Re-examination of the cranial osteology of the Arctic Alaskan hadrosaurine with implications for its taxonomic status

Abstract: Hadrosaurid fossils from the Liscomb Bonebed (Prince Creek Formation, North Slope, Alaska) were the first dinosaur bones discovered from the Arctic. While the Prince Creek Formation hadrosaurids were long identified as Edmontosaurus, a member of the sub-clade Hadrosaurinae, they were recently assigned to a newly-erected taxon, Ugrunaaluk kuukpikensis. However, taxonomic status of the new taxon is ambiguous largely due to the immature nature of the specimens upon which it was based. Here we reexamine cranial elements of the Prince Creek Formation hadrosaurine in order to solve its taxonomic uncertainties. The Prince Creek Formation hadrosaurine possesses a short dorsolateral process of the laterosphenoid, one of the diagnostic characters of Edmontosaurus. The Prince Creek Formation hadrosaurine also shows affinity to Edmontosaurus regalis in the presence of a horizontal shelf of the jugal. Our morphological comparisons with other North American Edmontosaurus specimens and our phylogenetic analyses demonstrate that the Prince Creek Formation hadrosaurine should be re-assigned to Edmontosaurus. Because the Prince Creek Formation Edmontosaurus shows differences with lower latitude Edmontosaurus in a dorsoventrally short maxilla, presence of a secondary ridge on the dentary teeth, and the absence of the transverse ridge between basipterygoid processes of the basisphenoid, we consider that the Prince Creek Formation Edmontosaurus should be regarded as Edmontosaurus sp. until further discoveries of mature hadrosaurines from the Prince Creek Formation Bonebed and/or equivalently juvenile Edmontosaurus specimens from the lower latitudes allow direct comparisons. The retention of the Prince Creek Formation hadrosaurine as Edmontosaurus re-establishes a significant latitudinal distribution for this taxon. Despite the large latitudinal distribution of the taxon, the morphological disparity of Edmontosaurus is small within Hadrosaurinae. The small morphological disparity may be related to the relatively low latitudinal temperature gradient during the latest Cretaceous compared to present day, a gradient which might not have imposed significant pressure for much morphological adaptations across a broad latitudinal range.


Effects of taphonomic deformation on geometric morphometric analysis of fossils: a study using the dicynodont Diictodon feliceps (Therapsida, Anomodontia)

Abstract: Taphonomic deformation, the distortion of fossils as a result of geological processes, poses problems for the use of geometric morphometrics in addressing paleobiological questions. Signal from biological variation, such as ontogenetic trends and sexual dimorphism, may be lost if variation from deformation is too high. Here, we investigate the effects of taphonomic deformation on geometric morphometric analyses of the abundant, well known Permian therapsid Diictodon feliceps. Distorted Diictodon crania can be categorized into seven typical styles of deformation: lateral compression, dorsoventral compression, anteroposterior compression, “saddle-shape” deformation (localized collapse at cranial mid-length), anterodorsal shear, anteroventral shear, and right/left shear. In simulated morphometric datasets incorporating known “biological” signals and subjected to uniform shear, deformation was typically the main source of variance but accurate “biological” information could be recovered in most cases. However, in empirical datasets, not only was deformation the dominant source of variance, but little structure associated with allometry and sexual dimorphism was apparent, suggesting that the more varied deformation styles suffered by actual fossils overprint biological variation. In a principal component analysis of all anomodont therapsids, deformed Diictodon specimens exhibit significant dispersion around the “true” position of this taxon in morphospace based on undistorted specimens. The overall variance associated with deformation for Anomodontia as a whole is minor, and the major axes of variation in the study sample show a strong phylogenetic signal instead. Although extremely problematic for studying variation in fossil taxa at lower taxonomic levels, the cumulative effects of deformation in this study are shown to be random, and inclusion of deformed specimens in higher-level analyses of morphological disparity are warranted. Mean morphologies of distorted specimens are found to approximate the morphology of undistorted specimens, so we recommend use of species-level means in higher-level analyses when possible.

taphonomic deformation
anomodontia
variance
low taxonomic level


Taxonomic reassessment of Clevosaurus latidens Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian Phylogeny based on Parsimony and Bayesian inference

Abstract:
Abstract.
 The Late Triassic rhynchocephalian Clevosaurus latidens Fraser, 1993 is known from the fissure deposits of Cromhall Quarry, England. Many studies have questioned its referral to the genus Clevosaurus Swinton, 1939 and some phylogenetic analyses suggest a close relationship with herbivorous rhynchocephalians. We re-examine the type specimens and referred material of C. latidens to elucidate its taxonomic identity. Additionally, we provide new phylogenetic analyses of the Rhynchocephalia using both parsimony and Bayesian approaches. Our taxonomic review and both phylogenetic analyses reveal that C. latidens is not referable to Clevosaurus, but represents a new genus. We reassess C. latidens and provide an amended diagnosis for Fraserosphenodon new genus. Both parsimony and Bayesian analyses recover similar topologies and we propose formal names for two higher clades within Rhynchocephalia: Eusphenodontia new infraorder and Neosphenodontia new clade.


Ecological niche modelling does not support climatically-driven dinosaur diversity decline before the Cretaceous/Paleogene mass extinction

Abstract: In the lead-up to the Cretaceous/Paleogene mass extinction, dinosaur diversity is argued to have been either in long-term decline, or thriving until their sudden demise. The latest Cretaceous (Campanian–Maastrichtian [83–66 Ma]) of North America provides the best record to address this debate, but even here diversity reconstructions are biased by uneven sampling. Here we combine fossil occurrences with climatic and environmental modelling to quantify latest Cretaceous North American dinosaur habitat. Ecological niche modelling shows a Campanian-to-Maastrichtian habitability decrease in areas with present-day rock-outcrop. However, a continent-wide projection demonstrates habitat stability, or even a Campanian-to-Maastrichtian increase, that is not preserved. This reduction of the spatial sampling window resulted from formation of the proto-Rocky Mountains and sea-level regression. We suggest that Maastrichtian North American dinosaur diversity is therefore likely to be underestimated, with the apparent decline a product of sampling bias, and not due to a climatically-driven decrease in habitability as previously hypothesised.

dinosaur diversity
paleogene mass extinction
sampling
ecological niche modelling
possibility


Single-stranded DNA library preparation from highly degraded DNA using T4 DNA ligase

Abstract: Abstract DNA library preparation for high-throughput sequencing of genomic DNA usually involves ligation of adapters to double-stranded DNA fragments. However, for highly degraded DNA, especially ancient DNA, library preparation has been found to be more efficient if each of the two DNA strands are converted into library molecules separately. We present a new method for single-stranded library preparation, ssDNA2.0, which is based on single-stranded DNA ligation with T4 DNA ligase utilizing a splinter oligonucleotide with a stretch of random bases hybridized to a 3΄ biotinylated donor oligonucleotide. A thorough evaluation of this ligation scheme shows that single-stranded DNA can be ligated to adapter oligonucleotides in higher concentration than with CircLigase (an RNA ligase that was previously chosen for end-to-end ligation in single-stranded library preparation) and that biases in ligation can be minimized when choosing splinters with 7 or 8 random nucleotides. We show that ssDNA2.0 tolerates higher quantities of input DNA than CircLigase-based library preparation, is less costly and better compatible with automation. We also provide an in-depth comparison of library preparation methods on degraded DNA from various sources. Most strikingly, we find that single-stranded library preparation increases library yields from tissues stored in formalin for many years by several orders of magnitude.

dna
library preparation
single-stranded dna


Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny

Abstract: Fire may have been a crucial component in the evolution of the Cape flora of South Africa, a region characterized by outstanding levels of species richness and endemism. However, there is, to date, no critical assessment of the age of the modern fire regime in this biome. Here, we exploit the presence of two obligate post-fire flowering clades in the orchid genus Disa, in conjunction with a robust, well-sampled and dated molecular phylogeny, to estimate the age by which fire must have been present. Our results indicate that summer drought (winter rainfall), the fire regime and the fynbos vegetation are several million years older than currently suggested. Summer drought and the fynbos vegetation are estimated to date back to at least the Early Miocene (ca 19.5 Ma). The current fire regime may have been established during a period of global cooling that followed the mid-Miocene Climatic Optimum (ca 15 Ma), which led to the expansion of open habitats and increased aridification. The first appearance of Disa species in the grassland biome, as well as in the subalpine habitat, is in striking agreement with reliable geological and palaeontological evidence of the age of these ecosystems, thus corroborating the efficacy of our methods. These results change our understanding of the historical mechanisms underlying botanical evolution in southern Africa, and confirm the potential of using molecular phylogenies to date events for which other information is lacking or inconclusive.


Diversity of gobioid fishes in the late middle Miocene of northern Moldova, Eastern Paratethys – part I: an extinct clade of Lesueurigobius look-alikes

Abstract: Studies of otoliths suggest that Gobioidei, which are among the most species-rich groups of modern-day vertebrates, were prominent elements of late middle Miocene (early Sarmatian sensu lato) fish faunas in Europe and Western Asia. However, few complete skeletons have come to light. Here, we report an assemblage of six marine gobiid species, based on skeletons preserved with otoliths in situ, from the lower Volhynian (lower Sarmatian s.l.) of Karpov Yar, Naslavcea, northern Moldova (Eastern Paratethys). Previously only one of these species had been reported from the Central Paratethys, based on its otoliths alone. Five new species representing four new genera are described: †Katyagobius prikryli gen. et sp. nov., Pseudolesueurigobius manfredi gen. et sp. nov., †Sarmatigobius compactus gen. et sp. nov., †Yarigobius decoratus gen. et sp. nov., and †Y. naslavcensis gen. et sp. nov. All six species share the following set of characters, suggesting that they represent a monophyletic clade: 27-29 vertebrae (of which 10 are abdominal); spines of first dorsal fin distally filamentous; second dorsal fin with spine and 14-16 soft rays; anal fin with spine and 13-15 soft rays; caudal fin longish-to-lanceolate; otoliths (sagittae) with rounded, trapezoid-to-squarish shape. Their skeletal features suggest that they are closely related to Lesueurigobius Whitley, 1950, but the otoliths preserved in situ do not support such a classification. The new fossils most likely represent a stem lineage of the European Aphia lineage, and indicate that the diversity of gobiid lineages 12 million years ago differed clearly from that observed today.

middle miocene
northern moldova
eastern paratethys


A new family of diprotodontian marsupials from the latest Oligocene of Australia and the evolution of wombats, koalas, and their relatives (Vombatiformes)

Abstract: We describe the partial cranium and skeleton of a new diprotodontian marsupial from the late Oligocene (~26–25 Ma) Namba Formation of South Australia. This is one of the oldest Australian marsupial fossils known from an associated skeleton and it reveals previously unsuspected morphological diversity within Vombatiformes, the clade that includes wombats (Vombatidae), koalas (Phascolarctidae) and several extinct families. Several aspects of the skull and teeth of the new taxon, which we refer to a new family, are intermediate between members of the fossil family Wynyardiidae and wombats. Its postcranial skeleton exhibits features associated with scratch-digging, but it is unlikely to have been a true burrower. Body mass estimates based on postcranial dimensions range between 143 and 171 kg, suggesting that it was ~5 times larger than living wombats. Phylogenetic analysis based on 79 craniodental and 20 postcranial characters places the new taxon as sister to vombatids, with which it forms the superfamily Vombatoidea as defined here. It suggests that the highly derived vombatids evolved from wynyardiid-like ancestors, and that scratch-digging adaptations evolved in vombatoids prior to the appearance of the ever-growing (hypselodont) molars that are a characteristic feature of all post-Miocene vombatids. Ancestral state reconstructions on our preferred phylogeny suggest that bunolophodont molars are plesiomorphic for vombatiforms, with full lophodonty (characteristic of diprotodontoids) evolving from a selenodont morphology that was retained by phascolarctids and ilariids, and wynyardiids and vombatoids retaining an intermediate selenolophodont condition. There appear to have been at least six independent acquisitions of very large (>100 kg) body size within Vombatiformes, several having already occurred by the late Oligocene.

new family
diprotodontian marsupial
late oligocene
vombatoidea


A Molecular Phylogeny of Plesiorycteropus Reassigns the Extinct Mammalian Order ‘Bibymalagasia’

Abstract: Madagascar is well known for its diverse fauna and flora, being home to many species not found anywhere else in the world. However, its biodiversity in the recent past included a range of extinct enigmatic fauna, such as elephant birds, giant lemurs and dwarfed hippopotami. The ‘Malagasy aardvark’ (Plesiorycteropus) has remained one of Madagascar’s least well-understood extinct species since its discovery in the 19th century. Initially considered a close relative of the aardvark (Orycteropus) within the order Tubulidentata, more recent morphological analyses challenged this placement on the grounds that the identifiably derived traits supporting this allocation were adaptations to digging rather than shared ancestry. Because the skeletal evidence showed many morphological traits diagnostic of different eutherian mammal orders, they could not be used to resolve its closest relatives. As a result, the genus was tentatively assigned its own taxonomic order ‘Bibymalagasia’, yet how this order relates to other eutherian mammal orders remains unclear despite numerous morphological investigations. This research presents the first known molecular sequence data for Plesiorycteropus, obtained from the bone protein collagen (I), which places the ‘Malagasy aardvark’ as more closely related to tenrecs than aardvarks. More specifically, Plesiorycteropus was recovered within the order Tenrecoidea (golden moles and tenrecs) within Afrotheria, suggesting that the taxonomic order ‘Bibymalagasia’ is obsolete. This research highlights the potential for collagen sequencing in investigating the phylogeny of extinct species as a viable alternative to ancient DNA (aDNA) sequencing, particularly in cases where aDNA cannot be recovered.


Million-year-old DNA sheds light on the genomic history of mammoths

Abstract: Temporal genomic data hold great potential for studying evolutionary processes, including speciation. However, sampling across speciation events would in many cases require genomic time series that stretch well into the Early Pleistocene (>1 million years). Although theoretical models suggest that DNA should survive on this timescale1, the oldest genomic data recovered so far is from a 560-780 ka old horse specimen2. Here we report the recovery of genome-wide data from three Early and Middle Pleistocene mammoth specimens, two of which are more than one million years old. We find that two distinct mammoth lineages were present in eastern Siberia during the Early Pleistocene. One of these gave rise to the woolly mammoth, whereas the other represents a previously unrecognised lineage that was ancestral to the first mammoths to colonise North America. Our analyses reveal that the North American Columbian mammoth traces its ancestry to a Middle Pleistocene hybridisation between these two lineages, with roughly equal admixture proportions. Finally, we show that the majority of protein-coding changes associated with cold adaptation in woolly mammoths were present already a million years ago. These findings highlight the potential of deep time palaeogenomics to expand our understanding of speciation and long-term adaptive evolution.


Nitrogen isotope evidence for expanded ocean suboxia in the early Cenozoic

Abstract: Circulation more than temperature Changes in continental configuration and sea level affected the ocean's oxygen levels and the rate of denitrification between 70 and 30 million years ago. That finding by Kast et al. shows a fundamental difference from the modern ocean, in which the extent of suboxia is controlled primarily by global temperature. Changes in the nitrogen isotopic composition of marine organic matter correlate with the collision of India and Asia and the circulation changes that occurred as a result. Later, isotopic composition changed further in response to a fall in sea level as global cooling caused ice sheets to grow. Science, this issue p. 386 The ocean was more suboxic and supported greater denitrification in the Paleocene than during the Oligocene. The million-year variability of the marine nitrogen cycle is poorly understood. Before 57 million years (Ma) ago, the 15N/14N ratio (δ15N) of foraminifera shell-bound organic matter from three sediment cores was high, indicating expanded water column suboxia and denitrification. Between 57 and 50 Ma ago, δ15N declined by 13 to 16 per mil in the North Pacific and by 3 to 8 per mil in the Atlantic. The decline preceded global cooling and appears to have coincided with the early stages of the Asia-India collision. Warm, salty intermediate-depth water forming along the Tethys Sea margins may have caused the expanded suboxia, ending with the collision. From 50 to 35 Ma ago, δ15N was lower than modern values, suggesting widespread sedimentary denitrification on broad continental shelves. δ15N rose at 35 Ma ago, as ice sheets grew, sea level fell, and continental shelves narrowed.

foraminifera
shell-bound organic matter
nitrogen


High Plant Diversity in Eocene South America: Evidence from Patagonia

Abstract: Tropical South America has the highest plant diversity of any region today, but this richness is usually characterized as a geologically recent development (Neogene or Pleistocene). From caldera-lake beds exposed at Laguna del Hunco in Patagonia, Argentina, paleolatitude ∼47°S, we report 102 leaf species. Radioisotopic and paleomagnetic analyses indicate that the flora was deposited 52 million years ago, the time of the early Eocene climatic optimum, when tropical plant taxa and warm, equable climates reached middle latitudes of both hemispheres. Adjusted for sample size, observed richness exceeds that of any other Eocene leaf flora, supporting an ancient history of high plant diversity in warm areas of South America.


Strong coupling of predation intensity and diversity in the Phanerozoic fossil record

Abstract: The importance of ecological interactions in driving the evolution of animals has been the focus of intense debate among paleontologists, evolutionary biologists, and macroecologists. To test whether the intensity of such interactions covaries with the secular evolutionary trend in global biodiversity, we compiled a species-level database of predation intensity, as measured by the frequency of common predation traces (drillings and repair scars ranging in age from Ediacaran to Holocene). The results indicate that the frequency of predation traces increased notably by the Ordovician, and not in the mid-Paleozoic as suggested by multiple previous studies. Importantly, these estimates of predation intensity and global diversity of marine metazoans correlate throughout the Phanerozoic fossil record regardless of corrections and methods applied. This concordance may represent (i) an ecological signal: long-term coupling of diversity and predation; (ii) a diversity-driven diffusion of predatory behaviors: an increased probability of more complex predatory strategies to appear at higher diversity levels; or (iii) a spurious concordance in signal capture: an artifact where rare species and less-frequent (e.g., trace-producing) predatory behaviors are both more detectable at times when sampling improves. The coupling of predation and diversity records suggests that macroevolutionary and macroecological patterns share common causative mechanisms that may reflect either historical processes or sampling artifacts.

coupling
predation intensity
phanerozoic fossil record
ediacaran


Body mass predicts isotope enrichment in herbivorous mammals

Abstract: Carbon isotopic signatures recorded in vertebrate tissues derive from ingested food and thus reflect ecologies and ecosystems. For almost two decades, most carbon isotope-based ecological interpretations of extant and extinct herbivorous mammals have used a single diet–bioapatite enrichment value (14‰). Assuming this single value applies to all herbivorous mammals, from tiny monkeys to giant elephants, it overlooks potential effects of distinct physiological and metabolic processes on carbon fractionation. By analysing a never before assessed herbivorous group spanning a broad range of body masses—sloths—we discovered considerable variation in diet–bioapatite δ13C enrichment among mammals. Statistical tests (ordinary least squares, quantile, robust regressions, Akaike information criterion model tests) document independence from phylogeny, and a previously unrecognized strong and significant correlation of δ13C enrichment with body mass for all mammalian herbivores. A single-factor body mass model outperforms all other single-factor or more complex combinatorial models evaluated, including for physiological variables (metabolic rate and body temperature proxies), and indicates that body mass alone predicts δ13C enrichment. These analyses, spanning more than 5 orders of magnitude of body sizes, yield a size-dependent prediction of isotopic enrichment across Mammalia and for distinct digestive physiologies, permitting reconstruction of foregut versus hindgut fermentation for fossils and refined mean annual palaeoprecipitation estimates based on δ13C of mammalian bioapatite.

enrichment
herbivorous mammal
δ13c enrichment
digestive physiology
body mass


A Cretaceous chronostratigraphic database: construction and applications

Abstract: Timing and rates of tectonic events, evolutionary processes, and oceanographic and paleoclimatic changes must be based on high-precision numerical age calibration of stages defined in Global Stratotype and Section Points (GSSPs). The Cretaceous Chronostratigraphic Database (CRETCSDB3) is an objective, testable database that calibrates select Cretaceous events and enables high-resolution chronostratigraphic correlations. CRETCSDB3 is a compilation of more than 3500 taxa and marker beds in nearly 300 published sections calibrated to a mega-annum (Ma) scale. The database spans the Jurassic/Cretaceous and the Cretaceous/Paleogene boundaries. Construction of CRETCSDB3 began by plotting bioevents in the Kalaat Senan, Tunisia, Cenomanian-Turonian section to the 1989 time scale. The sedimentology, sequence stratigraphy, and biostratigraphy of this section were precisely documented and stage boundaries defined biostratigraphically. Additional sections with radiometrically dated beds were graphed to constrain the accuracy of the numerical scale. Ranges of first and last occurrences are calibrated to mega-annums of Cretaceous stages defined by GSSPs or reference sections. This database serves as a look-up table for interpolation and age calibration of other stratigraphic sections. The age ranges of some taxa and marker beds are preliminary and may be extended as new sections are added to the database. CRETCSDB3 tested the numeric age calibration of the Albian/Cenomanian boundary. This boundary in North Texas accurately correlates with the GSSP in France by ammonites, planktic foraminifers and dinoflagellates. This stage boundary in North Texas correlates with the 97.88±0.69 Ma Clay Spur Bentonite in Wyoming by sequence stratigraphy and cosmopolitan dinoflagellates. The inconsistency between this age and the current 100.5 Ma date of the 2012 Geologic Time Scale remains to be evaluated independently.

chronostratigraphic database
database
ammonites
dinoflagellate


The Climate and Landscape of the Middle Part of the Weichselian Glaciation in Europe: The Stage 3 Project

Abstract: Abstract Oxygen isotope stage 3 (OIS 3) was a mild interval between the two cold maxima of the last (Weichselian) glaciation marked by climate changes oscillating on a 100–1000 yr time scale between near-interglacial and peak-glacial conditions. During OIS 3, modern humans entered Europe, and somewhat later their Neanderthal predecessors became extinct. Our understanding of this momentous event depends on an answer to the question, Did the unstable environmental conditions of the time play a significant role in early human history? The Stage 3 Project is an interdisciplinary study with two main goals: (1) to describe with existing data and to simulate the climates and landscapes of typical warm and cold phases between 45,000 and 30,000 yr ago and (2) to compare the results with the spatial and temporal distribution of human beings in this context. This paper introduces the Stage 3 Project and provides background to a set of papers on the climate and landscape aspects of the Project that will appear in Quaternary Research and to studies of their relevance to the Early Upper Paleolithic of Europe to appear in journals yet to be determined.


A new stem-neopterygian fish from the Middle Triassic of China shows the earliest over-water gliding strategy of the vertebrates

Abstract: Flying fishes are extraordinary aquatic vertebrates capable of gliding great distances over water by exploiting their enlarged pectoral fins and asymmetrical caudal fin. Some 50 species of extant flying fishes are classified in the Exocoetidae (Neopterygii: Teleostei), which have a fossil record no older than the Eocene. The Thoracopteridae is the only pre-Cenozoic group of non-teleosts that shows an array of features associated with the capability of over-water gliding. Until recently, however, the fossil record of the Thoracopteridae has been limited to the Upper Triassic of Austria and Italy. Here, we report the discovery of exceptionally well-preserved fossils of a new thoracopterid flying fish from the Middle Triassic of China, which represents the earliest evidence of an over-water gliding strategy in vertebrates. The results of a phylogenetic analysis resolve the Thoracopteridae as a stem-group of the Neopterygii that is more crown-ward than the Peltopleuriformes, yet more basal than the Luganoiiformes. As the first record of the Thoracopteride in Asia, this new discovery extends the geographical distribution of this group from the western to eastern rim of the Palaeotethys Ocean, providing new evidence to support the Triassic biological exchanges between Europe and southern China. Additionally, the Middle Triassic date of the new thoracopterid supports the hypothesis that the re-establishment of marine ecosystems after end-Permian mass extinction is more rapid than previously thought.

middle triassic
gliding strategy
vertebrate
asymmetrical caudal fin
thoracopteridae


Hierarchical biota-level and taxonomic controls on the chemistry of fossil melanosomes revealed using synchrotron X-ray fluorescence

Abstract: Fossil melanosomes, micron-sized granules rich in melanin in vivo, provide key information for investigations of the original coloration, taxonomy and internal anatomy of fossil vertebrates. Such studies rely, in part, on analysis of the inorganic chemistry of preserved melanosomes and an understanding of melanosome chemical taphonomy. The extent to which the preserved chemistry of fossil melanosomes is biased by biotic and abiotic factors is, however, unknown. Here we report the discovery of hierarchical controls on the inorganic chemistry of melanosomes from fossil vertebrates from nine biotas. The chemical data are dominated by a strong biota-level signal, indicating that the primary taphonomic control is the diagenetic history of the host sediment. This extrinsic control is superimposed by a biological, tissue-level control; tissue-specific chemical variation is most likely to survive in fossils where the inorganic chemistry of preserved melanosomes is distinct from that of the host sediment. Comparative analysis of our data for fossil and modern amphibians reveals that most fossil specimens show tissue-specific melanosome chemistries that differ from those of extant analogues, strongly suggesting alteration of original melanosome chemistry. Collectively, these findings form a predictive tool for the identification of fossil deposits with well-preserved melanosomes amenable to studies of fossil colour and anatomy.

control
chemistry
melanosome
fossil vertebrate


Anatomy and relationships of the fuxianhuiid euarthropod Guangweicaris from the early Cambrian Guanshan Biota in Kunming, Yunnan, Southwest China revisited

Abstract: The Guanshan Biota (Cambrian Series 2, Stage 4) is a typical Burgess Shale-type biota, and is one of the most significant Cambrian Konservat-Lagerstätten from China, in addition to the well-known Chengjiang and Kaili biotas. Guangweicaris is a relatively common arthropod from the Guanshan Biota. Based on new specimens from the Longbaoshan and Xinglongcun sections of the Wulongqing Formation in Kunming, we herein report novel features of Guangweicaris spinatus , such as the eye-bearing anterior sclerite that is associated with the head shield, as well as revised details of the trunk, including the ventral appendages. A detailed comparison of Guangweicaris, Fuxianhuia and Shankouia suggests a close affinity between Guangweicaris and Fuxianhuia , which improves our understanding of the evolution of early Cambrian euarthropods.

euarthropod
guanshan biota
guangweicaris spinatus


The global Hangenberg Crisis (Devonian–Carboniferous transition): review of a first-order mass extinction

Abstract: Abstract The global Hangenberg Crisis near the Devonian–Carboniferous boundary (DCB) represents a mass extinction that is of the same scale as the so-called ‘Big Five’ first-order Phanerozoic events. It played an important role in the evolution of many faunal groups and destroyed complete ecosystems but affected marine and terrestrial environments at slightly different times within a short time span of c. 100–300 kyr. The lower crisis interval in the uppermost Famennian started as a prelude with a minor eustatic sea-level fall, followed rather abruptly by pantropically widespread black shale deposition (Hangenberg Black Shale and equivalents). This transgressive and hypoxic/anoxic phase coincided with a global carbonate crisis and perturbation of the global carbon cycle as evidenced by a distinctive positive carbon isotope excursion, probably as a consequence of climate/salinity-driven oceanic overturns and outer-shelf eutrophication. It is the main extinction level for marine biota, especially for ammonoids, trilobites, conodonts, stromatoporoids, corals, some sharks, and deeper-water ostracodes, but probably also for placoderms, chitinozoans and early tetrapods. Extinction rates were lower for brachiopods, neritic ostracodes, bryozoans and echinoderms. Extinction patterns were similar in widely separate basins of the western and eastern Prototethys, while a contemporaneous marine macrofauna record from high latitudes is missing altogether. The middle crisis interval is characterized by a gradual but major eustatic sea-level fall, probably in the scale of more than 100 m, that caused the progradation of shallow-water siliciclastics (Hangenberg Sandstone and equivalents) and produced widespread unconformities due to reworking and non-deposition. The glacio-eustatic origin of this global regression is proven by miospore correlation with widespread diamictites of South America and South and North Africa, and by the evidence for significant tropical mountain glaciers in eastern North America. This isolated and short-lived plunge from global greenhouse into icehouse conditions may follow the significant drawdown of atmospheric CO2 levels due to the prior massive burial of organic carbon during the global deposition of black shales. Increased carbon recycling by intensified terrestrial erosion in combination with the arrested burial of carbonates may have led to a gradual rise of CO2 levels, re-warming, and a parallel increase in the influx of land-derived nutrients. The upper crisis interval in the uppermost Famennian is characterized by initial post-glacial transgression and a second global carbon isotope spike, as well as by opportunistic faunal blooms and the early re-radiation of several fossil groups. Minor reworking events and unconformities give evidence for continuing smaller-scale oscillations of sea-level and palaeoclimate. These may explain the terrestrial floral change near the Famennian–Tournaisian boundary and contemporaneous, evolutionarily highly significant extinctions of survivors of the main crisis. Still poorly understood small-scale events wiped out the last clymeniid ammonoids, phacopid trilobites, placoderms and some widespread brachiopod and foraminiferan groups. The post-crisis interval in the lower Tournaisian is marked by continuing eustatic rise (e.g. flooding of the Old Red Continent), and significant radiations in a renewed greenhouse time. But the recovery had not yet reached the pre-crisis level when it was suddenly interrupted by the global, second-order Lower Alum Shale Event at the base of the middle Tournaisian.

hangenberg crisis
mass extinction
famennian


Dietary constraints of phytosaurian reptiles revealed by dental microwear textural analysis

Abstract: Phytosaurs are a group of large, semi‐aquatic archosaurian reptiles from the Middle–Late Triassic. They have often been interpreted as carnivorous or piscivorous due to their large size, morphological similarity to extant crocodilians and preservation in fluvial, lacustrine and coastal deposits. However, these dietary hypotheses are difficult to test, meaning that phytosaur ecologies and their roles in Triassic food webs remain incompletely constrained. Here, we apply dental microwear textural analysis to the three‐dimensional sub‐micrometre scale tooth surface textures that form during food consumption to provide the first quantitative dietary constraints for five species of phytosaur. We furthermore explore the impacts of tooth position and cranial robustness on phytosaur microwear textures. We find subtle systematic texture differences between teeth from different positions along phytosaur tooth rows, which we interpret to be the result of different loading pressures experienced during food consumption, rather than functional partitioning of food processing along tooth rows. We find rougher microwear textures in morphologically robust taxa. This may be the result of seizing and processing larger prey items compared to those captured by gracile taxa, rather than dietary differences per se. We reveal relatively low dietary diversity between our study phytosaurs and that individual species show a lack of dietary specialization. Species are predominantly carnivorous and/or piscivorous, with two taxa exhibiting slight preferences for ‘harder’ invertebrates. Our results provide strong evidence for higher degrees of ecological convergence between phytosaurs and extant crocodilians than previously appreciated, furthering our understanding of the functioning and evolution of Triassic ecosystems.

reptile
dental microwear
tooth position
dietary constraint


Understanding modern extinctions in marine ecosystems: the role of palaeoecological data

Abstract: Because anthropogenic impacts on ecological systems pre-date the oldest scientific observations, historical documents and archaeological records, understanding modern extinctions requires additional data sources that extend further back in time. Palaeoecological records, which provide quantitative proxy records of ecosystems prior to human impact, are essential for understanding recent extinctions and future extinction risks. Here we critically review the value of the most recent fossil record in contributing to our understanding of modern extinctions and illustrate through case studies how naturally occurring death assemblages and Holocene sedimentary records provide context to the plight of marine ecosystems. While palaeoecological data are inherently restricted censuses of past communities (manipulative experiments are not possible), they yield quantitative records over temporal scales that are beyond the reach of ecology. Only by including palaeoecological data is it possible to fully assess the role of long-term anthropogenic processes in driving modern extinction risk.

modern extinction
marine ecosystem
palaeoecological data
ecology
perspective


Growth and patterning in the conodont skeleton

Abstract: Recent advances in our understanding of conodont palaeobiology and functional morphology have rendered established hypotheses of element growth untenable. In order to address this problem, hard tissue histology is reviewed paying particular attention to the relationships during growth of the component hard tissues comprising conodont elements, and ignoring a priori assumptions of the homologies of these tissues. Conodont element growth is considered further in terms of the pattern of formation, of which four distinct types are described, all possibly derived from a primitive condition after heterochronic changes in the timing of various developmental stages. It is hoped that this may provide further means of unravelling conodont phylogeny. The manner in which the tissues grew is considered homologous with other vertebrate hard tissues, and the elements appear to have grown in a way similar to the growing scales and growing dentition of other vertebrates.

tissue
conodont element
matter
lamellar crown tissue
white matter


Impact of biomineralization processes on the Mg content of foraminiferal shells: A biological perspective

Abstract: The Mg/Ca ratio in foraminiferal shells is widely used as a proxy for paleotemperatures. Nevertheless, it seems that the basic Mg content of foraminifera is determined by biological factors, as can be concluded from the large inter species and intrashell variability and the frequent deviations from inorganic behavior. This paper discusses three possible ways by which foraminifera can control or modify the Mg content in their shell: (1) involvement of organic matrix in the precipitation process that may alter the partition coefficient of Mg in biogenic calcite, (2) controlled conversion of transient amorphous phases to calcite, and (3) modification of the Mg concentration in the parent solution from which the crystals precipitate. The first two mechanisms are probably responsible for the precipitation of high‐Mg calcite phases (whole shell or sublayers), while the third mechanism leads to the formation of low‐Mg calcite phases. We propose a model adapted from epithelial cells that allows massive Mg2+ removal from the biomineralization site. This model is especially relevant to the planktonic and deep benthic low‐Mg foraminifera that are frequently used for paleotemperature reconstructions. We discuss the possible biological roles of Mg in the shell in terms of the calcite polymorph conservation, the in vivo chemical stability of the shell, the functions of Mg as a stabilizer of transient phases and as a controlling agent of the precipitation process. Several temperature sensitive biological processes that may influence the Mg/Ca ratio of the shell are suggested and a model that combines biogenic and inorganic considerations is presented. The model uses Mg heterogeneity in the shell together with temperature response (biologic and inorganic) of biomineralization processes, to account for the deviation of planktonic foraminifera from inorganic calcite at equilibrium with seawater.

biomineralization process
foraminiferal shell
deviation
precipitation


Early Triassic Marine Biotic Recovery: The Predators' Perspective

Abstract: Examining the geological past of our planet allows us to study periods of severe climatic and biological crises and recoveries, biotic and abiotic ecosystem fluctuations, and faunal and floral turnovers through time. Furthermore, the recovery dynamics of large predators provide a key for evaluation of the pattern and tempo of ecosystem recovery because predators are interpreted to react most sensitively to environmental turbulences. The end-Permian mass extinction was the most severe crisis experienced by life on Earth, and the common paradigm persists that the biotic recovery from the extinction event was unusually slow and occurred in a step-wise manner, lasting up to eight to nine million years well into the early Middle Triassic (Anisian) in the oceans, and even longer in the terrestrial realm. Here we survey the global distribution and size spectra of Early Triassic and Anisian marine predatory vertebrates (fishes, amphibians and reptiles) to elucidate the height of trophic pyramids in the aftermath of the end-Permian event. The survey of body size was done by compiling maximum standard lengths for the bony fishes and some cartilaginous fishes, and total size (estimates) for the tetrapods. The distribution and size spectra of the latter are difficult to assess because of preservation artifacts and are thus mostly discussed qualitatively. The data nevertheless demonstrate that no significant size increase of predators is observable from the Early Triassic to the Anisian, as would be expected from the prolonged and stepwise trophic recovery model. The data further indicate that marine ecosystems characterized by multiple trophic levels existed from the earliest Early Triassic onwards. However, a major change in the taxonomic composition of predatory guilds occurred less than two million years after the end-Permian extinction event, in which a transition from fish/amphibian to fish/reptile-dominated higher trophic levels within ecosystems became apparent.

mass extinction
marine reptile
early triassic
sauropterygian
thalattosaur


Diptera of the middle Eocene Kishenehn Formation. I. Documentation of diversity at the family level

Abstract: The Coal Creek Member of the Kishenehn Formation in northwestern Montana, USA, is an emerging middle Eocene Lagerstätte. While fish, plant, mammal and molluscan fossils are present, the most numerous and well-preserved fossils are those of insects. In this study, we initiate an effort to enumerate, at the family level, the diversity of flies (Insecta: Diptera) at this locality. Seventeen specimens from 17 different families (15 families with Limoniinae and Cylindrotominae within Tipulidae s.l.), 15 new species and three new genera are described. These include Tipula fji sp. nov. (Tipulidae), Ellipteroides kishenehn sp. nov. (Limoniidae), Cyttaromyia lynnae sp. nov. (Cylindrotomidae), Sylvicola silibrarius sp. nov. (Anisopodidae), Efcookella nigra sp. nov. (first fossil known in the genus) (Scatopsidae), Bibiodes kishenehnensis sp. nov. (Bibionidae), Eosciarites hermes gen. et sp. nov. (Sciaridae), Rymosia hypnolithica sp. nov. (Mycetophilidae), Litoleptis araeostylus sp. nov. (Rhagionidae), Kishenehnoasilus bhl gen. et sp. nov. (Asilidae), Drapetis adelomedos sp. nov. (Hybotidae), Salishomyia eocenica gen. et sp. nov. (Dolichopodidae), Agathomyia eocenica sp. nov. (first known fossil in genus) (Platypezidae), Lonchoptera eocenica sp. nov. (Lonchopteridae) and Aenigmatias kishenehnensis sp. nov. (Phoridae). Two specimens in the families Psychodidae and Pipunculidae are described but not assigned to a genus. In addition, we revise several related fossil species housed at the NMNH. Asilopsis fusculus Cockerell, 1921, formerly described in Asilidae, is transferred to Cyttaromyia (Cylindrotomidae) as C. fuscula, Sciara florissantensis Cockerell, 1917 is assigned to Sciaroidea incertae sedis, and Sciara gurnetensis Cockerell, 1916, Sciara lacoei Cockerell, 1915 and Sciara protoberidis Cockerell, 1915, are assigned to Sciaridae incertae sedis. Given their diversity and high degree of preservation, continued characterization of the Coal Creek Member fossils may help elucidate the Eocene radiation of Diptera in North America. GREENWALT ET AL.: KISHENEHN FORMATION DIPTERA 2 Dale E. Greenwalt. Department of Paleobiology, National Museum of Natural History MRC 121, Smithsonian Institution, 10th & Constitution Ave. NW, Washington, D.C. 20013-7012, USA. GreenwaltD@si.edu Daniel J. Bickel. Entomology, Australian Museum, 1 William Street, Sydney NSW 2010, Australia. dan.bickel@austmus.gov.au Peter H. Kerr. Plant Pest Diagnostics Branch, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, California 95832-1448, USA. pkerr@cdfa.ca.gov Gregory R. Curler. Mississippi Entomological Museum, Mississippi State University, 100 Old Highway 12, Box 9775, Mississippi 39762-9775, USA. gcurler@gmail.com Brian V. Brown. Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA. bbrown@nhm.org Herman de Jong. Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands. Herman.deJong@naturalis.nl Scott J. Fitzgerald. Pacific Northwest Diptera Research Lab, 1460 SW Allen St., Corvallis, Oregon, 97333, USA. woodyfitz@gmail.com Torsten Dikow, Smithsonian Institution, National Museum of Natural History, 10th & Constitution Ave. NW, Washington, DC 20560-0169, USA. dikowt@si.edu Michal Tkoč. Department of Entomology, National Museum, Cirkusová 1740, CZ-193 00 Praha 9 – Horní Počernice, Czech Republic. michaltkoc@gmail.com Christian Kehlmaier. Senckenberg Natural History Collections Dresden, Museum of Zoology, Königsbrücker Landstrasse 159, 01109 Dresden, Germany. kehlmaier@web.de Dalton De Souza Amorim. Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Av. Bandeirantes, 3900, 14040-901, Ribeirão Preto, SP, Brazil. dsamorim@usp.br

kishenehn formation
family level
northwestern montana
diptera


Rodents of the Caribbean: origin and diversification of hutias unravelled by next-generation museomics

Abstract: The Capromyidae (hutias) are endemic rodents of the Caribbean and represent a model of dispersal for non-flying mammals in the Greater Antilles. This family has experienced severe extinctions during the Holocene and its phylogenetic affinities with respect to other caviomorph relatives are still debated as morphological and molecular data disagree. We used target enrichment and next-generation sequencing of mitochondrial and nuclear genes to infer the phylogenetic relationships of hutias, estimate their divergence ages, and understand their mode of dispersal in the Greater Antilles. We found that Capromyidae are nested within Echimyidae (spiny rats) and should be considered a subfamily thereof. We estimated that the split between hutias and Atlantic Forest spiny rats occurred 16.5 (14.8–18.2) million years ago (Ma), which is more recent than the GAARlandia land bridge hypothesis (34–35 Ma). This would suggest that during the Early Miocene, an echimyid-like ancestor colonized the Greater Antilles from an eastern South American source population via rafting. The basal divergence of the Hispaniolan Plagiodontia provides further support for a vicariant separation between Hispaniolan and western islands (Bahamas, Cuba, Jamaica) hutias. Recent divergences among these western hutias suggest Plio-Pleistocene dispersal waves associated with glacial cycles.


The origin of pterygotid eurypterids (Chelicerata: Eurypterida)

Abstract: Abstract:  The Silurian and Lower Devonian pterygotid eurypterids were the largest of all arthropods, but their origin and early evolution are poorly understood. Pterygotus? ventricosus from Kokomo, Indiana, is shown to be the most basal of the pterygotids and constitutes the sister taxon to the rest of the clade. P.? ventricosus has walking appendages similar to those of Slimonia, which is the sister taxon of pterygotids. There is no evidence in P.? ventricosus of the dorsal median pretelson carina characteristic of other pterygotids, but the taxon does have the typical pterygotid undivided genital appendage. The status of the iconic pterygotid character (and presumed synapomorphy), the enlargement of the chelicerae, is uncertain in the Kokomo pterygotid. All the diagnostic characters of pterygotids did not develop simultaneously. A new genus Ciurcopterus is erected to accommodate the Kokomo pterygotid and Pterygotus? sarlei Ciurca and Tetlie.

pterygotid eurypterid
silurian
chelicerae


Temporal Patterns of Nucleotide Misincorporations and DNA Fragmentation in Ancient DNA

Abstract: DNA that survives in museum specimens, bones and other tissues recovered by archaeologists is invariably fragmented and chemically modified. The extent to which such modifications accumulate over time is largely unknown but could potentially be used to differentiate between endogenous old DNA and present-day DNA contaminating specimens and experiments. Here we examine mitochondrial DNA sequences from tissue remains that vary in age between 18 and 60,000 years with respect to three molecular features: fragment length, base composition at strand breaks, and apparent C to T substitutions. We find that fragment length does not decrease consistently over time and that strand breaks occur preferentially before purine residues by what may be at least two different molecular mechanisms that are not yet understood. In contrast, the frequency of apparent C to T substitutions towards the 5′-ends of molecules tends to increase over time. These nucleotide misincorporations are thus a useful tool to distinguish recent from ancient DNA sources in specimens that have not been subjected to unusual or harsh treatments.

dna fragment
ancient dna
fragment length
frequency
cytosine deamination


Evolution of morphological disparity in pterosaurs

Abstract: Pterosaurs were important flying vertebrates for most of the Mesozoic, from the Late Triassic to the end of the Cretaceous (225–65 Ma). They varied enormously through time in overall size (with wing spans from about 250 mm to about 12 m), and in features of their cranial and postcranial skeletons. Comparisons of disparity based on discrete cladistic characters show that the basal paraphyletic rhamphorhynchoids (Triassic–Early Cretaceous) occupied a distinct, and relatively small, region of morphospace compared to the derived pterodactyloids (Late Jurassic–Late Cretaceous). This separation is unexpected, especially in view of common constraints on anatomy caused by the requirements of flight. Pterodactyloid disparity shifted through time, with different, small portions of morphospace occupied in the Late Jurassic and Late Cretaceous, and a much larger portion in the Early Cretaceous. This explosion in disparity after 100 Ma of evolution is matched by the highest diversity of the clade: evidently, pterosaurs express a rather ‘top heavy’ clade shape, and this is reflected in delayed morphological evolution, again an unexpected finding. The expansion of disparity among pterodactyloids was comparable across subclades: pairwise comparisons among the four pterodactyloid superfamilies show that, for the most part, these clades display significant morphological separation, except in the case of Dsungaripteroidea and Azhdarchoidea. Finally, there is no evidence that rhamphorhynchoids as a whole were outcompeted by pterodactyloids, or that pterosaurs were driven to extinction by the rise of birds.


Morphological convergence obscures functional diversity in sabre-toothed carnivores

Abstract: The acquisition of elongated, sabre-like canines in multiple vertebrate clades during the last 265 Myr represents a remarkable example for convergent evolution. Due to striking superficial similarities in the cranial skeleton, the same or similar skull and jaw functions have been inferred for sabre-toothed species and interpreted as an adaptation to subdue large-bodied prey. However, although some sabre-tooth lineages have been classified into different ecomorphs (dirk-tooths and scimitar-tooths) the functional diversity within and between groups and the evolutionary paths leading to these specializations are unknown. Here, we use a suite of biomechanical simulations to analyse key functional parameters (mandibular gape angle, bending strength, bite force) to compare the functional performance of different groups and to quantify evolutionary rates across sabre-tooth vertebrates. Our results demonstrate a remarkably high functional diversity between sabre-tooth lineages and that different cranial function and prey killing strategies evolved within clades. Moreover, different biomechanical adaptations in coexisting sabre-tooth species further suggest that this functional diversity was at least partially driven by niche partitioning.


Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum.

Abstract: The aims of this paper are to review previously published palaeovegetation and independent palaeoclimatic datasets together with new results we present from dynamic vegetation model simulations and modern pollen rain studies to: (i) determine the responses of Amazonian ecosystems to changes in temperature, precipitation and atmospheric CO2 concentrations that occurred since the last glacial maximum (LGM), ca. 21 000 years ago; and (ii) use this long-term perspective to predict the likely vegetation responses to future climate change. Amazonia remained predominantly forested at the LGM, although the combination of reduced temperatures, precipitation and atmospheric CO2 concentrations resulted in forests structurally and floristically quite different from those of today. Cold-adapted Andean taxa mixed with rainforest taxa in central areas, while dry forest species and lianas probably became important in the more seasonal southern Amazon forests and savannahs expanded at forest-savannah ecotones. Net primary productivity (NPP) and canopy density were significantly lower than today. Evergreen rainforest distribution and NPP increased during the glacial-Holocene transition owing to ameliorating climatic and CO2 conditions. However, reduced precipitation in the Early-Mid-Holocene (ca. 8000-3600 years ago) caused widespread, frequent fires in seasonal southern Amazonia, causing increased abundance of drought-tolerant dry forest taxa and savannahs in ecotonal areas. Rainforests expanded once more in the Late Holocene owing to increased precipitation caused by greater austral summer insolation, although some of this forest expansion (e.g. in parts of the Bolivian Beni) is clearly caused by palaeo Indian landscape modification. The plant communities that existed during the Early-Mid-Holocene may provide insights into the kinds of vegetation response expected from similar increases in temperature and aridity predicted for the twenty-first century. We infer that ecotonal areas near the margins of the Amazon Basin are liable to be most sensitive to future environmental change and should therefore be targeted with conservation strategies that allow 'natural' species movements and plant community re-assortments to occur.

amazonian ecosystem
glacial maximum
lgm
forest


Axial growth gradients across the postprotaspid ontogeny of the Silurian trilobite Aulacopleura koninckii

Abstract:
Abstract.
 Recent morphometric analysis revealed a juvenile (meraspid) axial growth gradient in the trunk of the ∼429 Myr old trilobite Aulacopleura koninckii that resulted from growth control based on positional specification, as is common among extant organisms. Here we explore axial growth gradients in the more anterior body region, the cephalon, and in the cephalon and trunk during subsequent development in the holaspid period. We detected an axial growth gradient in the cephalon in the meraspid period, flatter and opposite in direction to that of the trunk, which also persisted during the holaspid period. We also found an holaspid trunk growth gradient, with a different distribution of growth rates among segments than that of the meraspid period. These newly observed growth gradients are compatible with the mechanism of growth control inferred for the meraspid trunk. Thus, the same kind of growth control may have operated in both body regions and during thewhole ontogeny of A. koninckii. This study, alongwith others on the same species that preceded it, show that morphometric analysis of appropriate data sets can address questions of high interest for evolutionary developmental biology using data fromfossils. By revealing developmental features at deep nodes of the phylogenetic tree, these studies will elucidate both how developmental processes evolved and how they themselves affected the evolution of organismal body patterning.

cephalon
meraspid period
axial growth gradient


The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)

Abstract: The extinct dromornithids, gastornithids and phorusrhacids are among the most spectacular birds to have ever lived, with some giants exceeding 500 kg. The affinities and evolution of these and other related extinct birds remain contentious, with previous phylogenetic analyses being affected by widespread convergence and limited taxon sampling. We address these problems using both parsimony and tip-dated Bayesian approaches on an expansive taxon set that includes all key extinct flightless and flighted (e.g. Vegavis and lithornithids) forms, an extensive array of extant fowl (Galloanseres), representative Neoaves and palaeognaths. The Paleogene volant Lithornithidae are recovered as stem palaeognaths in the Bayesian analyses. The Galloanseres comprise four clades inferred to have diverged in the Late Cretaceous on Gondwana. In addition to Anseriformes and Galliformes, we recognize a robust new clade (Gastornithiformes) for the giant flightless Dromornithidae (Australia) and Gastornithidae (Eurasia, North America). This clade exhibits parallels to ratite palaeognaths in that flight presumably was lost and giant size attained multiple times. A fourth clade is represented by the Cretaceous Vegavis (Antarctica), which was strongly excluded from Anseriformes; thus, a crucial molecular calibration point needs to be reconsidered. The presbyornithids Wilaru (Australia) and Presbyornis (Northern Hemisphere) are robustly found to be the sister group to Anatoidea (Anseranatidae + Anatidae), a relatively more basal position than hitherto recognized. South America's largest bird, Brontornis, is not a galloansere, but a member of Neoaves related to Cariamiformes; therefore, giant Galloanseres remain unknown from this continent. Trait analyses showed that while gigantism and flightlessness evolved repeatedly in groups, diet is constrained by phylogeny: all giant Galloanseres and palaeognaths are herbivores or mainly herbivorous, and giant neoavians are zoophagous or omnivorous.


The last terror birds (Aves, Phorusrhacidae): new evidence from the late Pleistocene of Uruguay

Abstract:
We report new fossil evidence of terror bird survival until the end of the Pleistocene in Uruguay. The new specimens comprise the distal portion of right tarsometatarsus and a left humerus; the latter is assigned to the genus Psilopterus. The sedimentary context of the remains yields a characteristic Pleistocene mammalian association along with numerical age dating giving an undoubted late Pleistocene age (OSL 96,040 ± 6300 years). We also revise and discuss the systematic placement of late Pleistocene phorusrhacid material previously published. The trophic role of terror birds and other South American carnivorous birds in late Pleistocene ecosystems should be revised based on the increasing findings of avianfossil materials.
Phorusrhacids • Psilopterinae • Late Pleistocene • Uruguay Kurzfassung Wir berichten von neuen fossilen Beweisen für das Überleben von Terrorvögeln bis zum Ende des Pleistozäns in Uruguay. Die neuen Exemplare umfassen den distalen Teil des rechten Tarsometatarsus sowie eines linken Humerus; letzterer wird der Gattung Psilopterus zugeordnet. Der sedimentäre Kontext dieser Reste liefert eine charakteristische pleistozäne Säugetiergemeinschaft und zusammen mit numerischer Altersdatierung ein unzweifelhaft spät-pleistozänes Alter (OSL 96.040 ± 6.300 Jahre). Wir revidieren und diskutieren außerdem die systematische Stellung des bisher publizierten spätpleistozänen Phorusrhaciden-Materials. Die trophische Rolle von Terrorvögeln und anderen südamerikanischen carnivoren Vögeln im Ökosystem des Spät-Pleistozäns sollte, basierend auf der Zunahme der Funde von Vogel-Fossilmaterial, revidiert werden.

terror bird
pleistocene
uruguay
carnivorous bird


PALEONTOLOGY, PHYLOGENY, AND CLASSIFICATION: AN EXAMPLE FROM THE MAMMALIAN FOSSIL RECORD

Abstract: Gingerich, P. D. (Museum of Paleontology, The University of Michigan, Ann Arbor, Michigan 48109) 1979. Paleontology, phylogeny, and classification: an example from the mammalian fossil records. Syst. Zool. 28:451-464.-The purpose of a general formal classification of animals is to foster effective communication within and between zoological subdisciplines. Cladistics is a method of classification based only on phylogeny, but knowledge of phylogeny requires a relatively dense and continuous fossil record not available for many groups. Cladistics presumes phylogeny, and it cannot be used to reconstruct phylogeny. Furthermore, phylogenetic relationships are not the only attributes of organisms, and classifications based solely on phylogeny ignore important attributes related to adaption, biogeography, and age. Effective communication requires that a classification be as simple as possible, balanced, and relatively stable. In practice, organisms are grouped into species, and species are grouped into successively higher taxa primarily on the basis of (1) overall similarity (including morphology, geography, and age), with the boundaries between taxa being drawn in such a way that (2) each group is monophyletic in the general sense of this term, with (3) traditional groupings being maintained to the maximum extent consistent with (1) and (2). Classification is basically phenetic, with phylogeny (where known) providing an important constraint on acceptable phenetic arrangements. Traditional and cladistic classifications of North American Eocene primates of the family Adapidae are compared, with a slightly modified traditional classification being favored because of its simplicity, balance, and stability. Phylogeny itself is better expressed in a diagram than in a formal classification. [Classification; phylogeny; stratophenetics; cladistics; Adapidae.] Classification occupies a curious position in zoology. The most interesting aspects of animal biology are develop

phylogeny
mammalian fossil record
paleontology


A reassessment of the cranial morphology of Neoepiblema acreensis (Rodentia: Chinchilloidea), a Miocene rodent from South America

Abstract: The rodent Neoepiblema acreensis (Chinchilloidea: Neoepiblemidae) is member of a lineage that reached gigantic dimensions during the Late Miocene of South America—the Neoepiblemidae. In this paper, the cranial anatomy of this rodent is reviewed. Noninvasive imaging is used to reveal internal structures. Our review is based mainly on an almost complete cranium from the Upper Miocene deposits of the western Amazonia of Brazil. The cranium has an elongated rostrum, large frontal sinuses, a deep temporal fossa, well‐developed sagittal, nuchal, medial occipital, and secondary crests, and a tympanic fenestra connected to the external acoustic meatus by a thin ventral cleft. Remarkably, the cranium shows the presence of fossae on the posterior region of the frontal and parietal bones, and a “W‐shaped” fronto‐parietal suture, which are not present in other analyzed chinchilloids. This study contributes to the knowledge of the morphology of this extinct rodent as well as to the phylogenetic relationships and paleobiology of neoepiblemids.

morphology
neoepiblema acreensis
chinchilloidea


Breeding Young as a Survival Strategy during Earth’s Greatest Mass Extinction

Abstract: Studies of the effects of mass extinctions on ancient ecosystems have focused on changes in taxic diversity, morphological disparity, abundance, behaviour and resource availability as key determinants of group survival. Crucially, the contribution of life history traits to survival during terrestrial mass extinctions has not been investigated, despite the critical role of such traits for population viability. We use bone microstructure and body size data to investigate the palaeoecological implications of changes in life history strategies in the therapsid forerunners of mammals before and after the Permo-Triassic Mass Extinction (PTME), the most catastrophic crisis in Phanerozoic history. Our results are consistent with truncated development, shortened life expectancies, elevated mortality rates and higher extinction risks amongst post-extinction species. Various simulations of ecological dynamics indicate that an earlier onset of reproduction leading to shortened generation times could explain the persistence of therapsids in the unpredictable, resource-limited Early Triassic environments, and help explain observed body size distributions of some disaster taxa (e.g., Lystrosaurus). Our study accounts for differential survival in mammal ancestors after the PTME and provides a methodological framework for quantifying survival strategies in other vertebrates during major biotic crises.

survival strategy
lystrosaurus
early onset


The oldest record of ornithuromorpha from the early cretaceous of China

Abstract: Ornithuromorpha is the most inclusive clade containing extant birds but not the Mesozoic Enantiornithes. The early evolutionary history of this avian clade has been advanced with recent discoveries from Cretaceous deposits, indicating that Ornithuromorpha and Enantiornithes are the two major avian groups in Mesozoic. Here we report on a new ornithuromorph bird, Archaeornithura meemannae gen. et sp. nov., from the second oldest avian-bearing deposits (130.7 Ma) in the world. The new taxon is referable to the Hongshanornithidae and constitutes the oldest record of the Ornithuromorpha. However, A. meemannae shows few primitive features relative to younger hongshanornithids and is deeply nested within the Hongshanornithidae, suggesting that this clade is already well established. The new discovery extends the record of Ornithuromorpha by five to six million years, which in turn pushes back the divergence times of early avian lingeages into the Early Cretaceous.

ornithuromorpha
early cretaceous
old record
furcula


A new developmental mechanism for the separation of the mammalian middle ear ossicles from the jaw

Abstract: Multiple mammalian lineages independently evolved a definitive mammalian middle ear (DMME) through breakdown of Meckel's cartilage (MC). However, the cellular and molecular drivers of this evolutionary transition remain unknown for most mammal groups. Here, we identify such drivers in the living marsupial opossum Monodelphis domestica, whose MC transformation during development anatomically mirrors the evolutionary transformation observed in fossils. Specifically, we link increases in cellular apoptosis and TGF-BR2 signalling to MC breakdown in opossums. We demonstrate that a simple change in TGF-β signalling is sufficient to inhibit MC breakdown during opossum development, indicating that changes in TGF-β signalling might be key during mammalian evolution. Furthermore, the apoptosis that we observe during opossum MC breakdown does not seemingly occur in mouse, consistent with homoplastic DMME evolution in the marsupial and placental lineages.

mammalian middle ear
jaw
cartilage
mandible


Triggering of the largest Deccan eruptions by the Chicxulub impact

Abstract: New constraints on the timing of the Cretaceous-Paleogene mass extinction and the Chicxulub impact, together with a particularly voluminous and apparently brief eruptive pulse toward the end of the “main-stage” eruptions of the Deccan continental fl ood basalt province suggest that these three events may have occurred within less than about a hundred thousand years of each other. Partial melting induced by the Chicxulub event does not provide an energetically plausible explanation for this coincidence, and both geochronologic and magnetic-polarity data show that Deccan volcanism was under way well before Chicxulub/Cretaceous-Paleogene time. However, historical data document that eruptions from existing volcanic systems can be triggered by earthquakes. Seismic modeling of the ground motion due to the Chicxulub impact suggests that the impact could have generated seismic energy densities of order 0.1–1.0 J/m 3 throughout the upper ~200 km of Earth’s mantle, suffi cient to trigger volcanic eruptions worldwide based upon comparison with historical examples. Triggering may have been caused by a transient increase in the effective permeability of the existing deep magmatic system beneath the Deccan province, or mantle plume “head.” It is therefore reasonable to hypothesize that the Chicxulub impact might have triggered the enormous Poladpur, Ambenali, and Mahabaleshwar (Wai Subgroup) lava fl ows, which together may account for >70% of the Deccan Traps main-stage eruptions. This hypothesis is consistent with independent stratigraphic, geochronologic, geochemical, and tectonic constraints, which combine to indicate that at approximately Chicxulub/Cretaceous-Paleogene time, a huge pulse of mantle plume–derived magma passed through the crust with little interaction and erupted to form the most extensive and voluminous lava fl ows known on Earth. High-precision radioisotopic dating of the main-phase Deccan fl ood basalt formations may be able either to confi rm or reject this hypothesis, which in turn might help to determine whether this singular outburst within the Deccan Traps (and possibly volcanic eruptions worldwide) contributed signifi cantly to the CretaceousPaleogene extinction.


Spatiotemporal variability in the δ18O-salinity relationship of seawater across the tropical Pacific Ocean.

Abstract: The relationship between salinity and the stable oxygen isotope ratio of seawater (δ18Osw) is of utmost importance to the quantitative reconstruction of past changes in salinity from δ18O values of marine carbonates. This relationship is often considered to be uniform across water masses, but the constancy of the δ18Osw-salinity relationship across space and time remains uncertain, as δ18Osw responds to varying atmospheric vapor sources and pathways, while salinity does not. Here we present new δ18Osw-salinity data from sites spanning the tropical Pacific Ocean. New data from Palau, Papua New Guinea, Kiritimati, and Galápagos show slopes ranging from 0.09 ‰/psu in the Galápagos to 0.32‰/psu in Palau. The slope of the δ18Osw-salinity relationship is higher in the western tropical Pacific versus the eastern tropical Pacific in observations and in two isotope-enabled climate models. A comparison of δ18Osw-salinity relationships derived from short-term spatial surveys and multi-year time series at Papua New Guinea and Galápagos suggests spatial relationships can be substituted for temporal relationships at these sites, at least within the time period of the investigation. However, the δ18Osw-salinity relationship varied temporally at Palau, likely in response to water mass changes associated with interannual El Niño-Southern Oscillation (ENSO) variability, suggesting nonstationarity in this local δ18Osw-salinity relationship. Applying local δ18Osw-salinity relationships in a coral δ18O forward model shows that using a constant, basin-wide δ18Osw-salinity slope can both overestimate and underestimate the contribution of δ18Osw to carbonate δ18O variance at individual sites in the western tropical Pacific.

variability
tropical pacific
pacific ocean


Shape, Drag, and Power in Ammonoid Swimming

Abstract: This study assesses swimming potential in a variety of ammonoid shell shapes on the basis of coefficients of drag (Cd) and the power needed to maintain a constant velocity. Reynolds numbers (Re) relevant to swimming ammonoids, and lower than those previously studied, were examined. Power consumption was scaled to a range of sizes and swimming velocities. Estimates of power available derived from studies of oxygen consumption in modern cephalopods and fish were used to calculate maximum sustainable swimming velocities (MSV). Laterally compressed, small thickness ratio (t. r.) ammonoids, previously assumed to be the most efficient swimmers, do not experience the lowest drag or power consumption at all sizes and velocities. At low values of size and velocity associated with Reynolds numbers below 104, less compressed forms have smaller drag coefficients and reduced power requirements. At hatching a roughly spherical shell shape would have minimized drag in ammonoids; with increasing size, hydrodynamic optima shift toward compressed morphologies. The high energetic cost of ammonoid locomotion may have limited dispersal and excluded ammonoids from high current velocity environments.

drag
power
shell shape
ammonoid locomotion


Skull anatomy of Dakosaurus andiniensis (Thalattosuchia: Crocodylomorpha) and the phylogenetic position of Thalattosuchia

Abstract: Synopsis New information on Dakosaurus andiniensis from the Latest Jurassic and Early Cretaceous is reported here. One of the specimens described herein consists of an almost complete skull and lower jaw found in the uppermost levels of the Vaca Muerta Formation (Tithonian) of Neuquén Province, Argentina. The new material allows a more complete understanding and diagnosis of this form, previously known only from the fragmentary type specimen. The new remains show that D. andiniensis had an unusual morphology for a marine crocodyliform,namely a remarkably short, high snout with ziphodont dentition. This new information allows testing of the phylogenetic relationships of this taxon, which is depicted as deeply nested within Metriorhynchidae, a clade of marine crocodyliforms with derived adaptations to the marine environment (e.g. paddle‐like forelimbs, hypertrophied nasal salt glands). In particular, D. andiniensis is inferred to be the sister taxon of D. maximus from the Jurassic of Europe. This relationship repeats the phylogenetic pattern seen in other Jurassic marine crocodyliforms from South America and Europe (e.g. Geosaurus), demonstrating the close faunal relationship between these two distant marine basins. The phylogenetic analysis reported here results in a most parsimonious hypothesis that depicts Thalattosuchia nested within Neosuchia and the strength and character evidence supporting this position is presented. In addition, several characters traditionally postulated as dependent upon the longirostrine morphology are critically examined under the light of new evidence. Most of them show a character state distribution that is not strictly compatible with that of the longirostrine condition and, therefore, are interpreted as independent units of phylogenetic evidence.

dakosaurus andiniensis
thalattosuchia
early cretaceous


The anatomy, taphonomy, taxonomy and systematic affinity of Markuelia: Early Cambrian to Early Ordovician scalidophorans

Abstract: Abstract:  Markuelia is a vermiform, annulated introvertan animal known as embryonic fossils from the Lower Cambrian to Lower Ordovician. Analysis of an expanded and revised dataset for Introverta shows that the precise position of Markuelia within this clade is dependent on the taxa included. As a result, Markuelia is assigned to the scalidophoran total group to reflect uncertainty as to whether it is a stem‐scalidophoran or a stem‐priapulid. The taxonomy of the genus is revised to provide an improved taxonomic framework for material assigned to Markuelia. Five species are recognized: M. secunda Val’kov, M. hunanensis Dong and Donoghue, M. lauriei Haug et al., M. spinulifera sp. nov. and M. waloszeki sp. nov. Finally, the preservation of Markuelia is evaluated in the light of both the taphonomy of the fossil embryos themselves and the experimental taphonomy of the priapulid Priapulus caudatus, which has been proposed as both a close relative and an anatomical analogue of Markuelia.

taphonomy markuelia ordovician scalidophoran


The monophyly of Euparkeriidae (Reptilia: Archosauriformes) and the origins of Archosauria: a revision of Dorosuchus neoetus from the Mid‐Triassic of Russia

Abstract: Euparkeria capensis is resolved as the sister taxon to Archosauria in many cladistic phylogenies and provides a key outgroup which may approximate the ancestral archosaur morphology. Several other taxa have been referred to the family Euparkeriidae, but the monophyly of this taxon remains doubtful and largely untested. To test this monophyly, the archosauriform and putative euparkeriid Dorosuchus neoetus from the Mid‐Triassic of Russia is re‐examined in the light of recent work on the evolution of stem archosaurs. Dorosuchus neoetus is found to possess a number of morphological features that place it close to Archosauria, including a sigmoidal femur with a clear attachment region for the m. caudifemoralis musculature, but no unambiguous archosaurian apomorphies. Dorosuchus neoetus is included for the first time in a numerical cladistic analysis and is recovered as the sole sister taxon to Archosauria + Phytosauria. A monophyletic Euparkeriidae including D. neoetus and E. capensis is slightly less parsimonious. In addition, a mandible and pterygoid that were previously referred to D. neoetus subsequent to the original description of the species are also included separately within the phylogenetic analysis and are recovered within Archosauria, possibly raising questions as to their correct taxonomic referral. However, this phylogenetic placement is based primarily on the absence of palatal teeth, but the presence or absence of palatal teeth exhibits considerable homoplasy within Archosauriformes. Based on other aspects of their morphology, we do not reject the referral of these elements to D. neoetus.

monophyly euparkeriidae archosauria sister


Injured trilobites within a collection of dinosaurs: Using the Royal Tyrrell Museum of Palaeontology to document Cambrian predation

Abstract: Biomineralised exoskeletons of trilobites are ideal for recording abnormalities, documenting injuries from failed attacks, and understanding predation in the fossil record. The illustration of new injured Cambrian-aged trilobites is particularly important for elucidating aspects of early Paleozoic predator-prey systems and identifying groups that were targets of this early predation. We document six new abnormal trilobite specimens from the Royal Tyrrell Museum of Palaeontology, including the first evidence for injuries on the corynexochid trilobite Hemirhodon amplipyge. These new data highlight the importance of museum collections in uncovering unique and informative specimens for studies of predation on early animals. Russell D. C. Bicknell. Palaeoscience Research Centre, School of Environmental and Rural Science, University of New England, Armidale, New South Wales, 2351, Australia. rdcbicknell@gmail.com Brayden Holland. Palaeoscience Research Centre, School of Environmental and Rural Science, University of New England, Armidale, New South Wales, 2351, Australia. blongle2@myune.edu.au

royal tyrrell museum
palaeontology
predation
injury


The Braincase of the Basal Sauropod Dinosaur Spinophorosaurus and 3D Reconstructions of the Cranial Endocast and Inner Ear

Abstract: Background Sauropod dinosaurs were the largest animals ever to walk on land, and, as a result, the evolution of their remarkable adaptations has been of great interest. The braincase is of particular interest because it houses the brain and inner ear. However, only a few studies of these structures in sauropods are available to date. Because of the phylogenetic position of Spinophorosaurus nigerensis as a basal eusauropod, the braincase has the potential to provide key evidence on the evolutionary transition relative to other dinosaurs. Methodology/Principal Findings The only known braincase of Spinophorosaurus (‘Argiles de l'Irhazer’, Irhazer Group; Agadez region, Niger) differs significantly from those of the Jurassic sauropods examined, except potentially for Atlasaurus imelakei (Tilougguit Formation, Morocco). The basisphenoids of Spinophorosaurus and Atlasaurus bear basipterygoid processes that are comparable in being directed strongly caudally. The Spinophorosaurus specimen was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. The endocast resembles that of most other sauropods in having well-marked pontine and cerebral flexures, a large and oblong pituitary fossa, and in having the brain structure obscured by the former existence of relatively thick meninges and dural venous sinuses. The labyrinth is characterized by long and proportionally slender semicircular canals. This condition recalls, in particular, that of the basal non-sauropod sauropodomorph Massospondylus and the basal titanosauriform Giraffatitan. Conclusions/Significance Spinophorosaurus has a moderately derived paleoneuroanatomical pattern. In contrast to what might be expected early within a lineage leading to plant-eating graviportal quadrupeds, Spinophorosaurus and other (but not all) sauropodomorphs show no reduction of the vestibular apparatus of the inner ear. This character-state is possibly a primitive retention in Spinophorosaurus, but due the scarcity of data it remains unclear whether it is also the case in the various later sauropods in which it is present or whether it has developed homoplastically in these taxa. Any interpretations remain tentative pending the more comprehensive quantitative analysis underway, but the size and morphology of the labyrinth of sauropodomorphs may be related to neck length and mobility, among other factors.

braincase
spinophorosaurus
cranial endocast
inner ear


New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography

Abstract: Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian–Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

sauropod
cretaceous dinosaur palaeobiogeography
australia
palaeolatitude
dorsal vertebrae


Estimating Latest Cretaceous and Tertiary atmospheric CO2 from stromatal indices

Abstract: A quantitative understanding of the levels of atmospheric CO2 in the geologic past sheds light on the operation of the carbon cycle and the biosphere, and aids in the prediction of future climate change. Here I present a CO2 reconstruction for the very latest Cretaceous to early Eocene and middle Miocene based on the stomatal distributions in fossil and extant Ginkgo and Metasequoia cuticles. Although both of these intervals are representative of globally warm climates, my CO2 reconstruction indicates near present-day values (300–450 ppmV) for both times. Although these data do not cast doubt on the theory of the greenhouse effect, they do suggest that other thermal forcings were more important during these intervals than they are today. 79 Royer, D.L., 2003, Estimating latest Cretaceous and Tertiary atmospheric CO2 from stomatal indices, in Wing, S.L., Gingerich, P.D., Schmitz, B., and Thomas, E., eds., Causes and Consequences of Globally Warm Climates in the Early Paleogene: Boulder, Colorado, Geological Society of America Special Paper 369, p. 79–93. © 2003 Geological Society of America. *E-mail: droyer@psu.edu. Present address: Department of Geosciences, Pennsylvania State University, University Park, Pennsylvania 16802, USA. Royer, 2001). Stomatal index (SI) is the percentage of epidermal cells that are stomatal complexes (guard cells + stomatal pore), and is defined as the following (Salisbury, 1927): SD SI = ________ × 100 SD + ED where SD = stomatal density (mm–2) and ED = epidermal cell density (mm–2). The mechanism(s) linking CO2 to SI is not fully understood, but probably involves the strong selective pressure in plants to maximize carbon fixation per unit of water transpired (water-use efficiency) (e.g., Woodward, 1987). For example, in a rising CO2 regime plants can increase their wateruse efficiency by reducing their stomatal conductance, thereby reducing water loss. Stomatal conductance is largely a function of stomatal pore area, which in turn is controlled by the size of individual stomatal pores and the density of those pores. On the time scale of several growing seasons or longer, stomatal density is typically more sensitive to changes in atmospheric CO2 than individual pore size (e.g., Royer, 2001). Furthermore, on multimillion year time scales stomatal density and SI across all plant taxa inversely respond to CO2 (Beerling and Woodward, 1997; McElwain, 1998; Royer, 2001). In contrast to stomatal density, SI is area independent, which normalizes for the effects of cell expansion. Stomatal index is essentially a measure of the number of stomatal complexes that develop per unit number of epidermal cells. Plant water-potential strongly influences cell expansion (e.g., water stressed plants generally reduce the size of their epidermal cells [Royer, 2001]), but not the rate of stomatal initialization. Thus, changes in a plant’s water budget can affect stomatal density, but not stomatal index. Both experimental and field studies show that SI is largely independent of water potential, irradiance, and temperature, and is primarily a function of CO2 (e.g., Kürschner et al., 1998; Beerling, 1999; Royer, 2001). Stomatal indices (and stomatal densities) have been experimentally shown to respond to the partial pressure of CO2, not concentration (as expressed as mole fraction, e.g., parts per million) (Woodward and Bazzaz, 1988). This finding is corroborated by a positive correlation in many plants between elevation and stomatal density (Körner and Cochrane, 1985; Woodward, 1986; Woodward and Bazzaz, 1988; Beerling et al., 1992) and SI (Rundgren and Beerling, 1999). It is important, therefore, to control for elevation when estimating CO2 from stomatal properties. The relationship between elevation and partial pressure in the lower atmosphere is roughly as follows (linear fit to equation 11.8 in Jones, 1992, p. 308):

tertiary atmospheric co2
stomatal index
index


Oceanic oxygenation events in the anoxic Ediacaran ocean

Abstract: The ocean‐atmosphere system is typically envisioned to have gone through a unidirectional oxygenation with significant oxygen increases in the earliest (ca. 635 Ma), middle (ca. 580 Ma), or late (ca. 560 Ma) Ediacaran Period. However, temporally discontinuous geochemical data and the patchy metazoan fossil record have been inadequate to chart the details of Ediacaran ocean oxygenation, raising fundamental debates about the timing of ocean oxygenation, its purported unidirectional rise, and its causal relationship, if any, with the evolution of early animal life. To better understand the Ediacaran ocean redox evolution, we have conducted a multi‐proxy paleoredox study of a relatively continuous, deep‐water section in South China that was paleogeographically connected with the open ocean. Iron speciation and pyrite morphology indicate locally euxinic (anoxic and sulfidic) environments throughout the Ediacaran in this section. In the same rocks, redox sensitive element enrichments and sulfur isotope data provide evidence for multiple oceanic oxygenation events (OOEs) in a predominantly anoxic global Ediacaran–early Cambrian ocean. This dynamic redox landscape contrasts with a recent view of a redox‐static Ediacaran ocean without significant change in oxygen content. The duration of the Ediacaran OOEs may be comparable to those of the oceanic anoxic events (OAEs) in otherwise well‐oxygenated Phanerozoic oceans. Anoxic events caused mass extinctions followed by fast recovery in biologically diversified Phanerozoic oceans. In contrast, oxygenation events in otherwise ecologically monotonous anoxic Ediacaran–early Cambrian oceans may have stimulated biotic innovations followed by prolonged evolutionary stasis.

oxygenation event
anoxic
ediacaran ocean
oceanic oxygenation event


Craniodental and humeral morphology of a new species of Masrasector (Teratodontinae, Hyaenodonta, Placentalia) from the late Eocene of Egypt and locomotor diversity in hyaenodonts

Abstract: Hyaenodonta is a diverse clade of carnivorous mammals that were part of terrestrial faunas in the Paleogene of Eurasia and North America, but the oldest record for the group is Afro-Arabian, making the record there vital for understanding the evolution of this wide-spread group. Previous studies show an ancient split between two major clades of hyaenodonts that converged in hypercarnivory: Hyainailourinae and Hyaenodontinae. These clades are each supported by cranial characters. Phylogenetic analyses of hyaenodonts also support the monophyly of Teratodontinae, an Afro-Arabian clade of mesocarnivorous to hypercarnivorous hyaenodonts. Unfortunately, the cranial anatomy of teratodontines is poorly known, and aligning the clade with other lineages has been difficult. Here, a new species of the phylogenetically controversial teratodontine Masrasector is described from Locality 41 (latest Priabonian, late Eocene) from the Fayum Depression, Egypt. The hypodigm includes the most complete remains of a Paleogene teratodontine, including largely complete crania, multiple dentaries, and isolated humeri. Standard and “tip-dating” Bayesian analyses of a character-taxon matrix that samples cranial, postcranial, and dental characters support a monophyletic Masrasector within Teratodontinae, which is consistently placed as a close sister group of Hyainailouridae. The cranial morphology of Masrasector provides new support for an expanded Hyainailouroidea (Teratodontinae + Hyainailouridae), particularly characters of the nuchal crest, palate, and basicranium. A discriminant function analysis was performed using measurements of the distal humerus from a diverse sample of extant carnivorans to infer the locomotor habits of Masrasector. Masrasector was assigned to the “terrestrial” locomotor category, a result consistent with the well-defined medial trochlear ridges, and moderately developed supinator crests of the specimens. Masrasector appears to have been a fast-moving terrestrial form with a diverse diet. These specimens considerably improve our understanding of Teratodontinae, an ancient member of the Afro-Arabian mammalian fauna, and our understanding of hyaenodont diversity before the dispersal of Carnivora to the continent near the end of the Paleogene.

morphology
hyaenodonta
late eocene
phylogenetic analysis


Intraspecific variability through ontogeny in early ammonoids

Abstract: Abstract Mollusks in general and ammonoids in particular are known to display a sometimes profound morphological intraspecific variability of their shell. Although this phenomenon is of greatest importance, it has rarely been investigated and quantified. It is especially crucial for taxonomy and incidentally for biodiversity analyses to account for it, because otherwise, the number of described species might exceed that of actual species within any group. Early ammonoids (Early Devonian, Paleozoic) typically suffer from this bias. For instance, most specimens from the same layer and the same region (e.g., the Erbenoceras beds of the Moroccan eastern Anti-Atlas studied here) differ morphologically from each other. Depending on the importance given to certain morphological characters, therefore, one could create a new species for almost every specimen. In this study, we measured nearly 100 such specimens from a restricted stratigraphic interval and quantified their intraspecific variability. There is a variable but strong overlap of the quantified shell characters at most ontogenetic stages, and only two species are here separated rather than the four previously recognized in Morocco. When ontogenetic trajectories of the Moroccan specimens are compared with coeval faunas from other regions (assigned to other species), a strong overlap between the morphospaces occupied by these taxa becomes apparent. The justification of some of these latter species is thus questionable even if their mean values in some conch parameters differ considerably from the mean values of the Moroccan species. Hence, the number of currently valid species of these loosely coiled early ammonoids is probably much too high. Extreme caution must therefore be taken when examining the diversity of groups in which the intraspecific variability is poorly known.

ontogeny
ammonoid
overlap
intraspecific variability


Breakup of Pangaea and Isolation of Relict Mammals in Australia, South America, and Madagascar

Abstract: The composition of aboriginal land mammal faunas in Australia and New Guinea (prototherians and metatherians), South America (metatherians and eutherians) and Madagascar (eutherians only) is reconsidered in light of continental drift reconstructions of Mesozoic-Tertiary world paleogeography It is proposed that these three faunas represent successively detached samples of the evolving world mammal fauna as it existed when each of these land masses became faunally isolated from the rest of the world as a result of the progressive fragmentation of Pangaea. Isolation of aboriginal prototherians and metatherians in Australia and New Guinea may date from the Upper JurassicLower Cretaceous; isolation of aboriginal metatherians and eutherians in South America may date from the Middle Cretaceous-Upper Cretaceous; isolation of aboriginal eutherians in Madagascar may date from the Paleocene-Eocene.


Sedimentary factories and ecosystem change across the Permian-Triassic Critical Interval (P-TrCI) – insights from the Xiakou area (South China)

Abstract: The Permian-Triassic mass extinction included a potentially catastrophic decline of biodiversity, but ecosystem change across this event remains poorly characterized. Here we reconstruct sedimentary factories and ecosystem change across the Permian-Triassic Critical Interval (P-TrCI) in the Xiakou area (South China). Six microfacies (MF) were classified. The succession begins with a eukaryote-controlled carbonate factory (MF-1) that passes upward into an organomineralizationdominated carbonate factory (MF-2-3). Organic-rich marls atop these units reflect carbonate factory collapse (MF-4). Organomineralization-driven carbonate formation restarts prior to the Permian-Triassic boundary (MF-5) and subsequently develops into a mixed carbonate factory where organomineralization and biomineralization are almost equally important (MF-6). MF-1 reflects oxygenated shallow water environments. In contrast, MF-2-6 were all deposited in somewhat deeper environments, some of which episodically exhibited elevated salinities, oxygen depletion, and, possibly, euxinic conditions. Our results demonstrate that distinct changes in carbonate production styles, biodiversity, and environmental conditions are not synchronous at Xiakou. Furthermore, the Xiakou record is strikingly different to that of other localities, even from the same area (e.g., the Global Stratotype Section and Point section at Meishan). Together, these findings highlight the enormous complexity of the P-TrCI and calls simplified views of the Permian-Triassic mass extinction into question.

ecosystem change
south china
permian-triassic boundary
sedimentary factory