examined.
Remarks.
Protoconch finely granulate, without spiral striation; R1 with regularly spaced strong ribs, R2 very short, consisting of ca. 15 lighter stripes, R3 finely ribbed close to the peristome. No spiral striation mentioned in original description, but some spiral lines visible on R1.
Tonkin, Van-Bu, leg. Dr. R. Bavay, MNHN-IM-2000-31798 (1 syntype).
Remarks.
Shell shape globular; protoconch matte, R1 with regular, low, but clearly visible ribs and extremely fine spiral striation; R2 relatively long (ca. 90°), light slimmer and darker thicker stripes alternate.
“Indonesia, Sumatera Barat (West Sumatra), Solok Selatan (South Solok Regency), Koto Parik Gadang Diateh subdistrict, Goa [=cave] Pinti Kayu, near Sungai [=river] Dareh, approximate GPS position: 1.3027°S, 101.1164°E”.
Material examined.
Holotype (MZB.Gst. 18.970), and several paratypes: see Páll-Gergely (2017).
Remarks.
Protoconch glossy, first 1.75 whorls of R1 with low, relatively dense, regular ribs and weak, dense spiral striation; this sculpture gradually changes to a more sparsely ribbed region, which spans ca. 0.5 a whorl; R2 and tube very short, with eight or nine ribs very narrow, only slightly elevated from the surface; for description of the fine structure of the microtunnels, see original description.
Shell small to very large (D: 3–13 mm), usually brightly coloured yellow or orange; shell shape triangular or depressed triangular; protoconch smooth without spiral striation; R1 usually finely reticulated, without prominent ribs; R2 very short, usually roughly, weakly wrinkled, R3 long, with blunt swelling. Operculum thin, outer surface smooth or with outer, elevated, trumpet-like projection. The outer operculum surface can also be finely granulated, flaky or have short calcareous spikes (Foon and Liew 2017). Central radular tooth elongated, usually with a single central cusp only, or central cusps with one or two small cusp at each side.
Differential diagnosis.
Most species classified in this genus were previously described in Alycaeus. However, Alycaeus species are characterised by a very long R2, whereas the R2 is extremely short is Stomacosmethis. Pincerna species possess a globular shell with more widely spaced ribs, and they are usually not colourful. The distinction of Stomacosmethis and Pincerna requires further examination. See also under Pincerna (p. 147).
Distribution.
This genus is distributed in the tropical forests of the Malay Archipelago (Malay Peninsula, Borneo, Java, Sumatra, Sulawesi).
Remarks.
This genus was originally diagnosed based on the calcareous, pipe, tongue or cup-shaped structure on the outer side of the operculum. Since the circular opercular structures have evolved multiple times, this genus must be re-diagnosed. The type species belongs to a clearly delineable group of the Alycaeidae having triangular, colourful shells, dense, fine ribs and short tube, which is now called Stomacosmethis. See also under S.balingensis, which is exceptional in terms of shell shape.
Protoconch very finely granulated, without spiral lines; R1 with irregular, low ribs and very weak spiral striation in-between, approximately same strength; R2 very short, with darker thick and fewer than ten lighter narrower stripes very slightly elevated from the surface.
This taxon was described as a subspecies of A.perakensis. We agree with Foon and Liew (2017) that it should be handled as a species due to the characteristic flat whorls and expanded peristome.
AnimaliaGastropodaCyclophoridae92CA0759-75AA-5E0D-9364-6B535431B48BStomacosmethisbalingensis(Tomlin, 1948)AlycaeusbalingensisTomlin, 1948: 224–225, pl. 1, fig. 2 (incorrectly cited as fig. 3).Alycaeusbalingensis– Foon and Liew 2017: 21–23, figs 7A, B, 10, 31A.Type locality.
Protoconch matte, without spiral striation; R1 with regular, dense riblets which become sparser anteriorly; R2 extremely short, with low, blunt ribs; R3 with widely spaced, regular, strong but low ribs.
The globular shell shape suggests that this species belongs to Pincerna. However, the dense ribbing would be unusual in Pincerna, and the radular morphology is identical with that of S.perakensis.
AnimaliaGastropodaCyclophoridaeD36413FC-D06F-5524-B53B-865BDF07E770Stomacosmethiscalopoma(E. von Martens, 1908)AlycaeuscalopomaE. von Martens, 1908: 279, pl. 5, fig. 16.Alycaeuscallopoma(sic) – Tielecke 1940: 345.Type locality.
“Gunung Sekerat, Sekuran, an Kalkfelsen”.
Material examined.
Gunung Sekurat, Sangana, Fanso, leg. M. Schmidt, ZMB/MOLL 109931.
Remarks.
We received photographs of the operculum of the type specimen but were unable to examine the shell. This species is classified in Stomacosmethis based on the illustrations in the original description.
Malaysia, Pahang, Bukit Tengkek (nw of Kuantan), Ptani, October 1998, ex coll. Hemmen, coll. PGB (2 paratypes).
Remarks.
Protoconch elevated (although the spire is low), matte, no spiral lines visible; R1 very roughly wrinkled with weak spiral striation; R2 extremely short, with alternating darker/thicker and lighter/narrower stripes; lighter stripes elevated from surface, overall surface roughly wrinkled.
“Thailand, Krabi Province, at km 117.6 of road # 4 (Krabi – Phang Nga)”, 1.5 km on unpaved road, 08°09.574'N, 098°51.761'E”.
Material examined.
Malaysia, Krabi Prov., at km 117.6 off rd. #4 (Krabi to Phang Nga), 1.5 km on unpaved road, 14.09.2000, ex coll. Hemmen, coll. PGB (1 paratype; note that Malaysia is erroneous on the label of this specimen).
Remarks.
Protoconch finely granulated without spiral lines; R1 regularly and finely ribbed with even finer spiral striations; R2 short, with low sharp ribs (cross sectional view is triangular).
AnimaliaGastropodaCyclophoridae0A0701BB-FA2E-5DC2-873B-016A58C27C19Stomacosmethiscongener(E. A. Smith, 1895)Fig. 42AAlycaeus (Orthalycaeus) congenerE. A. Smith, 1895: pl. 3, fig. 26.Alycaeus (Alycaeus) congener– Kobelt 1902: 342.Type locality.
“Mulu and Barit Mountains, Sarawak”.
Material examined.
Barriet Mt, N. Borneo, NHMUK 1892.7.20.30–32 (3 syntypes); Molu, N. Borneo, coll. Dr. A. Oberwimmer, NHMW 71640/O/12201 (1 shell).
Remarks.
Protoconch matte, without spiral lines; R1 with rather irregular ribs and much finer spiral striae; R2 very short, with alternating wider/darker and narrower/lighter stripes; lighter stripes somewhat elevate from the surface.
Shells of Stomacosmethis Bollinger, 1918 species AStomacosmethiscongener (E. A. Smith, 1895), syntype (NHMUK 1892.7.20.30–32) BStomacosmethisdohrni (O. Boettger, 1893), lectotype (SMF 109278). Figures: Barna Páll-Gergely, courtesy Ronald Janssen (B) and Harold Taylor (A).
https://binary.pensoft.net/fig/468687AnimaliaGastropodaCyclophoridaeB080F238-1B1D-5084-9435-A2F7802CB0B7Stomacosmethisdohrni(O. Boettger, 1893)Fig. 42BAlycaeushochstetteri(partim) – E. von Martens, 1867: 152.Alycaeusdohrni– Boettger 1893: 195–196.Alycaeus (Alycaeus) dohrni– Kobelt 1902: 343; Zilch 1957: 146–147, pl. 6, fig. 28.Type locality.
“Borneo”.
Material examined.
Borneo, leg. H. Dohrn 1889, coll. O. Boettger, SMF 109278 (lectotype, designated by Zilch 1957); Same data, SMF 109279 (paralectotype).
Remarks.
Protoconch very finely granulated, without spiral striae; R1 with irregular, rather strong ribs with very slight indication of spiral striation; R2 very short, with alternating darker/thicker and lighter/slimmer stripes; lighter stripes are somewhat elevated from the surface, the entire area is finely wrinkled.
AnimaliaGastropodaCyclophoridae8CA91F61-C068-5FB8-8FA1-DD619FA65108Stomacosmethisfultoni(Möllendorff, 1896)Fig. 43AAlycaeus (Orthalycaeus) fultoniMöllendorff in E. A. Smith, 1895: 117–118, pl. 3, fig. 28.Alycaeus (Alycaeus) fultoni– Kobelt 1902: 343–344; Zilch 1957: 146–147, pl. 6, fig. 29.Type locality.
“prope Gomanton insulæ Borneo”.
Material examined.
N-Borneo, Gomanton, coll. Möllendorff, SMF 109280 (lectotype, designated by Zilch 1957); Same data, SMF 109281 (2 paralectotypes); Borneo, ex coll. Fulton, NHMW 21002 (2 shells).
Remarks.
Protoconch without spiral lines, rather matte; R1 with irregular ribs and very weak spiral lines; R2 very short, with darker/wider and lighter/narrower stripes; lighter stripes are elevated from the surface, the overall surface is seemingly irregularly wrinkled.
Shells of Stomacosmethis Bollinger, 1918 species AStomacosmethisfultoni (Möllendorff, 1896), lectotype (SMF 109280) BStomacosmethisfultonidegenerans (Fulton, 1901), lectotype (SMF 109282). All figures: Barna Páll-Gergely, courtesy Ronald Janssen.
N-Borneo, Gomanton, leg. Fulton, coll. Möllendorff, SMF 109280 (lectotype, designated by Zilch 1957); Same data, SMF 109281 (2 paralectotypes).
Remarks.
Protoconch without spiral lines, rather matte; R1 with rather regular, strong, sharp ribs and very weak spiral lines; R2 very short, with alternating darker/wider and lighter/narrower stripes; lighter stripes are very slightly elevated from the surface.
Type specimens not found in the NHM. Niah-Berge, Borneo, coll. Rušnov ex coll. W. Blume, NHMW 71770/R/20 (2 shells).
Remarks.
Protoconch without spiral lines, matte; R1 with irregular, rather strong ribs and very weak spiral lines; R2 very short, with alternating darker/wider and lighter/narrower stripes; lighter stripes are elevated from the surface, the overall surface is seemingly irregularly wrinkled.
AnimaliaGastropodaCyclophoridaeD79E2F3B-EBF9-5859-B4FF-BD01FEBD5487Stomacosmethishosei(Godwin-Austen, 1889)AlycaeushoseiGodwin-Austen, 1889: 347, pl. 37, fig. 2.Alycaeushosei– E. A. Smith 1895: 117.Alycaeus (Alycaeus) hosei– Kobelt 1902: 346.Type locality.
“Busan Hills”.
Material examined.
Borneo, ex coll. Fulton, NHMW 21004 (3 shells).
Remarks.
Protoconch without spiral lines; R1 with rather regular, weak ribs and some indication of weak spiral striae; R2 very short, with darker wider and lighter slimmer stripes alternating; lighter stripes are slightly elevated from the surface.
AnimaliaGastropodaCyclophoridae618A9F76-EDBB-5934-BF8D-EF084494A29AStomacosmethisjagori(E. von Martens, 1860)AlycaeusjagoriE. von Martens, 1860: 208.AlycaeusHochstetteriL. Pfeiffer, 1861: 215, pl. 3, figs 1–4.Alycaeusjagori– L. Pfeiffer 1861: 215, pl. 3, figs 5–7; E. von Martens 1867: 152–153; Sarasin P. & Sarasin F. 1899: 61–62, pl. 4, figs 46, 46a, pl. 5, fig. 66, pl. 8, fig. 91; Tielecke 1940: 345; Benthem Jutting 1948: 568–570, fig. 25.Alycaeusfugori[sic] – Reeve 1878: pl. 6, species 50.Alycaeusjagorivar.minorE. von Martens, 1867: 152.Alycaeushochstetteri(partim) – E. von Martens 1867: 152.Alycaeus (Alycaeus) jagori– Kobelt 1902: 346; Bollinger 1918: 314; Zilch 1957: 147.Alycaeus (Alycaeus) hochstetteri– Kobelt 1902: 345–346; Bollinger 1918: 314, pl. 11, fig. 7.Type locality.
“Jagor aus Java” (A.jagori); “in montibus Nungnang insulae Javae” (A.hochstetteri); “südl. Celebes, bei Maros unweit Makassar” (A.jagorivar.minor).
Material examined.
Java. Jagor, coll. Möllendorff ex coll. Martens, SMF 109304 (1 syntype of A.jagori, photographs examined); Java. coll. Rušnov ex coll. W. Blume, NHMW/71770/R/21 (18 shells).
Remarks.
Benthem-Jutting (1948) claimed that Alycaeushochstetteri is a synonym of A.jagori. Neither she, nor we had a chance to examine the type specimens of A.hochstetteri, but we follow Benthem-Jutting’s view, which is based on that of Pfeiffer (1861). Hendriks et al. (2019) studied the phylogeography of S.jagori from Borneo. Given that the species is described from Java, the identity of the Bornean samples needs to be revised.
Protoconch glossy, without spiral lines; R1 with irregular, low ribs, very weak spiral striation in-between; R2 very short, with alternating darker/thicker and lighter/narrower stripes somewhat elevated from the surface; overall surface of R2 irregularly wrinkled.
“PHG 77 Bukit Mengapur, Pahang (3°44'42"N, 102°50'16"E)”.
Remarks.
According to the original description this taxon differs from A.charasensis, A.costacrassa, A.selangoriensis, and A.kapayanensis in the formation of the body whorl and in slight differences in the shell sculpture. However, the differences in shell shape are not obviously visible on the photographs in Foon and Liew (2017). Instead, only very slight differences (if any) are to be seen, which we feel do not represent distinctive character traits on a species level. Therefore, we treat this taxon as a subspecies of S.kapayanensis.
“PHG 73 Bukit Charas, Pahang (3°54'35"N, 103°08'48"E)”.
Remarks.
This taxon differs in minor traits (extent of the body whorl and strength of the sculpture) from S.kapayanensis and some other “species” described by Foon and Liew, therefore it is treated as a subspecies of S.kapayanensis.
“Mykarst-065 Batu Balong, Pahang (3°42'41"N, 101°51'25"E)”.
Remarks.
This taxon differs from Alycaeuskapayanensisselangoriensis and Stomacosmethiskapayanensiskapayanensis in characters of the operculum and shell sculpture, formation of the peristome and the body whorl. We believe however, that such characters are of minor taxonomic importance, and therefore, Alycaeuscostacrassa is here considered a subspecies of S.kapayanensis.
Alycaeusikanensis is similar to the other taxa described by Foon and Liew, which are treated here as subspecies of Stomacosmethiskapayanensis. The original description states that A.ikanensis is most similar to A.costacrassa and differs from that species in its smaller and more slender shell. Again, these traits are enough for subspecific distinction, and slight differences in size and shell shape are insufficient for differences at the species level.
Protoconch extremely finely granulated, matte; R1 rather irregularly, finely ribbed with weaker spiral striation; R2 short, with alternating darker/thicker and lighter/slimmer stripes, which are somewhat elevated from the surface, the whole area is rather irregularly sculptured.
Foon and Liew (2017) described six new species from Peninsular Malaysia, which are similar to S.kapayanensis in all the important characters, and show only minor differences, such as fine sculpture (e.g., strength of radial vs. spiral striation), operculum thickness, and shell size and shape. Here we maintain those species (alticola, charasensis, costacrassa, ikanensis, kurauensis, selangoriensis and virgogravida) as subspecies of S.kapayanensis.
“PRK 59 Bukit Batu Kurau, Perak (4°55'45"N, 100°49'02"E)”.
Remarks.
According to the original description, Alycaeuskurauensis differs from Stomacosmethiskapayanensis in “having a wider ultimate whorl, thicker operculum with smooth exterior and more widely spaced radial ribs.” These traits, in our opinion, are sufficient for distinction on subspecific level, but do not justify the distinctness at the species level. Therefore, this taxon is handled as a subspecies of S.kapayanensis.
AnimaliaGastropodaCyclophoridaeD60A0AB4-4CB4-5AA4-9EBA-FD2DE18D0A53Stomacosmethiskapayanensisselangoriensis(Foon & Liew, 2017)AlycaeusselangoriensisFoon & Liew, 2017: 64–66, figs 7U, 27, 31O.Alycaeuskapayanensis– Venmans 1956: 83–84, figs 4, 5 (radula, see Results on radula, p. 12).Type locality.
“SGR 01 Batu Caves, Selangor (3°14'17"N, 101°41'02"E)”.
Remarks.
This taxon differs from S.kapayanensis in the larger shell, more expanded body whorl, thicker operculum, and the formation of the upper palatal section of the peristome. These differences seem to justify differentiation at the subspecific level only.
“Limestone hill at east side of Pulau Dayang Bunting, off Langkawi Island, Kedah (6°12'26"N, 99°47'04"E)”.
Remarks.
This taxon differs from S.kapayanensis and its subspecies in only minor traits (shell and operculum sculpture, formation of whorls). Although described as a species in its own right, we suggest using it as a subspecies of S.kapayanensis.
“Gua Kelam, PRS 64 Wang Ulu, Perlis (6°38'41"N, 100°12'09"E)”.
Remarks.
Alycaeusclementsi is very similar to Stomacosmethiskelantanensis in general shells shape and sculpture. The differences mentioned in the original description (shell size, peristome thickness, outer surface of the operculum, animal colour) are sufficient to distinguish them at the subspecific level only.
Alycaeusexpansus Foon & Liew is most similar to A.clementsi and Stomacosmethiskelantanensis. Foon and Liew (2017) do not explain the differences between A.expansus and the other two taxa, but mention that the characteristic features of this species are the obtuse ultimate whorl, the strongly expanded peristome, the thick operculum and the animal colouration. Based on the photographs in the original description it is difficult to understand what the authors meant by “obtuse ultimate whorl”, because the general shell and body whorl shape do not differ conspicuously from the other two taxa; it was supposed to mean the more rounded-looking ultimate whorl, especially for shells from the type locality (Junn Kitt Foon, pers. comm. 2020 June). Contrary to the original description, the peristome of S.kelantanensis is even more expanded than that of A.expansus. The animal colouration and the operculum thickness are characters which should not be used as distinguishing characters at the species level, especially because thickness of shell or operculum is highly dependent on the environment. Therefore, this taxon is treated as a subspecies of S.kelantanensis.
Alycaeusexpansus Foon & Liew, 2017 is a primary homonym of Alycaeusexpansus Heude, 1890. To our knowledge, the older name has not been used in this combination since 1899, thus no replacement name for Alycaeusexpansus Foon & Liew, 2017 is necessary.
AnimaliaGastropodaCyclophoridae63537C59-1324-5827-8065-C427D80F71C0Stomacosmethiskelantanensis(Sykes, 1902)AlycaeuskelantanenseSykes, 1902: 62, pl. 3, figs 13, 14.Alycaeus (Orthalycaeus) kelantanensis– Möllendorff 1902: 145–146.Alycaeuskelantanensis– Foon and Liew 2017: 49–52, figs 7P, Q, 21, 31K.Type locality.
“Kelantan, Malay Peninsula” (from the title).
Material examined.
Kelantan, NHMUK 20160061 (1 syntype); Kelantan, coll. Möllendorff, SMF 158410 (6 shells); Malaysia, Kelantan, rd. #8 Gua Musang, Kuala Lipis, left side off km19, ex coll. Hemmen, coll. PGB (4 shells; they are similar to the syntype but narrower).
Remarks.
Protoconch spirally striated at its very beginning and this sculpture turns to a rather finely granulated surface until the end of the protoconch (in case of Metalycaeus species the first 0.5–1.0 whorl is usually without spiral lines); R1 rather irregularly ribbed with somewhat weaker spiral striation; R2 very short, with ca. seven low ribs. The SMF 158410 sample contains three shells with prominent spiral structure and another three shells with only very slight indication of spiral lines.
AnimaliaGastropodaCyclophoridae9245C1E2-8595-5B85-A2BE-42524A6DCC8FStomacosmethiskuekenthali(P. Sarasin & F. Sarasin, 1899)Fig. 44AlycaeuskuekenthaliP. Sarasin & F. Sarasin, 1899: 62–63, pl. 4, figs 47, 47a, pl. 5, figs 67a, 68, pl. 8, fig. 92.Alycaeus (Chamalycaeus) kuekenthali– Kobelt 1902: 357–358.Type locality.
“Kalkgrotten von Barabatuwa, nördlich von Maros, Süd-Celebes”.
Material examined.
Selatan Barabatuwa, Insel Sulawesi (=Celebes), in lime cave (Kalkgrotten), leg. Sarasin, Fritz + Paul, 1893–1896, NMB 02266a (lectotype, designated herein, photographs examined); S. Celebes, E.R. Sykes coll. 1954, Acc. 1825, NHMUK (1 shell, agrees with the original description and figure).
Remarks.
Protoconch matte; R1 with irregular, low wrinkles and extremely fine spiral striation on the first 1–1.5 whorls; R2 very short, with alternating lighter/slimmer, and darker/thicker stripes, the lighter stripes slightly elevated from the surface; operculum with a very strongly elevated, expanded, trumpet-like structure.
Lothar Forcart selected a specimen (NMB 02266a) and labelled it as lectotype, but never published this action (Ambros Hänggi, pers. comm. 2020 June). Thus, here we designate the specimen selected by him as the lectotype.
“Thailand, Khao Pratun Cave, an isolated limestone hill in Royong Province at 13°07'19"N, 101°36'03"E, 55 meters elevation”.
Remarks.
We had no opportunity to examine shells of Alycaeusmatchacheepiorum. However, the original description is sufficient for correct generic placement. The protoconch is smooth, R2 very short and the entire teleoconch is finely ribbed.
Protoconch glossy, without spiral lines; R1 with irregular, rather low ribs and very weak spiral striation in-between, but at the end of the region the ribs are more elevated, sharp; R2 very short, with alternating darker/thicker and lighter/narrower stripes which are somewhat elevated from the surface.
Stomacosmethisroebeleni may be a subspecies of A.perakensis although we refrain from treating it as such without further evaluation.
“Am Gumung-Sekerat, nahe Tandjong Kutei, O.-Borneo”.
Material examined.
O. Borneo, Tandjong, Kutei, don.: Prof. C. Schmidt, 1902, NHMB 2412-a (syntype [the number of available shells were not mentioned in the original description], labelled as holotype and also, lectotype, photographs examined).
Remarks.
Protoconch matte, no notable sculpture visible; R1 with rather regular, low ribs and very fine spiral striation; R2 short, with ca. 11 white, low, regular ribs. No operculum was found.
Protoconch very finely granulated, matte; R1 with rather irregular ribs and somewhat weaker spiral striation; R2 very short, with alternating thicker/darker and narrower/lighter stripes, which are slightly elevated from the surface.
Shells of Stomacosmethis Bollinger, 1918 species AS.praetextus (van Benthem Jutting, 1959), paratype (SMF 186532) BS.rimatus (O. Boettger, 1893), lectotype (SMF 109315) CStomacosmethissadongensis (E. A. Smith, 1895), syntype (NHMUK 1893.6.7.73–76). Photographs: Barna Páll-Gergely, courtesy Ronald Janssen (A, B), Harold Taylor (C).
“PHG 02 Gunung Senyum, Pahang (3°41'50"N, 102°26'04"E)”.
Remarks.
Seems to be a “good” species based on the original description, morphologically distinct from all other congeners. The short tube and triangular, coloured (bright yellow) shell fits with the relevant characters for this genus.
N-Borneo, Brunei, leg. O. Staudinger, coll. O. Boettger, SMF 109315 (lectotype, designated by Zilch 1957); Same data, SMF 109316 (2 paralectotypes).
Remarks.
Protoconch very finely granulated, matte; R1 with irregular, strong ribs and much weaker spiral striation; R2 very short, with alternating darker/thicker and lighter/slimmer stripes, which are slightly elevated from the surface.
AnimaliaGastropodaCyclophoridaeB2510A99-9483-5883-8995-7ADEE863E775Stomacosmethisroebeleni(Möllendorff, 1894)AlycaeusroebeleniMöllendorff, 1894: 154, pl. 16, figs 20, 21.Alycaeus (Alycaeus) roebeleni– Kobelt 1902: 350; Zilch 1957: 147, pl. 6, fig. 27.Alycaeusroebeleni– Foon and Liew 2017: 62–64, figs 26, 31W.Type locality.
Protoconch elevated, no spiral lines visible; R1 with rather irregular, fine ribs, and somewhat weaker, dense spiral striae; R2 short, with roughly ten thicker/darker stripes, and much narrower/lighter stripes in-between, which are elevated from the surface.
AnimaliaGastropodaCyclophoridae30CA21A1-4705-59A7-9C68-28B6484D73D4Stomacosmethissadongensis(E. A. Smith, 1895)Fig. 45CAlycaeus (Orthalycaeus) sadongensisE. A. Smith, 1895: 117, pl. 3, fig. 27.Alycaeus (Alycaeus) sadongensis– Kobelt 1902: 350–351.Type locality.
Protoconch elevated, matte, without spiral striation; R1 with dense, regular, rather low riblets and very weak spiral striae; R2 very short, with 6–7 darker/thicker stripes and very narrow/light stripes in-between, the ribs are very slightly elevated from the surface.
“Malawa-Quelle: nördl. des Bowonglangi in S.-Celebes”.
Material examined.
Malawa-Quelle, Celebes, don.: Dr. P. u. F. Sarasin, 1918, NHMB 2411.a (lectotype [designated in Egorov 2013]; photographs examined).
Remarks.
Protoconch matte, without notable sculpture; R1 with fine, regular, low ribs and weaker spiral striation; R2 extremely short, with slightly elevated, white ribs. No operculum was found.
No specimens of Stomacosmethissomnueki were examined by us, but the original description and the published figures are sufficient for generic placement. Protoconch smooth, R2 short, with rather regular, low ribs.
Protoconch rather matte, without spiral lines; R1 with dense, low, rather regular ribs and very weak spiral striation; R2 extremely short, with alternating darker/thicker and lighter/narrower stripes, which are elevated from the surface.
“Indonesia, Sumatra, Aceh Besar, westcoast near Pasi, 10 km S. of Lhong, on limestone slope”.
Remarks.
We had no opportunity to examine the holotype, but the original description and the photographs published therein provide sufficient information regarding the generic status of this species. The protoconch of the weathered holotype is smooth and R2 is short and regularly ribbed. Maassen (2006) questioned the generic status of A.wilhelminae based on its closed umbilicus; further investigation should focus on the value of this character. However, at the moment we see no need to classify it in a genus other than Stomacosmethis.
Nomina nuda
“Alycaeusscepticus Theobald, 1863”
Remarks.
The name Alycaeusscepticus appeared in Theobald (1863: 377), but it was not made available. Blanford (1965: 101) mentioned that it turned out to be a variety of A.ingrami. Thus, Gude (1921) treated this name as a synonym of that species.
“Alycaeusanapetes Panha & Burch, 1998”
Remarks.
The name Alycaeusanapetes Panha & Burch, 1998 appeared in the comparisons section under Alycaeussomnueki (see Panha and Patamakanthin 2001: 189). However, this taxon has never been formally described.
This name was mentioned by Kobelt and Möllendorff (1897: 150) but has never been made available.
“Alycaeusbroti Aldrich, 1889”
Remarks.
See under Chamalycaeuseveretti (Godwin-Austen, 1889).
“Alycaeus (Orthalycaeus) jagorivar.trigonostoma”
Remarks.
Kobelt (1902) mentioned that the taxon Alycaeus (Orthalycaeus) jagorivar.trigonostoma was listed by Kobelt and Möllendorff (1897, Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 29) on page 150, but we did not find that taxon listed there. This name has never made available.
Manuscript names
“Alycaeuspomatiaeformis Ancey”
Remarks.
A sample in the RBINS (Borneo, coll. Ancey, 28.VI.08) was labelled in this manner. The shell had a very wide peristome, and may be a new species.
Chamalycaeussatsumanuskiiensis Kuroda
Remarks.
Museum samples from the Sakurai and Inuba collections in the NSMT bear this name, but was never been formally published (Hanshin Shell Club 1986).
Concluding remarks
After examining specimens of virtually all alycaeid taxa and the relevant literature, here we propose a novel classification of 320 accepted species and 43 subspecies into seven genera. The following conclusions can be made:
(1) While the genera Dicharax, Dioryx, and Metalycaeus have unique character states which serve as diagnostic traits, the other four genera are defined based on a combination of shell characters. Distinctness between Pincerna and Alycaeus and between Pincerna and Stomacosmethis remain to be verified.
(2) The most important achievements of this study are the separation of Stomacosmethis from Alycaeus, and the identification of Dicharax and Metalycaeus species. We also found a common anatomical trait of the Alycaeidae (bursa copulatrix originates from the lateral side of the ovarium) besides the unique sutural tube-microtunnel system, which is probably a device for gas exchange.
(3) The species-level diversity (i.e., number of described species) is largely dependent on whether the splitting or lumping approaches were employed by the taxonomists who classified specimens from particular areas, which resulted in peculiar geographic differences in species diversity.
(4) Various lines of evidence suggest the presence of three subgroups within the Alycaeidae (Alycaeus-Dioryx, Chamalycaeus-Dicharax-Metalycaeus, Pincerna-Stomacosmethis), which can be tested by future studies.
Relationships between genera
Morphological traits (shell, genital system, radula) presented here and unpublished molecular phylogenetic pattern suggest the presence of three groups within Alycaeidae. Firstly, the genera Alycaeus and Dioryx form a distinct group based on the blunt central cusp of radula and the position of the bursa copulatrix that starts posterior to the middle section of the ovarium. Their similarly rather large shell sizes (at least for many Dioryx species and all Alycaeus species) and overlapping distribution suggest these two genera might be closely related. Secondly, the genera Chamalycaeus, Dicharax, and Metalycaeus generally possess the depressed shell and have a plesiomorphic radula type with a central, round tooth that has pointed cusps. The bursa copulatrix does not extend beyond the ovarium, which can be a synapomorphic characteristic for this group. Thirdly, Pincerna and Stomacosmethis are both characterised by having a very short tube and the elevated spire and inhabit a largely overlapping geographical area. However, the genital traits of the single known Pincerna and Stomacosmethis species are strikingly different, and the radula of Stomacosmethis with an elongated central tooth and few cusps is unique among Alycaeidae. Thus, dependable grouping of the recognised alycaeid genera is not yet possible because the morphology of radula and genital system, and DNA sequences have been available only in a limited number of alycaeid species.
Molecular phylogeny could be a powerful tool to obtain testable hypotheses of evolutionary relationships between species and/or higher taxa. In our attempts, direct PCR amplification of DNA extracted from Alycaeidae snails was not sufficiently successful as that from other families (Junn Kitt Foon, pers. comm.), although few alycaeid taxa have been included in preceding studies (Webster et al. 2012; Hendricks et al. 2019). Conventionally used primers may need to be specifically modified for several key alycaeids to resolve the generic phylogeny of this family. Our analysis of generic relationships including the outcome of molecular phylogeny without Dioryx suggests that the genus Alycaeus is distant from Stomacosmethis and Pincerna, which are closely related and may not be mutually monophyletic. Chamalycaeus, Metalycaeus and Dicharax are also close relatives of one another within the family Alycaeidae (Páll-Gergely et al. in prep.).
Biogeography and fossil records
The Alycaeidae is a characteristic family of the Oriental Biogeographic Region, with a few genera expanding their distributions to the Sino-Japanese/Palaearctic realms (see Holt et al. 2013) (Fig. 46). Isolated occurrence of alycaeids in the Western Ghats supports a hypothesis that the fauna of that region is mostly of Southeast Asian origin (Raheem et al. 2014, Fred Naggs, pers. comm.).
There are three areas that can be considered as biodiversity centres for this family, namely the southeastern Himalaya, northern Vietnam/southern China, and peninsular Malaysia/Sumatra (Table 5), each inhabited by five genera. The diversity in terms of the number of genera decreases towards the periphery of these areas, e.g., southwestern Himalaya (1 genus), southern India (1 genus), Japan (2 genera), and the Philippines (2 genera). The current classification suggests that alycaeids expanded their distributions to Japan and the Philippines in two independent events (Páll-Gergely and Auffenberg 2019). Budha et al. (2015) (and also Kuznetsov 1996 and Kuznetsov and Schileyko 1997) reported several alycaeid species from Nepal, but these were not examined by us. Identification of alycaeids is only possible by careful comparison with the type material, which was not done in these studies. Thus, we did not include their identification data in Nepal.
Distribution of all alycaeid genera showing regions with relatively high generic diversity.
https://binary.pensoft.net/fig/468691
Distribution of genera in the three regions with high diversity. Double asterisks indicate relatively high taxon diversity of that given genus.
Peninsular Malaysia and Sumatra
Northern Vietnam, northern Laos and southern China
Southeastern Himalaya
Alycaeus
*
*
Chamalycaeus
*
*
Dicharax
*
**
**
Dioryx
**
*
Metalycaeus
*
*
Pincerna
**
*
**
Stomacosmethis
**
Although Japan is not rich in the number of genera, unique features of conchology are found in a few of the Japanese species, such as the extremely expanded aperture of D.expanstoma, detached initial whorls of D.miyazakii, and uniquely shaped peristome with weak parietal callus in D.okamurai.
The genus Pincerna has a conspicuously disjunct distribution. One set of species inhabits the Himalaya, northern Thailand, northern Laos, northern Vietnam, and southern China, whereas the other inhabits peninsular Malaysia, Sumatra, and Borneo.
The only fossil alycaeid is 23–21 million years old Dicharax(?)sonlaensis (see Raheem and Schneider 2018) from northern Vietnam, which fits the range of morphological variation of an extant Dicharax species of the same region. In recent years several terrestrial caenogastropods were described from 99 million year-old Burmese ambers, and there is more to come. All cyclophoroidean families of Southeast Asia (Cyclophoridae, Diplommatinidae, Pupinidae) have been found from mid-Cretaceous Burmese amber (Hirano et al. 2019; Neubauer et al. 2019) with the exception of the Alycaeidae. The absence of alycaeids in Burmese amber may suggest that this is a younger group than the other related families, which is in agreement with the molecular dating of Hirano et al. (2019). Nevertheless, special attention should be paid on alycaeid-like fossil shells since they would not be recognised as alycaeids without the sutural tube and the R2 region that is differently ribbed than the other parts of the body whorl.
Acknowledgements
We are very grateful to all colleagues who in various ways contributed to our revision: Zoltán Fehér (HNHM), Anita Eschner (NHMW), Tom White (NHM), Sigrid Hof and Ronald Janssen (SMF), Kazunori Hasegawa (NSMT), Thierry Backeljau (RBINS), and Philippe Bouchet and Virginie Héros (MNHN) granted access to their museum collections; Christine Zorn (ZMB), Ambros Hänggi (NHMB), John Slapcinsky (UF), Taehwan Lee (UMMZ), Richard Preece (UMZC), Adam J. Baldinger (MCZ), Lily Berniker (AMNH), Paul Callomon (ANSP), Harold Taylor (NHM), Kevin Webb (NHM), Bram van der Bijl (RMNH), Hiroshi Minato, Junn Kitt Foon, András Hunyadi, Philippe Bouchet, Yansen Chen, Takahiro Hirano, Jamen U. Otani, Kanjo Okubo, Kenjo Ohara, and Christa Hemmen all sent information, specimens or photographs of museum specimens; J.U. Otani and Hukaya Sitosi translated Japanese literature and label data; Ruud Bank corrected publication dates. We are also grateful to three reviewers (Junn Kitt Foon, Chirasak Sutcharit, Bernhard Hausdorf) for reviewing this manuscript and to the handling editor, Frank Köhler for his useful comments and professional editorial work.
This study was supported by the MTA (Hungarian Academy of Sciences) Premium Post Doctorate Research Program and the SYNTHESYS Project (GB-TAF-2523) for BP-G, and by Grants-in-Aid for Scientific Research (KAKENHI) from Japan Society for the Promotion of Science to TA. SS and BT are grateful to the Director of the Zoological Survey of India, Kolkata for providing the necessary facilities for this study. We are indebted to The Biodiversity Heritage Library for the multitude of rare literature made available to us (www.biodiversitylibrary.org).
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Figure S1. Tukik, Central Aceh, specimen 1 (HNHM 104858)
image file
https://binary.pensoft.net/file/468692This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Barna Páll-Gergely