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The plesiosaur specimen NHMUK 36184 from the Lower Jurassic of Whitby, Yorkshire and kept in the Natural History Museum of London, comprises an almost entire skeleton with nearly complete skull. It was described as one of two syntypes of Plesiosaurus homalospondylus by Owen, and selected as the lectotype by Lydekker. Extensive preparation of the skull has revealed it as one of the most complete and best-preserved Jurassic plesiosaurian skulls known, and its description adds much-needed data to our knowledge of the cranial osteology of the Plesiosauria. The three-dimensional preservation permits a relatively reliable reconstruction of its form. Microcleidus homalospondylus displays an interesting combination of cranial characters present in Jurassic plesiosauroids and Cretaceous Elasmosauridea. Its snout presents a very distinctive sculpture; the first pair of premaxillae teeth are extremely reduced; the frontal is partially overlain by the premaxillae, contacts the pineal foramen but does not contact the temporal fenestra; the jugal does not contact the orbit nor the temporal fenestra; the squamosal contacts the postorbital but not the maxilla and presents a bulb; the postfrontal contacts the posterolateral orbit margin; the anterior interpterygoid vacuity is absent; the pterygoids meet posterior to the posterior interpterygoid vacuities and are pierced by a foramen at this level; the quadrate ramus of the pterygoid presents a ventromedial flange; the parasphenoid is crested; the epipterygoid contacts the parietal; the paroccipital process is spatulate distally; the prootic presents an anteroventral process; the mandibular symphysis is keeled and bears four pairs of teeth. Microcleidus appears very similar to Hydrorion and Occitanosaurus, and the three taxa share a great number of plesiomorphic characters with basal plesiosaurians and pliosauroids.
Rhynchonellida is the stratigraphically oldest and phylogenetically most basal of the extant rhynchonelliform brachiopod orders, yet phylogenetic relationships among rhynchonellides are poorly known. The fourteen named rhynchonellide superfamilies (four of which have extant representatives) were defined primarily on the basis of features of the dorsal cardinalia, particularly crural morphology, but their homology and polarity have not been investigated rigorously. Superfamily monophyly is unclear, as is the evolution of several distinctive rhynchonellide morphological features, such as crura.
The purpose of this study is to investigate the phylogenetic relationships among extant rhynchonellide genera using skeletal characters, and to compare the results with the current classification, elucidating the evolution of morphological features in the process. We completed parsimony-based and Bayesian analyses using fifty-eight characters of the interior and exterior of the shell that vary among the nineteen extant genera. Our results are readily interpretable with respect to the classification, and indicate that Hemithiridoidea, Dimerelloidea, and (in some analyses) Pugnacoidea appear to be monophyletic. Species classified in Dimerelloidea and Pugnacoidea, and in certain cases Hemithiridoidea, each form derived subclades that evolve from within a paraphyletic Norelloidea at the base of each subclade. Raduliform crura appear to be the most basal, phylogenetically; five other crural morphologies evolve from the raduliform state. However, morphological characters currently uniting genera in rhynchonellide superfamilies are not clearly diagnostic and exhibit a relatively high degree of homoplasy overall, suggesting that consistency with the classification may be based on a false sense of confidence in rhynchonellide morphology to clearly elucidate evolutionary relationships. Published molecular phylogenetic hypotheses conflict with the morphological topologies, further supporting this possibility.
The evolutionary trends among diagnostic characters of Recent rhynchonellides appear to reflect successive juvenilization in adult morphology in several subclades, suggesting that heterochrony may have played an important role in the evolution of the group.
Two amphibian taxa are reported for the first time from the lower to middle Miocene shallow-marine Calvert Formation. These are Batrachosauroides aff. B. dissimulans (a large proteoid salamander) and cf. Notophthalmus robustus (a small newt). Four kinds of identifiable nonmarine turtles were reported previously from this formation. These are Bairdemys miocenica (a pleurodire), Kinosternon sp. (a musk turtle), Hesperotestudo ducateli and Hesperotestudo wilsoni (tortoises). Four additional taxa reported here are Chrysemys isoni n. sp. (a painted turtle), Trachemys sp. (a pond turtle), Floridemys hurdi n. sp. (a small tortoise), and Apalone lima (a softshell turtle). Most of these taxa are known from elsewhere in the Atlantic and Gulf Coastal Plains from New Jersey to Texas. The regional widespread occurrence of many of these taxa indicates that they represent a modestly diverse community of amphibians, fresh water turtles, and land turtles that were endemic to the Atlantic and Gulf coastal regions of the United States during the late early and early middle Miocene. Their pattern of distribution is similar to that of eastern American land mammals during this time interval, indicating that the Atlantic and Gulf coastal plains of the southeastern United States then lay within a single climatic zone that constituted a distinctive and long-lived faunal province throughout the early and middle Miocene (Hemingfordian through Barstovian land mammal ages).
Bathymodiolin mussels are a group of bivalves associated with deep-sea hydrothermal vents and other reducing deep-sea habitats, and they have a particularly rich early Cenozoic fossil record in western Washington State, U.S.A. Here we recognize six species from middle Eocene to latest Oligocene deep-water methane seep deposits in western Washington. Two of them are new: Vulcanidas? goederti from the middle Eocene Humptulips Formation and Bathymodiolus (sensu lato) satsopensis from the late Oligocene part of the Lincoln Creek Formation. Very similar to the latter but more elongate are specimens from the early Oligocene Jansen Creek Member of the Makah Formation and are identified as B. (s.l.) aff. satsopensis. Bathymodiolus (s.l.) inoueiAmano and Jenkins, 2011 is reported from the Lincoln Creek Formation. Idas? olympicusKiel and Goedert, 2007 was previously known from late Eocene to Oligocene whale and wood falls in western Washington and is here reported from Oligocene seep deposits of the Makah and Pysht Formations. Vulcanidas? goederti occurs at a seep deposit from a paleodepth possibly as great as 2000 m, suggesting that its living relative, Vulcanidas insolatusCosel and Marshall, 2010, which lives at depths of only 150–500 m, is derived from a deep-water ancestor. The bathymodiolins in western Washington indicate that the group originated at least in the middle Eocene and underwent a first diversification in the late Eocene to Oligocene. Early ontogenetic shells of all fossil species investigated so far, including the middle Eocene Vulcanidas? goederti, reflect planktotrophic larval development indicating that this developmental mode is an ancestral trait of bathymodiolins.
A new penaeoid shrimp collected from the Middle Triassic Member II of the Guanling Formation in the vicinity of the city of Luxi, Yunnan, southwest China, is a new species, Aeger luxii n. sp. The new species possesses prominent spinose third maxillipeds, which is one of the typical characteristics of Aeger. The new species differs from the type species, Aeger tipularius from the Jurassic Solnhofen Plattenkalk, in having a long, smooth rostrum with no subrostral spines. The new taxon increases the diversity of Chinese decapods, and further expands our knowledge of the phylogeny and evolution of the Mesozoic decapods. The find is the first complete specimen of Aeger in the Middle Triassic, and reveals a close biogeographic connection of the marine ecosystem between Eastern and Western Tethys.
Turtles have served as a model system for molecular divergence dating studies using fossil calibrations. However, because some parts of the fossil record of turtles are very well known, divergence age estimates from molecular phylogenies often do not differ greatly from those observed directly from the fossil record alone. Also, the phylogenetic position and age of turtle fossil calibrations used in previous studies have not been adequately justified. We provide the first explicitly justified minimum and soft maximum age constraints on 22 clades of turtles following best practice protocols. Using these data we undertook a Bayesian relaxed molecular clock analysis establishing a timescale for the evolution of crown Testudines that we exploit in attempting to address evolutionary questions that cannot be resolved with fossils alone. Some of these questions, such as whether the turtle crown originated in the Triassic or Jurassic, cannot be resolved by our analysis. However, our results generate novel age-of-origination estimates for clades within crown Testudines. Finally, we compare our fossil calibrations and posterior age estimates to those from other studies, revealing substantial differences in results and interpretation.
A set of 127 binary and multistate characters, weighted by the number of derived character states, degree of covariation, and level of homoplasy, was used in a cladistic analysis of type species representing 12 genera previously assigned to families Monticuliporidae and Mesotrypidae. The most parsimonious tree consisted of a 10-genus monophyletic crown group with the remaining two genera forming a basal paraphyletic stem group. The composition of the monticuliporid crown group is broadly similar to two earlier classifications while stem group membership matches the family Mesotrypidae. Phenetic groupings, based on overall morphological similarity, have memberships that are similar to those of clades but provide no means of determining the polarity of evolutionary relationships either within or between them. Finally, only the observed stratigraphic ranges of the type species of genera provide a statistically significant match with cladistic branching sequence, perhaps because current composite generic ranges reflect the mixing of species belonging to different genera. Based on cladogram topology, we propose the placement of all 12 genera into a single family Monticuliporidae.
The first two rove beetle fossils discovered from the Late Jurassic Talbragar Fish Bed in New South Wales, Australia are described and illustrated. Juroglypholoma talbragarense n. sp. is the second fossil record for one of the smallest and latest recognized staphylinid subfamily Glypholomatinae. The other staphylinid, Protachinus minor n. gen. n. sp., is an unusual member of extant subfamily Tachyporinae (tribe Tachyporini). It significantly retains several distinct features, including entire epistomal suture, and abdominal tergites III–VI each with a pair of basolateral ridges. The discovery of a new glypholomatine in Australia, together with recently reported one from the Middle Jurassic Daohugou biota of China, suggests the subfamily Glypholomatinae was probably much more widespread in the Jurassic than previously thought.
Fossil evidence of barnacle encrustation of vertebrate bones is reported from the middle Pleistocene Port Orford Formation of southern coastal Oregon. This material includes two associated thoracic vertebrae and a femur referable to the extinct sea lion Proterozetes ulysses that are encrusted by 1400 individual barnacles (cf. Hesperibalanus hesperius), and a scapula of Zalophus californianus with barnacle attachment scars. In areas, the encrusting barnacles exhibit a roughly bimodal size range, and small barnacles are observed directly encrusting other larger individuals. The size, probable age, and lifespan of extant Hesperibalanus hesperius indicates a minimum period of four to seven months of seafloor exposure between decomposition and burial, although this estimate must be longer because at least two colonization events are represented. Barnacle attachment traces are identified as Anellusichnus circularis. The wide distribution of barnacles on some of these bones suggests these were regularly overturned by bottom currents, which would prevent barnacles from being smothered by prolonged contact with the sediment. Detailed study of barnacle-induced trace fossils on these specimens suggests that episkeletozoans and their traces can be useful sources of data regarding the biostratinomic history of vertebrate fossils.
A large and abundant columnar stromatoporoid, Quasiaulacera n. gen., from the Ellis Bay Formation, up to 3 m long and 40 cm in diameter, marks the Hirnantian (latest Ordovician) of Anticosti Island. Two species are present: Quasiaulacera stellata n. sp. from the basal Ellis Bay Formation (basal Prinsta Member, lower Hirnantian) along the northeastern coast of the island, and the type species Q. occidua n. sp. from the upper Ellis Bay Formation (Lousy Cove Member, upper Hirnantian) in the western carbonate facies of the island. Quasiaulacera is rare or absent in the reefal Laframboise Member (uppermost Hirnantian) of the formation. The new genus differs from Aulacera in the underlying Vaureal Formation (upper Katian) in having a large central axial zone marked by a single stack of large, convex-up cyst-plates, that is surrounded by a middle layer of small, concentric microcyst-plates, in places denticulate, and an outer layer composed of concentric laminae with dense pillars, in which microcyst-plates are either absent or rare. The outer two layers are defined by longitudinal fluting; there are no branching forms. Both species demonstrate a ball-like holdfast system, some with diameters of 30 to 70 cm, microbially cemented into the substrate. Quasiaulacera “gigantism” in the paleotropical Anticosti Basin evolved at a time of global cooling associated with the Hirnantian glaciation in south polar Gondwana, but terminated in mass extinction of the aulaceratids at the O/S boundary in Laurentia. This supports other evidence that the Hirnantian featured not only generic loss, but also innovation and migration in tropical latitudes.
The scorpionfly (Mecoptera) superfamily Panorpoidea underwent an Eocene radiation, replacing the extinct Mesozoic orthophlebiid grade and reaching its greatest family-level diversity: Panorpidae, Panorpodidae, Austropanorpidae, Holcorpidae, Dinopanorpidae, and a new family proposed here, the Eorpidae. Only the Panorpidae and Panorpodidae survived the Eocene and persist to the present day. This cluster of family extinctions is exceptional within Cenozoic insects. The Eorpidae includes at least one new genus and three new species described here from four localities of the early Eocene Okanagan Highlands of British Columbia, Canada, and Washington, U.S.A.: Eorpa ypsipeda n. gen. n. sp. (McAbee and possibly Falkland, BC, Canada; and Republic, WA, U.S.A.), Eorpa elverumi n. gen. n. sp. (Republic), and Eorpa jurgeni n. gen. n. sp. (Quilchena, BC). Some of the other fragmentary and poorly preserved specimens might represent further new species. We propose that the apex of Panorpoid family diversity ended by pressures from post-Eocene icehouse world climatic stress and the rise to ecological dominance of ants, some of which would have provided strong competition in scavenging for dead arthropods.
The Cenomanian hexactinellid assemblages are relatively rare and are known only from England and France. The phosphatized sponges from the Pokuttian Podillia in western Ukraine seem to be taxonomically the most diverse Cenomanian hexactinellid assemblage known from Europe and the world. Several new hexactinellid taxa (Class Hexactinellida) are described from the Cenomanian of Pokuttian Podillia: Myliusia rakovensis n. sp., Verrucocoelia regularis n. sp., Tretochone cretacea n. sp., Heterochone boguckii, Dunikowskiella nezvyskensis n. gen. n. sp., and Moretiella foliacea n. sp. A new lychniscosidan genus Dunikowskiella is established within the family Callodictyonidae. Until now, genus Moretiella was reported only from the Aptian of Spain and the Albian of France. The extant genera Heterochone and Myliusia are also rare as fossils. Heterochone was known from the Campanian of Germany. Myliusia, noted in the Miocene of Spain, is well documented in Late Cretaceous for the first time. Tretochone was known only from recent times. The sponges studied lived in the shelf sea, shallower than most recent hexactinellids occur.
The feather-shaped PlumalinaHall, 1858 is revised on the basis of new and reexamined specimens from New York. Previously described from Givetian through Famennian deposits, a single compression of P. tenera n. sp. from the Rochester Shale extends the range into the Wenlock, and provides new information regarding Plumalina's biology. We assess the utility of morphologic characters in diagnoses of taxa, and present the first quantitative analysis of fossil hydroids to distinguish P. brevis n. sp. (Frasnian) from other Devonian species.
Plumalina has been compared to plants, graptolites, and octocorals. Some interpretations have proposed affinities among hydrozoans based on colony form and the presence of putative polyp bases. Our analysis shows that, like extant thecate hydrozoans, Plumalina had a delicate, chitinous hydrocaulus with weakly articulated hydrocladia. An assemblage of in situ specimens, steeply inclined relative to the bedding plane in an Ithaca Formation (Frasnian Stage) turbidite, indicates that Devonian species produced sessile, erect colonies attached to a hard substrate, comparable to extant hydroids that feed in currents. Morphometric comparisons between putative Plumalina polyp bases and polyp bases of modern analogues reveal similarities to hydroids in the superfamily Plumularioidea McCrady, 1859. Plumalina is the most abundant fossil hydroid so far reported, and is pertinent to interpretations of the hydrozoan record.
A Maastrichtian rudist fauna composed of the radiolitids Biradiolites aguilerae Böse, B. Cárdenasensis Böse, Huasteca ojanchalensis (Myers), Tampsia floriformis Myers, and Trechmannites rudissimus (Trechmann), the hippuritids Caribbea muellerriedi (Vermunt) and Praebarrettia sparcilirata (Whitfield) sensu lato, and the plagioptychids Coralliochama gbohemi Böse and Mitrocaprina tschoppi (Palmer) is described from the Cárdenas Formation in San Luis Potosí State, Mexico. Abundant fossil material and excellent preservation of a number of specimens allowed observation of both the internal and external shell characters and their ontogenetic and eco-phenotypic variability. The description of some hitherto insufficiently known species has been enhanced and/or completed, making easier their subsequent identification and allowing their unequivocal generic assignation. Two new genera, Huasteca and Trechmannites, are proposed for two already known species of radiolitids. The Cárdenas Formation exhibits a continuously exposed sequence in the vicinity of Cárdenas. Thus, precise stratigraphic location of all fossil localities and their rudist associations, ranging from the early to the early late Maastrichtian, has been possible.
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