The evolutionary origin of the durophagous pelagic stingray ecomorph

Studies of the origin of evolutionary novelties (novel traits, feeding modes, behaviours, ecological niches, etc.) have considered a number of taxa experimenting with new body plans, allowing them to occupy new habitats and exploit new trophic resources. In the marine realm, colonization of pelagic environments by marine fishes occurred recurrently through time. Stingrays (Myliobatiformes) are a diverse clade of batoid fishes commonly known to possess venomous tail stings. Current hypotheses suggest that stingrays experimented with a transition from a benthic to a pelagic/benthopelagic habitat coupled with a transition from a non-durophagous diet to extreme durophagy. However, there is no study detailing macroevolutionary patterns to understand how and when habitat shift and feeding specialization arose along their evolutionary history. A new exquisitely preserved fossil stingray from the Eocene Konservat-Lagerstätte of Bolca (Italy) exhibits a unique mosaic of plesiomorphic features of the rajobenthic ecomorph, and derived traits of aquilopelagic taxa, that helps to clarify the evolutionary origin of durophagy and pelagic lifestyle in stingrays. A scenario of early evolution of the aquilopelagic ecomorph is proposed based on new data, and the possible adaptive meaning of the observed evolutionary changes is discussed. The body plan of †Dasyomyliobatis thomyorkei gen. et sp. nov. is intermediate between the rajobenthic and more derived aquilopelagic stingrays, supporting its stem phylogenetic position and the hypothesis that the aquilopelagic body plan arose in association with the evolution of durophagy and pelagic lifestyle from a benthic, soft-prey feeder ancestor.     

Eocene handfishes from Monte Bolca,with description of a new genus and species,and a phylogeny of the family Brachionichthyidae (Teleostei: Lophiiformes)

The family Brachionichthyidae, commonly known as the handfishes, is a small group of lophiiform fishes, the living species of which are restricted in distribution mostly to shallow temperate and subtropical waters of Tasmania and southern and eastern Australia. Despite their narrow present‐day distribution, and the extreme rarity of lophiiforms in the fossil record, handfishes are well represented in the Eocene of Monte Bolca, Italy. A revision of the known fossil material shows the presence of two fossil species in two monotypic genera, ?Histionotophorus and ? Orrichthys gen. nov. Diagnoses of the family Brachionichthyidae, the two fossil genera, as well as two recognized extant genera Brachionichthys and Sympterichthys are provided. An osteological analysis of ?Histionotophorus bassani revealed many new features as well as reinterpretations of some previously described skeletal parts. A phylogenetic analysis of brachionichthyid genera and representatives of the antennarioid families Antennariidae, Tetrabrachiidae, and Lophichthyidae, using 36 morphological characters, strongly supported monophyly of brachionichthyids and antennarioids, the former taxon representing the sister group of the other families of the latter. Within the Brachionichthyidae, the two extant genera Brachionichthys and Sympterichthys form a species pair, as do the extinct genera ?Histionotophorus and ? Orrichthys gen. nov. Biogeographical considerations suggest that the present geographical range of handfishes can be considered a residual distribution of a temporally and spatially dynamic range shift. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 621–647.     

AN EOCENE FROGFISH FROM MONTE BOLCA, ITALY: THE EARLIEST KNOWN SKELETAL RECORD FOR THE FAMILY

Abstract:  Eophryne  barbutii gen. et sp. nov., a new frogfish from the Ypresian of Monte Bolca (Italy), is described. This taxon is the oldest member of the family Antennariidae known to date based on articulated skeletal remains. Eophryne  barbutii gen. et sp. nov. possesses a unique combination of features that supports its recognition as a new genus of the family Antennariidae. It is characterized by the presence of tiny bifurcated dermal spinules on the skin between and on the surface of the second and third dorsal-fin spines, a naked illicium, outermost caudal-fin rays simple, caudal peduncle present, 11 dorsal-fin rays, and seven anal-fin rays. Eophryne gen. nov. shows a certain degree of phenetic resemblance to the extant genera Antennarius , Histrio and Nudiantennarius . Palaeoecological and evolutionary implications are also discussed.     

A small, “wader-like” bird from the Early Eocene of Messel (Germany)

A small bird with a shorebird-like beak and a very long hind toe is described from the Early Eocene of Messel in Germany. Vanolimicola longihallucis, gen. et sp. nov. is one of the few candidate species for a “wading bird” from Messel, that is, a bird that may have foraged along the shoreline of the ancient lake. A few features indicate a relationship to charadriiform birds and Jacanidae (jacanas) in particular, but charadriiform affinities are only weakly supported. In the preserved skeletal elements, V. longihallucis also resembles the taxon Songzia from the Early Eocene of China, which is considered to be a representative of the Ralloidea (rails and allies). A potential biogeographic significance of the new Messel fossil is strengthened by similar fossils from the North American Green River Formation.     

Phylogenetic implications of the some cranial features of the porcupine pufferfish <Emphasis Type="Italic">Pshekhadiodon</Emphasis> (Tetraodontiformes,Diodontidae) from the Eocene of the Northern Caucasus

The well-preserved skull of the diodontid fish Pshekhadiodon parini Bannikov et Tyler, 1997 from the Eocene of the North Caucasus has plesiomorphic conditions for four character that are apomorphic in all extant taxa, and the family Diodontidae is herein separated into two subfamilies, the Diodontinae for all of the extant taxa and the Pshekhadiodontinae for Pshekhadiodon and very tentatively for the three other Eocene taxa of the family, all from Monte Bolca, Italy.     

The taxonomy,chronostratigraphy and paleobiogeography of glyptosaurine lizards (Glyptosaurinae,Anguidae)

Glyptosaurine lizards (Glyptosaurinae, Anguidae) are an extinct group of heavily armored lizards known from North America, Europe and Asia. Glyptosaurine lizards, taxa that possess fully developed tuberculated dermal armor, appear to have been established in North America by late early Puercan time (To3). “Proxestops,” a taxon distinguished by a combination of vermiculate and tuberculated osteoderm sculpturing, is considered to be a non-glyptosaurine, a sister taxon of the Glyptosaurinae. Known from only fragmentary remains, its wide chronostratigraphic distribution suggests that “Proxestops” is a form genus that, in all probability, represents more than one taxon, that ranges from the middle Paleocene to the early Eocene of North America. Moreover, the taxa Odaxosaurus piger, Parodaxosaurus sanjuanensis and “Proxestops” are best considered “proto-glyptosaurines”. “Melanosaurins” and glyptosaurins were well-established by the early Eocene, especially in North America, and are here documented by their type species and chronostratigraphic levels. Both tribes are present in Europe (MP7), too, but the record is not as estensive as that of North America. The North American taxon Gaultia silvaticus (Wa0) is transitional between a “melanosaurin” and glyptosaurin. Because it lacks the well-defined hexagonal osteoderms that characterize the Glyptosaurini, it is removed from that group and considered to be a “melanosaurin”. The “melanosaurin” taxon “Xestops” savagei (Wa4–Wa6) cannot be referred to Xestops (Br2) based on non-corresponding elements and because superficial similarity does not justify assignment to this taxon. Arpadosaurus sepulchralis (Wa6?), whose holotype is a fragmentary right frontal, is considered a subjective junior synonym of A. gazinorum, based on minor differences in the epidermal scale pattern that probably represent individual variation. “Glyptosaurus” agmodon (Wa6?), based on a partial right maxilla, cannot be referred to Glyptosaurus (sensu stricto), and the material upon which this taxon is based bears strong resemblance to material identified as cf. “?Paraglyptosaurus” yatkolai (Wa5–Wa6). “Glyptosaurus” rhodinos (Wa5) is based on an incomplete parietal, and its reference to Glyptosaurus is considered problematic. Eoglyptosaurus donohoei (Wa7) is probably valid and is re-established here. Glyptosaurus (sensu stricto) is known solely from the middle Eocene (Br2) by G. sylvestris. Dimetoposaurus wyomingensis (Br3) is removed from Xestops vagans because its synonymy was based on superficial similarities. Helodermoides tuberculatus, the largest and last glyptosaurin (Ch3), is restricted to the Chadronian of North America. Only the “melanosaurin” Peltosaurus granulosus (Or2–Or3), which includes the species P. abbotti, seems to have crossed the Eocene-Oligocene boundary, and appears to be largely restricted to the Orellan, but extended into the Arikareean. European glyptosaurines are also represented by both glyptosaurins and “melanosaurins” early in the Eocene (MP7). Placosauriops-like “melanosaurins” are known from Dormaal (MP7), and the glyptosaurin taxon?Placosaurus ragei occurs at the same level. “Placosauriops abderhaldeni” has been identified from the Grube Messel (MP11), but this assignment remains dubious because the species has not been adequately diagnosed, and the holotype species is from the Geiseltal (MP13), which is some 4.5 million years younger. Placosauriops weigelti (MP13) is the only valid species of this genus. Paraxestops stehlini (MP14) is not referable to the North American taxon Xestops, and its relationship to Placosauriops has not been studied. The late Eocene glyptosaurins Placosaurus estesi (MP17) and P. rugosus (MP18) are the last glyptosaurines known from Europe and appear to have gone extinct at the Eocene-Oligocene boundary, casulties perhaps of the “Grande Coupure”. Asian glyptosaurines are known solely from one species, Stenoplacosaurus mongoliensis, from the middle Eocene (Sharamurunian) of China. Glyptosaurines most likely originated in North America, diversified by late Paleocene time, and rapidly spread across the North Atlantic into Europe by the early Eocene. Both “melanosaurins” and glyptosaurins took a foothold in Europe by the early Neustrian, but the glyptosaurins, aside from one occurrence (Dormaal, MP7), were conspicuously absent for most of Neustrian through early Robiacian time. In North America, glyptosaurins diversified during the early and middle Eocene, while in Europe small “melanosaurins” were a prominent part of the paleoherpetofauna, and glyptosaurins are unknown for most of the Neustrian through the Geiseltalian, in both the fossilferous Lagerstätten of Messel and Geiseltal. Stenoplacosaurus is the only known glyptosaurin glyptosaurine from Asia, and its abrupt appearance during the late Eocene suggests the possiblity of a Beringian dispersal from North America into Asia.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

A new genus and species of fossil scorpion (?Euscorpiidae) from the Early–Middle Eocene of Pesciara (Bolca,Italy)

Fossil scorpions are among the oldest terrestrial arthropods known from the fossil record. They have a worldwide distribution and a rich fossil record, especially for the Paleozoic. Fossil scorpions from Mesozoic and Cenozoic deposits are usually rare (except in amber-deposits). Here, we describe the only fossil scorpion from the Early to Middle Eocene Pesciara Lagerstätte in Italy. Eoeuscorpius ceratoi gen. et sp. nov. is probably a genus and species within the family Euscorpiidae. This may be the first fossil record of the Euscorpiidae, which are so far only known from four extant genera. Eoeuscorpius ceratoi gen. et sp. nov. was found in the “Lower Part” of the Pesciara Limestone, which is actually dated Late Ypresian stage (between 49.5 and 49.7 Ma). Besides a possible pseudoscorpion, the here-described fossil scorpion is the second arachnid species known from the Bolca Locality.     

New observations on the anatomy and paleobiology of the Eocene requiem shark 2Eogaleus bolcensis (Carcharhiniformes,Carcharhinidae) from Bolca Lagerstätte,Italy

Here we provide new information about the anatomy of the extinct shark 2Eogaleus bolcensis from the early Eocene Bolca Konservat-Lagerstätte. The combination of morphological and dental characters of the three known articulated specimens and a single isolated tooth collected from the Pesciara and Monte Postale sites (e.g., head bulky with rounded snout, pectoral fins of semiplesodic type, cutting-clutching dentition, dermal denticles shell-shaped and of drag-reducing type, caudal fin accounting for one fourth of the entire body length, 135–153 vertebrae) supports the assignment of 2Eogaleus to the family Carcharhinidae and allows us to confirm definitively its distinct taxonomic status from the sympatric carcharhiniform, the triakid 2Galeorhinus cuvieri. Moreover, the analysis of the dermal denticle morphology and the size and maturity age estimates of the studied specimens provide new paleobiological information about 2Eogaleus, suggesting a close association of this small top predator with the tropical shallow-water marine context hypothesized for the Bolca paleobiotopes.     

Ein Archaeotrogon (Aves: Archaeotrogonidae) aus dem Mittel-Eoz?n der Grube Messel (Hessen, Deutschland)?

Zusammenfassung Ein Vogel aus dem Mittel-Eozän der Grube Messel (Hessen, Deutschland) wird als neue Gattung und Art der Archaeotrogonidae Mourer-Chauviré 1980 beschrieben (incertae sedis). Die Exemplare aus Messel wären die ersten artikulierten Skelette dieser Familie.Hassiavis laticauda n. gen. n. sp. unterscheidet sich vor allem im Bau des Coracoids von der GattungArchaeotrogon Milne-Edwards 1892. Besonders bemerkenswert ist die ausgezeichnete Erhaltung der Flügel- und Schwanzbefiederung einiger Exemplare. Einige Schwanzfedern des Holotypus zeigen eine Querbänderung, welche möglicherweise auf die ursprüngliche Pigmentierung dieser Federn zurückzuführen ist. Die phylogenetische Stellung der Archaeotrogonidae wird diskutiert. Zur Zeit sind keine Synapomorphien bekannt, welche diese Familie mit einer der bestehenden Ordnungen verbinden.

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An Archaeotrogon (Aves: Archaeotrogonidae) from the Middle Eocene of the Grube Messel (Hessen, Germany)?

Summary A bird from the Middle Eocene of the Grube Messel (Hessen, Deutschland) is described as a new genus and species of the Archaeotrogonidae Mourer-Chauviré 1980 (incertae sedis). The specimens from Messel would be the first articulated skeletons of this family.Hassiavis laticauda n. gen. n. sp. is distinguished from the genusArchaeotrogon Milne-Edwards 1892 in the morphology of the coracoid. The excellent preservation of the wing feathers and those of the tail in several of the specimens is exceptional. Some tail feathers of the holotype are barred, which might be traced back to the original pigmentation of these feathers. The phylogenetic position of the Archaeotrogonidae is discussed. At present no synapomorphies are known, which could set up a closer relationship between this family and one of the existing orders.

     

Ontogeny and homology of the skeletal elements that form the sucking disc of remoras (Teleostei,Echeneoidei, Echeneidae)

The sucking disc of the sharksuckers of the family Echeneidae is one of the most remarkable and most highly modified skeletal structures among vertebrates. We studied the development of the sucking disc based on a series of larval, juvenile, and adult echeneids ranging from 9.3 mm to 175 mm standard length. We revisited the question of the homology of the different skeletal parts that form the disc using an ontogenetic approach. We compared the initial stages of development of the disc with early developmental stages of the spinous dorsal fin in a representative of the morphologically basal percomorph Morone. We demonstrate that the “interneural rays” of echeneids are homologous with the proximal‐middle radials of Morone and other teleosts and that the “intercalary bones” of sharksuckers are homologous with the distal radials of Morone and other teleosts. The “intercalary bones” or distal radials develop a pair of large wing‐like lateral extensions in echeneids, not present in this form in any other teleost. Finally the “pectinated lamellae” are homologous with the fin spines of Morone and other acanthomorphs. The main part of each pectinated lamella is formed by bilateral extensions of the base of the fin spine just above its proximal tip, each of which develops a row of spinous projections, or spinules, along its posterior margin. The number of rows and the number of spinules increase with size, and they become autogenous from the body of the lamellae. We also provide a historical review of previous studies on the homology of the echeneid sucking disc and demonstrate that the most recent hypotheses, published in 2002, 2005 and 2006, are erroneous. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

A new psittaciform bird from the London Clay (Lower Eocene) of England

A new psittaciform bird from the Lower Eocene (Ypresian) London Clay of England is described. This taxon, Pulchrapollia gracilis gen. et sp. nov., is assigned to the order Psittaciformes (parrots) on the basis of several distinctive structures of the tarsometatarsus, namely the trochlea for metatarsal III (trochlea metatarsi III) bearing a tubercle on its lateral side and the trochlea for metatarsal IV (trochlea metatarsi IV) completely retroverted (fully zygodactyl foot). Comparisons with other fossil and Recent taxa further support this conclusion. Cladistic analysis shows that Pulchrapollia is the sister-taxon of the single extant family within Psittaciformes, the Psittacidae. Palaeopsittacus georgei, a taxon previously described from the London Clay, is most likely based on some unassociated material and is regarded here as incertae sedis.     

The new family Caucasichthyidae (Pisces,Perciformes) from the Eocene of the North Caucasus

Caucasichthys kumaensis gen. et sp. nov., a representative of a new monotypic perciform family Caucasichthyidae, from the Middle Eocene (Bartonian, Kuma Horizon) of the North Caucasus (Gorny Luch locality) is described. The new family is characterized by elongated body, strong preopercular spine in adults, absence of supraneurals, large pelvic fins, long caudal peduncle, and anal fin longer at the base than soft dorsal fin. Scales vary from cycloid to spinoid on different parts of the body. Caucasichthys shares a number of apomorphic features with members of certain percoid families, most notably the Priacanthidae. However, because of its unique combination of features, the new family cannot be properly placed within any existing perciform suborder and it is placed incertae sedis among the Perciformes.     

New acrodont lizards (Lacertilia) from the Middle Eocene of southern Mongolia

The problematic lizard family Changjiangosauridae, representatives of which inhabited Asia in the Early Paleogene, is discussed. Six new species of this group, including Acrodontopsis robustus gen. et sp. nov., Agamimus gracilis gen. et sp. nov., Graminisaurus interruptus gen. et sp. nov., Khaichinsaurus reshetovi gen. et sp. nov., Lavatisaurus elegans gen. et sp. nov., and Lentisaurus giganteus gen. et sp. nov., from the Middle Eocene of the Khaichin Uul 2 locality (southern Gobi, Mongolia) are described. It is shown that Changjiangosauridae are probably related to the Late Cretaceous Isodontosauridae and recent Uromastycidae; independent development of a number of dental features in different lineages of Acrodonta (Iguania) is corroborated.     

A New Eocene Dasypodid with Caniniforms (Mammalia,Xenarthra, Cingulata) from Northwest Argentina

The aim of this work is to present a new genus and species of Dasypodidae from the Lumbrera Formation (“lower Lumbrera”), early-middle Eocene of Salta Province, northwest Argentina. The new taxon, documented by one specimen, consists of an incomplete skull and jaw with teeth housed in their alveoli, postcranial remains, and isolated osteoderms. Lumbreratherium oblitum, gen. et sp. nov., is characterized by heterodont dentition, with a caniniform as the first tooth, diastema between caniniform and first molariform, teeth with closed roots, and a peculiar morphology of the osteoderms. A phylogenetic analysis indicates that Lumbreratherium oblitum, gen. et sp. nov., and Pucatherium parvum belong to a monophyletic clade in a basal position within the Cingulata. The singularity of the morphological characteristics of these Paleogene armadillos of northwest Argentina reinforces the hypothesis of an intertropical origin of mammal clades different from those of the Paleogene in more austral regions of Argentina.     

A new basal raoellid artiodactyl (Mammalia) from the middle Eocene Subathu Group of Rajouri District,Jammu and Kashmir,northwest Himalaya,India

A new artiodactyl of moderate size, Rajouria gunnelli nov. gen., nov. sp., is described on the basis of several dentaries, maxillae and isolated teeth from the middle Eocene Subathu Group of the Kalakot area, Rajouri District, Jammu and Kashmir, India. Despite its general resemblance with the family Dichobunidae by the retention of a paraconid on m1-2 and a simple P4 where endocristids do not form an anterior loph, this taxon shares with Raoellidae two unambiguous characters: the presence of a hypoconid on p4, and an asymmetrical P4. The phylogenetic position of the new taxon within the Cetacea–Raoellidae clade is strongly supported by seven non ambiguous synapomorphies, among which a cristid obliqua on lower molars anteriorly pointing towards the postectoprotocristid, and a P3 with only two roots. The presence of a new basal raoellid in the middle Eocene Subathu Group sheds new light on the phylogeny and paleobiogeography of raoellid artiodactyls.     

New stoneflies (Insecta: Perlida = Plecoptera) from Eocene Rovno amber,Ukraine

A new genus, Rovnocapnia gen. nov., with new species R. atra sp. nov. and R. ambita sp. nov., from the family Capniidae, and a new genus and species Palaeoleuctra acuta gen. sp. nov., from the family Leuctridae, of the suborder Nemourina (Euholognatha), are described from Late Eocene Rovno amber. Familial assignment of one additional specimen (suborder Perlina, infraorder Perlomorpha (Systelognatha)), represented by a nymphal skin, is obscure due to its incomplete preservation.     

New Paleocene large cassiduline foraminifera (Cassidinidae fam. nov., Miscellanacea) from Egypt and its phylogeny

Larger foraminifera of the Late Paleocene–Early Eocene lower slope to toe-of-slope deposits of the southern Galala, Egypt, are described. Sulcocassidina nakkadyi n. gen. et sp. (base) and Vacuolicassidella dakhlensis n. gen. et sp. (top) represent a phylogenetic line related to the family Cassidinidae (new family), superfamily Miscellanacea (new superfamily) in the latest Paleocene (shallow benthic zone 4) of the southern Galala, Egypt. In addition, Oscucassidella cassis n. gen. et sp., Pellatispiroides youssefi n. gen. et sp. and Chordcassidella ainshamsiana n. gen. et sp. are described and illustrated. Genus: Pomerolina n. gen. (Type species: Alveolina meandrina Carter, 1861, is newly erected and it belongs to Cassidinidae. In this family, the chambers are initially arranged streptospirally, whereas in later stages they are planispirally enrolled. Lateral chamberlets (retral process) are separated from the laminae by a passage and a marginal crest is added during growth.     

Eolacertidae: a new extinct clade of lizards from the Palaeogene; with comments on the origin of the dominant European reptile group – Lacertidae

We describe a new lizard taxon, Stefanikia siderea gen. et sp. nov., from the early-middle Eocene locality of Messel in Germany based on a nearly complete skeleton, which we studied using μCT methods. It shares many characters with the Eocene taxon Eolacerta, which is broadly distributed in the Eocene of central and Western Europe, but is much smaller and shows several important anatomical differences. The new discovery sheds light on the paleodiversity of these lizards in the Eocene of Europe, and the new family name Eolacertidae is proposed to encompass Eolacerta and Stefanikia. The relationships of Eolacerta have been intractable. Our phylogenetic analyses confirm that Eolacertidae is a member of the clade Lacertiformes and provide strong support for a sister-group relationship to Lacertidae. In some places, skin impressions are preserved, displaying the body scalation. As such, the exquisitely preserved specimens of Eolacertidae from Messel provide new insight into the morphology and ecology of lizards on the stem of Lacertidae, Europe’s dominant group of living reptiles.

http://zoobank.org/urn:lsid:zoobank.org:pub:DFD01563-A913-4286-B64B-E0912474FD08