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Monograph
Genus-level revision of the Alycaeidae (Gastropoda, Cyclophoroidea), with an annotated species catalogue
expand article infoBarna Páll-Gergely, Sheikh Sajan§|, Basudev Tripathy§, Kaibaryer Meng, Takahiro Asami#, Jonathan D. Ablett¤
‡ Centre for Agricultural Research, Budapest, Hungary
§ Zoological Survey of India, Kolkata, India
| Wildlife Institute of India, Dehradun, India
¶ Institute of Zoology, Chinese Academy of Sciences, Beijing, China
# Shinshu University, Matsumoto, Japan
¤ Natural History Museum, London, United Kingdom
Open Access

Abstract

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: Dioryx urnula niosiensis Páll-Gergely, nom. nov. for Alycaeus urnula var. daflaensis Godwin-Austen, 1914; Dioryx urnula rotundus Páll-Gergely, nom. nov. for Alycaeus urnula var. globosus Godwin-Austen, 1914; Pincerna crenilabris juttingae Páll-Gergely, nom. nov. for Alycaeus crenilabris laevis van Benthem Jutting, 1959; Pincerna crenilabris korintjiensis Páll-Gergely, nom. nov. for Alycaeus crenilabris latecostatus van Benthem Jutting, 1959; Dicharax conicus jatingaensis Páll-Gergely, nom. nov. for Alycaeus conicus var. nanus Godwin-Austen, 1914; Metalycaeus godwinausteni Páll-Gergely, nom. nov. for Alycaeus neglectus Godwin-Austen, 1914; and finally Metalycaeus suhajdai Páll-Gergely, nom. nov. for Alycaeus varius Godwin-Austen, 1914.

Keywords

land snail, museum collections, systematics, taxonomy

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 Alycaeus jousseaumei 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 Alycaeus gibbus 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).

Specimens used for anatomical study

Alycaeus eydouxi Venmans, 1956: Vietnam, Marble Mountain, Da Nang, coll. No. V142, NHM 2008 VN expedition, 26.05.2008, NHMUK 20160702.

Alycaeus gibbosulus 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.

Dioryx messageri (Bavay & Dautzenberg, 1900): Vietnam, Ninh Bình, Cuc Phuong National Park (site 3), 20°18.141'N, 105°39.240'E, NHM 2013 VN expedition, 09.09.2013, NHMUK 20140343.

Dicharax tokunoshimanus principalis (Pilsbry & Y. Hirase, 1909): Japan, Kagoshima Prefecture, Amamioshima, Akina, southern edge of the village, 28°26.623'N, 129°33.381'E, 15 m, leg. Hunyadi, A., Miyai, T., Nakahara, Y., Otani, J.U., Páll-Gergely, B. & Yano, Sh., 01.10.2016, 2016.10.01B, spec2, HNHM 104428.

Metalycaeus minatoi Páll-Gergely, 2017: 20151214A, Japan, Kagoshima Pref., Tanegashima, Kumage-gun, Minamitane-chō, Kukinaga hōmanjinja, 30°23.051'N, 130°56.108'E, leg. Nakahara, Y., Otani, J.U. & Páll-Gergely, B., 14.12.2015, HNHM 104427.

Stomacosmethis dohrni (O. Boettger, 1893): Indonesia, Kalimantan Selatan, Beramban, leg. Yansen Chen, Apr 2012, HNHM 104426.

Stomacosmethis balingensis (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.

Specimens used for examining the radula

Alycaeus eydouxi: Vietnam, Marble Mountain, Da Nang, coll. No. V142, NHM 2008 VN expedition, 26.05.2008, NHMUK 20160702.

Alycaeus gibbosulus 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.

Dioryx messageri (Bavay & Dautzenberg, 1900): Vietnam, Hòa Bình Province, 20°21.329'N, 105°38.005'E, NHM 2013 VN expedition, NHMUK 20140281.

Chamalycaeus sp.: Indonesia, Tukik River, Central Aceh, GPS n.a., leg. Yansen Chen, (specimen1, ALY17 in molecular study), HNHM 104858.

Chamalycaeus everetti (Godwin-Austen, 1889): Indonesia, Kalimantan Selatan, Beramban, leg. Yansen Chen, Apr 2012, HNHM 104859.

Dicharax itonis (Kuroda, 1943): Japan, Hiroshima, Mihai city, Kui-cho, ex coll. K. Ohara, 24.10.2015, HNHM 104860.

Dicharax (?) okinawaensis (Uozumi, Yamamoto & Habe, 1979): Japan, Okinawa, Ogimi, Mt. Nekumachiji, 26°40.977'N, 128°8.332'E, 304 m, leg. Hirano, T. 09.09.2015, HNHM 104431.

Stomacosmethis balingensis (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.

Stomacosmethis perakensis (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

AMNH American Museum of Natural History (New York, USA);

ANSP Academy 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);

HNHM Hungarian Natural History Musem (Budapest, Hungary);

IZCAS National Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences (Beijing, China);

MCZ Museum of Comparative Zoology (Massachusetts, USA);

MNHN Muséum National d’Histoire Naturelle (Paris, France);

MZB Museum Zoologicum Bogoriense (Bogor, Indonesia);

NC Nishinomiya Shell Museum (Hyogo, Japan);

NHMB Naturhistorisches Museum, Basel (Basel, Switzerland);

NHM The Natural History Museum (London, UK);

NHMUK When citing lots deposited in the NHM;

NHMW Museum of Natural History of Vienna (Vienna, Austria);

NZSI National Zoological Collection of the Zoological Survey of India (when cited specimens deposited in the ZSI);

NSMT National Museum of Nature and Science, Tsukuba, Japan;

PGB Collection Barna Páll-Gergely (Mosonmagyaróvár, Hungary);

RBINS Royal Belgian Institute of Natural Sciences (Brussels, Belgium);

RMNH National Museum of Natural History Naturalis (Leiden, The Netherlands);

SMF Senckenberg Forschungsinstitut und Naturmuseum (Frankfurt am Main, Germany);

UF Florida Museum of Natural History, University of Florida (USA);

UMMZ University of Michigan, Museum of Zoology (Ann Arbor, USA);

UMZC University Museum of Zoology (Cambridge, United Kingdom);

USNM Smithsonian National Museum of Natural History (Washington, USA);

ZMB Museum für Naturkunde (Berlin, Germany);

ZSI Zoological Survey of India (Kolkata, India).

Description and assessment of morphological characters

Shell morphology

Protoconch 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 (Alycaeus conformis 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).

Figure 1. 

Traits of alycaeid protoconch A, B smooth: Dicharax cristatus (Möllendorff, 1886) (2010.05.08A, coll. PGB) C, D pitted. C: Chamalycaeus sculptilis (Benson, 1856) (NHMW 71770/R/17); D: Dicharax itonis (Kuroda, 1943) (NSMT 78866) E, F spirally striated: Metalycaeus muciferus (Heude, 1885) (2013/7, coll. PGB). All images: Barna Páll-Gergely.

Figure 2. 

Protoconch sculpture of Alycaeus Gray, 1850 species A Alycaeus eydouxi Venmans, 1956 (Cochinchina, coll. V.W. MacAndrew, NHMUK) B Alycaeus rolfbrandti Maassen, 2006 (MNHN-IM-2012-27321) C Alycaeus conformis Fulton, 1902 (HNHM 99714, spec3). Scale bars: 1 mm. All images: Barna Páll-Gergely.

Protoconch shape

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, Metalycaeus vinctus 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 Dicharax moellendorffi 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 Chamalycaeus microconus, C. mixtus, Dicharax akioi, and Alycaeus conformis, 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 (Dicharax bison, Metalycaeus nipponensis). 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.

Alycaeus eydouxi 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).

Alycaeus gibbosulus 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).

Figure 3. 

Female genital anatomy of Alycaeidae A positioning of females during anatomical examination B, C Alycaeus eydouxi Venmans, 1956 (NHMUK 20160702, V142, specimen5) D, E Alycaeus gibbosulus Stoliczka, 1872 (ALY03) F, G Dioryx messageri (Bavay & Dautzenberg, 1900), NHMUK 20140343 H, I Dicharax tokunoshimanus principalis (Pilsbry & Y. Hirase, 1909) HNHM 104428 (2016.10.01B, spec2) J, K Metalycaeus minatoi Páll-Gergely, 2017 HNHM 104427 (2015.12.14A, female28) L, M Stomacosmethis dohrni (O. Boettger, 1893) HNHM 104426 N, O Stomacosmethis balingensis (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.

Dioryx messageri (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).

Dicharax tokunoshimanus principalis (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).

Metalycaeus minatoi 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).

Stomacosmethis dohrni (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).

Stomacosmethis balingensis (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: Alycaeus eydouxi Venmans, 1956 (Fig. 4A), Alycaeus gibbosulus Stoliczka, 1872 (Fig. 4B), Dioryx messageri (Bavay & Dautzenberg, 1900) (Fig. 4C), Chamalycaeus sp. (Fig. 4D, Suppl. material 1: Fig. S1), Chamalycaeus everetti (Godwin-Austen, 1889) (Fig. 4E), Dicharax itonis (Kuroda, 1943) (Fig. 4F), Dicharax (?) okinawaensis (Uozumi, Yamamoto & Habe, 1979) (Fig. 4G), Stomacosmethis balingensis (Tomlin, 1948) (Fig. 4H), Stomacosmethis perakensis (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.

Figure 4. 

Radulae of Alycaeidae A Alycaeus eydouxi Venmans, 1956 NHMUK 20160702 B Alycaeus gibbosulus Stoliczka, 1872, HNHM 104857 C Dioryx messageri (Bavay & Dautzenberg, 1900) NHMUK 2014.02.81 D Chamalycaeus sp., HNHM 104858 E Chamalycaeus everetti (Godwin-Austen, 1889), HNHM 104859 F Dicharax itonis (Kuroda, 1943) HNHM 104860 G Dicharax okinawaensis (Uozumi, Yamamoto & Habe, 1979), HNHM 104431 H Stomacosmethis balingensis (Tomlin, 1948) HNHM 104861 I Stomacosmethis perakensis (Crosse, 1879) HNHM 104430. Scale bars: 50 μm. All images: Barna Páll-Gergely.

Table 1.

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
Alycaeus conformis see remarks Venmans 1956
Alycaeus eydouxi see remarks Venmans 1956
Alycaeus eydouxi 5 cusps, broad, central cusp blunt this study
Alycaeus gibbosulus 5 cusps, broad, central cusp blunt this study
Chamalycaeus everetti 5 cusps, broad, central cusp pointed this study
Dicharax alticola 5 cusps, broad, central cusp pointed Páll-Gergely et al. 2017
Dicharax ananensis 5 cusps, broad, central cusp pointed Yano et al. 2013
Dicharax bicrenatus 5 cusps, broad, central cusp pointed Godwin-Austen 1884 (1882–1920)
Dicharax cristatus 5 cusps, broad, central cusp pointed Páll-Gergely et al. 2017
Dicharax depressus 5 cusps, broad, central cusp pointed Páll-Gergely et al. 2017
Dicharax fimbriatus 5 cusps, broad, central cusp pointed Páll-Gergely et al. 2017
Dicharax immaculatus 5 cusps, broad, central cusp pointed Páll-Gergely et al. 2017
Dicharax itonis 5 cusps, broad, central cusp pointed this study
Dicharax longituba 5 cusps, broad, central cusp pointed Benthem-Jutting 1948
Dicharax okinawaensis 5 cusps, broad, central cusp pointed this study
Dicharax planorbulus 5 cusps, broad, central cusp pointed Heude 18821890
Pincerna maolanensis 7 cusps, broad, central cusp pointed Luo et al. 2009
Dioryx messageri 7 cusps, broad, central cusp blunt this study
Dioryx setchuanensis 5 cusps, broad, central cusp blunt Heude 18821890
Metalycaeus minatoi 5 cusps, broad, central cusp pointed Páll-Gergely & Asami, 2017
Metalycaeus vinctus 5 cusps, broad, central cusp pointed Páll-Gergely & Asami, 2017
Metalycaeus zayuensis 5 cusps, broad, central cusp pointed Zhang et al. 2008
Pincerna thieroti see remarks Venmans 1956
Pincerna yanseni 5 cusps, broad, central cusp pointed Páll-Gergely 2017
Stomacosmethis balingensis 3 cusps, elongated, central cusp pointed this study
Stomacosmethis hochstetteri (synonym of jagori) 3 cusps, elongated, central cusp pointed Bollinger 1918
Stomacosmethis jagori 3 cusps, elongated, central cusp pointed Thiele 1929
Stomacosmethis jagori 1 cusp, elongated, central cusp pointed Sarasin & Sarasin, 1899
Stomacosmethis kapayanensis selangoriensis see remarks Venmans 1956
Stomacosmethis kuekenthali 1 cusp, elongated, central cusp pointed Sarasin & Sarasin, 1899
Stomacosmethis perakensis 3 cusps elongated, central cusp pointed this study
Stomacosmethis porcilliferus 3 cusps elongated, central cusp pointed Bollinger 1918
Stomacosmethis sarasinorum 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 Alycaeus eydouxi and S. kapayanensis (specimens were collected as Batu Caves, which is the type locality of S. kapayanensis selangoriensis, see Foon and Liew 2017) are especially interesting. Venmans (1956) published drawings of the radula of a specimen of Alycaeus eydouxi 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.

Table 2.

Genus-group taxa of the Alycaeidae. Abbreviations: M: monotypy, OD: original designation, SD: subsequent designation. Valid genera are marked with an asterisk.

Genus Type species Mode of designation Remarks
*Alycaeus Gray, 1850 Cyclostoma gibbum Eydoux, 1838 (= Alycaeus eydouxi Venmans, 1956) SD accepted
Awalycaeus Kuroda, 1951 Awalycaeus abei Kuroda, 1951 M synonym of Dicharax
*Chamalycaeus Möllendorff 1897 Alycaeus (Chamalycaeus) fruhstorferi Möllendorff, 1897 M accepted
Charax Benson, 1859 Alycaeus hebes Benson, 1857 SD accepted name is Dicharax
Cipangocharax Kuroda, 1943 Alycaeus biexcisus Pilsbry, 1902 M synonym of Dicharax
Cycloryx Godwin-Austen, 1914 Cyclostoma constrictum Benson, 1851 OD synonym of Pincerna
*Dicharax Kobelt & Möllendorff, 1900 Alycaeus hebes Benson, 1857 SD accepted
*Dioryx Benson, 1859 Alycaeus amphora Benson, 1856 SD accepted
*Metalycaeus Pilsbry, 1900 Alycaeus (Metalycaeus) melanopoma Pilsbry, 1900 SD accepted
Orthalycaeus L. Pfeiffer, 1876 Cyclostoma gibbum Eydoux, 1838 (= Alycaeus eydouxi Venmans, 1956) SD synonym of Alycaeus
*Pincerna Preston, 1907 Alycaeus (Pincerna) liratula Preston, 1907 M accepted
Raptomphalus Godwin-Austen, 1914 Alycaeus (Raptomphalus) magnificus Godwin-Austen, 1914 M synonym of Metalycaeus
Sigmacharax Kuroda, 1943 Chamalycaeus (Sigmacharax) itonis Kuroda, 1943 M synonym of Dicharax
*Stomacosmethis Bollinger, 1918 Alycaeus (Stomacosmethis) sarasinorum Bollinger, 1918 SD accepted
Table 3.

Important traits of Alycaeid genera.

Genus Protoconch sculpture Tube (R2) length Shell diameter (mm) Central tooth Key trait Unclear relationship with
Alycaeus smooth to obliquely striated very long (ca. 1/2 whorl) 8–15 5 cusps, broad, central cusp blunt shell very large, R2 ca. half whorl long Pincerna
Chamalycaeus smooth, usually elevated variable 2–5 5 cusps, broad, central cusp pointed 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 Alycaeus conformis 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 (Pincerna yanseni), 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 Pincerna liratula 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). Alycaeus mouhoti has a smoother and larger shell with the simple (not double) peristome than otherwise very similar A. vanbuensis and is also similar to Alycaeus eydouxi (type species of Alycaeus) possessing commonly large and similarly shaped shell with a rather long tube. Alycaeus mouhoti 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. Pincerna crenilabris 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.

Figure 5. 

Relationships between alycaeid genera. Solid lines indicate partly unclear generic borders. See explanation in the text.

Annotated 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.

Table 4.

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 Dioryx urnula niosiensis 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
Pincerna latecostata 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. granum var. 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 Alycaeus daflaensis 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 pilsbryi Kobelt 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 Alycaeus varius 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 Dicharax cristatus, D. fimbriatus or Metalycaeus muciferus (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.

Alycaeidae W. T. Blanford, 1864

Alycaeinae W.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.

Alycaeus Gray, 1850

Alycaeus Gray, 1850: 27.

Orthalycaeus L. Pfeiffer, 1876: 57 (partim).

Alycaeus (Alycaeus)Thiele 1929: 108; Wenz 1938: 478; Egorov 2013: 33.

Type species

Cyclostoma gibbum Eydoux, 1838 (= Alycaeus eydouxi Venmans, 1956)

(Fig. 6A), SD Nevill (1878: 290). Cyclostoma gibbum Eydoux, 1838, is a junior homonym of Cyclostoma gibbum Draparnaud, 1805. Thus, Venmans (1856) proposed Alycaeus eydouxi 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.

Figure 6. 

Type species of alycaeid genus-group taxa A Alycaeus eydouxi Venmans, 1956 (SMF 109290; type species of Alycaeus) B Chamalycaeus fruhstorferi (Möllendorff, 1897), lectotype (SMF 109481; type species of Chamalycaeus). Photographs: Barna Páll-Gergely (A) and Frank Walther (B).

Diagnosis

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).

Figure 7. 

Distribution of Alycaeus Gray, 1850 (dark shaded area) and Dioryx Benson, 1859 (light shaded area).

Alycaeus conformis Fulton, 1902

Fig. 2C

Alycaeus conformis Fulton, 1902: 68–69.

Alycaeus conformisVenmans 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.

Alycaeus conformis 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.

Alycaeus eydouxi Venmans, 1956

Figs 2A, 6A, 8

Cyclostoma gibbum Eydoux, 1838: 6, pl. 117, fig 1. (non Cyclostoma gibbum Draparnaud, 1805)

Alycaeus gibbusReeve 1878: pl. 1, species 3.

Alycaeus (Alycaeus) gibbusKobelt 1902: 344–345.

Alycaeus (Orthalycaeus) gibbus – Godwin-Austen 1914: 427, pl. 156, figs 5, 5a.

Alycaeus eydouxi Venmans, 1956: 87, figs 6, 7 (radula). (nom. nov. pro Cyclostoma gibbum Eydoux, 1838, non Cyclostoma gibbum Draparnaud, 1805)

Alycaeus eydouxiEgorov 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”.

Material examined

Annam, Touranne, leg. Frühstorfer, coll. Möllendorff, SMF 109290 (5 shells); Same data, NHMW 43182 (4 shells); Cochinchina, coll. V.W. MacAndrew, NHMUK.

Remarks

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.

Figure 8. 

Living specimens of Alycaeus eydouxi 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.

Alycaeus gibbosulus Stoliczka, 1872

Alycaeus gibbosulus Stoliczka, 1872: 268–269, pl. 10, fig. 14.

Alycaeus chaperi de Morgan, 1885a: 70.

Alycaeus (Orthalycaeus) gibbosulusMöllendorff 1891: 342.

Alycaeus (Alycaeus) gibbosulusKobelt 1902: 344.

Alycaeus gibbosulusBerry 1963: 17, pl. 4, fig. 25; Foon and Liew 2017: 38–41, figs 7F, G, 17, 31E.

Dioryx pyramidalisHabe 1965: 111–112, pl. 2, figs 3, 4.

Type locality

“Penang island” (from the title).

Material examined

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 Alycaeus gibbosulus.

Alycaeus jousseaumei de Morgan, 1885

Alycaeus jousseaumei de Morgan, 1885b: 402, pl. 8, fig. 4.

Alycaeus jousseaumi [sic] – Möllendorff 1891: 343.

Alycaeus (Chamalycaeus) jousseaumeiKobelt 1902: 357.

Chamalycaeus jousseaumeiBerry 1963: 17, pl. 4, fig. 25.

Alycaeus jousseaumeiEgorov 2013: 33, fig. 58d; Foon and Liew 2017: 43–46, figs 7H, I, 19, 31F.

Type locality

“sur le mont Lano, pres de Campong Kapayan”.

Material examined

Mont Lano, Perak, MNHN-IM-2000-31800 (1 syntype), MNHN-IM-2000-31800 (4 syntypes); Perak, NHMW 41001 (1 shell).

Remarks

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.

Alycaeus pyramidalis Benson, 1856

Alycaeus pyramidalis Benson, 1856: 225.

Alycaeus pyramidalisReeve 1878: pl. 1, species 6; Godwin-Austen 1914: 427, pl. 156, figs 6, 6a; Gude 1921: 216.

Alycaeus (Alycaeus) pyramidalisKobelt 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 Alycaeus gibbosulus.

Alycaeus rolfbrandti Maassen, 2006

Fig. 2B

Alycaeus rolfbrandti Maassen, 2006: 136–137, figs 6–9.

Alycaeus rolfbrandtiPáll-Gergely et al. 2017: 10, fig. 3B; Inkhavilay et al. 2019: 13, fig. 3F.

Type locality

“Laos, limestone Hills 20 km E of Takek”.

Material examined

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.

Alycaeus somwangi Dumrongrojwattana & Maassen, 2008

Alycaeus somwangi Dumrongrojwattana & Maassen, 2008: 1–3, figs 1–6.

Type locality

“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 Alycaeus somwangi, but the original description provides enough information to allow for generic placement. Protoconch without spiral striae, R2 very long, with regular, low ribs.

Chamalycaeus Möllendorff, 1897

Alycaeus (Chamalycaeus) Möllendorff, 1897b: 93.

Chamalycaeus Kobelt and Möllendorff 1897: 148; Páll-Gergely et al. 2017: 5–7.

Chamalycaeus (Chamalycaeus)Thiele 1929: 107–108; Wenz 1938: 477–478; Egorov 2013: 35.

Type species

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).

Figure 9. 

Distribution of Chamalycaeus Möllendorff, 1897 (dark shaded area) and Metalycaeus Pilsbry, 1900 (light shaded area).

Remarks

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 Alycaeus andamaniae 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 Alycaeus andamaniae (incorrectly selected as the type species for Chamalycaeus). No detailed description of Alycaeus andamaniae has ever been published, and our revision suggests that most authors who described Chamalycaeus species did not examine samples of Alycaeus andamaniae. 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.

Chamalycaeus andamaniae (Benson, 1861)

Alycaeus andamaniae Benson, 1861: 28–29.

Alycaeus andamaniaeReeve 1878: pl. 2, species 10; Godwin-Austen, 1914: 430–431.

Alycaeus (Chamalycaeus) andamaniaeKobelt 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.

Chamalycaeus (?) armillatus (Benson, 1856)

Alycaeus armillatus Benson, 1856: 227.

Alycaeus armillatusReeve 1878: pl. 5, species 38; Godwin-Austen 1914: 406, pl. 151, figs 3, 3a.

Alycaeus (Dicharax) armillatusKobelt 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.

Chamalycaeus busbyi (Godwin-Austen, 1893)

Alycaeus busbyi Godwin-Austen, 1893: 595.

Alycaeus busbyi – Godwin-Austen 1897: 5, pl. 63, figs 1, 1a, b; Godwin-Austen 1914: 431.

Alycaeus (Chamalycaeus) busbyiKobelt 1902: 353; Gude 1921: 225.

Chamalycaeus busbyiSubba Rao and Mitra 1991: 26–27, pl. 3, fig. 3.

Chamalycaeus (Chamalycaeus) busbyi – Ramakrishna et al. 2010: 52.

Type locality

“Nicobars”.

Material examined

Nicobars, NHMUK 1894.5.23.2 (1 syntype).

Remarks

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.

Chmamalycaeus canaliculatus (Möllendorff, 1894)

Alycaeus canaliculatus Möllendorff, 1894: 154–155, pl. 16, figs 22, 23.

Alycaeus (Chamalycaeus) canaliculatusKobelt 1902: 353.

Chamalycaeus (Chamalycaeus) canaliculatusZilch 1957: 142, pl. 5, fig. 4.

Dicharax canaliculatusPáll-Gergely 2017: 25, fig. 15C.

Type locality

“Samui Islands, Gulf of Siam” (from the title).

Material examined

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.

Chamalycaeus celebensis (E. von Martens, 1891)

Fig. 10A

Alycaeus celebensis E. von Martens, In: Weber, 1891: 217–218.

Alycaeus (Chamalycaeus) celebensisKobelt 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.

Figure 10. 

Shells of Chamalycaeus Möllendorff, 1897 species A Chamalycaeus celebensis (E. von Martens, 1891), possible syntype (ZMB/MOLL 44738) B Chamalycaeus kessneri Vermeulen, 1996, paratype (SMF 311351). Photographs: Barna Páll-Gergely (B) and Christine Zorn (A).

Chamalycaeus everetti (Godwin-Austen, 1889)

Alycaeus everetti Godwin-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).

Alycaeus everetti – E. A. Smith 1895: 116.

Alycaeus (Dicharax) everettiKobelt 1902: 369.

Chamalycaeus (Dicharax) everettiZilch 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 Alycaeus broti 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.

Chamalycaeus excisus (Möllendorff, 1887)

Alycaeus excisus Möllendorff, 1887b: 287.

Alycaeus (Chamalycaeus) excisusKobelt 1902: 355.

Chamalycaeus (Chamalycaeus) excisusZilch 1957: 142, pl. 6, fig. 20.

Chamalycaeus excisus excisusPáll-Gergely and Auffenberg 2019: 378, figs 2B, 3, 4C, D.

Type locality

“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.

Chamalycaeus excisus sublimus Páll-Gergely & Auffenberg, 2019

Chamalycaeus excisus sublimus Páll-Gergely & Auffenberg, 2019: 381, figs 4A, B, E, F, 6A, B.

Type locality

“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

Holotype (UF 115862) and paratypes, see Páll-Gergely and Auffenberg (2019).

Remarks

Same as the nominotypical subspecies.

Chamalycaeus fruhstorferi (Möllendorff, 1897)

Fig. 6B

Alycaeus (Chamalycaeus) fruhstorferi Möllendorff, 1897b: 93–94.

Alycaeus (Chamalycaeus) fruhstorferiKobelt 1902: 356.

Chamalycaeus fruhstorferi – van Benthem Jutting 1948: 571–572, fig. 26; Páll-Gergely et al. 2017: 7, fig. 46D–F.

Chamalycaeus (Chamalycaeus) fruhstorferiZilch 1957: 142, pl. 6, fig. 21.

Type locality

“Java” (from the title).

Material examined

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.

Chamalycaeus kessneri Vermeulen, 1996

Fig. 10B

Chamalycaeus kessneri Vermeulen, 1996: 150, fig. 2a–c.

Chamalycaeus kessneriVermeulen and Whitten 1998: 46, fig. 23.

Type locality

“Nusa Penida”.

Material examined

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.

Chamalycaeus microconus (Möllendorff, 1887)

Fig. 11A

Alycaeus microconus Möllendorff, 1887a: 311–312.

Alycaeus microconusMöllendorff 1891: 343, pl. 30, figs 12, 12a, 12b.

Alycaeus (Chamalycaeus) microconusKobelt 1902: 358.

Chamalycaeus (Chamalycaeus) microconusZilch 1957: 143, pl. 5, fig. 6.

Type locality

“Ad Buket Pondong”.

Material examined

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.

Figure 11. 

Shells of Chamalycaeus Möllendorff, 1897 species A Chamalycaeus microconus (Möllendorff, 1887), lectotype (SMF 109493) B C. mixtus Zilch, 1957, holotype (SMF 109510). All images: Barna Páll-Gergely, courtesy Ronald Janssen.

Chamalycaeus mixtus Zilch, 1957

Fig. 11B

Chamalycaeus (Chamalycaeus) mixtus Zilch, 1957: 143–144, pl. 5, fig. 7.

Type locality

“Malakka, Bukit Pondong, (Perak)”.

Material examined

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.

Chamalycaeus oglei (Godwin-Austen, 1914)

Alycaeus oglei Godwin-Austen, 1914: 362, pl. 148, fig. 2.

Alycaeus ogleiGude 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.

Chamalycaeus perplexus (Godwin-Austen, 1914)

Alycaeus perplexus Godwin-Austen, 1914: 380, pl. 155, fig. 11.

Alycaeus perplexusGude 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.

Chamalycaeus rarus Páll-Gergely & Auffenberg, 2019

Chamalycaeus rarus Páll-Gergely & Auffenberg, 2019: 382, fig. 6C.

Type locality

“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).

Chamalycaeus reinhardti (Mörch, 1872)

Alycaeus (Charax) reinhardi (sic) Mörch, 1872a: 22.

Alycaeus (Charax) reinhardi (sic) – Mörch 1872b: 315.

Alycaeus nicobaricus Reeve, 1878: pl. 4, species 29.

Alycaeus reinhardtiGodwin-Austen 1895: 455; Godwin-Austen 1914: 431; Gude 1921: 216–217.

Alycaeus (Alycaeus) reinhardtiKobelt 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).

Chamalycaeus reinhardti sabangensis (B. Rensch, 1933)

Alycaeus reinhardti sabangensis B. Rensch, 1933: 200–201.

Alycaeus (Alycaeus) reinhardti sabangensisZilch 1957: 147.

Type locality

“aus dem Walde bei Sabang”.

Material examined

Sumatra: Wald b. Sabang, Pulu Weh., exp. Rensch, 1927, SMF 6241 (1 paratype).

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.

Chamalycaeus reticulatus (Möllendorff, 1897)

Alycaeus (Orthalycaeus) reticulatus Möllendorff, 1897b: 93.

Alycaeus (Alycaeus) reticulatusKobelt 1902: 349; Zilch 1957: 147, pl. 6, fig. 26.

Alycaeus reticulatus – van Benthem Jutting 1948: 570–571.

Type locality

“Java” (from the title).

Material examined

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.

Chamalycaeus richthofeni (W. T. Blanford, 1863)

Alycaeus richthofeni W.T. Blanford, 1863: 324.

Alycaeus richthofeniReeve 1878: pl. 3, species 23; Godwin-Austen 1914: 428, pl. 151, fig. 9.

Alycaeus (Dicharax) richthofeniKobelt 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.

Chamalycaeus sculptilis (Benson, 1856)

Alycaeus sculptilis Benson, 1856: 226–227.

Alycaeus sculptilisReeve 1878: pl. 4, species 32, figs a, b; Godwin-Austen 1914: 398, 412, pl. 139, figs 7, 7a; pl. 155, fig. 8.

Alycaeus margarita Theobald in Hanley & Theobald, 1874: pl. 97, fig. 7 (renamed A. microstoma by Reeve 1878)

Alycaeus microstoma Reeve, 1878: pl. 4, species 28.

Alycaeus (Chamalycaeus) sculptilisKobelt 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 Alycaeus microstoma, 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 Alycaeus sculptilis Benson, 1856 by Godwin-Austen (1914).

Chamalycaeus specus (Godwin-Austen, 1889)

Alycaeus specus Godwin-Austen, 1889: 347, pl. 37, figs 4, 4a.

Alycaeus (Alycaeus) specusKobelt 1902: 351.

Type locality

“In limestone caves at Jambusan”.

Material examined

Caves, Borneo, leg. A. Everett, coll. Godwin-Austen, NHMUK 1889.12.7.26 (1 syntype).

Remarks

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.

Chamalycaeus subfossilis (P. Sarasin & F. Sarasin, 1899)

Alycaeus subfossilis P. Sarasin & F. Sarasin, 1899: 63–64, pl. 4, figs 46, 46a, pl. 5, fig. 66, pl. 8, fig. 91.

Alycaeus (Chamalycaeus) subfossilisKobelt 1902: 363–364.

Chamalycaeus subfossilisPáll-Gergely and Auffenberg 2019: 378, fig. 2A.

Type locality

“Geröllbank am Limbotto-See”.

Material examined

Limbotto See, NHMB 2265a (lectotype, designated herein), NHMB 2265a' (3 paralectotypes).

Remarks

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.

Chamalycaeus sumatranus (E. von Martens, 1900)

Fig. 12A

Alycaeus (Orthalycaeus) sumatranus E. von Martens, 1900: 6–7.

Alycaeus (Alycaeus) sumatranusKobelt 1902: 351.

Alycaeus sumatranus – van Benthem Jutting 1959: 77–78, fig. 5.

Type locality

“Unter-Lankat”.

Material examined

Unter-Lankat, coll. Schneider, ZMB/MOLL 51748 (1 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 (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.

Figure 12. 

Shells of Chamalycaeus Möllendorff, 1897 species A Chamalycaeus sumatranus (Martens, 1900), syntype (ZMB/MOLL 51748) B Chamalycaeus troglodytes (Rensch, 1934), syntype (ZMB/MOLL 76101). Photographs: Christine Zorn.

Chamalycaeus tanghali (Godwin-Austen, 1914)

Alycaeus tanghali Godwin-Austen, 1914: 401, pl. 137, figs 3, 3a, 3b.

Type locality

“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.

Chamalycaeus troglodytes (B. Rensch, 1934)

Fig. 12B

Alycaeus (Chamalycaeus) troglodytes B. Rensch, 1934: 743–744, fig. 3.

Type locality

“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.

Chamalycaeus vulcani (W. T. Blanford, 1863)

Alycaeus vulcani W.T. Blanford, 1863: 323.

Alycaeus vulcaniReeve 1878: pl. 2, species 17; Godwin-Austen 1914: 413–414, pl. 151, figs 5, 5a; Gude 1921: 221–222.

Alycaeus (Alycaeus) vulcaniKobelt 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.

Dicharax Kobelt & Möllendorff, 1900

Charax Benson, 1859: 177.

Dicharax Kobelt & 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

Alycaeus hebes Benson, 1857 (Fig. 13A), SD Gude (1921: 236); Awalycaeus abei Kuroda, 1951 (Fig. 13B), by monotypy (Awalycaeus); Alycaeus biexcisus Pilsbry, 1902 (Fig. 13C), by monotypy (Cipangocharax); Chamalycaeus (Sigmacharax) itonis Kuroda, 1943 (Fig. 13D), by monotypy (Sigmacharax).

Figure 13. 

Type species of alycaeid genus-group taxa A Dicharax hebes (Benson, 1857) (SMF 109244; type species of Dicharax) B D. (?) abei (Kuroda, 1951) (NSMT 50125; type species of Awalycaeus) C D. (?) biexcisus (Pilsbry, 1902) (NSMT 263; type species of Cipangocharax) D D. (?) itonis (Kuroda, 1943) (NSMT 78866; type species of Sigmacharax). Close-up images of the aperture are not to scale. All photographs: Barna Páll-Gergely.

Diagnosis

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).

Figure 14. 

Distribution of Dicharax Kobelt & Möllendorff, 1900.

Remarks

Cipangocharax, introduced as the subgenus of Chamalycaeus, was described for a single species, Alycaeus biexcisus. The diagnosis of Cipangocharax was in fact the abbreviated description of Alycaeus biexcisus. 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 Cipangocharax kiuchii is relatively slim, whereas that of “Chamalycaeusmiyazakii 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 Chamalycaeus nipponensis and Dicharax simplicilabris, 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.

Figure 15. 

R2 ribs of Japanese Dicharax Kobelt & Möllendorff, 1900 species A, B Dicharax (?) abei (Kuroda, 1951), NSMT 50125 C, D Dicharax (?) biexcisus (Pilsbry, 1902), NSMT 263 E, F Dicharax (?) itonis (Kuroda, 1943), NSMT 78866. All images: Barna Páll-Gergely.

Awalycaeus is a peculiar group of alycaeids due to the reduced (short, un-swollen) R3. However, in Awalycaeus yanoshokoae 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).

Typical Dicharax

Dicharax anonymus (Godwin-Austen, 1914)

Alycaeus anonymus Godwin-Austen, 1914: 405–406, pl. 139, figs 1, 1a.

Alycaeus anonymusGude 1921: 205.

Type locality

“Akouk-toung, Pegu: Type; also Thoudaung and Yenandoung, Pegu”.

Material examined

Akouktoung, Pegu, NHMUK 1906.4.4.67a (2 syntypes).

Remarks

Protoconch slightly elevated, no spiral lines visible; R1 with rather regular, moderately elevated ribs without spiral lines inbetween; R2 with ribs being lamella-like.

Dicharax anthostoma (Möllendorff, 1885)

Alycaeus anthostoma Möllendorff, 1885: 162.

Alycaeus pentagonus Heude, 1886: 211.

Alycaeus (Charax) anthostomaMöllendorff 1886: 166, pl. 5, fig. 4.

Alycaeus anthostomaGredler 1891: 79. (considered A. pentagonus as synonym)

Alycaeus (Charax) anthostomaKobelt and Möllendorff 1897: 149.

Alycaeus (Dicharax) anthostomaKobelt 1902: 364.

Chamalycaeus (Dicharax) anthostomaZilch 1957: 145, pl. 6, fig. 22.

Dicharax anthostomaPáll-Gergely et al. 2017: 30–32, figs 19A, B.

Type locality

“in regione Badung provinciae sinensis Hubei”.

Material examined

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.

Dicharax asaluensis (Godwin-Austen, 1914)

Alycaeus crispatus var. – Godwin-Austen 1874: 93, pl. 4, fig. 2.

Alycaeus asaluensis Godwin-Austen, 1914: 385–386, pl. 145, figs 2, 2a, 2b.

Alycaeus (Dicharax) asaluensisGude 1921: 237.

Chamalycaeus (Dicharax) asaluensis – Ramakrishna et al. 2010: 56.

Type locality

“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.

Dicharax avae (W. T. Blanford, 1863)

Alycaeus avae W. T. Blanford, 1863: 323–324.

Alycaeus avaeReeve 1878: pl. 3, species 20; Godwin-Austen 1914: 406–407, pl. 151, fig. 6.

Alycaeus (Dicharax) avaeKobelt 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.

Dicharax bison Páll-Gergely & Hunyadi, 2017

Dicharax bison Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 32–34, figs 21A, 22, 23.

Type locality

“China, Sichuan, Dujiangyan Shi, Taianzhen, Qingchenghoushan, Sanlongshuijing Rongdong, 942 m, 30°55.15418'N, 103°29.72375'E”.

Material examined

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.

Dicharax caudapiscis Páll-Gergely & Hunyadi, 2018

Dicharax caudapiscis Páll-Gergely & Hunyadi, 2018: 60, fig. 1A–E.

Type locality

“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.

Dicharax chennelli (Godwin-Austen, 1886)

Alycaeus chennelli Godwin-Austen, 1886: 192–193, pl. 48, fig. 2.

Alycaeus chennelli and chennelli var. – Godwin-Austen 1914: 387.

Alycaeus (Dicharax) chennelliKobelt 1902: 366; Gude 1921: 240–241.

Chamalycaeus (Dicharax) chennelli – Ramakrishna et al. 2010: 57.

Type locality

“Piknúi, Naga Hills”; “Lhota Naga Hills” (chennelli var.).

Material examined

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.

Dicharax conicus (Godwin-Austen, 1871)

Alycaeus conicus Godwin-Austen, 1871: 87–88, pl. 3, fig. 1.

Alycaeus conicusReeve 1878: pl. 1, species 9; Godwin-Austen 1914: 387–388, pl. 143, figs 4, 4a, 4b; Gude 1921: 208.

Alycaeus (Alycaeus) conicusKobelt 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.

Dicharax conicus jatingaensis Páll-Gergely, nom. nov.

Alycaeus conicus var. nanus Godwin-Austen, 1914: 388, pl. 138, figs 6, 6a, 6b. (non Alycaeus nanus Möllendorff, 1886)

Alycaeus conicus var. nanaGude 1921: 208.

Type locality

“Jatinga Valley, North Cachar Hills”.

Material examined

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.

Alycaeus conicus var. nanus Godwin-Austen, 1914 is a primary homonym of Alycaeus nanus 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.

Dicharax crenatus (Godwin-Austen, 1871)

Alycaeus crenatus Godwin-Austen, 1871: 90–91, pl. 3, fig. 5.

Alycaeus crenatusReeve 1878: pl. 1, species 1, figs a, b; Godwin-Austen 1914: 388–389, pl. 143, figs 8, 8a, 8b.

Alycaeus (Dicharax) crenatusKobelt 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.

Dicharax cristatus (Möllendorff, 1886)

Alycaeus cristatus Möllendorff, 1886: 168, pl. 5, fig. 6.

Alycaeus (Charax) cristatusKobelt and Möllendorff 1897: 150.

Alycaeus (Dicharax) cristatusKobelt 1902: 367.

Alycaeus (Chamalycaeus) smithi Fulton, 1907: 157, pl. 10, fig. 5.

Alycaeus (Charax) fimbriatus var. simplicilabris Bavay & Dautzenberg, 1912: 53–54, pl. 6, fig. 18.

Chamalycaeus (Dicharax) cristatusZilch 1957: 146, pl. 6, fig. 23.

Dicharax cristatusPáll-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.

Dicharax cucullatus (Theobald, 1870)

Alycaeus cucullatus Theobald, 1870: 396–397, pl. 18, fig. 2.

Alycaeus cucullatusReeve 1878: pl. 2, species 12; Godwin-Austen 1914, Vol. II: 407, pl. 155, fig. 5.

Alycaeus (Dicharax) cucullatusKobelt 1902: 367–368; Gude 1921: 244–245.

Type locality

“Shan States”.

Material examined

Shan States, NHMUK 1888.12.4.951–952 (2 syntypes).

Remarks

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.

Dicharax damsangensis (Godwin-Austen, 1886)

Alycaeus damsangensis Godwin-Austen, 1886: 192, pl. 43, figs 3, 3a–c.

Alycaeus (Dicharax) damsangensisKobelt 1902: 368; Gude 1921: 246–247.

Alycaeus (Charax) damsangensis – Godwin-Austen 1914: 339.

Type locality

“Damsang Peak, Western Bhutan Hills”.

Material examined

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.

Dicharax davisi (Godwin-Austen, 1914)

Alycaeus davisi Godwin-Austen, 1914: 408, pl. 148, figs 9, 9a.

Alycaeus (Chamalycaeus) davisiGude 1921: 226.

Type locality

“Siam and Shan boundary”.

Material examined

Shan States, leg. Woodthorpe, NHMUK 1903.7.1.1630 (4 syntypes).

Remarks

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.

Dicharax depressus (Bavay & Dautzenberg, 1912)

Alycaeus depressus Bavay & Dautzenberg, 1912: 51–52, pl. 4, figs 10–13.

Dicharax depressusPáll-Gergely et al. 2017: 43–45, figs 12E, F, 13D, 28E–H, 29E, F, 31A–C; Inkhavilay et al. 2019: 14, fig. 5A.

Type locality

“Pac-Kha, Tonkin”

Material examined

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)

Dicharax diagonius (Godwin-Austen, 1871)

Alycaeus diagonius Godwin-Austen, 1871: 88–89, pl. 3, fig. 2.

Alycaeus diagonus [sic] – Reeve 1878: pl. 1, species 2.

Alycaeus (Dicharax) diagoniusKobelt 1902: 368–369; Gude 1921: 247–248.

Alycaeus diagonius – 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.

Dicharax digitatus (H. F. Blanford, 1871)

Alycaeus digitatus H. F. Blanford, 1871: 41–42, pl. 2, fig. 4.

Alycaeus (Dicharax) digitatusKobelt 1902: 369; Gude 1921: 248.

Alycaeus digitatus – 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.

Dicharax diminutus (Heude, 1885)

Alycoeus [sic] diminutus Heude, 1885: 96, pl. 24, figs 5, 5a.

Alycaeus diminutusHeude 1886: 210.

Alycaeus (Orthalycaeus) diminutusMöllendorff 1886: 170.

Alycaeus (Orthalycaeus) nanus Möllendorff 1886: 170, pl. 5, fig. 8.

Alycaeus (Chamalycaeus) diminutusKobelt and Möllendorff 1897: 148; Kobelt 1902: 354.

Alycaeus nanusTarruella and Domènech 2011: 72.

Dicharax diminutusPáll-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.

Dicharax diplochilus (Möllendorff, 1887)

Alycaeus diplochilus Möllendorff, 1887a: 310.

Alycaeus diplochilusMöllendorff 1891: 342, pl. 30, figs 8, 8a, 8b.

Alycaeus (Chamalycaeus) diplochilusKobelt 1902: 354–355.

Chamalycaeus (Chamalycaeus) diplochilusZilch 1957: 142, pl. 5, fig. 5.

Dicharax diplochilusPáll-Gergely et al. 2017: 45, fig. 31D.

Type locality

“Ad Buket Pondong”.

Material examined

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.

Dicharax draco Páll-Gergely & Hunyadi, 2017

Dicharax draco Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 52–54, fig. 34A.

Type locality

“China, Yunnan, Wenshanzhuang Zumiaozu Zizhizho, Guangnan Xian, Liji, 1611 m, 23°45.54175'N, 104°59.55567'E”.

Material examined

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.

Dicharax elevatus (Heude, 1886)

Fig. 16

Alycaeus elevatus Heude, 1886: 210.

Alycaeus elevatusHeude 1890: 129, pl. 36, fig. 19.

Alycaeus (Chamalycaeus) elevatusKobelt and Möllendorff 1897: 148; Kobelt 1902: 355.

Metalycaeus (?) elevatusPáll-Gergely et al. 2017: 104–105, fig. 69I.

Type locality

“Tchen K’eou”.

Material examined

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.

Figure 16. 

Dicharax (?) elevatus (Heude, 1886), syntype (HMT-218a). Photographs: Kaibaryer Meng.

Dicharax fimbriatus (Bavay & Dautzenberg, 1912)

Alycaeus (Charax) fimbriatus Bavay & Dautzenberg, 1912: 52–53, pl. 6, figs 14–17.

Chamalycaeus plicilabris multidentatus Yen, 1939: 29, pl. 2, fig. 33.

Dicharax fimbriatusPáll-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.

Dicharax fraterculus (Bavay & Dautzenberg, 1900)

Alycaeus (Charax) fraterculus Bavay & Dautzenberg, 1900a: 120.

Alycaeus (Charax) fraterculusBavay and Dautzenberg 1900b: 457–458, pl. 11, figs 11–14.

Alycaeus (Dicharax) fraterculusKobelt 1902: 370.

Dicharax fraterculusPáll-Gergely et al. 2017: 62–64, fig. 41.

Type locality

“Haut-Tonkin”.

Material examined

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.

Dicharax generosus (Godwin-Austen, 1914)

Alycaeus generosus Godwin-Austen, 1914: 374, pl. 138, figs 8, 8a, 8b.

Alycaeus (Cycloryx) generosusGude 1921: 279.

Cycloryx generosus – Ramakrishna et al. 2010: 71.

Type locality

“Khasi Hills”.

Material examined

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.

Dicharax globulus (Godwin-Austen, 1874)

Alycaeus globulus Godwin-Austen, 1874: 147–148, pl. 3, fig. 4.

Alycaeus (Dicharax) globulusKobelt 1902: 371; Gude 1921: 254.

Alycaeus globulus – Godwin-Austen 1914: 392, pl. 144, figs 4, 4a, 4b.

Chamalycaeus (Dicharax) globulus – Ramakrishna et al. 2010: 60.

Type locality

“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.

Dicharax habiangensis (Godwin-Austen, 1914)

Alycaeus habiangensis Godwin-Austen, 1914: 374, pl. 138, figs 2, 2a, 2b.

Alycaeus (Dicharax) habiangensisGude 1921: 254.

Chamalycaeus (Dicharax) habiangensis – Ramakrishna et al. 2010: 61.

Type locality

“Habiang Garo, on the West Khasi border”.

Material examined

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).

Dicharax hebes (Benson, 1857)

Alycaeus hebes Benson, 1857: 204–205.

Alycaeus hebesReeve 1878: pl. 6, species 52; Godwin-Austen 1886: 191, pl. 43, figs 1, 1a–c.

Alycaeus (Dicharax) hebesKobelt 1902: 371; Gude 1921: 255.

Alycaeus hebes – Godwin-Austen 1914: 374–375, pl. 145, figs 5, 5a, 5b.

Chamalycaeus (Dicharax) hebes – Ramakrishna et al. 2010: 61.

Dicharax hebesPáll-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.

Dicharax humilis (W. T. Blanford, 1862)

Alycaeus humilis W. T. Blanford, 1862: 136–137.

Alycaeus humilisReeve 1878: pl. 5, species 40; Godwin-Austen 1914: 408–409, pl. 151, fig. 8.

Alycaeus (Dicharax) humilisKobelt 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.

Dicharax imitator Páll-Gergely & Hunyadi, 2017

Dicharax imitator Páll-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.

Dicharax jaintiacus (Godwin-Austen, 1871)

Alycaeus Jaintiacus Godwin-Austen, 1871: 92–93, pl. 5, fig. 3.

Alycaeus (Dicharax) jaintiacusKobelt 1902: 372; Gude 1921: 256.

Alycaeus jaintiacus – Godwin-Austen 1914: 375, pl. 143, figs 3, 3a, 3b.

Chamalycaeus (Dicharax) jaintiacus – Ramakrishna et al. 2010: 61.

Type locality

“in Nongjinghi, Jiantia”.

Material examined

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.

Dicharax jaintiacus crassus (Godwin-Austen, 1914)

Alycaeus jaintiacus var. crassus Godwin-Austen, 1914: 375, pl. 137, figs 5, 5a.

Alycaeus (Dicharax) jaintiacus Var. crassaGude 1921: 256–257.

Type locality

“in Nongjinghi, Jiantia, 4563 feet”.

Material examined

Nonjinghi, Jiantia, coll. Godwin-Austen, NHMUK 1903.7.1.2752 (4 syntypes in 2 vials).

Remarks

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.

Dicharax kezamaensis (Godwin-Austen, 1914)

Alycaeus kezamaensis Godwin-Austen, 1914: 393, pl. 149, fig. 1.

Alycaeus (Dicharax) kezamaensisGude 1921: 258.

Chamalycaeus (Dicharax) kezamaensis – Ramakrishna et al. 2010: 62.

Type locality

“Kezama, Aughami-Naga Hills”.

Material examined

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).

Dicharax lahupaensis (Godwin-Austen, 1914)

Alycaeus lahupaensis Godwin-Austen, 1914: 394, pl. 141, figs 3, 3a.

Alycaeus (Raptomphalus) lahupaensisGude 1921: 287–288.

Chamalycaeus (Raptomphalus) lahupaensis – Ramakrishna et al. 2010: 68.

Type locality

“Gaziphimi, Lahupa Naga Hills, Munipur”.

Material examined

Gaziphimih, N.E. Munipur, NHMUK 1903.7.1.2655 (10 syntypes).

Remarks

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.

Dicharax longituba (E. von Martens, 1864)

Alycaeus longituba E. von Martens, 1864: 1 20.

Alycaeus longituba – E. von Martens 1867: 151, pl. 4, fig. 8.

Alycaeus (Dicharax) longitubaKobelt 1902: 373.

Chamalycaeus longituba – 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).

Dicharax maosmaiensis (Godwin-Austen, 1922)

Alycaeus maosmaiensis Godwin-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).

Dicharax microcostatus Páll-Gergely, 2017

Dicharax microcostatus Páll-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.

Dicharax microdiscus (Möllendorff, 1887)

Alycaeus microdiscus Möllendorff, 1887a: 311.

Alycaeus microdiscusMöllendorff 1891: 343.

Alycaeus (Chamalycaeus) microdiscusKobelt 1902: 358.

Chamalycaeus (Chamalycaeus) microdiscusZilch 1957: 143, pl. 5, fig. 8; Egorov 2013: 35, fig. 62b.

Dicharax microdiscusPáll-Gergely 2017: 25, fig. 15D.

Type locality

“Ad Buket Pondong”.

Material examined

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).

Dicharax micropolitus Páll-Gergely & Hunyadi, 2017

Dicharax micropolitus Páll-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.

Dicharax nitidus (W. T. Blanford, 1862)

Alycaeus nitidus W. T. Blanford, 1862: 141.

Alycaeus nitidusReeve 1878: pl 3, species 25; Godwin-Austen 1914: 421–422, pl. 151, figs 4, 4a.

Alycaeus (Chamalycaeus) nitidusKobelt 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.

Dicharax notatus (Godwin-Austen, 1876)

Alycaeus notatus Godwin-Austen, 1876: 176, pl. 7, figs 9, 9a, 9b.

Alycaeus notatus – Godwin-Austen 1886: 191–192, pl. 43, figs 2, 2a–c; Godwin-Austen 1914: 358–359, pl. 145, figs 8, 8a.

Alycaeus (Dicharax) notatusKobelt 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.

Dicharax notus (Godwin-Austen, 1914)

Alycaeus notus Godwin-Austen, 1914: 411, pl. 155, fig. 12.

Alycaeus (Dicharax) notusGude 1921: 262.

Type locality

“Fort Stedman, Burma”.

Material examined

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.

Dicharax nowgongensis (Godwin-Austen, 1914)

Alycaeus nowgongensis Godwin-Austen, 1914: 397, pl. 137, figs 4, 4a, 4b.

Alycaeus nowgongensisGude 1921: 213.

Alycaeus (Alycaeus) nowgongensis – Ramakrishna et al. 2010: 49.

Type locality

“Koliaghur or Koliahur, Nowgoug District, Assam”.

Material examined

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.

Dicharax ochraceus (Godwin-Austen, 1893)

Alycaeus ochraceus Godwin-Austen, 1893: 594–595.

Alycaeus ochraceus – Godwin-Austen 1897: 3, pl. 63, figs 7, 7a, 7b; Godwin-Austen 1914: 411.

Alycaeus (Dicharax) ochraceusKobelt 1902: 374; Gude 1921: 263.

Type locality

“Ruby Mines District, Upper Burmah”.

Material examined

Ruby mine Disr., Up. Burma, leg. Doherty, NHMUK 1903.7.1.2684 (2 syntypes).

Remarks

Protoconch low, moderately glossy, without spiral lines; R1 finely, regularly ribbed, without spiral striae; R2 moderately long, with regular ribs; ribs curved towards the aperture (typical Dicharax).

Dicharax oligopleuris (Möllendorff, 1887)

Alycaeus oligopleuris Möllendorff, 1887a: 310–311.

Alycaeus oligopleurisMöllendorff 1891: 342, pl. 30, figs 9, 9b.

Alycaeus (Chamalycaeus) oligopleurisKobelt 1902: 360–361.

Chamalycaeus (Chamalycaeus) oligopleurisZilch 1957: 144, pl. 5, fig. 9.

Dicharax oligopleurisPáll-Gergely et al. 2017: 45, fig. 31E.

Type locality

“Ad Buket Pondong”.

Material examined

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.

Dicharax parvulus (Möllendorff, 1887)

Alycaeus parvulus Möllendorff, 1887a: 311.

Alycaeus parvulusMöllendorff 1891: 343, pl. 30, figs 11, 11b.

Alycaeus (Chamalycaeus) parvulusKobelt 1902: 361.

Chamalycaeus (Chamalycaeus) parvulusZilch 1957: 144, pl. 5, fig. 10; Egorov 2013: 35, fig. 62a.

Dicharax parvulusPáll-Gergely 2017: 25, fig. 15E.

Type locality

“Ad Buket Pondong”.

Material examined

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.

Dicharax planorbulus (Heude, 1885)

Alycoeus [sic] planorbulus Heude, 1885: 96, pl. 24, figs 2, 2a–c.

Alycaeus (Chamalycaeus) planorbulusKobelt and Möllendorff 1897: 149; Kobelt 1902: 361.

Dicharax planorbulusPáll-Gergely et al. 2017: 68, figs 21B, C, 42E, F, 44.

Type locality

“in ditione Tchen-k’eou”.

Material examined

China, Tchen-K’eou, MCZ 167136 (10 syntypes).

Remarks

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.

Dicharax plicilabris (Möllendorff, 1886)

Alycaeus plicilabris Möllendorff, 1886: 167, pl. 5, fig. 5.

Alycaeus (Chamalycaeus) plicilabrisKobelt and Möllendorff 1897: 149; Kobelt 1902: 361.

Chamalycaeus plicilabris plicilabrisYen 1939: 29, pl. 2, fig. 32.

Chamalycaeus (Chamalycaeus) plicilabris plicilabrisZilch 1957: 145, pl. 5, fig. 14.

Dicharax plicilabrisPáll-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.

Dicharax politus (W. T. Blanford, 1865)

Alycaeus politus W. T. Blanford, 1865: 83–84.

Alycaeus politusReeve 1878: pl. 5, species 39; Godwin-Austen 1914: 422, pl. 139, figs 5, 5a; Gude, 1921: 214–215.

Alycaeus (Alycaeus) politusKobelt 1902: 348.

Type locality

“Phuong do, near Cape Negrais, Arakan”.

Material examined

Phungdo, Arakan, coll. Blanford, NHMUK 1906.4.4.178 (3 probable syntypes).

Remarks

Protoconch low, glossy, no spiral lines visible; R1 glossy, without notable sculpture; R2 short, with regular ribs, which are curved towards the aperture.

Dicharax pratatensis (Panha & Burch, 1997)

Alycaeus pratatensis Panha & Burch, 1997: 119–122, figs 2a–c.

Type locality

“Pratat cave, Erawan Natural Park, Karnchanaburi Province at 14°27'58"N, 99°49'49"E, 230 meters elevation.”

Material examined

Pratat cave, Erawan N. P., Karnchanaburi Province, Thailand, 26.10.1996, ex coll. S. Panha, 2008, SMF 331452 (2 paratypes).

Remarks

Protoconch low, without spiral lines; R1 irregularly wrinkled, without spiral lines; R2 short, with ribs, which are curved towards the aperture.

Dicharax robustus Páll-Gergely & Hunyadi, 2017

Dicharax micropolitus Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 70–71, figs 19C, 29A, B, 45.

Type locality

“China, Yunnan, Kunming Shi, Yuqiqu, Bijianshan, Guanyinsi (temple), approximate GPS data: 24°16.271'N, 102°49.726'E”.

Material examined

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.

Dicharax stuparum Páll-Gergely & Hunyadi, 2018

Dicharax stuparum Páll-Gergely & Hunyadi, 2018: 62, figs 1F–K.

Type locality

“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.

Dicharax sylheticus (Godwin-Austen, 1914)

Alycaeus sylheticus Godwin-Austen, 1914: 382, pl. 154, figs 4, 4a.

Alycaeus sylheticusGude 1921: 220.

Type locality

“South Sylhet Hills”.

Material examined

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).

Dicharax tangmaiensis (Chen & Zhang, 2001)

Fig. 17

Chamalycaeus tangmaiensis Chen & Zhang, 2001: 184–185, 188–189, figs 1–4.

Dicharax (?) tangmaiensisPáll-Gergely et al. 2017: 107–108.

Type locality

“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 Dicharax tangmaiensis 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.

Figure 17. 

Dicharax tangmaiensis (Chen & Zhang, 2001), holotype (CASIZ TM 0010054). Photographs: Kaibaryer Meng.

Dicharax theobaldi (W. T. Blanford, 1862)

Alycaeus Theobaldi W. T. Blanford, 1862: 142–143.

Alycaeus theobaldiReeve 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) theobaldiKobelt 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.

Dicharax theobaldi diyungensis (Godwin-Austen, 1914)

Alycaeus theobaldi var. diyungensis Godwin-Austen, 1914: 401–402, pl. 138, fig. 4.

Alycaeus (Dicharax) theobaldi Var. diyungensisGude 1921: 274.

Type locality

“ad Darjiling”.

Material examined

Diyung Valley, N of Asalu, NHMUK 1903.7.1.2546 (12 syntypes).

Remarks

Same as in theobaldi solidus, but R2 is longer.

Dicharax theobaldi solidus (Godwin-Austen, 1914)

Alycaeus theobaldi var. solidus Godwin-Austen, 1914: 383–384, pl. 155, fig. 10.

Alycaeus (Dicharax) theobaldi Var. solidaGude 1921: 273–274.

Type locality

“Garo Hills”.

Material examined

Garo Hills, NHMUK 1903.7.1.2560 (4 syntypes).

Remarks

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.

Dicharax vestitus (W. T. Blanford, 1862)

Alycaeus vestitus W. T. Blanford, 1862: 138–139.

Alycaeus vestitus var. minor W. T. Blanford, 1862: 138.

Alycaeus vestitusReeve 1878: pl. 1, species 3; Godwin-Austen 1914: 424–425, pl. 139, figs 2, 2a; Gude 1921: 220–221.

Alycaeus (Alycaeus) vestitusKobelt 1902: 352.

Dicharax vestitusPáll-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.

Dicharax vestitus akyabensis (Godwin-Austen, 1914)

Alycaeus vestitus var. akyabensis Godwin-Austen, 1914: 425–426, pl. 155, fig. 7.

Alycaeus vestitus var. akyabensisGude 1921: 221.

Type locality

“Baumi, Akyab”.

Material examined

Baumi, Akyab, NHMUK 1888.12.4.251–252 (2 syntypes).

Remarks

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 species

Dicharax (?) abdoui Páll-Gergely, 2017

Dicharax abdoui Páll-Gergely in Páll-Gergely et al., 2017: 14, fig. 6.

Dicharax abdouiInkhavilay 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).

Dicharax (?) akhaensis (Godwin-Austen, 1914)

Alycaeus akhaensis Godwin-Austen, 1914: 352, pl. 141, figs 1, 1a, 1b.

Alycaeus (Raptomphalus) akhaensisGude 1921: 286.

Chamalycaeus (Raptomphalus) akhaensis – Ramakrishna et al. 2010: 68.

Type locality

“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.

Dicharax (?) alticola Páll-Gergely & Hunyadi, 2017

Dicharax alticola Páll-Gergely & Hunyadi in Páll-Gergely et al., 2017: 14–20, figs 8A, 9A–D, 10A, B, 11, 12A, B, 13A, B.

Type locality

“China, Sichuan, Liangshan Yizu Zizhizhou, Yanyuan Xian, Bainiao Zhen, Kedeng Rongdong (cave), 2618 m, 27°43.103'N, 101°31.021'E”.

Material examined

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.

Dicharax (?) ataranensis (Godwin-Austen, 1914)

Fig. 18

Alycaeus ataranensis Godwin-Austen, 1914: 426, pl. 148, figs 4, 4a, 4b.

Alycaeus (Dicharax) ataranensisGude 1921: 237.

Type locality

“1 Ataran”.

Material examined

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.

Figure 18. 

Dicharax ataranensis (Godwin-Austen, 1914), holotype (NZSI M.8073). All images: Sheikh Sajan.

Dicharax (?) barowliensis (Godwin-Austen, 1914)

Alycaeus barowliensis Godwin-Austen, 1914: 352, pl. 141, fig. 4.

Alycaeus barowliensisGude 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.

Dicharax (?) bawai Aravind & Páll-Gergely, 2018

Dicharax (?) bawai Aravind & Páll-Gergely, 2018: 56, figs 1A, 2, 3.

Type locality

“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.

Dicharax (?) bicrenatus (Godwin-Austen, 1874)

Alycaeus bicrenatus Godwin-Austen, 1874: 148, pl. 3, fig. 5.

Alycaeus (Dicharax) bicrenatusKobelt 1902: 365; Gude 1921: 238–239.

Alycaeus bicrenatus – 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.

Dicharax (?) bifrons (Theobald, 1870)

Alycaeus bifrons Theobald, 1870: 396, pl. 18, fig. 1.

Alycaeus bifronsReeve 1878: pl. 6, species 48; Godwin-Austen 1914: 407, pl. 139, figs 3, 3a.

Alycaeus (Dicharax) bifronsKobelt 1902: 365–366; Gude 1921: 239.

Type locality

“Shan States”.

Material examined

Shan States, NHMUK 1888.12.4.956–958 (3 syntypes).

Remarks

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.

Dicharax (?) birugosus (Godwin-Austen, 1893)

Fig. 19

Alycaeus bi-rugosus Godwin-Austen, 1893: 593.

Alycaeus bi-rugosus – Godwin-Austen 1897: 387, pl. 63, figs 5, 5a.

Alycaeus (Dicharax) birugosusKobelt 1902: 366.

Alycaeus birugosus – Godwin-Austen 1914: 370; Godwin-Austen 1914: 387.

Alycaeus canaliculus Godwin-Austen, 1914: 371, pl. 154, fig. 11. syn. nov.

Alycaeus birugosus var. – Godwin-Austen 1914: 370, pl. 154, figs 7, 7a.

Alycaeus birugosus var. minor Godwin-Austen, 1914: 370, pl. 155, figs 9, 9a.

Alycaeus (Chamalycaeus) canaliculusGude 1921: 225.

Alycaeus (Dicharax) birugosusGude 1921: 239–240.

Alycaeus (Dicharax) birugosus var. minorGude 1921: 240.

Chamalycaeus (Chamalycaeus) canaliculus – Ramakrishna et al. 2010: 53.

Chamalycaeus (Dicharax) birugosus – Ramakrishna et al. 2010: 57.

Type locality

“Khasi Hills and Manipur” (A. birugosus); “Garo Hills” (A. birugosus var. minor); “Teria Ghat, foot of the Khasi Hills” (A. canaliculus).

Material examined

Khasi Hills, leg. Godwin-Austen, NHMUK 1903.7.1.2628 (2 syntypes of A. birugosus, Fig. 19B); Jawai, Jiantia Hills, 282a, leg. Godwin-Austen, NHMUK 1903.7.1.2571 (7 shells, labelled as “birugosus var.”); Garo Hills, NHMUK 1903.7.1.2755 (1 syntype of Alycaeus birugosus var. minor, labelled as duorugosus var. minor); Teria Ghat, Khasi, leg. Godwin-Austen, NHMUK 1903.7.1.2764 (1 syntype of A. canaliculus, Fig. 19A).

Figure 19. 

Shells of Dicharax (?) birugosus (Godwin-Austen, 1893) A Alycaeus canaliculus Godwin-Austen, 1914, (NHMUK 1903.7.1.2764) B A. birugosus Godwin-Austen, 1893, syntype (NHMUK 1903.7.1.2628). Photographs: Kevin Webb (NHM).

Remarks

Alycaeus birugosus 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.

Dicharax (?) blanfordi (Godwin-Austen, 1914)

Alycaeus blanfordi Godwin-Austen, 1914: 418, pl. 148, fig. 3.

Alycaeus blanfordiGude 1921: 206.

Type locality

“Chwegali, Arakan Hills”.

Material examined

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.

Dicharax (?) burroiensis (Godwin-Austen, 1914)

Alycaeus burroiensis Godwin-Austen, 1914: 354, pl. 141, figs 6, 6a.

Alycaeus (Cycloryx) burroiensisGude 1921: 277.

Type locality

“Burroi Gorge, Dafla Hills”.

Material examined

Burroi Rr., Dafla, NHMUK 1903.7.1.2653 (1 syntype).

Remarks

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.

Dicharax (?) candrakirana Nurinsiyah & Hausdorf, 2017

Dicharax (?) candrakirana Nurinsiyah & Hausdorf, 2017: 589–591, fig. 1.

Type locality

“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.

Dicharax (?) crassicollis (Benthem-Jutting, 1959)

Chamalycaeus crassicollis van Benthem Jutting, 1959: 76–77, fig. 4.

Type locality

“Sirung Galing, Karo Highlands”.

Remarks

No specimens were examined. The general shape and the sculpture of the species is similar to Dicharax longituba according to the original description. Therefore, Alycaeus crassicollis is tentatively classified in Dicharax.

Dicharax (?) crispatus (Godwin-Austen, 1871)

Alycaeus crispatus Godwin-Austen, 1871: 91–92, pl. 4, fig. 1.

Alycaeus crispatusGodwin-Austen 1875: 8, pl. 4, fig. 3.

Alycaeus (Dicharax) crispatusKobelt 1902: 367; Gude 1921: 242–243.

Alycaeus crispatus – 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 Alycaeus crispatus 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 cristatus minimus shells have only slightly elevated protoconchs, whereas it is characteristically Chamalycaeus-like (strongly elevated) in D. crispatus makarsae specimens.

Dicharax (?) crispatus makarsae (Godwin-Austen, 1914)

Alycaeus crispatus var. makarsae Godwin-Austen, 1914: 372, pl. 158, fig. 13.

Alycaeus (Dicharax) crispatus var. makarsaeGude 1921: 243.

Type locality

“Makarsa, N. Khasi Hills (or more correct, Maokarsa; the common Khasi prefix “Mao” meaning a stone”.

Material examined

Makarsa, Khasi, leg. Godwin-Austen, NHMUK 1903.7.1.2638 (8 syntypes).

Remarks

See under Chamalycaeus crispatus.

Dicharax (?) crispatus minimus (Godwin-Austen, 1914)

Alycaeus crispatus var. minimus Godwin-Austen, 1914: 373, pl. 148, figs 5, 5a.

Alycaeus (Dicharax) crispatus var. minimaGude 1921: 243–244.

Type locality

“Habiang Garo Hills, West Khasi”.

Material examined

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.

Dicharax (?) crispatus rywukensis (Godwin-Austen, 1914)

Alycaeus crispatus var. rywukensis Godwin-Austen, 1914: 373–374, pl. 154, figs 3, 3a.

Alycaeus (Dicharax) crispatus var. rywukensisGude 1921: 244.

Type locality

“Rywuk Valley of the Garo Hills”.

Material examined

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.

Dicharax (?) daflaensis (Godwin-Austen, 1876)

Alycaeus daflaensis Godwin-Austen, 1876: 176–177, pl. 7, figs 12, 12a, 12b.

Alycaeus (Dicharax) daflaensisKobelt 1902: 368; Gude 1921: 245–246.

Alycaeus daflaensis – 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.

Dicharax (?) daflaensis subdigitatus (Godwin-Austen, 1876)

Alycaeus sub-digitatus Godwin-Austen, 1876: 177.

Alycaeus (Dicharax) daflaensis var. subdigitataKobelt 1902: 368.

Alycaeus (Dicharax) daflaensis var. subdigitatusGude 1921: 246.

Type locality

“Shengorh Peak” and “Tánir ridge at 4000 feet”.

Material examined

Shengorh Peak, Dafla Hills, coll. Godwin-Austen, NHMUK 1903.7.1.2498 (3 syntypes).

Remarks

As in the nominotypical subspecies.

Dicharax (?) dalingensis (Godwin-Austen, 1914)

Alycaeus dalingensis Godwin-Austen, 1914: 338–339, pl. 134, figs 3, 3a–c.

Alycaeus (Dicharax) dalingensisGude 1921: 246.

Type locality

“Rechila Peak, Daling District, on Sikkim and Bhutan Boundary (10,300 ft.)”.

Material examined

Rechila Pk, Sikkim, leg. W. Robert, NHMUK 1903.7.1.1251 (7 syntypes).

Remarks

Protoconch low, glossy; R1 glossy, no spiral and radial lines visible; R2 short, also glossy, alternating thick/dark and narrow/lighter stripes.

Dicharax (?) dohertyi (Godwin-Austen, 1893)

Alycaeus dohertyi Godwin-Austen, 1893: 595.

Alycaeus dohertyi – Godwin-Austen 1897: 3, pl. 63, figs 3, 3a; Godwin-Austen 1914: 408.

Alycaeus (Dicharax) dohertyiKobelt 1902: 369; Gude 1921: 248–249.

Type locality

“Momeit, Burmah”.

Remarks

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.

Dicharax (?) dolichodeiros (Heude, 1890)

Alycaeus dolichodeiros Heude, 1890: 129, pl. 38, fig. 3.

Alycaeus (Chamalycaeus) dolichodeirus (sic) – Kobelt and Möllendorff 1897: 148.

Alycaeus (Chamalycaeus) dolichodirus (sic) – Kobelt 1902: 355.

Chamalycaeus (Chamalycaeus) dolichodirus (sic) – Zilch 1957: 142.

Dicharax (?) dolichodeirosPáll-Gergely et al. 2017: 22–23, figs 8D, E, 9E, F, 10E, F.

Type locality

“Tchen k’eou”.

Remarks

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 “Chamalycaeushelicodes (= synonym of Metalycaeus muciferus, see Páll-Gergely et al. 2017). This species was putatively classified into the genus Dicharax by Páll-Gergely et al. (2017).

Dicharax (?) duoculmen (Godwin-Austen, 1914)

Alycaeus duoculmen Godwin-Austen, 1914: 365, pl. 157, figs 2, 2a.

Alycaeus (Raptomphalus) duoculmenGude 1921: 286–287.

Chamalycaeus (Raptomphalus) duoculmen – Ramakrishna et al. 2010: 68.

Type locality

“Tsanspu Valley”.

Material examined

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.

Dicharax (?) duorugosus (Godwin-Austen, 1914)

Alycaeus duorugosus Godwin-Austen, 1914: 391.

Alycaeus (Dicharax) duorugosusGude 1921: 249.

Chamalycaeus (Dicharax) duorugosus – Ramakrishna et al. 2010: 59.

Type locality

“Burrail Range, Naga”, “Also Angaoluo Trigonometrical Station, No. 2572; South Barak, No. 2629, and Munipur, No. 2654 B.M.”.

Material examined

Burrail, coll. Godwin-Austen, NHMUK 1903.7.1.2771 (1 syntype).

Remarks

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.

Dicharax (?) edei (Godwin-Austen, 1914)

Alycaeus edei Godwin-Austen, 1914: 391–392, pl. 149, figs 2, 2a.

Alycaeus (Chamalycaeus) edeiGude 1921: 227.

Chamalycaeus (Chamalycaeus) edei – Ramakrishna et al. 2010: 53.

Type locality

“Naraindhur, Cachar, No. 1665 B.M.”.

Material examined

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.