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Tubular carbonate concretions of up to 1 m in length and perpendicular to bedding, occur abundantly in the Upper Pliensbachian (upper Amaltheus margaritatus Zone, Gibbosus Subzone) in outcrops (Fontaneilles section) in the vicinity of Rivière-sûr-Tarn, southern France. Stable isotope analyses of these concretions show negative delta 13C values that decrease from the rim to the center from - 18.8‰ to - 25.7‰ (V-PDB), but normal marine delta 18 O values (- 1.8‰). Carbon isotope analyses of Late Pliensbachian bulk carbonate (matrix) samples from the Fontaneilles section show clearly decreasing C-isotope values across the A. margaritatus Zone, from +1‰ to - 3‰ (V-PDB). Isotope analyses of coeval belemnite rostra do not document such a negative C-isotope trend with values remaining stable around +2‰ (V-PDB). Computer tomographic (CT) scanning of the tubular concretions show multiple canals that are lined or filled entirely with pyrite. Previously, the formation of these concretions with one, two, or more central tubes, has been ascribed to the activity of an enigmatic organism, possibly with annelid or arthropod affinities, known asTisoa siphonalis. Our results suggest tisoan structures are abiogenic. Based on our geochemical analyses and sedimentological observations we suggest that these concretions formed as a combination of the anaerobic oxidation of methane (AOM) and sulfate reduction within the sediment. Fluids rich in methane and/or hydrocarbons likely altered local bulk rock carbon isotope records, but did not affect the global carbon cycle. Interestingly, Tisoa siphonalis has been described from many locations in the Grands Causses Basin in southern France, and from northern France and Luxemburg, always occurring at the same stratigraphic level. Upper Pliensbachian authigenic carbonates thus possibly cover an area of many thousand square kilometers. Greatly reduced sedimentation rates are needed to explain the stabilization of the sulfate-methane transition zone in the sedimentary column in order for the tubular concretions to form. Late Pliensbachian cooling, reducing run-off, and/or the influx of colder water and more vigorous circulation could be responsible for a halt in sedimentation. At the same time (thermogenic) methane may have destabilized during a major phase of Late Pliensbachian sea level fall. As such Tisoa siphonalis is more than a geological curiosity, and its further study could prove pivotal in understanding Early Jurassic paleoenvironmental change.
The siliceous claystone and chert lithologic units of the Triassic-Jurassic chert-clastic sequence are well exposed in the Inuyama, Mt. Kinkazan and Hisuikyo areas of the southeastern Mino Terrane. Twenty-one continuous sections from those areas were investigated in order to establish comprehensive radiolarian biozones and clarify the successive lithologic changes through the Triassic and lowest Jurassic. Twenty new radiolarian zones are established; the lowest two are assemblage zones and the others are defined by the first or last occurrence of index taxa. The definitions are as follows in chronological order: TR 0, Follicucullus Assemblage Zone (early Spathian or older); TR 1, Parentactinia nakatsugawaensis Assemblage Zone (late Spathian); TR 2A, Eptingium nakasekoi Lowest-occurrence Zone (early Anisian); TR 2B, Triassocampe coronata group Lowest-occurrence Zone (early Anisian); TR 2C, Triassocampe deweveri Lowest-occurrence Zone (late Anisian); TR 3A, Spine A2 (possiblly derived from Oertlispongus inaequispinosus) Lowest occurrence Zone (late Anisian) ; TR 3B, Yeharaia elegans group Lowest-occurrence Zone (early Ladinian); TR 4A, Muelleritortis cochleata Lowest-occurrence Zone (late Ladinian); TR 4B, Spongoserrula dehli Lowest-occurrence Zone (late Ladinian to early Carnian); TR 5A, Capnuchosphaera Lowest-occurrence Zone (early Carnian); TR 5B, Poulpus carcharus sp. nov. Lowest-occurrence Zone (early to late Carnian); TR 6A, Capnodoce- Trialatus Concurrentrange Zone (late Carnian to early Norian), TR 6B, Trialatus robustus-Lysemelas olbia gen. et sp. nov. Partial-range Zone (early Norian); TR 7, Lysemelas olbia gen. et sp. nov. Lowest-occurrence Zone (early to late Norian); TR 8A: Praemesosaturnalis multidentatus group Lowest-occurrence Zone (late Norian); TR 8B: Praemesosaturnalis pseudokahleri sp. nov. Lowest-occurrence Zone (late Norian) ; TR 8C: Skirt F (possiblly derived from Haeckelicyrtium takemurai) Lowest-occurrence Zone (late Norian to early Rhaetian); TR 8D: Haeckelicyrtium breviora sp. nov. Taxon-range Zone (early to late Rhaetian) ; JR OA: Haeckelicyrtium breviora sp. nov.-Bipedis horiae sp. nov. Partial-range Zone (Hettangian); and JR OB: Bipedis horiae sp. nov. Lowest-occurrence Zone (Hettangian/Sinemurian) . These zones are correlated to previousy established radiolarian assemblages and zones in Japan and other regions. Age assignment of the zones is also discussed on the basis of the correlation and other available chronological data. The original stratigraphic succession of the Triassic in the studied area, which ranges in age from Early Triassic to Early Jurassic, is more than 100 m in thickness and can be reconstructed in detail. The succession is subdivided into seven units based on lithologic features. Each unit was probably accumulated under a particular sedimentary condition, thus successive changes of paleoceanographic environments during Triassic time can be traced continuously. Nine new genera including Ayrtonius, Blonzella, Braginella, Bulbocampe, Enoplocampe, Lysenzelas, Parvibrachiale, Spongoxystris and Veles, and 47 new species are described herein. A comprehensive list of identified taxa is presented.
Agglutinated foraminiferal assemblages from the Oligocene section of an exploration well drilled in the distal part of the Congo Fan are fully documented and interpreted for palaeoenvironment. A total of 65 ditch cutting samples were analysed at 10 m intervals, from 3630 to 4270 m below rotary table. An average of 170 specimens were extracted per sample, with over 100 species being documented and described using SEM and light photography. The results reveal the most taxonomically diverse deepsea Oligocene fauna yet described. Six assemblages have been defined and analysed with Correspondence and 'Morphogroup' Analysis. These are 1. Nothia robusta / Reticulophragmium Assemblage (4110-4270 m), 2. Nothia robusta / Scherochorella congoensis / Discammilloides sp. 1 Assemblage (4000-4100 m), 3. High diversity Reticulophragmium Assemblage (3870-3990 m), 4. Portatrochammina profunda Assemblage (3790-3860 m), 5. Nothia latissima Assemblage (3730-3780 m) and 6. Low abundance Assemblage (3630-3720 m). Palaeobathymetric estimates range from middle -lower bathyal based on comparison with living taxa and morphogroup distributions. These results extend the known stratigraphic range (last occurrences) of Reticulophragmium amp/eetens into the Oligocene in the Atlantic, and possibly also Paratrochamminoides gorayskii, Paratrochamminoides olszewskii, Trochamminoides aff. proteus, Trochamminoides subcoronatus, Haplophragmoides horridus and Haplophragmoides walteri, although reworking is documented with these species. Results also extend the known first occurrences of Recurvoides azuaensis, Spiropsammina primula, Cyclammina aff. orbicularis, Discamminoides sp. and Glaphyrammina americana into the Oligocene. Large scale variations within faunas are largely assigned to documente d variations in sand content, where higher proportions of sand generally coincide with reduced diversity and abundance along with a dominance of opportunistic species such as Nothia robusta, Nothia latissima and Ammodiscus latus. A major excursion in the infaunal morpho group, suspension-feeding morpho group and diversity and abundance within Assemblage 2 is termed the 'Scherochorella Event', and does not correlate with an increase in sand. This fauna is thought to be the result of lower oxygen conditions allowing the dominance of the low oxygen morphotype Scherochorella congoensis and the opportunistic species Nothia robusta. Deep-water circulation in the Atlantic at this time is generally thought to have been strong, and this event suggests that there may have been a temporary expansion of the oxygen minimum zone during the Late Oligocene, coinciding with increased benthic 8180 values, global cooling, and increased upwelling associated with a stronger polar front. The otherwise high diversity of the fauna in the well supports the interpretation of well-oxygenated conditions.
Homology of virtually all major components of facial anatomy is assessed in Archosauria in order to address the function of the antorbital cavity, an enigmatic structure that is diagnostic for the group. Proposed functions center on its being a housing for a gland, a muscle, or a paranasal air sinus. Homology is approached in the context of the Extant Phylogenetic Bracket method of reconstructing unpreserved aspects of extinct organisms. Facial anatomy and its ontogeny was studied in extant archosaurs (birds and crocodilians) to determine the osteological correlates of each soft-tissue component; resemblances between birds and crocodilians comprised the similarity test of homology. The congruence test of homology involved surveying phyiogenetically relevant fossil archosaurs for these bony signatures. The facial anatomy of extant birds and crocodilians is examined in detail to provide background and to discover those apomorphic aspects that contribute to the divergent specialization of these two groups and thus obscure homologies. Birds apomorphically show enlarged eyeballs, expanded nasal vestibules, and reduced maxillae, whereas crocodilian faces are dorsoventrally flattened (due to nasal rotation) and elongated. Most facial attributes of archosaurs are demonstrably homologous and in fact characterize much more inclusive groups. Special emphasis has been placed on the nasal conchae and paranasal air sinuses. Within Amniota, the following conchal structures are homologous, and all others are neomorphs: avian caudal concha, crocodilian concha + preconcha, Sphenodon caudal concha, squamate concha, and probably the mammalian crista semicircularis. The avian antorhital paranasal air sinus is homologous with the crocodilian caviconchal sinus; the maxillary sinus of placental mammals is not homologous with the archosaurian paranasal sinus. With regard to the function of the antorbital cavity, archosaurs possess homologous nasal glands, dorsal pterygoideus muscles, and paranasal air sinuses, but the osteological correlates of only the paranasal sinus involve the antorbital fenestrae and fossae. Thus, the antorbital cavity is best interpreted as principally a pneumatic structure.
Carnian (Upper Triassic) fishes from Polzberg bei Lunz have been known since 1886 but no comprehensive account has been published. Eleven species are described nine of which, Saurichthys calcaratus, Polzbergia brochatus, Peltoplellrus dinlmptus, Habroichthys gregarius, Nannolepis elegans, Phaidrosoma lunzensis, Elpistoichthys pectinatus, E. striolatus and Pholidophoretes salvus are new, and two others, Thoracopterus niederristi Bronn and Gigantopterus telleri Abel, previously little-known. New supraspecific taxa defined are: the order Polzbergiiformes, the family Thoracopteridae and the genera Polzbergia, Nannolepis, Phaidrosoma, Elpistoichthys and Pholidophoretes. Habroichthys. Thoracopterus, Gigantopterus and Nannolepis show an unusual skull-roof pattern and are included in the re-defined order Luganoiiformes. Two new ichthyokentemids considerably extend the known time-range of this family. The genus Pholidophoretes is intermediate between the Archaeomenidae Goodrich 1909, and the Pholidophoridae sensu stricto Nybelin 1966. The Polzberg assemblage was probably mainly marine with a small freshwater contribution; it shows less similarity to the Besano and Raibl assemblages than these do to each other. The Luganoiiformes are probably, but not certainly, monophyletic; relationships within the order are analyzed and a cladogram constructed. The Platysiagiformes, Peltopleuriformes, Luganoiiformes and Cephaloxeniformes could all have been derived from a common ancestor at the Perleidus level and are probably offshoots of the perleidid radiation.
Delthyridoid spiriferids are characterized by a global abundance and fast evolution during Silurian and Devonian, and, therefore, are used as important biostratigraphical and palaeobiogeographical tools. In this work, delthyridoid brachiopod faunas from different regions of today’s world, resp., of different palaeobiogeographical units, are compared side-by-side to investigate their phylogenetic relationships and to improve, in a second step, the palaeobiogeography from Late Silurian to Early Eifelian time. A new systematics of Delthyridoidae is established which is more complicated than hitherto assumed. The results of this study are mainly based on direct comparison of articulated and isolated brachiopod shells, external and internal moulds, as well as latex casts and serial sections. The computer supported cladistic analyses have turned out not to be useful due to different kinds of preservation resulting in an incomplete matrix which is insufficient for reliable cladograms. A further problem in terms of cladistical analyses are various convergences during the evolution of spiriferids. Many characters evolved independently from each other at different times in each lineage so that autapomorphies are hardly or not at all recognizable. As a result, families and genera are only definable by a combination of characters rather than by a single or a few autapomorphies. As a new method, 3D reconstruction from serial sections is introduced which made it possible for the first time to compare directly mouldic and shelly material. Preliminary results are presented herein. Statistical analyses of measurements taken from new taxa are made but regarded as a descriptive argument rather than a deciding factor for taxonmy due to incomplete preservation and/or tectonic deformation. Brachiopods, especially type material, from collections of different institutions and museums are studied as well as personal material, whenever possible collected from topotype outcrops. Emended diagnoses, if necessary, from family to species level are given. During this work several new taxa have been erected: 7 new families: Australospiriferidae, Murchisonispiriferidae, Orientospiriferidae, Otospiriferidae, Patriaspiriferidae, Rostrospiriferidae, and Trigonospiriferidae; 6 new genera, 1 of these in open nomenclature: Cyclopterospirifer, Hallispirifer, Parlinispirifer, Murchisonispirifer, Shujiapingensispirifer, and gen. nov. B; and 3 new species: Patriaspirifer merriami, Patriaspirifer johnsoni, and Murchisonispirifer feldmani; 1 taxon is defined as nomen novum: Orientospirifer nakaolingensis wani. In the framework of this project, 2 families: Filispiriferidae and Multispiriferidae; 1 subfamily: Multiplicatispiriferinae, 6 genera, 1 of them in open nomenclature: Frequentispirifer, Leonispirifer, Multiplicatispirifer, Ovetensispirifer, Turcispirifer, and Gen. A; and 9 new species, 3 of them in open nomenclature: Filispirifer hamadae, Leonispirifer leonensis, Multiplicatispirifer foumzguidensis, Oventensispirifer novascotianus, Quiringites arensentiae, Turcispirifer turciae, Multiplicatispirifer cf. foumzguidensis, Quiringites cf. arensentiae, and ?Turcispirifer sp. A which have already been established are also described in this work. The brachiopod faunas studied consist of externally very similar spiriferids which have been identified as same genera, species, or even subspecies in earlier times. These forms are considered as 6 distinct morphotypes Howellella-, Arduspirifer-, Acrospirifer-, Euryspirifer-, Paraspirifer-, and Multiplicatispirifer-like morphotypes, which are briefly introduced. The new systematics is characterized by different clades, the European/North African delthyridoid spiriferid clade, the North American delthyridoid spiriferid clade, the Asian delthyridoid spiriferid clade, the Malvinokaffric delthyridoid spiriferid clade, and the delthyridoid multiplicated spiriferid clade. Each of them is described in a cladistic and in a phylogenetic way. Their phylogenetic relationship sheds new light on palaeobiogeographical interpretations for the different stages from Late Silurian to early Middle Devonian time. A tendency for increasing endemicity is seen until the end of the Early Emsian, which is interrupted by short term regional faunal exchange within a province or within a realm, followed by a loss of endemicity resulting in global distribution of brachiopod genera until the end of Givetian time. The Old World Realm is re-defined due to the lack of phylogenetic relationship between its faunas and subdivided into the European Realm, consisting of the Gondwanan and Avalonian provinces, and the Asian Realm, consisting of the Siberian, Sino, and Mongolian provinces. A reconstruction of Lower Devonian palaeobiographical map is introduced.
Glyptostrobus Endlicher is well represented in early Early Cretaceous to Pleistocene deposits in the middle to high latitudes of North America and Eurasia. Although the taxonomy and nomenclature of the genus is complicated, the fossil record indicates Glyptostrobus was represented by a small number of species. The genus first appears in Aptian age deposits from western Canada and Greenland, and achieved a wide distribution early in its evolutionary history. Exchange of Glyptostrobus between Asia and North America occurred across the Spitsbergen and Beringian corridors, which were functional about 110 and 100 million years ago, respectively The Late Cretaceous fossil record of Glyptostrobus shows that the genus had spread into Russia, China and the shores of the Turgai Strait. By the early Tertiary, Glyptostrobus was a prominent constituent of the polar broad-leaved deciduous forests. Paleocene age deposits across western Canada and the United States indicate the genus was present in great abundance in the lowland warm temperate and subtropical forests east of the Rocky Mountains. The broad distribution in North America and Russia during the Paleocene and Eocene indicates that Glyptostrobus grew and reproduced under a diverse range of climatic and environmental conditions, including the cold and unique lighting conditions of the polar latitudes. The presence of Glyptostrobus in Europe indicates the North Atlantic land bridges that extended between North America and Eurasia (Fennoscandia) and Europe during the early Tertiary were used. In Europe, extensive Glyptostrobus dominated swan1ps occupied the Central European Depression during the late Tertiary. Increasing global aridity and cooling, as well as landscape stabilization together with increasing competition for resources and habitat by representatives of the Pinaceae, seem to have forced the genus out of North America, Europe and most of Asia during the Miocene and Pliocene. In Japan, Glyptostrobus persisted until the early Pleistocene. After the early Pleistocene extinction in Japan, Glyptostrobus reappeared in southeastern China. Details of the taxonomic and biogeographic history of Glyptostrobus are examined.
Very little is known about the occlusal wear pattern in the Neanderthal posterior dentition. Usually dental wear is closely related to the physical properties of the ingested food, and consequently can be used to obtain information about diet. Neanderthal dietary reconstructions have been mostly based on the analysis of accompanying faunal remains and isotopic signatures of bones and tooth enamel, suggesting that they exploited larger portions of animal proteins from large and medium-sized herbivores. Probably these studies may do not reflect the bulk diet, tending to underestimate plant consumption and to overestimate meat consumption. In the present work the occlusal wear pattern of maxillary molars of Homo neanderthalensis (N=19) and early Homo sapiens (N=12)have been analyzed, applying non-destructive methods based on virtual three-dimensional polygonal models generated from surface scanning of dental casts. The sample groups occupied different geographical areas at different chronological times. The 3D digital tooth models were analyzed using the “Occlusal Fingerprint Analysis” (OFA) method (Kullmer et al. 2009), describing and quantifying the occlusal wear pattern derived from two wear facet angles (dip and dip direction), wear facet area and occlusal relief index (ORI). The OFA method provides information about the dynamics of the occlusal relationships and their function, permitting the reconstruction of the mandibular movements responsible for the contacts created during the chewing cycle. Since jaw movements and diet are closely related, the results obtained, can be used to interpret the diet of the two Pleistocene hominin species. In order to evaluate how dietary differences influence the occlusal wear pattern, upper molars of modern hunter-gatherers (N=42) with known diet and different dietary habits, have been included in the sample and compared with those of Neanderthals and early Homo sapiens. Results show that within the modern hunter-gatherers sample, the occlusal wear pattern of carnivorous populations differs from those who relied on a mixed-diet. In particular, the study of relative facet areas clearly distinguish meat-eaters from mixed-diet hunter-gatherers, while ORI results and wear facet inclinations (dip angle) seem to reflect directly the abrasiveness of the diet, including the influence of exogenous materials during food preparation. The Neanderthal occlusal wear pattern is characterized by an ecogeographic variation, suggesting the exploitation of different food resources. In particular Neanderthals who inhabited relatively warm environments of southern Europe and the Near East exhibit an occlusal wear pattern different from those of meat-eaters hunter-gatherers from tempered and cooler regions, displaying some features similar to those of Bushmen. These results suggest the exploitation of a broad variety of food sources. The analysis of the occlusal wear pattern in Neanderthals and early Homo sapiens who inhabited Europe during the cooler Oxygen Isotope Stage 3 (OIS3) shows many similarities between the two hominid species. These results indicate the exploitation of similar and low-diversified food sources, based mostly on the consumption of animal proteins, as suggested through the clear similarities with the wear patterns found in modern meat-eaters hunter-gatherers. In both studied groups, Neanderthals and early Homo sapiens the occlusal wear pattern is characterized by high ORI and dip angle values, suggesting the intake of a low-abrasive diet, probably due to the absence of sophisticated food preparation techniques introducing external silica grains, e.g. from soil (grinding of seeds) or plant cells, as those, seen in modern hunter-gatherer populations. The analysis of the occlusal fingerprints in Neanderthal and early European Homo sapiens upper molars suggests that both species followed very similar adaptive dietary strategies, based on a distinctive versatility and flexibility in the daily diet, depending on availability of resources according to environmental circumstances.
The 20 species of Neogene Scleractinia in the suborders Caryophylliina and Dendrophylliina known from the Dominican Republic are revised and illustrated. This research was based on 1590 specimens obtained primarily from the collections of the Naturhistorisches Museum, Basel, Switzerland; National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.; and Tulane University, New Orleans, LA, U.S.A. Eight new records are reported for the Neogene of the Dominican Republic, including four new species: Antillocyathus alatus, Trochocyathus chevalieri, T. duncani, and Paracyathus sinuosus. Special attention is given to the genus Asterosmilia, since half (five) of the known species in this genus occur in the Dominican Republic. Most species described herein are assumed to constitute a deep-water fauna by analogy to depth ranges of the same or similar species known from the Recent. Certain localities and parts of formations are inferred to represent deep-water (> 200 m) facies. These inferences may aid in the paleoecological interpretation of other fossils collected from these areas.
Multivariate statistical procedures are used to distinguish species in the reef-coral genus Stephanocoenia through a continuous Neogene sequence (five-million year time interval) in the Cibao Valley of the northern Dominican Republic. This genus is the only member of the family Astrocoeniidae that occurs in the sequence. The material consists of 56 colonies (17 of which are measured) from 24 localities in four river sections, the most important being Rio Gurabo and Rio Cana. Ten characters are measured on each of 10 corallites per colony. The data are analyzed using cluster and canonical discriminant analysis to group colonies into clusters representing species. Identical measurements on modem colonies collected near Discovery Bay, Jamaica are included for comparison. Two fossil species are defined in the analysis, one of which is new (Stephanocoenia duncani, n. sp.). Both species are significantly distinct from the single modem species (S. intersepla) that is the sole living representative of the genus. Study of collections from other reef localities shows that both fossil species occur only during Neogene time and only at a limited number of localities. Patterns within each species are traced up a composite stratigraphic section using nonparametric statistical analyses. One of the two fossil species (S. spongiformis) is found to remain stable through time, whereas the other (S. duncam) changes its morphology in a direction approaching the cluster for the modem species. Further study of patterns of variation within the one modern and two fossil clusters shows that intraspecific variation is unusually complicated in this genus. The clusters overlap, and colonies within each cluster differ widely. Variation between populations within the modem species occurs in the same characters as those which distinguish the modem species from the fossil species converging with it (S. duncam). However, these two species form a morphologic continuum that cannot be explained by environment alone. Therefore, they may represent two gradually intergrading chronospecies within one lineage. Of the two fossil species of Stephanocoenia defined, one species (s. spongiformis) exhibits an evolutionary pattem similar to that observed in the family Poritidae. In this pattern, species were found to have short durations and stable morphologies and to have become extinct during the mid- to late Pliocene through early Pleistocene mass extinction. In contrast, the second species of Slephanocaenia (S. duncam) may have evolved over a long time period, possibly forming chronospecies that survived the mass extinction. Unlike genera in the Poritidae, however, no radiation of taxa occurred in the genus after the extinction event. Since no consistent relationship has been discovered between morphology and environment in these corals with the data at hand, their paleoecologic value can only be determined after data on more taxa are collected, and their associations with other corals are studied. This study represents part of a multidisciplinary project on the stratigraphy and paleontology of the northern Dominican Republic, coordinated by P. Jung and J. B. Saunders of the Naturhistorisches Museum Basel, Switzerland.