A nomenclator of extant and fossil taxa of the Melanopsidae (Gastropoda, Cerithioidea)

Abstract This nomenclator provides details on all published names in the family-, genus-, and species-group, as well as for a few infrasubspecific names introduced for, or attributed to, the family Melanopsidae. It includes nomenclaturally valid names, as well as junior homonyms, junior objective synonyms, nomina nuda, common incorrect subsequent spellings, and as far as possible discussion on the current status in taxonomy. The catalogue encompasses three family-group names, 79 genus-group names, and 1381 species-group names. All of them are given in their original combination and spelling (except mandatory corrections requested by the Code), along with their original source. For each family- and genus-group name, the original classification and the type genus and type species, respectively, are given. Data provided for species-group taxa are type locality, type horizon (for fossil taxa), and type specimens, as far as available.


Introduction
The family Melanopsidae (Caenogastropoda: Cerithioidea) is one of the most diverse groups of non-marine gastropods in Earth history (Strong et al. 2008). Today, the family occurs in southern to eastern Europe, northern Africa, and the Middle East, as well as New Zealand and some south Pacific islands (Bănărescu 1990, Glaubrecht 1993, 1996, Altaba 1998, Strong et al. 2008. Its record dates back at least into the late Cretaceous of Europe (Bandel 2000, Neubauer et al. 2016a. While the fossil record of New Zealand species is fairly restricted, the record for Europe, northern Africa and the Middle East yielded many hundreds of species-group taxa. Early species were pure brackish-water dwellers, which they remained until the early Miocene, when they started to conquer freshwater (Glaubrecht 1996, Neubauer et al. 2016a). This change in life style coincided with a series of adaptive radiations in the middle and late Miocene, producing hundreds of species, and forming the basis for the modern clades and present diversity and distribution (Neubauer et al. 2016a).
The first detailed listing of names of living "Melanidae" -at that time the Melanopsidae were considered a subfamily of that group -was published by Brot (1874Brot ( -1879. Soon later, Bourguignat (1884) followed with his "Histoire des mélaniens du système européen", in which he described numerous new species, but few of them are currently still accepted. Pallary (1926a, b) provided a list of all published names of fossil and recent Melanopsidae, yet without any comments. Wenz (1929) summarized all Cenozoic (Paleocene-Pleistocene) names and gave extensive synonymy lists. Besides, there are papers on the fossil and recent Melanopsidae of Italy (Pantanelli 1886b) and Spain (Pallary 1924, Azpeitia Moros 1929. In addi-tion, Pallary (1916aPallary ( , b, 1920Pallary ( , 1925 provided a detailed account of nomenclatural ocene species of the Stomatopsinae Stache, 1889(genera Stomatopsis Stache in Sandberger, 1871, Stomatopsella Stache, 1889, and Megastomopsis Stache, 1889 which was considered a subfamily of Melanopsidae by Bouchet & Rocroi (2005) but probably does not belong there; species introduced in the genus Coptostylus Sandberger, 1872, which has occasionally been classified within the Melanopsidae, but is currently assigned to the Thiaridae (Pacaud and Le Renard 1995: 156).
Basic sources used for the present assembly were the catalogues of Brot (1874Brot ( -1879, Bourguignat (1884), Pallary (1926a, b) and Wenz (1929), as well as internet resources such as the Biodiversity Heritage Library (http://biodiversitylibrary.org/), the Global Names Index (http://gni.globalnames.org/) and the Worldwide Mollusc Species Data Base (http:// www.bagniliggia.it/WMSD/WMSDhome.htm). However, many names listed in those online repositories are obvious misspellings deriving from automatic digitization procedures that have not been critically reviewed (compare discussion in Haszprunar 2014). Such errors are not included here because they do not come from the published literature. Some of the names currently listed in those databases could neither be traced back to any publication, nor could they be linked to an evident misspelling. These names are on purpose excluded from the present paper to avoid introducing potential misspellings or nomina nuda into the literature. Nevertheless, I will be grateful to receive information on names presently not included here, as well as corrections, explanations and additions.
Some of the publications consulted for this work display problematic cases, regarding their actual publication dates or the nomenclatural or taxonomic concepts applied therein. These issues require careful examinations, which are provided below.

Melanopsis species introduced by André Etienne Justin Pascal Joseph Francois d'Audebard de Férussac
Férussac described several new species of Melanopsis in his "Histoire naturelle générale et particulière des mollusques terrestres et fluviatiles", as well as in the "Monographie des espèces vivantes et fossiles du genre mélanopside". The problem is to determine the exact publication dates of those works and, therefore, when and in which work which species was described.
The "Histoire naturelle" is an extremely comprehensive work, containing two volumes of text and 210 plates, which were published between March 1819 and August 1851 in 35 "livraisons". It has been subject of intensive debates regarding the exact publication dates -for details see Kennard (1942a-c). The work was commenced by Férussac, who issued the first 28 livraisons between 1819 and 1832. After his death in 1836, Gérard Paul Deshayes finished the work by adding another seven livraisons between 1839 and 1851, including new explanations for the plates. The work includes two plates of "Mélanopsides fossiles". The first plate was published in livraison 15 on 13 April 1822, but its captions came along with livraison 17, which was published on 2 November 1822 (Kennard 1842b: 106). The second plate appeared in livraison 21 on 27 September 1823 including its captions (Kennard 1842a: 16).
To increase confusion, there is a disagreement between the original captions provided by Férussac and the ones supplied by Deshayes with livraison 29 in 1839 (Table  1; see also Kennard 1842b: 106). Deshayes made several corrections and synonymizations and even introduced two new names in the altered captions for plate 2, i.e., Melanopsis subcarinata and Melanopsis nodosa var. longa. While the original captions for plate 2 are available to me, I could not find the captions for plate 1. Both plates were also published par for par in Férussac's (1823) monograph on the Melanopsidae. The captions for plate 2 are the same in both works (except minor differences regarding the details provided), so we can assume the same for plate 1. However, the exact wording of the original captions of 1822 needs to be seen in the future, in order to finally ascertain the availability of the associated names. A different problem appears for plate 2. While the year of publication of the species illustrated there is undoubtedly 1823, the original publication itself is uncertain -either the livraison 21 of the "Histoire naturelle" [27 September 1823] or the "Monographie" [exact date unknown]. This matter could unfortunately not be solved.

Validity of names mentioned in Férussac's and Deshayes' works
Apart from the uncertainties detailed above, the inconsistent formatting of Férussac's (1823) monograph led later authors to adopt names as valid that were certainly not intended as such by Férussac himself. He introduced several new Melanopsis species and varieties, latter of which he denoted by Greek letters followed by Latin descriptions. Quite often, he used the same Latin words to characterize different varieties and apparently did not mean to introduce available names (e.g., Férussac 1823: 154). New names (or new combinations) were always denoted by him with "nobis". Table 2 summarizes the available species-group names and unnamed varieties introduced and described by Férussac (1823).
The variety M. buccinoidea var. "γ) Antiquua" [sic] (Férussac 1823: 149) poses a special case. It appeared first in the legend of plate 1 of the "Mélanopsides fossiles" in the "Histoire naturelle" but -if the format was the same as in the monograph -not in italic font as are the other names. In the monograph, it is the only term on the subspecific level Férussac gave in small caps, a format he otherwise used only for vernacular names of species. Because of this and the fact that he did not add "nobis" it is unlikely that Férussac wanted to introduce a new name; rather, he may have intended to indicate its status as a fossil (Latin antiqua = old). If it had been introduced as a distinct taxon, the name would have been invalid as an objective junior synonym of Melanopsis fusiformis Sowerby, 1822, which Férussac listed in synonymy.
Férussac (1822) also introduced two additional names (inflata, elongata) in the captions of plate 1 of the "Mélanopsides fossiles", ranked below var. γ ["Antiquua"]. In the monograph (1823) they are clearly marked with "nobis". If "antiquua" [= fusiformis] was considered as a distinct taxon, these names would be of infrasubspecfic rank, which is not governed by the Code. Both names would have nonetheless become Table 1. Férussac's original figure legends of the two plates of "Mélanopsides fossiles" (1822/1823) in the "Histoire naturelle", compared with the altered legends supplied by Deshayes in 1839 when finalizing the work. Note that Férussac's original figure legends for plate 1 are unavailable to me and thus based on the legends for plate 7 in Férussac (1823), which are supposed to be identical (see text for details).

Names introduced by Jean-Pierre Sylvestre de Grateloup in 1828 and 1838
A similar problem as for Férussac's monograph appears in works by Grateloup (1828Grateloup ( , 1838. He labelled varieties with Latin letters, occasionally followed by Latin terms (e.g., "olivula") and Latin descriptions and sometimes marked with "Nob.". In his 1828-work he attributed the Latin terms minor and major to varieties of two different species. Apparently, most of these terms were meant as keywords rather than real names. This becomes obvious also from Grateloup (1838), where he introduced the new name Melanopsis gibbosula for specimens treated as "var. b. minor" in Grateloup (1828).
In order to bring stability to the problem, I propose to use only the Latin terms marked by Grateloup with "Nob." as available names.

The taxonomic concepts applied by Jules René Bourguignat, Paul Pallary, Jean-Marie Pérès and Ferdinand Starmühlner
The works by Pallary between 1853 and extended the list of melanopsid names enormously ( Figure 1). Both authors applied very detailed taxonomic concepts and introduced dozens of species and varieties for even minor morphological deviations or differences in shell color. This procedure artificially inflated present-day melanopsid diversity, and few of those names are actually used today (Glaubrecht 2011). Nonetheless, most of the names are nomenclaturally available.
Unfortunately, Bourguignat and Pallary did not foresee that subspecies (as well as forma and variety names published before before 1961, Art. 10.2) would all become included by the Code in the species group, with the Principle of Homonymy applying throughout. Both malacologists introduced many varieties, such as minor and major, often several times for different species within the very same work. Pallary apparently considered some of the variety names he introduced as self-explanatory (e.g., Latin minor means "small") and left them undescribed (see, e.g., Pallary 1904: 37). In order to avoid inflating the present catalogus with anyway unavailable and taxonomically hardly useful names, I chose not to include these nomina nuda.
Finally, Pérès (1938aPérès ( , b, 1939Pérès ( , 1946 and Starmühlner (1970Starmühlner ( , 1973 applied elaborate systems to categorize morphologies and coloration patterns of recent melanopsids. Pérès introduced several "formae" and "modes" that he obviously did not use as valid names but rather as descriptive terms to fit existing species into his morphological concept (see, e.g., Pérès 1939: 135-137, Pérès 1946. He even assigned "types" to his morpho-units (Pérès 1939: 137). Although hardly any of his new names have any meaning in nomenclature or taxonomy, the available names are included and discussed herein. Starmühlner (1970Starmühlner ( , 1973 in a similar way classified New Caledonian specimens of Melanopsis frustulum according to form and size of the shell ("forma"), type and height of the spire ("modus") and coloration ("coloratus"). None of the names used by Starmühlner are available in nomenclature because they were introduced after 1961 and are of infrasubspecific rank (Art. 10.2). They are not included in this catalogus.

Note on the information provided
The catalogue lists all names in alphabetical order in the original spelling and combination, with the necessary amendments required by the Code. The status of taxa that are invalid, unavailable or unresolved is denoted in square brackets after the taxon name; those without status declaration are available and nomenclaturally valid, irrespective of their taxonomic status. The first description, or alternatively the basis of record for unavailable names, is always indicated. Taxa solely found as fossils are marked by a dagger. Taxon authorities attributed to a person other than one of the authors in the original source are only accepted as such if there is clear evidence that the description derived from that person (Art. 50.1.1). In such cases, the notation is given in accordance with Recommendation 51E. Information on type locality, type horizon (for fossils only) and type specimens are indicated as far as available. The exact spelling or phrase (given in quotes) provided for the type locality in the original source is denoted, along with an English translation if required. If the localities have been indicated indirectly (e.g., "same as for the previous species"), the phrase is given in square brackets. Old locality names have been matched with today's geographic names as far as possible, mostly using the GeoNames geographical database (http://www.geonames.org/v3/). Places that could not be found on the map or where the matched name is uncertain are marked by a postposed question mark in the translation.
Note that Wenz (1929) in his Fossilium Catalogus quite often misstated type localities, which he indicated in spaced letters. For taxa where no type has been fixed and several localities have been mentioned in the original publication, he apparently preferred localities mentioned first or those indicated for illustrated specimens. After Art. 76.1 of the Code, in absence of a holotype or lectotype the type locality encompasses the localities of all syntypes. Illustrated specimens do not automatically qualify as holotypes or have any priority in this respect.
Information on type horizons follows the most recent age classifications found in the literature (e.g., Neubauer et al. 2015a for the Neogene and Quaternary freshwater deposits of Europe). As far as possible, the present taxonomic status of a taxon is indicated, i.e., whether it has been considered a junior synonym by later authors. Data on type specimens are provided for all names where information is available, particularly for those published after 1999, when a type fixation became mandatory (Art. 16.4). If not indicated otherwise, the information provided regards the storage of the holotype. For names published by Spiridion Brusina and Petar S. Pavlović the inventory catalogues of the Croatian Natural History Museum in Zagreb (Milan et al. 1974) and the Natural History Museum in Belgrade (Milošević 1962) formed the basis. Note, however, that Milan et al. (1974) often wrongly indicated holotypes from syntype series, apparently considering illustrated specimens preferable to the rest of the material (see also discussion in Neubauer et al. 2016b). Milošević (1962) provided inventory numbers of the specimens illustrated by Pavlović; neither of both applied a type concept.

Nomenclatural notes
The following list of species-group names comprises several recurrent nomenclatural issues. In order to save space, to avoid multiple elaborate repetitions of the same rules, and to prevent that the reader needs to consult the Code constantly, I refer in text to two nomenclatural notes which are defined as follows: Note 1: Because of the Principle of Coordination (Art. 46), homonymy in the speciesgroup does not depend on a taxon's original rank in the species-group. This also encompasses variety and forma names published before 1961 (Art. 10.2). Many authors have been unaware that species names can constitute junior homonyms of subspecies, variety or forma names. Only in the case of simultaneously published names the taxon of higher rank takes precedence (Art. 24.1). Note 2: Several names of Melanopsidae first occurred in synonymy lists of other names (see, e.g., Brot 1874Brot -1879. According to Art. 11.6 of the Code, "a name which when first published in an available work was treated as a junior synonym of a name then used as valid is not thereby made available". Some of the names, however, have been made indeed available following the provisions of Art. 11.6.1: "However, if such a name published as a junior synonym had been treated before 1961 as an available name and either adopted as the name of a taxon or treated as a senior homonym, it is made available thereby but dates from its first publication as a synonym [...]". † Melanopsis aegyptiaca Blanckenhorn, 1901 Original source. Blanckenhorn 1901: 414, pl Types. Papp (1955: pl. 20, fig. 9) designated a specimen from the type material of Neumayr (1880a) as lectotype, but did not indicate whether it had been illustrated by Neumayr or not. A comparison of the images in both works was inconclusive. However, since it is likely that Neumayr (1880a) used the material reported in 1876 for the description, this designation is considered to be based on the original material and is thus valid. The specimen is stored in the Institute of Paleontology, University of Vienna; no number indicated.
Remarks. The name became available from Neumayr (1876), where he briefly described the species. He nevertheless described the species as "new" in Neumayr (1880a: 126), providing a detailed description and illustrations (pl. 6, figs 13-17; note that the plate is inserted at the end of the previous article of the same volume). † Melanopsis affinis Handmann, 1882 Original source. Handmann 1882: 558.
Remarks. The name often appears as "M. affinis Férussac" in the literature (e.g., Wenz 1929Wenz : 2653Esu and Girotti 1975: 251). It was first mentioned in Férussac (1814) as nomen nudum for a specimen from the Euphrates River. Later, Férussac (1823) listed the name in synonymy of M. nodosa Férussac, 1822. Unlike Pallary (1916) claimed and followed by Wenz (1929), affinis is not to be preferred over nodosa, because at that time affinis was not an available name -it has never been described, indicated or illustrated (as, for example, wrongly presumed by Esu and Girotti 1975) and so does not fulfill the requirements of availability (Art. 11 and 12; see also Art. 11.6). Pallary (1916) was the first to affiliate the name with an illustration, namely the one of M. nodosa provided by Férussac, 1823 (pl. 7, fig. 13), and made the name thereby available. The illustrated specimen is, however, the holotype (by monotypy) of M. nodosa Férussac, 1822 (Férussac used the same plates in both of his works and thus Pallary actually referred to the specimen illustrated as M. nodosa in the "Histoire naturelle"; see also introduction and Table 1 for details of Férussac's publications). Consequently, Melanopsis affinis Pallary, 1916 is a junior objective synonym of M. nodosa. Moreover, the name is a junior homonym of M. affinis Handmann, 1882. Rossmässler, 1839 Original source. Rossmässler 1838Rossmässler -1844. Type locality. "Aus der Ringelsza bei Laibach" [Ringelsza (?) brook near Ljubljana], Slovenia.
Remarks. Based on an "in schedis" name from Hazay.
Remarks. Nomen nudum, found only in species lists by Paetel (1888) and Pallary (1926a). Perhaps confused with "Bulimus bondeensis" from the same locality, which Gassies (1871: 203)  Remarks. In the old literature the name appears frequently as "M. bonellii Sismonda". The name is not available from Sismonda (1847: 55), however, who solely referred to a misidentified name (M. carinata non Sowerby) in the unpublished museum catalogue of Bonelli (1827) but did not supply a description. It appears as a nomen nudum also in d'Orbigny (1852: 28) and Doderlein (1863: 16). Hörnes (1856: 595) mentioned the name along with two other melanopsids from Italy and gave a collective description for all of them (translated from German: "all individuals are smaller as the Viennese [M. martiniana], as well as more elongate, but they have the characteristic marginal bulge and the typical keel below the suture"). Following Art. 12.1, every name [...] must be accompanied by a description or a definition of the taxon that it denotes [...], which is not the case for the mentioning of M. bonellii in Hörnes (1856) because the description given there refers to a group of taxa and not a single taxon. Manzoni (1870) made the name available by providing an illustration (Art. 12.2.7).
Types. Museum of Palaeontology and Geology of the University of Athens; no number indicated (Papp 1955).
Remarks. Justified emendation of M. cosmanni Pallary, 1916by Neubauer et al. (2014a. Original source. Olujić 1999: 20, 48, pl. 1, fig. 5. Type horizon. Langhian, middle Miocene. Type locality. It is unclear from the original work in which of the studied localities/sections along the valleys of the Sutina, Batarelov and Vojskava rivers (4 km W of Sinj) the taxon occurred and in which not, Croatia.
Types. Milan et al. (1974: 89) indicated a holotype, but it is uncertain whether the specimen actually derives from the original type series and whether it was the only specimen Brusina had at hand. The specimen is stored in the Croatian Natural History Museum, Zagreb, coll. no. 2988-634. Remarks. Introduced for M. costata sensu Neumayr, 1969, non Olivier, 1804. Wenz (1929 considered all records of "M. costata" from the Pliocene of Slavonia to represent the same species and synonymized them with M. abbreviata cosmanni. Obviously, he was unaware that M. croatica Brusina, 1884 is the first available name for them (see also discussion of M. pseudocostata Oppenheim, 1890 Gassies, 1870. Wenz (1929: 2764 considered the variety as a junior synonym of Melanopsis kleini Kurr, 1856. "Melanopsis praemorsa f. curta" mentioned in Pérès (1939) [unavailable] Locality. "Station 119. Aïn Attig. Source près de la route de Rabat à Casablanca à 13 kilomètres de Rabat" [station 119 at Ain Attig. A spring near the road from Rabat to Casablanca, 13 km from Rabat], Morocco.
Types. Milan et al. (1974: 89) stated that only one of the specimens illustrated by Brusina (1902: pl. 29, figs 19-20) has been preserved, which they designated as lectotype. The specimen is stored in the Croatian Natural History Museum, Zagreb, coll. no Remarks. The species epithet is a noun in apposition and needs not to agree in gender with the generic name (Art. 31.2.1). The name "daphne" as mentioned in Neumayr (1869: 369) and Hoernes (1876: 16) is an incorrect subsequent spelling. Wenz (1929Wenz ( : 2772 considered the taxon as a junior synonym of Melanopsis longa Deshayes in Férussac, 1839.
Remarks. There is considerable uncertainty about the correct authority and spelling of this species in the literature. It was first mentioned and validly described in the year 1821 as "Melanopsis Daudebartii" in an article in the Bulletin des Sciences, par la Société philomatique de Paris. Constant Prevost was often considered to be the author of this article, but from the title and text it is obvious that the article is an "Extrait" of a talk given by Prevost earlier and summarized by an anonymous author. According to Art. 50.2 and Recommendation 51D, the correct citation should be M. daudebartii [Prevost], 1821. The names "daudebarti", "audebarti" or "audebardi", each occurring multiple times in the literature, are incorrect subsequent spellings. Currently, the species is classified within the genus Microcolpia (see also Neubauer et al. 2014d: 126 Remarks. The name was also marked as new taxon by Brusina (1902: vii). It is uncertain whether Brusina really intended to introduce a new subspecies or actually referred to the species of Penecke (1884), which he had ranked as subspecies of M. croatica earlier (Brusina 1897 Brusina, 1902, non Klika, 1891. Jekelius (1944 considered this species as a junior synonym of M. sturii Fuchs, 1873. Pallary, 1936 Original source. Pallary 1936: 55, pl. 3, fig. 3. Type locality. "Dans les canaux de l'Aguedal, à Marrakech" [in the channels of Aguedal in Marrakech], Morocco. Pallary, 1939 Original source. Pallary 1939: 85, textfig. 11, pl. 6 Remarks. Heller et al. (1999: 56) considered the species as a junior synonym of Melanopsis buccinoidea (Olivier, 1801). In Heller et al. (2005: 248) in turn it is treated as a junior synonym of Melanopsis saulcyi Bourguignat, 1853.

Melanopsis (Sistaniana) drangianensis Starobogatov & Izzatullaev, 1985
Original source. Starobogatov and Izzatullaev 1985: 34, fig. 4 Remarks. The name first appeared in 1822 on the captions for plate 1 of the "Mélanopsides fossiles" in Férussac's "Histoire naturelle" (see also introduction for details). While Férussac (1823) included also recent specimens under that name in his monograph on the Melanopsidae, the only specimen illustrated in 1822 in the "Histoire naturelle" was a fossil one from the Miocene of France. This fact remained widely unknown to biologists and paleontologists alike. Only few malacologists, such as Grateloup (1828Grateloup ( , 1838Grateloup ( , 1840, used the name correctly and attributed it to the French fossils. Consequently, none of the specimens referred to as M. dufourii in the biological literature actually corresponds to the real M. dufourii. Very likely, some of the other melanopsids from Dax (see, e.g., Grateloup 1838, d'Orbigny 1852, Pallary 1916 are unrecognized junior synonyms of this species. The names "dufouri" as mentioned by numerous authors and databases and "dufourei" as given by Morelet (1853: 297) and Bourguignat (1884: 114)  Remarks. Wenz (1929Wenz ( : 2637 considered the taxon as a junior synonym of Faunus cerithiformis (Watelet, 1851) (Pachychilidae).

Melanopsis duveyrieri Bourguignat in Letourneux & Bourguignat, 1887
Original source. Letourneux  Remarks. Based on material from the Rhône Basin, not Italy as claimed by Wenz (1929); probably the taxon was mixed up by Italian authors cited by Wenz (1929). Harzhauser et al. (2015: 9) considered the taxon as a junior synonym of Melanopsis narzolina d'Archiac in Viquesnel, 1846. Pallary, 1918 Original source.  (1854), to whom Pallary referred, because Rossmässler did not use the term "elatior" to denote a species-group taxon but only cited Férussac's (1823) description of an unnamed variety of M. dufourii. Pallary briefly described the taxon (by indicating its size) and therefore made the name available.
Remarks. Unavailable according to Art. 13.1, because it lacks a verbal description or definition. Heller et al. (2005: 244) considered it as a junior synonym of Melanopsis costata (Olivier, 1804). Remarks. Introduced within the var. γ, to which Férussac attributed the term "antiquua", which was probably not intended as species-group name (see introduction for a detailed discussion of the names introduced by Férussac 1823). Note that for figs 6 and 7 of pl. 7 in Férussac (1823), which is the same as pl. 1 of the "Mélanopsides fossiles" in Férussac (1819-1851), Pallary (1916) introduced two new names, i.e., M. depressa and M. sparnacensis.
Type locality. Not explicitly stated but probably the same as for the species ("L'Oued Sous, au pont des Aït Melloul, sur la route d'Agadir à Tiznit, à 14 kil. S. O. d'Agadir" [in the Oued Sous, at the bridge of Ait Melloul, at the road from Agadir to Tiznit, 14 km southwest of Agadir], Morocco).
Type locality. "Wady Um Bagkek, between Sebbeh and Jebel Usdum, at the south-west corner of the Dead Sea", Israel.
Remarks. The name was first mentioned by Brot (1862: 63) as "etrusca Villa (v. min)" from the locality "Toscane", without description or bibliographical reference. Brot listed it under the name "Melanopsis Dufourii Fér.", for which a bibliographical reference (to "Rossm. Icon. 835-839") and locality "Espagne" was given. Since etrusca was neither described by Brot nor by Rossmässler, the name etrusca is not available from this case (Art. 12.1). Brot (1879: 434) listed the name ("Melanopsis etrusca Villa MSS.") as synonym of Melanopsis Dufourii var. β, which he introduced there with a short Latin description. On p. 435 Brot even associated etrusca with an illustration of a specimen from Toscana (pl. 47, fig. 3). However, the name was explicitly referred to as synonym of the (unnamed) variety β, which is not a valid name, and thus the requirements of Art. 11.6 (and therefore 12.1) are not met (see Note 2). Westerlund (1886: 128) used the name as "[Melanopsis lorcana] var. etrusca Villa ap. Brot (Cat. syst. Melan. 1862)" and provided a description and thus made it available as species-group taxon. Since he referred to Brot's catalogue, the type locality comprises both his and Brot's records. The authority of Melanopsis etrusca remains with Westerlund. Art. 11.6.1 does not apply here as Brot (1879) had listed the name as synonym of "Var. β", which is not a valid name (see Note 2). † Melanopsis eulimopsis Brusina, 1902 Original source. Brusina 1902: pl. 5, figs 42-44. Type horizon. Middle Pannonian, late Miocene. Type locality. "Kúp", Hungary.
Férussac (1823), who was the first to combine the species with Melanopsis, listed the records by Walch, Martini and Gmelin but apparently did not consider "fossile" a valid name and introduced "Melanopsis martiniana" as new name. That name is permanently invalid as it is an objective synonym (see also Pallary 1916: 81).

Melanopsis germaini
Type locality. "Sources du Nahr es Sine, au Sud de Lattaquié, sur la route de Beyrouth" [Nahr as Sinn, south of Latakia, at the road to Beirut], Syria.
Type horizon. Langhian, middle Miocene. Type locality. It is unclear from the original work in which of the studied localities/sections along the valleys of the Sutina, Batarelov and Vojskava rivers (4 km W of Sinj) the taxon occurred and in which not, Croatia.

Melanopsis heliophila Bourguignat, 1884
Original source. Remarks. Introduced for M. costata sensu Kobelt, 1880, non Olivier, 1804. Bourguignat attributed the authority to Letourneux, but there is no evidence that the description really derived from that author. Heller et al. (2005: 243) listed the species as a junior synonym of Melanopsis costata (Olivier, 1804) (Turkey). Alive in the lower Orontes (Turkey/Syria), as well as in the Karasu, a northern tributary of the Lake Anuk (Turkey), and in the Sājūr river near Aleppo (Syria)].
Remarks. Junior homonym of M. hoernesi Sandberger, 1872 (note that latter is a junior objective synonym of M. subbuccinoides and thus invalid). Heller et al. (2005: 244) considered the species as a junior synonym of Melanopsis costata (Olivier, 1804).
Type locality. "Dans la source et la rivière de la Moulouiah, à l'ouest de Lalla-Maghnia" [in the source and the river Moulouya, west of Maghnia], Morocco or Algeria.

Melanopsis (Sistaniana) lutensis Starobogatov & Izzatullaev, 1985
Original source. Starobogatov and Izzatullaev 1985 Remarks. Pallary (1916) erroneously treated "magna" as available name attributed to Férussac (1823), who solely used the word as descriptive term (Latin "big"), which he applied to many of his varieties. The name became nevertheless available from Pallary (1916) who associated the name with an illustration in Férussac (1823: pl. 7, fig.  16). The illustrated specimen is, however, the holotype (by monotypy) of M. dufourii Férussac, 1822 (Férussac used the same plates in both of his works and thus Pallary actually referred to the specimen illustrated as M. dufourii in the "Histoire naturelle"; see also introduction and Table 1 for details of Férussac's publications). Therefore, Melanopsis magna Pallary, 1916 is a junior objective synonym of M. dufourii.
Type horizon. Plio-Pleistocene. Type locality. "Coo: molto frequente in tutte e due le zone fossilifere" [Kos island: very common in both areas rich in fossils, i.e., between Antimáchei and Pýli and in the northeast of the island, near Ágios Fokás], Greece.
Type locality. Not explicitly stated but probably the same as for the species ("La Zousfana, à la hauteur de Figuig et à Beni Ounif" [Oued Zousfana, at the height of Figuig, and at Beni Ounif], Algeria).
Type locality. Not indicated, but probably in Morocco.
Remarks. Junior homonym of Melanopsis buccinoidea major Grateloup, 1838 (fig. 16). Note, moreover, that Morelet's synonymization is likely based on a wrong concept of M. dufourii, which is a fossil species described from the Miocene of France.
Remarks. Replacement name for M. brevis Parreyss in Mousson, 1854, non Sowerby, 1826 (see also Pallary 1916). Blanckenhorn (1897) denoted the authority as "Bourg.", but from the discussion it is clear that Blanckenhorn must be the author.
Type locality. Not indicated, but probably as for the species ("Dans la Save, audessous d'Agram" [in the Sava river below Zagreb], Croatia).

Melanopsis minutula
Types. Museum of Palaeontology and Geology of the University of Athens; no number indicated.
Types. Museum of Palaeontology and Geology of the University of Athens, coll. no. 1963/83. Remarks. The status of this species is unclear. First of all, Kühn did not at all refer to the senior homonym Melanopsis (Melanosteira) mitzopoulosi Papp, 1955 but clearly described the species as new. Neither did Papp (1979: 668), who must have been aware of the issue and who ranked it as a subspecies of M. longa Deshayes in Férussac, 1839. Moreover, Papp stated that Kühn's species was based on the material collected by Papp (1947) and considered the type locality mentioned by Kühn a mistake. Kühn (1963), however, did not refer to Papp's material at all.
Type locality. Not explicitly stated but probably the same as for the species ("Au cours de l'Euphrate et du Tigre" [from the Euphrates and Tigris rivers], Iraq).
Remarks. Dollfus mentioned the name as nomen nudum without explanation alongside the discussion of [M. costata] "Var. Biliottii Buk.". Apparently, he considered it a potential new name should biliottii be elevated to the species level. If not already a nomen nudum, M. monolithos would therefore be a junior objective synonym of M. biliottii Bukowski, 1892, which actually was introduced as a distinct species (see also Willmann 1981: 105 Remarks. Obviously unaware of the fact that variety names are available in nomenclature as species-group names, Kormos (1903) stated in the description of his new variety M. sikorai carinata that, if raised to species, he suggests "M. mucronifera" as name for the taxon. Both names were simultaneously published and are objective synonyms. Since M. carinata Kormos is a junior homonym of M. carinata Sowerby, 1826, M. mucronifera is the valid name of the taxon. Neubauer et al. (2014d) considered the taxon as a junior synonym of Microcolpia parreyssii sikorai (Brusina, 1903 (Brusina, 1903) by Neubauer et al. (2014d: 125). † Melanopsis doboi multifilosa Schréter, 1975 [invalid] Original source. Schréter 1975: 10, pl. 2, figs 11-12, pl. 3, fig. 13. Type horizon. Riss/Würm end to early Würm Ice Age, Pleistocene. Type locality. "Eger, az egri vár Zárkándy bástyájának vasúti átmetszése" [Eger, section at the railway at the Zarkandy bastion of the fortress Eger], Hungary.
Types. Milan et al. (1974: 95) indicated collection numbers for "syntypes" illustrated in Brusina (1902), but it is uncertain whether the specimens were part of the original type series. They are stored in the Croatian Natural History Museum, Zagreb, coll. no. 2533-179/1-4.
Remarks. Note that Bourguignat denoted the authority as " Bourguignat, 1879". Starobogatov et al. (1992  Remarks. The name first appeared in 1822 on the captions for plate 1 of the "Mélanopsides fossiles" in Férussac's "Histoire naturelle" (see also introduction for details). While Férussac (1823) included also recent specimens under that name in his monograph on the Melanopsidae, the only specimen illustrated in 1822 in the "Histoire naturelle" was a fossil one from the Villafranchian of Italy. This fact remained widely unknown to biologists and paleontologists alike. The consequence, however, is that probably none of the specimens referred to as M. nodosa in the biological literature (e.g., Pallary 1939, Glaubrecht 1993, 1996 or the IUCN Red List (Van Damme 2014), which reports it from Anatolia and the Middle East, actually correspond to the real M. nodosa.
Remarks. Bourguignat (1884) attributed the authority to himself, but even listed Gassies' record in synonymy. Apparently, he did not consider a variety as a valid name that can result in homonymy (see also the introduction).
Type horizon. Pliocene? Type locality. "D'un puits, profond de 25 mètres, situé dans la propriété Lamur, sur la rive gauche d'un ravinement creusé par les eaux pluviales dans une dépression de terrain, perpendiculaire au chemin d'Aïn Beïda" (p. 284) [from a well, 25 m deep, located in the village Lamur (near Oran) on the left bank of a ravine carved by rainwater in a depression in the ground, perpendicular to the path from 'Aïn el Beïda], Algeria.
Remarks. Pallary cited in the synonymy list his paper on the fauna of the "Berbérie", which appeared in the Journal de Conchyliologie. That work, however, was not published before March 1928. † Melanopsis orientalis Bukowski, 1892 Original source. Bukowski 1892: 249.

Melanopsis praemorsa f. perbrevis Pérès, 1946 [invalid]
Original source. Pérès 1946: 113. Type locality. Not indicated. Remarks. First mentioned as nomen nudum in Pérès (1939), but described and made available by Pérès (1946). In both works Pérès obviously used the name not as a separate taxon but rather as descriptive term to fit existing species into his morphological concept. The name is in fact a junior objective synonym of several different species. † Melanopsis percallosa Sandberger, 1875 Original source. Sandberger 1870Sandberger -1875 Type horizon. Middle Miocene. Type locality. "Locle", Switzerland.

Melanopsis douttei var. perornata
Remarks. Appeared first as a nomen nudum in Pavlović (1922: 49 (1974: 84) invalid: it is uncertain whether the specimen illustrated by Brusina (1902) was part of the original type series. The specimen is stored in the Croatian Natural History Museum, Zagreb, coll. no. 2484-130.
Type locality. "Rivières de Carniole; la Save à Steinbrück" [rivers in Carniola, a historical region that comprised parts of present-day Slovenia; in the Sava river near Zidani Most], Slovenia.
Type locality. "Dans une source tiède à proximité de la Moulouïa" [in a warm source close to the river Moulouya], Morocco.

Remarks.
Coptostylus is at present considered to belong in the family Thiaridae (Pacaud and Le Renard 1995: 156).
Remarks. The name "pseudaustriaca" as mentioned in Lueger (1980: 117) is an incorrect subsequent spelling. † Melanopsis pseudobesa Bandel, 2000 Original source. Remarks. Different concepts of this species were applied in the literature and created considerable confusion. Oppenheim introduced the name clearly for the misidentified record of M. costata sensu Neumayr in Neumayr & Paul, 1875 (p. 41), non Olivier, 1804 from Slavonia, despite referring about Greek material. Later, however, he considered the name as rectification also for M. costata sensu Fuchs, 1877 from Megara and sensu Tournouër, 1876 from Kos Island (Oppenheim 1891: 465). This fact remained unnoticed by Wenz (1929Wenz ( : 2808, who treated M. pseudocostata as replacement name of the latter two records (Fuchs, Tournouër) but not of Neumayr and Paul's species.
Wenz obviously considered all erroneous "M. costata" records from Slavonia to represent the same species and synonymized them with M. cosmanni, but was unaware that M. croatica Brusina, 1884 is the first available name for them. Although the four names (croatica, pseudocostata, cosmanni, permutabilis) likely refer to the very same species they are based on different specimens and thus no objective synonyms.
Remarks. Introduced to replace -for whatever reason -the variety "gracilis" mentioned by Locard (1883a: 288), making pulchella its junior objective synonym. M. gracilis Locard, 1883 is a junior homonym of M. gracilis Brusina, 1874, which makes pulchella the next available name. However, M. costata pulchella is a homonym of the simultaneously introduced M. seignetti pulchella Bourguignat, 1884 (see Note 1). The action of a First Reviser is required to determine which of both pulchella is to be treated as valid.  Brusina, 1902 Original source. Brusina 1902: pl. 5, figs 37-38. Type horizon. Middle Pannonian, late Miocene. Type locality. "Markuševec", Croatia.
Types. The syntypes are stored in the Croatian Natural History Museum, Zagreb; no number indicated (Milan et al. 1974: 96).
Remarks. Pallary (1939) indicated an illustration (" Fig. 4") of this taxon, which, however, does not appear as such in the plate captions. Junior homonym of Melanopsis seurati semilaevigata Pallary, 1928 and M. eximia semilaevigata Pallary, 1928 (simultaneously published; no priority fixed yet; see Note 1). Remarks. Introduced as infrasubspecific taxon ("mode"), which is not governed by the provisions of the Code. Moreover, the name is a nomen nudum. Pérès referred to an earlier publication of his (Pérès 1938a), claiming that he had described the taxon there already. In that work, however, he only mentioned a subfossil and a "forme modifiée" of M. magnifica Bourguignat, 1884. † Melanopsis lanzae senilis Olujić, 1999 Original source. Olujić 1999: 22, 50, pl. 3, figs 30-32. Type horizon. Langhian, middle Miocene. Type locality. It is unclear from the original work in which of the studied localities/sections along the valleys of the Sutina, Batarelov and Vojskava rivers (4 km W of Sinj) the taxon occurred and in which not, Croatia.
Type locality. Rossmässler, to whom Bourguignat referred, indicated the locality only as "Palästina" [Palestine]. As the species was explicitly introduced for Rossmässler's material only this is the type locality.
Type locality. "New Zealand, Bay of Islands, Waitanga Falls", New Zealand.
Type locality. "Dans les aqueducs de Séville et dans le Guadalquivir" [in the aqueducts of Sevilla and in the Guadalquivir river], Spain.
Remarks. Homonym of the simultaneously published name Melanopsis costata ventrosa Bourguignat, 1884. This case requires the action of a First Reviser.
Type locality. Not indicated, but apparently the specimens derive from somewhere along the Jordan river or the Sea of Galilee.
Types. Milan et al. (1974: 99) indicated a holotype, but it is uncertain whether the specimen was the only one Brusina had at hand (holotype by monotypy, Art. 73.1.2). The specimen is stored in the Croatian Natural History Museum, Zagreb, coll. no. 2531-177/1.
For some journals and books, the dates given on the cover or the first page of the article do not necessarily reflect the real publication dates. Examples are the Journal de Conchyliologie and the Bulletin de la Société Géologique de France, which occur many times in the reference list and where the dates deviated considerably (mostly regarding volumes published in the second half of the 19 th century). Publication dates of the Bulletin were usually supplied in the appendix; for the Journal de Conchyliologie, the dates are based on Fischer-Piette (1937).
The special case of Matheron (1842Matheron ( , 1843 should be discussed here briefly because it led to erroneous conclusions on homonymy issues and invalid replacement names. The two references refer to two, slightly different versions of the same publication by Philippe Matheron. The first version was published in the Répertoire des travaux de la Société de statistique de Marseille, vol. 6, pp. 81-341, the second as book in May 1843. In a note at the end of the book (p. 269), Matheron stated: "Circumstances beyond my control have delayed the release of the discussion above, the work that I presented at the Statistical Society of Marseille at the beginning of the year 1842 and part of which was already printed in August of the same year. The time that has elapsed since the original wording of the catalogue until the moment I write this note (in May 1843) allowed me to mention many species recently published by M. d'Orbigny and make my job more complete than it would have been if it had been published a year ago." Hence, we deal with two different publications that need to be cited separately. The melanopsids in question already appeared in the 1842-version and no changes were made for the 1843-version.