Keratin Durability Has Implications for the Fossil Record: Results from a 10 Year Feather Degradation Experiment

Abstract: Keratinous ‘soft tissue’ structures (i.e. epidermally derived and originally non-biomineralized), include feathers, skin, claws, beaks, and hair. Despite their relatively common occurrence in the fossil record (second only to bone and teeth), few studies have addressed natural degradation processes that must occur in all organic material, including those keratinous structures that are incorporated into the rock record as fossils. Because feathers have high preservation potential and strong phylogenetic signal, in the current study we examine feathers subjected to different burial environments for a duration of ~10 years, using transmission electron microscopy (TEM) and in situ immunofluorescence (IF). We use morphology and persistence of specific immunoreactivity as indicators of preservation at the molecular and microstructural levels. We show that feather keratin is durable, demonstrates structural and microstructural integrity, and retains epitopes suitable for specific antibody recognition in even the harshest conditions. These data support the hypothesis that keratin antibody reactivity can be used to identify the nature and composition of epidermal structures in the rock record, and to address evolutionary questions by distinguishing between alpha- (widely distributed) and beta- (limited to sauropsids) keratin.

feather
high preservation potential
keratin


"Dinosaur" petroglyphs at Kachina Bridge site, Natural Bridges National Monument, southeastern Utah: not dinosaurs after all

Abstract: Bridge is what appears to be an unambiguous depiction of a sauropod dinosaur, herein called Dinosaur 1. Because mainstream science has produced no alternate explanation for Dinosaur 1, it has become an important weapon in the arsenal of the anti-evolution movement. The movement's proponents claim that it demonstrates the coexistence of humans and dinosaurs, thus casting doubt on the geological time scale of millions of years. Until now that claim has gone unchallenged. The hypothesis that a given petroglyph depicts a dinosaur predicts that the image is not a composite; depicts an animal; has features that cannot be reconciled with non-dinosaurian local fauna; has features of a specific, identifiable dinosaur; and is entirely human-made. These predictions were tested for Dinosaur 1 and three other alleged dinosaur petroglyphs at Kachina Bridge by on-site visual examination under varying light conditions. Examination reveals that the "neck" and "back" of Dinosaur 1 are a composite of two separate petroglyphs, and its "legs" are a natural mud or mineral stain. A second alleged sauropod petroglyph is a mere mud stain. The other two alleged dinosaur petroglyphs are human-made, but neither depicts an animal. The four Kachina Bridge "dinosaurs" are illusions produced by pareidolia. None of them satisfy the predictions of the hypothesis that a dinosaur is depicted. Dinosaur 1-heretofore a creationist poster child-and its counterparts now join the plethora of discredited "evidence" for the ancient coexistence of humans and dinosaurs.

petroglyph
kachina bridge
leg
mineral stain


Chronostratigraphy and “Land-Mammal Ages” in the Cenozoic of southern South America: principles, practices, and the “Uquian” problem

Abstract: The concept of “land-mammal age” as developed in South America is examined. The “Uquian Land-mammal age” is used as a study case. “Land-mammal age” parataxonomy is here considered methodologically but not conceptually different from chronostratigraphic taxonomy. “Land-mammal ages” in South America are based on stages. However, we consider that accurate biostratigraphic studies must be done in South America for establishing the biostratigraphy and precise boundary stratotypes of most stages-ages. The Uquia outcrops are here considered inadequate as a stratotype. A new South American continental stage-age is proposed. This stage is based on a biostratigraphic scheme. The stratotypes of the stage and biozones are located in the fossiliferous southeastern Buenos Aires Province marine cliffs. The lower boundary stratotype is proposed. The stage-age is probably correlated with the Gauss Chron and the lower Matuyama Chron. Additionally, some major units of Ameghino are validated and a different timing for the arrival of North American mammals to southern South America is presented.


Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)

Abstract: Machimosaurus was a large-bodied genus of teleosaurid crocodylomorph, considered to have been durophagous/chelonivorous, and which frequented coastal marine/estuarine ecosystems during the Late Jurassic. Here, we revise the genus based on previously described specimens and revise the species within this genus. We conclude that there were three European Machimosaurus species and another taxon in Ethiopia. This conclusion is based on numerous lines of evidence: craniomandibular, dental and postcranial morphologies; differences in estimated total body length; geological age; geographical distribution; and hypothetical lifestyle. We re-diagnose the type species Machimosaurus hugii and limit referred specimens to only those from Upper Kimmeridgian–Lower Tithonian of Switzerland, Portugal and Spain. We also re-diagnose Machimosaurus mosae, demonstrate that it is an available name and restrict the species to the uppermost Kimmeridgian–lowermost Tithonian of northeastern France. We re-diagnose and validate the species Machimosaurus nowackianus from Harrar, Ethiopia. Finally, we establish a new species, Machimosaurus buffetauti, for the Lower Kimmeridgian specimens of France and Germany (and possibly England and Poland). We hypothesize that Machimosaurus may have been analogous to the Pliocene–Holocene genus Crocodylus in having one large-bodied taxon suited to traversing marine barriers and additional, geographically limited taxa across its range.


Fossil Dolphin Otekaikea marplesi (Latest Oligocene, New Zealand) Expands the Morphological and Taxonomic Diversity of Oligocene Cetaceans

Abstract: The Oligocene Epoch was a time of major radiation of the Odontoceti (echolocating toothed whales, dolphins). Fossils reveal many odontocete lineages and considerable structural diversity, but whether the clades include some crown taxa or only archaic groups is contentious. The New Zealand fossil dolphin “Prosqualodon” marplesi (latest Oligocene, ≥23.9 Ma) is here identified as a crown odontocete that represents a new genus, Otekaikea, and adds to the generic diversity of Oligocene odontocetes. Otekaikea marplesi is known only from the holotype, which comprises a partial skeleton from the marine Otekaike Limestone of the Waitaki Valley. Otekaikea marplesi was about 2.5 m long; it had procumbent anterior teeth, and a broad dished face for the nasofacial muscles implicated in production of echolocation sounds. The prominent condyles and unfused cervical vertebrae suggest a flexible neck. A phylogenetic analysis based on morphological features places Otekaikea marplesi in the extinct group Waipatiidae, within the clade Platanistoidea. The phylogeny implies an Oligocene origin for the lineage now represented by the endangered Ganges River dolphin (Platanista gangetica), supporting an Oligocene history for the crown Odontoceti.

otekaikea marplesi
new zealand
prosqualodon
new genus
phylogenetic analysis


Dynamics of dental evolution in ornithopod dinosaurs

Abstract: Ornithopods were key herbivorous dinosaurs in Mesozoic terrestrial ecosystems, with a variety of tooth morphologies. Several clades, especially the ‘duck-billed’ hadrosaurids, became hugely diverse and abundant almost worldwide. Yet their evolutionary dynamics have been disputed, particularly whether they diversified in response to events in plant evolution. Here we focus on their remarkable dietary adaptations, using tooth and jaw characters to examine changes in dental disparity and evolutionary rate. Ornithopods explored different areas of dental morphospace throughout their evolution, showing a long-term expansion. There were four major evolutionary rate increases, the first among basal iguanodontians in the Middle-Late Jurassic, and the three others among the Hadrosauridae, above and below the split of their two major clades, in the middle of the Late Cretaceous. These evolutionary bursts do not correspond to times of plant diversification, including the radiation of the flowering plants, and suggest that dental innovation rather than coevolution with major plant clades was a major driver in ornithopod evolution.

ornithopod
hadrosaurid
dental disparity
dynamic


Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny

Abstract: The fossil record suggests a rapid radiation of placental mammals following the Cretaceous–Paleogene (K–Pg) mass extinction 65 million years ago (Ma); nevertheless, molecular time estimates, while highly variable, are generally much older. Early molecular studies suffer from inadequate dating methods, reliance on the molecular clock, and simplistic and over-confident interpretations of the fossil record. More recent studies have used Bayesian dating methods that circumvent those issues, but the use of limited data has led to large estimation uncertainties, precluding a decisive conclusion on the timing of mammalian diversifications. Here we use a powerful Bayesian method to analyse 36 nuclear genomes and 274 mitochondrial genomes (20.6 million base pairs), combined with robust but flexible fossil calibrations. Our posterior time estimates suggest that marsupials diverged from eutherians 168–178 Ma, and crown Marsupialia diverged 64–84 Ma. Placentalia diverged 88–90 Ma, and present-day placental orders (except Primates and Xenarthra) originated in a ∼20 Myr window (45–65 Ma) after the K–Pg extinction. Therefore we reject a pre K–Pg model of placental ordinal diversification. We suggest other infamous instances of mismatch between molecular and palaeontological divergence time estimates will be resolved with this same approach.

molecular study
fossil calibration
placentalia


Evolutionary Patterns Among Permo-Triassic Therapsids*

Abstract: ▪ Abstract A rich fossil record documents nonmammalian evolution. In recent years, the application of cladistic methodology has shed valuable light on the relationships within the therapsid clades Biarmosuchia, Dinocephalia, Anomodontia, and Cynodontia. Recent discoveries from South Africa suggest that Gondwana, rather than Laurasia, was the center of origin and radiation for many early therapsids. Because of their relative abundance and global distribution, therapsids have enjoyed widespread use in biostratigraphy, basin analysis, and paleo-environmental and -continental reconstructions. Synapsids (including therapsids) form the bulk of tetrapod diversity (in terms of both number of species and abundance) from Early Permian to Middle Triassic times and thus can provide critical information on the nature of the Permo-Triassic extinction in the terrestrial realm. Quantitative techniques have produced headway into understanding the relative importance of homoplasy and convergent evolution in the origin of m...


A New Stem Odontocete from the Late Oligocene Pysht Formation in Washington State, U.S.A.

Abstract:
ABSTRACT
 Odontocetes first appear in the fossil record during the early Oligocene and soon after show a near-worldwide distribution. However, little is still known of their early diversity, especially in the North Pacific Region. A new taxon of stem odontocete with heterodont dentition, Olympicetus avitus, gen. et sp. nov., is described herein from the upper Oligocene Pysht Formation in Washington State, U.S.A. The material consists of two partial skulls belonging to a juvenile and a neonate and includes part of the dentition and a tympanic bulla. Olympicetus shares many similarities with the Oligocene odontocetes Simocetus rayi and Ashleycetus planicapitis but displays a combination of characters that distinguish it from these and other coeval taxa. A phylogenetic analysis shows Olympicetus in a polytomy with Xenorophidae, a clade of early high-frequency-hearing odontocetes mainly known from the western Atlantic. Furthermore, Olympicetus displays characters that can be considered as structural proxies for echolocation, such as having premaxillary sac fossae and the maxilla expanded over the frontal. In addition to Olympicetus, other marine tetrapods from the Pysht Formation include marine birds, desmostylians, pinnipeds, and early mysticetes, giving us a unique insight into late Oligocene marine tetrapod faunas of the North Pacific.

oligocene pysht formation
phylogenetic analysis
xenorophidae


Evolution of Grasses and Grassland Ecosystems

Abstract: The evolution and subsequent ecological expansion of grasses (Poaceae) since the Late Cretaceous have resulted in the establishment of one of Earth's dominant biomes, the temperate and tropical grasslands, at the expense of forests. In the past decades, several new approaches have been applied to the fossil record of grasses to elucidate the patterns and processes of this ecosystem transformation. The data indicate that the development of grassland ecosystems on most continents was a multistage process involving the Paleogene appearance of (C3 and C4) open-habitat grasses, the mid-late Cenozoic spread of C3 grass-dominated habitats, and, finally, the Late Neogene expansion of C4 grasses at tropical-subtropical latitudes. The evolution of herbivores adapted to grasslands did not necessarily coincide with the spread of open-habitat grasses. In addition, the timing of these evolutionary and ecological events varied between regions. Consequently, region-by-region investigations using both direct (plant fossil...


The phylogeny of Tetanurae (Dinosauria: Theropoda)

Abstract: Tetanuran theropods represent the majority of Mesozoic predatory dinosaur diversity and the lineage leading to extant Aves. Thus their history is relevant to understanding the evolution of dinosaur diversity, Mesozoic terrestrial ecosystems, and modern birds. Previously, the fragmentary and poorly sampled fossil record of basal (non-coelurosaur) tetanurans led to uncertainties regarding their basic interrelationships. This in turn prevented determining the relationships of many incompletely known taxa that nonetheless document a global radiation spanning more than 120 million years. We undertook an exhaustive examination of all basal tetanurans and all existing character data, taking advantage of recent discoveries and adding new morphological, temporal and geographic data. Our cladistic analysis of 61 taxa achieved significantly improved phylogenetic resolution. These results position several ‘stem’ taxa basal to a succession of monophyletic clades (Megalosauroidea, Allosauroidea and Coelurosauria). Megalosauroids include nearly 20 taxa arrayed amongst a basalmost clade (Piatnitzkysauridae, fam. nov.) and the sister taxa Spinosauridae and Megalosauridae; the latter includes two subfamilies, Megalosaurinae and Afrovenatorinae subfam. nov. Allosauroidea contains a diverse Metriacanthosauridae (= Sinraptoridae), Neovenatoridae, Carcharodontosauridae and a reduced Allosauridae. Finally, we assessed more than 40 fragmentary forms and hundreds of additional reported tetanuran occurrences. Tetanuran evolution was characterized by repeated acquisitions of giant body size and at least two general skull forms, but few variations in locomotor morphology. Despite parallel diversification of multiple lineages, there is evidence for a succession of ‘dominant’ clades. Tetanurae first appeared by the Early Jurassic and was globally distributed by the Middle Jurassic. Several major clades appeared prior to the breakup of Pangaea; as such their absence in specific regions, and at later times, must be due to poor sampling, dispersal failure and/or regional extinction. Finally, we outline a general perspective on Mesozoic terrestrial biogeography that should apply to most clades that appeared before the Late Jurassic.

tetanurae
tetanurans
megalosauroidea
major clade


The revival of a so-called rotten fish: the ontogeny of the Devonian acanthodian Triazeugacanthus

Abstract: Since its original description as a chordate, the Late Devonian Scaumenella mesacanthi has been interpreted alternately as a prochordate, a larval ostracoderm and an immature acanthodian. For the past 30 years, these minute specimens were generally considered as decayed acanthodians, most of them belonging to Triazeugacanthus affinis. Among the abundant material of ‘Scaumenella’, we identified a size series of 188 specimens of Triazeugacanthus based on otolith characteristics. Despite taphonomic alteration, we describe proportional growth and progressive appearance of skeletal elements through size increase. Three ontogenetic stages are identified based on squamation extent, ossification completion and allometric growth. We demonstrate that what has been interpreted previously as various degrees of decomposition corresponds to ontogenetic changes.

ontogeny
triazeugacanthus
skeletal element


Horseshoe crab phylogeny and independent colonizations of fresh water: ecological invasion as a driver for morphological innovation

Abstract: Xiphosurids are an archaic group of aquatic chelicerate arthropods, generally known by the colloquial misnomer of ‘horseshoe crabs’. Known from marine environments as far back as the early Ordovician, horseshoe crabs are generally considered ‘living fossils’ – descendants of a bradytelic lineage exhibiting little morphological or ecological variation throughout geological time. However, xiphosurids are known from freshwater sediments in the Palaeozoic and Mesozoic; furthermore, the contention that xiphosurids show little morphological variation has never been tested empirically. Attempts to test this are hampered by the lack of a modern phylogenetic framework with which to explore different evolutionary scenarios. Here, I present a phylogenetic analysis of Xiphosurida and explore patterns of morphospace and environmental occupation of the group throughout the Phanerozoic. Xiphosurids are shown to have invaded non‐marine environments independently at least five times throughout their evolutionary history, twice resulting in the radiation of major clades – bellinurines and austrolimulids – that occupied novel regions of morphospace. These clades show a convergent ecological pattern of differentiation, speciation and subsequent extinction. Horseshoe crabs are shown to have a more dynamic and complex evolutionary history than previously supposed, with the extant species representing only a fraction of the group's past ecological and morphological diversity.

independent colonization
mesozoic
phylogenetic analysis
xiphosurida
horseshoe crab


Bolide impact triggered the Late Triassic extinction event in equatorial Panthalassa

Abstract:
Extinctions within major pelagic groups (e.g., radiolarians and conodonts) occurred in a stepwise fashion during the last 15 Myr of the Triassic. Although a marked decline in the diversity of pelagic faunas began at the end of the middle Norian, the cause of the middle Norian extinction is uncertain. Here we show a possible link between the end-middle Norian radiolarian extinction and a bolide impact. Two palaeoenvironmental events occurred during the initial phase of the radiolarian extinction interval:
(1) a post-impact shutdown of primary and biogenic silica production within a time span of 10 4 -10 5 yr, and (2) a sustained reduction in the sinking flux of radiolarian silica for ~0.3 Myr after the impact. The catastrophic collapse of the pelagic ecosystem at this time was probably the dominant factor responsible for the end-middle Norian conodont extinction.

late triassic
bolide impact
strong biotic turnover
extraterrestrial impact(s
conodont diversity change


Direct Comparisons of 2D and 3D Dental Microwear Proxies in Extant Herbivorous and Carnivorous Mammals

Abstract: The analysis of dental microwear is commonly used by paleontologists and anthropologists to clarify the diets of extinct species, including herbivorous and carnivorous mammals. Currently, there are numerous methods employed to quantify dental microwear, varying in the types of microscopes used, magnifications, and the characterization of wear in both two dimensions and three dimensions. Results from dental microwear studies utilizing different methods are not directly comparable and human quantification of wear features (e.g., pits and scratches) introduces interobserver error, with higher error being produced by less experienced individuals. Dental microwear texture analysis (DMTA), which analyzes microwear features in three dimensions, alleviates some of the problems surrounding two-dimensional microwear methods by reducing observer bias. Here, we assess the accuracy and comparability within and between 2D and 3D dental microwear analyses in herbivorous and carnivorous mammals at the same magnification. Specifically, we compare observer-generated 2D microwear data from photosimulations of the identical scanned areas of DMTA in extant African bovids and carnivorans using a scanning white light confocal microscope at 100x magnification. Using this magnification, dental microwear features quantified in 2D were able to separate grazing and frugivorous bovids using scratch frequency; however, DMTA variables were better able to discriminate between disparate dietary niches in both carnivorous and herbivorous mammals. Further, results demonstrate significant interobserver differences in 2D microwear data, with the microwear index remaining the least variable between experienced observers, consistent with prior research. Overall, our results highlight the importance of reducing observer error and analyzing dental microwear in three dimensions in order to consistently interpret diets accurately.

carnivorous mammal
pit
microwear texture analysis
dmta
dietary niche


Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Abstract: The mineral phase of the aragonite skeletal fibers of extant scleractinians (Favia, Goniastrea) examined with Atomic Force Microscope (AFM) consists entirely of grains ca. 50-100 nm in diameter separated from each other by spaces of a few nanometers. A similar pattern of nanograin arrangement was observed in basal calcite skeleton of extant calcareous sponges (Petrobiona) and aragonitic extant stylasterid coralla (Adelopora). Aragonite fibers of the fossil scleractinians: Neogene Paracyathus (Korytnica, Poland), Cretaceous Rennensismilia (Gosau, Austria), Trochocyathus (Black Hills, South Dakota, USA), Jurassic Isastraea (Ostromice, Poland), and unidentified Triassic tropiastraeid (Alpe di Specie, Italy) are also nanogranular, though boundaries between individual grains occasionally are not well resolved. On the other hand, in diagenetically altered coralla (fibrous skeleton beside aragonite bears distinct calcite signals) of the Triassic corals from Alakir Cay, Turkey (Pachysolenia), a typical nanogranular pattern is not recognizable. Also aragonite crystals produced synthetically in sterile environment did not exhibit a nanogranular pattern. Unexpectedly, nanograins were recognized in some crystals of sparry calcite regarded as abiotically precipitated. Our findings support the idea that nanogranular organization of calcium carbonate fibers is not, per se, evidence of their biogenic versus abiogenic origin or their aragonitic versus calcitic composition but rather, a feature of CaCO 3 formed in an aqueous solution in the presence of organic molecules that control nanograin formation. Consistent orientation of crystalographic axes of polycrystalline skeletal fibers in extant or fossil coralla, suggests that nanograins are monocrystalline and crystallographically ordered (at least after deposition). A distinctly granular versus an unresolvable pattern of nano-organization of CaCO 3 fibers seems to correspond, respectively, to an original versus a diagenetically depleted amount of organic matter bounding a mineral phase; this is consistent with qualitative and quantitative analyses of organic matter content in extant and fossil skeletons.


Trabecular Evidence for a Human-Like Gait in Australopithecus africanus

Abstract: Although the earliest known hominins were apparently upright bipeds, there has been mixed evidence whether particular species of hominins including those in the genus Australopithecus walked with relatively extended hips, knees and ankles like modern humans, or with more flexed lower limb joints like apes when bipedal. Here we demonstrate in chimpanzees and humans a highly predictable and sensitive relationship between the orientation of the ankle joint during loading and the principal orientation of trabecular bone struts in the distal tibia that function to withstand compressive forces within the joint. Analyses of the orientation of these struts using microCT scans in a sample of fossil tibiae from the site of Sterkfontein, of which two are assigned to Australopithecus africanus, indicate that these hominins primarily loaded their ankles in a relatively extended posture like modern humans and unlike chimpanzees. In other respects, however, trabecular properties in Au africanus are distinctive, with values that mostly fall between those of chimpanzees and humans. These results indicate that Au. africanus, like Homo, walked with an efficient, extended lower limb.

australopithecus ankle chimpanzee orientation


Tapeworm Eggs in a 270 Million-Year-Old Shark Coprolite

Abstract: Remains of parasites in vertebrates are rare from the Mesozoic and Paleozoic. Once most parasites that live in – or pass through – the gastrointestinal tract of vertebrates, fossil feces (coprolites) or even intestinal contents (enterolites) can eventually preserve their remains. Here we announce the discovery of a spiral shark coprolite from the Paleozoic bearing a cluster of 93 small oval-elliptical smooth-shelled structures, interpreted as eggs of a tapeworm.The eggs were found in a thin section of an elasmobranch coprolite. Most of the eggs are filled by pyrite and some have a special polar swelling (operculum), suggesting they are non-erupted eggs. One of the eggs contains a probable developing larva. The eggs are approximately 145–155 µm in length and 88–100 µm in width and vary little in size within the cluster. The depositional and morphological features of the eggs closely resemble those of cestodes. Not only do the individual eggs have features of extant tapeworms, but their deposition all together in an elongate segment is typical to modern tapeworm eggs deposited in mature segments (proglottids). This is the earliest fossil record of tapeworm parasitism of vertebrates and establishes a timeline for the evolution of cestodes. This discovery shows that the fossil record of vertebrate intestinal parasites is much older than was hitherto known and that the interaction between tapeworms and vertebrates occurred at least since the Middle-Late Permian.

egg
coprolite
parasitism
intestinal parasite
predation


FOSSIL VERTEBRATE TRACKS AS PALEOPENETROMETERS: CONFOUNDING EFFECTS OF FOOT MORPHOLOGY

Abstract: Abstract The depth to which a vertebrate track is indented can provide a wealth of information, being a direct result of the weight, duty factor, and limb kinematics of the animal as well as media ( =  substrate or sediment) consistency. In order to recreate the formation of the track and elucidate media consistency at the time of track formation, such factors as animal mass, duty factor, and foot morphology must be taken into consideration. This study uses Finite Element Analysis and physical modeling to demonstrate for the first time that the shape of the foot is an important factor that influences the depth to which the sediment is penetrated. In cohesive sediment, less compact morphology allows more sediment to move vertically upwards at the edges of the foot, dissipating force at the surface, and retarding transmission of load vertically down into the sediment. The reverse of this effect is seen in noncohesive sediment. Foot morphology, therefore, has a direct impact on preservation potential, both of surface tracks and undertracks, that is irrespective of the pressure exerted on the sediment surface by the foot and independent of mass and duty factor.

foot morphology
substrate
track formation


Quantifying the rise of the Himalaya orogen and implications for the South Asian monsoon

Abstract: We reconstruct the rise of a segment of the southern flank of the Himalaya-Tibet orogen, to the south of the Lhasa terrane, using a paleoaltimeter based on paleoenthalpy encoded in fossil leaves from two new assemblages in southern Tibet (Liuqu and Qiabulin) and four previously known floras from the Himalaya foreland basin. U-Pb dating of zircons constrains the Liuqu flora to the latest Paleocene (ca. 56 Ma) and the Qiabulin flora to the earliest Miocene (21- 19 Ma). The proto-Himalaya grew slowly against a high (similar to 4 km) proto-Tibetan Plateau from similar to 1 km in the late Paleocene to similar to 2.3 km at the beginning of the Miocene, and achieved at least similar to 5.5 km by ca. 15 Ma. Contrasting precipitation patterns between the Himalaya-Tibet edifice and the Himalaya foreland basin for the past similar to 56 m.y. show progressive drying across southern Tibet, seemingly linked to the uplift of the Himalaya orogen.

south
early miocene
vegetation change
himalayan orogeny


Phylogeny and biogeography of Caribbean mammals

Abstract: Vicariance and dispersal hypotheses have been proposed over the last two hundred years to explain the distribution, diversity, and faunal composition of the Caribbean biota. Despite great advances in understanding the geological history of the region, recent biogeographical reviews have not used historical biogeographical methods. In this paper I review the taxonomy, distribution and phylogeny of all Cenozoic Caribbean non-volant mammals and four bat lineages, and present reconciled trees for available phylogenies. Dates available from the fossil record and hypotheses of divergence based on molecular phylogenetic studies are also included in general assessments of fit between proposed geological models and Caribbean mammal diversification. The evidence posited in mammalian phylogenies does not add to the argument of dispersal vs. vicariance. One previously unidentified temporal pattern, the colonization of the Caribbean by South American mammals between the Palaeocene and the Middle Miocene, accounts for the distribution and phylogeny of the majority of lineages studied. Choloepodine and megalocnine sloths, hystricognath rodents, and primates all arrived during this window of colonization. Of these, megalocnine sloths, hystricognath rodents, Brachyphylla and allied bats, Stenodermatina bats, and primates fit the pattern of divergence from the mainland implied by the Gaarlandia hypothesis. Sloths, rodents and primates also roughly fit the timing of arrival to the Caribbean implied by Gaarlandia. The remaining taxa show contradictory dates of divergence according to molecular clock estimates, and no taxa fit the predicted timing and pattern of divergence among Antillean landmasses under the Gaarlandia model. Choloepodine sloths, murid rodents, insectivorans, mormoopids, and natalids show patterns of divergence from the mainland that are inconsistent with the Gaarlandia hypothesis and seem to require taxon-specific biogeographical explanations.


An exceptional record of Cambrian trilobite moulting behaviour preserved in the Emu Bay Shale, South Australia

Abstract: Trilobites dominate the Emu Bay Shale (EBS) assemblage (Cambrian Series 2, Stage 4, South Australia) in terms of numbers, with Estaingia bilobata Pocock 1964 being extremely abundant, and the larger Redlichia takooensis Lu 1950, being common. Many specimens within the EBS represent complete moulted exoskeletons, which is unusual for Cambrian fossil deposits. The abundance of complete moults provides an excellent record that has allowed the recognition of various recurrent moult configurations for both species, enabling the inference of movement sequences required to produce such arrangements. Moult configurations of E. bilobata are characterized by slight displacement of the joined rostral plate and librigenae, often accompanied by detachment of the cranidium, suggesting ecdysis was achieved by anterior withdrawal via opening of the cephalic sutures. Moulting in R. takooensis often followed the same method, but configurations show greater displacement of cephalic sclerites, suggesting more vigorous movement by the animal during moulting. Both species also show rare examples of Salter's configuration, with the entire cephalon anteriorly inverted, and several other unusual configurations. These results indicate that moulting in trilobites was a more variable process than originally thought. In contrast, other Cambrian Konservat-Lagerst€ atten with an abundance of trilobites (e.g. Wheeler Shale, USA, and Mount Stephen Trilobite Beds, Canada) show larger numbers of 'axial shields' and isolated sclerites, often interpreted as disarticulated exuviae. This points to a higher level of disturbance from factors, such as animal activity, depositional processes or water movement, compared to that of the EBS, where quiescent conditions and intermittent seafloor anoxia contributed to an unparalleled trilobite moulting record. □ Burgess Shale, Cambrian, ecdysis, Emu Bay Shale, Lagerst€ atten, moult configuration, trilobite.

trilobite
emu bay shale
exoskeleton
moult configuration
cranidium


An Early Neogene Elasmobranch fauna from the southern Caribbean (Western Venezuela)

Abstract: The Cantaure Formation (Burdigalian to ?early Langhian) is located in the Falcón Basin, North Western Venezuela, and includes one of the most diverse Neogene teleostean and benthonic invertebrate faunas in Tropical America. The paleoenvironmental preferences of the members of this fauna, as well as published paleogeographic reconstructions, suggest that the Cantaure Formation was deposited in a highly-productive shallow water environment, associated with coastal upwelling. We documented a paleodiversity of 39 shark and ray species, including 15 previously unreported taxa for Venezuela and six for Tropical America. We performed a bathymetric analysis of the fossil assemblage based on the distribution of closely-related extant chondrichthyan relatives of fossil taxa and discuss the ecological role and stratigraphic significance of the latter. Our results support the hypothesis that the Cantaure Formation was deposited in an insular inner-middle shelf environment. The elasmobranch fauna is characterized by a predominance of benthopelagic sharks with piscivorous feeding preferences (e.g., †Paratodus, Galeorhinus, Hemipristis, Rhizoprionodon, Carcharhinus, Isogomphodon, Negaprion, †Physogaleus and Sphyrna) followed by durophagous/cancritrophic feeders (e.g., Heterodontus, Nebrius, Mustelus, Rhynchobatus, Pristis, Dasyatis, cf. Pteroplatytrygon, cf. Taeniurops, Aetobatus, Aetomylaeus and Rhinoptera). Filter (e.g., Mobula and †Plinthicus), eurytrophic/sarcophagous (e.g., †Carcharocles and Galeocerdo) and teuthitrophic (e.g., Alopias) feeder species were also found. Teeth of Carcharocles megalodon found in Burdigalian sediments of the Cantaure Formation support the presence of this species already in the early Miocene. Some taxa (Nebrius, Carcharhinus cf. C. macloti and Rhynchobatus) are absent from the extant Caribbean and Western Atlantic fauna, but were present in the region before the closure of the Central American Seaway.

shark
aetomylaeus
carcharocles megalodon
cetacean bone
otodontidae


The gladiuses in coleoid cephalopods: homology, parallelism, or convergence?

Abstract: In the cephalopod subclass Coleoidea, several homology problems exist, mainly owing to unsolved phylogenetic relationships between decabrachian orders. The present contribution reviews the ''similarity'' of the gladius, the chitinous shell rudiment in the dorsal mantle that provides rigid attachment sites for the locomotory-relevant musculature. As a secretion product of the shell sac epithelium as well as in the light of a common three-layered construction, both the octobrachian and the decabrachian gladius types most probably represent homologues with identical developmental mechanisms; ''similarities'' in gladius shapes in unrelated lineages therefore should be considered as the result of parallelism. Ultrastructural comparisons with Mesozoic coleoids suggest that an organic gladius is actually embedded in every proostracum-bearing phragmocone. It is therefore generally accepted that a gladius evolved through decalcification of a proostracum-bearing phragmocone. The character ''gladius'' accordingly represents a plesiomorphy within pro-ostracum-bearing coleoids. Whereas the gladius of Vampyroteuthis as well as the octopod fin supports indirectly derived from phragmoteuthid-like phragmocone via Mesozoic gladius types, the decabrachian gladius types can morphogenetically be linked with various ancestral groups (Belemnitida, Diplobelida, Groenlandibelidae, Vasseuria, Belosepiella). Experimental decalcification of a sepiid cuttlebone demonstrates furthermore that a gladius might have also evolved from a secondarily proostracum-less phragmocone. Life styles and habitats of living and Mesozoic gladius-bearing octobrachians are finally discussed in the light of our conclusions.


The fossil record of the Neogene Carnivore Mammals from Spain

Abstract: Carnivore mammals (Carnivora, Mammalia) constitute a significant component of the Spanish Neogene faunas, not so much due to their fossil abundance, which is generally low, but rather because of their high degree of taxonomic diversity. We assessed their evolutionary dynamics from the fossil record of Iberian carnivores using per-taxon rates of origination, extinctions and turnover combined with a recent approach for removing the sampling signal from diversity curves. Our analysis interval covers most of the Neogene and the early Pleistocene (MN 2 to MN 17), spanning from around 21.6 to 1.8 Ma. We identified six carnivore associations by applying factor analysis to our data. The diversification pattern shows four origination episodes: MN4, MN7/8, MN12 and MN14. We also identified four extinction episodes. The first two are coincident with the onset of the MN 4 and MN7/8 faunas. The last two extinction episodes take place during MN9, coinciding with the Mid Vallesian Crisis (MVC), and MN13, co-occurring with the Messinian Salinity Crisis (MSC). Two major turnover pulses are recognised during MN4 and MN14, the turnover rate remaining moderately high between MN6 and MN13. We suggest that the pattern observed might be primarily triggered by the biogeographic and climatic shifts that took place during the Neogene.

mid-vallesian faunal turnover
major extinction event
iberian carnivora


Taxonomy, Phylogeny, and Functional Morphology of the Foraminiferal Genus Involutina

Abstract: Early Jurassic aragonitic foraminifers are outstandingly well-preserved in the Marmorea crust, a multiphased ferromanganese layer limiting the Schnöll and Adnet formations (Adnet, Northern Calcareous Alps, Austria). This remarkable preservation, related to the pervasive impregnation of aragonitic tests prior to their recrystallization, allowed observing unknown diagnostic features of the genus Involutina, which typifies the Suborder Involutinina. Thanks to a detailed examination of the Adnet specimens, this paper clarifies the taxonomy, systematic position, and phylogeny of Involutina. A new diagnosis, structural model, and lineage are introduced for the group. Involutina is the direct descendant of Aulotortus and the two taxa probably showed a parallel evolution. As Aulotortus, Involutina presents a high intraspecific variability and its diversity must be revised downward. Current phylogenetic and taxonomic frames of the Suborder Involutinina are firmly questioned as, contrary to previous schemes, the type-genus possesses more than one lamellar deposit per whorl. In Involutina, the height and distribution of papillae on the test surface is not random and probably related to a biological function. We here propose that the papillose lamellae and tube infoldings that characterize representatives of the genus were rudimentary features for light catching and symbiont positioning, respectively.

phylogeny
involutina
foraminifer
marmorea crust
adnet


Why are there so few evolutionary transitions between aquatic and terrestrial ecosystems

Abstract: Abstract Insects and flowering plants have rarely invaded the sea. Explanations for this have traditionally centered on the unique shortcomings of these groups in the marine environment. We show, however, that transitions among terrestrial, freshwater, and marine environments are infrequent in all major plant and animal clades except tetrapod vertebrates. In general, well-adapted incumbents are at a competitive advantage over would-be invaders from a physically different habitat. Data on the times and places of transition are consistent with our contention that evolutionary transitions among physically different environments are most likely when incumbents in the recipient environment exist in a regime of low-intensity competition and predation, as in terrestrial communities of the middle Paleozoic or the land biotas of oceanic islands. Freshwater environments, in which inferred intensities of predation are lower than in most marine and terrestrial environments, offer less biotic resistance to invaders than do communities in the sea or on land. Most invaders respond to novel physical circumstances by shutting down their metabolic machinery, and therefore add to their subordinate status as competitors with active incumbents. Only active tetrapods, particularly those with high and endothermically driven rates of metabolism, have successfully overcome this limitation.

evolutionary transition
freshwater
major plant
tetrapod
land


Global perturbation of the marine calcium cycle during the Permian-Triassic transition

Abstract: A negative shift in the calcium isotopic composition of marine carbonate rocks spanning the end-Permian extinction horizon in South China has been used to argue for an ocean acidification event coincident with mass extinction. This interpretation has proven controversial, both because the excursion has not been demonstrated across multiple, widely separated localities, and because modeling results of coupled carbon and calcium isotope records illustrate that calcium cycle imbalances alone cannot account for the full magnitude of the isotope excursion. Here, we further test potential controls on the Permian-Triassic calcium isotope record by measuring calcium isotope ratios from shallow-marine carbonate successions spanning the Permian-Triassic boundary in Turkey, Italy, and Oman. All measured sections display negative shifts in δ44/40Ca of up to 0.6‰. Consistency in the direction, magnitude, and timing of the calcium isotope excursion across these widely separated localities implies a primary and global δ44/40Ca signature. Based on the results of a coupled box model of the geological carbon and calcium cycles, we interpret the excursion to reflect a series of consequences arising from volcanic CO2 release, including a temporary decrease in seawater δ44/40Ca due to short-lived ocean acidification and a more protracted increase in calcium isotope fractionation associated with a shift toward more primary aragonite in the sediment and, potentially, subsequently elevated carbonate saturation states caused by the persistence of elevated CO2 delivery from volcanism. Locally, changing balances between aragonite and calcite production are sufficient to account for the calcium isotope excursions, but this effect alone does not explain the globally observed negative excursion in the δ13C values of carbonate sediments and organic matter as well. Only a carbon release event and related geochemical consequences are consistent both with calcium and carbon isotope data. The carbon release scenario can also account for oxygen isotope evidence for dramatic and protracted global warming as well as paleontological evidence for the preferential extinction of marine animals most susceptible to acidification, warming, and anoxia.

calcium cycle
ocean acidification
global warming
anoxia


Using experimental decay of modern forms to reconstruct the early evolution and morphology of fossil enteropneusts

Abstract: Abstract. Decay experiments are becoming a more widespread tool in evaluating the fidelity of the fossil record. Character interpretations of fossil specimens stand to benefit from an understanding of how decay can result in changes in morphology and, potentially, total character loss. We performed a decay experiment for the Class Enteropneusta to test the validity of anatomical interpretations of the Burgess Shale enteropneust Spartobranchus tenuis and to determine how the preservation of morphological features compares with the sequence of character decay in extant analogues. We used three species of enteropneust (Saccoglossus pusillus, Harrimania planktophilus, and Balanoglossus occidentalis) representing the two major families of Enteropneusta. Comparisons between decay sequences suggest that morphological characters decay in a consistent and predictable manner within Enteropneusta, and do not support the hypothesis of stemward slippage. The gill bars and nuchal skeleton were the most decay resistant, whereas the gill pores and pre-oral ciliary organ were unequivocally the most decay prone. Decay patterns support the identification of the nuchal skeleton, gill bars, esophageal organ, trunk, and proboscis in Spartobranchus tenuis and corroborate a harrimaniid affinity. Bias due to the taphonomic loss of taxonomically informative characters is unlikely. The morphologically simple harrimaniid body plan can be seen, therefore, to be plesiomorphic within the enteropneusts. Discrepancies between the sequence of decay in a laboratory setting and fossil preservation also exist. These discrepancies are highlighted not to discredit the use of modern decay studies but rather to underline their non-actualistic nature. Paleoenvironmental variables besides decay, such as the timeframe between death and early diagenesis as well as postmortem transport, are discussed relative to decay data. These experiments reinforce the strength of a comprehensive understanding of decay sequences as a benchmark against which to describe fossil taxa and understand the conditions leading to fossilization.


Dietary Change and Evolution of Horses in North America

Abstract: A survey of horse-teeth wear during the past 55 million years implies that evolutionary pressures were generally low. The evolution of high-crowned molars among horses (Family Equidae) is thought to be an adaptation for abrasive diets associated with the spread of grasslands. The sharpness and relief of the worn cusp apices of teeth (mesowear) are a measure of dietary abrasion. We collected mesowear data for North American Equidae for the past 55.5 million years to test the association of molar height and dietary abrasion. Mesowear trends in horses are reflective of global cooling and associated vegetation changes. There is a strong correlation between mesowear and crown height in horses; however, most horse paleopopulations had highly variable amounts of dietary abrasion, suggesting that selective pressures for crown height may have been weak much of the time. However, instances of higher abrasion were observed in some paleopopulations, suggesting intervals of stronger selection for the evolution of dentitions, including the early Miocene shortly before the first appearance of Equinae, the horse subfamily in which high-crowned dentitions evolved.


Evidence for a Grooming Claw in a North American Adapiform Primate: Implications for Anthropoid Origins

Abstract: Among fossil primates, the Eocene adapiforms have been suggested as the closest relatives of living anthropoids (monkeys, apes, and humans). Central to this argument is the form of the second pedal digit. Extant strepsirrhines and tarsiers possess a grooming claw on this digit, while most anthropoids have a nail. While controversial, the possible presence of a nail in certain European adapiforms has been considered evidence for anthropoid affinities. Skeletons preserved well enough to test this idea have been lacking for North American adapiforms. Here, we document and quantitatively analyze, for the first time, a dentally associated skeleton of Notharctus tenebrosus from the early Eocene of Wyoming that preserves the complete bones of digit II in semi-articulation. Utilizing twelve shape variables, we compare the distal phalanges of Notharctus tenebrosus to those of extant primates that bear nails (n = 21), tegulae (n = 4), and grooming claws (n = 10), and those of non-primates that bear claws (n = 7). Quantitative analyses demonstrate that Notharctus tenebrosus possessed a grooming claw with a surprisingly well-developed apical tuft on its second pedal digit. The presence of a wide apical tuft on the pedal digit II of Notharctus tenebrosus may reflect intermediate morphology between a typical grooming claw and a nail, which is consistent with the recent hypothesis that loss of a grooming claw occurred in a clade containing adapiforms (e.g. Darwinius masillae) and anthropoids. However, a cladistic analysis including newly documented morphologies and thorough representation of characters acknowledged to have states constituting strepsirrhine, haplorhine, and anthropoid synapomorphies groups Notharctus tenebrosus and Darwinius masillae with extant strepsirrhines rather than haplorhines suggesting that the form of pedal digit II reflects substantial homoplasy during the course of early primate evolution.


A Field Guide to Finding Fossils on Mars

Abstract: The Martian surface is cold, dry, exposed to biologically harmful radiation and apparently barren today. Nevertheless, there is clear geological evidence for warmer, wetter intervals in the past that could have supported life at or near the surface. This evidence has motivated National Aeronautics and Space Administration and European Space Agency to prioritize the search for any remains or traces of organisms from early Mars in forthcoming missions. Informed by (1) stratigraphic, mineralogical and geochemical data collected by previous and current missions, (2) Earth's fossil record, and (3) experimental studies of organic decay and preservation, we here consider whether, how, and where fossils and isotopic biosignatures could have been preserved in the depositional environments and mineralizing media thought to have been present in habitable settings on early Mars. We conclude that Noachian‐Hesperian Fe‐bearing clay‐rich fluvio‐lacustrine siliciclastic deposits, especially where enriched in silica, currently represent the most promising and best understood astropaleontological targets. Siliceous sinters would also be an excellent target, but their presence on Mars awaits confirmation. More work is needed to improve our understanding of fossil preservation in the context of other environments specific to Mars, particularly within evaporative salts and pore/fracture‐filling subsurface minerals.


THE DEMOSPONGE GENUS LEPTOMITUS and A NEW SPECIES FROM THE MIDDLE CAMBRIAN OF SPAIN

Abstract: Abstract:  Forty‐five specimens of Leptomitus conicus sp. nov. have been collected from the base of the Murero Formation (Caesaraugustan, Middle Cambrian) in the Murero area of Zaragoza, north‐east Spain. The new species is a small obconical sponge, which has coarse oxeas forming parallel rods that run the whole length of the specimens in the manner typical of the genus. The biostratigraphy and facies of the levels with Leptomitus conicus have been analysed. Palaeoecological information derived from associated trilobites, brachiopods and ichnofossils shows that L. conicus lived in a soft substrate of a sublittoral environment of low to low/moderate energy. According to established Cambrian palaeogeographical reconstructions, a tropical distribution is inferred for the genus Leptomitus. Based on body shape, wall structure and stratigraphical distribution, a phylogenetic tree of the various Leptomitus species is proposed, with a central stock of simple cylindrical sponges branching out to other morphs.

middle cambrian
sponge
trilobite
substrate


Observation of the dissolution process of Globigerina bulloides tests (planktic foraminifera) by X‐ray microcomputed tomography

Abstract: We performed a 9 day dissolution experiment with tests of the planktic foraminifer Globigerina bulloides at pH 6.7 ± 0.1 in water undersaturated with respect to calcite. The initial stage of the dissolution process, which is not recognizable from the surface structure of the tests, was quantitatively evaluated by X-ray microcomputed tomography (XMCT). XMCT revealed three distinct test structures: early-developed calcite formed during the juvenile stage of G. bulloides, an inner calcite layer, and an outer calcite layer. The test ultrastructure was observed by scanning electron microscopy, and CT number evaluated the density distribution in the test. The early-developed calcite and inner calcite layer had low CT numbers (500–1300; low density, porous) and were sensitive to dissolution, whereas the outer calcite layer had high CT numbers (<1300; high density) and resisted dissolution. Both the modes and the frequencies of the CT numbers decreased with progress of dissolution. Changes in the CT number histogram with progress of dissolution were quantified in terms of the percentage of calcite volume accounted for by low-density calcite (% Low-CT-number calcite). A clear linear relationship (R2 = 0.87) between % Low-CT-number calcite and % Test weight loss was found. This relationship indicates that the amount of test dissolution can be estimated from the distribution of CT numbers. We propose that XMCT measurements will be useful for quantitatively estimating the amount of carbonate loss from foraminiferal tests by dissolution.


Paleontological and developmental evidence resolve the homology and dual embryonic origin of a mammalian skull bone, the interparietal

Abstract: The homologies of mammalian skull elements are now fairly well established, except for the controversial interparietal bone. A previous experimental study reported an intriguing mixed origin of the interparietal: the medial portion being derived from the neural crest cells, whereas the lateral portion from the mesoderm. The evolutionary history of such mixed origin remains unresolved, and contradictory reports on the presence or absence and developmental patterns of the interparietal among mammals have complicated the question of its homology. Here we provide an alternative perspective on the evolutionary identity of the interparietal, based on a comprehensive study across more than 300 extinct and extant taxa, integrating embryological and paleontological data. Although the interparietal has been regarded as being lost in various lineages, our investigation on embryos demonstrates its presence in all extant mammalian “orders.” The generally accepted paradigm has regarded the interparietal as consisting of two elements that are homologized to the postparietals of basal amniotes. The tabular bones have been postulated as being lost during the rise of modern mammals. However, our results demonstrate that the interparietal consists not of two but of four elements. We propose that the tabulars of basal amniotes are conserved as the lateral interparietal elements, which quickly fuse to the medial elements at the embryonic stage, and that the postparietals are homologous to the medial elements. Hence, the dual developmental origin of the mammalian interparietal can be explained as the evolutionary consequence of the fusion between the crest-derived “postparietals” and the mesoderm-derived “tabulars.”

homology
interparietal
extant taxa
postparietal
tabular


Three-Dimensional Synchrotron Virtual Paleohistology: A New Insight into the World of Fossil Bone Microstructures

Abstract: Abstract The recent developments of phase-contrast synchrotron imaging techniques have been of great interest for paleontologists, providing three-dimensional (3D) tomographic images of anatomical structures, thereby leading to new paleobiological insights and the discovery of new species. However, until now, it has not been used on features smaller than 5–7 μm voxel size in fossil bones. Because much information is contained within the 3D histological architecture of bone, including an ontogenetic record, crucial for understanding the paleobiology of fossil species, the application of phase-contrast synchrotron tomography to bone at higher resolutions is potentially of great interest. Here we use this technique to provide new 3D insights into the submicron-scale histology of fossil and recent bones, based on the development of new pink-beam configurations, data acquisition strategies, and improved processing tools. Not only do the scans reveal by nondestructive means all of the major features of the histology at a resolution comparable to that of optical microscopy, they provide 3D information that cannot be obtained by any other method.


Upper Rhine Graben: Role of preexisting structures during rift evolution

Abstract: The evolution of the Cenozoic Upper Rhine Graben was controlled by a repeatedly changing stress field and the reactivation of a complex set of crustal discontinuities that had come into evidence during Permo‐Carboniferous times. A comparison of the spatial and temporal thickness distribution of synrift deposits with preexisting fault patterns permits to infer a sequence of distinct basin subsidence phases that can be related to changes in the ambient stress field. Reactivation of a system of late Palaeozoic fault systems, outlining troughs and highs, controlled the nucleation of initially separated middle and late Eocene basins, the depocenters of which coincided with a preexisting WSW‐ENE trend. During Oligocene crustal extension the individual basins coalesced, resulting in the development of the SSW‐NNE striking Upper Rhine Graben. During the late Oligocene (Chattian) change in stress field, the Upper Rhine Graben was probably reactivated as a dextral strike‐slip system with the central graben segment forming a releasing bend. During the early Miocene (Aquitanian), a major reorientation of the regional stress field is held responsible for the main subsidence phase of the northern parts of the Upper Rhine Graben. This is reflected by a counterclockwise rotation and northeastward shift of the depocenter axis and later by the middle Miocene uplift and erosion of the southern parts of the Upper Rhine Graben. During the Plio‐Quaternary, the Upper Rhine Graben was reactivated as a sinistral strike‐slip system with the central graben segment forming a restraining bend.

upper rhine graben
north sea
mediterranean
urg


Southern hemisphere biogeography inferred by event-based models: plant versus animal patterns.

Abstract: The Southern Hemisphere has traditionally been considered as having a fundamentally vicariant history. The common trans-Pacific disjunctions are usually explained by the sequential breakup of the supercontinent Gondwana during the last 165 million years, causing successive division of an ancestral biota. However, recent biogeographic studies, based on molecular estimates and more accurate paleogeographic reconstructions, indicate that dispersal may have been more important than traditionally assumed. We examined the relative roles played by vicariance and dispersal in shaping Southern Hemisphere biotas by analyzing a large data set of 54 animal and 19 plant phylogenies, including marsupials, ratites, and southern beeches (1,393 terminals). Parsimony-based tree fitting in conjunction with permutation tests was used to examine to what extent Southern Hemisphere biogeographic patterns fit the breakup sequence of Gondwana and to identify concordant dispersal patterns. Consistent with other studies, the animal data are congruent with the geological sequence of Gondwana breakup: (Africa(New Zealand(southern South America, Australia))). Trans-Antarctic dispersal (Australia <--> southern South America) is also significantly more frequent than any other dispersal event in animals, which may be explained by the long period of geological contact between Australia and South America via Antarctica. In contrast, the dominant pattern in plants, (southern South America(Australia, New Zealand)), is better explained by dispersal, particularly the prevalence of trans-Tasman dispersal between New Zealand and Australia. Our results also confirm the hybrid origin of the South American biota: there has been surprisingly little biotic exchange between the northern tropical and the southern temperate regions of South America, especially for animals.

disjunction
breakup
dispersal
biogeographic pattern
southern hemisphere


Internal conulariid structures unveiled using µCT

Abstract: An extensive sample of well-preserved conulariids from the Pennsylvanian of the North American Midcontinent (Texas and Oklahoma, USA) have been studied using X-ray micro-Computed Tomography (µCT) and have shown structures identified as longitudinal muscle bundles and a potential gastric cavity. These unequivocal structures appear in several specimens coming from different sites. Their preservation varies from a gastric cavity with muscle bundles in some individuals to only longitudinal muscle bundles in others. The muscle bundles fuse apically or medially, normally forming V-shaped pairs, and they extend along the theca/exoskeleton, parallel to the corner, towards the aperture. Longitudinal bundles have predominant perradial positions. Although there have been some articles on conulariid soft parts, most of them refer to relic soft parts. This is the first time that these structures are shown using µCT. Discovery of conulariid soft parts contributes to knowledge of metazoan evolutionary history.

conulariid
cavity
corner
longitudinal bundle
soft part


Land to sea transitions in vertebrates: the dynamics of colonization

Abstract:
Abstract.
 Vertebrates with terrestrial or freshwater ancestors colonized the sea from the Early Triassic onward and became competitively dominant members of many marine ecosystems throughout the Mesozoic and Cenozoic eras. The circumstances that led to initial marine colonization have, however, received little attention. One hypothesis is that mass extinction associated with ecosystem collapse provided opportunities for clades of amphibians, reptiles, birds, and mammals to enter marine environments. Another is that competitive pressures in donor ecosystems on land and in freshwater, coupled with abundant food in nearshore marine habitats, favored marine colonization. Here we test these hypotheses by compiling all known secondarily marine amniote clades and their times of colonization. Marine amniotes are defined as animals whose diet consists primarily of marine organisms and whose locomotion includes swimming, diving, or wading in salt water. We compared the number of clades entering during recovery phases from mass extinctions with the rate of entry of clades during nonrecovery intervals of the Mesozoic and Cenozoic. We conservatively identify 69 marine colonizations by amniotes. The only recovery interval for which prior mass extinction could have been a trigger for marine entry is the Early Triassic, when four clades colonized the sea over 7 Myr, significantly above the rates at which clades entered during other intervals. High nearshore productivity was a greater enticement to colonization than was a low diversity of potential marine competitors or predators in nearshore environments of a highly competitive terrestrial or freshwater donor biota. Rates of marine entry increased during the Cenozoic, in part because of rising productivity and in part thanks to the participation of warm-blooded birds and mammals, which broadened the range of thermal environments in which initial colonization of the sea became possible.

early triassic
marine ecosystem
predator
land


Fossil Giraffes from the Miocene of Africa and a Revision of the Phylogeny of the Giraffoidea

Abstract: Specimens of Climacoceras africanus are described from Maboko, Kenya. The new species Climacoceras gentryi is established on the basis of ossicones, mandibles, and upper and lower dentitions from Fort Ternan and Baringo, Kenya. By interpretation of its lower canines Climacoceras is identified as a giraffoid and is placed in the new family Climacoceridae. Canthumeryx sirtensis is identified from Muruarot and Rusinga, Kenya. A dentition and associated partial skeleton of this species are described. The teeth agree closely with specimens of the same species from Gebel Zelten, Libya. Zarafa zelteni from Gebel Zelten is synonymized with Canthumeryx sirtensis. Again on the basis of its lower canines Canthumeryx is identified as a giraffoid and is placed in the new family Canthumerycidae. Specimens of Palaeotragus primaevus are described from Baringo, Kenya. This material includes a cranium with the ossicones, skull roof, occipital and basicranial regions preserved. Palaeotragus primaevus specimens from Fort Ternan are used in this description and some of these are redescribed. The relations of the giraffoids are assessed by methods of phylogenetic systematics. Palaeomeryx, Prolibytherium and Propalaeoryx are excluded from the Giraffoidea as their lower canines are not known. The Palaeotraginae is shown to be an invalid polyphyletic grouping and the genus Palaeotragus is also shown to be polyphyletic. Palaeotragus microdon is probably synonymous with Palaeotragus rouenii and the three species Palaeotragus rouenii (P. microdon), Palaeotragus coelophrys and Palaeotragus quadricornis are retained in the genus Palaeotragus. It is suggested that 'Palaeotragus' expectans and 'Palaeotragus' decipiens are closely related to Samotherium. Palaeotragus primaevus is probably synonymous with Palaeotragus tungurensis and this species is closely related to the giraffines. With slight changes the subfamilies Sivatheriinae and Giraffinae are valid monophyletic groups. Hydaspitherium is synonymized with Bramatherium and the Sivatheriinae includes the genera Giraffokeryx, Birgerbohlinia, Bramatherium and Sivatherium while the Giraffinae includes the genera Honanotherium, Bohlinia and Giraffa and the species 'Palaeotragus' tungurensis (P. primaevus). Okapia is identified as the sister-group of the other giraffids. Triceromeryx is the sister-group of the Giraffidae. Canthumeryx is the sister-group of Triceromeryx plus the Giraffidae while Climacoceras is the sister-group of the other giraffoids.


An Australian multituberculate and its palaeobiogeographic implications

Abstract: A dentary fragment containing a tiny left plagiaulacoid fourth lower premolar from the Early Cretaceous (Aptian) of Victoria provides the first evidence of the Multituberculata from Australia. This unique specimen represents a new genus and species, Corriebaatar marywaltersae, and is placed in a new family, Corriebaataridae. The Australian fossil, together with meagre records of multituberculates from South America, Africa, and Madagascar, reinforces the view that Multituberculata had a cosmopolitan distribution during the Mesozoic, with dispersal into eastern Gondwana probably occurring prior to enforcement of climatic barriers (indicated by marked differentiation in regional floras) in the Early Cretaceous.

multituberculate
early cretaceous
corriebaatar marywaltersae
madagascar
gondwana


A Relict Stem Salamander: Evidence from the Early Cretaceous of Siberia

Abstract: The early evolution of salamanders, which are one of the three living groups of lissamphibians, is not well known. Both stem- and crown-group salamanders first appeared in the Middle Jurassic (Bathonian), but subsequently had different evolutionary histories: stem salamanders were thought to have gone extinct in the Late Jurassic, while crown salamanders persist to the present day. Here, I report the discovery of an indeterminate stem salamander in the Lower Cretaceous (Aptian—Albian) Ilek Formation of Western Siberia. This is new evidence that the most basal salamanders survived beyond the Jurassic—Cretaceous boundary and co-existed with crown-group salamanders during approximately the first 40 million years of the known history of salamanders. The recognition of stem salamanders in the Early Cretaceous of Western Siberia adds to the inventory of taxa that suggest this area was a refugium for various groups of vertebrates with Jurassic affinities.


ULTRASTRUCTURAL AND FUNCTIONAL MORPHOLOGY OF EGGSHELLS SUPPORTS THE IDEA THAT DINOSAUR EGGS WERE INCUBATED BURIED IN A SUBSTRATE

Abstract: Abstract:  The reproductive biology of dinosaurs is of great interest, particularly in light of the many fossil eggs assigned to this group. The ultrastructural characteristics of dinosaur eggshells are examined in order to calculate water vapour conductance, which indicates the nesting environment. Data were mainly derived from the literature but new values are also presented. Allometric analyses were carried out on a variety of shell parameters against predicted egg mass, and comparison was made with allometric equations for bird eggs. Shell thickness was generally larger than seen for extant birds. Total pore number and pores per unit area were similar to values predicted from bird eggs. Total pore area showed an isometric increase with egg mass, parallel to the relationship for birds, but the constant value was an order to magnitude higher than the bird values. Pore radius was unaffected by egg mass. Water vapour conductance showed an allometric increase with egg mass, parallel to the bird values, but for any given egg mass values for dinosaurs were an order of magnitude higher. Mass‐specific water vapour conductance was unaffected by egg mass but was an order of magnitude higher than the bird values. Water vapour conductance per pore showed an allometric decrease with egg mass but again the predicted values were an order of magnitude higher than for bird eggs. The ultrastructural characteristics of dinosaur eggshells indicate that the nesting environment had to be saturated with water vapour and that dinosaur eggs had to be fully buried in a substrate. In this sense, therefore, dinosaur eggs resemble more those of modern reptiles than those of birds. As a consequence, maintenance of incubation conditions would have depended on the prevailing environment.

egg
substrate
nesting environment
extant bird


The cranial endocast of the Middle Devonian dipnoan Dipterus valenciennesi and a fossilized dipnoan otoconial mass

Abstract: The well known Middle Devonian (Eifelian–Givetian) lungfish Dipterus valenciennesi from Scotland, UK, has been studied for more than one hundred years though our understanding of the neurocranium and cranial cavity is incomplete. Micro‐CT scanning demonstrates that the internal cast of the cranial cavity, the endocast, possesses a mix of primitive and derived characters. The olfactory bulbs are sessile, as in the derived extant Lepidosirenidae. However, Dipterus valenciennesi possesses the primitive condition of a shallow telencephalon unlike Upper Devonian forms such as Rhinodipterus kimberleyensis and extant lungfish taxa. Further information revealed by micro‐CT scanning has allowed coding for characters pertaining to the neurocranial cristae, previously unobservable in Dipterus valenciennesi, allowing hypotheses of relationships between Devonian Dipnoi and the position of Dipterus valenciennesi to be reassessed. New analyses do not refine the phylogenetic position of Dipterus valenciennesi but do increase support for most recently established Devonian dipnoan phylogeny. The first record of a three‐dimensionally preserved fossilized otoconial mass from the utricular recess in a fossil dipnoan is also described. Comparison of morphometric data of the endosseous and soft tissue manifestations of the labyrinth system in extant dipnoan taxa demonstrates that there is largely good correspondence between the two. When compared to Dipterus valenciennesi, extant taxa exhibit semi‐circular canals that are reduced in length. Furthermore, compared to Dipterus, in extant dipnoan taxa the size of the otoconial mass has increased relative to the utricular and sacculolagenar pouches containing them. The functional implications for these observations suggest that the Dipnoi have evolved towards a more sedentary lifestyle and behaviour at least since the Middle Devonian.

middle devonian
dipterus
otoconial mass
rhinodipterus kimberleyensis


Using X-ray computed tomography analysis tools to compare the skeletal element morphology of fossil and modern frog (Anura) species

Abstract: Computed tomography (CT) analysis software has predominantly been developed for medical and industrial use. Thus far there have not been detailed reports in the palaeontological literature of CT analysis tools utilizing automatic analysis functions for visual 3D comparisons between different taxa/species in order to assess and quantify differences. This paper presents the methodology for the general morphological comparison and analysis of fossil and modern bones–in this case frog femora are showcased. Industrial X-ray computed tomography analysis functions are applied to the analysis, and specifically, the comparison, of bone morphologies and micro-anatomical differences (e.g., cortical wall thickness) of differentially sized frog taxa. In contrast to traditional landmark analysis, this method makes use of the higher detail of the full surface data in the form of a CAD (Computer-aided drafting) data set output from the microCT scan. The application of these industrial analysis functions is shown to be useful for viewing of qualitative, as well as quantitative, differences. Differences are highlighted using 3D nominal/actual comparisons, and wall thickness is compared using the wall thickness analysis function applied to both samples side by side. Single bones as well as complete frog specimens were scanned and, in the case of the latter, femora were virtually extracted (segmented) for the purposes of comparison with other specimens. These techniques may be used for the effective identification and quantification of morphological differences between fossil and extant taxa. Thalassa Matthews. Iziko South African Museum, 25 Queen Victoria Street, Cape Town 8000, South Africa and DST-NRF Centre of Excellence in Palaeosciences, Evolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, Wits, 2050, South Africa. tmatthews.matthews@gmail.com Anton du Plessis. CT Scanner Facility, Central Analytical Facilities, Paul Sauer Building, Bosman Street, University of Stellenbosch, Stellenbosch, 7602, South Africa/Physics Department, University of Stellenbosch, Stellenbosch, 7602, South Africa. anton2@sun.ac.za

x-ray
whole anatomy
lissamphibian
new technology


Functional Morphology of Stylophoran Echinoderms

Abstract: The life orientation and mode of life of stylophorans are a subject of much ongoing debate. Examination of the ornamentation occurring both on the arm and theca in several cornutes and mitrates strongly supports the view that the life orientation was similar in all stylophorans and was ‘flat‐surface down’. The presence of an asymmetrical ornamentation adapted to hinder, or minimize, back slippage of the organism in all stylophorans gives strong support to their interpretation as mostly sessile organisms, feeding with the arm facing the current and the theca downstream. The examination of a wide array of thecal morphologies and sculpture patterns displayed by the various groups of cornutes and mitrates allows the identification of three main modes of life in stylophorans: (1) an epibenthic mode of life, with the theca as main anchor to the substrate (e.g. asymmetrical cornutes, Diamphidiocystis); (2) an epibenthic mode of life, with the arm as main anchor to the sediment (e.g. symmetrical cornutes, Peltocystis, primitive Mitrocystitida, some Kirkocystidae); (3) an infaunal mode of life, with the theca buried in a slightly inclined attitude (e.g. some Kirkocystidae, Mitrocystitida with cuesta‐shaped ribs). The partially buried mode of life of Lagynocystis is intermediate between 2 and 3.


Oldest Known Pantherine Skull and Evolution of the Tiger

Abstract: The tiger is one of the most iconic extant animals, and its origin and evolution have been intensely debated. Fossils attributable to extant pantherine species-lineages are less than 2 MYA and the earliest tiger fossils are from the Calabrian, Lower Pleistocene. Molecular studies predict a much younger age for the divergence of modern tiger subspecies at <100 KYA, although their cranial morphology is readily distinguishable, indicating that early Pleistocene tigers would likely have differed markedly anatomically from extant tigers. Such inferences are hampered by the fact that well-known fossil tiger material is middle to late Pleistocene in age. Here we describe a new species of pantherine cat from Longdan, Gansu Province, China, Panthera zdanskyi sp. nov. With an estimated age of 2.55–2.16 MYA it represents the oldest complete skull of a pantherine cat hitherto found. Although smaller, it appears morphologically to be surprisingly similar to modern tigers considering its age. Morphological, morphometric, and cladistic analyses are congruent in confirming its very close affinity to the tiger, and it may be regarded as the most primitive species of the tiger lineage, demonstrating the first unequivocal presence of a modern pantherine species-lineage in the basal stage of the Pleistocene (Gelasian; traditionally considered to be Late Pliocene). This find supports a north-central Chinese origin of the tiger lineage, and demonstrates that various parts of the cranium, mandible, and dentition evolved at different rates. An increase in size and a reduction in the relative size of parts of the dentition appear to have been prominent features of tiger evolution, whereas the distinctive cranial morphology of modern tigers was established very early in their evolutionary history. The evolutionary trend of increasing size in the tiger lineage is likely coupled to the evolution of its primary prey species.


A New Crurotarsan Archosaur From the Late Triassic of South Wales

Abstract: ABSTRACT We report a third archosaur from the Late Triassic cavity-fill sediments of Pant-y-ffynnon Quarry in South Wales, in addition to the basal crocodylomorph Terrestrisuchus and the basal sauropodomorph dinosaur Pantydraco. The new taxon, Aenigmaspina pantyffynnonensis, previously referred to informally as ‘Edgar,’ was recovered in 1952 and is preserved as a tight cluster of vertebrae, ribs, and a scapula in a single block, as well as numerous associated, but now isolated, elements, including skull bones, vertebrae, further putative elements of the forelimb, and a partial pelvis. We present a digital model based on computed tomography (CT) scans of the main associated blocks, which shows exquisite detail of an associated series of 12 cervical and anterior dorsal vertebrae, with ribs and a scapula. The well-developed spine tables and eight matching osteoderms confirm that this is a crurotarsan archosaur, but it lacks apomorphies of all major crurotarsan clades. Phylogenetic analysis gives equivocal results, showing weak relationships with Proterochampsidae and Ornithosuchidae; the new taxon might lie close to Erpetosuchidae, based on the ‘Y'-shaped spine tables.

archosaur
south wales
aenigmaspina pantyffynnonensis
skull bone


Global patterns in leaf 13C discrimination and implications for studies of past and future climate

Abstract: Fractionation of carbon isotopes by plants during CO2 uptake and fixation (Δleaf) varies with environmental conditions, but quantitative patterns of Δleaf across environmental gradients at the global scale are lacking. This impedes interpretation of variability in ancient terrestrial organic matter, which encodes climatic and ecological signals. To address this problem, we converted 3,310 published leaf δ13C values into mean Δleaf values for 334 woody plant species at 105 locations (yielding 570 species-site combinations) representing a wide range of environmental conditions. Our analyses reveal a strong positive correlation between Δleaf and mean annual precipitation (MAP; R2 = 0.55), mirroring global trends in gross primary production and indicating stomatal constraints on leaf gas-exchange, mediated by water supply, are the dominant control of Δleaf at large spatial scales. Independent of MAP, we show a lesser, negative effect of altitude on Δleaf and minor effects of temperature and latitude. After accounting for these factors, mean Δleaf of evergreen gymnosperms is lower (by 1–2.7‰) than for other woody plant functional types (PFT), likely due to greater leaf-level water-use efficiency. Together, environmental and PFT effects contribute to differences in mean Δleaf of up to 6‰ between biomes. Coupling geologic indicators of ancient precipitation and PFT (or biome) with modern Δleaf patterns has potential to yield more robust reconstructions of atmospheric δ13C values, leading to better constraints on past greenhouse-gas perturbations. Accordingly, we estimate a 4.6‰ decline in the δ13C of atmospheric CO2 at the onset of the Paleocene-Eocene Thermal Maximum, an abrupt global warming event ∼55.8 Ma.

leaf
annual precipitation
altitude
functional type


Postcranial material of Nothosaurus marchicus from the Lower Muschelkalk (Anisian) of Winterswijk, The Netherlands, with remarks on swimming styles and taphonomy

Abstract:
The postcranial morphology of Nothosaurus from Winterswijk is described on the basis of three partially preserved skeletons. Because of the association with a diagnostic cranium, two of these can be confidently assigned to N. marchicus. Preserved humeri are recognized as morphotype II, which confirms the relationship between this humeral morphotype and N. marchicus. The occurrence of an additional nothosaur taxon in the Lower Muschelkalk, strong sexual dimorphism, or a broader size range of N. marchicus is evidenced again. The postcranial morphology of Nothosaurus is compared to published and new data on the basal pistosauroid (cf. Cymatosaurus) from the same locality. Numerous shared morphological characters of N. marchicus, Nothosaurus sp., and the basal pistosauroid from Winterswijk hamper assignment of isolated bones, which is presently only possible if a combination of features on multiple bones from a single individual can be assessed. Ontogenetic stage of the described skeletons is discussed as well. Differences between N. marchicus, the basal pistosauroid, and the morphologically similar, but smaller pachypleurosaur A. heterodontus are mainly related to the morphologies of the humerus, ulna, and the clavicle-interclavicle complex, and thereby indicate different modes of locomotion i.e., swimming styles. The pachypleurosaur and the basal pistosauroid were most likely anguilliform swimmers, whereas Nothosaurus also employed its forelimbs during swimming (paraxial swimming). Taphonomical observations on material of the Winterswijk locality indicate the presence of scavengers, although most disarticulated skeletons were transported by water and decayed and disintegrated in shallow water.
We dedicate this paper to Wim Berkelder, Winterswijk, The Netherlands a passionate private collector of Winterswijk fossils.

swimming style
taphonomy
sexual dimorphism
winterswijk fossil
skull


Adaptive radiation of the comatulid crinoids

Abstract: Modern crinoids are dominated by the comatulids (unstalked forms) which range from the intertidal to abyssal depths. Modern stalked crinoids are restricted to depths greater than about 100 m. In the geologic past some stalked crinoids lived at depths of a few tens of meters or less in reef and bank environments. The primary vehicles postulated for the post-Triassic radiation of comatulids are lack of permanent fixation to the substratum and the capacity for mobility. Development of complex muscular articulations has enabled crawling or swimming which serve in habitat selection and avoidance of stress and predators. These and other adaptations may have bestowed on comatulids a higher survival capacity in shallow-water environments compared to stalked crinoids. Modern stalked crinoids lack mobility and complex behavioral adaptations seen in comatulids. Possibly, stalked crinoids in shallow water were unable to cope with the radiation of abundant, predaceous bony fishes in the late Mesozoic and became restricted to greater depths while the more adaptable comatulids gained ascendancy in shallow water.


Calcareous nannoplankton in the Upper Jurassic marine deposits of the Bohemian Massif: new data concerning the Boreal–Tethyan communication corridor

Abstract: Calcareous nannoplankton assemblages from the Jurassic relict deposits in the northern part of the Bohemian Massif are described here for the first time. They are generally of low diversity and dominated by watznaueriaceans. Some of them are diagenetically affected, probably due to dolomitisation. Calcareous nannoplankton enables the stratigraphical range of the Northern Bohemia Jurassic succession to be extended to the Tithonian by reference to the stratigraphical range of Jurassic platform sequences in Central Poland and the eastern part of the Bohemian Massif. The Oxfordian–Kimmeridgian nannofossil assemblages indicate a generally oligotrophic condition of the restricted sea with episodic fluvial input containing terrestrial nutrients. The character of the upper part of the water column was generally uniform and did not reflect variability at the sea-floor expressed by lithofacies diversity. The palaeoenvironment interpreted for the famous former palaeontological locality “Sternberk Quarry” was characterized by a higher nutrient content and more stable environment. The Tithonian nannofossil assemblages contain warm-water Tethyan taxa which suggest south-north migration of nannoplankton due to warming during the Jurassic–Cretaceous boundary interval.

bohemian massif
permo-carboniferous deposit
tithonian stage
calcareous nannofossil


Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medicine Formation of northwestern Montana; evidence for drought-related mortality

Abstract: Two bone beds, Canyon Bone Bed and Dino Ridge Quarry, have yielded the near-exclusive remains of a new species of Styracosaurus (Family Ceratopsidae); the third bone bed, Westside Quarry, is dominated by a new species of Prosaurolophus (Family Hadrosauridae). Evidence supporting a drought hypothesis includes: 1) a seasonal, semiarid paleoclimate, 2) associated caliche horizons, 3) aqueous depositional settings, 4) apparent age distributions characteristic of modern drought mortality (CBB and DRQ), and 5) the intraformational recurrence of low-diversity bone beds. Several alternative scenarios were considered, but drought proved most reasonable in light of the enhanced probability of preserving drought assemblages, and the species-selective and recurrent nature of modern drought mortality


Osteology of the sauropod embryos from the Upper Cretaceous of Patagonia

Abstract: Exceptionally well-preserved embryonic skulls of Upper Cretaceous (Campanian Anacleto Formation) sauropods from Auca Mahuevo (Neuquén Province, Argentina) provide important insights into the ontogeny and evolution of titanosaurian neosauropods. The most important cranial modifications occurring during titanosaurian ontogeny appear to be centered on the infraorbital and narial regions, which exhibit a substantial degree of "mosaic" evolution. On one hand, the Auca Mahuevo embryos show a large jugal that forms part of the lower margin of the skull and unretracted external nares, as indicated by the position and orientation of the lacrimals as well as the anterior extension of the frontals. Both of these features are ancestral for neosauropods, being present in prosauropods. On the other hand, the embryonic skull exhibits a large ventral notch, tentatively interpreted as homologous to the neosauropod preantorbital fenestra, that opens ventral to the jugal and between the maxilla and the quadratojugal, and a temporal region that closely resembles the adult neosauropod condition. This mosaic of character states indicates that different regions of the skull of titanosaurian neosauropods acquired their characteristic morphology at substantially different rates during their ontogenetic development.

patagonia
skull
campanian anacleto formation


Chronological reassessment of the Middle to Upper Paleolithic transition and Early Upper Paleolithic cultures in Cantabrian Spain

Abstract: Methodological advances in dating the Middle to Upper Paleolithic transition provide a better understanding of the replacement of local Neanderthal populations by Anatomically Modern Humans. Today we know that this replacement was not a single, pan-European event, but rather it took place at different times in different regions. Thus, local conditions could have played a role. Iberia represents a significant macro-region to study this process. Northern Atlantic Spain contains evidence of both Mousterian and Early Upper Paleolithic occupations, although most of them are not properly dated, thus hindering the chances of an adequate interpretation. Here we present 46 new radiocarbon dates conducted using ultrafiltration pre-treatment method of anthropogenically manipulated bones from 13 sites in the Cantabrian region containing Mousterian, Aurignacian and Gravettian levels, of which 30 are considered relevant. These dates, alongside previously reported ones, were integrated into a Bayesian age model to reconstruct an absolute timescale for the transitional period. According to it, the Mousterian disappeared in the region by 47.9–45.1ka cal BP, while the Châtelperronian lasted between 42.6k and 41.5ka cal BP. The Mousterian and Châtelperronian did not overlap, indicating that the latter might be either intrusive or an offshoot of the Mousterian. The new chronology also suggests that the Aurignacian appears between 43.3–40.5ka cal BP overlapping with the Châtelperronian, and ended around 34.6–33.1ka cal BP, after the Gravettian had already been established in the region. This evidence indicates that Neanderthals and AMH co-existed <1,000 years, with the caveat that no diagnostic human remains have been found with the latest Mousterian, Châtelperronian or earliest Aurignacian in Cantabrian Spain.

cantabrian spain
aurignacian
châtelperronian


Nest substrate reflects incubation style in extant archosaurs with implications for dinosaur nesting habits

Abstract: Dinosaurs thrived and reproduced in various regions worldwide, including the Arctic. In order to understand their nesting in diverse or extreme environments, the relationships between nests, nesting environments, and incubation methods in extant archosaurs were investigated. Statistical analyses reveal that species of extant covered nesters (i.e., crocodylians and megapodes) preferentially select specific sediments/substrates as a function of their nesting style and incubation heat sources. Relationships between dinosaur eggs and the sediments in which they occur reveal that hadrosaurs and some sauropods (i.e., megaloolithid eggs) built organic-rich mound nests that relied on microbial decay for incubation, whereas other sauropods (i.e., faveoloolithid eggs) built sandy in-filled hole nests that relied on solar or potentially geothermal heat for incubation. Paleogeographic distribution of mound nests and sandy in-filled hole nests in dinosaurs reveals these nest types produced sufficient incubation heat to be successful up to mid latitudes (≤47°), 10° higher than covered nesters today. However, only mound nesting and likely brooding could have produced sufficient incubation heat for nesting above the polar circle (>66°). As a result, differences in nesting styles may have placed restrictions on the reproduction of dinosaurs and their dispersal at high latitudes.

incubation
extant archosaur
sauropod
nest


Using Time-of-Flight Secondary Ion Mass Spectrometry to Study Biomarkers

Abstract: Time-of-flight secondary ion mass spectrometry (ToF-SIMS) is a technique designed to analyze the composition and spatial distribution of molecules and chemical structures on surfaces. These capabilities have generated much interest in its use in geobiology, in particular for the characterization of organic biomarkers (molecular biosignatures) at the microscopic level. We here discuss the strengths, weaknesses, and potential of ToF-SIMS for biomarker analyses with a focus on applications in geobiology, including biogeochemistry, organic geochemistry, geomicrobiology, and paleobiology. After describing the analytical principles of ToF-SIMS, we discuss issues of biomarker spectral formation and interpretation. Then, key applications of ToF-SIMS to soft (microbial matter, cells), hard (microbial mineral precipitates), and liquid (petroleum) samples relevant in geobiology are reviewed. Finally, we examine the potential of ToF-SIMS in biomarker research and the current limitations and obstacles for which furthe...

ion mass spectrometry
tof-sims
application


A new damsel-dragonfly from the Lower Jurassic of northwestern China and its paleobiogeographic significance

Abstract:
Abstract.
 A new species of the Lower Jurassic genus Dorsettia Whalley, 1985 is described from the Lower Jurassic Badaowan Formation of the Junggar Basin, northwestern China, as Dorsettia sinica new species. It provides additional morphological characters for this genus and is the earliest Jurassic dragonfly in China after the end-Triassic extinction. The occurrence of Dorsettia in England and northwestern China indicates that the end-Triassic extinction probably did not have a drastic influence on damsel-dragonflies, or that the dispersal of damsel-dragonflies was relatively quick during the earliest Jurassic.

damsel-dragonflie
northwestern china
end-triassic extinction
dispersal


A New Notosuchian from the Late Cretaceous of Brazil and the Phylogeny of Advanced Notosuchians

Abstract: A new notosuchian crocodyliform from the Late Cretaceous Bauru Group found in the southeastern State of São Paulo (Brazil) is described here. The new taxon, Caipirasuchus stenognathus, is referred as a new species of the recently erected genus Caipirasuchus within the clade Sphagesauridae based on a phylogenetic analysis of basal mesoeucrocodylians. Caipirasuchus stenognathus is represented by an almost complete skull and lower jaw that has autapomorphic characters that distinguish it from other species of Sphagesauridae. These autapomorphies include: maxilla forming part of the orbital margin (absence of lacrimal-jugal contact), nasal with smooth depressions on the posterior region close to the contact with the maxilla and lacrimal, postorbital with posterior palpebral facet that extends posteriorly underneath the ear-flap groove, and a distinct anterior process of the medial flange of the retroarticular process. Additionally, the new taxon lacks autapomorphic features described in other sphagesaurids. The phylogenetic analysis results in a monophyletic genus Caipirasuchus, that is the sister group of a clade fomed by Sphagesaurus huenei, Caryonosuchus pricei, and Armadillosuchus arrudai. Sphagesaurids also include a basal clade formed by Adamantinasuchus navae and Yacarerani boliviensis. Other notosuchian taxa, such as Mariliasuchus amarali, Labidiosuchus amicum, Notosuchus terrestris, and Morrinhosuchus luziae are successive sister taxa of Sphagesauridae, forming a clade of advanced notosuchians that are restricted to the Late Cretaceous of South America. These results contrast with most previous phylogenetic hypotheses of the group that depicted some members of Sphagesauridae as more closely related to baurusuchids, or found Asian (e.g., Chimaerasuchus) or African (Malawisuchus, Pakasuchus) forms nested within advanced notosuchians that are, according to our analysis, endemic of the Late Cretaceous of South America.

late cretaceous
brazil
sphagesaurid
south america
baurusuchid


METHODOLOGIES FOR THE VISUALIZATION AND RECONSTRUCTION OF THREE-DIMENSIONAL FOSSILS FROM THE SILURIAN HEREFORDSHIRE LAGERSTÄTTE

Abstract: The dominantly soft-bodied fauna of the Silurian (Wenlock) Herefordshire Lagerstätte of England is preserved as three-dimensional calcitic fossils within calcareous nodules. These fossils are difficult to interpret from study of sections produced by splitting, and are difficult also to isolate from their matrix. CT scanning of nodules is feasible, but is expensive and unlikely to be able to image the fossils at a sufficiently high resolution. We describe a technique based on serial grinding and digital photography which enables the capture of three-dimensional morphological information from the fossils. Computer video files are used to package the resulting datasets for inspection . Three-dimensional reconstructions can be created by a method involving the manual tracing of outlines on each image, but we consider this approach inappropriate in this instance. Datasets from grinding are treated instead as volumes, which are rendered either directly to produce virtual X-ray images, or by the computation of isosurfaces which are then visualized by ray-tracing. Rendering is performed from sequential angles to produce video files of rotating three-dimensional models. The method is illustrated with reconstructions of a trilobite and graptolite associated with the Herefordshire soft-bodied fauna, and of the worm-like mollusc Acaenoplax hayae. Reconstructions produced in this way can be retouched easily and manipulated to remove noise and preservational artifacts, to dissect out structures for detailed study, or to combine part and counterpart into a single reconstruction. The approach will provide the primary means of study of the Herefordshire fauna, and has the potential to provide a method of obtaining and/or communicating morphological information in a variety of palaeontological applications.

image
serial grinding
graptolite
virtual fossil
spiers software suite