Citation: Neubauer TA, Harzhauser M, Kroh A, Georgopoulou E, Mandic O (2014) Replacement names and nomenclatural comments for problematic species-group names in Europe's Neogene freshwater Gastropoda. Part 2. ZooKeys 429: 13–46. doi: 10.3897/zookeys.429.7420
In the course of a new database project on Miocene to Recent freshwater gastropods of Europe, a great many of primary and secondary homonyms were revealed. Such nomenclatural issues need clarification in order to avoid misunderstandings and wrong statements about geographical distributions and temporal ranges. The following 16 new names are introduced to replace existing homonyms: Theodoxus militaris jurisicpolsakae nom. n., Viviparus stevanovici nom. n., Melanopsis haueri ripanjensis nom. n., Melanopsis wolfgangfischeri nom. n., Micromelania ramacanensis nom. n., Pseudamnicola welterschultesi nom. n., Muellerpalia haszprunari nom. n., Muellerpalia pseudovalvatoides nom. n., Lithoglyphus gozhiki nom. n., Valvata heidemariae willmanni nom. n., Radix macaleti nom. n., Gyraulus okrugljakensis nom. n., Gyraulus rasseri nom. n., Gyraulus vrapceanus nom. n., Planorbarius halavatsi nom. n., and Segmentina mosbachensis nom. n. Additionally, six cases of homonyms are discussed that are not replaced by new names, because they are considered junior synonyms.
Homonyms, synonyms, nomina nova, fossil freshwater gastropods
The latest estimate of living freshwater gastropod species involves about 4, 000 described and valid species world-wide (
As to the fossil part, there are several publications dealing explicitly with this problem. In four subsequent works
The newly established FreshGEN (Freshwater Gastropods of the European Neogene) database project, an initiative aimed at a pan-European reconstruction of the Neogene and Quaternary biodiversity of lacustrine gastropods, successively uncovered nomenclatural mistakes that have not yet been detected and/or revised. Following the first part of the resulting nomenclatural amendment (
The systematics follows
Theodoxus lutetianus Montfort, 1810 [currently considered as a synonym of Theodoxus fluviatilis (Linnaeus, 1758)]. Recent, Europe. Type by original designation (
In honor of Zlata Jurišić-Polšak (Croatian Natural History Museum), who contributed to our knowledge of Neogene Neritidae.
Malino, Croatia.
Late Pliocene to Early Pleistocene ("Paludina Beds").
Croatian Natural History Museum, coll. no. 9454.
Viviparus fluviorum Montfort, 1810 [currently considered as a synonym of Viviparus viviparus (Linnaeus, 1758)]. Recent, Northern Eurasia, Europe, Anatolia and Northern America. Type by original designation (
In honor of Petar M. Stevanović (Belgrade), who greatly contributed to our knowledge of the mollusc fauna and biostratigraphy of the Late Miocene of Serbia.
Kostolac opencast mine, Serbia.
Late Miocene to Early Pliocene (Late Pannonian, Late Portaferrian).
Natural History Museum, Belgrade, coll. no. 5683.
Since Paludina Férussac, 1812 is a junior objective synonym of Viviparus Montfort, 1810 (
Melania costata Olivier, 1804. Recent, Europe. Subsequent designation by
Named after the type locality.
Ripanj, Serbia.
Late Miocene (Early-Middle Pannonian;
Croatian Natural History Museum, Zagreb, coll. no. 2530-176/1-2 (
Obviously unaware of the fact that also subspecific or variety names can constitute homonyms,
In honor of Wolfgang Fischer (Vienna), who greatly contributed to nomenclature and taxonomy of fossil and Recent melanopsids.
Wittmannsdorf near Leobersdorf, Austria (
Late Miocene (Early Pannonian, Slavonian;
Geological Survey Austria, Vienna, no number indicated (
This taxon is a primary homonym of Melanopsis rugosa Matheron, 1842 (p. 293, pl. 37, fig. 11), a fossil species from SE France. Melanopsis rugosa Handmann, 1887 is a member of the complexly evolving Melanopsis impressa-species lineage in the Late Miocene Lake Pannon (
Additionally, there exists another primary homonym of Melanopsis rugosa, i.e. Melanopsis lanzaeana rugosa Brusina, 1897 from the Middle Miocene deposits of the Sinj Basin. It was synonymized with Melanopsis lanzaeana by
The taxonomic status of the Micromelaniinae is currently under discussion. The rank as subfamily follows
Micromelania cerithiopsis Brusina, 1874. Late Miocene, Croatia. Subsequent designation by
Named after the type locality.
Ripanj, Ramača hamlet (also read Ramaća), Serbia.
Late Miocene (Early-Middle Pannonian;
This is a classic case of a primary homonym requiring a replacement name according to
Paludina macrostoma Küster, 1853. Recent, Europe. Subsequent designation by
In honor of Francisco W. Welter-Schultes (University of Göttingen), a great expert for the living non-marine mollusks of Europe.
Megara, Greece.
Pliocene (
Natural History Museum Vienna, coll no. 1878/0020/0023 (designation by
This species is a primary homonym of Valvata minima Hislop, 1859 (p. 170, pl. 5, fig. 13) from the Tertiary of East India (see also
Planorbis bicincta Fuchs, 1870 in
In honor of Gerhard Haszprunar (Bavarian State Collection of Zoology Munich and Ludwig Maximilians University Munich), who summarized all existing names of living and fossil valvatids in a comprehensive nomenclator (
Tihany at Lake Balaton, Veszprém, Hungary.
Late Miocene (Late Pannonian, Transdanubian sensu
According to the inventory books of the Natural History Museum Vienna the material should be stored there, but despite great effort it could not be located.
This species is a primary homonym of the extant taxon Valvata tricarinata var. simplex Gould, 1841 (p. 226) from Massachusetts, USA. The American taxon was elevated to species level by
Already
The following subspecies have been introduced or ranked within Valvata simplex Fuchs, 1870:
1) Valvata bicincta Fuchs, 1870 [erroneously "bicinata" on p. 536; from captions and description there is no doubt about the correct name] from Tihany: It was considered a subspecies of Valvata simplex by
2) Valvata carinata Fuchs, 1870 (p. 536) from Tihany: It was considered as subspecies of Valvata simplex by
3) Valvata simplex öcsensis Soós, 1934 (p. 189) from Öcs:
4) Valvata octonaria Brusina, 1902 (pl. 13) from Tihany: It was also ranked as subspecies of Valvata simplex by
5) Valvata simplex polycincta Lőrenthey, 1906 (p. 167) from Tihany: It was synonymized with Valvata simplex octonaria by
6) Valvata simplex unicincta Lőrenthey, 1906 (p. 165) from Tihany (Fehérpart): The status of this taxon is doubtful. It was not mentioned by
To denote that it is despite its similar shape not a member of the genus Valvata.
Tihany at Lake Balaton, Veszprém, Hungary.
Late Miocene (Late Pannonian, Transdanubian sensu
According to the inventory books of the Natural History Museum Vienna the material should be stored there, but despite great effort it could not be located.
Up to now it has been overlooked by several authors, including ourselves (
Paludina naticoides Pfeiffer, 1828. Recent, Europe. Subsequent designation by
In honor of Piotr F. Gozhik (Kiev), who intensively studied the Neogene deposits of Ukraine and southern Russia.
Nizhniy Dnepr (= lower Dnieper), Ukraine.
Late Miocene (Early Maeotian, Oltenian).
Institute of Geological Sciences, National Academy of Sciences of Ukraine, Kiev, coll. no. 2174.
A classic case of a primary homonym. Probably as a result of prolonged publication times, Gozhik had no chance to become aware of this problem. However, the taxonomic status of Lithoglyphus maeoticus Papaianopol, 2006 from the Early Maeotian of the Dacian Basin is doubtful. It greatly resembles and might be a synonym of the Dacian species Lithoglyphus acutus Cobălcescu, 1883 (p. 145, pl. 14, fig. 10; see also
The here applied suprageneric systematics of Valvata follows
Valvata cristata Müller, 1774. Recent, Europe. Type by subsequent monotypy (
In honor of Rainer Willmann (University of Kiel), who intensively studied the Plio-Pleistocene deposits and freshwater mollusks of Greece.
Vokasia Valley 3 km SE of Kos City, Kos Island, Greece.
Early Pleistocene (Middle Irakli Formation).
Geological-Paleontological Institute, University of Kiel, no number indicated;
The species-group name bicarinata in combination with Valvata is preoccupied by the Recent species Valvata bicarinata Lea, 1841 from Pennsylvania, USA. The taxonomic separation from Valvata heidemariae Willmann, 1981 seems plausible, given the presence of a strong median keel on the upper whorl surface that is lacking in the nominal species.
Helix Auricularia Linnaeus, 1758. Recent, Europe. Type by original designation (for details see
In honor of Rodica Macaleț (Bucharest), who studied the mollusk fauna of the Dacian Basin.
Butuci near Sângeru, Prahova, Romania.
Latest Miocene to earliest Pliocene (Pontian, Portaferrian-Bosphorian).
Collection of the Geological Institute of Romania, coll. no. 19.546.
This species is a secondary homonym of Limnaea socialis von Zieten, 1832, of which the presently accepted and widely used combination is Radix socialis (e.g.,
Planorbis albus Müller, 1774. Recent, Europe. Subsequent designation by
Named after the type locality.
Okrugljak (today within the city limits of Zagreb), Croatia.
Late Miocene (Late Pannonian, Portaferrian;
Croatian Natural History Museum, Zagreb, coll. no. 2953-599/1 (
This species represents a primary homonym of Planorbis (Helisoma) clathratus Sandberger, 1880 from the Pleistocene of West Runton, Norfolk, United Kingdom. We follow
In honor of Michael W. Rasser (State Museum of Natural History Stuttgart), who studied the Gyraulus species flock of Lake Steinheim.
Orahovac (= Rahovec), Kosovo.
Early Pliocene (Late Dacian to Early Romanian;
Natural History Museum, Belgrade, coll. no. 1176 (
The name Planorbis discoideus as established by
Named after the type locality.
Vrapče (also read as Vrabče; today within the city limits of Zagreb), Croatia.
Late Miocene (Early Pannonian, Slavonian).
Croatian Natural History Museum, Zagreb, coll. no. 5195-360/2 (
The name Planorbis dubius was first used by
Helix cornea Linnaeus, 1758. Recent, Europe. Subsequent monotypy by
In honor of Gyula von Halaváts (Budapest), who greatly contributed to our knowledge of Pannonian mollusks.
Balatonfőkajár, Veszprém, Hungary.
Late Miocene (Late Pannonian, ?Transdanubian sensu
Hungarian Geological Institute, Budapest, coll. no. Pl. 121 (
As both taxa were introduced within Planorbis, the species described by Halaváts is a primary homonym. Both are today unambiguously assigned to the genus Planorbarius (for the Pannonian species see, e.g.,
Nautilus lacustris Lightfoot, 1786 [currently considered as a synonym of Segmentina nitida (Müller, 1774)]. Recent, Europe. Type by monotypy (
Named after the type locality.
Mosbach, Hessen, Germany.
Early Pleistocene.
No storage or types indicated.
The species name established by Sandberger is a primary homonym of Planorbis micromphalus Fuchs, 1870, although he was apparently aware of the existence of this name (compare
In the following, we present six cases of primary and secondary homonyms that seem not to be in use anymore (e.g., are unambiguously considered junior synonyms). We were unable to find any recommendation in the Code regarding the necessity of replacement names for disused junior homonyms. Following the intent expressed in Article 23.9.5, which seems to discourage the proposal of unnecessary replacement names, we choose not to introduce new names for these cases. In addition, the statuses of two taxa apparently constituting homonyms are discussed.
Gjurakovc (= Đurakovac), Kosovo.
Late Pliocene to Early Pleistocene (= "Levantin").
Collection de l'Institut de recherches géologiques et géophysiques de Belgrade, no number indicated.
Still we refrain from introducing a replacement name, because the taxonomic status of this subspecies is highly doubtful. It greatly resembles the nominal species Viviparus lomejkoi Pavlović, 1932 from Crmljan and Orahovac (like the type locality Gjurakovc in the Metohia Basin). The only difference is the stronger degree of whorl stepping, which is not documented by Pavlović's original description and illustrations. This is regarded to fall into intraspecific variability, why we suggest synonymizing Viviparus lomejkoi brevis with Viviparus lomejkoi. If, however, another author keeps both forms separate, a replacement name has to be introduced.
Puilor Valley, Buzău, Romania.
Early Pliocene (Late Dacian, Parscovian).
Institut de Géologie et Géophysique, Bucharest, coll. no. 17055.
Vivipara, as given by Neumayr in
No locus typicus given; occurs in Stegersbach, Litzelsdorf, Olbendorf, and Oberdorf in the Styrian Basin, Austria.
Late Miocene (Pannonian, Serbian, biozones E–F).
No storage or types indicated (material derived from Sauerzopf's private collections).
There are several issues with the name Melanopsis inflata. First, the name introduced by Sauerzopf definitely constitutes a primary homonym of Melanopsis pygmaea inflata Handmann, 1882.
The second problem regards the availability of Melanopsis inflata Handmann, 1882. This name was already introduced as subordinate taxon by
In the biological and palaeontological literature several species-group taxa were introduced as "Melanopsis elongata". The first mention traces back to
Consequently, all later introduced taxa also named "Melanopsis elongata" are primary homonyms of Melanopsis elongata Férussac, 1923. According issues are provided by
The names introduced by Locard, Jooss and Gillet and Marinescu have not yet undergone nomenclatural revision. Although primary homonyms are invalid, it is, however, not expedient to introduce new names for taxa that are not used anymore. This particularly regards Melanopsis narzolina elongata Locard, 1878 from the Late Miocene of Tersanne, which was apparently not used at all by subsequent authors and synonymized by
A more complicated case in terms of synonymy is presented by Melanopsis callosa elongata Jooss, 1911 from the Aquitanian of the Mainz Basin.
The remaining two homonyms are still in usage and thus require a more detailed assessment.
Ueken, Aargau, Switzerland.
Middle to Late Burdigalian ("Helvetian").
Paleontological Institute and Museum, University of Zurich, no number indicated.
Unlike the case of Melanopsis narzolina elongata Locard, 1878, this taxon was not synonymized by
Rădmănești, Romania.
Late Miocene (Late Pannonian, Transdanubian sensu
This case represents another primary homonym of Melanopsis elongata Férussac, 1823. Here some specific notes are necessary to elucidate the history of this taxon.
A part of the material of Melanopsis defensa defensa in
Helix tentaculata Linnaeus, 1758. Recent, Europe. Subsequent designation by
Bengeşti, Gorj, Romania.
Early Pliocene (Early Dacian, Getian).
Collection of the Geological Institute of Romania, coll. no. 18.906.
The genus-group name Bulimus Scopoli, 1777 was suppressed under Plenary Powers and placed on the Official Index of Rejected and Invalid Generic Names in Zoology by
The status of Bithynia socialis (Papaianopol & Macaleț 2006), however, is doubtful. The bithyniids of the Dacian Basin are quite well studied (e.g.,
Cyclostoma acutum Draparnaud, 1805. Recent, France. Type by monotypy.
No type locality indicated; occurs in several places in Frankfurt am Main, Germany.
Early Miocene (Aquitanian, upper Corbicula beds = Rüssingen Formation).
Only one of the syntypes is stored in the Senckenberg Research Institute and Natural History Museum, coll. no. SMF 245299/1 (pers. comm. R. Janssen, Frankfurt).
This tricky case requires a careful assessment of the original literature.
Șimleu Silvaniei (= Szilágy-Somlyó), Sălaj, Romania.
Late Miocene (Middle Pannonian, Serbian).
No storage or types indicated; probably stored in the Hungarian Geological Institute, Budapest.
Both involved taxa were originally combined with different genera, but have been attributed to Pseudamnicola in the second half of the 20th century. Lőrenthey's species was recombined with Amnicola by
Our sincere thanks go to Francisco W. Welter-Schultes (University of Göttingen) for sharing thoughts on nomenclatural issues. We thank Sonja Herzog-Gutsch (Natural History Museum Vienna) for assistance with literature research. We are grateful to Ronald Janssen (Senckenberg Research Institute and Natural History Museum, Frankfurt) and Imre Magyar (Hungarian Natural History Museum, Budapest) for providing information about type material. Michael Rasser (State Museum of Natural History, Stuttgart) is kindly thanked for his constructive review. This work was conducted within the project "Freshwater systems in the Neogene and Quaternary of Europe: Gastropod biodiversity, provinciality, and faunal gradients" financed by the Austrian Science Fund (FWF project no. P25365-B25).