Decaying in different clays: implications for soft‐tissue preservation

Abstract: Lagerstätten, places where soft‐bodied organisms became mineralized, provide a substantial bulk of palaeobiological information, but the detailed mechanisms of how soft‐tissue preservation takes place remain debatable. An experimental taphonomy approach, which allows for direct study of decay and mineralization, offers a means to study the preservational potential of different soft‐bodied organisms under controlled conditions. Here we compare the preservational capacity of two types of clay (kaolinite and montmorillonite) through a long‐term (24 month) experiment involving the burial and decay of small crustaceans. Our experimental design is innovative in that it models catastrophic sedimentation in fine‐grained colloidal suspension, which is believed to form Lagerstätten deposits. We demonstrated better preservation of buried organisms in clays compared to water, and in kaolinite compared to montmorillonite. As aluminium cations were present in high concentrations in kaolinite sediment but not in montmorillonite, the better preservation in kaolinite is attributed to the tanning properties of aluminium, which catalyses cross‐linking in proteins, protecting them from bacterial degradation. Anaerobic environments and acidification also slow down decay, but they are less effective than tanning. Kaolinite and montmorillonite replaced the crustacean integuments differently: in the remains buried in kaolinite, Al and Si were detected in equal proportions, while in those buried in montmorillonite, the Si content appeared to be much higher even in comparison with the initial sample of the clay. These variations probably arose from the different dynamics of acidic hydrolysis in the two clays associated with anaerobic decomposition of organic matter. Our results show that the preservation mechanism includes multi‐component interactions between the solution, mineral, sediment and organic remains; taken separately, any single component explains little. The specific conditions that occur within the colloidal clay sediments can facilitate conservation and start fast mineralization according to chemical properties and elemental content.

clay
soft‐tissue preservation
mineralization
decay


New evidence from China for the nature of the pterosaur evolutionary transition

Abstract: Pterosaurs are extinct flying reptiles, the first vertebrates to achieve powered flight. Our understanding of the evolutionary transition between basal, predominantly long-tailed forms to derived short-tailed pterodactyloids remained poor until the discovery of Wukongopterus and Darwinopterus in western Liaoning, China. In this paper we report on a new genus and species, Douzhanopterus zhengi, that has a reduced tail, 173% the length of the humerus, and a reduced fifth pedal digit, whose first phalange is ca. 20% the length of metatarsal III, both unique characters to Monofenestra. The morphological comparisons and phylogenetic analysis presented in this paper demonstrate that Douzhanopterus is the sister group to the ‘Painten pro-pterodactyloid’ and the Pterodactyloidea, reducing the evolutionary gap between long- and short-tailed pterosaurs.

china
pterosaur
evolutionary transition
douzhanopterus zhengi


Estimating the timing of early eukaryotic diversification with multigene molecular clocks

Abstract: Although macroscopic plants, animals, and fungi are the most familiar eukaryotes, the bulk of eukaryotic diversity is microbial. Elucidating the timing of diversification among the more than 70 lineages is key to understanding the evolution of eukaryotes. Here, we use taxon-rich multigene data combined with diverse fossils and a relaxed molecular clock framework to estimate the timing of the last common ancestor of extant eukaryotes and the divergence of major clades. Overall, these analyses suggest that the last common ancestor lived between 1866 and 1679 Ma, consistent with the earliest microfossils interpreted with confidence as eukaryotic. During this interval, the Earth's surface differed markedly from today; for example, the oceans were incompletely ventilated, with ferruginous and, after about 1800 Ma, sulfidic water masses commonly lying beneath moderately oxygenated surface waters. Our time estimates also indicate that the major clades of eukaryotes diverged before 1000 Ma, with most or all probably diverging before 1200 Ma. Fossils, however, suggest that diversity within major extant clades expanded later, beginning about 800 Ma, when the oceans began their transition to a more modern chemical state. In combination, paleontological and molecular approaches indicate that long stems preceded diversification in the major eukaryotic lineages.


Dinosaur teeth from the Jurassic Qigu and Shishugou Formations of the Junggar Basin (Xinjiang/China) and their paleoecologic implications

Abstract: The Middle and early Late Jurassic Qigu and Shishugou Formations of the southern and central Junggar Basin yielded teeth of theropods (Theropoda indet.), sauropods (Eusauropoda indet.), and stegosaurs. The dinosaur assemblage of the southern Junggar Basin is less diverse and is represented by smaller forms than in the central part of the basin. The microwear of the teeth of Eusauropoda indet. resembles that observed in Camarasaurus and may have formed as a result of biting through resistant woody materials. Carbon and oxygen isotope data of the sauropod and theropod teeth indicate feeding within a C 3 -plant ecosystem in a continental setting. Differences in enamel d 13 C and d 18 O values between Eusauropoda indet. and the theropod teeth are comparable to those observed in other herbivorous and carnivorous vertebrates, and suggest at least partial preservation of original dietary signals.

jurassic qigu
shishugou formations
junggar basin
sauropod


Diversification events and the effects of mass extinctions on Crocodyliformes evolutionary history

Abstract: The rich fossil record of Crocodyliformes shows a much greater diversity in the past than today in terms of morphological disparity and occupation of niches. We conducted topology-based analyses seeking diversification shifts along the evolutionary history of the group. Our results support previous studies, indicating an initial radiation of the group following the Triassic/Jurassic mass extinction, here assumed to be related to the diversification of terrestrial protosuchians, marine thalattosuchians and semi-aquatic lineages within Neosuchia. During the Cretaceous, notosuchians embodied a second diversification event in terrestrial habitats and eusuchian lineages started diversifying before the end of the Mesozoic. Our results also support previous arguments for a minor impact of the Cretaceous/Palaeogene mass extinction on the evolutionary history of the group. This argument is not only based on the information from the fossil record, which shows basal groups surviving the mass extinction and the decline of other Mesozoic lineages before the event, but also by the diversification event encompassing only the alligatoroids in the earliest period after the extinction. Our results also indicate that, instead of a continuous process through time, Crocodyliformes diversification was patchy, with events restricted to specific subgroups in particular environments and time intervals.

crocodyliformes
neosuchia
mesozoic
diversification event


Decoding the Evolution of Melanin in Vertebrates.

Abstract:
Melanins are widespread pigments in vertebrates, with important roles in visual signaling, UV protection, and homeostasis. Fossil evidence of melanin and melanin-bearing organellesmelanosomesin ancient vertebrates may illuminate the evolution of melanin and its functions, but macroevolutionary trends are poorly resolved. Here, we integrate fossil data with current understanding of melanin function, biochemistry, and genetics. Mapping key genes onto phenotypic attributes of fossil vertebrates identifies potential genomic controls on melanin evolution. Taxonomic trends in the anatomical location, geometry, and chemistry of vertebrate melanosomes are linked to the evolution of endothermy. These shifts in melanin biology suggest fundamental links between melanization and vertebrate ecology. Tissue-specific and taxonomic trends in melanin chemistry support evidence for evolutionary tradeoffs between function and cytotoxicity.
Melanins (see Glossary) are dark to rufous pigments that are widespread in vertebrates and underpin critical functions in physiology and behavior [1] . Fossil evidence of melanin extending to over 300 million years ago has triggered a paradigm shift in paleobiology, prompting remarkable reconstructions of the coloration and behavior of extinct vertebrates [2] [3] [4] [5] [6] . New discoveries of internal melanins in vertebrate fossils have broadened our understanding of the functional diversity of ancient melanins [7-9] and invite a re-evaluation of the macroevolutionary history of melanin and its functions. Here, we synthesize trends in the fossil record of melanin and explore fossil evidence for the evolution of melanin function and the genetic basis of melanization. This highlights the value of the fossil record as a resource for tracking melanin evolution through deep time.
In extant vertebrates, melanin occurs as micron-sized organelles, melanosomes, in the integument, eyes and internal tissues and functions in photoprotection, visual signaling, thermoregulation, immunity, antioxidation, mechanical strengthening, and abrasion resistance [6, 10, 11] (Figure 1 , Box 1). It is unclear which functions evolved first and which selection pressures dominate [11, 12] . Fossils preserving evidence of melanin offer a unique temporal perspective.
Melanin has been reported from fossil vertebrates from >25 localities from the Carboniferous to the Pliocene (Table S1 in the supplemental information online). The fossils include cyclostomes, fish, frogs, lizards, and other squamates, ichthyosaurs, plesiosaurs, turtles, pterosaurs, feathered and nonfeathered dinosaurs, birds, and mammals. This phylogenetically and temporally broad dataset yields evidence for ancient functions of melanin (Figure 2 ).
In extant vertebrates melanin fulfils diverse roles including visual communication, photoprotection, antioxidation, and mechanical strengthening of tissues, but the evolution of these functions is debated.
The discovery that melanosomes in fossil and modern vertebrates are associated with tissue-specific suites of trace metals supports hypotheses that melanin has ancient functions in metal homeostasis and antioxidant regulation.
Shifts in melanosome biology across the dinosaur-bird transition reveal intimate links between adaptive and pleiotropic processes relating to the evolution of endothermy, metal homeostasis, photoprotection and the lymphatic system. Key genes can be mapped onto color pattern phenotypes in fossil vertebrates.
Melanin-based coloration in vertebrates is dominated by melanin forms associated with low cytotoxicity, possibly reflecting selective adaptation against forms linked with greater oxidative stress or co-option of melanin forms with specific metal binding behavior for coloration.


An acid-free method of microfossil extraction from clay-rich lithologies using the surfactant Rewoquat

Abstract: Marine rocks characterized by high clay content provide excellent conditions for fossil preservation, particularly for organic-walled microfossils such as retiolitid graptolites and chitinozoans. Nevertheless, the phyllosilicate minerals, which constitute the clay component, make microfossil extractions difficult. The problem results from the tendency of phyllosilicates to form aggregates in low pH values, as standard methods of microfossil extraction employ acids for rock digestion. Consequently, the use of acids for clay-rich rocks is often inefficient and time-consuming. We propose a method of rock disintegration using the surfactant Rewoquat and compare it with two commonly applied approaches: digestion in buffered acetic acid and in HCl-HF. Using examples from the Mulde Brick-clay Member from the Silurian of Gotland and the Daleje Shale from the Devonian of the Prague Basin we observed that disintegration in Rewoquat was faster (days) than digestion in acid (months), and allowed to recover calcareous in addition to organic-walled fossils. The yield and preservation was comparably good, except for conodonts, which were strongly etched after using HCl-HF. Retiolitid graptolites recovered using Rewoquat were preserved in 3D and showed a lower degree of fragmentation. The fossil content of the residue obtained using Rewoquat was higher due to dispersion of clay aggregates. For observation of delicate fossils we recommend to coat the sample with the surfactant. Application of Rewoquat can reveal the most delicate forms and growth stages, and thus provide a better insight into the ontogeny, autecology, and body size distributions of a number of fossil groups.

microfossil extraction
surfactant rewoquat
disintegration


Understanding long‐term carbon cycle trends: The late Paleocene through the early Eocene

Abstract: (1) The late Paleocene to the early Eocene (� 58-52 Ma) was marked by significant changes in global climate and carbon cycling. The evidence for these changes includes stable isotope records that reveal prominent decreases in i 18 Oa ndi 13 C, suggesting a rise in Earth's surface temperature (� 4 i C) and a drop in net carbon output from the ocean and atmosphere. Concurrently, deep-sea carbonate records at several sites indicate a deepening of the calcite compensation depth (CCD). Here we investigate possible causes (e.g., increased volcanic degassing or decreased net organic burial) for these observations, but from a new perspective. The basic model employed is a modified version of GEOCARB III. However, we have coupled this well-known geochemical model to LOSCAR (Long-term Ocean-atmosphere Sediment CArbon cycle Reservoir model), which enables simulation of seawater carbonate chemistry, the CCD, and ocean i 13 C. We have also added a capacitor, in this case represented by gas hydrates, that can store and release 13 C-depleted carbon to and from the shallow geosphere over millions of years. We further consider accurate input data (e.g., i 13 C of carbonate) on a currently accepted timescale that spans an interval much longer than the perturbation. Several different scenarios are investigated with the goal of consistency amongst inferred changes in temperature, the CCD, and surface ocean and deep ocean i 13 C. The results strongly suggest that a decrease in net organic carbon burial drove carbon cycle changes during the late Paleocene and early Eocene, although an increase in volcanic activity might have contributed. Importantly, a drop in net organic carbon burial may represent increased oxidation of previously deposited organic carbon, such as stored in peat or gas hydrates. The model successfully recreates trends in Earth surface warming, as inferred from i 18 O records, the CCD, and i 13 C. At the moment, however, our coupled modeling effort cannot reproduce the magnitude of change in all these records collectively. Similar problems have arisen in simulations of short-term hyperthermal events during the early Paleogene (Paleocene-Eocene Thermal Maximum), suggesting one or more basic issues with data interpretation or geochemical modeling remain.


What nymphal morphology can tell us about parental investment -a group of cockroach hatchlings in Baltic amber documented by a multi-method approach

Abstract:
We describe a piece of Baltic amber, about 50 million years old, which contains a group of 13 small cockroach nymphs. These specimens were documented with different methods to explore the advantages and limitations of certain imaging techniques:
(1) light-based methods, such as stereo-macro photography, composite imaging under polarised light, combined with virtual surface reconstruction, and (2) X-ray micro-computed tomography, processed as volume renderings and surface reconstructions. All nymphs within the amber piece are of the same size and do not exhibit any noticeable morphological variance. Their developmental state and the way in which they are arranged indicate that these nymphs represent hatchlings. Dictyopterans (including Mantodea and Blattodea with Isoptera as ingroup) exhibit a wide range of different types of social and brood care behaviour. The evolution of this complex set of characters has been addressed repeatedly in extant-based approaches, yet deep-time aspects of this evolutionary process have rarely been addressed. The specimens described here could represent a case of a group of blattodean nymphs hatching from an ootheca, which would represent the first fossil record of such a process, or even possibly provide the first indirect evidence of social behaviour in fossil non-termite dictyopterans, indicating that it was already developed 50 million years ago.


Evolutionary development of the cephalopod arm armature: a review

Abstract: The cephalopod arm armature is certainly one of the most important morphological innovations responsible for the evolutionary success of the Cephalopoda. New palaeontological discoveries in the recent past afford to review and reassess origin and homology of suckers, sucker rings, hooks, and cirri. Since a priori character state reconstructions are still ambiguous, we suggest and discuss three different evolutionary scenarios. Each of them is based on the following assumptions: (1) Neocoleoidea uniting extant Decabrachia and Octobrachia is monophyletic (= proostracumbearing coleoids); (2) extinct Belemnitida and Diplobelida are stem decabrachians; (3) proostracum-less coleoids (Hematitida, Donovaniconida, Aulacoceratida) represent stem-neocoleoids; (4) Ammonoidea and Bactritoidea are stem coleoids. We consider a scenario where belemnoid hooks derived from primitive suckers as well-supported. Regarding belemnoid hooks and suckers as homologues implies that belemnoid, oegopsid, and probably ammonoid arm hooks arose through parallel evolution. Our conclusions challenge the widespread opinion, whereupon belemnoid hooks evolved de novo, and instead support earlier ideas formulated by Sigurd von Boletzky.


A New Centrosaurine from the Late Cretaceous of Alberta, Canada, and the Evolution of Parietal Ornamentation in Horned Dinosaurs

Abstract: In 1916, a centrosaurine dinosaur bonebed was excavated within the Campanian-aged deposits of what is now Dinosaur Provincial Park, Alberta, Canada. Specimens from this now-lost quarry, including two parietals, a squamosal, a skull missing the frill, and an incomplete dentary, were purchased by The Natural History Museum, London. The material was recently reprepared and identified herein as a previously unknown taxon, Spinops sternbergorum gen. et sp. nov. Based upon the available locality data and paleopalynology, the quarry lies in either the upper part of the Oldman Formation or the lower part of the Dinosaur Park Formation. The facial region of the partial skull is similar to putative mature specimens of Centrosaurus spp. and Styracosaurus albertensis, with short, rounded postorbital horncores and a large, erect nasal horncore. Parietal ornamentation is consistent on both known parietals and is unique among ceratopsids. Bilateral, procurved parietal hooks occupy the P1 (medial-most) position on the dorsal surface of the parietal and are very similar to those seen in Centrosaurus apertus. Epiparietals in the P2 or possibly P3 position (lateral to P1) manifest as extremely elongate, caudally directed spikes, unlike the condition in C. apertus, S. albertensis, or any other “derived” centrosaurine. Cladistic analysis suggests that S. sternbergorum is closely related to Centrosaurus and Styracosaurus. Historically, based upon the condition in Styracosaurus and related centrosaurines, it was assumed that the medial-most elongated spikes on centrosaurine parietals correspond to the P3 epiparietal position. The exception illustrated in the new taxon suggests that homologies of epiparietals among basal centrosaurines (e.g., Albertaceratops and Diabloceratops) and derived centrosaurines (e.g., Styracosaurus and “pachyrhinosaurs”) should be reconsidered. The medially-placed, caudally-directed “P3” process of basal centrosaurines may, in fact, be homologous with P2.

centrosaurine
alberta
parietal ornamentation


The diverse dietary profiles of MIS 3 cave bears from the Romanian Carpathians: insights from stable isotope (δ13C and δ15N) analysis

Abstract: Late Pleistocene European cave bears (Ursus spelaeus) have been considered to be largely vegetarian, although stable isotope data (δ13C and δ15N values) from the Romanian Carpathians has suggested considerable dietary variation. Here we evaluate previous and additional adult cave bear isotopic data from four Marine Isotope Stage 3 (MIS 3) sites in the Carpathians. Peştera Urşilor (N = 35), Peştera Cioclovina (N = 32), Peştera Muierilor (N = 8), and Peştera cu Oase (N = 72) provide both a dichotomy between samples suggesting vegetarian diets (from Cioclovina and Muierilor) and more omnivorous diets (from Urşilor and Oase), and considerable isotopic variation within samples from each site. While an inference of a strictly vegetarian diet may apply to groups that lived in ecosystems which restricted the available animal protein for these large ursids, the within and between sample isotopic variation among the Carpathian cave bears indicates considerable flexibility in their sources of protein and hence in their dietary regimes. In addition, developmental assessment of Cioclovina isotopic profiles (neonates, juveniles, sub‐adults and adults) provides patterns of transfer of stable isotope signatures throughout immature life for both δ13C and δ15N (increase and decrease, respectively), whereas those from Urşilor show little developmental shift.


Foraminiferal Mg/Ca increase in the Caribbean during the Pliocene: Western Atlantic Warm Pool formation, salinity influence, or diagenetic overprint?

Abstract: We constructed a high‐resolution Mg/Ca record on the planktonic foraminifer Globigerinoides sacculifer in order to explore the change in sea surface temperature (SST) due to the shoaling of the Isthmus of Panama as well as the impact of secondary factors like diagenesis and large salinity fluctuations. The study covers the latest Miocene and the early Pliocene (5.6–3.9 Ma) and was combined with δ18O to isolate changes in sea surface salinity (SSS). Before 4.5 Ma, SSTMg/Ca and SSS show moderate fluctuations, indicating a free exchange of surface ocean water masses between the Pacific and the Atlantic. The increase in δ18O after 4.5 Ma represents increasing salinities in the Caribbean due to the progressive closure of the Panamanian Gateway. The increase in Mg/Ca toward values of maximum 7 mmol/mol suggests that secondary influences have played a significant role. Evidence of crystalline overgrowths on the foraminiferal tests in correlation with aragonite, Sr/Ca, and productivity cyclicities indicates a diagenetic overprint on the foraminiferal tests. Laser ablation inductively coupled plasma–mass spectrometry analyses, however, do not show significantly increased Mg/Ca ratios in the crystalline overgrowths, and neither do calculations based on pore water data conclusively result in significantly elevated Mg/Ca ratios in the crystalline overgrowths. Alternatively, the elevated Mg/Ca ratios might have been caused by salinity as the δ18O record of Site 1000 has been interpreted to represent large fluctuations in SSS, and cultivating experiments have shown an increase in Mg/Ca with increasing salinity. We conclude that the Mg/Ca record <4.5 Ma can only reliably be considered for paleoceanographical purposes when the minimum values, not showing any evidence of secondary influences, are used, resulting in a warming of central Caribbean surface water masses after 4.5 Ma of ∼2°C.

caribbean
diagenetic overprint
late miocene


A lower jaw of Palaeoxonodon from the Middle Jurassic of the Isle of Skye, Scotland, sheds new light on the diversity of British stem therians

Abstract: The Middle Jurassic was a key interval of mammalian evolutionary history that witnessed the diversification of the therian stem group. Great Britain has yielded a significant record of mammalian fossils from this interval, represented by numerous isolated jaws and teeth from the Bathonian of Oxfordshire and the Isle of Skye. This record captures a key period in early cladotherian evolution, with amphitheriids, peramurans and ‘stem zatherians’ displaying intermediate talonid morphologies that document the evolutionary assembly of tribosphenic molars. We present a mandible with near‐complete dentition from the late Bathonian (c. 167.4–166.5 Ma) Kilmaluag Formation, near Elgol, Skye, representing the amphitheriid Palaeoxonodon ooliticus, previously known only from isolated teeth. The specimen sheds new light on the taxonomic diversity of British Middle Jurassic stem therians, as the morphological variation within the preserved tooth row encompasses that previously ascribed to three distinct species within two genera: Palaeoxonodon ooliticus, P. freemani and Kennetheridium leesi. Thus, both P. freemani and K. leesi are subjective junior synonyms of P. ooliticus. The dental formula of P. ooliticus (i4:c1:p5:m5) is intermediate between the primitively larger postcanine count (p5:m6–7) of Amphitherium and the reduced number in peramurans and tribosphenidans (p5:m3). Phylogenetic analyses of P. ooliticus generally confirm a close affinity with Amphitherium, but highlight the lack of strong empirical support for hypothesized patterns of divergences among early cladotherians.

jaw
palaeoxonodon
early cladotherian
phylogenetic analysis


Drowning of the −150 m reef off Hawaii: A casualty of global meltwater pulse 1A?

Abstract: We present evidence that the drowning of the 2150 m coral reef around Hawaii was caused by rapid sea-level rise associated with meltwater pulse 1A (MWP-1A) during the last deglaciation. New U/Th and 14 C accelerator mass spectrometry dates, combined with reinterpretation of existing radiometric dates, constrain the age of the coral reef to 15.2-14.7 ka (U/Th age), indicating that reef growth persisted for 4.3 k.y. following the end of the Last Glacial Maximum at 19 ka. The drowning age of the reef is roughly synchronous with the onset of MWP-1A between 14.7 and 14.2 ka. Dates from coralline algal material range from 14 to 10 cal ka (calibrated radiocarbon age), 1-4 k.y. younger than the coral ages. A paleoenvironmental reconstruction incorporating all available radiometric dates, high-resolution bathymetry, dive observations, and coralgal paleobathymetry data indicates a dramatic rise in sea level around Hawaii ca. 14.7 ka. Paleowater depths over the reef crest increased rapidly above a critical depth (30-40 m), drowning the shallow reef-building Porites corals and causing a shift to deep- water coralline algal growth, preserved as a crust on the drowned reef crest.


Saws, Scissors, and Sharks: Late Paleozoic Experimentation with Symphyseal Dentition

Abstract: Sharks of Late Paleozoic oceans evolved unique dentitions for catching and eating soft bodied prey. A diverse but poorly preserved clade, edestoids are noted for developing biting teeth at the midline of their jaws. Helicoprion has a continuously growing root to accommodate >100 crowns that spiraled on top of one another to form a symphyseal whorl supported and laterally braced within the lower jaw. Reconstruction of jaw mechanics shows that individual serrated crowns grasped, sliced, and pulled prey items into the esophagus. A new description and interpretation of Edestus provides insight into the anatomy and functional morphology of another specialized edestoid. Edestus has opposing curved blades of teeth that are segmented and shed with growth of the animal. Set on a long jaw the lower blade closes with a posterior motion, effectively slicing prey across multiple opposing serrated crowns. Further examples of symphyseal whorls among Edestoidae are provided from previously undescribed North American examples of Toxoprion, Campyloprion, Agassizodus, and Sinohelicoprion. The symphyseal dentition in edestoids is associated with a rigid jaw suspension and may have arisen in response to an increase in pelagic cephalopod prey during the Late Paleozoic. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:363–376, 2020. © 2018 American Association for Anatomy


Splendid Innovation: The Extinct South American Native Ungulates

Abstract: A remarkable diversity of plant-eating mammals known as South American native ungulates (SANUs) flourished in South America for most of the Cenozoic. Although some of these species likely filled ecological niches similar to those of modern hoofed mammals, others differed substantially from extant artiodactyls and perissodactyls in their skull and limb anatomy and probably also in their ecology. Notoungulates and litopterns were the longest-lived and most diverse SANU clades and survived into the Quaternary; astrapotheres went extinct in the late Miocene, whereas other SANU groups were restricted to the Paleogene. Neogene notoungulates were quite specialized in craniodental structure, but many were rather unspecialized postcranially; in contrast, litopterns evolved limb specializations early in their history while maintaining more conservative dentitions. In this article, we review the current understanding of SANU evolutionary relationships and paleoecology, provide an updated compilation of genus temporal ranges, and discuss possible directions for future research. ▪  South American native ungulates (SANUs) were a diverse, long-lived, and independent radiation of mammals into varied terrestrial plant-eater niches. ▪  We review origins, evolution, and paleoecology of the major SANU clades: Notoungulata, Litopterna, Astrapotheria, Xenungulata, and Pyrotheria. ▪  At their peak, during the Eocene and Oligocene, more than 40 genera of native ungulates inhabited South America at any one time. ▪  SANUs ranged from <1 kg to several tons and evolved many combinations of diet and locomotor adaptations not seen in living ungulates.

sanu
notoungulate
litopterna
late pleistocene
evolutionary history


Diversity changes in lycopsid and aquatic fern megaspores through geologic time

Abstract: Quantitative data on lycopsid and aquatic fern megaspore taxa recovered from Carboniferous, Mesozoic, and Tertiary strata have been compiled in order to analyze the changes in diversity of the two groups of fossil plants that produced them. Numbers of species of lycopsid megaspores are similar in the Carboniferous and Mesozoic, whereas the diversity of megafossils is much lower in post-Paleozoic deposits. Our data suggest that lycopsids were more diverse in the Mesozoic than previously thought and that there is a preservational bias against the megafossils, because the plants were probably mainly herbaceous. Heterosporous aquatic ferns first appeared in the Neocomian and gradually diversified until the early Late Cretaceous, after which their numbers remained relatively stable, whereas the variety of lycopsids declined dramatically during the Late Cretaceous. These changes occurred at a time of rapid angiosperm diversification. The reduced diversity of the lycopsids may have been caused by the invasion of their aquatic and damp forest-floor habitats by heterosporous ferns and by aquatic and herbaceous angiosperms. These diversity changes do not seem to be directly related to the global events at the Cretaceous-Tertiary boundary, but the relatively few samples available and the resulting range truncation would make detection of such correlations difficult.


Large-scale phylogeny of chameleons suggests African origins and Eocene diversification

Abstract: Oceanic dispersal has emerged as an important factor contributing to biogeographic patterns in numerous taxa. Chameleons are a clear example of this, as they are primarily found in Africa and Madagascar, but the age of the family is post-Gondwanan break-up. A Malagasy origin for the family has been suggested, yet this hypothesis has not been tested using modern biogeographic methods with a dated phylogeny. To examine competing hypotheses of African and Malagasy origins, we generated a dated phylogeny using between six and 13 genetic markers, for up to 174 taxa representing greater than 90 per cent of all named species. Using three different ancestral-state reconstruction methods (Bayesian and likelihood approaches), we show that the family most probably originated in Africa, with two separate oceanic dispersals to Madagascar during the Palaeocene and the Oligocene, when prevailing oceanic currents would have favoured eastward dispersal. Diversification of genus-level clades took place in the Eocene, and species-level diversification occurred primarily in the Oligocene. Plio-Pleistocene speciation is rare, resulting in a phylogeny dominated by palaeo-endemic species. We suggest that contraction and fragmentation of the Pan-African forest coupled to an increase in open habitats (savannah, grassland, heathland), since the Oligocene played a key role in diversification of this group through vicariance.

phylogeny
chameleon
madagascar
palaeocene
eastward dispersal


Widespread mermithid nematode parasitism of Cretaceous insects

Abstract: Mermithid nematodes are obligate invertebrate parasites dating back to the Early Cretaceous. Their fossil record is sparse, especially before the Cenozoic, thus little is known about their early host associations. This study reports 16 new mermithids associated with their insect hosts from mid-Cretaceous Kachin amber, 12 of which include previously unknown hosts. These fossils indicate that mermithid parasitism of invertebrates was already widespread and played an important role in the mid-Cretaceous terrestrial ecosystem. Remarkably, three hosts (bristletails, barklice and perforissid planthoppers) were previously unknown to be parasitized by mermithids both past and present. Furthermore, our statistical analyses show that in contrast to their Cenozoic counterparts, Cretaceous nematodes including mermithids are more abundant in heterometabolous insect hosts. This result suggests that nematodes have not completely exploited the dominant Holometabola as their hosts until the Cenozoic. This study reveals what appears to be a vanished history of nematodes that parasitized Cretaceous insects.

mermithid
nematode
cretaceous insect
invertebrate


First Evidence of Reproductive Adaptation to “Island Effect” of a Dwarf Cretaceous Romanian Titanosaur, with Embryonic Integument In Ovo

Abstract: Background The Cretaceous vertebrate assemblages of Romania are famous for geographically endemic dwarfed dinosaur taxa. We report the first complete egg clutches of a dwarf lithostrotian titanosaur, from Toteşti, Romania, and its reproductive adaptation to the “island effect”. Methodology/Findings The egg clutches were discovered in sequential sedimentary layers of the Maastrichtian Sânpetru Formation, Toteşti. The occurrence of 11 homogenous clutches in successive strata suggests philopatry by the same dinosaur species, which laid clutches averaging four ∼12 cm diameters eggs. The eggs and eggshells display numerous characters shared with the positively identified material from egg-bearing level 4 of the Auca Mahuevo (Patagonia, Argentina) nemegtosaurid lithostrotian nesting site. Microscopic embryonic integument with bacterial evidences was recovered in one egg. The millimeter-size embryonic integument displays micron size dermal papillae implying an early embryological stage at the time of death, likely corresponding to early organogenesis before the skeleton formation. Conclusions/Significance The shared oological characters between the Haţeg specimens and their mainland relatives suggest a highly conservative reproductive template, while the nest decrease in egg numbers per clutch may reflect an adaptive trait to a smaller body size due to the “island effect”. The combined presence of the lithostrotian egg and its embryo in the Early Cretaceous Gobi coupled with the oological similarities between the Haţeg and Auca Mahuevo oological material evidence that several titanosaur species migrated from Gondwana through the Haţeg Island before or during the Aptian/Albian. It also suggests that this island might have had episodic land bridges with the rest of the European archipelago and Asia deep into the Cretaceous.

reproductive adaptation
island effect
embryonic integument
egg
auca mahuevo


Two new horned dinosaurs from the upper Cretaceous Two Medicine Formation of Montana; with a phylogenetic analysis of the Centrosaurinae (Ornithischia: Ceratopsidae)

Abstract: ABSTRACT Two new ceratopsid dinosaurs, Einiosaurus procurvicornis and Achelousaurus horneri, are described from the Two Medicine Formation (Upper Cretaceous) of Montana. E. procurvicornis is known from three skulls and numerous cranial and postcranial elements from two bonebed assemblages. A. horneri is based on three skulls, one with associated postcranial elements. A phylogenetic review of the subfamily Centrosaurinae reveals two clades, one containing Centrosaurus and Styracosaurus and the other Pachyrhinosaurus plus the two new taxa from Montana. Diagnostic traits for resolving within-group relationships are found only in the skull roof in association with what appear to be secondary sexual characters, probably the result of sexual selection. In addition to illuminating the pattern of ceratopsian evolution, these taxa suggest an increased rate of evolution that may correlate with the late Campanian transgression of the Bearpaw Sea.

montana
centrosaurinae
achelousaurus horneri


Hypoanthraconaia: a new genus of non-marine bivalve molluscs from the Early Permian of Far East Russia

Abstract: Early Permian continental deposits include a large number of localities containing anthracosiid-like nonmarine bivalves traditionally assigned to Anthraconaia Trueman and Weir, 1946, Palaeanodonta Amalitzky, 1895, and Palaeomutela Amalitzky, 1892. In most cases, these classifications are only tentative due to insufficient preservation in which the shells are missing their main characteristics: the ligament and the hinge. Non-marine bivalves from the Early Permian Upper Pospelovka Subformation of Russky Island (South Primorye, Far East Russia), described here as Hypoanthraconaia gen. nov., differ morphologically from the above genera by a set of external features including the initial shell, the mode of intersection of the growth lines with the dorsal margin, and the details of the sculpture. Hypoanthraconaia gen. nov. shows the most external similarity with ''atypical'' anthracosiid-like morphotypes of Anthraconaia that are widespread in the Late Pennsylvanian and Early Permian of eastern North America, and the Stephanian and Early Permian Lower Rotliegend of northwestern Europe. On this basis, the new genus is conventionally assigned to the family Naiaditidae Scarlato and Starobogatov, 1979.


Spatial niche partitioning in dinosaurs from the latest cretaceous (Maastrichtian) of North America

Abstract: We examine patterns of occurrence of associated dinosaur specimens (n = 343) from the North American Upper Cretaceous Hell Creek Formation and equivalent beds, by comparing their relative abundance in sandstone and mudstone. Ceratopsians preferentially occur in mudstone, whereas hadrosaurs and the small ornithopod Thescelosaurus show a strong association with sandstone. By contrast, the giant carnivore Tyrannosaurus rex shows no preferred association with either lithology. These lithologies are used as an indicator of environment of deposition, with sandstone generally representing river environments, and finer grained sediments typically representing floodplain environments. Given these patterns of occurrence, we argue that spatial niche partitioning helped reduce competition for resources between the herbivorous dinosaurs. Within coastal lowlands ceratopsians preferred habitats farther away from rivers, whereas hadrosaurs and Thescelosaurus preferred habitats in close proximity to rivers, and T. rex, the ecosystem's sole large carnivore, inhabited both palaeoenvironments. Spatial partitioning of the environment helps explain how several species of large herbivorous dinosaurs coexisted. This study emphasizes that different lithologies can preserve dramatically dissimilar vertebrate assemblages, even when deposited in close proximity and within a narrow window of time. The lithology in which fossils are preserved should be recorded as these data can provide unique insights into the palaeoecology of the animals they preserve.

maastrichtian
north america
dinosaur specimen
lithology


A high-resolution, multiproxy stratigraphic analysis of the Devonian–Carboniferous boundary sections in the Moravian Karst (Czech Republic) and a correlation with the Carnic Alps (Austria)

Abstract: Abstract A multidisciplinary correlation of the Devonian–Carboniferous (D–C) boundary sections from the Moravian Karst (Czech Republic) and the Carnic Alps (Austria), based on conodont and foraminifer biostratigraphy, microfacies analysis, field gamma-ray spectroscopy (GRS), carbon isotopes and element geochemistry, is presented in this paper. The study is focused on the interval from the Middle Palmatolepis gracilis expansa Zone (Late Famennian) to the Siphonodella sandbergi Zone (Early Tournaisian). In Lesní lom (Moravian Karst), a positive δ13C excursion in the Bisphatodus costatus – Protognathodus kockeli Interregnum from a distinct laminated carbonate horizon is correlated with a carbon isotope excursion from the Grüne Schneid section of the Carnic Alps and is interpreted as the equivalent of the Hangenberg black shales and a local expression of the global Hangenberg Event sensu stricto. Higher up at both sections, a significant increase in the terrigenous input, which is inferred from the GRS signal and elevated concentrations of terrigenous elements (Si, Ti, Zr, Rb, Al, etc.), provides another correlation tieline and is interpreted as the equivalent of the Hangenberg sandstone. Both horizons are discussed in terms of relative sea-level fluctuations and palaeoceanographic changes. Recent studies show that conodont biostratigraphy is facing serious problems associated with the taxonomy of the first siphonodellids, their dependence on facies and discontinuous occurrences of protognathodids at the D–C boundary. Therefore, the correlative potential of geochemical and petrophysical signatures is high and offers an alternative for the refining of the problematic biostratigraphic division of the D–C boundary.

boundary section
moravian karst
black shale


Filling the gap in knowledge of early Miocene continental molluscs of southwest Europe: gastropods from Ribesalbes-Alcora Basin (Spain)

Abstract: such as those of Vallés-Penedés, Ebro, Calatayud-Daroca, Teruel, Duero and Tajo, as well as smaller ones such as Cabriel, Fortuna, Guadix-Baza and Granada (Agustí 2018). Continental sediments of Aragonian age (end of the early Miocene to middle Miocene; mammal biozones MN4 to MN7/8) are represented in many of these basins (Agustí 2018). While mammal associations have been studied extensively in several palaeontological sites, the malacological record is comparatively poor documented. The few studies carried out on the malacological assemblages recorded in sediments today attributed to an Aragonian age are to some extent outdated (Vilanova y Piera 1859; Royo Gómez 1922, 1926, 1928; Jodot 1959). Also, many of sites and faunas documented in these works could not be precisely located. The available references accurately located and dated correspond to sites attributed to the middle Aragonian of the Tajo Basin (Truc 1977) and the upper Aragonian of the Duero Basin (González Delgado et al. 1986, Civis et al. 1989). The references from the early Aragonian are practically inexistent and correspond to isolated cites or recent multidisciplinary studies in the Ribesalbes-Alcora Basin (Peñalver et al. 2016, ÁlvarezParra et al. 2021). In line with the above, the available data on the mala cological content of the sediments of the Ribesalbes-Alcora Basin (Fig. 1A) are mostly old and very scarce. The first one corresponds to Vilanova y Piera (1859) who studied the materials in the “Balsa de Fanzaraˮ (the classical name for the Cenozoic sediments of the basin) and the nearby area of Alcalá. He cited the generic presence of “lymneasˮ, “planorbis” and “helicesˮ (Vilanova y Piera 1859: p. 44), and more specifically a single specimen of Planorbis rotundatus Brongniart, 1810 from the “Balsa de Fanzara”, which is shown on a plate alongside with four species belonging to the Alcalá area. Royo Gómez (1922) included a brief reference to the sediments of the studied area correcting the determination of Planorbis rotundatus to

early miocene
ribesalbes-alcora basin
mollusk group
insectivore


Systematics and Evolution of Paleozoic And Mesozoic Damselfly-Like Odonatoptera of the ‘Protozygopteran’ Grade

Abstract: Abstract The Paleozoic to Mesozoic grade ‘Protozygoptera’ is revised. It appears to be composed of two main lineages, namely the superfamily Permagrionoidea, and the Archizygoptera. The latter taxon forms a monophyletic group together with Panodonata ( = crown-Odonata plus their closest stem-relatives). Therefore, the ‘Protozygoptera’ as previously understood is paraphyletic. Diagnostic characters of the ‘Protozygoptera’, Permagrionoidea, and Archizygoptera are re-evaluated. The Permolestidae is considered as a junior synonym of the Permagrionidae. The following new taxa are described: Permolestes sheimogorai new species, Permolestes soyanaiensis new species, Epilestes angustapterix new species, Solikamptilon pectinatus new species (all in Permagrionidae); Lodeviidae new family (for Lodevia); Luiseiidae new family (including Luiseia breviata new genus and species); Kennedya azari new species, Kennedya pritykinae new species, Kennedya ivensis new species, Progoneura grimaldii new species (all in Kennedyidae); Engellestes chekardensis new genus and species (in Bakteniidae); and Azaroneura permiana new genus and species (in Voltzialestidae). The Kaltanoneuridae and Oboraneuridae are revised. The evolution of protozygopteran Odonatoptera during the transition from the Permian to the Triassic is discussed. The larger taxa of the permagrionoid lineage apparently did not cross through the Permian–Triassic boundary, unlike the more gracile Archizygoptera. This last group shows a remarkable longevity from the late Carboniferous to the Early Cretaceous. It also presents a great taxonomic and morphological stability, with genera ranging from the Permian to the Triassic, and a wing venation pattern nearly unchanged from the late Carboniferous to the Late Triassic. The mass extinction at the end of the Permian period seemingly had a minor effect on these tiny and delicate insects.


Chronology of Miocene-Pliocene Deposits at Split Mountain Gorge, Southern California: A Record of Regional Tectonics and Colorado River Evolution

Abstract: Late Miocene to early Pliocene deposits at Split Mountain Gorge, California, preserve a record of basinal response to changes in regional tectonics, paleogeography, and evolution of the Colorado River. The base of the Elephant Trees Formation, magnetostratigraphically dated as 8.1 ± 0.4 Ma, provides the earliest well-dated record of extension in the southwestern Salton Trough. The oldest marine sediments are ca. 6.3 Ma. The nearly synchronous timing of marine incursion in the Salton Trough and northern Gulf of California region supports a model for localization of Pacifi c‐North America plate motion in the Gulf ca. 6 Ma. The fi rst appearance of Colorado River sand at the Miocene-Pliocene boundary (5.33 Ma) suggests rapid propagation of the river to the Salton Trough, and supports a lake-spillover hypothesis for initiation of the lower Colorado River.

split mountain gorge
colorado river sand
miocene-pliocene boundary


Giant lizards occupied herbivorous mammalian ecospace during the Paleogene greenhouse in Southeast Asia

Abstract: Mammals dominate modern terrestrial herbivore ecosystems, whereas extant herbivorous reptiles are limited in diversity and body size. The evolution of reptile herbivory and its relationship to mammalian diversification is poorly understood with respect to climate and the roles of predation pressure and competition for food resources. Here, we describe a giant fossil acrodontan lizard recovered with a diverse mammal assemblage from the late middle Eocene Pondaung Formation of Myanmar, which provides a historical test of factors controlling body size in herbivorous squamates. We infer a predominately herbivorous feeding ecology for the new acrodontan based on dental anatomy, phylogenetic relationships and body size. Ranking body masses for Pondaung Formation vertebrates indicates that the lizard occupied a size niche among the larger herbivores and was larger than most carnivorous mammals. Paleotemperature estimates of Pondaung Formation environments based on the body size of the new lizard are approximately 2–5°C higher than modern. These results indicate that competitive exclusion and predation by mammals did not restrict body size evolution in these herbivorous squamates, and elevated temperatures relative to modern climates during the Paleogene greenhouse may have resulted in the evolution of gigantism through elevated poikilothermic metabolic rates and in response to increases in floral productivity.


Mechanical modelling of tooth wear

Abstract: Different diets wear teeth in different ways and generate distinguishable wear and microwear patterns that have long been the basis of palaeodiet reconstructions. Little experimental research has been performed to study them together. Here, we show that an artificial mechanical masticator, a chewing machine, occluding real horse teeth in continuous simulated chewing (of 100 000 chewing cycles) is capable of replicating microscopic wear features and gross wear on teeth that resemble wear in specimens collected from nature. Simulating pure attrition (chewing without food) and four plant material diets of different abrasives content (at n = 5 tooth pairs per group), we detected differences in microscopic wear features by stereomicroscopy of the chewing surface in the number and quality of pits and scratches that were not always as expected. Using computed tomography scanning in one tooth per diet, absolute wear was quantified as the mean height change after the simulated chewing. Absolute wear increased with diet abrasiveness, originating from phytoliths and grit. In combination, our findings highlight that differences in actual dental tissue loss can occur at similar microwear patterns, cautioning against a direct transformation of microwear results into predictions about diet or tooth wear rate.


Cenozoic environmental change in South America as indicated by mammalian body size distributions (cenograms)

Abstract: A cenogram is a rank‐ordered body size distribution of non‐predatory terrestrial mammal species within a community. Studies of cenograms for modern faunas have shown that certain quantifiable attributes of cenograms are correlated with environmental variables such as rainfall and vegetation structure. Based on these correlations, cenograms of fossil communities have been used to infer palaeoenvironments and palaeoenvironmental variables. The present study uses cenogram statistics to interpret palaeoenvironmental conditions for eight Cenozoic South American mammal faunas, ranging from Eocene to Pleistocene in age. Body sizes for fossil taxa were taken either from the literature or were estimated using regressions of body size on molar length (or femoral bicondylar width) for modern mammals. Cenogram statistics are calculated for the eight fossil faunas and compared to similar statistics calculated for 16 modern South American mammal faunas, allowing palaeoenvironmental interpretations to be made. The palaeoenvironmental interpretations based on cenogram analyses sometimes support and sometimes contradict interpretations based on herbivore craniodental morphology (e.g. levels of hypsodonty). Simulations of expected errors in body size estimates for fossil taxa suggest that the discrepancies do not result primarily from erroneous body size estimates. It is possible that some of the incongruity in interpretations results from certain non‐analogue attributes of South American faunas during much of the Cenozoic (e.g. the relatively depauperate mammalian predator diversity prior to the Great American Biotic Interchange).

body size distribution
cenogram
regression
bicondylar width


Elevation‐induced climate change as a dominant factor causing the late Miocene C4 plant expansion in the Himalayan foreland

Abstract: During the late Miocene, a dramatic global expansion of C4 plant distribution occurred with broad spatial and temporal variations. Although the event is well documented, whether subsequent expansions were caused by a decreased atmospheric CO2 concentration or climate change is a contentious issue. In this study, we used an improved inverse vegetation modeling approach that accounts for the physiological responses of C3 and C4 plants to quantitatively reconstruct the paleoclimate in the Siwalik of Nepal based on pollen and carbon isotope data. We also studied the sensitivity of the C3 and C4 plants to changes in the climate and the atmospheric CO2 concentration. We suggest that the expansion of the C4 plant distribution during the late Miocene may have been primarily triggered by regional aridification and temperature increases. The expansion was unlikely caused by reduced CO2 levels alone. Our findings suggest that this abrupt ecological shift mainly resulted from climate changes related to the decreased elevation of the Himalayan foreland.

climate change
himalayan foreland
deciduous forest taxa
plant megafossil evidence
vegetation shift


Minimum convex hull mass estimations of complete mounted skeletons

Abstract: Body mass is a critical parameter used to constrain biomechanical and physiological traits of organisms. Volumetric methods are becoming more common as techniques for estimating the body masses of fossil vertebrates. However, they are often accused of excessive subjective input when estimating the thickness of missing soft tissue. Here, we demonstrate an alternative approach where a minimum convex hull is derived mathematically from the point cloud generated by laser-scanning mounted skeletons. This has the advantage of requiring minimal user intervention and is thus more objective and far quicker. We test this method on 14 relatively large-bodied mammalian skeletons and demonstrate that it consistently underestimates body mass by 21 per cent with minimal scatter around the regression line. We therefore suggest that it is a robust method of estimating body mass where a mounted skeletal reconstruction is available and demonstrate its usage to predict the body mass of one of the largest, relatively complete sauropod dinosaurs: Giraffatitan brancai (previously Brachiosaurus) as 23200 kg.

mounted skeleton
alternative approach
body mass estimation
convex hull volume


Impact of a Tethyan circumglobal passage on ocean heat transport and “equable” climates

Abstract: [1] The presence of low-latitude circumglobal passage from the late Jurassic (∼160 Ma) through the Miocene (∼14 Ma) provides a possible mechanism for increased poleward ocean heat transport during periods of warm climate and may help explain low meridional temperature gradients of the past. Experiments using an ocean general circulation model (GCM) with an energy-balance atmosphere and idealized bathymetry reveal that, like the modern Drake Passage, a circumglobal Tethyan Passage might have induced high rates of wind-driven upwelling of relatively cold and deep water, but at low latitudes. With no change in radiative forcing, a low-latitude circumglobal passage increases simulated northern high-latitude temperatures by 3°–7°C, while tropical temperatures cool by up to 2°C relative to a scenario with solid meridional boundaries. Combining this mechanism of heat transport with increased radiative forcing allows substantial warming of northern high latitudes by 7°–11°C, while tropical temperatures remain within 3°C of present-day temperatures.

ocean heat transport
warming
high latitude


A Paleopolar Dinosaur Track Site in the Cretaceous (Maastrichtian) Prince Creek Formation of Arctic Alaska: Track Characteristics and Probable Trackmakers

Abstract: ABSTRACT For the first time a dinosaur track site is identified in Maastrichtian paleopolar coastal plain deposits of the Prince Creek Formation (PCF) along the Colville River, North Slope of Alaska. Tracks were made and preserved by trampling of an ash-covered swamp margin, subsequent filling of tracks with alluvium from nearby rivers, and modification of sediments by pedogensis. Tracks are grouped into three classes based on track width and depth, with the largest tracks (>800 mm wide) recording overstepping by multiple individuals. As no bedding plane views of the tracks are present, the true shapes of the tracks were not available and, thus, a high probability of identification is not achievable. The tracks can be interpreted, however, using hypothetical-deductive reasoning by integrating paleontological and ichnological data from local and regional outcrops. The tracks likely represent the presence of hadrosaurs based on the overwhelming percentage of hadrosaur fossils that comprise nearby bonebeds, dominated by juvenile hadrosaurs (∼ 99%); to date no adult hadrosaur bone has been documented in the PCF. This interpretation is also supported by comparison of PCF hadrosaur track dimensions to exquisitely preserved (three-dimensional tracks with skin impressions) trackways of the coeval Cantwell Formation in Denali National Park (DENA), central Alaska. PCF track size dimensions, in comparison to DENA tracks, also represent a series of growth stages including both juvenile and adult hadrosaurs, and indicate that multiple generations and sizes of individuals lived and traveled together on the Arctic Alaska coastal plain. This is the first evidence for adult hadrosaurs in the PCF. This track site also preserves the highest latitude Maastrichtian footprints known.

track
maastrichtian
prince creek formation
arctic alaska
colville river


Was Frozen Mammoth or Giant Ground Sloth Served for Dinner at The Explorers Club?

Abstract: Accounts of woolly mammoths (Mammuthus primigenius) preserved so well in ice that their meat is still edible have a long history of intriguing the public and influencing paleontological thought on Quaternary extinctions and climate, with some scientists resorting to catastrophism to explain the instantaneous freezing necessary to preserve edible meat. Famously, members of The Explorers Club purportedly dined on frozen mammoth from Alaska, USA, in 1951. This event, well received by the press and general public, became an enduring legend for the Club and popularized the notorious annual tradition of serving rare and exotic food at Club dinners that continues to this day. The Yale Peabody Museum holds a sample of meat preserved from the 1951 meal, interestingly labeled as a South American giant ground sloth (Megatherium), not mammoth. We sequenced a fragment of the mitochondrial cytochrome-b gene and studied archival material to verify its identity, which if genuine, would extend the range of Megatherium over 600% and alter our views on ground sloth evolution. Our results indicate that the meat was not mammoth or Megatherium but green sea turtle (Chelonia mydas). The prehistoric dinner was likely an elaborate publicity stunt. Our study emphasizes the value of museums collecting and curating voucher specimens, particularly those used for evidence of extraordinary claims.

frozen mammoth
explorers club
catastrophism


Acheulean technology and landscape use at Dawadmi, central Arabia

Abstract: Despite occupying a central geographic position, investigations of hominin populations in the Arabian Peninsula during the Lower Palaeolithic period are rare. The colonization of Eurasia below 55 degrees latitude indicates the success of the genus Homo in the Early and Middle Pleistocene, but the extent to which these hominins were capable of innovative and novel behavioural adaptations to engage with mid-latitude environments is unclear. Here we describe new field investigations at the Saffaqah locality (206–76) near Dawadmi, in central Arabia that aim to establish how hominins adapted to this region. The site is located in the interior of Arabia over 500 km from both the Red Sea and the Gulf, and at the headwaters of two major extinct river systems that were likely used by Acheulean hominins to cross the Peninsula. Saffaqah is one of the largest Acheulean sites in Arabia with nearly a million artefacts estimated to occur on the surface, and it is also the first to yield stratified deposits containing abundant artefacts. It is situated in the unusual setting of a dense and well-preserved landscape of Acheulean localities, with sites and isolated artefacts occurring regularly for tens of kilometres in every direction. We describe both previous and recent excavations at Saffaqah and its large lithic assemblage. We analyse thousands of artefacts from excavated and surface contexts, including giant andesite cores and flakes, smaller cores and retouched artefacts, as well as handaxes and cleavers. Technological assessment of stratified lithics and those from systematic survey, enable the reconstruction of stone tool life histories. The Acheulean hominins at Dawadmi were strong and skilful, with their adaptation evidently successful for some time. However, these biface-makers were also technologically conservative, and used least-effort strategies of resource procurement and tool transport. Ultimately, central Arabia was depopulated, likely in the face of environmental deterioration in the form of increasing aridity.


A new wing skeleton of the Jehol tapejarid Sinopterus and its implications for ontogeny and paleoecology of the Tapejaridae

Abstract: The tapejarid pterosaurs flourished in the Jehol Biota with an abundance of immature individuals and a rarity of individuals at skeletal maturity. Most of these individuals plot well on an ontogenetic series based on the proportions of limb elements, but this has lacked histological evidence until now. Here, a new wing skeleton of Sinopterus was thin-sectioned to provide the first histological data about the ontogeny of the Jehol tapejarids. Histologically, the new specimen is an immature individual at a late juvenile stage prior to sexual maturity. It is grouped with medium-sized and medium-crested individuals, which are distinct from the small-sized and crestless individuals as well as the rare large-sized and large-crested individuals at skeletal maturity, supporting the presence of the premaxillary crest as an ontogenetic feature in the Jehol tapejarids. Furthermore, this histology indicates that the largest skeletally immature individuals might have reached the sexual maturity. Enigmatically, there is a size gap between sexual and skeletal maturity, which is at about 79% of the large size, implying a ontogenetic strategy comparable with Pteranodon and possibly with the Brazilian tapejarid Caiuajara. This size gap is consistent with lack of the larger sexually mature individuals in the Jehol Biota, which is hypothesized to be a migratory habitat for the Jehol tapejarids.

new wing skeleton
jehol tapejarid
ontogeny


Juvenile spinosaurs (Theropoda: Spinosauridae) from the middle Cretaceous of Morocco and implications for spinosaur ecology

Abstract: The Spinosauridae is a specialised clade of theropod dinosaurs known from the Berriasian to the Cenomanian of Africa, South America, Europe and Asia. Spinosaurs were unusual among non-avian dinosaurs in exploiting a piscivorous niche within riverine and estuarine habitats, and they include the largest known theropod. Although fossils of giant spinosaurs are increasingly well-represented in the fossil record, little juvenile material has been described. Here, we describe new examples of juvenile spinosaurines from the middle Cretaceous (Cenomanian) Kem Kem beds of Morocco. The fossils include material from a range of sizes and are relatively common within the Kem Kem deposits, suggesting that juveniles exploited the same semiaquatic niche as the adults throughout ontogeny. This implies that the Cenomanian delta habitats supported an age-inclusive population of spinosaurs that was neither geographically or environmentally separated, though some ecological separation between juveniles and adults is likely based on the large variation in size. Bones or teeth of very small (<2 m) spinosaurs have not been found, however. This could represent a taphonomic bias, or potentially an ecological signal that the earliest ontogenetic stages inhabited distinct environments. Skeletal remains include individuals referable to Sigilmassasaurus brevicollis and a second spinosaurine taxon. Consistent with this, we also identify two distinct cranial morphs of Spinosauridae present within the Kem Kem, supporting previous recognition of distinct taxa in the assemblage.

spinosaur
spinosauridae
non-avian dinosaur
juvenile material


A Jurassic stem pleurodire sheds light on the functional origin of neck retraction in turtles

Abstract: Modern turtles are composed of two monophyletic groups, notably diagnosed by divergent neck retraction mechanisms. Pleurodires (side-necked turtles) bend their neck sideways and protect their head under the anterior margin of the carapace. Cryptodires (hidden-necked turtles) withdraw their neck and head in the vertical plane between the shoulder girdles. These two mechanisms of neck retraction appeared independently in the two lineages and are usually assumed to have evolved for protective reasons. Here we describe the neck of Platychelys oberndorferi, a Late Jurassic early stem pleurodire, and find remarkable convergent morphological and functional similarities with modern cryptodires. Partial vertical neck retraction in this taxon is interpreted to have enabled fast forward projection of the head during underwater prey capture and offers a likely explanation to the functional origin of neck retraction in modern cryptodires. Complete head withdrawal for protection may therefore have resulted from an exaptation in that group.

jurassic
functional origin
neck retraction
turtle
platychelys oberndorferi


The ammonite septum is not an adaptation to deep water: re-evaluating a centuries-old idea

Abstract: The shells of ammonoid cephalopods are among the most recognizable fossils, whose fractally folded, internal walls (septa) have inspired many hypotheses on their adaptive value. The enduring explanation for their iterative evolution is that they strengthen the shell against pressure at increasing water depths. The fossil record does not definitively support this idea and much of the theoretical mechanical work behind it has suffered from inaccurate testing geometries and conflicting results. By using a different set of mathematical methods compared with previous studies, I generate a system of finite-element models that explore how different parameters affect the shell's response to water pressure. Increasing the number of initial folds of the septa ultimately has little to no effect on the resulting stress in the shell wall or the septum itself. The introduction of higher-order folds does reduce the tensile stress in the shell wall; however, this is coupled with a higher rate of increase of tensile stress in the septum itself. These results reveal that the increase in complexity should not be expected to have a significant effect on the shell's strength and suggests that the evolution of ammonitic septa does not reflect a persistent trend towards deeper-water habitats.


Appendages of an early Cambrian metadoxidid trilobite from Yunnan, SW China support mandibulate affinities of trilobites and artiopods

Abstract: Abstract Appendage anatomy contributes crucial data for understanding the evolution and ecology of Euarthropoda. The Palaeozoic trilobites show a great diversity of exoskeletons in the fossil record. However, soft parts, especially appendages, have only been discovered from a few trilobite species. Here we report extraordinarily preserved appendages in the trilobite species Hongshiyanaspis yiliangensis Zhang & Lin in Zhang et al. 1980 (Redlichiida, Metadoxididae) from a single mudstone layer of the Xiazhuang fossil assemblage within the Hongjingshao Formation (Cambrian Series 2, Stage 3) near Kunming, Yunnan, SW China. The appendages exhibit the common architecture revealed by other trilobites and artiopods by consisting of a pair of uniramous antennae followed by a series of paired homonomous biramous limbs. The antennae in holaspid individuals comprise up to 27 spinous podomeres and their ontogeny occurs by lengthening of the podomeres. The post-antennal biramous limbs are similar to those in other polymerid trilobites and artiopods by having a single-segmented protopodite and an endopodite comprising seven segments, but possess a unique wide tripartite exopodite with long setae. Sophisticated appendage anatomy, including the body–limb junction, fine setae, putative muscle bundles and duct-type tissues, are also revealed. Appendages of trilobites, artiopods and other upper stem-group euarthropods are compared and summarized. The H. yiliangensis appendages highlight the high morphological disparity of exopodites and the conservativeness of endopodites in trilobites and artiopods. This morphological pattern, together with similar body patterning seen in crustaceans but not in chelicerates, supports the mandibulate affinities of trilobites and at least some artiopods.

trilobite
mandibulate affinity
euarthropod


Osteology of Eubiodectes libanicus (Pictet & Humbert, 1866) and some other ichthyodectiformes (Teleostei): phylogenetic implications

Abstract: The osteology of the ichthyodectiform Eubiodectes libanicus (Pictet & Humbert, 1866) from the Cenomanian of Lebanon is described on the basis of acid-prepared specimens. For comparative purposes, the osteology of other putative ichthyodectiforms is described or discussed. We focus on the description of hitherto poorly described species such as Chiromystus mawsoni Cope, 1885, Ghrisichthys bardacki (Cavin, 1997a), ‘Saurodon' intermedius (Newton, 1878) and Thrissops sp. from Kimmeridge, England. Sixty-nine morphological characters are defined and analysed to infer the phylogenetic relationships within ichthyodectiforms. The main results are: (1) exclusion of Ascalabothrissops and Pachythrissops from the ichthyodectiforms; (2) redesignation of ‘Ichthyodectes' bardacki, from the Turonian of Morocco, to Ghrisichthys gen. nov.; (3) resolution of a monophyletic family Cladocyclidae grouping mid-Cretaceous ichthyodectiforms that lived mainly along the southern margin of Tethys (Chirocentrites, Cladocyclus, Eubiodectes, Chiromystus); (4) resolution of a Late Cretaceous family Ichthyodectidae containing Ichthyodectes, Ghrisichthys and Xiphactinus from mid-northern latitude seas; and (5) a better resolved Late Cretaceous family Saurodontidae in the northern hemisphere, that includes the genus Gillicus.


Inference of facultative mobility in the enigmatic Ediacaran organism Parvancorina

Abstract: Establishing how Ediacaran organisms moved and fed is critical to deciphering their ecological and evolutionary significance, but has long been confounded by their non-analogue body plans. Here, we use computational fluid dynamics to quantitatively analyse water flow around the Ediacaran taxon Parvancorina, thereby testing between competing models for feeding mode and mobility. The results show that flow was not distributed evenly across the organism, but was directed towards localized areas; this allows us to reject osmotrophy, and instead supports either suspension feeding or detritivory. Moreover, the patterns of recirculating flow differ substantially with orientation to the current, suggesting that if Parvancorina was a suspension feeder, it would have been most efficient if it was able to re-orient itself with respect to current direction, and thus ensure flow was directed towards feeding structures. Our simulations also demonstrate that the amount of drag varied with orientation, indicating that Parvancorina would have greatly benefited from adjusting its position to minimize drag. Inference of facultative mobility in Parvancorina suggests that Ediacaran benthic ecosystems might have possessed a higher proportion of mobile taxa than currently appreciated from trace fossil studies. Furthermore, this inference of movement suggests the presence of musculature or appendages that are not preserved in fossils, but which would noneltheless support a bilaterian affinity for Parvancorina.

facultative mobility
osmotrophy
suspension feeding
inference


Thermally‐induced structural and chemical alteration of organic‐walled microfossils: an experimental approach to understanding fossil preservation in metasediments

Abstract: The identification and confirmation of bona fide Archean–Paleoproterozoic microfossils can prove to be a challenging task, further compounded by diagenetic and metamorphic histories. While structures of likely biological origin are not uncommon in Precambrian rocks, the search for early fossil life has been disproportionately focused on lesser thermally altered rocks, typically greenschist or lower‐grade metamorphism. Recently, however, an increasing number of inferred micro‐ and macrofossils have been reported from higher‐grade metasediments, prompting us to experimentally test and quantify the preservability of organic‐walled microfossils over varying durations of controlled heating and under two differing redox conditions. Because of their relatively low‐intensity natural thermal alteration, acritarchs from the Mesoproterozoic Ruyang Group were chosen as subjects for experimental heating at approximately 500°C, with durations ranging from 1 to 250 days and in both oxic (normal present day conditions) and anoxic conditions. Upon extraction, the opacity, reflectivity, color, microchemistry, and microstructures of the heated acritarchs were characterized using optic microscopy, scanning electron microscopy, Raman spectroscopy, and X‐ray photoelectron spectroscopy. The results differ for acritarchs prepared under oxic vs. anoxic conditions, with the anoxic replicates surviving experimental heating longer and retaining biological morphologies better, despite an increasing degree of carbonization with continuous heating. Conversely, the oxic replicates show aggressive degradation. In conjunction with fossils from high‐grade metasediments, our data illustrate the preservational potential of organic‐walled microfossils subjected to metamorphism in reducing conditions, offer insights into the search for microfossils in metasediments, and help to elucidate the influence of time on the carbonization/graphitization processes during thermal alteration.

alteration
organic‐walled microfossil
metasediment
redox condition
laboratory experiment


The first Silurian trilobite with three‐dimensionally preserved soft parts reveals novel appendage morphology

Abstract: The first Silurian trilobite known with soft parts preserved, a Dalmanites species, is described from the Herefordshire Lagerstätte. Biramous appendages and much of the alimentary system are evident. High‐fidelity three‐dimensional preservation reveals a novel arrangement of the exopod, in which successive filaments are connected by a presumed membrane. This morphology explains a misinterpretation of the exopod as supporting spiral structures, originally reported nearly 150 years ago. Comparison with other trilobite limbs indicates that the exopod morphology of Dalmanites is present in other members of Phacopida. The function of the exopod is considered to be primarily respiratory.

silurian trilobite
soft part
morphology
filament


Phasing of millennial-scale climate variability in the Pacific and Atlantic Oceans

Abstract: Calving cousins Walczak et al. report that increases in Pacific Ocean ventilation and periods of rapid production of icebergs from the Cordilleran Ice Sheet during the last glacial period preceded episodic iceberg discharges into the Atlantic Ocean (see the Perspective by Jaeger and Shevenell). Marine sediments from the Gulf of Alaska show that increases in vertical mixing of the ocean there correspond with intense iceberg calving from the ice sheet that covered much of high-latitude western North America and that these changes occurred before the analogous Heinrich events in the North Atlantic. Thus, these Pacific climate system reorganizations may have been an early part of a cascade of dynamic climate events with global repercussions. Science, this issue p. 716; see also p. 662 Pacific iceberg discharge events preceded their Atlantic counterparts during the last glacial period. New radiocarbon and sedimentological results from the Gulf of Alaska document recurrent millennial-scale episodes of reorganized Pacific Ocean ventilation synchronous with rapid Cordilleran Ice Sheet discharge, indicating close coupling of ice-ocean dynamics spanning the past 42,000 years. Ventilation of the intermediate-depth North Pacific tracks strength of the Asian monsoon, supporting a role for moisture and heat transport from low latitudes in North Pacific paleoclimate. Changes in carbon-14 age of intermediate waters are in phase with peaks in Cordilleran ice-rafted debris delivery, and both consistently precede ice discharge events from the Laurentide Ice Sheet, known as Heinrich events. This timing precludes an Atlantic trigger for Cordilleran Ice Sheet retreat and instead implicates the Pacific as an early part of a cascade of dynamic climate events with global impact.

pacific
pacific ocean ventilation
millennial-scale episode


Asmodochelys parhami, a new fossil marine turtle from the Campanian Demopolis Chalk and the stratigraphic congruence of competing marine turtle phylogenies

Abstract: Resolving the phylogeny of sea turtles is uniquely challenging given the high potential for the unification of convergent lineages due to systematic homoplasy. Equivocal reconstructions of marine turtle evolution subsequently inhibit efforts to establish fossil calibrations for molecular divergence estimates and prevent the accurate reconciliation of biogeographic or palaeoclimatic data with phylogenetic hypotheses. Here we describe a new genus and species of marine turtle, Asmodochelys parhami, from the Upper Campanian Demopolis Chalk of Alabama and Mississippi, USA represented by three partial shells. Phylogenetic analysis shows that A. parhami belongs to the ctenochelyids, an extinct group that shares characteristics with both pan-chelonioids and pan-cheloniids. In addition to supporting Ctenochelyidae as a sister taxon of Chelonioidea, our analysis places Protostegidae outside of the Chelonioidea crown group and recovers Allopleuron hofmanni as a stem dermochelyid. Gap excess ratio (GER) results indicate a strong stratigraphic congruence of our phylogenetic hypothesis; however, the highest GER value is associated with the phylogenetic hypothesis of marine turtles which excludes Protostegidae from the Cryptodira crown group. Ancestral range estimations derived from our phylogeny imply a European or North American origin of Chelonioidea in the middle-to-late Campanian, approximately 20 Myr earlier than current molecular divergence studies suggest.

turtle
stratigraphic congruence
protostegidae
asmodochelys parhami


The phylogenetic position of the Tyrannosauridae: implications for theropod systematics

Abstract: Tyrannosaurids are a well-supported clade of very large predatory dinosaurs of Late Cretaceous Asiamerica. Traditional dinosaurian systematics place these animals within the infraorder Carnosauria with the other large theropods (allosaurids, megalosaurids). A new cladistic analysis indicates that the tyrannosaurs were in fact derived members of the Coelurosauria, a group of otherwise small theropods. Despite certain gross cranial similarities with the large predators of the Jurassic and Early Cretaceous, the Late Cretaceous tyrannosaurids are shown to be the sister group to ornithomimids and troodontids, which share a derived condition of the metatarsus. This clade is found to be nested within Maniraptora, which is a more inclusive taxon than previously recognized. The atrophied carpal structure found in tyrannosaurids and ornithomimids is derived from a maniraptoran condition with a large semilunate carpal, rather than from the plesiomorphic theropod morphology. The taxa “Carnosauria” and “Deinonychosauria” (Dromaeosauridae plus Troodontidae) are shown to be polyphyletic, and the Late Jurassic African form Elaphrosaurus is found to be the sister taxon to Abelisauridae rather than a primitive ornithomimosaur. Purported allosaurid-tyrannosaurid synapomorphies are seen to be largely size-related, present in the larger members of both clades, but absent in smaller members of the Tyrannosauridae. The remaining giant tetanurine theropods (Megalosaurus and Torvosaurus) were found to be progressively distant outgroups to an allosaurid-coelurosaur clade. The inclusion of the Tyrannosauridae within Maniraptora suggests a major adaptive radiation of coelurosaurs within Cretaceous Asiamerica comparable to contemporaneous radiations in various herbivorous dinosaurian clades.


Spontaneous formation and base pairing of plausible prebiotic nucleotides in water

Abstract: The RNA World hypothesis presupposes that abiotic reactions originally produced nucleotides, the monomers of RNA and universal constituents of metabolism. However, compatible prebiotic reactions for the synthesis of complementary (that is, base pairing) nucleotides and mechanisms for their mutual selection within a complex chemical environment have not been reported. Here we show that two plausible prebiotic heterocycles, melamine and barbituric acid, form glycosidic linkages with ribose and ribose-5-phosphate in water to produce nucleosides and nucleotides in good yields. Even without purification, these nucleotides base pair in aqueous solution to create linear supramolecular assemblies containing thousands of ordered nucleotides. Nucleotide anomerization and supramolecular assemblies favour the biologically relevant β-anomer form of these ribonucleotides, revealing abiotic mechanisms by which nucleotide structure and configuration could have been originally favoured. These findings indicate that nucleotide formation and selection may have been robust processes on the prebiotic Earth, if other nucleobases preceded those of extant life.

base pairing
nucleotide
melamine
barbituric acid
aqueous solution


Plant and insect herbivore community variation across the Paleocene–Eocene boundary in the Hanna Basin, southeastern Wyoming

Abstract: Ecosystem function and stability are highly affected by internal and external stressors. Utilizing paleobotanical data gives insight into the evolutionary processes an ecosystem undergoes across long periods of time, allowing for a more complete understanding of how plant and insect herbivore communities are affected by ecosystem imbalance. To study how plant and insect herbivore communities change during times of disturbance, we quantified community turnover across the Paleocene–Eocene boundary in the Hanna Basin, southeastern Wyoming. This particular location is unlike other nearby Laramide basins because it has an abundance of late Paleocene and Eocene coal and carbonaceous shales and paucity of well-developed paleosols, suggesting perpetually high water availability. We sampled approximately 800 semi-intact dicot leaves from five stratigraphic levels, one of which occurs late in the Paleocene–Eocene thermal maximum (PETM). Field collections were supplemented with specimens at the Denver Museum of Nature & Science. Fossil leaves were classified into morphospecies and herbivore damage was documented for each leaf. We tested for changes in plant and insect herbivore damage diversity using rarefaction and community composition using non-metric multidimensional scaling ordinations. We also documented changes in depositional environment at each stratigraphic level to better contextualize the environment of the basin. Plant diversity was highest during the mid-late Paleocene and decreased into the Eocene, whereas damage diversity was highest at the sites with low plant diversity. Plant communities significantly changed during the late PETM and do not return to pre-PETM composition. Insect herbivore communities also changed during the PETM, but, unlike plant communities, rebound to their pre-PETM structure. These results suggest that insect herbivore communities responded more strongly to plant community composition than to the diversity of species present.

paleocene–eocene boundary
hanna basin
southeastern wyoming
fossil leave


The Triassic of Traill Ø and Geographical Society Ø, East Greenland: Implications for North Atlantic palaeogeography

Abstract: The continental Triassic succession north of 72° in East Greenland has seen little investigation but is key in understanding how facies belts vary towards the East Greenland Shelf, and more widely in the North Atlantic region, through this period. This study presents sedimentological analysis of exposures in northern Traill Ø (the Mols Bjerge) and further north in northern Geographical Society Ø (Laplace Bjerg). These sections are correlated with the more widely studied succession which lies to the south, in Jameson Land and broad scale palaeogeographical reconstructions of the North Atlantic are presented. A largely continental Triassic succession of 1445 m was recorded from the Mols Bjerge. However, a 125 m thick clean sand unit (the Vega Sund Member), which includes in its upper part the Gråklint Beds, is re‐interpreted as of shallow marine origin. Over 1200 m of Triassic strata were recognized on Laplace Bjerg. Within this succession 300 m of clean, cross‐bedded sandstones are recorded and are correlated with the Vega Sund Member recorded in the Mols Bjerge. The northward increase in thickness of marine strata developed at this time provides evidence for a more extensive marine influence in the north. This work has important implications for regional palaeogeographies and the character of the Triassic succession in adjacent basins, in particular, constraining the southern extent of the Boreal Ocean during the Mid to Late Triassic. Furthermore, the correlations made places bitumen staining, reported from Laplace Bjerg, within Triassic strata, suggesting the viability of a Triassic play at depth in adjacent offshore basins.


A SPALACOLESTINE SPALACOTHERIID (MAMMALIA, TRECHNOTHERIA) FROM THE EARLY CRETACEOUS (BARREMIAN) OF SOUTHERN ENGLAND AND ITS BEARING ON SPALACOTHERIID EVOLUTION

Abstract: Abstract:  Bulk screening of Early Cretaceous (Barremian) Wealden Group strata of the Wessex Formation exposed on the south‐west and south‐east coasts of the Isle of Wight, southern England, has resulted in the recovery of fragmentary remains pertaining to a new spalacolestine spalacotheriid mammal, Yaverlestes gassoni gen. et sp. nov. These represent the first European record of the Spalacolestinae. The remains comprise a dentulous incomplete dentary and isolated upper and lower molariforms, the former representing the most substantial mammal remains yet recovered from the Wealden Group. Hitherto, six species of spalacotheriid mammal were known from the Lower Cretaceous of Europe. All are referred to the genus Spalacotherium but in the case of taxa diagnosed on the basis of isolated lower teeth and other specimens where the post‐canine dentition is incompletely known, it is now evident that these referrals should be treated with caution. Furthermore, the new Wessex Formation spalacotheriid and recently described spalacotheriids from the ?Barremian of Japan, and the Barremian and Aptian of China exhibit combinations of characters that suggest that spalacotheriids were more diverse and that their evolution was more complex than previously recognized. The systematic position of an isolated tooth from the basal Cretaceous Lourinhã Formation of Portugal is discussed and the tooth reassigned to the Spalacotheriidae. Together with the new Wessex Formation taxon, eight species are now known from the Lower Cretaceous of Europe. The discovery of a spalacolestine in the Barremian Wessex Formation supports the concept of faunal interchange between Europe, Asia and North America during the Early Cretaceous. It also supports derivation of North American spalacotheriids from a European or Eurasian ancestor.

barremian
southern england
mammal remain
spalacotheriidae


Whence the beardogs? Reappraisal of the Middle to Late Eocene ‘Miacis’ from Texas, USA, and the origin of Amphicyonidae (Mammalia, Carnivora)

Abstract: The Middle to Late Eocene sediments of Texas have yielded a wealth of fossil material that offers a rare window on a diverse and highly endemic mammalian fauna from that time in the southern part of North America. These faunal data are particularly significant because the narrative of mammalian evolution in the Paleogene of North America has traditionally been dominated by taxa that are known from higher latitudes, primarily in the Rocky Mountain and northern Great Plains regions. Here we report on the affinities of two peculiar carnivoraforms from the Chambers Tuff of Trans-Pecos, Texas, that were first described 30 years ago as Miacis cognitus and M. australis. Re-examination of previously described specimens and their inclusion in a cladistic analysis revealed the two taxa to be diminutive basal amphicyonids; as such, they are assigned to new genera Gustafsonia and Angelarctocyon, respectively. These two taxa fill in some of the morphological gaps between the earliest-known amphicyonid genus, Daphoenus, and other Middle-Eocene carnivoraforms, and lend additional support for a basal caniform position of the beardogs outside the Canoidea. The amphicyonid lineage had evidently given rise to at least five rather distinct forms by the end of the Middle Eocene. Their precise geographical origin remains uncertain, but it is plausible that southern North America served as an important stage for a very early phase of amphicyonid radiation.


Tectonic models for accretion of the Central Asian Orogenic Belt

Abstract: The Central Asian Orogenic Belt (c. 1000–250 Ma) formed by accretion of island arcs, ophiolites, oceanic islands, seamounts, accretionary wedges, oceanic plateaux and microcontinents in a manner comparable with that of circum-Pacific Mesozoic–Cenozoic accretionary orogens. Palaeomagnetic and palaeofloral data indicate that early accretion (Vendian–Ordovician) took place when Baltica and Siberia were separated by a wide ocean. Island arcs and Precambrian microcontinents accreted to the active margins of the two continents or amalgamated in an oceanic setting (as in Kazakhstan) by roll-back and collision, forming a huge accretionary collage. The Palaeo-Asian Ocean closed in the Permian with formation of the Solonker suture. We evaluate contrasting tectonic models for the evolution of the orogenic belt. Current information provides little support for the main tenets of the one- or three-arc Kipchak model; current data suggest that an archipelago-type (Indonesian) model is more viable. Some diagnostic features of ridge–trench interaction are present in the Central Asian orogen (e.g. granites, adakites, boninites, near-trench magmatism, Alaskan-type mafic–ultramafic complexes, high-temperature metamorphic belts that prograde rapidly from low-grade belts, rhyolitic ash-fall tuffs). They offer a promising perspective for future investigations.

accretion
tectonic model
subduction
late palaeozoic
metamorphic rock


Thirty years on the “Broad Spectrum Revolution” and paleolithic demography

Abstract: All Paleolithic hominids lived by hunting and collecting wild foods, an aspect of existence that began to disappear only with the emergence of the farming and herding societies of the Neolithic ≤10,000 years ago (10 KYA). What are the roots of this remarkable economic transformation? The answer lies in equally revolutionary changes that took place within certain stone age cultures several millennia before. In 1968, Lewis R. Binford noted what appeared to be substantial diversification of human diets in middle- and high-latitude Europe at the end of the Paleolithic, roughly 12–8 KYA (1). Rapid diversification in hunting, food processing, and food storage equipment generally accompanied dietary shifts, symptoms of intensified use of habitats, and fuller exploitation of the potential foodstuffs they contained. Some of this behavior was directed to grinding, drying, and storing nuts, but it also involved small animals (2–6). Kent Flannery pushed these observations further in 1969 with his “Broad Spectrum Revolution” (BSR) hypothesis, proposing that the emergence of the Neolithic in western Asia was prefaced by increases in dietary breadth in foraging societies just before this period (7). He argued that subsistence diversification, mainly by adding new species to the diet, raised the carrying capacity of an environment increasingly constrained by climate instability at the end of the Pleistocene.

broad spectrum revolution
human diet
environment
pleistocene


Effects of the early Toarcian Oceanic Anoxic Event on ichthyosaur body size and faunal composition in the Southwest German Basin

Abstract:
Abstract.
 The Early Jurassic Toarcian Oceanic Anoxic Event is considered one of the most dramatic environmental perturbations of the Mesozoic. An elevated extinction rate among marine invertebrates accompanied rapid environmental changes, but effects on large vertebrates are less understood. We examined changes in ichthyosaur body size in the Posidonia Shale of the Southwest German Basin spanning the extinction interval to assess how environmental changes and biotic crisis among prey species affected large reptiles. We report no species-level extinction among the ichthyosaurs coinciding with peak invertebrate extinction. Large ichthyosaurs were absent from the fauna during the extinction interval, but became more abundant in the immediate aftermath. Stenopterygius quadriscissus, the most abundant species during the extinction interval, increased in body size after the biotic event. Rapid invasion by large taxa occurred immediately following the extinction event at the end of the first ammonite zone of the early Toarcian. Greater mobility permitting exploitation of ephemeral resources and opportunistic feeding behavior may minimize the impacts of environmental change on large vertebrates.

ichthyosaur body size
southwest german basin
invertebrate
vertebrate


Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia.

Abstract: Were molecular data available for extinct taxa, questions regarding the origins of many groups could be settled in short order. As this is not the case, various strategies have been proposed to combine paleontological and neontological data sets. The use of fossil dates as node age calibrations for divergence time estimation from molecular phylogenies is commonplace. In addition, simulations suggest that the addition of morphological data from extinct taxa may improve phylogenetic estimation when combined with molecular data for extant species, and some studies have merged morphological and molecular data to estimate combined evidence phylogenies containing both extinct and extant taxa. However, few, if any, studies have attempted to estimate divergence times using phylogenies containing both fossil and living taxa sampled for both molecular and morphological data. Here, I infer both the phylogeny and the time of origin for Lissamphibia and a number of stem tetrapods using Bayesian methods based on a data set containing morphological data for extinct taxa, molecular data for extant taxa, and molecular and morphological data for a subset of extant taxa. The results suggest that Lissamphibia is monophyletic, nested within Lepospondyli, and originated in the late Carboniferous at the earliest. This research illustrates potential pitfalls for the use of fossils as post hoc age constraints on internal nodes and highlights the importance of explicit phylogenetic analysis of extinct taxa. These results suggest that the application of fossils as minima or maxima on molecular phylogenies should be supplemented or supplanted by combined evidence analyses whenever possible.

lissamphibia
calibration
phylogenetic analysis
divergence time estimation
fossil specie


The evolution of feeding within Euchelicerata: data from the fossil groups Eurypterida and Trigonotarbida illustrate possible evolutionary pathways

Abstract: When the evolution of Euarthropoda is discussed, often the lineage of Chelicerata s. str. is assumed to be the more ‘primitive’ or ‘basal’ part of the tree, especially when compared to the other major lineage, Mandibulata. This claimed primitiveness is (at least partly) based on the assumption that different morphological structures are still in an ancestral state and did not evolve any further. One of these sets of structures is the feeding apparatus, which has been stated to be highly advanced in Mandibulata, but not ‘properly’ developed, or at least not to such a high degree, within Chelicerata s. str. In this study, I reinvestigate the feeding apparatus of different ingroups of Euchelicerata, with a focus on assumed ‘primitive’ groups such as Eurypterida and Trigonotarbida. The basis of this study is a large amount of material from different museum collections, with fossils with the entire feeding apparatuses being exceptionally well preserved. Based on high-resolution micro-photography and three-dimensional imaging, it is possible to resolve fine details of the feeding apparatuses. The results make clear that the feeding apparatuses of different ingroups of Euchelicerata are highly specialised and often possess morphological structures comparable to those of the feeding apparatuses of representatives of Mandibulata, apparently convergently evolved. Though the reconstruction of the evolution of the feeding apparatus within Euchelicerata is to a certain degree hampered by unclear phylogenetic relationships, there was clearly a shortening of the feeding apparatus from posterior (i.e. only the anterior appendages being involved in the feeding apparatus), probably linked to the colonisation of land in Arachnida. Subjects Evolutionary Studies, Paleontology, Zoology