The Interatheriinae (Mammalia, Notoungulata) of the Friasian sensu stricto and Mayoan (middle to late Miocene), and the fossils from Cerro Zeballos, Patagonia, Argentina

Abstract: The middle to late Miocene Friasian sensu stricto, Colloncuran and Mayoan South American Land Mammal Ages (SALMAs) were established based on the mammals recovered from the Río Frías, Collón Curá and Río Mayo formations, respectively (southern Argentina and Chile). Records of Interatheriinae (Notoungulata) from these sediments have been known since the first contributions of Ameghino and Roth, but most of this material has only been superficially revised or is almost unstudied. Based on these old collections and new interatheriine records from Cerro Zeballos (Chubut Province), we improve the diagnosis of Caenophilus tripartitus, describe new and peculiar characters for this taxon, consider Epipatriarchus bifidens as Protypotherium cf. colloncurensis, and propose Epipatriarchus as a junior synonym of Protypotherium. In turn, the holotype of Epipatriarchus innexus is here identified as the mesotheriid Trachytypotherium sp. Therefore, only two genera of Interatheriinae, Protypotherium and Caenophilus, are recognized in the Friasian sensu stricto and Mayoan. We identify Protypotherium cf. endiadys and a probable new species of Protypotherium for the Friasian sensu stricto, P. cf. colloncurensis and Caenophilus tripartitus for the Mayoan, and C. tripartitus, P. colloncurensis and P. cf. endiadys for Cerro Zeballos. An extended phylogenetic analysis is performed, including species from low latitudes and new data for P. colloncurensis and Caenophilus tripartitus, with the latter recovered as a sister taxon to Miocochilius federicoi. Both Protypotherium and Miocochilius are recovered as paraphyletic. The updated information we present provides a better knowledge of Interatheriinae for the middle to late Miocene of southern South America.

interatheriinae
notoungulata
late miocene


The Shibantan Lagerstätte: insights into the Proterozoic–Phanerozoic transition

Abstract: The Shibantan Lagerstätte (551–543 Ma) in the Yangtse Gorges area in South China is one of the best-known examples of terminal Ediacaran fossil assemblages preserved in marine carbonate rocks. Taxonomically dominated by benthic organisms, the Shibantan Lagerstätte preserves various photoautotrophs, biomineralizing tubular fossils, Ediacara-type macrofossils (including rangeomorphs, arboreomorphs, erniettomorphs, palaeopascichnids, a possible dickinsoniomorph, the mobile bilaterian Yilingia and soft-bodied tubular fossils), abundant ichnofossils and a number of problematic and dubious fossils. Shibantan fossils provide intriguing insights into ecological interactions among mobile bilaterians, sessile benthic Ediacara-type organisms and microbial mats, thus offering important data to test various hypotheses accounting for the decline of the Ediacara biota and the concurrent expansion of bilaterian bioturbation and mobility across the Proterozoic–Phanerozoic transition.

shibantan lagerstätte
proterozoic–phanerozoic transition
mobility


Ontogeny of the trilobite Elrathia kingii (Meek) and comparison of growth rates between Elrathia kingii and Aulacopleura koninckii (Barrande)

Abstract: Trilobites offer almost unparalleled insight into the growth and development of fossil ecdysozoans. Here I use newly collected material of Elrathia kingii (Meek) to estimate growth rates and describe shape change over the ontogeny of E. kingii. Well‐preserved, articulated specimens from all post‐embryonic stages were collected from a 1.5 m interval of the upper Wheeler Formation (Miaolingian Series, Cambrian) in western Utah (USA), and size and landmark‐based shape data were digitized from photographs. Growth rates were estimated and compared with previously published data on the Silurian trilobite, Aulacopleura koninckii (Barrande). Like A. koninckii, the cephalic growth rate in E. kingii was constant and of similar magnitude to the minimum growth rate along the trunk, and growth rates in the trunk were lower during the holaspid (‘adult’) period than during the meraspid (‘juvenile’) period. However, body length at the onset of meraspis was smaller, the growth gradient along the trunk during meraspis was shallower, and the terminal number of thoracic tergites was smaller in E. kingii than in A. koninckii. Despite these differences, these two species had similar maximum body lengths, because higher overall growth rates in E. kingii compensated for other differences. The rate of cephalic shape change in E. kingii decreased at the transition from meraspis to holaspis, while the pygidium became more morphologically distinct from the thorax during holaspis. I also provide an emended diagnosis for E. kingii, descriptions of the ontogeny and ventral morphology, and evidence that E. kingii holaspids had an invariant number of tergites.


Homologies of the longissimus, iliocostalis, and hypaxial muscles in the anterior presacral region of extant diapsida

Abstract: Homologies of muscles of the m. longissimus and m. iliocostalis groups in the dorsal and cervical regions, as well as those of the subvertebral muscles and mm. intercostales externi that continue from the dorsal into the cervical regions, in extant Diapsida are proposed based on detailed dissections and published accounts of lepidosaurs, crocodylians, and birds. The morphology of tendons and innervation patterns suggest that the avian “m. iliocostalis” in the dorsal region include the homologs of both m. longissimus and m. iliocostalis in non‐avian diapsids. The conserved nature of the morphology of tendons in palaeognath birds also revealed that the avian mm. intertransversarii in the cervical region consist of muscles of the both m. longissimus and m. iliocostalis groups despite having been treated as a single series of muscles, and thus are not homologous with muscles of the same name in Lepidosauria or Crocodylia. The avian mm. inclusi that lie medial to mm. intertransversarii are homologous with mm. intercostales externi in Lepidosauria and mm. intercostales externi and m. scalenus combined in Crocodylia. Innervation patterns suggest that a muscle (“m. iliocostalis capitis”) connecting the atlas rib and occiput in Crocodylia includes contributions from the subvertebral layer and m. cucullaris complex, and possibly m. iliocostalis as well. The present findings may serve as a basis for revising the currently used avian nomenclature so that it will reflect homologies of muscles with their non‐avian counterparts. J. Morphol., 2007. © 2007 Wiley‐Liss, Inc.


Competition and constraint drove Cope's rule in the evolution of giant flying reptiles

Abstract: The pterosaurs, Mesozoic flying reptiles, attained wingspans of more than 10 m that greatly exceed the largest birds and challenge our understanding of size limits in flying animals. Pterosaurs have been used to illustrate Cope’s rule, the influential generalization that evolutionary lineages trend to increasingly large body sizes. However, unambiguous examples of Cope’s rule operating on extended timescales in large clades remain elusive, and the phylogenetic pattern and possible drivers of pterosaur gigantism are uncertain. Here we show 70 million years of highly constrained early evolution, followed by almost 80 million years of sustained, multi-lineage body size increases in pterosaurs. These results are supported by maximum-likelihood modelling of a comprehensive new pterosaur data set. The transition between these macroevolutionary regimes is coincident with the Early Cretaceous adaptive radiation of birds, supporting controversial hypotheses of bird–pterosaur competition, and suggesting that evolutionary competition can act as a macroevolutionary driver on extended geological timescales.

wingspan
body size
competition
late triassic
small pterosaur


Identifying fossil Myrtaceae leaves: the first described fossils of Syzygium from Australia.

Abstract:
PREMISE OF THE STUDY
Although leaves of Myrtaceae are easily identified to family level, very few studies have convincingly identified fossil Myrtaceae leaves to living genera. We used a broadly comparative approach with a large data set of extant taxa to confidently assign the mummified remains of myrtaceous leaves from early Miocene sediments at Kiandra (New South Wales, Australia) to a living genus.

METHODS
Fossils were identified using a nearest living relative approach, against a database of 232 extant broadleaf rainforest species of Myrtaceae. Leaf cuticles were prepared from 106 species, sourced from herbarium specimens as well as some living individuals, and a further 127 records were assembled from the literature. A set of simple but phylogenetically informative cuticular characters were observed, described, and recorded under both scanning electron microscopy and standard light microscopy.

KEY RESULTS
A new fossil species of Syzygium Gaertn. is described from mummified remains found in early Miocene (21.5-21.7 Ma) sediments. The fossil taxon is here named Syzygium christophelii sp. nov., in honor of the late Australian paleobotanist David Christophel.

CONCLUSIONS
These fossils represent some of the most confidently described Myrtaceae leaf fossils published to date and are the first and oldest described fossil record of Syzygium from Australia. While several fossil parataxa have been illustrated from New Zealand, and several fossil species of Syzygium have previously been proposed in the literature, many of these fossils lack characters for a confident diagnosis.


The transition between Carcharocles chubutensis and Carcharocles megalodon (Otodontidae, Chondrichthyes): lateral cusplet loss through time

Abstract: ABSTRACT The teeth of two megatooth macro-predatory shark species (Carcharocles chubutensis and Carcharocles megalodon; Otodontidae, Chondrichthyes) occur within the Miocene Chesapeake Group of Maryland, U.S.A. Definitive separation between all the teeth of Carcharocles chubutensis and Carcharocles megalodon is impossible because a complex mosaic evolutionary continuum characterizes this transformation, particularly in the loss of lateral cusplets. The cuspleted and uncuspleted teeth of Carcharocles spp. are designated as chronomorphs because there is wide overlap between them both morphologically and chronologically. In the lower Miocene Beds (Shattuck Zones) 2–9 of the Calvert Formation (representing approximately 3.2 million years, 20.2–17 Ma, Burdigalian) both cuspleted and uncuspleted teeth are present, but cuspleted teeth predominate, constituting approximately 87% of the Carcharocles spp. teeth represented in our sample. However, in the middle Miocene Beds 10–16A of the Calvert Formation (representing approximately 2.4 million years, 16.4–14 Ma, Langhian), there is a steady increase in the proportion of uncuspleted Carcharocles teeth. In the upper Miocene Beds 21–24 of the St. Marys Formation (representing approximately 2.8 million years, 10.4–7.6 Ma, Tortonian), lateral cusplets are nearly absent in Carcharocles teeth from our study area, with only a single specimen bearing lateral cusplets. The dental transition between Carcharocles chubutensis and Carcharocles megalodon occurs within the Miocene Chesapeake Group. Although this study helps to elucidate the timing of lateral cusplet loss in Carcharocles locally, the rationale for this prolonged evolutionary transition remains unclear.

carcharocles megalodon
otodontidae
chondrichthyes
shark


Belgian substages as a basis for an international chronostratigraphic division of the Tournaisian and Viséan

Abstract: Abstract The Tournaisian and Viséan were formerly considered as series and in Belgium were divided into two (Hastarian and Ivorian) and three stages (Moliniacian, Livian and Warnantian), which are now considered as substages. The Belgian substages are based on conodonts and foraminifers, and incidentally on rugose corals, and are described here. Their boundaries, biostratigraphy and sequence stratigraphy are well detailed and clearly defined. The base of the Hastarian (lower Tournaisian) corresponds to the base of the Tournaisian (base of Carboniferous); the base of the Ivorian (upper Tournaisian) corresponds to the appearance of the conodont Polygnathus communis carina, a little above the last Siphonodella; the base of the Moliniacian (lower Viséan) corresponds to the base of the Viséan stage defined by the first occurrence of the foraminifer Eoparastaffella simplex; the Livian (middle Viséan) corresponds to the foraminiferal MFZ12 Zone and is marked by the appearance of Koskinotextularia and Pojarkovella nibelis; the base of the Warnantian (upper Viséan) is marked by the appearance of Neoarchaediscus, Vissariotaxis, Planospirodiscus, and Palaeotextularia with a bilaminar wall, the index taxa of the MFZ13-Neoarchaediscus Zone. The up-to-date chronostratigraphic subdivision of the Tournaisian and Viséan is not limited to Belgium and the surrounding areas. It can be applied through Eurasia as far as South China. The Belgian units could therefore be the basis for a future international division of the Tournaisian into two parts (Hastarian and Ivorian) and of the Viséan into three parts (Moliniacian, Livian and Warnantian), corresponding to time intervals of c. 5–8 Ma.


Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates

Abstract: The fossil record is our only direct means for evaluating shifts in biodiversity through Earth's history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic–Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent ‘Lagerstätten effects’. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous–Palaeogene extinction.

marine tetrapod diversity
geological megabiase
sampling
mesozoic


Correlation between investment in sexual traits and valve sexual dimorphism in Cyprideis species (Ostracoda)

Abstract: Assessing the long-term macroevolutionary consequences of sexual selection has been hampered by the difficulty of studying this process in the fossil record. Cytheroid ostracodes offer an excellent system to explore sexual selection in the fossil record because their readily fossilized carapaces are sexually dimorphic. Specifically, males are relatively more elongate than females in this superfamily. This sexual shape difference is thought to arise so that males carapaces can accommodate their very large copulatory apparatus, which can account for up to one-third of body volume. Here we test this widely held explanation for sexual dimorphism in cytheroid ostracodes by correlating investment in male genitalia, a trait in which sexual selection is seen as the main evolutionary driver, with sexual dimorphism of carapace in the genus Cyprideis. We analyzed specimens collected in the field (C. salebrosa, USA; C. torosa, UK) and from collections of the National Museum of Natural History, Washington, DC (C. mexicana). We digitized valve outlines in lateral view to obtain measures of size (valve area) and shape (elongation, measured as length to height ratio), and obtained several dimensions from two components of the hemipenis: the muscular basal capsule, which functions as a sperm pump, and the section that includes the intromittent organ (terminal extension). In addition to the assessment of this primary sexual trait, we also quantified two dimensions of the male secondary sexual trait—where the transformed right walking leg functions as a clasping organ during mating. We also measured linear dimensions from four limbs as indicators of overall (soft-part) body size, and assessed allometry of the soft anatomy. We observed significant correlations in males between valve size, but not elongation, and distinct structural parts of the hemipenis, even after accounting for their shared correlation with overall body size. We also found weak but significant positive correlation between valve elongation and the degree of sexual dimorphism of the walking leg, but only in C. torosa. The correlation between the hemipenis parts, especially basal capsule size and male valve size dimorphism suggests that sexual selection on sperm size, quantity, and/or efficiency of transfer may drive sexual size dimorphism in these species, although we cannot exclude other aspects of sexual and natural selection.

investment
sexual trait
dimorphism
indicator


Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks

Abstract: The biotic recovery from Earth’s most severe extinction event at the Permian-Triassic boundary largely reestablished the preextinction structure of marine trophic networks, with marine reptiles assuming the predator roles. However, the highest trophic level of today's marine ecosystems, i.e., macropredatory tetrapods that forage on prey of similar size to their own, was thus far lacking in the Paleozoic and early Mesozoic. Here we report a top-tier tetrapod predator, a very large (>8.6 m) ichthyosaur from the early Middle Triassic (244 Ma), of Nevada. This ichthyosaur had a massive skull and large labiolingually flattened teeth with two cutting edges indicative of a macropredatory feeding style. Its presence documents the rapid evolution of modern marine ecosystems in the Triassic where the same level of complexity as observed in today’s marine ecosystems is reached within 8 My after the Permian-Triassic mass extinction and within 4 My of the time reptiles first invaded the sea. This find also indicates that the biotic recovery in the marine realm may have occurred faster compared with terrestrial ecosystems, where the first apex predators may not have evolved before the Carnian.

ichthyosaur
middle triassic
trophic network
biotic recovery
mass extinction


Stratification of surface waters during the last glacial millennial climatic events: a key factor in subsurface and deep-water mass dynamics

Abstract: Abstract. The last glacial period was punctuated by abrupt climatic events with extrema known as Heinrich and Dansgaard–Oeschger events. These millennial events have been the subject of many paleoreconstructions and model experiments in the past decades, but yet the hydrological processes involved remain elusive. In the present work, high-resolution analyses were conducted on the 12–42 ka BP section of core MD99-2281 retrieved southwest of the Faeroe Islands, and combined with analyses conducted in two previous studies (Zumaque et al., 2012; Caulle et al., 2013). Such a multiproxy approach, coupling micropaleontological, geochemical and sedimentological analyses, allows us to track surface, subsurface, and deep hydrological processes occurring during these rapid climatic changes. Records indicate that the coldest episodes of the studied period (Greenland stadials and Heinrich stadials) were characterized by a strong stratification of surface waters. This surface stratification seems to have played a key role in the dynamics of subsurface and deep-water masses. Indeed, periods of high surface stratification are marked by a coupling of subsurface and deep circulations which sharply weaken at the beginning of stadials, while surface conditions progressively deteriorate throughout these cold episodes; conversely, periods of decreasing surface stratification (Greenland interstadials) are characterized by a coupling of surface and deep hydrological processes, with progressively milder surface conditions and gradual intensification of the deep circulation, while the vigor of the subsurface northward Atlantic flow remains constantly high. Our results also reveal different and atypical hydrological signatures during Heinrich stadials (HSs): while HS1 and HS4 exhibit a "usual" scheme with reduced overturning circulation, a relatively active North Atlantic circulation seems to have prevailed during HS2, and HS3 seems to have experienced a re-intensification of this circulation during the middle of the event. Our findings thus bring valuable information to better understand hydrological processes occurring in a key area during the abrupt climatic shifts of the last glacial period.

surface water
climatic event
stratification


Warming, euxinia and sea level rise during the Paleocene–Eocene Thermal Maximum on the Gulf Coastal Plain: implications for ocean oxygenation and nutrient cycling

Abstract: Abstract. The Paleocene–Eocene Thermal Maximum (PETM, ~ 56 Ma) was a ~ 200 kyr episode of global warming, associated with massive injections of 13C-depleted carbon into the ocean–atmosphere system. Although climate change during the PETM is relatively well constrained, effects on marine oxygen concentrations and nutrient cycling remain largely unclear. We identify the PETM in a sediment core from the US margin of the Gulf of Mexico. Biomarker-based paleotemperature proxies (methylation of branched tetraether–cyclization of branched tetraether (MBT–CBT) and TEX86) indicate that continental air and sea surface temperatures warmed from 27–29 to ~ 35 °C, although variations in the relative abundances of terrestrial and marine biomarkers may have influenced these estimates. Vegetation changes, as recorded from pollen assemblages, support this warming. The PETM is bracketed by two unconformities. It overlies Paleocene silt- and mudstones and is rich in angular (thus in situ produced; autochthonous) glauconite grains, which indicate sedimentary condensation. A drop in the relative abundance of terrestrial organic matter and changes in the dinoflagellate cyst assemblages suggest that rising sea level shifted the deposition of terrigenous material landward. This is consistent with previous findings of eustatic sea level rise during the PETM. Regionally, the attribution of the glauconite-rich unit to the PETM implicates the dating of a primate fossil, argued to represent the oldest North American specimen on record. The biomarker isorenieratene within the PETM indicates that euxinic photic zone conditions developed, likely seasonally, along the Gulf Coastal Plain. A global data compilation indicates that O2 concentrations dropped in all ocean basins in response to warming, hydrological change, and carbon cycle feedbacks. This culminated in (seasonal) anoxia along many continental margins, analogous to modern trends. Seafloor deoxygenation and widespread (seasonal) anoxia likely caused phosphorus regeneration from suboxic and anoxic sediments. We argue that this fueled shelf eutrophication, as widely recorded from microfossil studies, increasing organic carbon burial along many continental margins as a negative feedback to carbon input and global warming. If properly quantified with future work, the PETM offers the opportunity to assess the biogeochemical effects of enhanced phosphorus regeneration, as well as the timescales on which this feedback operates in view of modern and future ocean deoxygenation.

sea level rise
paleocene–eocene thermal maximum
gulf coastal plain
ocean deoxygenation


Dynamic and synchronous changes in metazoan body size during the Cambrian Explosion

Abstract: Many aspects of the drivers for, and evolutionary dynamics of, the Cambrian Explosion are poorly understood. Here we quantify high-resolution changes in species body size in major metazoan groups on the Siberian Platform during the early Cambrian (ca. 540–510 Million years ago (Ma)). Archaeocyath sponges, hyolith lophophorates, and helcionelloid mollusc species show dynamic and synchronous trends over million-year timescales, with peaks in body size during the latest Tommotian/early Atbadanian and late Atdabanian/early Botoman, and notably small body sizes in the middle Atdabanian and after the Sinsk anoxic extinction event, starting ca. 513 Ma. These intervals of body size changes are also mirrored in individual species and correlate positively with increased rates of origination and broadly with total species diversity. Calcitic brachiopods (rhynchonelliformeans), however, show a general increase in body size following the increase in species diversity through this interval: phosphatic brachiopods (linguliformeans) show a body size decrease that negatively correlates with diversity. Both brachiopod groups show a rapid recovery at the Sinsk Event. The synchronous changes in these metrics in archaeocyath, hyoliths and helcionelloids suggest the operation of external drivers through the early Cambrian, such as episodic changes in oxygenation or productivity. But the trends shown by brachiopods suggests a differing physiological response. Together, these dynamics created both the distinct evolutionary record of metazoan groups during the Cambrian Explosion and determined the nature of its termination.

synchronous change
cambrian explosion
archaeocyath


Late Mesoproterozoic – early Neoproterozoic organic‐walled microfossils from the Madhubani Group of the Ganga Valley, northern India

Abstract: The age of the sedimentary basement of the Ganga Valley in northern India, which is represented by the entirely subsurface Ganga Supergroup, is key for addressing issues related to the tectonic history of the Himalaya. However, the stratigraphic correlations between the Ganga Supergroup in the Ganga Valley, the Vindhyan Supergroup in cratonic India to the south, and Proterozoic successions in the Lesser Himalaya to the north have long been a matter of controversy. This is largely because of the poor age constraint of the Madhubani Group of the upper Ganga Supergroup, which has been variously interpreted as Proterozoic, lower Palaeozoic, or even Mesozoic. To address this issue, we used a low manipulation maceration technique to extract organic‐walled microfossils from the Ujhani and Tilhar formations of the lower Madhubani Group. Our study recovered a total of 24 taxa, including Devisphaera corallis gen. et sp. nov. The co‐occurrence of Trachyhystrichosphaera aimika, Caudosphaera expansa and Annulusia annulata in the lower Madhubani Group indicates a late Mesoproterozoic to early Neoproterozoic age. Thus, the biostratigraphical data suggest a >300 myr depositional gap between the Madhubani Group and the immediately underlying Bahraich Group, which has been independently constrained to be upper Palaeoproterozoic to lower Mesoproterozoic in age. Therefore, the first‐order stratigraphic architecture, with a Palaeoproterozoic–Mesoproterozoic succession unconformably overlain by a Mesoproterozoic–Neoproterozoic succession, is closely similar throughout the Vindhyan Basin, Ganga Valley and Lesser Himalaya, suggesting a shared sedimentary and tectonic history among them.

mesoproterozoic
organic‐walled microfossil
microfossil


First record of non-mineralized cephalopod jaws and arm hooks from the latest Cretaceous of Eurytania, Greece

Abstract: Due to the lower fossilization potential of chitin, non-mineralized cephalopod jaws and arm hooks are much more rarely preserved as fossils than the calcitic lower jaws of ammonites or the calcitized jaw apparatuses of nautilids. Here, we report such non-mineralized fossil jaws and arm hooks from pelagic marly limestones of continental Greece. Two of the specimens lie on the same slab and are assigned to the Ammonitina; they represent upper jaws of the aptychus type, which is corroborated by finds of aptychi. Additionally, one intermediate type and one anaptychus type are documented here. The morphology of all ammonite jaws suggest a desmoceratoid affinity. The other jaws are identified as coleoid jaws. They share the overall U-shape and proportions of the outer and inner lamellae with Jurassic lower jaws of Trachyteuthis (Teudopseina). We also document the first belemnoid arm hooks from the Tethyan Maastrichtian. The fossils described here document the presence of a typical Mesozoic cephalopod assemblage until the end of the Cretaceous in the eastern Tethys.

non-mineralized cephalopod jaw
arm hook
greece


Protracted growth impedes the detection of sexual dimorphism in non‐avian dinosaurs

Abstract: Evidence for sexual dimorphism is extremely limited in the non‐avian dinosaurs despite their high diversity and disparity, and despite the fact that dimorphism is very common in vertebrate lineages of all kinds. Using body‐size data from both Alligator mississippiensis and Rhea americana, which phylogenetically bracket the dinosaurs, we demonstrate that even when there is strong dimorphism in a species, random sampling of populations of individuals characterized by sustained periods of growth (as in the alligator and most dinosaurs) can result in the loss of this signal. Dimorphism may be common in fossil taxa but very hard to detect without ontogenetic age control and large sample sizes, both of which are hampered by the limitations of the fossil record. Signal detection may be further hindered by Type III survivorship, whereby increased mortality among the young favours the likelihood that they will be sampled (unless predation or taphonomic bias against small size acts against this). These, and other considerations relating to behaviour and ecology, provide powerful reasons to suggest that sexual dimorphism in dinosaurs may be very difficult to detect in almost all currently available samples. Similar issues are likely also to be applicable to many fossil reptiles, or animals more generally.

detection
sexual dimorphism
non‐avian dinosaur
fossil taxa
large sample size


Eggshell Porosity Provides Insight on Evolution of Nesting in Dinosaurs

Abstract: Knowledge about the types of nests built by dinosaurs can provide insight into the evolution of nesting and reproductive behaviors among archosaurs. However, the low preservation potential of their nesting materials and nesting structures means that most information can only be gleaned indirectly through comparison with extant archosaurs. Two general nest types are recognized among living archosaurs: 1) covered nests, in which eggs are incubated while fully covered by nesting material (as in crocodylians and megapodes), and 2) open nests, in which eggs are exposed in the nest and brooded (as in most birds). Previously, dinosaur nest types had been inferred by estimating the water vapor conductance (i.e., diffusive capacity) of their eggs, based on the premise that high conductance corresponds to covered nests and low conductance to open nests. However, a lack of statistical rigor and inconsistencies in this method render its application problematic and its validity questionable. As an alternative we propose a statistically rigorous approach to infer nest type based on large datasets of eggshell porosity and egg mass compiled for over 120 extant archosaur species and 29 archosaur extinct taxa/ootaxa. The presence of a strong correlation between eggshell porosity and nest type among extant archosaurs indicates that eggshell porosity can be used as a proxy for nest type, and thus discriminant analyses can help predict nest type in extinct taxa. Our results suggest that: 1) covered nests are likely the primitive condition for dinosaurs (and probably archosaurs), and 2) open nests first evolved among non-avian theropods more derived than Lourinhanosaurus and were likely widespread in non-avian maniraptorans, well before the appearance of birds. Although taphonomic evidence suggests that basal open nesters (i.e., oviraptorosaurs and troodontids) were potentially the first dinosaurs to brood their clutches, they still partially buried their eggs in sediment. Open nests with fully exposed eggs only became widespread among Euornithes. A potential co-evolution of open nests and brooding behavior among maniraptorans may have freed theropods from the ground-based restrictions inherent to covered nests and allowed the exploitation of alternate nesting locations. These changes in nesting styles and behaviors thus may have played a role in the evolutionary success of maniraptorans (including birds).

nesting
behavior
egg
crocodylian
eggshell porosity


The dentition of Carodnia vieirai (Mammalia: Xenungulata): enamel microstructure and mastication pattern

Abstract:
Post canine teeth of the Paleogene Carodnia from Brazil were investigated in regard to the enamel microstructure and the mastication pattern. The enamel is entirely prismatic, and the interprismatic matrix (IPM) is oriented parallel to the prisms, which consist of Boyde's patterns 1 and 2. The prominent vertical Hunter-Schreger-bands (HSB) are restricted to an outer zone, whereas the inner zone is formed by transverse but irregular HSB. Such a combination is very rare in mammalian teeth, but shows some similarities to the schmelzmuster of specific perissodactyls, where it evolved independently. Significant differences to Astrapotheria were found. The mastication pattern can be derived from the orientation of the wear facets on the transverse lophs. In stages of moderate tooth wear, the lophs have one-sided planar facets that are inclined in the upper and lower teeth antagonistically. Enamel crests on the leading sides cut like scissors. Subsequently, the antagonistic shear-cutting crests of the lophs pass each other and compress the food items until central occlusion. Thus, the jaw movement during phase I is predominantly mesial with a distinct inclination upwards.
No traces indicate a phase II of the power stroke. Neither the enamel nor the widelydistributed mastication pattern offer convincing arguments to support any of proposed phylogenetic relationship of Carodnia. The rare schmelzmuster contributes to the diversity of enamel microstructures in South American ungulates.

enamel microstructure
mastication pattern
phylogenetic relationship


THE HEAD AND NECK MUSCLES ASSOCIATED WITH FEEDING IN SPHENODON (REPTILIA: LEPIDOSAURIA: RHYNCHOCEPHALIA)

Abstract:
Feeding in Sphenodon, the tuatara of New Zealand, is of interest for several reasons. First, the modern animal is threatened by extinction, and some populations are in competition for food with Pacific rats. Second, Sphenodon demonstrates a feeding apparatus that is unique to living amniotes: an enlarged palatine tooth row, acrodont dentition, enlarged incisor-like teeth on the premaxilla, a posterior extension of the dentary and an elongate articular surtace that permits prooral shearing. Third, Sphenodon has a skull with two complete lateral temporal bars and is therefore structurally analogous to the configuration hypothesised for the ancestral diapsid reptile. Furthermore, the fossil relatives of Sphenodon demonstrate considerable variation in terms of feeding apparatus and skull shape. Lastly, as Sphenodon is the only extant rhynchocephalian it represents a potentially useful reference taxon for both muscle reconstruction in extinct reptile taxa and determination of muscle homology in extant taxa.
Here we provide an up-to-date consensus view of osteology and musculature in Sphenodon that is relevant to feeding. Discrepancies within previous descriptions are evaluated and synthesised with new observations. This paper displays the complex muscle arrangement using a range of different imaging techniques and a variety of different angles. This includes photographs, illustrations, schematic diagrams, and microcomputed tomography (micro-CT) slice images.

sphenodon
rhynchocephalian
muscle arrangement


Differential influences of allometry, phylogeny and environment on the rostral shape diversity of extinct South American notoungulates

Abstract: Understanding the mechanisms responsible for phenotypic diversification, and the associated underlying constraints and ecological factors represents a central issue in evolutionary biology. Mammals present a wide variety of sizes and shapes, and are characterized by a high number of morphological convergences that are hypothesized to reflect similar environmental pressures. Extinct South American notoungulates evolved in isolation from northern mammalian faunas in highly disparate environments. They present a wide array of skeletal phenotypes and convergences, such as ever-growing dentition. Here, we focused on the origins of the rostral diversity of notoungulates by quantifying the shape of 26 genera using three-dimensional geometric morphometric analysis. We tested the influence of allometry and phylogeny on rostral shape and evaluated rates of evolutionary change in the different clades. We found strong allometric and phylogenetic signals concerning the rostral shape of notoungulates. Despite convergent forms, we observed a diffuse diversification of rostral shape, with no significant evidence of influence by large-scaled environmental variation. This contrasts with the increase in dental crown height that occurred in four late-diverging families in response to similar environmental pressures. These results illustrate the importance of considering both biological components and evolutionary rates to better understand some aspects of phenotypic diversity.


Finding a home for hyoliths

Abstract: Manyof the animal lineages that aroseduring theCambrian evolutionary radiation, 540 million years ago, are difficult to relate to living taxa. Constraining the phylogenetic position of these highly disparate taxa allows their idiosyncratic morphologies to illuminate the stepwise establishment of modern body plans [1–3]. Hyoliths are one such lineage; their operculate conical shells are common Palaeozoic fossils. Hyoliths have recently been identified as brachiopods based on the description of an attachment stalk [4] and a tentaculate feeding apparatus [5]. Liu et al. [6] present an alternative view, interpreting the putative pedicle as a damaged shell apex, and questioning whether the feeding apparatus ought to be termed a lophophore.

hyolith
phylogenetic position
brachiopod


Metabolic dominance of bivalves predates brachiopod diversity decline by more than 150 million years

Abstract: Brachiopods and bivalves feed in similar ways and have occupied the same environments through geological time, but brachiopods were far more diverse and abundant in the Palaeozoic whereas bivalves dominate the post-Palaeozoic, suggesting a transition in ecological dominance 250 Ma. However, diversity and abundance data alone may not adequately describe key changes in ecosystem function, such as metabolic activity. Here, we use newly compiled body size data for 6066 genera of bivalves and brachiopods to calculate metabolic rates and revisit this question from the perspective of energy use, finding that bivalves already accounted for a larger share of metabolic activity in Palaeozoic oceans. We also find that the metabolic activity of bivalves has increased by more than two orders of magnitude over this interval, whereas brachiopod metabolic activity has declined by more than 50%. Consequently, the increase in bivalve energy metabolism must have occurred via the acquisition of new food resources rather than through the displacement of brachiopods. The canonical view of a mid-Phanerozoic transition from brachiopod to bivalve dominance results from a focus on taxonomic diversity and numerical abundance as measures of ecological importance. From a metabolic perspective, the oceans have always belonged to the clams.

dominance
bivalve
brachiopod
metabolic rate


Bayesian Analysis Using a Simple Likelihood Model Outperforms Parsimony for Estimation of Phylogeny from Discrete Morphological Data

Abstract: Despite the introduction of likelihood-based methods for estimating phylogenetic trees from phenotypic data, parsimony remains the most widely-used optimality criterion for building trees from discrete morphological data. However, it has been known for decades that there are regions of solution space in which parsimony is a poor estimator of tree topology. Numerous software implementations of likelihood-based models for the estimation of phylogeny from discrete morphological data exist, especially for the Mk model of discrete character evolution. Here we explore the efficacy of Bayesian estimation of phylogeny, using the Mk model, under conditions that are commonly encountered in paleontological studies. Using simulated data, we describe the relative performances of parsimony and the Mk model under a range of realistic conditions that include common scenarios of missing data and rate heterogeneity.


A NEW PTEROSAUR FROM THE LIAONING PROVINCE OF CHINA, THE PHYLOGENY OF THE PTERODACTYLOIDEA, AND CONVERGENCE IN THEIR CERVICAL VERTEBRAE

Abstract: Abstract:  The largest known flying organisms are the azhdarchid pterosaurs, a pterodactyloid clade previously diagnosed by the characters of their extremely elongate middle‐series cervical vertebrae. The named species of the Azhdarchidae are from the Late Cretaceous. However, isolated mid‐cervical vertebrae with similar dimensions and characters have been referred to this group that date back to the Late Jurassic, implying an almost 60 million year gap in the fossil record of this group and an unrecorded radiation in the Jurassic of all the major clades of the Pterodactyloidea. A new pterosaur from the Early Cretaceous of Liaoning Province of China, Elanodactylus prolatus gen. et sp. nov., is described with mid‐cervical vertebrae that bear these azhdarchid characters but has other postcranial material that are distinct from the members of this group. Phylogenetic analysis of the new species and the Pterodactyloidea places it with the Late Jurassic vertebrae in the Late Jurassic–Early Cretaceous Ctenochasmatidae and reveals that the characters of the elongate azhdarchid vertebrae appeared independently in both groups. These results are realized though the large taxon sampling in the analysis demonstrating that the homoplastic character states present in these two taxa were acquired in a different order in their respective lineages. Some of these homoplastic characters were previously thought to appear once in the history of pterosaurs and may be correlated to the extension of the neck regions in both groups. Because the homoplastic character states in the Azhdarchidae and Ctenochasmatidae are limited to the mid‐cervical vertebrae, these states are termed convergent based on a definition of the term in a phylogenetic context. A number of novel results from the analysis presented produce a reorganization in the different species and taxa of the Pterodactyloidea.

pterosaur
china
pterodactyloidea
phylogenetic analysis
ctenochasmatidae


Exceptionally well-preserved early Eocene fossil reveals cranial and vertebral features of a stem group roller (Aves: Coraciiformes)

Abstract: Three-dimensionally preserved skulls of small Paleogene land birds are very rare. Here, we describe a cranium and associated partial postcranial remains of an early Eocene stem group roller (Aves: Coraciiformes) from the London Clay of the Isle of Sheppey (England). The fossil shows features of the skull and vertebral column in great detail. It is distinguished from extant Coraciidae and Brachypteraciidae in several presumably plesiomorphic characteristics, which are likely to reflect differences in diet and/or foraging strategy between Eocene and extant rollers. Preserved stomach contents in other early Eocene fossils indicate that fruits were a regular part of the diet of stem group rollers. The extant Coraciidae and Brachypteraciidae, by contrast, almost exclusively feed on larger-sized invertebrates and small vertebrates, which are usually dispatched by beating before being swallowed. Stronger biting forces as well as the characteristic prey manipulation behavior of extant rollers may account for some of the observed differences in the cranial and vertebral morphology of the fossil and extant taxa, but the exact functional correlations remain elusive. We furthermore identify a previously undescribed cranial feature of rollers: a very large foramen for the ramus occipitalis of the arteria ophthalmica externa, which is of unknown functional significance and constitutes a potentially promising research target for future studies.

eocene fossil
stem group roller
coraciiformes
skull
london clay


Late Quaternary paleoenvironmental changes in East Africa: a review of multiproxy evidence from palynology, lake sediments, and associated records

Abstract: This paper presents an overview of paleoenvironmental changes in East Africa during the late Quaternary based on evidence from pollen, diatoms, microscopic charcoal, and lake level records and associated proxies. The paleoenvironmental records derived from different proxies complement each other to provide a more accurate and complete assessment of the paleoenvironmental changes in East Africa. The records show that the period prior to c. 42,000 14C yr BP was characterized by warm climatic conditions similar to the present. This was followed by a change to cold dry conditions from 42,000 to 30,000 14C yr BP, and cold and moist conditions from 30,000 to 21,000 14C yr BP. Temperatures during the latter period leading to the Last Glacial Maximum (LGM) were probably 2 to 4.1°C lower than the present. Between c. 21,000 and 12,500 14C yr BP East Africa's environment was generally cool, punctuated by two significant episodes of prolonged desiccation. Warm and moist conditions punctuated by rapid climatic changes prevailed in the region during the deglacial and middle Holocene period. Ice core records document two significant and abrupt drought events in the region, one at 8300 14C yr BP and the other at 5200 14C yr BP. The onset of a longer and more extensive desiccation period commencing 4000 14C yr BP was registered in nearly all sites. The climate of East Africa was generally drier than present during the Medieval Warm Period (MWP) while fairly wet conditions prevailed during the Little Ice Age (LIA) interrupted by three episodes of aridity, more severe than those of more recent times. Whereas this review advances our understanding of climate and vegetational changes in East Africa beyond the Last Glacial Maximum, it also highlights limitations of the paradigms that explain the forcing mechanisms behind the changes. However, unequivocal interpretation of the multiproxy data from East Africa with respect to paleoenvironmental changes becomes extremely complex and challenging especially when the anthropogenic input is considered.

paleoenvironmental change
east africa
review


A Pliocene‐Pleistocene stack of 57 globally distributed benthic δ18O records

Abstract: [1] We present a 5.3-Myr stack (the “LR04” stack) of benthic δ18O records from 57 globally distributed sites aligned by an automated graphic correlation algorithm. This is the first benthic δ18O stack composed of more than three records to extend beyond 850 ka, and we use its improved signal quality to identify 24 new marine isotope stages in the early Pliocene. We also present a new LR04 age model for the Pliocene-Pleistocene derived from tuning the δ18O stack to a simple ice model based on 21 June insolation at 65°N. Stacked sedimentation rates provide additional age model constraints to prevent overtuning. Despite a conservative tuning strategy, the LR04 benthic stack exhibits significant coherency with insolation in the obliquity band throughout the entire 5.3 Myr and in the precession band for more than half of the record. The LR04 stack contains significantly more variance in benthic δ18O than previously published stacks of the late Pleistocene as the result of higher-resolution records, a better alignment technique, and a greater percentage of records from the Atlantic. Finally, the relative phases of the stack's 41- and 23-kyr components suggest that the precession component of δ18O from 2.7–1.6 Ma is primarily a deep-water temperature signal and that the phase of δ18O precession response changed suddenly at 1.6 Ma.


A Total-Evidence Approach to Dating with Fossils, Applied to the Early Radiation of the Hymenoptera

Abstract: Abstract Phylogenies are usually dated by calibrating interior nodes against the fossil record. This relies on indirect methods that, in the worst case, misrepresent the fossil information. Here, we contrast such node dating with an approach that includes fossils along with the extant taxa in a Bayesian total-evidence analysis. As a test case, we focus on the early radiation of the Hymenoptera, mostly documented by poorly preserved impression fossils that are difficult to place phylogenetically. Specifically, we compare node dating using nine calibration points derived from the fossil record with total-evidence dating based on 343 morphological characters scored for 45 fossil (4--20 complete) and 68 extant taxa. In both cases we use molecular data from seven markers (∼5 kb) for the extant taxa. Because it is difficult to model speciation, extinction, sampling, and fossil preservation realistically, we develop a simple uniform prior for clock trees with fossils, and we use relaxed clock models to accommodate rate variation across the tree. Despite considerable uncertainty in the placement of most fossils, we find that they contribute significantly to the estimation of divergence times in the total-evidence analysis. In particular, the posterior distributions on divergence times are less sensitive to prior assumptions and tend to be more precise than in node dating. The total-evidence analysis also shows that four of the seven Hymenoptera calibration points used in node dating are likely to be based on erroneous or doubtful assumptions about the fossil placement. With respect to the early radiation of Hymenoptera, our results suggest that the crown group dates back to the Carboniferous, ∼309 Ma (95% interval: 291--347 Ma), and diversified into major extant lineages much earlier than previously thought, well before the Triassic. [Bayesian inference; fossil dating; morphological evolution; relaxed clock; statistical phylogenetics.]

dating
early radiation
hymenoptera
morphological character
phylogenetic analysis


Early evolution of limb regeneration in tetrapods: evidence from a 300-million-year-old amphibian

Abstract: Salamanders are the only tetrapods capable of fully regenerating their limbs throughout their entire lives. Much data on the underlying molecular mechanisms of limb regeneration have been gathered in recent years allowing for new comparative studies between salamanders and other tetrapods that lack this unique regenerative potential. By contrast, the evolution of animal regeneration just recently shifted back into focus, despite being highly relevant for research designs aiming to unravel the factors allowing for limb regeneration. We show that the 300-million-year-old temnospondyl amphibian Micromelerpeton, a distant relative of modern amphibians, was already capable of regenerating its limbs. A number of exceptionally well-preserved specimens from fossil deposits show a unique pattern and combination of abnormalities in their limbs that is distinctive of irregular regenerative activity in modern salamanders and does not occur as variants of normal limb development. This demonstrates that the capacity to regenerate limbs is not a derived feature of modern salamanders, but may be an ancient feature of non-amniote tetrapods and possibly even shared by all bony fish. The finding provides a new framework for understanding the evolution of regenerative capacity of paired appendages in vertebrates in the search for conserved versus derived molecular mechanisms of limb regeneration.

limb regeneration
tetrapod
amphibian
micromelerpeton


Evolution of High Tooth Replacement Rates in Sauropod Dinosaurs

Abstract: Background Tooth replacement rate can be calculated in extinct animals by counting incremental lines of deposition in tooth dentin. Calculating this rate in several taxa allows for the study of the evolution of tooth replacement rate. Sauropod dinosaurs, the largest terrestrial animals that ever evolved, exhibited a diversity of tooth sizes and shapes, but little is known about their tooth replacement rates. Methodology/Principal Findings We present tooth replacement rate, formation time, crown volume, total dentition volume, and enamel thickness for two coexisting but distantly related and morphologically disparate sauropod dinosaurs Camarasaurus and Diplodocus. Individual tooth formation time was determined by counting daily incremental lines in dentin. Tooth replacement rate is calculated as the difference between the number of days recorded in successive replacement teeth. Each tooth family in Camarasaurus has a maximum of three replacement teeth, whereas each Diplodocus tooth family has up to five. Tooth formation times are about 1.7 times longer in Camarasaurus than in Diplodocus (315 vs. 185 days). Average tooth replacement rate in Camarasaurus is about one tooth every 62 days versus about one tooth every 35 days in Diplodocus. Despite slower tooth replacement rates in Camarasaurus, the volumetric rate of Camarasaurus tooth replacement is 10 times faster than in Diplodocus because of its substantially greater tooth volumes. A novel method to estimate replacement rate was developed and applied to several other sauropodomorphs that we were not able to thin section. Conclusions/Significance Differences in tooth replacement rate among sauropodomorphs likely reflect disparate feeding strategies and/or food choices, which would have facilitated the coexistence of these gigantic herbivores in one ecosystem. Early neosauropods are characterized by high tooth replacement rates (despite their large tooth size), and derived titanosaurs and diplodocoids independently evolved the highest known tooth replacement rates among archosaurs.

tooth
replacement rate
sauropod dinosaur


Archaeological evidence for two separate dispersals of Neanderthals into southern Siberia

Abstract: Significance Neanderthals once inhabited Europe and western Asia, spreading as far east as the Altai Mountains in southern Siberia, but the geographical origin and time of arrival of the Altai populations remain unresolved. Excavations at Chagyrskaya Cave in the Altai foothills have yielded 90,000 stone artifacts, numerous bone tools, 74 Neanderthal fossils, and animal and plant remains recovered from 59,000- to 49,000-year-old deposits. The Chagyrskaya Neanderthals made distinctive stone tools that closely resemble Micoquian artifacts from eastern Europe, whereas other Altai sites occupied by earlier Neanderthal populations lack such artifacts. This suggests at least two dispersals of Neanderthals into southern Siberia, with the likely ancestral homeland of the Chagyrskaya toolmakers located 3,000 to 4,000 kilometers to the west, in eastern Europe. Neanderthals were once widespread across Europe and western Asia. They also penetrated into the Altai Mountains of southern Siberia, but the geographical origin of these populations and the timing of their dispersal have remained elusive. Here we describe an archaeological assemblage from Chagyrskaya Cave, situated in the Altai foothills, where around 90,000 Middle Paleolithic artifacts and 74 Neanderthal remains have been recovered from deposits dating to between 59 and 49 thousand years ago (age range at 95.4% probability). Environmental reconstructions suggest that the Chagyrskaya hominins were adapted to the dry steppe and hunted bison. Their distinctive toolkit closely resembles Micoquian assemblages from central and eastern Europe, including the northern Caucasus, more than 3,000 kilometers to the west of Chagyrskaya Cave. At other Altai sites, evidence of earlier Neanderthal populations lacking associated Micoquian-like artifacts implies two or more Neanderthal incursions into this region. We identify eastern Europe as the most probable ancestral source region for the Chagyrskaya toolmakers, supported by DNA results linking the Neanderthal remains with populations in northern Croatia and the northern Caucasus, and providing a rare example of a long-distance, intercontinental population movement associated with a distinctive Paleolithic toolkit.


Pros and cons of methylation-based enrichment methods for ancient DNA

Abstract: The recent discovery that DNA methylation survives in fossil material provides an opportunity for novel molecular approaches in palaeogenomics. Here, we apply to ancient DNA extracts the probe-independent Methylated Binding Domains (MBD)-based enrichment method, which targets DNA molecules containing methylated CpGs. Using remains of a Palaeo-Eskimo Saqqaq individual, woolly mammoths, polar bears and two equine species, we confirm that DNA methylation survives in a variety of tissues, environmental contexts and over a large temporal range (4,000 to over 45,000 years before present). MBD enrichment, however, appears principally biased towards the recovery of CpG-rich and long DNA templates and is limited by the fast post-mortem cytosine deamination rates of methylated epialleles. This method, thus, appears only appropriate for the analysis of ancient methylomes from very well preserved samples, where both DNA fragmentation and deamination have been limited. This work represents an essential step toward the characterization of ancient methylation signatures, which will help understanding the role of epigenetic changes in past environmental and cultural transitions.


Extinction, survival and recovery of corals from the Triassic to Middle Jurassic time

Abstract: Recognizing extinction events and determining their cause at the Triassic/Jurassic (T/J) transition and near the Pliensbachian–Toarcian (Lower Jurassic) boundary is a field of growing interest. We provide arguments for these events through a literature based new evaluation of coral diversity from Triassic to Dogger and a new palaeobiogeographical map. The T/J extinction of corals is clearly related to the breakdown of reef environments. Origination curves show that Hettangian (the lowest Jurassic stage) was not only a survival phase but already rather a recovery phase. Post‐extinction evolution of reefs and their survival only in the northernmost margin of the Tethys support the hothouse hypothesis for the T/J extinction event. During Pliensbachian, many new taxa appear, but mostly solitary corals, not really framebuilders. Many of these taxa do not occur anymore during the following stages. The new increase in diversity is related to the development of Bajocian (Middle Jurassic) reefs.


How old is the Isthmus of Panama

Abstract: The Standard Model of the formation of the Isthmus of Panama proposes that final closure occurred at 4-3 Ma. The model is based on evidence from studies of marine stratigraphy, fossil sequences, divergent molecular phylogenies, the timing of the Great American Biological Interchange (GABI), and proxies for marine paleosalinity, paleobathymetry, productivity, and paleotemperatures. The new model uses cooling of magmas in the Cretaceous to Early Miocene Central American Volcanic arc to propose Eocene emergence of the discrete structural blocks of the arc and then U/Pb dating, paleomagnetic pole rotations, and atlantic sea-floor anomalies to reconfigure the blocks for different time slices back to 25 Ma. Closure is proposed at 15 Ma, because by this time the alignment of the blocks leaves no space for trans-isthmian marine passages. We propose that the Indonesian Australian Archipelago (IAA) provides a model for the Central American arc between 15 and 3 Ma because it accounts for the extensive marine interchange between the Pacific and Indian oceans through few and narrow passages while maintaining a complete separation of the terrestrial faunas (Wallace and Lydekker lines) of the two continental platforms of Sunda and Sahul. Using the Indonesian Volcanic arc as a model, we can now accommodate the general tectonic configuration and much of the emergence of the new model, while accounting for the marine fossil record, the molecular evidence of rapid recent speciation, and the delayed Great American Biotic Interchange that the New Model fails to explain.


Neuroanatomy of the mekosuchine crocodylian Trilophosuchus rackhami Willis, 1993

Abstract: Although our knowledge on crocodylomorph palaeoneurology has experienced considerable growth in recent years, the neuroanatomy of many crocodylomorph taxa has yet to be studied. This is true for Australian taxa, where thus far only two crocodylian crocodylomorphs have had aspects of their neuroanatomy explored. Here, the neuroanatomy of the Australian mekosuchine crocodylian Trilophosuchus rackhami is described for the first time, which significantly increases our understanding on the palaeoneurology of Australian crocodylians. The palaeoneurological description is based on the taxon's holotype specimen (QMF16856), which was subjected to a μCT scan. Because of the exceptional preservation of QMF16856, most neuroanatomical elements could be digitally reconstructed and described in detail. Therefore, the palaeoneurological assessment presented here is hitherto the most in‐depth study of this kind for an extinct Australian crocodylomorph. Trilophosuchus rackhami has a brain endocast with a distinctive morphology that is characterized by an acute dural peak over the hindbrain region. While the overall morphology of the brain endocast is unique to T. rackhami, it does share certain similarities with the notosuchian crocodyliforms Araripesuchus wegeneri and Sebecus icaeorhinus. The endosseous labyrinth displays a morphology that is typical for crocodylians, although a stand‐out feature is the unusually tall common crus. Indeed, the common crus of T. rackhami has one of the greatest height ratios among crocodylomorphs with currently known endosseous labyrinths. The paratympanic pneumatic system of T. rackhami is greatly developed and most similar to those of the extant crocodylians Osteolaemus tetraspis and Paleosuchus palpebrosus. The observations on the neuroanatomy of T. rackhami are also discussed in the context of Crocodylomorpha. The comparative palaeoneurology reinforces previous evaluations that the neuroanatomy of crocodylomorphs is complex and diverse among species, and T. rackhami has a peculiar neuromorphology, particularly among eusuchian crocodyliforms.

mekosuchine
trilophosuchus rackhami
qmf16856
neuroanatomy
ristevski


Life history, sexual dimorphism and ‘ornamental’ feathers in the mesozoic bird Confuciusornis sanctus

Abstract: The life history of Confuciusornis sanctus is controversial. Recently, the species’ body size spectrum was claimed to contradict osteohistological evidence for a rapid, bird-like development. Moreover, sexual size dimorphism was rejected as an explanation for the observed bimodal size distribution since the presence of elongated rectrices, an assumed ‘male’ trait, was uncorrelated with size. However, this interpretation (i) fails to explain the size spectrum of C. sanctus which is trimodal rather than bimodal, (ii) requires implausible neonate masses and (iii) is not supported by analogy with sexual dimorphisms in modern birds, in which elongated central rectrices are mostly sex-independent. Available information on C. sanctus is readily reconciled if we assume a bird-like life history, as well as a pronounced sexual size dimorphism and sexually isomorphic extravagant feathers as frequently observed in extant species.

sexual dimorphism
feather
confuciusornis sanctus
bird-like life history
life history


The Antiquity of the Rhine River: Stratigraphic Coverage of the Dinotheriensande (Eppelsheim Formation) of the Mainz Basin (Germany)

Abstract: Background Mammalian fossils from the Eppelsheim Formation (Dinotheriensande) have been a benchmark for Neogene vertebrate palaeontology since 200 years. Worldwide famous sites like Eppelsheim serve as key localities for biochronologic, palaeobiologic, environmental, and mammal community studies. So far the formation is considered to be of early Late Miocene age (∼9.5 Ma, Vallesian), representing the oldest sediments of the Rhine River. The stratigraphic unity of the formation and of its fossil content was disputed at times, but persists unresolved. Principal Findings Here we investigate a new fossil sample from Sprendlingen, composed by over 300 mammalian specimens and silicified wood. The mammals comprise entirely Middle Miocene species, like cervids Dicrocerus elegans, Paradicrocerus elegantulus, and deinotheres Deinotherium bavaricum and D. levius. A stratigraphic evaluation of Miocene Central European deer and deinothere species proof the stratigraphic inhomogenity of the sample, and suggest late Middle Miocene (∼12.5 Ma) reworking of early Middle Miocene (∼15 Ma) sediments. This results agree with taxonomic and palaeoclimatic analysis of plant fossils from above and within the mammalian assemblage. Based on the new fossil sample and published data three biochronologic levels within the Dinotheriensand fauna can be differentiated, corresponding to early Middle Miocene (late Orleanian to early Astaracian), late Middle Miocene (late Astaracian), and early Late Miocene (Vallesian) ages. Conclusions/Significance This study documents complex faunal mixing of classical Dinotheriensand fauna, covering at least six million years, during a time of low subsidence in the Mainz Basin and shifts back the origination of the Rhine River by some five million years. Our results have severe implications for biostratigraphy and palaeobiology of the Middle to Late Miocene. They suggest that turnover events may be obliterated and challenge the proposed ‘supersaturated’ biodiversity, caused by Middle Miocene superstites, of Vallesian ecosystems in Central Europe.

dinotheriensande
eppelsheim formation
mainz basin
late miocene


A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina

Abstract: Titanosaurian sauropod dinosaurs were the most diverse and abundant large-bodied herbivores in the southern continents during the final 30 million years of the Mesozoic Era. Several titanosaur species are regarded as the most massive land-living animals yet discovered; nevertheless, nearly all of these giant titanosaurs are known only from very incomplete fossils, hindering a detailed understanding of their anatomy. Here we describe a new and gigantic titanosaur, Dreadnoughtus schrani, from Upper Cretaceous sediments in southern Patagonia, Argentina. Represented by approximately 70% of the postcranial skeleton, plus craniodental remains, Dreadnoughtus is the most complete giant titanosaur yet discovered, and provides new insight into the morphology and evolutionary history of these colossal animals. Furthermore, despite its estimated mass of about 59.3 metric tons, the bone histology of the Dreadnoughtus type specimen reveals that this individual was still growing at the time of death.

titanosaurian sauropod dinosaur
southern patagonia
argentina
dreadnoughtus schrani


New teeth of allotherian mammals from the English Bathonian, including the earliest multituberculates

Abstract:
Twenty one isolated multituberculate-like teeth are described from the Forest Marble (late Bathonian) of Oxfordshire and Dorset, England. Eighteen are additional to the teeth described as Eleutherodon oxfordensis by Kermack et al. (1998), and three of those are placed in new taxa. Six new molars of Eleutherodon provide further information on variation in size, proportion and root pattern. Millsodon superstes gen. et sp. nov. (family indeterminate), based on first and last lower molars and a referred upper molar, has resemblances to Haramiyidae and Theroteinidae. Kirtlingtonia catenata gen. et sp. nov. (family indeterminate), based on last upper molars and a probable upper premolar, has a slight resemblance to Eleutherodon, and also to M2 of some paulchoffatiid multituberculates. Kermackodon multicuspis gen. et sp. nov. (family Kermackodontidae nov.) and Hahnotherium antiquum gen. et sp. nov. (family Hahnotheriidae nov.
) are based on second upper molars, recognised as multituberculate by their horizontal wear and inferred occlusal displacement with respect to m2. A lower molar referred to H. antiquum confirms this. A blade-like lower premolar and an upper premolar with conical cusps, referred to Kermackodon, are multituberculate-like, but distinctive. Divergence between the two Bathonian multituberculates indicates that the order originated much earlier, more probably from a haramiyid than from a morganucodontid source. Mojo is regarded as probably a haramiyid. The Hahnodontidae, which have basined wear, are removed from the Multituberculata to the "Haramiyida".

multituberculate haramiyidae tooth haramiyidan


First turtle from the Ipubi Formation (Early Cretaceous), Santana Group, Araripe Basin, Brazil

Abstract: The Araripe Basin is located in northeastern Brazil between the states of Ceara, Piaui and Pernambuco (Figure 1). The Crato and Romualdo formations (Santana Group, see details in Neumann & Cabrera, 1999; Valenca et al., 2003) that comprise the most fossiliferous strata of this basin, are famous worldwide for the diverse and exquisitely well preserved fossil assemblages (e.g. Maisey, 1991). These two different lagerstatten were formed during the Lower Cretaceous, Aptian/Albian (Pons et al., 1990; ABSTRACT – Up to date turtle remains from the Early Cretaceous Santana Group are known only to the Crato and Romualdo formations (Aptian-Albian) that have yielded five species: Araripemys barretoi Price, 1973; Santanachelys gaffneyi Hirayama, 1998; Brasilemys josai Lapparent de Broin, 2000; Cearachelys placidoi Gaffney, Campos & Hirayama, 2001 and Euraxemys essweini Gaffney, Tong & Meylan, 2006. Except for A. barretoi, known from both, the Romualdo and Crato formations, all other turtle taxa are restricted to the Romualdo Formation. Fragments of the skull, an incomplete lower jaw and carapace of the first turtle remains from the Ipubi Formation are reported here. This material was collected in the shales above the gypsum layer, close to the contact with the Romualdo Formation (Aptian-Albian). Compared to other turtles from the Santana Group, this specimen is referred to Pelomedusoides, excluding Araripemys and Brasilemys, based on the following characteristics: absence of a cavum pterygoidei, absence of cervical scute, absence of fontanels in the carapace, contact between nuchal and first peripheral, and absence of nuchal emargination.


A LONG PROCESS TOWARDS AGRICULTURE IN THE MIDDLE YELLOW RIVER VALLEY, CHINA: EVIDENCE FROM MACRO- AND MICRO-BOTANICAL REMAINS

Abstract: Macro- and micro-botanical remains dating from the Upper Paleolithic through early Neolithic periods in North China have provided significant information for reconstructing the changing subsistence patterns as human groups evolved from mobile hunting-gathering societies to sedentary farming communities. Starch analysis on grinding stones, in particular, has revealed much new data that supplement the inventory of carbonized remains recovered by flotation methods. This paper reviews some recent research projects which have documented a long tradition of processing various plants with grinding stones in the Middle Yellow River valley, including tubers, beans, nuts, and cereals. Exploitation of wild millet can be traced back to 23,000-19,500 cal. BP, more than 10,000 years before its domestication. Several species of tuber, acorn, and wild grasses made up significant proportions of staple food during the early Neolithic, when millet domestication was already underway. These new data help us to better understand the extended transitional process to agriculture in the Middle Yellow River region. Archaeobotany is in an early stage of development in China; it is important to employ an interdisciplinary approach for a more complete documentation of plant use in the past and a better understanding of subsistence practices then.


Gryposuchus (Crocodylia, Gavialoidea) from the early Miocene of Venezuela

Abstract: Here, a fragment of a mandible recently discovered in the Cerro Zamuro site (Castillo Formation, Lara State, northwestern Venezuela) is assigned to the giant gavialoid Gryposuchus. This specimen, recovered from putative brackish environments of the early Miocene (*18 Ma) age, is unequivocally the earliest record of the genus in South America. Gryposuchus, together with the other gryposuchine previously recognized from the Castillo Formation, Siquisiquesuchus venezuelensis, increases the early Miocene taxonomic diversity of the group in the northern Neotropics. This new information from the Castillo Formation supports the conclusion that early gryposuchine evolutionary stages were in coastal, shallow marine or brackish environments, while the presence of some genera, such as Gryposuchus, in middle to late Miocene freshwater environments, is secondary habitat colonization late in the evolution of the clade. Freshwater colonization is probably the result of the gradual adaptation of early marine-adapted gryposuchines to the extensive estuarine-like environments of northern South America lowlands associated with marine transgressions that systematically occurred during the middle Eocene to early Oligocene. This new record is evidence of the wide chronological distribution of Gryposuchus in northern South America, highlighting the importance of this area as the center of origin and radiation of this successful Miocene gavialoid.

early miocene
castillo formation
northern south america
gryposuchus


When bivalves took over the world

Abstract: Abstract The end-Permian mass extinction is commonly portrayed not only as a massive biodiversity crisis but also as the time when marine benthic faunas changed from the Paleozoic Fauna, dominated by rhynchonelliform brachiopod taxa, to the Modern Fauna, dominated by gastropod and bivalve taxa. After the end-Permian mass extinction, scenarios involving the Mesozoic Marine Revolution portray a steady increase in numerical dominance by these benthic molluscs as largely due to the evolutionary effects of an “arms race.” We report here a new global paleoecological database from study of shell beds that shows a dramatic geologically sudden earliest Triassic takeover by bivalves as numerical dominants in level-bottom benthic marine communities, which continued through the Early Triassic. Three bivalve genera were responsible for this switch, none of which has any particular morphological features to distinguish it from many typical Paleozoic bivalve genera. The numerical success of these Early Triassic bivalves cannot be attributed to any of the well-known morphological evolutionary innovations of post-Paleozoic bivalves that characterize the Mesozoic Marine Revolution. Rather, their ability to mount this takeover most likely was due to the large extinction of rhynchonelliform brachiopods during the end-Permian mass extinction and aided by their environmental distribution and physiological characteristics that enabled them to thrive during periods of oceanic and atmospheric stress during the Permian/Triassic transition.


Decoupled taxonomic and ecological recoveries from the Permo-Triassic extinction

Abstract: After the enormous Permian-Triassic mass extinction, marine animals high in the food chain recovered the most quickly. The Permian-Triassic mass extinction was the worst crisis faced by life; it killed >90% of marine species in less than 0.1 million years (Ma). However, knowledge of its macroecological impact over prolonged time scales is limited. We show that marine ecosystems dominated by non-motile animals shifted to ones dominated by nektonic groups after the extinction. In Triassic oceans, animals at high trophic levels recovered faster than those at lower levels. The top-down rebuilding of marine ecosystems was still underway in the latest Triassic, ~50 Ma after the extinction, and contrasts with the ~5-Ma recovery required for taxonomic diversity. The decoupling between taxonomic and ecological recoveries suggests that a process of vacant niche filling before reaching the maximum environmental carrying capacity is independent of ecosystem structure building.


Competition structured a Late Cretaceous megaherbivorous dinosaur assemblage

Abstract: Modern megaherbivore community richness is limited by bottom-up controls, such as resource limitation and resultant dietary competition. However, the extent to which these same controls impacted the richness of fossil megaherbivore communities is poorly understood. The present study investigates the matter with reference to the megaherbivorous dinosaur assemblage from the middle to upper Campanian Dinosaur Park Formation of Alberta, Canada. Using a meta-analysis of 21 ecomorphological variables measured across 14 genera, contemporaneous taxa are demonstrably well-separated in ecomorphospace at the family/subfamily level. Moreover, this pattern is persistent through the approximately 1.5 Myr timespan of the formation, despite continual species turnover, indicative of underlying structural principles imposed by long-term ecological competition. After considering the implications of ecomorphology for megaherbivorous dinosaur diet, it is concluded that competition structured comparable megaherbivorous dinosaur communities throughout the Late Cretaceous of western North America.

dinosaur assemblage
north america
competition


Experimental disintegration of regular echinoids: roles of temperature, oxygen, and decay thresholds

Abstract: Laboratory experiments on regular echinoids indicate that low water temperatures retard organic decomposition far more effectively than anoxia, and that the primary role of anoxia in the preservation of articulated multi-element calcareous skeletons may be in excluding scavenging organisms. When tumbled at 20 rpm, specimens that were first allowed to decay for two days in warm seawater (30°C) disintegrated more than six times faster than specimens treated at room temperature (23°C) and more than an order of magnitude faster than specimens treated in cool water (11°C). In contrast, the effects of aerobic versus anerobic decay on disintegration rates were insignificant. The longer the period that specimens were allowed to decay before tumbling, the greater the rate at which specimens disintegrated, until a threshold time that appears to mark the decomposition of collagenous ligaments. This required a few days at 30°C, about two weeks at 23°C, and more than 4 weeks at 11°C for Strongylocentrotus. Up until this threshold, coronas disintegrate by a combination of cross-plate fractures and separation along plate sutures; cross-plate fractures thus can be taphonomic in origin and are not necessarily related to predation. Specimens decayed for longer-than-threshold periods of time disintegrate virtually instantaneously upon tumbling by sutural separation only. Undisturbed coronas can remain intact for months, sufficient time for epibiont occupation. Rates of disintegration were documented semi-quantitatively by recognizing seven stages of test disarticulation, and quantitatively by tensometer measures of test strength and toughness. The effects of temperature and oxygen on decay and the existence of a decay threshold in disintegration should apply at least in a qualitative sense to many other animals whose skeletons consist of multiple, collagen-bound elements. Regular echinoids should still be perceived as taphonomically fragile organisms, but our results suggest the potential for latitudinal as well as bathymetric gradients in the preservation of fossil echinoid faunas. Echinoid preservation under any given set of conditions should also be a function of taxonomic differences in test construction (particularly stereom interlocking along plate sutures) as suggested by previous workers, although our experiments indicate that these effects should only be significant among post-threshold specimens. A survey of regular echinoids from Upper Cretaceous white chalk facies of Britain substantiates the basic experimental patterns, yielding examples of all disarticulation stages and significant taxonomic differences in quality of preservation. A diverse array of borers and encrusters on fossil coronas also corroborates the post-mortem persistence of some tests on mid-latitude seafloors.

disintegration
regular echinoid
oxygen
plate suture


Temporal labyrinths of eastern Eurasian Pleistocene humans

Abstract: Significance The assessment of the paleobiology and morphological affinities of the Neandertals and other Late Pleistocene archaic humans is central to resolving issues regarding the emergence and establishment of modern human morphology and diversity. One feature, which has been used as a distinctive Neandertal feature in this context, is the apparently derived shape of their temporal labyrinths (especially semicircular canals). Analysis of East Asian labyrinths documents the “Neandertal” pattern in the Xujiayao 15 temporal bone, although none of the Xujiayao human remains exhibits other distinctly Neandertal features. It therefore raises questions regarding possible biological correlates of labyrinthine morphology, distinctive Neandertal features, and the nature of late archaic human variation across Eurasia. One of the morphological features that has been identified as uniquely derived for the western Eurasian Neandertals concerns the relative sizes and positions of their semicircular canals. In particular, they exhibit a relatively small anterior canal, a relatively larger lateral one, and a more inferior position of the posterior one relative to the lateral one. These discussions have not included full paleontological data on eastern Eurasian Pleistocene human temporal labyrinths, which have the potential to provide a broader context for assessing Pleistocene Homo trait polarities. We present the temporal labyrinths of four eastern Eurasian Pleistocene Homo, one each of Early (Lantian 1), Middle (Hexian 1), and Late (Xujiayao 15) Pleistocene archaic humans and one early modern human (Liujiang 1). The labyrinths of the two earlier specimens and the most recent one conform to the proportions seen among western early and recent modern humans, reinforcing the modern human pattern as generally ancestral for the genus Homo. The labyrinth of Xujiayao 15 is in the middle of the Neandertal variation and separate from the other samples. This eastern Eurasian labyrinthine dichotomy occurs in the context of none of the distinctive Neandertal external temporal or other cranial features. As such, it raises questions regarding possible cranial and postcranial morphological correlates of Homo labyrinthine variation, the use of individual “Neandertal” features for documenting population affinities, and the nature of late archaic human variation across Eurasia.


The Basal Penguin (Aves: Sphenisciformes) Perudyptes devriesi and a Phylogenetic Evaluation of the Penguin Fossil Record

Abstract: Abstract We present the first detailed description of Perudyptes devriesi, a basal penguin from the middle Eocene (~42 Ma) Paracas Formation of Peru, and a new analysis of all published extinct penguin species as well as controversial fragmentary specimens. The Perudyptes devriesi holotype includes key regions of the skull and significant postcranial material, thus helping to fill a major phylogenetic and stratigraphic (~20 million year) gap between the earliest fossil penguins (Waimanu manneringi and Waimanu tuatahi, ~58–61.6 Ma) and the next oldest partial skeletons. Perudyptes devriesi is diagnosable by five autapomorphies: (1) an anteroventrally directed postorbital process, (2) marked anterior expansion of the parasphenoid rostrum, (3) posterior trochlear ridge of the humerus projecting distal to the middle trochlear ridge and conformed as a large, broadly curved surface, (4) convex articular surface for the antitrochanter of the femur, and (5) extremely weak anterior projection of the lateral condyle of the tibiotarsus. The skull of Perudyptes is characterized by deep temporal fossae and an elongate, narrow beak that differs from other reported stem penguins in its short mandibular symphysis. The wing skeleton of Perudyptes preserves a combination of plesiomorphic features also observed in the basal penguin Waimanu and derived features shared with more crownward penguins. Features of the wing optimized as primitive for Sphenisciformes include retention of a discrete dorsal supracondylar tubercle on the humerus and presence of a modestly projected pisiform process on the carpometacarpus. Derived features present in Perudyptes and all more crownward penguins, but absent in Waimanu, include a more flattened humerus, development of a trochlea for the tendon of m. scapulotriceps at the distal end of the humerus, and bowing of the anterior face of the carpometacarpus. A combined molecular and morphological dataset for Spheniciformes was expanded by adding 25 osteological and soft tissue characters as well as 11 taxa. In agreement with previous results, Perudyptes devriesi is identified as one of the most basal members of Sphenisciformes. This analysis also confirms the placement of the middle/late Miocene (~11–13 Ma) fossil Spheniscus muizoni as a member of the Spheniscus clade and places the late Miocene (~10 Ma) Madrynornis mirandus as sister taxon to extant Eudyptes. These two species, known from relatively complete partial skeletons, are the oldest crown clade penguin fossils and represent well-corroborated temporal calibration points for the Spheniscus-Eudyptula divergence and Megadyptes-Eudyptes divergence, respectively. Our results reaffirm that the Miocene penguin taxon Palaeospheniscus, recently proposed to represent a member of the crown radiation, belongs outside of the crown clade Spheniscidae. The phylogenetic positions of small Eocene Antarctic penguin taxa (Delphinornis, Marambiornis, and Mesetaornis) recently proposed as possible direct ancestors to crown Spheniscidae were further evaluated using alternate coding strategies for incorporating scorings from isolated elements that preserve critical morphologies and are thought to represent these taxa, although they cannot yet be reliably assigned to individual species. Under all scoring regimes, Delphinornis, Marambiornis, and Mesetaornis were recovered as distantly related to Spheniscidae. Using synapomorphies identified in the primary analysis, we evaluated the phylogenetic position of fragmentary specimens, including the holotypes of valid but poorly known species, specimens currently unassignable to the species level, and morphologically distinct specimens that have not yet been named. All pre-Miocene specimens can be excluded from Spheniscidae based on presence of plesiomorphies lost in all crown penguins, consistent with a recent radiation for the penguin crown clade. This study provides additional support for a scenario of penguin evolution characterized by an origin of flightlessness near the K-T boundary, dispersal throughout the Southern Hemisphere during the early Paleogene, and a late Cenozoic origin for the crown clade Spheniscidae. Stratigraphic distribution and phylogenetic relationships of fossil penguins are consistent with distinct radiations during the Eocene, Oligocene, and Miocene. While the Eocene and Oligocene penguin faunas are similar in many respects, the Miocene fauna is characterized by smaller average size and novel cranial morphologies, suggesting that an ecological shift in diet occurred close to the origin of crown Spheniscidae.

basal penguin
sphenisciforme
perudyptes devriesi


First Evidence of Dinosaurian Secondary Cartilage in the Post-Hatching Skull of Hypacrosaurus stebingeri (Dinosauria, Ornithischia)

Abstract: Bone and calcified cartilage can be fossilized and preserved for hundreds of millions of years. While primary cartilage is fairly well studied in extant and fossilized organisms, nothing is known about secondary cartilage in fossils. In extant birds, secondary cartilage arises after bone formation during embryonic life at articulations, sutures and muscular attachments in order to accommodate mechanical stress. Considering the phylogenetic inclusion of birds within the Dinosauria, we hypothesized a dinosaurian origin for this “avian” tissue. Therefore, histological thin sectioning was used to investigate secondary chondrogenesis in disarticulated craniofacial elements of several post-hatching specimens of the non-avian dinosaur Hypacrosaurus stebingeri (Ornithischia, Lambeosaurinae). Secondary cartilage was found on three membrane bones directly involved with masticatory function: (1) as nodules on the dorso-caudal face of a surangular; and (2) on the bucco-caudal face of a maxilla; and (3) between teeth as islets in the alveolar processes of a dentary. Secondary chondrogenesis at these sites is consistent with the locations of secondary cartilage in extant birds and with the induction of the cartilage by different mechanical factors - stress generated by the articulation of the quadrate, stress of a ligamentous or muscular insertion, and stress of tooth formation. Thus, our study reveals the first evidence of “avian” secondary cartilage in a non-avian dinosaur. It pushes the origin of this “avian” tissue deep into dinosaurian ancestry, suggesting the creation of the more appropriate term “dinosaurian” secondary cartilage.

dinosaurian
cartilage
hypacrosaurus stebingeri


Comparative dental microwear of ruminant and perissodactyl molars: Implications for paleodietary analysis of rare and extinct ungulate clades

Abstract: Abstract. Dental microwear analyses of ungulates and other large herbivores rely on correlations of diet and microwear among extant ungulates, primarily ruminants. Microwear is considered a ‘taxon-free’ method of paleodietary analysis. The properties of food are associated with causality of microwear, but the possibility that heritable properties of the consumer (tooth morphologies, masticatory dynamics, enamel mechanical properties, digestive physiologies) may introduce bias is not considered. Using an observer blind method of light microscopy, we examined the distribution of microwear features on the molars of eight species of ruminants and perissodactyls. Grazing and browsing ruminants had statistically different numbers of scratches forming discrete data clusters. Perissodactyls differ in the numbers of scratches and pits but without discrete browser and grazer clusters. Microwear features were distributed homogeneously across ruminantmolars and strongly predictive of diet fromthe labial edge of the molar to the lingual edge. Microwear was heterogeneously distributed across perissodactyl molars with more pits on the labial edge andmore scratches on the lingual edge. In perissodactyls, microwear sampled from the labial edge was strongly predictive of diet, while microwear sampled from other areas were not. Discriminant function analyses of microwear assigned individual molars to diets (browser and grazer) and clades (ruminant and perissodactyl) with similar success (70–73%) indicating that phylogeny and diet influencemicrowear equally. Rhino microwear was more sensitive to clade membership while other perissodactyl microwear was more sensitive to diet. Although it is not clear what heritable variables may phylogenetically bias dentalmicrowear, extant ruminants may not be appropriate models for themicrowear of other large herbivores.

microwear
perissodactyl molar
paleodietary analysis


Whipspiders (Arachnida: Amblypygi) in amber from the Early Eocene and mid-Cretaceous, including maternal care

Abstract: Two new genera and species of fossil whipspiders (Chelicerata: Arachnida: Amblypygi) are described from Tertiary and Cretaceous ambers of southern Asia.  Paracharonopsis cambayensis Engel & Grimaldi, new genus and species, preserved in Cambay amber of Ypresian age from western India is the first Tertiary and Asian fossil of the Paleoamblypygi, a highly relict taxon that includes the Late Carboniferous genus Graeophonus Scudder and the living West African species Paracharon caecus Hansen.  Paracharonopsis cambayensis is one of the few examples in Cambay amber of a biotic connection to Africa; most taxa show widespread or Laurasian distributions.  Kronocharon prendinii Engel & Grimaldi, new genus and species, is the first Cretaceous amber whipspider and putatively a sister group to the Phrynoidea (= Apulvillata).  The holotype female of K. prendinii is preserved with the remains of three nymphs near her, documenting the Early Cretaceous presence of the extended maternal care so distinctive for the order, and a behavioral repertoire widespread among arachnids.


An Oligo-Miocene Magpie Goose (Aves: Anseranatidae) from Riversleigh, Northwestern Queensland, Australia

Abstract: ABSTRACT The magpie goose Anseranas semipalmata (Aves: Anseranatidae), the sole modern representative of the family, is endemic to Australia. The fossil record of Anseranatidae in Australia was until now restricted to Pliocene and younger sediments. Here we describe an anseranatid from the Oligo-Miocene Carl Creek Limestone in the Riversleigh World Heritage Property, Boodjamulla (Lawn Hill) National Park, in northwestern Queensland, as a new species and genus, based on a coracoid and two scapulae. These fossils extend the known age of the Anseranatidae lineage in Australia to about 25 million years ago. The fossil distribution of Anseranatidae now includes the Paleocene in North America, Eocene and Late Oligocene of Europe, and the Late Oligocene to earliest Miocene in Australia, indicating a globally widespread distribution of the family during the early-mid Tertiary.

anseranatidae
riversleigh
northwestern queensland


Functional analysis of sabertooth cranial morphology

Abstract: Elongate canines evolved independently at least four times among mammalian carnivores, and each time skulls were modified in similar ways. We have compared the cranial morphology of sabertooths to that of their non-sabertoothed relatives, living and extinct, and applied simple biomechanical models to elucidate the functional significance of the morphological differences. Our analysis suggests that (1) sabertooth morphology represents modification for wider gape with retention of a powerful bite force at the carnassial; (2) sabertooths probably used a throat or ventral neck slash to kill prey; and (3) elongate canines and retractile claws may have facilitated the exploitation of relatively larger prey by sabertooths compared to non-sabertooth carnivores.

morphology
sabre-tooth morphology
similar ecomorphologie
functional difference


The oldest ammonoids of Morocco (Tafilalt, lower Emsian)

Abstract: During a fieldtrip of the 10th International Cephalopod Symposium to the Tafilalt (SE Morocco), a fauna with the so far oldest ammonoid of the region was discovered at the top of the basal Emsian Deiroceras Limestone of Jebel Mech Irdane. The new material confirms the record of a single, poorly preserved specimen of the same age from Ras El Kebbar in the northwestern Tafilalt. All specimens are assigned within the Chebbitinae n. subfam. (Mimosphinctidae) to Praechebbites debaetsi n. gen. n. sp., which differs from the slightly younger Chebbites reisdorfi mostly in the lack of a dorsal lobe and dorsal imprint zone. In the conodont succession, the new species and genus falls in the top of the Eolinguipolygnathus excacatus M114 Zone, which has been proposed to define in future the base of the Emsian. Since the new taxon is morphologically already advanced, it is likely that even older ammonoids with loosely coiled conchs will be discovered in the future. A block from a Lower Carboniferous olistostrome of the adjacent Tinerhir region contained the gyroconic Ivoites n. sp. only. The global comparison of oldest ammonoid faunas shows the rarity of assemblages assigned to Lower Devonian (LD) III-B, often dated by conodonts and dacryoconarids. These faunas display a high level of endemism, which suggests that ammonoid origin and early evolution took place in regional populations with a restricted dispersal of genera and species.


Disambiguating the scientific names of cockroaches

Abstract: Cockroaches are typically recognized as an order or a suborder in insects, sometimes rank-free. Many scientific names for cockroaches are in use, most of them are ambiguous because of frequent changes of the spelling and definition, and the discrepancy between phenetics and cladistics. Omission of the authorship of the name and of the definition further confuses readers, which include taxonomists. The underlying causes of the ambiguity are the paraphyly of cockroaches and various reproduction strategies (crucial in cockroach classification) of fossil cockroaches. This paper historically reviews the scientific names for cockroaches, and recommends the following names with phylogenetic definition (details and authorship are given in the text): 1) Blattodea are cockroaches with internal ovipositors (producing oothecae of the extant type) and all descendants including termites; 2) suborder Blattaria are Blattodea minus termites; 3) order Dictyoptera are cockroaches that produce genuine oothecae and all descendants including Blattodea, Mantodea and Alienoptera; 4) Holopandictyoptera are all insects that are recognized as cockroaches and their descendants; 5) Eoblattodea are the so-called ‘roachoids’, i.e., cockroaches bearing long external ovipositors, no genuine oothecae. Accordingly, the cockroaches in the broader sense fall into three taxa: Blattaria, basal Dictyoptera, and Eoblattodea.