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Concept: Nomen dubium


The split of our own clade from the Panini is undocumented in the fossil record. To fill this gap we investigated the dentognathic morphology of Graecopithecus freybergi from Pyrgos Vassilissis (Greece) and cf. Graecopithecus sp. from Azmaka (Bulgaria), using new ╬╝CT and 3D reconstructions of the two known specimens. Pyrgos Vassilissis and Azmaka are currently dated to the early Messinian at 7.175 Ma and 7.24 Ma. Mainly based on its external preservation and the previously vague dating, Graecopithecus is often referred to as nomen dubium. The examination of its previously unknown dental root and pulp canal morphology confirms the taxonomic distinction from the significantly older northern Greek hominine Ouranopithecus. Furthermore, it shows features that point to a possible phylogenetic affinity with hominins. G. freybergi uniquely shares p4 partial root fusion and a possible canine root reduction with this tribe and therefore, provides intriguing evidence of what could be the oldest known hominin.

Concepts: Phylogenetics, Hominidae, Greece, Miocene, Morphology, Homininae, Hominini, Nomen dubium


This study reports on a new ceratopsid, Spiclypeus shipporum gen et sp. nov., from the lower Coal Ridge Member of the Judith River Formation in Montana, USA, which dates to ~76 Ma (upper Campanian). The species is distinguished by rugose dorsal contacts on the premaxillae for the nasals, laterally projecting postorbital horncores, fully fused and anteriorly curled P1 and P2 epiparietals, and a posterodorsally projecting P3 epiparietal. The holotype specimen is also notable for its pathological left squamosal and humerus, which show varied signs of osteomyelitis and osteoarthritis. Although the postorbital horncores of Spiclypeus closely resemble those of the contemporaneous ‘Ceratops’, the horncores of both genera are nevertheless indistinguishable from those of some other horned dinosaurs, including Albertaceratops and Kosmoceratops; ‘Ceratops’ is therefore maintained as a nomen dubium. Cladistic analysis recovers Spiclypeus as the sister taxon to the clade Vagaceratops + Kosmoceratops, and appears transitional in the morphology of its epiparietals. The discovery of Spiclypeus adds to the poorly known dinosaur fauna of the Judith River Formation, and suggests faunal turnover within the formation.

Concepts: Cladistics, Clade, Cretaceous, Dinosaur, Taxon, Linnaean taxonomy, Ceratopsidae, Nomen dubium


Exosphaeromaamplicauda (Stimpson, 1857) from the west coast of North America is reviewed and redescribed and revealed to be a group of closely related species. A neotype is designated and the species redescribed based on the neotype and topotypic specimens. Exosphaeromaamplicauda is known only from the coast of California, at Marin, Sonoma and San Mateo Counties. Exosphaeromaaphrodita (Boone, 1923), type locality La Jolla, California and previously considered nomen dubium is taken out of synonymy and re-validated. A further three species: Exosphaeromapaydenae sp. n., Exosphaeromarussellhansoni sp. n., and Exosphaeromapentcheffi sp. n. are described herein. Sphaeromaoctonctum Richardson, 1899 is placed into junior synonymy with Exosphaeromaamplicauda. A key to the Pacific West Coast Exosphaeroma is provided.

Concepts: United States, California, North America, Los Angeles, San Diego, Zoological nomenclature, San Francisco, Nomen dubium


In 1941, Abel established Balaena belgica based on a series of fused cervical vertebrae and citing other cranial fragments from the late Neogene of the Antwerp harbor (northern Belgium). Later, Plisnier-Ladame & Quinet (1969) added a neurocranium and other skeletal remains from the same area to this species. Recently, the neurocranium was re-assigned to the genus Eubalaena thanks to newer phylogenetic analyses. Here, a new description is provided of materials previously assigned to “Balaena” belgica together with taxonomic revisions. Our work suggests that the cervical complex originally designated as the type of “Balaena” belgica is too poorly preserved to be used as such and is assigned to Balaenidae gen. et sp. indet., thus making “Balaena” belgica a nomen dubium. In addition to the neurocranium, the other remains consist in a fragment of maxilla assigned to Balaenidae gen. et sp. indet. and in a humerus assigned to Eubalaena sp. Discovered in the Kruisschans Sands Member of the Lillo Formation (3.2-2.8 Ma, Piacenzian, Late Pliocene), the neurocranium is designated as the holotype of the new species Eubalaena ianitrix. Our phylogenetic analysis supports a sister-group relationship of Eubalaena ianitrix and Eubalaena glacialis, and helps constraining the ages of origin for balaenid clades. Ecological and phylogenetic data suggest that Eubalaena ianitrix may represent the direct ancestor of Eubalaena glacialis, the latter having evolved through phyletic transformation including body size increase during the temperature decline of the Late Pliocene.

Concepts: Mammal, Phylogenetics, Cervical vertebrae, Pliocene, Cetacea, Right whale, Balaenidae, Nomen dubium


Two extremely-long necked elasmosaurids, AMNH 1495, holotype of Hydralmosaurus serpentinus, and AMNH 5835, previously referred to H. serpentinus, are here reviewed in detail. Unique features of the cervical vertebrae, which are only present on elasmosaurids from the Western Interior Seaway, are recognized based on these specimens and by comparison with penecontemporaneous taxa with biogeographic affinities. Phylogenetic analysis, bivariate graphic analysis of cervical vertebrae proportions, comparisons of different cervical vertebral types, paleobiogeographic distribution and study of the elasmosaurid axial evolution throughout the Cretaceous are here integrated. As a result, at least two separate lineages within the Elasmosauridae are identified by independently acquired extremely-long necks (over 60 cervical vertebrae). First, a still scarcely known lineage is so far represented by the lower Cenomanian Thalassomedon haningtoni, the Turonian Libonectes morgani and close relatives. A second lineage is here defined as a new clade, the Styxosaurinae, which groups the Campanian genera Terminonatator, Styxosaurus (=‘Hydralmosaurus’), Albertonectes and Elasmosaurus, the two latter forming a derived branch that includes the most extreme amniote necks known to date (more than 70 cervical vertebrae). Phylogenetic analysis supports AMNH 1495 and AMNH 5835 as being closely related to Styxosaurus snowii. Therefore, the species Styxosaurus browni is re-validated, while AMNH 1495 is here referred to Styxosaurus sp. This research also recognizes the ‘Cimoliasauridae’ (nomen dubium) as a paraphyletic group but informative of a plesiomorphic cervical vertebral morphology of elasmosaurids which was persistent throughout the whole Cretaceous and from whom aristonectines, styxosaurines and Thalassomedon and close relatives are derived. The genus Hydralmosaurus is recommended for being abandoned.

Concepts: Neck, Plesiosaur, Campanian, Plesiosauria, Nomen dubium, Elasmosauridae, Cretaceous plesiosaurs, Styxosaurus


The aphid genus Myzaphis van der Goot, 1913 from the tribe Macrosiphini is revised to include eight species. Apterous and alate viviparous females, known fundatrices and known sexual morphs (oviparous females and males) of Myzaphis bucktoni, M. juchnevitschae, M. rosarum, M. tianshanica and M. turanica are re-described and illustrated. Lectotype and paralectotypes of Myzaphis bucktoni and M. turanica are designated. The status of M. komatsubarae nomen dubium is discussed. Myzaphis avariolosa is regarded as a species belonging to the genus Ericaphis. Three new species: M. oezdemirae Kanturski & Barjadze sp. nov., M. tuatayae Kanturski & Barjadze sp. nov. from Turkey and M. rezwanii Kanturski & Barjadze sp. nov. from Iran are described and illustrated. Myzaphis bucktoni is recorded from Portugal for the first time. Diagnosis of the genus Myzaphis van der Goot, 1913 is redefined and a new genus Richardsaphis Kanturski & Barjadze gen. nov. is erected with the type species R. canadensis (Richards) comb. nov. Richardsaphis is for the first time recorded from the USA and hitherto unknown oviparous female and alate male are described and illustrated. Original keys to species of the genus Myzaphis and aphid genera of the tribe Macrosiphini with 2-2-2 first tarsal chaetotaxy are also provided.

Concepts: Male, Female, Insect, Sex, Genus, Aphid, Zoological nomenclature, Nomen dubium


The tardigrade class Mesotardigrada was erected on the basis of the description of Thermozodium esakii by Gilbert Rahm in 1937. In some characteristics, T. esakii is intermediate between members of the classes Eutardigrada and Heterotardigrada. The class Mesotardigrada is known only from Rahm’s published drawings of T. esakii; no voucher specimens are known, and subsequent attempts to collect it at the locus typicus have been unsuccessful. Among the possible explanations for this situation are that Rahm may have collected specimens of a more typical tardigrade, but misinterpreted what he saw. Alternatively, changes in habitat in the area may have led to the tardigrade’s extirpation. Perhaps T. esakii is a rare species, such that recent sampling efforts have been insufficient to rediscover it. Finally, Rahm’s 1937 description may be an attempt at deception. Until physical evidence of T. esakii is found, the species, and by extension the class Mesotardigrada, should be considered nomen dubium.

Concepts: Collected, Zoological nomenclature, Rare species, 2008 in film, Nomen dubium, Tardigrades, Nomen nudum, Mesotardigrada


A form of Plagioporus Stafford, 1904 is described from the intestine of three North American species of darters (Perciformes: Percidae) from River West Twin, Wisconsin, USA, that we consider to be conspecific with Plagioporus boleosomi (Pearse, 1924) Peters, 1957 based on similarities in the sucker ratio, extent of the forebody, shape and position of the testes, vitellarium distribution and terminal genitalia. Three new species of Plagioporus are described from the intestine of darters as follows: Plagioporus fonti n. sp. from Percina nigrofasciata Agassiz in Florida, USA, Plagioporus limus n. sp. from Etheostoma squamosum Distler in Arkansas, USA and Plagioporus aliffi n. sp. from Etheostoma blennioides newmanni Miller in Arkansas, USA. Morphologically Plagioporus fonti n. sp., Plagioporus limus n. sp. and Plagioporus aliffi n. sp. are most similar to one another and to P. boleosomi, Plagioporus lepomis Dobrovolny, 1939 and ‘P. etheostomae’, a nomen nudum for a species described from Etheostoma blennioides Rafinesque in Kentucky, USA, all of which are collectively distinguished from congeners in having a combination of confluent vitellarium in the post-testicular space and absence of vitelline follicles with their entire length distributed in the forebody. Plagioporus fonti n. sp., P. limus n. sp. and P. aliffi n. sp. are respectively distinguished from one another and their closest congeners in having the anterior extent of the vitellarium in the anterior half of forebody to slightly anterior to the ventral sucker as opposed to one approximately at the level of the posterior margin of the ventral sucker, possession of an excretory vesicle reaching the anterior testis as opposed to one only reaching the posterior testis and having a longer than wide oral sucker and a wider than long ventral sucker. A Bayesian inference (BI) analysis of partial 28S rDNA sequences was conducted using the three new species and 24 sequences of opecoelids retrieved from GenBank, including ten species of Plagioporus. Plagioporus aliffi n. sp., Plagioporus fonti n. sp. and P. boleosomi comprised a moderately supported sister group to a clade containing all species of Plagioporus except Plagioporus limus n. sp. and Plagioporus shawi (Mcintosh, 1939) Margolis, 1970. Plagioporus limus and in turn P. shawi were resolved as sister to all other congeners with high and moderate support, respectively.

Concepts: Testicle, Phylogenetics, Cladistics, Clade, Bayes' theorem, Percidae, Nomen dubium, Nomen nudum


Vespertilio oreias (generally known as Myotis oreias) has long been considered an endemic bat species to Singapore but its taxonomic status has been in doubt, and no specimens have been found since its description in 1840. Temminck formally described it based on a mounted skin (now in poor condition and accompanied by some skull fragments). The holotype was re-examined and we found it to be a composite, consisting of two separate individuals representing two distinct genera, the skin belonging to a Kerivoula whereas the skull fragments are of a Myotis. The mounted skin is accepted herewith as the name-bearing type, as the skull fragments were taken out after Temminck had published his description. Unfortunately, neither the skin nor the dental remains show enough anatomical details to identify the species unambiguously. Hence, the name Vespertilio oreias is considered a nomen dubium and the name oreias should be referred to the genus Kerivoula.

Concepts: Biology, Species, Binomial nomenclature, Genus, Zoological nomenclature, Botanical nomenclature, Nomen dubium, Holotype


Twenty-three new synonyms are introduced and one species is placed as nomen dubium in the conopid genera Conops, Myopa, Physocephala and Thecophora. Myopa fasciata is designated as the type species of Ischiodonta Lioy. Dipodium apiarium Bosc d'Antic is interpreted as an unrecognised species and therefore Dipodium Bosc d'Antic also has to be treated as an unrecognised genus. A key to the Western Palaearctic species of Physocephala is provided.

Concepts: Biology, Species, Genus, Order, Zoological nomenclature, Botanical nomenclature, Nomen dubium, Conopidae