Annotated nomenclator of extant and fossil taxa of the Paludomidae (Caenogastropoda, Cerithioidea)

Abstract This nomenclator provides bibliographic details on all names in the family-, genus-, and species-group of the the family Paludomidae. All nomenclaturally available names are discussed including junior homonyms and objective junior synonyms as well as unavailable names such as nomina nuda, infrasubspecific names and, to some extent, also incorrect subsequent spellings. In the family-group a total of 28 names are included in the nomenclator, of which 21 are available and seven unavailable names. Of the available names in the family-group, six are invalid for nomenclatural reasons. In the genus-group a total of 57 names are included in the catalogue. Of the available names in the genus-group, 11 are invalid for nomenclatural reasons. In the species-group a total of 499 names are included, of which 463 are available, but 21 are invalid for nomenclatural reasons. All names are given in their original combination and spelling (mandatory changes are discussed and corrected spellings are provided), along with the reference to the original publication. For each family- and genus-group name, the original classification and the type genus or type species, respectively, are given. For species-group taxa the type locality and type horizon (for fossil taxa) are provided, usually as given in the original publication. A new name, Cleopatraadami nom. nov., is proposed for the fossil Cleopatracylindrica (Adam, 1957), which is a homonym of Cleopatracridlandicylindrica Mandahl-Barth, 1954, and a lectotype for Cleopatradubia Adam, 1959 is designated. A new replacement name Leloupiella nom. nov. is proposed for Stormsia Leloup, 1953 which is a homonym of Stormsia Bourguignat, 1891.


Introduction
The Paludomidae Stoliczka, 1868 is currently understood as representing taxa with a general distribution including most of tropical sub-Saharan Africa and Madagascar as well as South and Southeast Asia (see Neiber and Glaubrecht 2019b who summarize the results of phylogenetic analyses presented by West and Michel 2000;Michel 2004;Wilson et al. 2004;Glaubrecht and Strong 2007;Strong et al. 2011; for a recent compilation on the diversity of family-group names available for this group and recent classifications, see also Bouchet and Rocroi 2005;Bouchet et al. 2017).
Species in this family of freshwater cerithioideans show a high diversity with regard to shell shape and ornamentation. The endemic paludomids of Lake Tanganyika have especially attracted much attention by taxonomists, early biogeographers, and evolutionary biologists because of their thalassoid (Bourguignat 1885) appearance, i.e. their resemblance to various marine gastropod families (Glaubrecht 1996(Glaubrecht , 2008; see also Glaubrecht 2009Glaubrecht , 2011. A large number of names have been introduced for Tanganyikan paludomids by the French malacologists J.-R. Bourguignat, the head of the so called 'Nouvelle École' (Dance 1970), on the basis of differences of the shell that are often hardly perceptible (Bourguignat 1885(Bourguignat , 1888. However, being among the first to work on the gastropod fauna of the lake, several names first introduced by this author are still in use to this day, although the far greater number certainly represent individual variations that do not merit to be accepted and are thus to a large extent a burden today for taxonomists, biogeographers, and evolutionary biologists working on the paludomid fauna of Lake Tanganyika. Although several synoptical works covering the regional diversity of Paludomidae in Africa or Asia have been published (e.g. Preston 1915b; Pilsbry and Bequaert 1927;Leloup 1953;Brown 1994;West et al. 2003), Asian members of the family are still especially in need of revision. The basic prerequisite of any taxonomic work on a specific group of organisms is the historical perspective of what is already known, or from a nomenclatural point of view, which names have already been proposed for the group in question. However, a comprehensive annotated list of paludomid names is currently unavailable. Therefore, the nomenclator presented here aims at filling this gap by providing information on all names for Recent and fossil family-, genus-, and species-group taxa introduced for Paludomidae. In addition, the present contribution is intended to facilitate future taxonomic work on Paludomidae by providing direct internet links to many important older taxonomic publications following the general outlines of the nomenclators of other groups of freshwater snails presented by Haszprunar (2014) on valvatids and Neubauer (2016) on melanopsids.
searcher, such names may be taxonomically invalid names. A proposal whether a name is currently accepted or not is not made as this is, in our opinion, the task for future revisionary work. The original reference for a name is always provided. An authorship of a name deviating from the authorship of the work in which it was published, is only accepted in cases where clear evidence exists that person(s) other than the author(s) of the work, in which the name was published, were responsible for the name (Code Art. 50.1.1 and 50.1.3). In such cases, the name(s) of the person(s) responsible for the name is/are listed directly behind the name and the author(s) of the work, in which the name was published, is/are cited under 'Original reference'. Information on type genera and type species, where relevant, are provided including references to the selection of type species (if subsequently designated). Names exclusively based on fossils are marked by a dagger. Information on type localities and type strata (for fossils only) are usually given exactly as provided in the original reference (in quotes); only in few cases also other sources are additionally provided. Additional information is given in square brackets (e.g., country, province). All other nomenclaturally relevant issues are discussed in the 'Remarks' under a name; however actions of a First Reviser in the sense of the Code are only discussed in cases where simultaneously published primary homonyms or different spellings of a name in the original publication are involved. Incorrect subsequent spellings are discussed under the respective correctly spelled names whenever these were noticed.

Part 1. Family-group names
In the family-group a total of 28 names are included in the nomenclator, of which 21 are available and seven unavailable names. Of the available names in the family-group, six are invalid for nomenclatural reasons. Rocroi (2005: 176) the name is interpreted as descriptive alluding to the resemblance of species from Lake Tanganyika to the Trochidae Rafinesque, 1815(see Rafinesque 1815; see also the remarks by Bouchet et al. (2017: 250-251).
Original spelling. Turbinidopsidae Nicolas, 1899. Remarks. The name is not based on a genus-group name. It was established as a series within Tanganikidae Nourry, 1898 (see Nicolas 1899). The name is interpreted here as descriptive alluding to the resemblance of species from Lake Tanganyika to the Turbinidae Rafinesque, 1815 (see Rafinesque 1815: 144).

Part 2. Genus-group names
In the genus-group a total of 57 available names are included in the catalogue. Of the available names in the genus-group, 11 are invalid for nomenclatural reasons.
The genus-group name Tanalia Gray, 1847, type species Nerita aculeata Gmelin, 1791 by original designation, which has often been used as a genus-group name in Paludomidae Stoliczka, 1868, is not included in the nomenclator because its type species belongs to the Neritidae Rafinesque, 1815 (Neiber and Glaubrecht 2019a).
Remarks. Gender: feminine. Hirtia as used by Van Damme and Pickford (2003) is an incorrect subsequent spelling.
Remarks. Gender: feminine, see Remarks under Hilacantha Ancey, 1886. Unjustified emendation of Hilacantha Ancey, 1886 as evidenced by the derivation of the name by Bourguignat (1890: 126, footnote on that page), see also Martens (1891: 7-8). The earlier use of Hylacantha by Bourguignat (1888: 23-24, pl. 9) is here interpreted as an incorrect subsequent spelling of Hilacantha Ancey, 1886 as there is no evidence of an intentional change of the name. Tylacanaeha (Nicolas 1899: 512) is here also regarded as an incorrect subsequent spelling, although hardly recognizable anymore, of Hylacantha Bourguignat, 1890. The work of Nicolas (1899) contains numerous typesetting errors and apparently that author was aware of the name Tiphobia introduced by Smith (1880a) and its replacement name Hilacantha introduced by Ancey (1886) in the emendated form Hylacantha by Bourguignat (1890) as suggested by the statement "L'énumeration se continue par un genre des plus caracteristques, Tiphobia ou Tylacanaeha, qui par l'opercule est une Melanidae, tandis que l'absence d'épiderme l'en sépare, d'eû une grande indecision pour cette espèce" (Nicolas 1899: 512) in combi-nation with the synonymous usage of the names Hylacantha longirostris and Tiphobia longirostris (Nicolas 1899: 515).
Type species. Lithoglyphus neritinoides Smith, 1880, by original designation. Remarks. Gender: feminine. The name Stanleya neritoides as used by Bourguignat (1885Bourguignat ( , 1888 has to be interpreted as an incorrect subsequent spelling of the name of the nominal taxon Lithoglyphus neritinoides Smith, 1880 and should not be regarded (as e.g. by Ancey 1906b: 254 or Baker 1923 as a newly introduced name by Bourguignat (1888) (although the author attributed the taxon to himself in that publication) because Bourguignat (1885: 87, footnote) explicitly referred to Smith (1880: 426) and Smith (1881: 287) and also used the misspelled name Lithoglyphus neritoides in the synonymy when first using the name Stanleya neritoides. Therefore, Stanleya is here interpreted as not having been introduced again by Bourguignat (1888) and the fixation of Stanleya neritoides with author Bourguignat (1888) by Baker (1923: 174) is not valid.
Type species. Tanalia stomatodon Benson, 1862, by monotypy. Remarks. Gender: masculine (Code Art. 30.1.2), from Greek στόμα, neuter (= mouth) and ὀδών, masculine (= tooth). The name Stomatodon Benson, 1862 was conditionally proposed by Benson (1862: 414). Benson (1862: 414) stated: "Unfortunately, all the specimens were deficient in the operculum, which, when examined, may possibly authorize its transfer to a new genus, in which case the specific name may fairly be employed to designate it." And further below on the page and on the next page: "Stomatodon seems partly to supply one of the absent links, inasmuch as its operculum must necessarily be provided with a basal projection, while its constriction is also likely to be unguiculate." Stomatodon Benson, 1862 is, however, a junior homonym of Stomatodon Seeley, 1861 (see Seeley 1861: 293; Gastropoda, Ringiculidae Philippi, 1853, see Philippi 1853. It has been replaced by Odontochasma Tomlin, 1930(see Tomlin 1930: 23 and Neiber and Glaubrecht 2019a.

Vinundu Michel, 2004
Original reference. Michel (2004: 2, 4 Michel & Todd, 2003in West et al. (2003 is unavailable because the work includes a disclaimer stating "this publication is not deemed to be valid for formal taxonomic/nomenclatural purposes". The name was also used by Van Damme and Pickford (2003: 2) as a nomen nudum because no diagnosis, definition, or description was given.

Part 3. Species-group names
In the species-group a total of 499 names are included, of which 463 are available but 21 are invalid for nomenclatural reasons.
Remarks. The species was classified as Cleopatra cylindrica (Adam, 1957) by Van Damme and Pickford (2003). This name is a secondary junior homonym of Cleopatra cridlandi cylindrica Mandahl-Barth, 1954 and because no junior synonyms exist, the new replacement name Cleopatra adami nom. nov. is introduced here (see also Remarks under Viviparus cylindricus Adam, 1957). The replacement name is chosen in honor of William Adam, the former Director of the Malacology Department of the Royal Belgian Institute of Natural Sciences, who was the first to describe the species (Adam 1957).
Type stratum. Not given.
Remarks. Attributed to L. Germain, but J. Roger alone was responsible for the description.
Remarks. Attributed to V. Giraud by Bourguignat (1885a: 73), but Bourguignat alone was responsible for the description. Therefore, Bourguignat is the author of the name (Code Art. 50.1).
Type locality. "Sur les rives nord et surtout occidentales du lac" [Democratic Republic of the Congo, northern and western shores of Lake Tanganyika].
Type locality. "Côtes occidentales et méridionales du lac" [Democratic Republic of the Congo/Zambia, western and southern shores of Lake Tanganyika].

Hauttecoeuria charmetanti Bourguignat, 1888
Original reference. Bourguignat (1888: 19, pl. 7, figs 15-17 Remarks. Nevill (1884) distinguished between 'subspecies', 'var.', and 'subvar.' Not every subvariety was referred to a subspecies or variety, however, but for example on p. 288 subvarieties were directly associated with a species without intermittent subspecies or variety. It seems that Nevill (1884) expressed by using the different terms to what extent a named taxon differed from the nominal species, with subspecies differing considerably, varieties slightly, and subvarieties only very slightly. Therefore, subspecific and varietal names are regarded here as subspecific names in accordance with the Code (Art. 45.6.4). The names denoted by 'subvar.' are, however, regarded as infrasubspecific. Made available by Kobelt in Pfeffer and Kobelt (1887: 422)
Type locality. "The Rungwa Sleeping Sickness Concentration Area, lies about eighty miles to the north-west of the present shore of the lake" [Tanzania, Rungwa].

Paludomus clavata Reeve, 1852
Original reference. Reeve (1852: 129 (Olivier, 1804) and attributed to Troschel who introduced Cleopatra as a genus-group name in 1857. Treated by Frauenfeld (1864: 588) as an available name, but not in a way that satisfies it to be adopted (Code, Glossary), i.e. not satisfying the Code (Art. 12.1). Therefore, the name was not made available under the Code (Art. 11.6.1) by Frauenfeld (1864).
Type locality. "Sur les rives nord et surtout occidentales du lac" [Democratic Republic of the Congo, northern and western shores of Lake Tanganyika].
Type locality. "Sur les rives nord et surtout occidentales du lac" [Democratic Republic of the Congo, northern and western shores of Lake Tanganyika].
Remarks. The species was classified as Cleopatra cylindrica (Adam, 1957) by Van Damme and Pickford (2003) in their revision of Albertine Rift Valley freshwater Cerithioidea. By that taxonomic act, the taxon becomes a secondary junior homonym of the nominal taxon Cleopatra cridlandi cylindrica Mandahl-Barth, 1954 and as no junior synonyms exist, the new replacement name Cleopatra adami nom. nov. is proposed here (see also Remarks under that name above).

Paludomus conica dihiriensis Nevill, 1884
Original reference. Nevill (1884: 288 Adam, 1959 is a nomen nudum is incorrect because W. Adam in Adam and Lepersonne (1959: 34-38) correctly introduced it, although not explicitly selecting a holotype and a type locality. Furthermore, the type selection of Van Damme and Pickford (2003: 49-50) does not constitute a valid lectotype designation in accordance with the Code Art. 74.7.1 because the term lectotype or an exact translation of that term was used not by these authors but only the term type and holotype. To avoid further confusion and nomenclatural instability, we here select the specimen proposed as type/holotype by Van Damme and Pickford (2003), i.e. the specimen figured by Adam (1959: pl. 5, fig. 1a), leg. J. Lepersonne, Lep. 464 III (M.C. 2382) from Nyamavi no 68, camp Nyamavi, as lectotype of Cleopatra dubia Adam, 1959.
Original spelling. Hauttecoeuria Duveyieri Germain, 1905. Remarks. Unjustified emendation of Hauttecoeuria duveyrieriana Bourguignat, 1885. Germain (1905a changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885, 1888). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Original spelling. Spekia Duveyieri Germain, 1906. Remarks. Unjustified emendation of Spekia duveyrieriana Bourguignat, 1885. Germain (1906 changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Remarks. The name was attributed to V. Giraud by Bourguignat (1885a: 79) but Bourguignat alone was responsible for the description and is therefore the author of the name alone (Code 50.1).
Remarks. Mentioned in the synonymy of Anceya (Anceya) giraudi as a manuscript name of Dupuis; unavailable (Code Art. 11.6).
Remarks. The name was attributed to V. Giraud by Bourguignat (1885a: 81) but Bourguignat alone was responsible for the description and therefore is the author of the name (Code Art. 50.1). Germain, 1906 Original reference. Germain (1906: 579).
Type locality. Not given.
Type locality. "De la côte occidentale" [Democratic Republic of the Congo, western shore of Lake Tanganyika].
Type locality. Not given.
Original spelling. Syrnolopsis Grandidieri Germain, 1905. Remarks. Unjustified emendation of Syrnolopsis grandidieriana Bourguignat, 1885. Germain (1905a changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885Bourguignat ( , 1888. The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Original spelling. Spekia Grandidieri Germain, 1906. Remarks. Unjustified emendation of Spekia grandidieriana Bourguignat, 1885. Germain (1906 changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Type locality. "Dans le sable d'une plage au nord de Mlilo" [Democratic Republic of the Congo, on a beach north of Mlilo (= Moliro), Lake Tanganyika].

Paludomus laevis Layard, 1855
Original reference. Layard (1855: 89). Original spelling. Paludomus laevis Layard, 1855. Type locality. "Ceylon, in slow-running streams on the northern side of the mountain zone extending into the flat country beyond Anarajahpoora [and] in a paddy field in the south of the island, near the village of Heneratgodde" [Sri Lanka].
Remarks. The name was attributed to V. Giraud by Bourguignat (1885a: 78), but Bourguignat alone was responsible for the description and therefore is the author of the name (Code Art 50.1).
Remarks. The name was attributed to V. Giraud by Bourguignat (1885a: 85), but Bourguignat alone was responsible for the description and therefore is the author of the name (Code Art. 50.1).
Original spelling. Hauttecoeuria Locardi Germain, 1905. Bourguignat, 1888. Germain (1905a changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885, 1888). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Original spelling. Paramelania Locardi Germain, 1905. Remarks. Unjustified emendation of Paramelania locardiana Bourguignat, 1885. Germain (1905a changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885, 1888). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Original spelling. Hauttecoeuria Maunoiri Germain, 1905. Bourguignat, 1885. Germain (1905a changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885, 1888). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Remarks. Unjustified emendation of Hauttecoeuria milneedwardsiana Bourguignat, 1885. Germain (1905a) changed the suffix of several species-group names ending in -iana or -ana introduced by Bourguignat (1885, 1888). The names thus formed are to be treated as emendations of the original names (Code Art. 33.2.1) and are available and have as unjustified emendations their own author and date (Code Art. 33.2.2).
Type locality. "Sur les plages de la partie septentrionale du lac" [Democratic Republic of the Congo/Burundi, northern shores of Lake Tanganyika].
Type locality. "In a brook about ten miles from Puerto Princesa in the Island of Palawan, Philippine Archipelago" [Philippines].
Remarks. Brot (1891: 17-18) used both, the spelling P. palawanicus and P. palawanica in the original description. The grammatical gender of the genus-group name Paludomus is feminine. Therefore, Paludomus palawanica Brot, 1891 is the correct original spelling.
Remarks. Published as a manuscript name under the unavailable name (introduced without description, definition, or indication) Paludomus microsculpta; unavailable (Code Art. 11.6).
Remarks. Cleopatra bulimoides var tschadiensis as used by Kobelt (1909) is an incorrect subsequent spelling.
Type locality. "In Travancor (p. 219) and more specifically: in streams traversing the plains between Trevandrum and the foot of the Aghastyamali hill (p. 220)" [India, Kerala].
Remarks. Attributed to Parreyss; published in the synonymy of Bithynia bulimoides (Olivier, 1804). Also mentioned by Martens (1865: 203) in the synonymy of Paludina (Cleopatra) bulimoides. Treated as an available name, Paludina trifasciata, of a taxon by Paetel (1873: 64). However, the name was not adopted, i.e. the unavailable name was not used as the valid name of a taxon in a way which establishes it as a new name with its own authorship and date in accordance with the Code (Art. 12.1), and therefore was not made available under the Code (Art. 11.6.1) by Paetel (1873).
Type locality. "Lake Tanganyika in Central Africa".