Paoliida, a Putative Stem-Group of Winged Insects: Morphology of New Taxa from the Upper Carboniferous of Poland

Abstract: New representatives of a stem group Paoliida attributed to family Paoliidae (Insecta: Protoptera) are described from the Upper Carboniferous (Langsettian) sphero-sideritic concretions of the Upper Silesian Coal Basin (USCB) in Poland. Zdenekia silesiensis sp. nov. is based on forewing venation and supplemented by material of isolated hindwing similar in venation pattern. Darekia sanguinea gen. et sp. nov. differs from all other paoliid genera by the presence of a short contact between veins MP and CuA behind the division CuA and CuP. Composition of insect fauna exhibits high abundance of paoliid insects in the early Late Carboniferous ecosystems known also from other European localities such as Hagen Vorhalle in Ruhr Basin (Germany), and South Limbourg (Belgium and the Netherlands). It is the first record of true paoliids from the Polish part of paralic USCB supplementing a single historical record of Stygne roemeri considered as a taxon closely related to Paoliidae. The high abundance of paoliid insects from sphero-sideritic concretions in Sosnowiec and coal deposits previously known from the Czech part of Upper Silesian Coal Basin indicates considerable similarity of both faunas supported as well by their close stratigraphical correlation. Morphology of basal wing parts with remnants of articular sclerites preserved supports neopteran relationships of paoliids. Discovery of the first paoliid immature wing is reported suggesting similar living habitat for larvae and adults.


Rapid ocean acidification and protracted Earth system recovery followed the end-Cretaceous Chicxulub impact

Abstract: Significance Debate lingers over what caused the last mass extinction 66 million years ago, with intense volcanism and extraterrestrial impact the most widely supported hypotheses. However, without empirical evidence for either’s exact environmental effects, it is difficult to discern which was most important in driving extinction. It is also unclear why recovery of biodiversity and carbon cycling in the oceans was so slow after an apparently sudden extinction event. In this paper, we show (using boron isotopes and Earth system modeling) that the impact caused rapid ocean acidification, and that the resulting ecological collapse in the oceans had long-lasting effects for global carbon cycling and climate. Our data suggest that impact, not volcanism, was key in driving end-Cretaceous mass extinction. Mass extinction at the Cretaceous–Paleogene (K-Pg) boundary coincides with the Chicxulub bolide impact and also falls within the broader time frame of Deccan trap emplacement. Critically, though, empirical evidence as to how either of these factors could have driven observed extinction patterns and carbon cycle perturbations is still lacking. Here, using boron isotopes in foraminifera, we document a geologically rapid surface-ocean pH drop following the Chicxulub impact, supporting impact-induced ocean acidification as a mechanism for ecological collapse in the marine realm. Subsequently, surface water pH rebounded sharply with the extinction of marine calcifiers and the associated imbalance in the global carbon cycle. Our reconstructed water-column pH gradients, combined with Earth system modeling, indicate that a partial ∼50% reduction in global marine primary productivity is sufficient to explain observed marine carbon isotope patterns at the K-Pg, due to the underlying action of the solubility pump. While primary productivity recovered within a few tens of thousands of years, inefficiency in carbon export to the deep sea lasted much longer. This phased recovery scenario reconciles competing hypotheses previously put forward to explain the K-Pg carbon isotope records, and explains both spatially variable patterns of change in marine productivity across the event and a lack of extinction at the deep sea floor. In sum, we provide insights into the drivers of the last mass extinction, the recovery of marine carbon cycling in a postextinction world, and the way in which marine life imprints its isotopic signal onto the geological record.

chicxulub impact
mass extinction
rapid ocean acidification


Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus)

Abstract: The cave bear (Ursus spelaeus) was one of several spectacular megafaunal species that became extinct in northern Eurasia during the late Quaternary. Vast numbers of their remains have been recovered from many cave sites, almost certainly representing animals that died during winter hibernation. On the evidence of skull anatomy and low δ15N values of bone collagen, cave bears appear to have been predominantly vegetarian. The diet probably included substantial high quality herbaceous vegetation. In order to address the reasons for the extinction of the cave bear, we have constructed a chronology using only radiocarbon dates produced directly on cave bear material. The date list is largely drawn from the literature, and as far as possible the dates have been audited (screened) for reliability. We also present new dates from our own research, including results from the Urals. U. spelaeus probably disappeared from the Alps and adjacent areas – currently the only region for which there is fairly good evidence –c. 24 000 radiocarbon years BP (c. 27 800 cal. yr BP), approximately coincident with the start of Greenland Stadial 3 (c. 27 500 cal. yr BP). Climatic cooling and inferred decreased vegetational productivity were probably responsible for its disappearance from this region. We are investigating the possibility that cave bear survived significantly later elsewhere, for example in southern or eastern Europe.


A new Late Cretaceous leaf mine Leucopteropsa spiralae gen. et sp. nov. (Lepidoptera: Lyonetiidae) represents the first confirmed fossil evidence of the Cemiostominae

Abstract: A new fossil leaf mine ichnogenus and species, Leucopteropsa spiralae gen. et sp. nov. (Lepidoptera: Lyonetiidae), from the Upper Cretaceous Kaiparowits Formation (Campanian age, c. 76.6 to 74.5 Ma) in Utah, USA, is the earliest record (75.6 ± 0.18 Ma) and only fossil evidence of a lyonetiid-like leaf-mining moth, as well as one of the oldest known fossils within the Yponomeutoidea-Gracillarioidea clade. The blotch-style mine consists of a central oviposition, or egg laying, site with a spiral trail packed with faecal pellets that concentrically spirals outwards. The trail increases in width from oviposition site to terminus, the trajectory of the trail does not cross itself, and there is no apparent pupation chamber present. The morphology of the fossil mine is reliably associated with Cemiostominae and is most similar to mines produced by extant members of the genus Leucoptera, such as the mountain ash bent-wing moth, Leucoptera malifoliella, and the laburnum leaf miner L. laburnella. The leaf-mining moth responsible for L. spiralae may be an early member of the genus Leucoptera or other Cemiostominae genera, and the new ichnogenus Leucopteropsa is erected because the morphology, and therefore the taxonomy, of the moth specifically responsible for L. spiralae mines was not preserved in association with the leaf mine. Despite the phylogenetic analysis of 15 lyonetiids undertaken in this study and the need for further phylogenetic work for the Lepidoptera, particularly the placement of the family Lyonetiidae and the subfamily Cemiostominae within the larger Yponomeutoidea-Gracillarioidea phylogeny, this fossil provides an important Late Cretaceous (∼76 Ma) calibration point for lepidopteran phylogeny and serves as an indicator for the antiquity of the most diverse lepidopteran group, Ditrysia. Because lepidopteran body fossils are extremely rare, owing to their small and lightly sclerotized bodies, this discovery also underscores the importance of ichnofossils in the lepidopteran fossil record. http://zoobank.org/urn:lsid:zoobank.org:pub:52B20B15-5C1A-4916-9096-A490DFED7D1C


Early Gnathostome Phylogeny Revisited: Multiple Method Consensus

Abstract: A series of recent studies recovered consistent phylogenetic scenarios of jawed vertebrates, such as the paraphyly of placoderms with respect to crown gnathostomes, and antiarchs as the sister group of all other jawed vertebrates. However, some of the phylogenetic relationships within the group have remained controversial, such as the positions of Entelognathus, ptyctodontids, and the Guiyu-lineage that comprises Guiyu, Psarolepis and Achoania. The revision of the dataset in a recent study reveals a modified phylogenetic hypothesis, which shows that some of these phylogenetic conflicts were sourced from a few inadvertent miscodings. The interrelationships of early gnathostomes are addressed based on a combined new dataset with 103 taxa and 335 characters, which is the most comprehensive morphological dataset constructed to date. This dataset is investigated in a phylogenetic context using maximum parsimony (MP), Bayesian inference (BI) and maximum likelihood (ML) approaches in an attempt to explore the consensus and incongruence between the hypotheses of early gnathostome interrelationships recovered from different methods. Our findings consistently corroborate the paraphyly of placoderms, all ‘acanthodians’ as a paraphyletic stem group of chondrichthyans, Entelognathus as a stem gnathostome, and the Guiyu-lineage as stem sarcopterygians. The incongruence using different methods is less significant than the consensus, and mainly relates to the positions of the placoderm Wuttagoonaspis, the stem chondrichthyan Ramirosuarezia, and the stem osteichthyan Lophosteus—the taxa that are either poorly known or highly specialized in character complement. Given that the different performances of each phylogenetic approach, our study provides an empirical case that the multiple phylogenetic analyses of morphological data are mutually complementary rather than redundant.

jawed vertebrate
antiarch
chondrichthyan
stem gnathostome
morphological data


Body Size Distribution of the Dinosaurs

Abstract: The distribution of species body size is critically important for determining resource use within a group or clade. It is widely known that non-avian dinosaurs were the largest creatures to roam the Earth. There is, however, little understanding of how maximum species body size was distributed among the dinosaurs. Do they share a similar distribution to modern day vertebrate groups in spite of their large size, or did they exhibit fundamentally different distributions due to unique evolutionary pressures and adaptations? Here, we address this question by comparing the distribution of maximum species body size for dinosaurs to an extensive set of extant and extinct vertebrate groups. We also examine the body size distribution of dinosaurs by various sub-groups, time periods and formations. We find that dinosaurs exhibit a strong skew towards larger species, in direct contrast to modern day vertebrates. This pattern is not solely an artefact of bias in the fossil record, as demonstrated by contrasting distributions in two major extinct groups and supports the hypothesis that dinosaurs exhibited a fundamentally different life history strategy to other terrestrial vertebrates. A disparity in the size distribution of the herbivorous Ornithischia and Sauropodomorpha and the largely carnivorous Theropoda suggests that this pattern may have been a product of a divergence in evolutionary strategies: herbivorous dinosaurs rapidly evolved large size to escape predation by carnivores and maximise digestive efficiency; carnivores had sufficient resources among juvenile dinosaurs and non-dinosaurian prey to achieve optimal success at smaller body size.

vertebrate group
herbivorous dinosaur
body size distribution


A perspective on the evidence for keratin protein preservation in fossils: An issue of replication versus validation

Abstract: The preservation potential of biomolecules within vertebrate integument through deep time has recently been subject to much research and controversy. In particular, the preservation potential of proteins, such as collagen and keratin, is currently debated. Here, we examine claims from a recent study (Schweitzer et al., 2018, PLoS One), which concludes that feather keratin has a high preservation potential. We argue that this work provides insufficient evidence for protein preservation due to issues of methodology and data interpretation. Additionally, we contrast their approach and claims to those of other recently published studies in relation to the question of keratin protein preservation in fossils. We worry that most of the perceived evidence for Mesozoic polypeptide survival stems from repeated replication of methods prone to false detection, rather than triangulation by validating these claims with alternative methods that provide independent lines of evidence. When alternative explanations exist for the evidence cited as support for dinosaur proteins far exceeding their predicted preservation limits, it is most parsimonious to reject the more extreme taphonomic hypotheses. The evidence is instead more consistent with a mode of preservation in which keratinous structures do not fossilize organically as polypeptides, but rather as largely pigment and/or calcium phosphate remnants, which were originally held within the keratin matrix that is now lost. Unsupported taphonomic models (e.g., keratin polypeptide preservation) have the potential to influence our interpretation of fossil data, potentially resulting in erroneous paleobiological or evolutionary conclusions, as illustrated in another recent paper (Pan et al., 2019 , PNAS) that we also discuss.

keratin protein preservation
issue
replication
biomolecule


Small carbonaceous fossils (SCFs) from North Greenland: new light on metazoan diversity in early Cambrian shelf environments

Abstract: The Sirius Passet Lagerstätte of North Greenland is one of the oldest records of soft‐bodied metazoan‐dominated ecosystems from the early Cambrian. The Lagerstätte site itself is restricted to just a single c. 1‐km‐long outcrop located offshore from the shelf margin, in an area affected by metamorphic alteration during the Ellesmerian Orogeny (Devonian – Early Carboniferous). The recent recovery of small carbonaceous fossils (SCFs) to the south, in areas that escaped the effects of this deformation, has substantially expanded the known coverage of organic preservation into shallower water depositional settings in this region. Here, we describe additional SCF assemblages from the siliciclastic shelf succession of the Buen Formation (Cambrian Series 2, stages 3–4; c. 515 Ma), expanding the previously documented SCF biota. Newly recovered material indicates a rich diversity of non‐mineralizing metazoans, chiefly represented by arthropod remains. These include the filtering and grinding elements of a sophisticated crustacean feeding apparatus (the oldest crustacean remains reported to date), alongside an assortment of bradoriid sclerites, including almost complete, 3D valves, which tie together a number of SCFs previously found in isolation. Other metazoan remains include various trilobite cuticles, diverse scalidophoran sclerites, and a range of metazoan fragments of uncertain affinity. This shallower water assemblage differs substantially from the Sirius Passet biota, which is dominated by problematic euarthropod stem‐group members and sponges. Although some of these discrepancies are attributable to taphonomic or temporal factors, these lateral variations in taxonomic composition also point to significant palaeoenvironmental and/or palaeoecological controls on early Cambrian metazoan communities.

north greenland
scf assemblage
crustacean
small carbonaceous fossil


Anthropologically introduced biases in natural history collections, with a case study on the invertebrate paleontology collections from the middle Cambrian Spence Shale Lagerstätte

Abstract: Natural history collections are critical for modern scientific investigations, which are greatly expanding on the potential data applications of historic specimens. However, using these specimens outside their original intent introduces biases and potential misinterpretations. Anthropogenic biases can be introduced at any point during the life of museum specimens, from collection, preparation, and accession, to digitization. These biases can cause significant effects when the user is unaware of the collection context, as specific collection biases are often known anecdotally, but not ubiquitously. In this case study, the University of Kansas collection of Spence Shale Lagerstätte material was examined for anthropogenic biases using a collections inventory, interviews with stakeholders, and a literature review. Biases were found related to collector interest, locality preference, and researcher interest and specialization. These biases create a distorted view on the diversity and ecology of the Spence Shale, and need to be considered in future research. Anna F. Whitaker. Department of Chemical and Physical Sciences, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, ON L5L 1C6, Canada. annawhitaker01@gmail.com Julien Kimmig. Earth and Mineral Sciences Museum & Art Gallery, Pennsylvania State University, University Park, Pennsylvania, 16802, USA. jkimmig@psu.edu

natural history collection
case study
anthropogenic bias


A review of the ecology, palaeontology and distribution of atlantid heteropods (Caenogastropoda: Pterotracheoidea: Atlantidae)

Abstract: Fewer than 1% of marine gastropod species live a holoplanktic life. Of these, the shelled heteropods of the family Atlantidae are among the most poorly understood. The atlantids potentially make up an important part of the ocean zooplankton, composing up to 69% of shelled holoplanktic gastropods in the Late Pleistocene to Recent fossil record. They are also likely to be at high risk from current and future global changes, including anthropogenic ocean acidification. However, due to their small size (<12 mm), difficulty of sampling and complicated morphology, we still lack key information about atlantid taxonomy and ecology. This makes it difficult to understand how important they are in the ocean foodweb and how they will be affected by environmental change. Although many studies have been carried out on the atlantids, these have generally been broad and unconnected. Here, we draw together this previous research, summarizing what is currently known about atlantid taxonomy, palaeontology, ecology and biogeography, and aiming to provide a foundation for future research on this group. The data indicate complex behaviours involving seasonal and vertical migration, and demonstrate extended geographical ranges, with implications for understanding the role of atlantids in the ocean foodweb and their sensitivity to environmental changes. This review highlights the urgent need for further taxonomic research on the atlantids, including molecular analysis, and for improved sampling techniques.

ecology
palaeontology
heteropod
atlantidae
recent fossil record


Trifoliolate Leaves of Platanus bella (Heer) comb. n. from the Paleocene of North America, Greenland, and Asia and Their Relationships among Extinct and Extant Platanaceae

Abstract: Trifoliolate leaves of Platanus bella (Heer) comb. n., a species previously known only from the Paleocene of western Greenland, are newly recognized from the Paleocene of northern Wyoming, U.S.A., and Altai of Xinjiang Province, northwestern China, indicating that the species was circumboreal in the early Tertiary. Epidermal anatomy preserved in specimens from all three areas confirms that these compound leaves belong to the Platanaceae. Platanus bella (Heer) comb. n. differs from modern species of Platanus but resembles the European Tertiary species Platanus fraxinifolia (Johnson & Gilmore) Walther and Platanus neptuni (Ettingshausen) Bůžek, Holý & Z. Kvaček, in the presence of large peltate glandular trichomes. We erect a new subgenus, Glandulosa, to accommodate P. bella, P. fraxinifolia, and P. neptuni. Each of these species possesses similar leaf epidermal structure, with the characteristic platanaceous stomatal apparatus and compound hair bases. In addition, the fossils bear peltate glandular trichomes on the epidermal surfaces that are not known among extant Platanus species. Reproductive structures linked to P. neptuni indicate that subgenus Glandulosa is properly placed in the Platanaceae but that it is a distinct clade from those of the extant subgenera Platanus and Castaneophyllum Leroy. We also review the status of the fossil genera Debeya and Dewalquea, to which some Cretaceous and Tertiary leaves of similar morphology have been placed, and reject the use of either of these names to accommodate leaves of subgenus Glandulosa.


Parasites in the Fossil Record: A Cretaceous Fauna with Isopod-Infested Decapod Crustaceans, Infestation Patterns through Time, and a New Ichnotaxon

Abstract: Parasites are common in modern ecosystems and are also known from the fossil record. One of the best preserved and easily recognisable examples of parasitism in the fossil record concerns isopod-induced swellings in the branchial chamber of marine decapod crustaceans. However, very limited quantitative data on the variability of infestation percentages at the species, genus, and family levels are available. Here we provide this type of data for a mid-Cretaceous (upper Lower Cretaceous, upper Albian) reef setting at Koskobilo, northern Spain, on the basis of 874 specimens of anomurans and brachyurans. Thirty-seven specimens (4.2%), arranged in ten species, are infested. Anomurans are more heavily infested than brachyurans, variability can be high within genera, and a relationship may exist between the number of specimens and infestation percentage per taxon, possibly suggesting host-specificity. We have also investigated quantitative patterns of infestation through geological time based on 88 infested species (25 anomurans, 55 brachyurans, seven lobsters, and one shrimp), to show that the highest number of infested species can be found in the Late Jurassic, also when corrected for the unequal duration of epochs. The same Late Jurassic peak is observed for the percentage of infested decapod species per epoch. This acme is caused entirely by infested anomurans and brachyurans. Biases (taphonomic and otherwise) and causes of variability with regard to the Koskobilo assemblage and infestation patterns through time are discussed. Finally, a new ichnogenus and -species, Kanthyloma crusta, are erected to accommodate such swellings or embedment structures (bioclaustrations).

decapod crustacean
chamber
late jurassic
kanthyloma crusta
parasite


Crystallographic orientations of structural elements in skeletons of Syringoporicae (tabulate corals, Carboniferous): implications for biomineralization processes in Palaeozoic corals

Abstract: The crystallographic orientation of structural elements in skeletons of representatives of Carboniferous Syringoporicae (Auloporida) has been analysed by scanning electron microscopy (SEM), petrographic microscopy and electron backscatter diffraction (EBSD) on specimens from the Iberian Peninsula. The skeletons of the tabulate corals of the Syringoporicae consist of biogenic calcite crystals, and their microstructure is composed of lamellae, fibres and granules, or of a combination of these. Independent of the microstructure, the c‐axis is oriented towards the lumen, quasi‐perpendicular to the growth direction of the skeleton (perpendicular to the morphological axis lamellae, parallel to fibres). Most phaceloid taxa have a turbostratic distribution, as a biogenic response to prevent the cleavage of crystals. Cerioid and some phaceloid corals, whose microstructure is conditioned by wall elements, do not exhibit turbostratic distribution. Wall elements are determined by the biology of each taxon. Holacanth septal spines are composed of fibres arranged in a cone‐shape structure, sometimes clamped to the external part of the corallite and show a complex crystallography. Monacanth septal spines are spindle shaped and composed of bundles of fibres. Tabulae are composed of lamellae. Their development and crystallographic orientation depends on the position of the epithelium in each case. Shared walls are formed by a combination of the walls of two independent corallites with a median lamina, composed of granules; these have a crystallographic orientation between that of the two corallites. The growth of the microstructure is derived by a coordinated stepping mode of growth, similar to other groups of organisms such as molluscs and scleractinians. The nucleation and formation of packages of co‐oriented microcrystals suggest a growth mode similar to mineral bridges with a competitive growth mode between each crystal. The growth pattern of corallites suggests that the growth direction is divided into two main components: a horizontal growth direction towards the lumen and a vertical direction towards the top.

structural element
coral
crystallographic orientation


The eomyid rodents (Mammalia) from the Oligocene and Miocene of the Valley of Lakes (Central Mongolia)

Abstract: We report new discoveries of eomyid rodents from the Valley of Lakes (Central Mongolia) yielded by diverse layers ranging in age from the Early Oligocene (local biozone A) to the Late Miocene (local biozone E). The remains of eomyid rodents are relatively rare compared to other groups of fossil rodents found in the same region. All together, eight taxa have been identified: Eomys cf. orientalis (biozone A), Eomys aff. orientalis and Eomys sp. (biozone B), cf. Asianeomys bolligeri (biozone C), Asianeomys dangheensis (biozones C1 and D), Eomyops/ Leptodontomys sp. and Keramidomys sp. (biozone D1/2), and Omboomys builstynensis gen. nov., sp. nov. (biozone E). A progressive change of dental morphology and size indicates that the genus Asianeomys represents a local evolution of the genus Eomys in Asia, but also suggests the possible existence of a lineage including Eomys cf. orientalis, Eomys aff. orientalis, cf. Asianeomys bolligeri, and Asianeomys dangheensis.


Cope's Rule and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates

Abstract: Gigantism is widespread among Palaeozoic arthropods, yet causal mechanisms, particularly the role of (abiotic) environmental factors versus (biotic) competition, remain unknown. The eurypterids (Arthropoda: Chelicerata) include the largest arthropods; gigantic predatory pterygotids (Eurypterina) during the Siluro-Devonian and bizarre sweep-feeding hibbertopterids (Stylonurina) from the Carboniferous to end-Permian. Analysis of family-level originations and extinctions among eurypterids and Palaeozoic vertebrates show that the diversity of Eurypterina waned during the Devonian, while the Placodermi radiated, yet Stylonurina remained relatively unaffected; adopting a sweep-feeding strategy they maintained their large body size by avoiding competition, and persisted throughout the Late Palaeozoic while the predatory nektonic Eurypterina (including the giant pterygotids) declined during the Devonian, possibly out-competed by other predators including jawed vertebrates.

gigantism
eurypterid
palaeozoic vertebrate
large body size
predator


Redescription of the phytosaurs Paleorhinus (‘Francosuchus’) angustifrons and Ebrachosuchus neukami from Germany, with implications for Late Triassic biochronology

Abstract: Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London

paleorhinus
ebrachosuchus neukami
germany


Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly

Abstract: ABSTRACT The Crocodylotarsi are a group erected by Benton and Clark (1988) for archosaurs that share the “crocodile-normal” ankle structure. In this study, the phylogeny of the Crocodylotarsi was re-examined based on study of most relevant fossil material of the early non–crocodyliform members of the clade. Relationships among the major archosaurian taxa (Ornithodira, Ornithosuchidae, and Crocodylotarsi) and their proximal outgroups (Proterochampsidae, Euparkeria) were also considered. The monophyly of the Archosauria, Crurotarsi, and Crocodylotarsi is supported by the current analysis. The Parasuchia are the most plesiomorphic clade within the Crocodylotarsi. The Suchia comprise the crocodylotarsans excluding the phytosaurs. The taxon Prestosuchidae is placed near the base of this clade, as are Lewisuchus and Turfanosuchus dabanensis. The new taxon Rauisuchiformes includes the last common ancestor of Aetosauria and Crocodylia and its descendants. Rauisuchia (new combination) includes Rauisuchidae, Gracil...

crocodylotarsi euparkeria archosauria phylogeny


Hypsodonty in ungulates: an adaptation for grass consumption or for foraging in open habitat?

Abstract: Hypsodont (i.e. high-crowned) teeth have been interpreted as an indicator of feeding preferences and habitat selection in ungulates. For this reason, the degree of hypsodonty has been used for estimating the diet of ancient taxa and in palaeoenvironmental reconstructions. The goal of this study is to elucidate the relative importance of grass consumption and open habitat foraging in the development of hypsodont teeth, using novel computer techniques of knowledge discovery applied to a dataset of 134 species of artiodactyls and perissodactyls distributed among thirteen families. The results obtained suggest that highcrowned teeth represent an adaptation for feeding in an open habitat, although the minimum threshold of hypsodonty seems to increase with the relative length of the anterior part of the jaw. On the contrary, there is no direct relationship between the degree of hypsodonty and the percentage of grass consumed, except for the correspondence between grazing and dwelling in open habitats. A relatively wide muzzle evidences an adaptation for grass foraging in open and mixed habitats, but there are some non-grazing species from a closed habitat that also show wide muzzles. Thus, the hypsodonty index, combined with the length of the anterior part of the jaw and the width of the muzzle, allows accurate inferences on the ecological preferences of extinct ungulates.

ungulate
grass consumption
open habitat
hypsodont


First bird footprints from the lower Miocene Lerín Formation, Ebro Basin, Spain

Abstract: A new tracksite with bird footprints, found in the Bardenas Reales de Navarra Natural Park (Navarre, Spain), is presented in this study. The footprints are preserved in four sandstone blocks of the Lerín Formation from the northwest sector of the Ebro Basin. According to the magnetostratigraphic data, the age of these blocks is 20.4 Ma (Agenian, lower Miocene). The footprints are more than 100 mm in length, mesaxonic, and tridactyl, and have a prominent central pad impression with the digit impressions not jointed proximally. These features allow classifying them as Uvaichnites riojana. Some of the studied footprints are better preserved than the type series of Uvaichnites, which were found also in the northwest sector of the Ebro Basin. Therefore, the original diagnosis has been emended. Available chronostratigraphic data for these localities as well as for other footprints from China indicate a latest Oligocene-earliest Miocene age (from about 23 to 20 Ma) for Uvaichnites-like footprints. Sedimentological data also indicate similar continental environments, namely perilacustrine deltaic systems and distal alluvial systems. The information about early Miocene avian remains (bones, eggs and footprints) in the Iberian Peninsula is scarce. The skeletal and oological record of this age has been included within the families Phoenicopteridae, Phaisanidae and Cathartidae (or incertae sedis), while the ichnological record was related with trackmakers belonging to Charadriiformes, Ardeidae and Gruidae taxa. For this scenario, in which there are few avian remains, the ichnological diversity shown in this paper complements and improves the knowledge about the Iberian avian diversity in the early Miocene.


Feeding biomechanics suggests progressive correlation of skull architecture and neck evolution in turtles

Abstract: The origin of turtles is one of the most long-lasting debates in evolutionary research. During their evolution, a series of modifications changed their relatively kinetic and anapsid skull into an elongated akinetic structure with a unique pulley system redirecting jaw adductor musculature. These modifications were thought to be strongly correlated to functional adaptations, especially to bite performance. We conducted a series of Finite Element Analyses (FEAs) of several species, including that of the oldest fully shelled, Triassic stem-turtle Proganochelys, to evaluate the role of force distribution and to test existing hypotheses on the evolution of turtle skull architecture. We found no support for a relation between the akinetic nature of the skull or the trochlear mechanisms with increased bite forces. Yet, the FEAs show that those modifications changed the skull architecture into an optimized structure, more resistant to higher loads while allowing material reduction on specific regions. We propose that the skull of modern turtles is the result of a complex process of progressive correlation between their heads and highly flexible necks, initiated by the origin of the shell.

progressive correlation
skull architecture
turtle
performance


Radiometric dating of the type-site for Homo heidelbergensis at Mauer, Germany

Abstract: The Mauer mandible, holotype of Homo heidelbergensis, was found in 1907 in fluvial sands deposited by the Neckar River 10 km southeast of Heidelberg, Germany. The fossil is an important key to understanding early human occupation of Europe north of the Alps. Given the associated mammal fauna and the geological context, the find layer has been placed in the early Middle Pleistocene, but confirmatory chronometric evidence has hitherto been missing. Here we show that two independent techniques, the combined electron spin resonance/U-series method used with mammal teeth and infrared radiofluorescence applied to sand grains, date the type-site of Homo heidelbergensis at Mauer to 609 ± 40 ka. This result demonstrates that the mandible is the oldest hominin fossil reported to date from central and northern Europe and raises questions concerning the phyletic relationship of Homo heidelbergensis to more ancient populations documented from southern Europe and in Africa. We address the paleoanthropological significance of the Mauer jaw in light of this dating evidence.


A revision of "pediomyid" marsupials from the Late Cretaceous of North America

Abstract: Pediomyids" are a diverse group of small-to medium-sized marsupials which comprise a significant portion of many Late Cretaceous North American mammalian faunas. Known almost exclusively from isolated teeth and jaw fragments, "pediomyids" exhibit far more diversity than any other contemporaneous group of North American mammals. This has led some to suggest that the family "Pediomyidae" is an artificial, polyphyletic assemblage composed of multiple lineages that independently acquired various traditionally-recognized "pediomyid" molar characters, such as a reduction of the anterior stylar shelf, reduction of the stylocone and a labial shift in the attachment of the cristid obliqua. The present study seeks to elucidate the interrelationships of "pediomyid" marsupials and test the monophyly of the group using cladistic methodology, including a broad sampling of Late Cretaceous North American taxa and a comprehensive set of qualitative molar characters. Results suggest that the family "Pediomyidae" and the genus "Pediomys" are both polyphyletic and are in need of systematic revision. Iqualadelphis lactea (Aquilan) appears to be unrelated to the "pediomyid" radiation, and rests as a stem taxon near the base of the cladogram. The large Aquilan Aquiladelphis nests in a trichotomy with a strictly-defined "Pediomyidae" and the enigmatic Lancian taxon Glasbius, suggesting the possibility of a distant relationship (above the familial level). Three clades are recognized within the "Pediomyidae": a restricted Pediomys, Leptalestes gen. nov. (containing the three smallest species), and Protolambda (containing the remaining three larger species). Results suggest that "Pediomys" exiguus is a stem taxon lacking a close relationship to Pediomyidae sensu stricto, and is removed to permit recognition of the family as monophyletic. The results carry implications for the role "pediomyids" might have played in the initial North American marsupial radiation sometime prior to the Campanian, and the pattern of molar evolution throughout major Late Cretaceous lineages.


Biotic interchange between the Indian subcontinent and mainland Asia through time

Abstract: Biotic interchange after the connection of previously independently evolving floras and faunas is thought to be one of the key factors that shaped global biodiversity as we see it today. However, it was not known how biotic interchange develops over longer time periods of several million years following the secondary contact of different biotas. Here we present a novel method to investigate the temporal dynamics of biotic interchange based on a phylogeographical meta-analysis by calculating the maximal number of observed dispersal events per million years given the temporal uncertainty of the underlying time-calibrated phylogenies. We show that biotic influx from mainland Asia onto the Indian subcontinent after Eocene continental collision was not a uniform process, but was subject to periods of acceleration, stagnancy and decrease. We discuss potential palaeoenvironmental causes for this fluctuation.

mainland asia
biota
biotic interchange
middle miocene
molecular phylogenetic data


The Lithic Assemblages of Xiaochangliang, Nihewan Basin: Implications for Early Pleistocene Hominin Behaviour in North China

Abstract: Xiaochangliang (XCL), located in the Nihewan Basin of North China, is a key archaeological locality for understanding the behavioural evolution of early humans. XCL dates to ca. 1.36 Ma, making it one of the earliest sites in Northeast Asia. Although XCL represents the first excavation of an Early Pleistocene site in the Nihewan Basin, identified and excavated in the 1970’s, the lithic assemblages have never been published in full detail. Here we describe the lithic assemblages from XCL, providing information on stone tool reduction techniques and the influence of raw materials on artefact manufacture. The XCL hominins used both bipolar and freehand reduction techniques to manufacture small flakes, some of which show retouch. Bipolar reduction methods at XCL were used more frequently than previously recognized. Comparison of XCL with other Early Pleistocene sites in the Nihewan Basin indicates the variable use of bipolar and freehand reduction methods, thereby indicating a flexible approach in the utilization of raw materials. The stone tools from XCL and the Nihewan sites are classifiable as Mode I lithic assemblages, readily distinguished from bifacial industries manufactured by hominins in Eastern Asia by ca. 800 ka.

lithic assemblage
nihewan basin
north china


A skull of a new pelecaniform bird from the Middle Eocene of Messel, Germany

Abstract: A skull of a new pelecaniform bird is described from the Middle Eocene of Messel (Germany). Masillastega rectirostris gen. et sp. nov. is tentatively referred to the Sulidae (boobies and gannets). If this assignment is correct, the new taxon would represent the earliest fossil record of the family, preceding Sula ronzoni Milne-Edwards, 1867 from the lowermost Oligocene of France by about 15 million years. Masillastega rectirostris most distinctly differs from extant Sulidae in the proportionally longer beak which indicates that the Eocene taxon was not adapted to plunge-diving. Contrary to extant Sulidae, which are exclusively marine birds, Masillastega rectirostris was found in a freshwater deposit. It is the first pelecaniform bird known from Messel and one of the few large birds discovered at this site.

skull
new pelecaniform bird
middle eocene
sulidae


Early Oligocene chinchilloid caviomorphs from Puerto Rico and the initial rodent colonization of the West Indies

Abstract: By their past and present diversity, rodents are among the richest components of Caribbean land mammals. Many of these became extinct recently. Causes of their extirpation, their phylogenetic affinities, the timing of their arrival in the West Indies and their biogeographic history are all ongoing debated issues. Here, we report the discovery of dental remains from Lower Oligocene deposits (ca 29.5 Ma) of Puerto Rico. Their morphology attests to the presence of two distinct species of chinchilloid caviomorphs, closely related to dinomyids in a phylogenetic analysis, and thus of undisputable South American origin. These fossils represent the earliest Caribbean rodents known thus far. They could extend back to 30 Ma the lineages of some recently extinct Caribbean giant rodents (Elasmodontomys and Amblyrhiza), which are also retrieved here as chinchilloids. This new find has substantial biogeographic implications because it demonstrates an early dispersal of land mammals from South America to the West Indies, perhaps via the emergence of the Aves Ridge that occurred ca 35–33 Ma (GAARlandia hypothesis). Considering both this new palaeontological evidence and recent molecular divergence estimates, the natural colonization of the West Indies by rodents probably occurred through multiple and time-staggered dispersal events (chinchilloids, then echimyid octodontoids (spiny rats/hutias), caviids and lastly oryzomyin muroids (rice rats)).


Intraskeletal histovariability, allometric growth patterns, and their functional implications in bird-like dinosaurs

Abstract:
With their elongated forelimbs and variable aerial skills, paravian dinosaurs, a clade also comprising modern birds, are in the hotspot of vertebrate evolutionary research. Inferences on the early evolution of flight largely rely on bone and feather morphology, while osteohistological traits are usually studied to explore life-history characteristics. By sampling and comparing multiple homologous fore-and hind limb elements, we integrate for the first time qualitative and quantitative osteohistological approaches to get insight into the intraskeletal growth dynamics and their functional implications in five paravian dinosaur taxa, Anchiornis, Aurornis, Eosinopteryx, Serikornis, and Jeholornis. Our qualitative assessment implies a considerable diversity in allometric/isometric growth patterns among these paravians. Quantitative analyses show that neither taxa nor homologous elements have characteristic histology, and that ontogenetic stage, element size and the newly introduced relative element precocity only partially explain the diaphyseal histovariability. Still, Jeholornis, the only avialan studied here, is histologically distinct from all other specimens in the multivariate visualizations raising the hypothesis that its bone tissue characteristics may be related to its superior aerial capabilities compared to the nonavialan paravians. Our results warrant further research on the osteohistological correlates of flight and developmental strategies in birds and bird-like dinosaurs.
Studying bone histology of extinct paravian dinosaurs (all dinosaurs closer to modern birds than to Oviraptor 1 ) is a powerful tool to infer the biology of these feathered theropods and to better understand the early evolution of birds. Previous studies focusing on ontogenetic stage, growth rate, age, and sexual maturation in dinosaur-bird transitional forms either based their conclusions on the histology of a single element, usually the femur 2-4 , or multiple, but often non-homologous bones were investigated across different studies 5-10 rendering interspecific comparisons difficult. As osteohistological traits vary extensively among different individuals as well as within the skeleton [11] [12] [13] , single-element analyses and comparison of non-homologous bones may lead to different interpretations on the overall growth patterns depending on the elements studied. By contrast, assessing the extent of intraskeletal histovariability gives a more complete picture on ontogenetic stages and growth trajectories, and provides new insights into allometric skeletal growth dynamics and potentially locomotor aspects. Limb morphometrics combined with growth dynamics-related osteohistological traits have been used to infer functional shifts among different elements during the ontogeny in Psittacosaurus 14 . However, up to now, no attempt was made for applying this approach to explore the interelemental dynamics of skeletal development in paravian dinosaurs.
Apart from lineages with secondarily reduced forelimbs, paravian dinosaurs are characterized by disproportionately long and robust arms compared with other theropods. Contrasting with other bipedal dinosaurs, the length of the humerus, radius and ulna in these paravians is comparable with, or may even exceed, that of the femur, and the total forelimb to hind limb length ratio is close to or exceeds one [15] [16] [17] [18] . This implies heterochronic allometry with the fore-and hind limb elements having differential growth rates in different phases of ontogeny, which may also reflect locomotor aspects of the precocial -altricial developmental spectrum of ontogenetic

growth pattern
bird-like dinosaur
ontogenetic stage
intraskeletal histovariability


Astragalus of Pondaungimys (Rodentia, Anomaluroidea) from the late middle Eocene Pondaung Formation, central Myanmar

Abstract: ABSTRACT A well-preserved astragalus of the anomaluroid rodent Pondaungimys anomaluropsis is described from the late middle Eocene Pondaung Formation of central Myanmar. This specimen is the first postcranial element of a rodent from the Pondaung Formation and the oldest postcranial fossil currently known for Anomaluroidea. It illuminates plesiomorphous postcranial conditions in early anomaluroids and provides a basis for reconstructing the evolution of arboreal locomotion in this group of rodents. In contrast to those of living anomaluroids, the astragalus of Pondaungimys bears features indicating a less mobile ankle, specifically including a reduced range of plantarflexion at the upper ankle joint and a diminished capacity for inversion at the lower ankle joint. Its anatomy suggests that early anomaluroids were generalized quadrupeds, with intermediate arboreal adaptations between those of paramyids and modern anomaluroids. A cladistic analysis based on astragalar characters corroborates a basal position for Pondaungimys among anomaluroids. The phylogenetic signal derived from astragalar morphology is consistent with recent assessments of relationships among living anomaluroids based on craniodental characters and molecular data. Stem anomaluroids such as Pondaungimys apparently lacked the ability to glide, a locomotor pattern that is retained in extant Zenkerella.


Molecular analysis of black coatings and anointing fluids from ancient Egyptian coffins, mummy cases, and funerary objects

Abstract: Significance Previous studies of ritual black coatings on coffins from ancient Egypt have taken single small samples from objects of wide-ranging time periods and have conflated different types of application. This study takes 100 samples of black ritual liquids identified by type of application and precisely located on the objects, to allow the results to be fully contextualized within the wider discipline. It shows that black coatings on coffins were made using a remarkably consistent selection of natural products. The molecules identified in these black coatings overlap with those used in Egyptian mummification balms, which may suggest a link among separate applications: preparation of the body for burial, decoration of the coffin, and rites performed during the funeral. Black organic coatings and ritual deposits on ancient Egyptian coffins and cartonnage cases are important and understudied sources of evidence about the rituals of funerary practice. Sometimes, the coatings were applied extensively over the surface of the coffin, resembling paint; in other cases, they were poured over the mummy case or wrapped body, presumably as part of a funerary ritual. For this study, multiple samples of black coatings and ritual liquids were taken from 20 Egyptian funerary items dating to a specific time period (c. 943 to 716 BC). Multiple sampling from each object enabled several comparisons to be made: the variability of the black coating within one application, the variability between two applications on one object, and the variability from object to object. All samples were analyzed for lipids using gas chromatography–mass spectrometry (GC-MS), and 51 samples from across the 20 items were further analyzed for the presence of bitumen using solid phase separation followed by selected ion monitoring GC-MS. The majority of the black substances were found to comprise a complex mixture of organic materials, including bitumen from the Dead Sea, conifer resin, and Pistacia resin, providing evidence for a continuation in international trade between Egypt and the eastern Mediterranean after the Late Bronze Age. Both the coating and the anointing liquid are very similar to mummification balms, pointing to parallels with Egyptian embalming rituals and raising questions about the practical aspects of Egyptian funerary practice.

black coating
ancient egyptian coffin
mummy case
lipid


A revision of European Plesiosminthus (Rodentia, Dipodidae), and new material from the upper Oligocene of Teruel (Spain)

Abstract: The European record of the rodent genus Plesiosminthus is revised, and two new species are described. Plesiosminthus promyarion is found to be a nomen dubium in the sense of the International Code of Zoological Nomenclature and that name should therefore no longer be used. Until now the genus was hardly known from localities older than MP28; for the first time we describe rich samples from the upper Oligocene (MP 27) of Spain. By now, 10 European species are known, one from the lower Miocene, and the other ones from the upper Oligocene. For most of them it is not possible to establish ancestor-descendant relationships.

european plesiosminthus
upper oligocene
stratigraphical range
mosaic history


Systematics, morphology, and appendages of Anacheirurus (Pilekiinae, Trilobita) from the Fezouata Shale and the early diversification of Cheiruridae

Abstract: Pilekiines are the earliest diverging members of the successful trilobite family Cheiruridae. The pilekiine genus Anacheirurus is characterized by sub-quadratic to sub-oval glabella, pitted genae, and a distinct trunk with elongated pleural spines in its posterior part. Anacheirurus adserai is a common component of the Fezouata Shale (Lower Ordovician, Morocco), where it was intially included into several species of the genus Lehua. This assignment and taxonomic over-splitting created confusion, overestimated cheirurid diversity at this locality, and simultaneously underestimated morphological variability within A. adserai. In this contribution we examine new material of A. adserai from the Fezouata Shale, clarifying its morphology and systematics. A detailed re-description of the species shows that Anacheirurus is distinct from Lehua, the latter being a more derived member of Cheiruridae. The comparison of Anacheirurus with other pilekiines shows that morphological variability within this subfamily is mostly constrained to the trunk region. Exceptionally preserved specimens of A. adserai from the Fezouata Shale show details of appendages, revealing the endopodite and exopodite morphologies in early members of Cheiruridae. The endopodite of A. adserai is unique among trilobites in possessing comparatively longer distal podomeres 5 and 6, but otherwise, it has the same general morphology as other described trilobite endopodites. The exopodite morphology of A. adserai shows characters typical of some Cambrian species but differs in several aspects from those known in post-Cambrian taxa. It is concluded that trilobite exopodite morphology was probably more variable than the endopodite morphology, which remains rather conservative across different taxa. Morphological diversity of trilobite exopodites in post-Cambrian taxa might be related to ecological escalations during the Ordovician biodiversification and the transition between Cambrian and Ordovician trilobite faunas.

anacheirurus
fezouata shale
cheiruridae
trilobite
long locomotory endopod


The skull and pectoral girdle of the parasemionotid fish Watsonulus eugnathoides from the Early Triassic Sakamena Group of Madagascar, with comments on the relationships of the holostean fishes

Abstract: ABSTRACT Watsonulus eugnathoides (Piveteau, 1935) is a parasemionotid fish from Early Triassic rocks of Madagascar. The skull and pectoral girdle of this holostean are described from new material. The braincase retains a number of primitive chondrostean-like characters such as an open lateral cranial fissure and frequently open vestibular fontanelle, presence of an endochondral intercalar without membranous outgrowths, and fusion between most of the the endochondral bones in the adult, but is otherwise similar to “caturids” such as Heterolepidotus. The dermal skull retains one major, putatively primitive, feature shared with chondrosteans, a preopercular with a broad dorsal edge. The dermal shoulder girdle retains a chondrostean-style clavicle, but the endochondral shoulder girdle is most similar to teleosts among actinopterygians. Watsonulus also has an Amia-style jaw joint. The non-reduced clavicle and dorsally expanded preoperculum are lost in all other non-parasemionotid neopte-rygians (the reductions...


The unexpected survival of an ancient lineage of anseriform birds into the Neogene of Australia: the youngest record of Presbyornithidae

Abstract: Presbyornithids were the dominant birds in Palaeogene lacustrine assemblages, especially in the Northern Hemisphere, but are thought to have disappeared worldwide by the mid-Eocene. Now classified within Anseriformes (screamers, ducks, swans and geese), their relationships have long been obscured by their strange wader-like skeletal morphology. Reassessment of the late Oligocene South Australian material attributed to Wilaru tedfordi, long considered to be of a stone-curlew (Burhinidae, Charadriiformes), reveals that this taxon represents the first record of a presbyornithid in Australia. We also describe the larger Wilaru prideauxi sp. nov. from the early Miocene of South Australia, showing that presbyornithids survived in Australia at least until ca 22 Ma. Unlike on other continents, where presbyornithids were replaced by aquatic crown-group anatids (ducks, swans and geese), species of Wilaru lived alongside these waterfowl in Australia. The morphology of the tarsometatarsus of these species indicates that, contrary to other presbyornithids, they were predominantly terrestrial birds, which probably contributed to their long-term survival in Australia. The morphological similarity between species of Wilaru and the Eocene South American presbyornithid Telmabates antiquus supports our hypothesis of a Gondwanan radiation during the evolutionary history of the Presbyornithidae. Teviornis gobiensis from the Late Cretaceous of Mongolia is here also reassessed and confirmed as a presbyornithid. These findings underscore the temporal continuance of Australia’s vertebrates and provide a new context in which the phylogeny and evolutionary history of presbyornithids can be examined.


Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony

Abstract: A “beautiful chimera” from the mid-Cretaceous sheds light on the evolution of novel forms and forces a rethink of what a crab is. Evolutionary origins of novel forms are often obscure because early and transitional fossils tend to be rare, poorly preserved, or lack proper phylogenetic contexts. We describe a new, exceptionally preserved enigmatic crab from the mid-Cretaceous of Colombia and the United States, whose completeness illuminates the early disparity of the group and the origins of novel forms. Its large and unprotected compound eyes, small fusiform body, and leg-like mouthparts suggest larval trait retention into adulthood via heterochronic development (pedomorphosis), while its large oar-like legs represent the earliest known adaptations in crabs for active swimming. Our phylogenetic analyses, including representatives of all major lineages of fossil and extant crabs, challenge conventional views of their evolution by revealing multiple convergent losses of a typical “crab-like” body plan since the Early Cretaceous. These parallel morphological transformations may be associated with repeated invasions of novel environments, including the pelagic/necto-benthic zone in this pedomorphic chimera crab.

marine arthropod
novel form
crab
exceptional preservation


The deforestation of Easter Island

Abstract: Easter Island deforestation has traditionally been viewed as an abrupt island‐wide event caused by the prehistoric Rapanui civilization, which precipitated its own cultural collapse. This view emerges from early palaeoecological analyses of lake sediments, which showed a sudden and total replacement of palm pollen by grass pollen shortly after Polynesian settlement (800–1200 CE). However, further palaeoecological research has challenged this view, showing that the apparent abruptness and island‐wide synchronicity of forest removal was an artefact due to the occurrence of a sedimentary gap of several millennia that prevented a detailed record of the replacement of palm‐dominated forests by grass meadows. During the last decade, several continuous (gap‐free) and chronologically coherent sediment cores encompassing the last millennia have been retrieved and analysed, providing a new picture of forest removal on Easter Island. According to these analyses, deforestation was not abrupt but gradual and occurred at different times and rates, depending on the site. Regarding the causes, humans were not the only factors responsible for forest clearing, as climatic droughts as well as climate–human–landscape feedbacks and synergies also played a role. In summary, the deforestation of Easter Island was a complex process that was spatially and temporally heterogeneous and took place under the actions and interactions of both natural and anthropogenic drivers. In addition, archaeological evidence shows that the Rapanui civilization was resilient to deforestation and remained healthy until European contact, which contradicts the occurrence of a cultural collapse. Further research should aim to obtain new continuous cores and make use of recently developed biomarker analyses to advance towards a holistic view of the patterns, causes and consequences of Easter Island deforestation.

deforestation
easter island
drought
european contact


Skeletal material from larger Eusauropterygia (Reptilia: Eosauropterygia) with nothosaurian and cymatosaurian affinities from the Lower Muschelkalk of Winterswijk, The Netherlands

Abstract: Eusauropterygian elements from the Lower Muschelkalk (Vossenveld Formation) of Winterswijk, The Netherlands, comprising three mandibles and a selection of postcranial material, are described. The new mandibles correspond to crania larger than eusauropterygian cranial material previously known from this locality. The postcranial material represents a selection of isolated and associated material of comparably large size. Two of the mandibles conform to the morphology commonly recognized in Nothosaurus from the Lower Muschelkalk, but the third mandible accommodates five fangs per ramus in the symphyseal domain and exhibits a symphyseal ratio of 1.04, a condition not yet described for Eusauropterygia from the Vossenveld Formation. Body length for the animals represented by the finds was estimated through dimensional comparison with MB.R.27, a nearly complete skeleton from the lower Middle Muschelkalk locality of Ru ¨dersdorf (Germany) assigned to Nothosaurus marchicus. Body length estimates for the individuals represented by the largest mandible and by a selection of associated postcranial material exceed 1,500 mm; approximately 50 % larger than the eusauropterygian taxa from Winterswijk described so far. Furthermore, certain morphological characters identified in one mandible and in the postcranial elements suggest a cymatosaurian affinity over a nothosaurian affinity. The present study indicates eusauropterygian diversity during the Lower Muschelkalk was greater than traditionally believed. This challenges the concept of a slow paced, gradual biotic response to the origination and expansion of the Muschelkalk Sea, which commenced only 5 million years after the Permian-Triassic mass extinction event.

eusauropterygia
cymatosaurian affinity
lower muschelkalk


Calcified Pulmonary Nodules Identified in a 350-Year-Old-Joseon Mummy: the First Report on Ancient Pulmonary Tuberculosis from Archaeologically Obtained Pre-modern Korean Samples

Abstract: We found calcified pulmonary nodules in a middle-aged female mummy discovered from 350-yr-old Joseon tomb of Korea. In the CT scan, we found six radiopaque nodules in right lung, through the levels of thoracic vertebrae 1 to 6. We also found presumptive pleural adhesions in right thoracic cavity of CT images. We re-confirmed radiological findings by our post-factum dissection on the same mummy. By the differential diagnosis, we speculate that the radiopaque calcification nodules and associated pleural adhesion could have been caused by tuberculosis. This is the first-ever report on the pulmonary tuberculosis identified in archaeologically obtained, pre-modern Korean samples.

pulmonary nodule
ancient pulmonary tuberculosis
pre-modern korean sample


A new odontocete (toothed cetacean) from the Early Miocene of Peru expands the morphological disparity of extinct heterodont dolphins

Abstract: A key step in the evolutionary history of Odontoceti (echolocating toothed cetaceans) is the transition from the ancestral heterodont condition – characterized by the presence of double-rooted cheek teeth bearing accessory denticles – to the homodont dentition displayed by most extant odontocete species. During the last few decades, new finds and the reassessment of specimens in collections revealed an increased morphological disparity amongst the Oligo–Miocene heterodont odontocetes. Based on a partly articulated skeleton from late Early Miocene (Burdigalian, 18.8–18.0 Ma) beds of the Chilcatay Formation (Pisco Basin, Peru), we describe a new genus and species of heterodont odontocete, Inticetus vertizi, in the new family Inticetidae. This large dolphin is characterized by, amongst other things, a long and robust rostrum bearing at least 18 teeth per quadrant; the absence of procumbent anterior teeth; many large, broad-based accessory denticles in double-rooted posterior cheek teeth; a reduced ornament of dental crowns; the styliform process of the jugal being markedly robust; a large fovea epitubaria on the periotic, with a correspondingly voluminous accessory ossicle of the tympanic bulla; and a shortened tuberculum of the malleus. Phylogenetic analyses (with and without molecular constraint; with and without down-weighting of homoplastic characters) yielded contrasting results, with Inticetus falling either as a stem Odontoceti or as an early branching member of a large Platanistoidea clade. With its large size, robust rostrum and unusual dental morphology, and the absence of conspicuous tooth wear, Inticetus increases the morphological and ecological disparity of Late Oligocene–Early Miocene heterodont odontocetes. Finally, this new taxon calls for caution when attempting to identify isolated cetacean cheek teeth, even at the suborder level. http://zoobank.org/urn:lsid:zoobank.org:pub:5B306B49-EB1B-42F9-B755-B0B05B4F938F

odontocete
cetacean
early miocene
phylogenetic analysis


Post-Boreotropical dispersals explain the pantropical disjunction in Paederia (Rubiaceae).

Abstract:
BACKGROUND AND AIMS
Pantropical intercontinental disjunction is a common biogeographical pattern in flowering plants exhibiting a discontinuous distribution primarily in tropical Asia, Africa and the Americas. Only a few plant groups with this pattern have been investigated at the generic level with molecular phylogenetic and biogeographical methods. Paederia (Rubiaceae) is a pantropical genus of 31 species of woody lianas, with the greatest species diversity in continental Asia and Madagascar and only two species from tropical America. The aim of this study was to reconstruct the biogeographical history of Paederia based on phylogenetic analyses to explore how the genus attained its pantropical distribution.

METHODS
Maximum parsimony and Bayesian inference were used for phylogenetic analyses using sequences of five plastid markers (the rbcL gene, rps16 intron, trnT-F region, atpB-rbcL spacer and psbA-trnH spacer). Biogeographical inferences were based on a Bayesian uncorrelated lognormal relaxed molecular clock together with both Bayesian and likelihood ancestral area reconstructions.

KEY RESULTS
The data suggest an early diverged Asian lineage sister to the clade of the remaining species consisting of a predominantly Asian sub-clade and a primarily Malagasy sub-clade. Paederia is inferred to have originated in the Oligocene in tropical continental Asia. It then reached Africa in the early to middle Miocene, most probably via long-distance dispersal across the Indian Ocean. The two Neotropical species are inferred to have derived independently in the late Miocene from ancestors of Asia and East Africa, respectively.

CONCLUSIONS
The results demonstrate the importance of post-Boreotropical long-distance dispersals (across three major oceans) in shaping the global pantropical disjunction in some plants, such as Paederia, with small, winged diaspores adapted to long-distance dispersal by various agents including wind, ocean currents or birds. Overland migration is less likely to explain its palaeotropical disjunction between Asia and Africa.


A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny

Abstract: Characterized by atypical frontalmost appendages, leanchoiliids are early arthropods whose phylogenetic placement has been much debated. Morphological interpretations have differed, some of which concern critical characters such as the number of eyes and head appendages, but methodological approaches also have diverged. Here, we describe a new leanchoiliid, Yawunik kootenayi gen. et sp. nov., based on 42 specimens from the newly discovered Marble Canyon locality of the Burgess Shale (Kootenay National Park, British Columbia; middle Cambrian). This new morphotype demonstrates the presence of a four‐segmented head in leanchoiliids, along with two small antero‐median eyes in addition to lateral eyes. Yawunik is characterized by a 12‐segmented trunk and a carinate, lanceolate telson adorned with minute spines. The ‘great appendages’ of the animal bear teeth on their two distal rami, which would have enhanced their ability to grasp prey. Attitudes of specimens, resulting from burial at multiple aspects of bedding, suggest the ‘great appendages’ were flexible and capable of antero‐posterior rotation. We also discuss the nature of intersegmental tissues and filaments present within the ‘great appendages’. Our phylogenetic analyses extend the monophyly of leanchoiliids to include Haikoucaris and Yohoia in a new clade, the Cheiromorpha nom. nov. (within Heptopodomera nom. nov.). Other nodes are poorly resolved unless implied weights are used, and in this case, the topology is critically sensitive to the coding prerogative of inapplicable states (NAs). Both the traditional ‘Arachnomorpha’ hypothesis (NAs as additional states) and the more recently favoured ‘Artiopoda + Crustacea’ (NAs as uncertainties) were obtained using the same data set and outgroup. This result stresses, first, the historical importance of polarization over data content in scenarios of early arthropod evolution, and second, a pressing need to investigate the impacts of coding inapplicables, especially given the inflating effect of implied weights.

new leanchoiliid
burgess shale
arthropod
phylogeny


Taphonomy and affinity of an enigmatic Silurian vertebrate, Jamoytius kerwoodi White

Abstract: Abstract:  The anatomy and affinities of Jamoytius kerwoodi White have long been controversial, because its complex taphonomy makes unequivocal interpretation impossible with the methodology used in previous studies. Topological analysis, model reconstruction and elemental analysis, followed by anatomical interpretation, allow features to be identified more rigorously and support the hypothesis that Jamoytius is a jawless vertebrate. The preserved features of Jamoytius include W‐shaped phosphatic scales, 10 or more pairs of branchial openings, optic capsules, a circular, subterminal mouth and a single terminal nasal opening. Interpretations of paired ‘appendages’ remain equivocal. Phylogenetic analysis places Jamoytius and Euphanerops together (Jamoytiiformes), as stem‐gnathostomes rather than lamprey related or sister taxon to Anaspida.

vertebrate
jamoytius kerwoodi white
taphonomy


Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: The avian brain shape left other dinosaurs behind

Abstract: Ecological or sensory system shifts associated with brain shape and size change may have contributed to unique avian survivorship. Birds today are the most diverse clade of terrestrial vertebrates, and understanding why extant birds (Aves) alone among dinosaurs survived the Cretaceous-Paleogene mass extinction is crucial to reconstructing the history of life. Hypotheses proposed to explain this pattern demand identification of traits unique to Aves. However, this identification is complicated by a lack of data from non-avian birds. Here, we interrogate survivorship hypotheses using data from a new, nearly complete skull of Late Cretaceous (~70 million years) bird Ichthyornis and reassess shifts in bird body size across the Cretaceous-Paleogene boundary. Ichthyornis exhibited a wulst and segmented palate, previously proposed to have arisen within extant birds. The origin of Aves is marked by larger, reshaped brains indicating selection for relatively large telencephala and eyes but not by uniquely small body size. Sensory system differences, potentially linked to these shifts, may help explain avian survivorship relative to other dinosaurs.

mass extinction
brain shape
other dinosaur
extant bird
ichthyornis


Ancient DNA from pollen: a genetic record of population history in Scots pine

Abstract: Assessments of plant population dynamics in space and time have depended on dated records of fossil pollen synthesized on a subcontinental scale. Genetic analyses of extant populations have revealed spatial relationships that are indicative of past spatial dynamics, but lack an explicit timescale. Synthesis of these data requires genetic analyses from abundant dated fossil material, and this has hitherto been lacking. Fossil pollen is the most abundant material with which to fill this data gap. Here we report genetic analyses of fossil pollen retrieved from Holtjärnen postglacial lake sediment in Sweden and show that plastid DNA is recoverable from Scots Pine and Norway spruce pollen grains that are 100 and 10 000 years old. By sequencing clones from two short plastid PCR products and by using multiple controls we show that the ancient sequences were endogenous to the fossil grains. Comparison of ancient sequences and those obtained from an extant population of Scots pine establishes the first genetic link between extant and fossil samples in this species, providing genetic continuity through time. The finding of one common haplotype present in modern, 100‐year old and 10 000‐year old samples suggests that it may have persisted near Holtjärnen throughout the postglacial period. This retrieval of ancient DNA from pollen has major implications for plant palaeoecology in conifer species by allowing direct estimates of population dynamics in space and time.

pollen
scots pine
sweden
common haplotype
ancient dna


Origins and early evolution of arthropods

Abstract: Phylogenomics reconstructs an arthropod tree in which a monophyletic Arthropoda splits into Pycnogonida + Euchelicerata and Myriapoda + Pancrustacea. The same chelicerate–mandibulate groups are retrieved with morphological data sets, including those encompassing most taxa known from Palaeozoic Konservat‐Lagerstätten. With respect to the interrelationships of the three extant clades of Panarthropoda, a sister group relationship between Onychophora and Arthropoda is endorsed by transcriptomics and microRNAs, although this hypothesis forces homoplasy in characters of the segmental ganglia that are shared by tardigrades and arthropods. Cambrian lobopodians, dinocaridids, bivalved arthropods and fuxianhuiids document the successive appearance of characteristic arthropod features in the stem lineage of Euarthropoda (crown‐group arthropods). Molecular dating suggests that arthropods had their origin and initial diversification in the Ediacaran, but no convincing palaeontological evidence for Panarthropoda is available until the earliest Cambrian.

arthropod
interrelationship
phylogeny
major pancrustacean group
debate


From monsoon to marine productivity in the Arabian Sea: insights from glacial and interglacial climates

Abstract: Abstract. The current-climate Indian monsoon is known to boost biological productivity in the Arabian Sea. This paradigm has been extensively used to reconstruct past monsoon variability from palaeo-proxies indicative of changes in surface productivity. Here, we test this paradigm by simulating changes in marine primary productivity for eight contrasted climates from the last glacial–interglacial cycle. We show that there is no straightforward correlation between boreal summer productivity of the Arabian Sea and summer monsoon strength across the different simulated climates. Locally, productivity is fuelled by nutrient supply driven by Ekman dynamics. Upward transport of nutrients is modulated by a combination of alongshore wind stress intensity, which drives coastal upwelling, and by a positive wind stress curl to the west of the jet axis resulting in upward Ekman pumping. To the east of the jet axis there is however a strong downward Ekman pumping due to a negative wind stress curl. Consequently, changes in coastal alongshore stress and/or curl depend on both the jet intensity and position. The jet position is constrained by the Indian summer monsoon pattern, which in turn is influenced by the astronomical parameters and the ice sheet cover. The astronomical parameters are indeed shown to impact wind stress intensity in the Arabian Sea through large-scale changes in the meridional gradient of upper-tropospheric temperature. However, both the astronomical parameters and the ice sheets affect the pattern of wind stress curl through the position of the sea level depression barycentre over the monsoon region (20–150° W, 30° S–60° N). The combined changes in monsoon intensity and pattern lead to some higher glacial productivity during the summer season, in agreement with some palaeo-productivity reconstructions.


Climate change and the selective signature of the Late Ordovician mass extinction

Abstract: Selectivity patterns provide insights into the causes of ancient extinction events. The Late Ordovician mass extinction was related to Gondwanan glaciation; however, it is still unclear whether elevated extinction rates were attributable to record failure, habitat loss, or climatic cooling. We examined Middle Ordovician-Early Silurian North American fossil occurrences within a spatiotemporally explicit stratigraphic framework that allowed us to quantify rock record effects on a per-taxon basis and assay the interplay of macrostratigraphic and macroecological variables in determining extinction risk. Genera that had large proportions of their observed geographic ranges affected by stratigraphic truncation or environmental shifts at the end of the Katian stage were particularly hard hit. The duration of the subsequent sampling gaps had little effect on extinction risk, suggesting that this extinction pulse cannot be entirely attributed to rock record failure; rather, it was caused, in part, by habitat loss. Extinction risk at this time was also strongly influenced by the maximum paleolatitude at which a genus had previously been sampled, a macroecological trait linked to thermal tolerance. A model trained on the relationship between 16 explanatory variables and extinction patterns during the early Katian interval substantially underestimates the extinction of exclusively tropical taxa during the late Katian interval. These results indicate that glacioeustatic sea-level fall and tropical ocean cooling played important roles in the first pulse of the Late Ordovician mass extinction in Laurentia.

late
ordovician mass extinction
thermal tolerance


A 28‐ka history of sea surface temperature, primary productivity and planktonic community variability in the western Arabian Sea

Abstract: [1] Uranium series radionuclides and organic biomarkers, which represent major groups of planktonic organisms, were measured in western Arabian Sea sediments that span the past 28 ka. Variability in the past strength of the southwest and northeast monsoons and its influence on primary productivity, sea surface temperature (SST), and planktonic community structure were investigated. The average alkenone-derived SST for the last glacial period was ∼3°C lower than that measured for the Holocene. Prior to the deglacial, the lowest SSTs coincide with the highest measured fluxes of organic biomarkers, which represent primarily a planktonic suite of diatoms, coccolithophorids, dinoflagellates, and zooplankton. We propose that intensification of winter northeast monsoon winds during the last glacial period resulted in deep convective mixing, cold SSTs and enhanced primary productivity. In contrast, postdeglacial (<17 ka) SSTs are warmer during times in which biomarker fluxes are high. Associated with this transition is a planktonic community structure change, in which the ratio of the average cumulative flux of diatom biomarkers to the cumulative flux of coccolithophorid biomarkers is twice as high during the deglacial and Holocene than the average ratio during the last glacial period. We suggest that this temporal transition represents a shift from a winter northeast monsoon-dominated (pre-17 ka) to a summer southwest monsoon-dominated (post-17 ka) wind system.

sea surface temperature
primary productivity
glacial period


Cranial Growth and Variation in Edmontosaurs (Dinosauria: Hadrosauridae): Implications for Latest Cretaceous Megaherbivore Diversity in North America

Abstract: The well-sampled Late Cretaceous fossil record of North America remains the only high-resolution dataset for evaluating patterns of dinosaur diversity leading up to the terminal Cretaceous extinction event. Hadrosaurine hadrosaurids (Dinosauria: Ornithopoda) closely related to Edmontosaurus are among the most common megaherbivores in latest Campanian and Maastrichtian deposits of western North America. However, interpretations of edmontosaur species richness and biostratigraphy have been in constant flux for almost three decades, although the clade is generally thought to have undergone a radiation in the late Maastrichtian. We address the issue of edmontosaur diversity for the first time using rigorous morphometric analyses of virtually all known complete edmontosaur skulls. Results suggest only two valid species, Edmontosaurus regalis from the late Campanian, and E. annectens from the late Maastrichtian, with previously named taxa, including the controversial Anatotitan copei, erected on hypothesized transitional morphologies associated with ontogenetic size increase and allometric growth. A revision of North American hadrosaurid taxa suggests a decrease in both hadrosaurid diversity and disparity from the early to late Maastrichtian, a pattern likely also present in ceratopsid dinosaurs. A decline in the disparity of dominant megaherbivores in the latest Maastrichtian interval supports the hypothesis that dinosaur diversity decreased immediately preceding the end Cretaceous extinction event.

dinosauria
north america
hadrosaurid
edmontosaurus


First report on the occurrence of Neseuretinus and Ovalocephalus trilobites in the Middle Ordovician of Iran

Abstract: In the Derenjal Mountains of east Central Iran, the upper part of the Shirgesht Formation (uppermost Darriwilian) contains a distinct trilobite assemblage that includes Neseuretinus birmanicus and Ovalocephalus aff. obsoletus among others. Both genera were previously unknown in Iran. The occurrence of Ovalocephalus represents the earliest sign of westward taxon migration from China towards higher latitudes along the West Gondwanan margin, which may be related to global warming, after a short episode of cooler climate in the early to mid Darriwilian. Patterns of biogeographical distribution of Ovalocephalus and Neseuretinus suggest that Central Iran was part of an "overlap zone" where tropical and high latitude benthic taxa mingled.

neseuretinus
ovalocephalus
middle ordovician


A NEW SPECIES OF MOERITHERIUM (PROBOSCIDEA, MAMMALIA) FROM THE EOCENE OF ALGERIA: NEW PERSPECTIVES ON THE ANCESTRAL MORPHOTYPE OF THE GENUS

Abstract: Abstract:  A new species of Moeritherium (Proboscidea, Mammalia), M. chehbeurameuri sp. nov., is described from remains discovered in the early late Eocene locality of Bir El Ater, Algeria. Although mainly represented by isolated teeth, it shows distinct synapomorphies which justify its attribution to the genus Moeritherium, together with exclusive features that led to the creation of the new species. The main characteristic of this new taxon is the almost complete lophodonty shown by its molars, while Moeritherium is commonly regarded as a bunolophodont to bunodont taxon. In addition to this lophodonty, this new taxon shows anatomical features as yet unknown for the genus, but often met within lophodont early proboscidean taxa such as Phosphatherium escuilliei and Numidotherium koholense. Although a revision of the whole genus Moeritherium is outside the scope of this paper, the main controversies and discussion about the definition of species within the genus Moeritherium are discussed. The surprising lophodonty of M. chehbeurameuri, together with its small size, its early late Eocene age and the weak molarization of its P3 support the hypothesis of a lophodont hypothetical ancestor for moeritheres, and therefore strengthen the growing hypothesis of a lophodont dental ancestral morphotype for proboscideans.

moeritherium
mammalia
ancestral morphotype
lophodont


Faunal turnover of marine tetrapods during the Jurassic–Cretaceous transition

Abstract: Marine and terrestrial animals show a mosaic of lineage extinctions and diversifications during the Jurassic–Cretaceous transition. However, despite its potential importance in shaping animal evolution, few palaeontological studies have focussed on this interval and the possible climate and biotic drivers of its faunal turnover. In consequence evolutionary patterns in most groups are poorly understood. We use a new, large morphological dataset to examine patterns of lineage diversity and disparity (variety of form) in the marine tetrapod clade Plesiosauria, and compare these patterns with those of other organisms. Although seven plesiosaurian lineages have been hypothesised as crossing the Jurassic–Cretaceous boundary, our most parsimonious topology suggests the number was only three. The robust recovery of a novel group including most Cretaceous plesiosauroids (Xenopsaria, new clade) is instrumental in this result. Substantial plesiosaurian turnover occurred during the Jurassic–Cretaceous boundary interval, including the loss of substantial pliosaurid, and cryptoclidid diversity and disparity, followed by the radiation of Xenopsaria during the Early Cretaceous. Possible physical drivers of this turnover include climatic fluctuations that influenced oceanic productivity and diversity: Late Jurassic climates were characterised by widespread global monsoonal conditions and increased nutrient flux into the opening Atlantic‐Tethys, resulting in eutrophication and a highly productive, but taxonomically depauperate, plankton. Latest Jurassic and Early Cretaceous climates were more arid, resulting in oligotrophic ocean conditions and high taxonomic diversity of radiolarians, calcareous nannoplankton and possibly ammonoids. However, the observation of discordant extinction patterns in other marine tetrapod groups such as ichthyosaurs and marine crocodylomorphs suggests that clade‐specific factors may have been more important than overarching extrinsic drivers of faunal turnover during the Jurassic–Cretaceous boundary interval.

jurassic–cretaceous transition
plesiosaurian lineage
faunal turnover
polycotylidae
marine reptile


Mammals as a Key to Evolutionary Theory

Abstract: Mammalogy provides exceptionally fertile grounds for advancing evolutionary theory, because its data spans from diverse researches on living forms to a rich fossil record. I illustrate this by integrating interdisciplinary evidence and hypotheses in the habitat theory, including : 1) the context of paleoclimatic changes, and how species' distributions responded to them; 2) geographical biases in turnover rates of species; 3) the turnover-pulse hypothesis; 4) breadth of resource use as a cause of phylogenetic turnover rates. Preliminary tests using the late Neogene records of the Americas and Africa suggest that major aspects of the Great American Interchange have parallels in the African record, as predicted by the habitat theory

evolutionary theory
habitat theory
turnover-pulse hypothesis


Retention of fish-like odontode overgrowth in Permian tetrapod dentition supports outside-in theory of tooth origins

Abstract: Teeth are often thought of as structures that line the margins of the mouth; however, tooth-like structures called odontodes are commonly found on the dermal bones of many Palaeozoic vertebrates including early jawless fishes. ‘Odontode’ is a generalized term for all tooth-like dentine structures that have homologous tissues and development. This definition includes true teeth and the odontodes of early ‘fishes’, which have been recently examined to gain new insights into the still unresolved origin of teeth. Two leading hypotheses are frequently referenced in this debate: the ‘outside-in’ hypothesis, which posits that dermal odontodes evolutionarily migrate into the oral cavity, and the ‘inside-out’ hypothesis, which posits that teeth originated in the oropharyngeal cavity and then moved outwards into the oral cavity. Here, we show that, unlike the well-known one-to-one replacement patterns of marginal dentition, the palatal dentition of the early Permian tetrapods, including the dissorophoid amphibian Cacops and the early reptile Captorhinus, is overgrown by a new layer of bone to which the newest teeth are then attached. This same overgrowth pattern has been well documented in dermal and oral odontodes (i.e. teeth) of early fishes. We propose that this pattern represents the primitive condition for vertebrates and may even predate the origin of jaws. Therefore, this pattern crosses the fish–tetrapod transition, and the retention of this ancestral pattern in the palatal dentition of early terrestrial tetrapods provides strong support for the ‘outside-in’ hypothesis of tooth origins.


The ‘Unicorn’ Dinosaur That Wasn’t: A New Reconstruction of the Crest of Tsintaosaurus and the Early Evolution of the Lambeosaurine Crest and Rostrum

Abstract: The lambeosaurine Tsintaosaurus spinorhinus has traditionally been reconstructed with an elevated, hollow, spike-like crest composed entirely of the nasal bones, although this has been disputed. Here, we provide a new reconstruction of the skull of this species based on reexamination and reinterpretation of the morphology and articular relationships of the type and Paratype skulls and a fragmentary crest. We confirm the presence of a supracranial crest composed of the elevated nasal bones, but also including the premaxillae. We hypothesize that the crest is a tall, lobate, hollow structure that projects dorsally and slightly caudally a distance greater than the height of the skull along the quadrate. In our reconstruction, the nasal passage passes through the crest, but enters the skull rostral to the tubular process of the nasals, not through it. Tsintaosaurus spinorhinus is rediagnosed on the basis of a suite of cranial autapomorphies including a circumnarial fossa subdivided into three accessory fossae, prefrontal with ascending rostral process and lateral flange, nasals fused sagittally to form elongate tubular process that rises dorsally from skull roof, each nasal being expanded rostrocaudally into a rhomboid distal process, and medial processes of premaxillae at the summit of the cranial crest inserted between rhomboid processes of nasals. Tsintaosaurus spinorhinus lacks characters that are present in more derived lambeosaurines (parasaurolophins and lambeosaurins), such as rotation of the caudal margin of the crest to an acute angle with the skull roof, lateral processes of the nasals that enclose part of the intracranial cavity and participate in the formation of the walls of the common median chamber, and a smooth narial fossa lacking ridges and accessory fossae. We hypothesize that ancestrally the rostrum of lambeosaurines may have been more similar to that in Saurolophinae, and became subsequently reduced in complexity during evolution of the group.


Frond orientations with independent current indicators demonstrate the reclining rheotropic mode of life of several Ediacaran rangeomorph taxa

Abstract: Abstract.— Fossils from the deep-sea Ediacaran biotas of Newfoundland are among the oldest architecturally complex soft-bodied macroorganisms on Earth. Most organisms in the Mistaken Point–type biotas of Avalonia—particularly the fractal-branching frondose Rangeomorpha— have been traditionally interpreted as living erect within the water column during life. However, due to the scarcity of documented physical sedimentological proxies associated with fossiliferous beds, Ediacaran paleocurrents have been inferred in some instances from the preferential orientation of fronds. This calls into question the relationship between frond orientation and paleocurrents. In this study, we present an integrated approach from a newly described fossiliferous surface (the “Melrose Surface” in the Fermeuse Formation at Melrose, on the southern portion of the Catalina Dome in the Discovery UNESCO Global Geopark) combining: (1) physical sedimentological evidence for paleocurrent direction in the form of climbing ripple cross-lamination and (2) a series of statistical analyses based on modified polythetic and monothetic clustering techniques reflecting the circular nature of the recorded orientation of Fractofusus misrai specimens. This study demonstrates the reclining rheotropic mode of life of the Ediacaran rangeomorph taxon Fractofusus misrai and presents preliminary inferences suggesting a similar mode of life for Bradgatia sp. and Pectinifrons abyssalis based on qualitative evidence. These results advocate for the consideration of an alternative conceptual hypothesis for position of life of Ediacaran organisms in which they are interpreted as having lived reclined on the seafloor, in the position that they are preserved.

reclining rheotropic mode
water column
frond orientation


Reconstruction of breastfeeding and weaning practices using stable isotope and trace element analyses: A review.

Abstract: Biogeochemical methods using stable isotopes and trace elements have been increasingly developed and applied to reconstruct modern and ancient breastfeeding and weaning practices of mammals, including humans, because they offer direct proxies for the dietary intake of subadults. Carbon, nitrogen, and oxygen isotopes have been used to evaluate breast milk lipid, protein, and water intake, respectively. Carbon and sulfur isotopes have been used to estimate the content of weaning foods. The elemental concentrations of Sr and Ba in subadult tissues differ because of the dietary change during the weaning process. For analyses, various tissues have been used, such as hair, nail, blood, and feces for modern mammals and bone and teeth for ancient ones. Of these, trace element analysis of tooth enamel offers a good opportunity for the reconstruction of breastfeeding and weaning practices of the more distant past at finer resolution, although further understanding of the metabolism of trace elements is necessary. There are various tissue- and element-specific advantages and disadvantages, and a combination of different proxies can illuminate practices from various viewpoints. Finally, applying the geochemical reconstruction of breastfeeding and weaning practices to human ecology, primatology, and paleoanthropology is important; basic studies of the underlying physiological mechanisms and technical improvements in the analyses will further highlight avenues for future research.

weaning practice
stable isotope
tissue
element analysis


Northern Eurasian rhinocerotines (Mammalia, Perissodactyla) by the Pliocene–Pleistocene transition: phylogeny and historical biogeography

Abstract: Pliocene and earliest Pleistocene Northern Eurasian rhinocerotines are poorly documented and understudied in comparison to Pleistocene and Miocene ones. However, they represent a key-group of species for understanding the phylogeny and historical biogeography of their Pleistocene relatives. In the present paper, we revise the abundant material from the late Pliocene locality of Kvabebi, Georgia from a systematic, phylogenetic and palaeobiogeographical perspective. The specimens from Kvabebi are documented by two partially preserved skulls, one mandible and several postcranial remains. Morphological and morphometric comparison with the type material assigned to Pliocene and earliest Pleistocene Northern Eurasian Rhinocerotinae reveal that the specimens from Kvabebi have close affinities with the poorly known Dicerorhinus miguelcrusafonti Guérin & Santafé-Llopis, 1978, described from Layna (Spain). The latter species, represented by scanty remains from the Iberian Peninsula, is usually excluded from morphological and morphometrical comparisons and no findings were reported after the 1990s. Pliocene rhinocerotine species have monotonous dental and postcranial morphologies and only a few features allow us to discern the different species. The material from Layna and Kvabebi is somewhat smaller than that of other Pliocene taxa, except for the largest representatives of Stephanorhinus etruscus (Falconer, 1868). Accordingly, the earliest specimens assigned to S. etruscus on morphometric grounds should be revised in the light of the new data here presented. A cladistic analysis performed on 280 characters and 30 species suggests that the emblematic early Pliocene European species, ‘Dihoplus’ megarhinus (de Christol, 1834), is sister taxon to the Layna and Kvabebi rhinoceroses. Accordingly, both species are here assigned to a new genus named Pliorhinus gen. nov. Although distinct, this clade has close affinities with the paraphyletic genus Stephanorhinus, therefore suggesting the co-occurrence of at least two distinct rhinocerotine lineages raised in the late Miocene interval in Northern Eurasia. http://zoobank.org/urn:lsid:zoobank.org:pub:D3A1AB6D-EFE7-4813-AE74-A27D204A18CA

phylogeny
historical biogeography
stephanorhinus etruscus
northern eurasian rhinocerotine