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Taxonomic analysis is provided for a Middle Pennsylvanian macrofloral assemblage collected from clastic wetland deposits in Clay County, Indiana, on the eastern margin of the Illinois Basin. Adpressed plant fossils were recovered from four distinct beds in the lowermost Staunton Formation, positioned above the Minshall Coal (uppermost Brazil Formation), part of a succession deposited near the Atokan-Desmoinesian boundary. The assemblage of 22 fossil-taxa is dominated by pteridosperms (including Neuropterisflexuosa, Macroneuropteris scheuchzeri, Alethopteris densinervosa, Neuropteris ovata, Eusphenopteris neuropteroides, and Neuropteris missouriensis) with lesser cordaitaleans (Cordaites spp. indet.) and sphenopsids (particularly Sphenophyllum cuneifolium). Lycopsids are uncommon, and ferns are rare. In contrast, the microfloral assemblage from the Minshall Coal and overlying clastic units is dominated by lycopsid and tree fern spores. Comparisons with established biozonation schemes yield different ages depending on the regional biostratigraphic framework used: (1) latest Bolsovian (Radiizonates difformis Biozone, American microfloras); (2) latest Bolsovian or earliest Asturian (‘Neuropteris’ rarinervis Biozone, Appalachian Basin macrofloras); or (3) earliest Asturian (Linopteris obliqua Biozone, European macrofloras). The placement and correlation of the Bolsovian-Asturian and Atokan-Desmoinesian boundaries, which have traditionally been equated by palynology, are evaluated in the context of this discordance. Several revised stratigraphic scenarios are proposed for this interval in the Illinois Basin, which is being increasingly recognized as a time of significant environmental change throughout Euramerica. Homotaxial comparisons with European macrofloral assemblages indicate that, of the 18 biological taxa recorded, between 14 and 17 (78–94%) also are common in coeval wetland deposits in Europe. The similarities exemplify the spatial conservatism and low diversity of wetland plant communities over vast areas of tropical Euramerica, a manifestation of the intrinsically stressful conditions that characterize such habitats, and indicates that neither the Laurentian Shield nor the Appalachian-Variscan Mountains were an insurmountable barrier to plant dispersal during the Middle Pennsylvanian.
From new and more complete material, which includes frond fragments with casts of tracheid remains of the rachis, it can be demonstrated that the putative liverwort Hepaticites iporangaeRicardi-Branco, Faria, Jasper, and Guerra-Sommer, 2011, from the early Permian Rio Bonito Formation (Sakmarian) of the Paraná Basin, Brazil, is not a bryophyte but a tracheophyte. The new material was collected from the same locality and layer as the type material, in the Quitéria outcrop in the municipality of Encruzilhada do Sul, state of Rio Grande do Sul, Brazil. From the morphology of the deeply dissected pinnatifid pinnules with narrow laminar lobes, the taxon is provisionally reassigned to the genus Rhodeopteridium. Thus we propose the new combination ‘Rhodeopteridium’ iporangae new combination for this taxon. This new systematic interpretation contributes to our understanding of early liverworts (by removing Hepaticites iporangae as a possible taxon thereof) and clarifies an issue of diversity of the flora of the early Permian Rio Bonito Formation resulting from the original misidentification.
Angiosperms first appeared in the fossil record as pollen during the Valanginian—Hauterivian; they spread out of the tropics in the Aptian and Albian, and radiated in the Late Cretaceous. Despite these general patterns, details of the taxonomic, geographic, and ecological evolution of Cretaceous angiosperms are relatively poorly known because only a handful of Early and mid-Cretaceous macrofloras have been reported. This is the first detailed report of a fossil leaf flora from the Cedar Mountain Formation from the mid-Cretaceous of the Western Interior. We describe a flora that is overwhelmingly dominated by angiosperms (152 of 153 identified specimens are angiosperms) from the Albian—Cenomanian transition that is preserved in a clay- and carbonate-rich, lacustrine mudstone from the uppermost Cedar Mountain Formation of Emery County, Utah. We recognize 18 leaf morphotypes, all of which are dicotyledonous angiosperms. The majority of the Cedar Mountain morphotypes have taxonomic affinities with forms of similar age described from the Atlantic and Gulf coastal plains and other localities from the Western Interior. From this, we infer that a relatively diverse angiosperm flora grew along the margins of a small pond on the coastal plain. Palynological preparations of the fossil matrix were barren; however, previous studies of other facies within the formation showed that both conifers and ferns were important components of the regional vegetation during Cedar Mountain time. The effective absence of conifers and ferns in this macroflora and low leaf mass per area values among the angiosperms measured suggests that even at the Early—Late Cretaceous transition, angiosperms had come to dominate some sites, particularly those that were disturbed or seasonally ephemeral, where fast-growth or seasonal deciduousness would have been favored.
The recently described radiodontan Lyrarapax unguispinusCong et al., 2014 from the Chengjiang biota (Cambrian Series 2, Stage 3) highlighted a new morphological type of frontal appendage and unique mouth structures, a functional combination reinforcing the diversification of feeding strategies of radiodontans during the early Cambrian. Here we describe Lyrarapax trilobus n. sp. from the same fossil Konservat-Lagerstätte. The new species differs from L. unguispinus in the morphology and distribution of endites on the frontal appendage and the strengthening structure of the body flaps. The two species resemble each other in body shape (pattern of flap size), neck segment number, cephalic plates, and most importantly a mouth characterized by concentric wrinkled furrows. The latter confirms that a soft mouth without sclerotized plates is a real feature of Lyrarapax and supports the idea that oral structures provide valid diagnostic characters within Radiodonta.
The late Cambrian-Early Ordovician sequences of the Cordillera Oriental in northwestern Argentina are extensive and highly fossiliferous. The olenid trilobite Parabolinella argentinensis Kobayashi sensu Harrington and Leanza, 1957 was reported from a great number of Tremadocian localities and includes a wide range of morphologies. Based on specimens collected from the type locality (Purmamarca, Jujuy) and the quebrada Moya (Huacalera, Jujuy), as well as on material of the Harrington and Leanza collections in the University of Buenos Aires, classic morphometry and geometric morphometry methods were used to evaluate the variability in the cranidium of P. argentinensis. The results obtained from the two methodologies are similar. Both analyses allowed the review of the diagnosis of P. argentinensis, and the distinction of two new morphologies: Parabolinella clarisae n. sp. and Parabolinella pompadouris n. sp. The results show how morphogeometrics distinguishes more clearly the three morphotypes and provides graphical representations of the differences between those groups and how from the representations plus the correlation between classical variables and principal components, new diagnostic characters that distinguish the three morphospecies, can be identified.
The adoral surface of a crinoid theca has traditionally been called the tegmen, despite a wide range of morphologies; and, unfortunately, this has obscured the potential to recognize homologies between blastozoans and crinoids. With present recognition of these homologies, the constructional morphology of crinoid oral regions is explored, herein. Two major types of oral regions exist among crinoids: (1) an oral surface with the mouth exposed; and (2) a tegmen, in a restricted definition, with the mouth covered beneath solid plating. A tegmen is constructed by exaptation of oral surface plating and, commonly, other thecal plating. A pseudo-tegmen is an exaptation of aboral cup plates (i.e., radial plates). Tegmens or pseudo-tegmens evolved in all major crinoid clades at least once as an exaptation of oral surface plating. Tegmens evolved iteratively both between and within clades. In some cases, tegmen plates can be homologized with oral surface plates, but in other cases this is not apparent. Examples of tegmens that evolved many times include tegmens with an appearance of oral surface plates cemented in place; tegmens with fixed ambulacral cover plates and primary peristomial cover plates disproportionately enlarged; tegmens composed exclusively, or nearly so, of greatly enlarged primary peristomial cover plates; tegmens with tessellate plating but presumably with some flexibility; and tegmens constructed of innumerable undifferentiated plates. Most tegmens have all ambulacral cover plates fixed; but in some instances, the abaxial ambulacral cover plates remain moveable. Additionally, some lineages that possessed a tegmen evolved an oral surface secondarily, likely as an atavism. Based on this restricted definition of a tegmen, the hemicosmitid blastozoan Caryocrinites also evolved a tegmen. As known, tegmens dominated among camerate crinoids; and oral surfaces were more common among cladids, hybocrinids, disparids, flexibles, and articulates. However, oral surfaces evolved in some camerate lineages; tegmens evolved in some cladid, disparid, and articulate lineages; and pseudo-tegmens evolved in some flexible and articulate lineages.The iterative evolution of tegmens in crinoids and blastozoans is thought to be an adaptive response to cover the mouth and proximal ambulacra to protect this portion of the digestive tract from predation, scavenging, parasites, and disease causing agents.
A well-preserved omission surface (sedimentary discontinuity) in an outcrop near Alcoi in southeastern Spain displays trace fossils and body fossils that reflect a dynamic benthic community during the Miocene (Langhian—Tortonian). This outcrop, besides being the type locality of Spongeliomorpha ibericaSaporta, 1887, exhibits other abundant trace fossils, such as Glossifungites saxicavaŁomnicki, 1886 and Gastrochaenolites ornatusKelly and Bromley, 1984. These trace fossils are restricted to a single stratigraphic horizon and constitute a typical firmground ichnoassemblage of the Glossifungites ichnofacies. The interiors of some of the Glossifungites and Spongeliomorpha burrows were occupied by encrusting balanomorph barnacles (Actinobalanus dolosusDarwin, 1854). This paper is the first report of cryptic barnacles colonizing the interior of open burrows that constitute a typical firmground ichnocoenose in the fossil record. Detailed ichnologic study demonstrates that the ichnospecies Glossifungites saxicava stands as a valid ichnotaxon and is not a synonym of the ichnogenus Rhizocorallium, as has been suggested by some previous workers.
Fossil fish are poorly represented in middle Lower Devonian deposits of the western United States. Here we describe vertebrate microremains from mid-Pragian levels in the Kobeh Member, McColley Canyon Formation, central Nevada. The species diversity of the assemblages is low and dominated by the acanthodian Nostolepis costataGoujet, 1976. This species is found in older (late Lochkovian—early Pragian) deposits in France and Spain and mid-Pragian or slightly younger deposits in Saudi Arabia. This distribution suggests dispersal to the west from the Armorica terrane around the south of Laurussia to the Nevada region, and southeastward to Arabia, on the northern margin of Gondwana.
Although Ptychodus teeth are well known in Late Cretaceous marine deposits in North America and Europe and a few specimens with jaw elements have been discovered, the taxonomic position of the shark genus Ptychodus is enigmatic due to the lack of preservation of diagnostic material other than teeth. These sharks possessed a pavement dentition suited to a diet of hard-shelled macroinvertebrates (durophagy), leading several studies to variously describe Ptychodus as a batoid, a hybodont shark, or a selachimorph. Members of the Selachimorpha share one dental synapomorphy, a triple-layered enameloid (TLE) consisting of an outer shiny-layered enameloid (SLE) of randomly oriented hydroxyapatite crystallites, a middle layer of parallel-bundled enameloid (PBE), and an inner layer of tangled-bundled enameloid (TBE). Batoids and hybodonts both have teeth with single crystallite enameloid (SCE). We examined teeth from Ptychodus collected from the Lincoln Limestone of the Greenhorn Formation of Barton County, Kansas, and compared their enameloid ultrastructure with that of a Carboniferous hybodontiform and the Cretaceous lamniform shark Squalicorax curvatusWilliston, 1900. Scanning electron microscopic examination of Ptychodus shows that crystallite bundling in the form of a TLE is evident in these teeth. The PBE is most apparent at transverse enameloid ridges of Ptychodus teeth. Columns of dentine penetrate into the tooth enameloid, and the crystallites near the dentine are randomly oriented. These observations bolster the argument that Ptychodus is a genus of highly specialized selachimorph shark, rather than a hybodont or batoid.
The Field Museum of Natural History collection contains several isolated hadrosaurid specimens collected by Charles H. Sternberg from the lower Kirtland Formation of the San Juan Basin, New Mexico, that have been previously overlooked. Cranial elements described herein consist of a dentary and three jugals while appendicular material is limited to two humeri and two pubes. Many of the specimens preserve taxonomically informative characters that show strong affinities with Kritosaurini but are distinct from Kritosaurus navajovius (Brown, 1910) suggesting that the saurolophine-dominated San Juan Basin diversity is greater than currently recognized. Future examination of currently unprepared material will add to our developing understanding of the ambiguous hadrosaurid diversity of the San Juan Basin.
The genus Neocricetodon represents one of the most diverse radiations of extinct cricetines. Its early evolution remains unclear, mainly owing to the incomplete fossil record. One of the earliest and least known species of this genus is N. moldavicus described from the early late Miocene localities Bujor 1 and Calfa, Moldova. We reexamined the type material of this species and compared it with other species of Neocricetodon. Despite its old geological age, N. moldavicus demonstrates rather advanced dental morphology including the anterocone of M1 being deeply split; mesolophs and mesolophids partially reduced; and the labial spur of the anterolophule, ectomesolophids, and lingual anterolophid of m2 lacking. The phylogenetic analysis of 15 Neocricetodon species based on 22 dental characters revealed three synapomorphies for the genus: presence of the M1 labial anterolophule, the four-rooted M2, and presence of the labial anterolophulid of m1. “Kowalskia cf. schaubi” from Rudabánya and “Kowalskia sp.” from Vösendorf, formerly considered as the earliest finds of Neocricetodon, do not possess these characters and rather represent the genus Democricetodon. Our study also suggests that Kowalskia complicidens is a member of Sinocricetus, and that N. neimengensis and N. zhengi are junior synonyms of N. polonicus. The recognition of N. moldavicus as the earliest member of the genus within the clade containing the species of Neocricetodon from France and Spain supports the hypothesis of the Eastern European origin of the Western European Neocricetodon species.
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