Fossil freshwater sponges: taxonomy, geographic distribution, and critical review

Abstract: Sponges are one of the most ancient animal phyla with about 8850 living species and about 5000 described fossil taxa. Most sponges are marine and live at all depths of all oceans. Freshwater bodies (lakes, rivers) are inhabited only by a small minority of species, ca. 240 (< 3%) comprising the order Spongillida (Demospongiae) most of which are able to produce specialized resting bodies to survive harsh terrestrial environmental conditions. This highly disproportionate ratio of marine and freshwater sponges is even more accentuated in the field of palaeontology with rare records reported up to the Miocene (< 0.4% of all known fossil sponges). Only a few fossil taxa were correctly supported by strong and convincing taxonomic morphotraits at genus and species level, thus we provide here an overview of fossil freshwater sponges focusing on their morphotraits and distribution in time and space. Each recorded taxon is described in detail following the modern taxonomy and nomenclature. All fossil data suggest a clear trend of long term conservative morphology in the evolutionary history of Spongillida, although some traits of Recent gemmules evolved in a wide array of adaptive morpho-functional novelties. The majority of accepted fossil species belongs to the cosmopolitan family Spongillidae. The genera Oncosclera and Potamophloios of the family Potamolepidae seem to have had, in the past, a much larger geographic range than today. A synthesis of fossil taxa morphotraits is also provided in an Appendix 1.

taxonomy
fossil specie
family spongillidae


Pliensbachian gastropods from Venetian Southern Alps (Italy) and their palaeobiogeographical significance

Abstract: Abstract:  The Pliensbachian gastropods described by De Toni in 1912, coming from an isolated boulder at the foot of Mt Vedana (eastern margin of Trento Platform, Venetian Southern Alps, Italy) are revised. The fauna consists of 13 species representing nine families and eight superfamilies. Despite the low number of species, the assemblage represents the most diverse Early Jurassic gastropod fauna known for the Venetian Southern Alps. The boulder yielding the material was thought to derive from the upper part of the Early Jurassic Calcari Grigi Group, a carbonate platform unit extensively cropping out in the Mt Vedana area. The sedimentological analysis indicates a prevalently bioclastic wackstone‐floatstone, reflecting a lime‐muddy deposit undergone to an early consolidation. This and the high content of ammonoids, which is unusual for the Calcari Grigi Group, are typical aspects of a condensed pelagic sediment, presumably a fissure filling at the top of the carbonate platform succession. Palaeobiogeographical comparisons show that the fauna is composed of species occurring exclusively in pelagic limestones of the western Tethys. By contrast, it shows no relationships with the coeval faunas of the adjacent Trento platform and of the other western Tethyan carbonate platforms. These lines of evidence and the facies analysis would testify to the Pliensbachian drowning of the eastern margin of the Trento platform. In the wider context of the palaeobiogeographical history of Early Jurassic western Tethyan gastropods, the species from Vedana belong to a faunal stock which is typical for pelagic, mainly postdrowning sediments. Thus, appearance and diffusion in space and time of this stock were probably regulated by the direction, rate and pattern of the Neotethyan rifting. A new subgenus, Proarcirsa (Schafbergia) subgen. nov., and three new species, namely Ataphrus (Ataphrus) cordevolensis sp. nov., Guidonia pseudorotula sp. nov. and Proarcirsa (Schafbergia) zirettoensis sp. nov. are erected.

venetian southern alps
western tethys
neotethyan rifting
pliensbachian gastropod


Brachiopod-dominated communities and depositional environment of the Guanshan Konservat-Lagerstätte, eastern Yunnan, China

Abstract: The Guanshan Biota is an unusual early Cambrian Konservat-Lagerstätte from China and is distinguished from all other exceptionally preserved Cambrian biotas by the dominance of brachiopods and a relatively shallow depositional environment. However, the faunal composition, overturn and sedimentology associated with the Guanshan Biota are poorly understood. This study, based on collections through the best-exposed succession of the basal Wulongqing Formation at the Shijiangjun section, Wuding County, eastern Yunnan, China recovered six major animal groups with soft tissue preservation; brachiopods vastly outnumbered all other groups. Brachiopods quickly replace arthropods as the dominant fauna following a transgression at the base of the Wulongqing Formation. A transition from a botsfordiid-, eoobolid- and acrotretid- to an acrotheloid-dominated brachiopod assemblage occurs up-section. Four episodically repeated lithofacies reveal a relatively low-energy, offshore to lower shoreface sedimentary environment at the Shijiangjun section, which is very different from the Wulongqing Formation in the Malong and Kunming areas. Multiple event flows and rapid obrution are responsible for faunal overturn and fluctuation through the section. A detailed lithofacies and palaeontological investigation of this section provides a better understanding of the processes and drivers of faunal overturn during the later phase of the Cambrian Explosion. Supplementary material: Composition and comparison of the Malong Fauna and the Guanshan Biota is are available at: https://doi.org/10.6084/m9.figshare.c.5080799

depositional environment
eastern yunnan
biota


A high‐resolution diatom record of the last deglaciation from the SE Norwegian Sea: Documentation of rapid climatic changes

Abstract: Although the pulsating nature and the abruptness of the last deglaciation are well documented in marine and land records, very few marine records have so far been able to capture the high-frequency climatic changes recorded in the Greenland ice core Dye 3. We studied high-resolution sediment cores from SE Norwegian Sea, which display a detailed climatic record during the last deglaciation comparable to that of Dye 3. Accelerator mass spectrometry age control of the cores enables us to correlate this record in detail with continental records. The results indicate that the surface waters of the SE Norwegian Sea were seasonally ice free after 13,400 B.P. The Bolling/Allerod interstadial complex (13,200–11,200 B.P.) was a climatically unstable period with changing Arctic-Subarctic conditions. This period was punctuated by four progressively more severe sea surface temperature (SST) minima: between 12,900–12,800 B.P. (BCP I); 12,500–12,400 B.P. (BCP II); 12,300–12,000 B.P. (OD I); and 11,800–11,500 B.P. (OD II). The Younger Dryas (YD) (11,200–10,200 B.P.) represents the severest and most prolonged cold episode of this series of climatic deteriorations. It was bounded by very rapid SST changes and characterized by Arctic-Polar conditions. The first true warm Atlantic water incursion to the SE Norwegian Sea took place around 10,100 B.P., followed by a brief cooler condition between 9900–9600 B.P. (YD II). The early Holocene climatic optimum occurred between 8000–5000 B.P. A conceptual model is proposed where meltwater fluxes are suggested to cause the observed instability in the SST record.

last deglaciation
climatic change
younger dryas


Evolutionary History of Lagomorphs in Response to Global Environmental Change

Abstract: Although species within Lagomorpha are derived from a common ancestor, the distribution range and body size of its two extant groups, ochotonids and leporids, are quite differentiated. It is unclear what has driven their disparate evolutionary history. In this study, we compile and update all fossil records of Lagomorpha for the first time, to trace the evolutionary processes and infer their evolutionary history using mitochondrial genes, body length and distribution of extant species. We also compare the forage selection of extant species, which offers an insight into their future prospects. The earliest lagomorphs originated in Asia and later diversified in different continents. Within ochotonids, more than 20 genera occupied the period from the early Miocene to middle Miocene, whereas most of them became extinct during the transition from the Miocene to Pliocene. The peak diversity of the leporids occurred during the Miocene to Pliocene transition, while their diversity dramatically decreased in the late Quaternary. Mantel tests identified a positive correlation between body length and phylogenetic distance of lagomorphs. The body length of extant ochotonids shows a normal distribution, while the body length of extant leporids displays a non-normal pattern. We also find that the forage selection of extant pikas features a strong preference for C3 plants, while for the diet of leporids, more than 16% of plant species are identified as C4 (31% species are from Poaceae). The ability of several leporid species to consume C4 plants is likely to result in their size increase and range expansion, most notably in Lepus. Expansion of C4 plants in the late Miocene, the so-called ‘nature’s green revolution’, induced by global environmental change, is suggested to be one of the major ‘ecological opportunities’, which probably drove large-scale extinction and range contraction of ochotonids, but inversely promoted diversification and range expansion of leporids.

global environmental change
ochotonid
extant pikas
late miocene
evolutionary history


The stratigraphic distribution of fossils

Abstract: In several increasingly realistic steps, a model of the stratigraphic distribution of fossils is presented. The first and simplest step assumes that if a taxon was extant it will have been preserved. The second step admits that if a taxon was extant, there is some probability less than one that it will have been preserved. This step produces randomly distributed gaps, and fossil ranges that are somewhat shorter than the span of time in which a taxon actually lived. The third step assumes facies-controlled taxa and parasequence-style cyclicity. This produces randomly and nonrandomly distributed gaps, including the anomalously long gaps recognized in confidence limit studies. The final model incorporates depositional sequences and indicates that first and last occurrences will cluster at sequence boundaries and at flooding surfaces in the transgressive systems tract. Across-shelf gradients in diversity, taphonomy, or eurytopy can control the strength, but not the stratigraphic position of these peaks. Comparison of the model to data from the Upper Ordovician suggests that these modeled features are present in the fossil record. Many previously studied paleobiologic patterns may be, at least in part, an artifact of facies control and sequence architecture. Such patterns include gradual or stepwise mass extinction, punctuated morphologic and taxonomic change, iterative evolution, and the replacement of shallow water faunas by deeper water faunas at biomere boundaries.

stratigraphic distribution
probability
flooding surface


Osteology and phylogeny of Robustichthys luopingensis, the largest holostean fish in the Middle Triassic

Abstract: The extinct ray-finned fish taxon Robustichthys luopingensis from Luoping, eastern Yunnan, China represents the largest holostean known in the Middle Triassic. Despite its potential significance for investigating the holostean phylogeny and reconstructing the Triassic marine ecosystems, Robustichthys has so far not been described in detail and its phylogenetic position within the Holostei was controversy. This study provides a redescription and revision of Robustichthys based upon a comparative study of eight type specimens and nine new specimens. Newly recognized information includes a toothed parasphenoid, a pair of premaxillae not pierced by the olfactory nerve, a splint-like quadratojugal, a hatchet-shaped hyomandibula, an hourglass-shaped symplectic, anterior and posterior ceratohyals, a complete series of branchiostegal rays, and sclerotic bones. A revised reconstruction of Robustichthys is presented. Results of a cladistic analysis confirmed Robustichthys as an ionoscopiform within the Halecomorphi; the previous placements of Robustichthys as a basal ginglymodian and Ionoscopidae as a basal amiiform clade are not supported. The sister group relationship between Sinamiinae (Sinamia and Ikechaoamia) and Amiinae (Amia and Cyclurus) within the Amiidae is newly recognized. This revised topology provides new insights into the evolution and historical paleoecology of halecomorph fishes.


Review of Devonian-Carboniferous Boundary sections in the Rhenish Slate Mountains (Germany)

Abstract: Thirty Devonian-Carboniferous Boundary sections of the Rhenish Slate Mountains and adjacent subsurface areas are reviewed with respect to litho-, event, conodont, ammonoid, sequence, and chemostratigraphy. In the interval from the base of the uppermost Famennian (Wocklum Beds, Wocklumian) to the base of the middle Tournaisian (base Lower Alum Shale), 11 conodont and 16 ammonoid (sub)zones are distinguished. The terminology of the Hangenberg Crisis Interval is refined, with an overall regressive Crisis Prelude below the main Hangenberg Extinction, which defines the base of the transgressive Lower Crisis Interval (Hangenberg Black Shale). The glacigenic and regressive Middle Crisis Interval (Hangenberg Shale/Sandstone) is followed by the overall transgressive Upper Crisis Interval that can be subdivided into three parts (I to III) with the help of conodont stratigraphy (upper costatus-kockeli Interregnum = upper ckI, Protognathodus kockeli Zone, and lower part of Siphonodella (Eosiphonodella) sulcata s.l./Pr. kuehni Zone). Protognathodus kockeli includes currently a wide range of forms, which variabilities and precise ranges need to be established before a precise GSSP level should be selected. Returning to its original definition, the former Upper duplicata Zone is re-named as Siphonodella (S.) mehli Zone. It replaces the S. (S.) jii Zone, which is hampered by taxonomic complications. The S. (S.) quadruplicata Zone of Ji (1985) is hardly supported by Rhenish data. The entry of typical S. (S.) lobata (M1) characterises an upper subdivision (subzone) of the S. (S.) sandbergi Zone; the new S. (S.) lobata M2 enters much earlier within the S. (S.) mehli Zone. The ammonoid-defined base of the Wocklum-Stufe (Upper Devonian = UD VI) begins with the Linguaclymenia similis Zone (UD VI-A 1 ). The oldest S. (Eosiphonodella) enter within the Muessenbiaergia bisulcata Zone (UD VI-A 2 ). The traditional Parawocklumeria paradoxa Zone of Schindewolf (1937) is divided into successive P. paprothae (VI-C 1 ), P. paradoxa (VI-C 2 ), and Mayneoceras nucleus (VI-C 3 ) Subzones. In the lower Tournaisian (Lower Carboniferous = LC I), the Gattendorfia subinvoluta Zone is subdivided into G. subinvoluta (LC I-A 2 ) and "Eocanites" nodosus (LC I-A 3 ) Subzones. The Paprothites dorsoplanus Zone (LC I-B) can be divided into Pap. dorsoplanus (LC I-B 1 ) and Paragattendorfia sphaeroides (LC I-B 2 ) Subzones. Potential subdivisions of the Pseudarietites westfalicus (LC I-C) and Parag. patens Zones (LC I-D) are less distinctive. The unfossiliferous or argillaceous upper part of the Hangenberg Limestone and the overlying Lower Alum Shale Event Interval remain regionally unzoned for ammonoids.

devonian-carboniferous boundary section
assumed hangenberg limestone
lithostratigraphic unit
northern rhenish massif


A new symmetrodont mammal (Trechnotheria: Zhangheotheriidae) from the Early Cretaceous of China and trechnotherian character evolution

Abstract: We report the discovery of Anebodon luoi, a new genus and species of zhangheotheriid symmetrodont mammal from the Lujiatun site of the Lower Cretaceous Yixian Formation, China. The fossil is represented by an associated partial skull and dentaries with a nearly complete dentition, and with a dental formula of I4/3 C1/1 P5/4 M3/4. This new taxon lacks the high molar count typical of derived symmetrodonts, differing from the well-represented zhangheotheriids Zhangheotherium and Maotherium in having a postcanine dental formula that resembles more primitive tinodontid symmetrodonts on the one hand, and sister taxa to therians such as Peramus on the other. Upper and lower distal premolars are strongly molariform and are captured undergoing replacement, clarifying positional homology among related taxa. We also describe the rostrum and, for the first time in a symmetrodont, much of the orbital mosaic. Importantly, our new taxon occupies a basal position within the Zhangheotheriidae and permits discussion of trechnotherian character evolution, ultimately shedding additional light on the evolution of therians.

symmetrodont
trechnotherian character evolution
yixian formation


A Simulation-Based Evaluation of Tip-Dating Under the Fossilized Birth–Death Process

Abstract: Abstract Bayesian molecular dating is widely used to study evolutionary timescales. This procedure usually involves phylogenetic analysis of nucleotide sequence data, with fossil-based calibrations applied as age constraints on internal nodes of the tree. An alternative approach is tip-dating, which explicitly includes fossil data in the analysis. This can be done, for example, through the joint analysis of molecular data from present-day taxa and morphological data from both extant and fossil taxa. In the context of tip-dating, an important development has been the fossilized birth–death process, which allows non-contemporaneous tips and sampled ancestors while providing a model of lineage diversification for the prior on the tree topology and internal node times. However, tip-dating with fossils faces a number of considerable challenges, especially, those associated with fossil sampling and evolutionary models for morphological characters. We conducted a simulation study to evaluate the performance of tip-dating using the fossilized birth–death model. We simulated fossil occurrences and the evolution of nucleotide sequences and morphological characters under a wide range of conditions. Our analyses of these data show that the number and the maximum age of fossil occurrences have a greater influence than the degree of among-lineage rate variation or the number of morphological characters on estimates of node times and the tree topology. Tip-dating with the fossilized birth–death model generally performs well in recovering the relationships among extant taxa but has difficulties in correctly placing fossil taxa in the tree and identifying the number of sampled ancestors. The method yields accurate estimates of the ages of the root and crown group, although the precision of these estimates varies with the probability of fossil occurrence. The exclusion of morphological characters results in a slight overestimation of node times, whereas the exclusion of nucleotide sequences has a negative impact on inference of the tree topology. Our results provide an overview of the performance of tip-dating using the fossilized birth–death model, which will inform further development of the method and its application to key questions in evolutionary biology.


Calibration uncertainty in molecular dating analyses: there is no substitute for the prior evaluation of time priors

Abstract: Calibration is the rate-determining step in every molecular clock analysis and, hence, considerable effort has been expended in the development of approaches to distinguish good from bad calibrations. These can be categorized into a priori evaluation of the intrinsic fossil evidence, and a posteriori evaluation of congruence through cross-validation. We contrasted these competing approaches and explored the impact of different interpretations of the fossil evidence upon Bayesian divergence time estimation. The results demonstrate that a posteriori approaches can lead to the selection of erroneous calibrations. Bayesian posterior estimates are also shown to be extremely sensitive to the probabilistic interpretation of temporal constraints. Furthermore, the effective time priors implemented within an analysis differ for individual calibrations when employed alone and in differing combination with others. This compromises the implicit assumption of all calibration consistency methods, that the impact of an individual calibration is the same when used alone or in unison with others. Thus, the most effective means of establishing the quality of fossil-based calibrations is through a priori evaluation of the intrinsic palaeontological, stratigraphic, geochronological and phylogenetic data. However, effort expended in establishing calibrations will not be rewarded unless they are implemented faithfully in divergence time analyses.

evaluation
time prior
divergence time
calibration


Thrusting and exhumation around the margins of the western Tarim basin during the India‐Asia collision

Abstract: The Cenozoic collision between India and Asia has deformed a large part of central Asia. To the north of Tibet around the margins of the western Tarim basin, major basin-vergent thrusting has uplifted and exhumed thick Jurassic to Neogene sedimentary sections; this presumably reflects the propagation of collision-induced deformation into the basin. Apatite fission track data from five sections involved in this thrusting record strong late Oligocene to middle Miocene exhumation and cooling. On the northwest margin of the basin on the piedmont of the Tian Shan, a section exhumed by thrusting yields an exhumation age of 13.6±2.2 Ma (±1σ). Four Miocene sandstones from a second section 40 km to the east yield detrital source area cooling ages which decrease upsection from 25.0±3.9 to 13.1±2.2 Ma. Landsat imagery suggests that probable sediment source areas were dominated by Neogene thrusting, so these ages likely record progressive unroofing in Tian Shan thrust systems. Deformed Miocene to Pleistocene strata indicate that thrusting has continued and propagated basinward up until the present. Previously published apatite data from the Junggar basin on the northern flank of the Tian Shan yield a similar age of 24.7±3.9 Ma. On the southwest margin of Tarim on the piedmont of the western Kunlun Shan, three sections yield cooling ages of 19.8±0.9 Ma, 20.0±3.1 Ma, and roughly 20 Ma. Farther south at Kudi, previous work has yielded apatite cooling ages of 17±2 Ma and a zircon cooling age of 22±2 Ma. These similar cooling ages over a ≈250 km long belt in the western Kunlun Shan are associated with the transpressional Kumtag fault and the Main Pamir Thrust (MPT). Geologic relations within the western Kunlun Shan suggest that the MPT-Kumtag fault system offsets the originally linear trend of the Paleozoic-early Mesozoic Kunlun arc system by 200–300 km, accommodating much of the Neogene northward indentation of the Pamir block. We propose that the ≈20 Ma ages slightly postdate the initiation of this indentation and consequent crustal thickening. Taken together, the Tian Shan and Kunlun Shan results indicate that crustal thickening, in part accommodated by strike-slip faulting, became the dominant mode of deformation by ≈25–20 Ma in a large region extending from the Pamir and west Kunlun Shan north to the Tian Shan.


Test of Martin’s overkill hypothesis using radiocarbon dates on extinct megafauna

Abstract: Significance Coincident with the human colonization of the Western Hemisphere, dozens of genera of Pleistocene megafauna were lost to extinction. Following Martin, we argue that declines in the record of radiocarbon dates of extinct genera may be used as an independent means of detecting the first presence of humans in the New World. Our results, based on analyses of radiocarbon dates from Eastern Beringia, the contiguous United States, and South America, suggest north to south, time, and space transgressive declines in megafaunal populations as predicted by the overkill hypothesis. This finding is difficult to reconcile with other extinction hypotheses. However, it remains to be determined whether these findings will hold with larger samples of radiocarbon dates from all regions. Following Martin [Martin PS (1973) Science 179:969–974], we propose the hypothesis that the timing of human arrival to the New World can be assessed by examining the ecological impacts of a small population of people on extinct Pleistocene megafauna. To that end, we compiled lists of direct radiocarbon dates on paleontological specimens of extinct genera from North and South America with the expectation that the initial decline of extinct megafauna should correspond in time with the initial evidence for human colonization and that those declines should occur first in eastern Beringia, next in the contiguous United States, and last in South America. Analyses of spacings and frequency distributions of radiocarbon dates for each region support the idea that the extinction event first commenced in Beringia, roughly 13,300–15,000 BP. For the United States and South America, extinctions commenced considerably later but were closely spaced in time. For the contiguous United States, extinction began at ca. 12,900–13,200 BP, and at ca. 12,600–13,900 BP in South America. For areas south of Beringia, these estimates correspond well with the first significant evidence for human presence and are consistent with the predictions of the overkill hypothesis.

overkill hypothesis
radiocarbon date
south america


True Dalmanella and taxonomic implications for some Late Ordovician dalmanellid brachiopods from North America

Abstract: Examination of topotype material of the type species of Dalmanella, D. testudinaria, from the Hirnantian strata of Borenshult, southern Sweden, revealed several diagnostic characters that distinguish the type species from some common North American species assigned previously to the genus. The typical Dalmanella testudinaria has a consistently developed, primary medial interspace in the dorsal valve, a cardinal process with an invariably bilobed myophore, punctae of two distinctly different sizes, and a lack of aditicules. In contrast, many Late Ordovician species from North America, such as ‘Dalmanella’ meeki and ‘Dalmanella’ multisecta from the Cincinnati type area, consistently have a dorsal medial costa, with a strong tendency to develop aditicules and a trilobate cardinal process. This study confirms the previous notion of many authors that true Dalmanella testudinaria, or the genus as a whole, is either rare or entirely absent in North America. Typical Dalmanella was predominant in cool-water depositional environments (such as the Hirnantia Fauna), whereas the North American dalmanellids (e.g. Cincinnati-type ‘Dalmanella’, Paucicrura, and Diceromyonia) were most abundant and diverse in tropical, epicontinental sea settings. A clear definition of the true Dalmanella, therefore, has important implications for the study of Late Ordovician–Early Silurian brachiopod evolution, paleoecology, and paleobiogeography.


New Insights into the Skull of Istiodactylus latidens (Ornithocheiroidea, Pterodactyloidea)

Abstract: The skull of the Cretaceous pterosaur Istiodactylus latidens, a historically important species best known for its broad muzzle of interlocking, lancet-shaped teeth, is almost completely known from the broken remains of several individuals, but the length of its jaws remains elusive. Estimates of I. latidens jaw length have been exclusively based on the incomplete skull of NHMUK R3877 and, perhaps erroneously, reconstructed by assuming continuation of its broken skull pieces as preserved in situ. Here, an overlooked jaw fragment of NHMUK R3877 is redescribed and used to revise the skull reconstruction of I. latidens. The new reconstruction suggests a much shorter skull than previously supposed, along with a relatively tall orbital region and proportionally slender maxilla, a feature documented in the early 20th century but ignored by all skull reconstructions of this species. These features indicate that the skull of I. latidens is particularly distinctive amongst istiodactylids and suggests greater disparity between I. latidens and I. sinensis than previously appreciated. A cladistic analysis of istiodactylid pterosaurs incorporating new predicted I. latidens skull metrics suggests Istiodactylidae is constrained to five species (Liaoxipterus brachyognathus, Lonchengpterus zhoai, Nurhachius ignaciobritoi, Istiodactylus latidens and Istiodactylus sinensis) defined by their distinctive dentition, but excludes the putative istiodactylids Haopterus gracilis and Hongshanopterus lacustris. Istiodactylus latidens, I. sinensis and Li. brachyognathus form an unresolved clade of derived istiodactylids, and the similarity of comparable remains of I. sinensis and Li. brachyognathus suggest further work into their taxonomy and classification is required. The new skull model of I. latidens agrees with the scavenging habits proposed for these pterosaurs, with much of their cranial anatomy converging on that of habitually scavenging birds.

skull
istiodactylus latiden
istiodactylid
liaoxipterus


Estimation of locomotory forces and stresses in the limb bones of Recent and extinct equids

Abstract: The locomotory stresses acting at the midshaft of the third metacarpal of Equus are estimated from a geometrical model of the manus during peak loading, which is derived from in vivo data. The stress results compare favorably with those from in vivo strain gage experiments. The model is then modified to account for the anatomical and inferred functional differences in the feet of the two extinct, tridactyl equids Merychippus and Mesohippus, incorporating assumptions from earlier functional studies. Estimates of the stresses in the fossil metacarpals are derived, and fall within the range of values obtained in vivo for limb bones of a wide size range of mammals. The results indicate that the third metacarpal of Merychippus was the most highly stressed and that of Mesohippus the least, with stresses for Equus falling between. Thus the anatomical and functional changes in the manus between Mesohippus and Equus are not simply a result of the difference in body size.

force
stress
limb bone
third metacarpal
merychippus


A mitogenomic timetree for Darwin’s enigmatic South American mammal Macrauchenia patachonica

Abstract: The unusual mix of morphological traits displayed by extinct South American native ungulates (SANUs) confounded both Charles Darwin, who first discovered them, and Richard Owen, who tried to resolve their relationships. Here we report an almost complete mitochondrial genome for the litoptern Macrauchenia. Our dated phylogenetic tree places Macrauchenia as sister to Perissodactyla, but close to the radiation of major lineages within Laurasiatheria. This position is consistent with a divergence estimate of ∼66 Ma (95% credibility interval, 56.64–77.83 Ma) obtained for the split between Macrauchenia and other Panperissodactyla. Combined with their morphological distinctiveness, this evidence supports the positioning of Litopterna (possibly in company with other SANU groups) as a separate order within Laurasiatheria. We also show that, when using strict criteria, extinct taxa marked by deep divergence times and a lack of close living relatives may still be amenable to palaeogenomic analysis through iterative mapping against more distant relatives.

perissodactyla
litopterna
phylogenetic analysis


Early development and coloniality in Oligophylloides from the Devonian of Morocco—Are Heterocorallia Palaeozoic octocorals?

Abstract: Heterocorals represent an enigmatic group of Palaeozoic corals, known from relatively short time intervals in the Devonian and Carboniferous periods. The major differences between Heterocorallia and other Palaeozoic corals are the lack of an external theca (epitheca), lack of calices and the presence of dichotomously dividing septa-like structures. Heterocoral skeleton was presumably externally covered by the soft tissue and each branch of their skeleton has, until now, been regarded as a corallite–a skeleton of a single polyp. We investigated upper Famennian Oligophylloides from Morocco, focussing on branching processes, wall structure, previously poorly known initial growth stages and the growing tip, described here for the first time. We demonstrate that Oligophylloides shows a unique colony development not known in any group of anthozoans possessing a septate-like architecture and suggest that the previously postulated homology between true septa in hexa- and rugose corals on one hand, and Oligophylloides on the other, must be rejected. Based on the skeleton structure and branching patterns, we postulate, contrary to former ideas, that the stem and branches of heterocorals represent the skeleton of a multi-polyp colonial coral, similar to many extant octocorals. We found numerous potential homologies with octocoral skeletons (notably the Keratoisidinae within the Isididae) and, as a result, we propose the inclusion of the order Heterocorallia within the subclass Octocorallia. This suggestion requires, however, further research on the other taxa of heterocorals. We also propose some changes to the morphological terminology for the Heterocorallia.


Impact of skeletal dissolution and secondary aragonite on trace element and isotopic climate proxies in Porites corals

Abstract: Restricted zones of recent dissolution and secondary aragonite infilling were identified in a coral core collected in 1986 from a living massive Porites colony from the central Great Barrier Reef, Australia. Secondary aragonite needles, ≥20 μm long, cover skeletal surfaces deposited from 1972 to late 1974 and increase bulk density by 10%. Dissolution is observed above this zone, whereas older skeleton is pristine. We investigate the impact of both types of early marine diagenesis on skeletal geochemistry and coral paleoclimate reconstructions by comparison with similar records from eight contemporary Porites colonies collected at nearby reefs. Secondary aragonite overgrowth causes anomalies in skeletal density, Mg/Ca, Sr/Ca, U/Ca, δ O, and δC. The secondary aragonite is consistently associated with a cool temperature anomaly for each of the sea surface temperature (SST) proxies (δO-SST -1.6°C; Sr/Ca-SST -1.7°C; Mg/Ca-SST -1.9°C; U/Ca-SST -2.8°C). Dissolution, through incongruent leaching, also causes cool SST artifacts but only for trace element ratios (Mg/Ca-SST -1.2°C; Sr/Ca-SST -1.2°C; U/Ca-SST -2.1°C). The sequence of preference with respect to dissolution of coral skeleton in seawater is Mg > Ca > Sr > U. Rigorous screening of coral material for paleoclimate reconstructions is therefore necessary to detect both dissolution and the presence of secondary minerals. The excellent agreement between apparent SST anomalies generated by different modes of diagenesis means that replication of tracers within a single coral cannot be used to validate climate-proxy interpretations. Poor replication of records between different coral colonies, however, provides a strong indication of nonclimatic artifacts such as dissolution and secondary aragonite. Copyright 2007 by the American Geophysical Union.


Silurian craniide brachiopods from Gotland

Abstract: Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.


Substrate affinity and diversity dynamics of Paleozoic marine animals

Abstract: Abstract Short-term fluctuations in the diversification rate of Paleozoic marine animal genera are more strongly correlated with extinction-rate variation than with origination-rate variation. Diversity dynamics are strikingly different in the Mesozoic and Cenozoic, when variation in origination is more important than extinction. Data on the lithologic context of taxonomic occurrences in the Paleobiology Database are used to assess the substrate affinities of Paleozoic genera. The greater role of extinction-rate variation in the Paleozoic is found to characterize genera with an affinity for carbonate substrates but not those that prefer terrigenous clastic substrates. It is therefore plausible that the Paleozoic to post-Paleozoic shift in diversity dynamics is underlain in part by the secular decline in the relative areal extent of carbonate environments, and the concomitant decline in the relative diversity of carbonate- versus clastic-loving taxa.

diversity dynamic
origination
substrate affinity


Reproductive innovations and pulsed rise in plant complexity

Abstract: Description Evolutionary pulses of complexity The evolution of land plants has been marked by an increase in the complexity of reproductive structures. Leslie et al. analyzed the temporal pattern of this increase in a study of fossil and extant land plants across the entire paleontological record. Reproductive complexity increased in two widely separated pulses corresponding to key innovations in reproductive biology: the origin of seeds in the very late Devonian and the origin of flowering plants in the mid-Cretaceous almost 250 million years later. After the origin of flowers, there was a rapid expansion in the morphological complexity of flowering plants. In contrast to many aspects of animal morphological diversity, which expanded early in evolutionary history, most complexity in plants was achieved relatively late. —AMS Increases in plant reproductive complexity in two widely separated pulses were associated with major evolutionary innovations. Morphological complexity is a notable feature of multicellular life, although whether it evolves gradually or in early bursts is unclear. Vascular plant reproductive structures, such as flowers, are familiar examples of complex morphology. In this study, we use a simple approach based on the number of part types to analyze changes in complexity over time. We find that reproductive complexity increased in two pulses separated by ~250 million years of stasis, including an initial rise in the Devonian with the radiation of vascular plants and a pronounced increase in the Late Cretaceous that reflects flowering plant diversification. These pulses are associated with innovations that increased functional diversity, suggesting that shifts in complexity are linked to changes in function regardless of whether they occur early or late in the history of vascular plants.

innovation
rise
reproductive structure
flower
vascular plant


Phylogenetic analyses with four new Cretaceous bristletails reveal inter‐relationships of Archaeognatha and Gondwana origin of Meinertellidae

Abstract: Based on fifteen Archaeognatha (=Microcoryphia) specimens from Myanmar (Burmese) amber, including males, females and immatures, two new genera and four species, Cretaceomachilis longa sp.n., Unimeinertellus abundus gen. et sp.n., U. bellus sp.n. and Nullmeinertellus wenxuani gen. et sp.n., are described. Phylogenetic analyses of taxa in Archaeognatha were conducted using Maximum parsimony and Bayesian inference based on morphological characters and DNA sequence data. Our results confirm the phylogenetic position of the new genera, clarify the monophyly of Meinertellidae and indicate that the ‘paleo‐types’ excluding Ditrigoniophthalmus are nested within the Machilidae group, but suggest that the three subfamilies within Machilidae may be artificial. The diversity of meinertellids with derived characters found from the Cretaceous indicate that the divergence time of Machilidae and Meinertellidae is much earlier than the Cretaceous. We propose the possibility that Meinertellidae might have originated on Gondwana.

archaeognatha
meinertellidae
phylogenetic analysis
tropical forest community
southeastern asia


The uncertain role of diversity dependence in species diversification and the need to incorporate time-varying carrying capacities

Abstract: There is no agreement among palaeobiologists or biologists as to whether, or to what extent, there are limits on diversification and species numbers. Here, we posit that part of the disagreement stems from: (i) the lack of explicit criteria for defining the relevant species pools, which may be defined phylogenetically, ecologically or geographically; (ii) assumptions that must be made when extrapolating from population-level logistic growth to macro-evolutionary diversification; and (iii) too much emphasis being placed on fixed carrying capacities, rather than taking into account the opportunities for increased species richness on evolutionary timescales, for example, owing to increased biologically available energy, increased habitat complexity and the ability of many clades to better extract resources from the environment, or to broaden their resource base. Thus, we argue that a more effective way of assessing the evidence for and against the ideas of bound versus unbound diversification is through appropriate definition of the relevant species pools, and through explicit modelling of diversity-dependent diversification with time-varying carrying capacities. Here, we show that time-varying carrying capacities, either increases or decreases, can be accommodated through changing intrinsic diversification rates (diversity-independent effects), or changing the effects of crowding (diversity-dependent effects).

time-varying carrying capacity
iii
diversification rate


Phloem sap in Cretaceous ambers as abundant double emulsions preserving organic and inorganic residues

Abstract: Fossilized remains preserved in amber provide abundant data on the paleobiota surrounding the resin-producing plants, but relatively scarcer information about the resinous sources themselves. Here, dark pseudoinclusions in kidney-shaped amber pieces from the Early Cretaceous (Albian) amber from Spain are studied. This type of fossilized remain, abundant in Cretaceous ambers, was first interpreted as fossilized vacuole-bearing microorganisms, but later regarded as artifactual and probably secreted by the resinous trees, although their origin remained unclear. Using complementary microscopy (light, electron, confocal), spectroscopy (infrared, micro-Raman), mass spectrometry and elemental analysis techniques, we demonstrate that the pseudoinclusions correspond to droplets of phloem sap containing amber spheroids and preserving both organic and inorganic residues consistent with degraded components from the original sap. The amber pieces containing pseudoinclusions are fossilized, resin-in-sap-in-resin double emulsions, showing banding patterns with differential content of resin-in-sap emulsion droplets. Our findings represent the first time fossilized phloem sap, 105 million years old, has been recognized and characterized, and open new lines of paleontological research with taxonomic, taphonomic, physiological and ecological implications.

double emulsion
microorganism
resinous tree
phloem sap


Neurocranial anatomy of the petalichthyid placoderm Shearsbyaspis oepiki Young revealed by X‐ray computed microtomography

Abstract: Stem‐group gnathostomes reveal the sequence of character acquisition in the origin of modern jawed vertebrates. The petalichthyids are placoderm‐grade stem‐group gnathostomes known from both isolated skeletal material and rarer articulated specimens of one genus. They are of particular interest because of anatomical resemblances with osteostracans, the jawless sister group of jawed vertebrates. Because of this, they have become central to debates on the relationships of placoderms and the primitive cranial architecture of gnathostomes. However, among petalichthyids, only the braincase of Macropetalichthys has been studied in detail, and the diversity of neurocranial morphology in this group remains poorly documented. Using X‐ray computed microtomography, we investigated the endocranial morphology of Shearsbyaspis oepiki Young, a three‐dimensionally preserved petalichthyid from the Early Devonian of Taemas‐Wee Jasper, Australia. We generated virtual reconstructions of the external endocranial surfaces, orbital walls and cranial endocavity, including canals for major nerves and blood vessels. The neurocranium of Shearsbyaspis resembles that of Macropetalichthys, particularly in the morphology of the brain cavity, nerves and blood vessels. Many characters, including the morphology of the pituitary vein canal and the course of the trigeminal nerve, recall the morphology of osteostracans. Additionally, the presence of a parasphenoid in Shearsbyaspis (previously not known with confidence outside of arthrodires and osteichthyans) raises some questions about current proposals of placoderm paraphyly. Our detailed description of this specimen adds to the known morphological diversity of petalichthyids, and invites critical reappraisal of the phylogenetic relationships of placoderms.


The origins and early elaboration of projectile technology

Abstract: The ability of Homo sapiens to kill prey at a distance is arguably one of the catalysts for our current ecological dominance. Many researchers have suggested its origins lie in the African Middle Stone Age or the European Middle Palaeolithic (∼300‐30 thousand years ago), but the perishable components of armatures rarely preserve. Most research on this subject therefore emphasises analysis of armature tip size, shape, and diagnostic impacts or residues. Other lines of evidence have included human skeletal anatomy or analyses of the species composition of faunal assemblages. Projectile Impact Marks (PIMs) on archaeofaunal remains offer an ideal complement to this work, but their potential has been restricted mainly to the later Eurasian zooarchaeological record. A review of current evidence and approaches shows that systematic PIM research could add much to our understanding of early projectile technology, especially in Africa.

projectile technology
archaeofaunal remain
review


Spatial simulation and modelling of the early Pleistocene site of DS (Bed I, Olduvai Gorge, Tanzania): a powerful tool for predicting potential archaeological information from unexcavated areas

Abstract: Spatial statistical models are powerful tools for creating simulation and prediction models. Here, we apply such models to the newly discovered 1.84 Ma site of DS (Bed I, Olduvai Gorge, Tanzania). Ongoing excavation has already exposed 370 m2 of the same discrete archaeological level. This is the biggest window into an Early Pleistocene anthropogenic site. With such a large area opened, modelling based on spatial trends (using coordinates) and on covariates (topography) has enabled the creation of predictions about where the densest concentrations of unexcavated materials may lie. Following this modelling, excavation has confirmed the predictions; the densest clusters of stone tools and fossils bones are palaeotopographically and palaeoecologically influenced. Spatial statistical analysis is, therefore, a powerful analytical tool to model and understand in‐site and off‐site hominin behaviour as an interaction between hominins and environments.


A skeleton of a Uintan machaeroidine ‘creodont’ and the phylogeny of carnivorous eutherian mammals

Abstract: An edentulous partial skeleton of a carnivorous mammal from the Uinta Formation (middle Eocene) of Utah is referred to the rare and enigmatic sabre-tooth clade Machaeroidinae primarily on the basis of alveolar patterns and cranial morphology. The newly recognized skeleton includes portions of both girdles, all long bones, and the first known tarsal and phalangeal material of a machaeroidine. The specimen permits a preliminary reconstruction of the locomotor habits of machaeroidines, which appear to have been adapted to scansorial or arboreal rather than terrestrial locomotion. The new material also prompts a review of machaeroidine affinities, which have been unresolved for over a century, with consensus opinion favouring a link to either oxyaenid or limnocyonine hyaenodontid ‘creodonts’. Postcranial evidence favours a link to oxyaenids, as machaeroidines share numerous features with oxyaenids that are lacking in hyaenodontids. To test this relationship machaeroidines were included in a phylogenetic analysis broadly sampling early carnivorous eutherians, including members of both ‘Creodonta’ and Carnivoramorpha. Results place Machaeroidinae within Oxyaenidae but fail to support either ‘creodont’ or carnivoramorphan monophyly. Instead, Oxyaenidae is linked with Carnivoraformes, while Viverravidae is basally positioned among carnivorous eutherians. Reconsideration of the character evidence cited in support of Carnivoramorpha indicates that many features are ambiguous in the context of a broad sample of ‘creodonts’ and early carnivoramorphans. Hyaenodontid monophyly is also not recovered but this likely reflects the influence of one morphologically divergent genus, Arfia.

creodont
carnivorous eutherian mammal
middle eocene
machaeroidinae
phylogenetic analysis


The Neogene transition from C3 to C4 grasslands in North America: assemblage analysis of fossil phytoliths

Abstract: Abstract The rapid ecological expansion of grasses with C4 photosynthesis at the end of the Neogene (8-2 Ma) is well documented in the fossil record of stable carbon isotopes. As one of the most profound vegetation changes to occur in recent geologic time, it paved the way for modern tropical grassland ecosystems. Changes in CO2 levels, seasonality, aridity, herbivory, and fire regime have all been suggested as potential triggers for this broadly synchronous change, long after the evolutionary origin of the C4 pathway in grasses. To date, these hypotheses have suffered from a lack of direct evidence for floral composition and structure during this important transition. This study aimed to remedy the problem by providing the first direct, relatively continuous record of vegetation change for the Great Plains of North America for the critical interval (ca. 12-2 Ma) using plant silica (phytolith) assemblages. Phytoliths were extracted from late Miocene–Pliocene paleosols in Nebraska and Kansas. Quantitative phytolith analysis of the 14 best-preserved assemblages indicates that habitats varied substantially in openness during the middle to late Miocene but became more uniformly open, corresponding to relatively open grassland or savanna, during the late Miocene and early Pliocene. Phytolith data also point to a marked increase of grass short cells typical of chloridoid and other potentially C4 grasses of the PACMAD clade between 8 and 5 Ma; these data suggest that the proportion of these grasses reached up to ∼50–60% of grasses, resulting in mixed C3-C4 and highly heterogeneous grassland communities by 5.5 Ma. This scenario is consistent with interpretations of isotopic records from paleosol carbonates and ungulate tooth enamel. The rise in abundance of chloridoids, which were present in the central Great Plains since the early Miocene, demonstrates that the “globally” observed lag between C4 grass evolution/taxonomic diversification and ecological expansion occurred at the regional scale. These patterns of vegetation alteration imply that environmental change during the late Miocene–Pliocene played a major role in the C3-C4 shift in the Great Plains. Specifically, the importance of chloridoids as well as a decline in the relative abundance of forest indicator taxa, including palms, point to climatic drying as a key trigger for C4 dominance.

grassland
north america
phytolith
miocene


A new neosuchian crocodylomorph (Crocodyliformes, Mesoeucrocodylia) from the Early Cretaceous of north‐east Brazil

Abstract: Abstract:  A new neosuchian crocodylomorph, Susisuchus jaguaribensis sp. nov., is described based on fragmentary but diagnostic material. It was found in fluvial‐braided sediments of the Lima Campos Basin, north‐eastern Brazil, 115 km from where Susisuchus anatoceps was found, in rocks of the Crato Formation, Araripe Basin. S. jaguaribensis and S. anatoceps share a squamosal–parietal contact in the posterior wall of the supratemporal fenestra. A phylogenetic analysis places the genus Susisuchus as the sister group to Eusuchia, confirming earlier studies. Because of its position, we recovered the family name Susisuchidae, but with a new definition, being node‐based group including the last common ancestor of Susisuchus anatoceps and Susisuchus jaguaribensis and all of its descendents. This new species corroborates the idea that the origin of eusuchians was a complex evolutionary event and that the fossil record is still very incomplete.

new neosuchian crocodylomorph
susisuchus jaguaribensis
lima campos basin


Skeletal homologies, phylogeny and classification of the earliest asterozoan echinoderms

Abstract: Synopsis A thorough reappraisal of extant and fossil asterozoan plate systems is used to construct working hypotheses of primary plate homologies for Ordovician Asterozoa. Results indicate that inferomarginal ossicles of asteroids, somasteroids and the primitive ophiuroid Phragmactis are likely to be homologous with edrioasteroid marginals. The intermediate virgals and coverplates of somasteroids are identified as modified actinals and the sub‐laterals of certain stenurids are axial, here termed podial plates, with laterals in these taxa homologous to true lateral plates and adambulacrals of ophiuroids and asteroids, respectively. The nature of the asterozoan mouth frame is reinvestigated as a result of new study of the mouth frames of key taxa, with a reappraisal of existing developmental and symmetry data. It is argued that primitive asterozoan mouth angle plates were entirely ambulacral and probably not spine‐bearing, but that mouth spines evolved in asteroids and ophiuroids either on the existing first ambulacrals or by the superposition of first adambulacrals adoral to first ambulacrals. These new data are used to undertake a cladistic analysis of all known genera of Ordovician asterozoan echinoderms (38 taxa), based on a dataset of 150 morphological characters, rooted on the Middle Cambrian edrioasteroid taxa Stromatocystites and Cambraster. The resulting tree is fully resolved although some deeper branches show poor support. It reveals that Ordovician asteroids form a monophyletic group (defined by the presence of an unpaired axillary inferomarginal, opposing ambulacrals and blocky abutting adambulacrals) and are a sister group to the remainingasterozoans, including the somasteroids. Somasteroids are monophyletic and, with the primitive taxon Phragmactis, are a sister group to the remaining ophiuroids. The class Ophiuroidea is diagnosed by the near‐simultaneous appearance of apomorphies including an interradial disc with highly separated arms and aboral and oral interambulacral longitudinal musculature. A phylogenetic tree constructed for asterozoans based on current stratigraphic occurrence data indicates that at the time of their first appearance, asteroids and ophiuroid body plans were already established. This suggests a significant but unobserved pre‐Ordovician history. Comparison of observed and phylogenetic diversity together with the appearance of relatively young taxa close to the base of the tree point to a poor asterozoan record, particularly in the Lower Ordovician. A revised classification for stem‐group asterozoans is proposed based on well‐supported clades with the new taxon Eopentaroida erected. Mapping of character states on the most parsimonious solution suggests that whilst asteroid morphological evolution was conservative in the Ordovician, ophiuroids underwent a profound morphological diversification. Functionally, these changes are linked to the adoption of carnivory by derived Ordovician ophiuroids. In contrast, asteroids probably remained dominantly deposit‐feeders throughout the period. Character distribution on the cladogram refutes none of the hypotheses of primary plate homology, although it suggests that podial ossicles were developed in two clades independently and that the marginals of eopentaroids may not be homologous to those of other asterozoans and edrioasteroids.

phylogeny
asterozoan echinoderm
ordovician asterozoa
ophiuroid
arm


Non‐traditional isotope perspectives in vertebrate palaeobiology

Abstract: The recent development of multi‐collector inductively coupled plasma mass spectrometry (MC‐ICP‐MS) notably in the disciplines of earth sciences, now allows the precise measurement of isotope ratios, even at low concentration. Non‐traditional isotope systems, such as alkaline earth (Ca, Mg) and transition (Cu, Fe, Zn) metals are now being measured in a variety of biological tissues, including bone and teeth. Although our understanding of the environmental and biological mechanisms behind the fractionation of such elements is still in its infancy, some of these isotopes are suspected to fractionate along the food chain, as has been reported in the literature for calcium, magnesium and zinc. Other geochemical methods, such as concentration analyses, permit a prior assessment of diagenesis in the fossils and such an approach indicates that in some circumstances, not only enamel but also dentine or bone can preserve its original biogenic composition. The aims here are to: review the current knowledge surrounding these various isotopic tools; address their potential preservation in biological apatite; and provide the palaeobiologist with a guide to the different toolkits available, including a discussion of their potential applications in vertebrate palaeobiology with a case study involving two mammal assemblages from the Pleistocene of Europe.

vertebrate palaeobiology
fractionation
calcium


On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness

Abstract: The size and flight mechanics of giant pterosaurs have received considerable research interest for the last century but are confused by conflicting interpretations of pterosaur biology and flight capabilities. Avian biomechanical parameters have often been applied to pterosaurs in such research but, due to considerable differences in avian and pterosaur anatomy, have lead to systematic errors interpreting pterosaur flight mechanics. Such assumptions have lead to assertions that giant pterosaurs were extremely lightweight to facilitate flight or, if more realistic masses are assumed, were flightless. Reappraisal of the proportions, scaling and morphology of giant pterosaur fossils suggests that bird and pterosaur wing structure, gross anatomy and launch kinematics are too different to be considered mechanically interchangeable. Conclusions assuming such interchangeability—including those indicating that giant pterosaurs were flightless—are found to be based on inaccurate and poorly supported assumptions of structural scaling and launch kinematics. Pterosaur bone strength and flap-gliding performance demonstrate that giant pterosaur anatomy was capable of generating sufficient lift and thrust for powered flight as well as resisting flight loading stresses. The retention of flight characteristics across giant pterosaur skeletons and their considerable robustness compared to similarly-massed terrestrial animals suggest that giant pterosaurs were not flightless. Moreover, the term ‘giant pterosaur’ includes at least two radically different forms with very distinct palaeoecological signatures and, accordingly, all but the most basic sweeping conclusions about giant pterosaur flight should be treated with caution. Reappraisal of giant pterosaur material also reveals that the size of the largest pterosaurs, previously suggested to have wingspans up to 13 m and masses up to 544 kg, have been overestimated. Scaling of fragmentary giant pterosaur remains have been misled by distorted fossils or used inappropriate scaling techniques, indicating that 10–11 m wingspans and masses of 200–250 kg are the most reliable upper estimates of known pterosaur size.

giant pterosaur
pterosaur
flightless
body plan
modification


An integrated approach to understanding the role of the long neck in plesiosaurs

Abstract: The evolution and function of the long neck in plesiosaurs, and how the problems associated with stiffness or flexibility were overcome during feeding, or rapid swimming during predator avoidance, are explored, and a new interpretation for the function of the plesiosaur neck is presented. Based on the anatomy of the articular faces of contiguous cervical vertebral centra, neural arches, and cervical ribs, the plesiosaur neck was mainly adapted for ventral bending, with dorsal, lateral and rotational movements all relatively restricted. Predominant ventral bending indicates the neck was adapted for use beneath the body, suggesting feeding in the water column, close to the sea floor, or within soft sediments on the sea floor. A new model is proposed for the plesiosaur bauplan, comprising the head as a filter, straining, sieve feeding or sediment raking apparatus, mounted on a neck which acted as a stiff but ventrally flexible feeding tube, attached to the body which acted as a highly mobile feeding platform. Numerous features of plesiosaurs, including cranial and dental form, cervical vertebral morphology, body shape and limb-based propulsion, conform to this model. Comparative data from modern organisms support this novel explanation for the structure and function of the plesiosaur long neck. This integrative analysis offers an explanation for the evolution of the plesiosaur long neck as a key evolutionary novelty, and why this apparently enigmatic feature remained a prominent feature of plesiosaurs throughout their long evolutionary history.

long neck
plesiosaur
neural arch
morphology


Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology

Abstract: Twelve different bones from the skeleton of the holotype specimen of the hadrosaurian dinosaur Hypacrosaurus stebingeri were thin-sectioned to evaluate the significance of lines of arrested growth (LAGs) in age assessments. The presence of an external fundamental system (EFS) at the external surface of the cortex and mature epiphyses indicate that the Hypacrosaurus specimen had reached adulthood and growth had slowed considerably from earlier stages. The number of LAGs varied from none in the pedal phalanx to as many as eight in the tibia and femur. Most elements had experienced considerable Haversian reconstruction that had most likely obliterated many LAGs. The tibia was found to have experienced the least amount of reconstruction, but was still not optimal for skeletochronology because the LAGs were difficult to count near the periosteal surface. Additionally, the numbers of LAGs within the EFS vary considerably around the circumference of a single element and among elements. Counting LAGs from a single bone to assess skeletochronology appears to be unreliable, particularly when a fundamental system exists. Because LAGs are plesiomorphic for tetrapods, and because they are present in over a dozen orders of mammals, they have no particular physiological meaning that can be generalized to particular amniote groups without independent physiological evidence. Descriptions of dinosaur physiology as “intermediate” between the physiology of living reptiles and that of living birds and mammals may or may not be valid, but cannot be based reliably on the presence of LAGs.

age assessment
physiology
different bone
circumference
growth mark


Odyssey in the evolution of a paleopathologist

Abstract: Abstract. A recent report suggesting perceived limitations of and opportunities in the study of paleopathology suggested the importance of incorporation of scientific methodologies. It seems reasonable to also explore how those methodologies are developed and, indeed, how one approaches paleopathology as a science. The development of one such paleopathologist is delineated from his serendipitous observations to application of hypothesis generation and subsequent testing approach developed during basic medical science education. This approach resulted in recognition of how much he thought he knew was actually contrary to the facts. A critical factor was the collaborative approach with specialists in other fields, wherein linguistic confusion was overcome and perspectives refined by point–counterpoint analysis of hypotheses. The limited reliability of tertiary information was clearly exposed through examination of primary sources – original articles rather than what might be referred to as "meta-analyses". It became clear that linguistics was not the only challenge; application of techniques had to be observed and validated. Without validation one might obtain precision (method repeatedly reveals same results) but at the expense of accuracy (assurance that the method actually assesses the question). Paleontological studies are generally limited to examination of organisms and their traces. Archeologically based studies incorporate additional sources of information (e.g., historic), but are no less subject to such semantic and methodological issues. Proof of concept studies provided new windows to recognition not only of disease but to previous anatomical challenges (e.g., localization of direct muscle attachment sites and distribution). Trans-phylogenetic representation of disease falsified speculation that "evolution" would preclude analysis through time. Pathology is an intrinsic component of life and transcends both species and time. Knowledge gained in a given species and time can be applied to similar disease manifestations in other species in modern time. Once speculations were tested and either verified or falsified, paleo-epidemiologic approach allowed identification of patterns of spread and even application of that knowledge to recognition of human migration patterns. Proof of concept studies provided new windows to recognition not only of disease but to previous anatomical challenges (e.g., localization of direct muscle attachment sites and distribution).


Infernovenator steenae, a new serpentine recumbirostran from the ‘Mazon Creek’ Lagerstätte further clarifies lysorophian origins

Abstract: The Carboniferous (Pennsylvanian; 309–307 Mya) ‘Mazon Creek’ Lagerstätte produces some of the earliest tetrapod fossils of major Palaeozoic lineages. Previously, the Mazon Creek record of lysorophians was known from a single poorly preserved specimen consisting only of a partial vertebral column. Here we describe a new, virtually complete lysorophian genus and species, Infernovenator steenae gen. & sp. nov. on the basis of a unique combination of characters, including a near complete circumorbital series and the retention of a postfrontal. Parsimony-based phylogenetic analysis placed the new taxon in the family Molgophidae, as sister to Brachydectes newberryi. Those results and the more generalized cranial morphology present in Infernovenator further support a recumbirostran origin of Molgophidae. Co-occurrence of two morphologically and functionally distinct molgophids in the Early Moscovian suggests a rapid and underappreciated diversification of this family in the Early Pennsylvanian.

recumbirostran
mazon creek
tetrapod fossil
infernovenator


Jurassic synorogenic basin filling in western Korea: sedimentary response to inception of the western Circum‐Pacific orogeny

Abstract: This is the first sedimentologic and stratigraphic attempt to demonstrate Jurassic subduction‐induced basin‐filling processes in the early stage of the western Circum‐Pacific orogeny. The Chungnam Basin in western Korea was filled with a Lower to Middle Jurassic nonmarine succession, the Nampo Group, whose deposition postdated the Triassic final assembly of Chinese continental blocks. The Nampo Group consists of two repeated, fining‐ to coarsening‐upward alluvio‐lacustrine sequences, separated by an interval of thick breccia–gravel progradation deposits and its related strong proximal unconformities. No temporal variation in the degree of chemical weathering, along with the predominance of coals and a tropic to subtropic paleoflora, reveals little or no climate fluctuations during deposition of the Nampo Group. The observed relationships provide a record of sedimentation most likely controlled by temporal variations of tectonically driven sediment flux. Such syntectonic sedimentation of the Chungnam Basin occurred at a convergent margin of continental‐arc setting during the Daebo orogeny, synchronous with the early subduction of the western paleo‐Pacific ocean that resulted in formation of an accretionary complex along the East Asian continental margin during Jurassic time. Hence, synorogenic deposition in the Chungnam Basin is interpreted as sedimentary response to subduction–accretion of the western paleo‐Pacific plate.

sedimentary response
western circum‐pacific orogeny
chungnam basin
daebo orogeny


Upper Palaeolithic genomes reveal deep roots of modern Eurasians

Abstract: We extend the scope of European palaeogenomics by sequencing the genomes of Late Upper Palaeolithic (13,300 years old, 1.4-fold coverage) and Mesolithic (9,700 years old, 15.4-fold) males from western Georgia in the Caucasus and a Late Upper Palaeolithic (13,700 years old, 9.5-fold) male from Switzerland. While we detect Late Palaeolithic-Mesolithic genomic continuity in both regions, we find that Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers B45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers B25 kya, around the Last Glacial Maximum. CHG genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe B3,000 BC, supporting a formative Caucasus influence on this important Early Bronze age culture. CHG left their imprint on modern populations from the Caucasus and also central and south Asia possibly marking the arrival of Indo-Aryan languages.

caucasus hunter-gatherer
chg
upper palaeolithic


THERAPSIDS FROM THE PERMIAN CHIWETA BEDS AND THE AGE OF THE KAROO SUPERGROUP IN MALAWI

Abstract: Exposures of the Karoo Supergroup occur in the northern and southern portions of Malawi, south central Africa. The section in southern Malawi contains Lower Permian coal-bearing shales and extends upward to the Chikwawa basalt of Jurassic age (179 Ma). The Chikwawa basalt represents the northern limit of the Karoo Large Igneous Province, which signals the rifting of Gondwana. The Chiweta Beds of northern Malawi contain a Late Permian (Cistecephalus Zone; 256-258 Ma) fauna, including the dicynodont Oudenodon and a new burnetiamorph biarmosuchian more derived than Lemurosaurus and sister to Proburnetia, Burnetia, and Bullacephalus.


The oldest known metriorhynchid super-predator: a new genus and species from the Middle Jurassic of England, with implications for serration and mandibular evolution in predacious clades

Abstract: The Oxford Clay Formation of England has yielded numerous sympatric species of metriorhynchid crocodylomorphs, although disagreement has persisted regarding the number of valid species. For over 140 years teeth reminiscent of the genus Dakosaurus have been known from the Oxford Clay Formation but these have never been properly described and their taxonomy and systematic affinity remain contentious. Furthermore, an enigmatic mandible and associated postcranial skeleton discovered by Alfred Leeds in the Fletton brick pits near Peterborough also remains undescribed. We show that this specimen, and several isolated teeth, represents the oldest known remains of a large-bodied predatory metriorhynchid. This material is described herein and referred to Tyrannoneustes lythrodectikos gen. et sp. nov. This species has a unique occlusal pattern: the dentition was arranged so that the posterior maxillodentary teeth interlock in the same plane and occlude mesiodistally. It is the first described crocodylomorph with microscopic denticles that are not contiguous along the carinae (forming short series of up to 10 denticles) and do not noticeably alter the height of the keel. Additionally, the dorsally expanded and curved posterior region of the mandible ventrally displaced the dentary tooth row relative to the jaw joint facilitating the enlargement of the dentition and increasing optimum gape. Therefore, Tyrannoneustes would have been a large-bodied marine predator that was well-suited to feed on larger prey than other contemporaneous metriorhynchids. A new phylogenetic analysis finds Tyrannoneustes to be the sister taxon to the subclade Geosaurini. An isolated tooth, humerus, and well-preserved mandible suggest a second species of metriorhynchid super-predator may also have lived in the Oxford Clay sea. Finally, we revise the diagnoses and descriptions of the other Oxford Clay metriorhynchid species, providing a guide for differentiating the many contemporaneous taxa from this exceptional fossil assemblage. http://zoobank.org/urn:lsid:zoobank.org:pub:C6222858-EC36-430D-8E28-1F636A6697CA

metriorhynchid super-predator
england
serration
denticle
geosaurini


A new cymbospondylid ichthyosaur (Ichthyosauria) from the Middle Triassic (Anisian) of the Augusta Mountains, Nevada, USA

Abstract: A new taxon, Cymbospondylus duelferi sp. nov., is described from the late Anisian of the Fossil Hill Member of the Favret Formation of the Augusta Mountains, Pershing County, Nevada, USA. The holotype and only specimen consists of a fairly complete skull associated with articulated and disarticulated postcranial material. Body length reconstruction revealed a medium-sized individual with an estimated body length of 4.3 m. Dorsal vertebrae as well as the left humerus of this specimen are associated with three strings of articulated tail and posterior dorsal vertebrae, which are on average 68% smaller than the vertebrae of the medium-sized individual. Due to the small size of these vertebrae and their position within the trunk region, they most likely represent fetuses. Cymbospondylus duelferi sp. nov. provides the second-oldest evidence for viviparity in ichthyosaurs. In the course of this study, the skull morphology of C. petrinus and C. nichollsi was reviewed, resulting in revised character scorings for both. Phylogenetic analyses demonstrate a sister-taxon relationship of the new taxon with C. petrinus, which was found in the same stratigraphic unit, the Fossil Hill Member, that also crops out at the classical Fossil Hill locality in the Humboldt Range, where the member is part of the Prida Formation. Cymbospondylus duelferi sp. nov. shares a very similar skull morphology with C. petrinus and C. nichollsi, whereas the shoulder girdle morphology differs. Because there is no evidence for ontogenetic differences or sexual dimorphism in the new specimen, a third species of Cymbospondylus – neglecting the type species C. piscosus that is only known from five vertebrae – is recognized from the Fossil Hill Member. Further occurrences of the genus in the Lower and Middle Triassic of Svalbard (Boreal Ocean) and Europe (Tethys) point to a very fast radiation and dispersal of cymbospondylids during the Middle Triassic. https://zoobank.org:pub:906F5F32-9090-496D-8FC7-680D25EED5EF

ichthyosaur
middle triassic
cymbospondylus


From body scale ontogeny to species ontogeny: Histological and morphological assessment of the Late Devonian acanthodian Triazeugacanthus affinis from Miguasha, Canada

Abstract: Growth series of Palaeozoic fishes are rare because of the fragility of larval and juvenile specimens owing to their weak mineralisation and the scarcity of articulated specimens. This rarity makes it difficult to describe early vertebrate growth patterns and processes in extinct taxa. Indeed, only a few growth series of complete Palaeozoic fishes are available; however, they allow the growth of isolated elements to be described and individual growth from these isolated elements to be inferred. In addition, isolated and in situ scales are generally abundant and well-preserved, and bring information on (1) their morphology and structure relevant to phylogenetic relationships and (2) individual growth patterns and processes relative to species ontogeny. The Late Devonian acanthodian Triazeugacanthus affinis from the Miguasha Fossil-Lagerstätte preserves one of the best known fossilised ontogenies of early vertebrates because of the exceptional preservation, the large size range, and the abundance of complete specimens. Here, we present morphological, histological, and chemical data on scales from juvenile and adult specimens (scales not being formed in larvae). Histologically, Triazeugacanthus scales are composed of a basal layer of acellular bone housing Sharpey’s fibers, a mid-layer of mesodentine, and a superficial layer of ganoine. Developmentally, scales grow first through concentric addition of mesodentine and bone around a central primordium and then through superposition of ganoine layers. Ontogenetically, scales form first in the region below the dorsal fin spine, then squamation spreads anteriorly and posteriorly, and on fin webs. Phylogenetically, Triazeugacanthus scales show similarities with acanthodians (e.g. “box-in-box” growth), chondrichthyans (e.g. squamation pattern), and actinopterygians (e.g. ganoine). Scale histology and growth are interpreted in the light of a new phylogenetic analysis of gnathostomes supporting acanthodians as stem chondrichthyans.

species ontogeny
vertebrate
phylogenetic analysis


The evolutionary history of polycotylid plesiosaurians

Abstract: Polycotylidae is a clade of plesiosaurians that appeared during the Early Cretaceous and became speciose and abundant early in the Late Cretaceous. However, this radiation is poorly understood. Thililua longicollis from the Middle Turonian of Morocco is an enigmatic taxon possessing an atypically long neck and, as originally reported, a series of unusual cranial features that cause unstable phylogenetic relationships for polycotylids. We reinterpret the holotype specimen of Thililua longicollis and clarify its cranial anatomy. Thililua longicollis possesses an extensive, foramina-bearing jugal, a premaxilla–parietal contact and carinated teeth. Phylogenetic analyses of a new cladistic dataset based on first-hand observation of most polycotylids recover Thililua and Mauriciosaurus as successive lineages at the base of the earliest Late Cretaceous polycotyline radiation. A new dataset summarizing the Bauplan of polycotylids reveals that their radiation produced an early burst of disparity during the Cenomanian–Turonian interval, with marked plasticity in relative neck length, but this did not arise as an ecological release following the extinction of ichthyosaurs and pliosaurids. This disparity vanished during and after the Turonian, which is consistent with a model of ‘early experimentation/late constraint’. Two polycotylid clades, Occultonectia clade nov. and Polycotylinae, survived up to the Maastrichtian, but with low diversity.

plesiosaurian
polycotylidae
phylogenetic analysis


Swimming capabilities of Mesozoic marine reptiles: implications for method of predation

Abstract: Body shape and mode of swimming were major factors that affected the swimming capabilities of Mesozoic marine reptiles. By estimating the total drag and the amount of energy available through metabolism, the maximum sustained swimming speed was calculated for 115 marine reptile specimens. Calculated sustained swimming speeds range from 1.8 to 2.7 m/sec, but are probably too high by as much as a factor of two. Mesozoic marine reptiles were probably much slower than modern toothed whales. The diversification of fast, agile teleost fish in the Cretaceous may have therefore contributed to the decline of the marine reptiles. Long-bodied reptiles appear to have had slower sustained swimming speeds than deep-bodied forms of the same length. For a given length, ichthyosaurs were probably faster sustained swimmers than plesiosaurs, and plesiosaurs were probably faster sustained swimmers than crocodiles and mosasaurs. This suggests that the long-bodied forms probably used an ambush technique to capture prey, to maximize the range of possible prey and to minimize competition with the faster pursuit predators.

marine reptile
predator
body shape
ichthyosaur
swimming capability


Miocene and Early Pleistocene Rodents (Mammalia) from Denizli Basin (Southwestern Turkey) and a New Species of Fossil Mus

Abstract:
ABSTRACT
 The terrestrial Neogene Basin of Denizli, located in southwestern Turkey, yields rodent faunas from nine localities dated between MN6 and MNQ18–19. Arabosminthus gansus, which is known in China and Mongolia, is recorded for the first time in Turkey at the Yenicekent localities (MN8–9), an older stratigraphical level than it is usually known from. Another species from the same localities, Microdyromys complicatus, is described for the first time in Anatolia in a younger locality than usual known in Western Europe. Myomimus dehmi, Myocricetodon eskihisarensis, Dryomys tosyaensis, and Dryomys sp. are the other species described in the present study. A new species of fossil Mus (Mammalia, Rodentia), Mus denizliensis, sp. nov. (MNQ18–19), has been found at the G€okpınar locality. This is the oldest fossil species of this genus known from Turkey or continental Europe.

southwestern turkey
fossil mus
mus denizliensis


Synchrotron Reveals Early Triassic Odd Couple: Injured Amphibian and Aestivating Therapsid Share Burrow

Abstract: Fossorialism is a beneficial adaptation for brooding, predator avoidance and protection from extreme climate. The abundance of fossilised burrow casts from the Early Triassic of southern Africa is viewed as a behavioural response by many tetrapods to the harsh conditions following the Permo-Triassic mass-extinction event. However, scarcity of vertebrate remains associated with these burrows leaves many ecological questions unanswered. Synchrotron scanning of a lithified burrow cast from the Early Triassic of the Karoo unveiled a unique mixed-species association: an injured temnospondyl amphibian (Broomistega) that sheltered in a burrow occupied by an aestivating therapsid (Thrinaxodon). The discovery of this rare rhinesuchid represents the first occurrence in the fossil record of a temnospondyl in a burrow. The amphibian skeleton shows signs of a crushing trauma with partially healed fractures on several consecutive ribs. The presence of a relatively large intruder in what is interpreted to be a Thrinaxodon burrow implies that the therapsid tolerated the amphibian’s presence. Among possible explanations for such unlikely cohabitation, Thrinaxodon aestivation is most plausible, an interpretation supported by the numerous Thrinaxodon specimens fossilised in curled-up postures. Recent advances in synchrotron imaging have enabled visualization of the contents of burrow casts, thus providing a novel tool to elucidate not only anatomy but also ecology and biology of ancient tetrapods.


New long-proboscid lacewings of the mid-Cretaceous provide insights into ancient plant-pollinator interactions

Abstract: Many insects with long-proboscid mouthparts are among the pollinators of seed plants. Several cases of the long-proboscid pollination mode are known between fossil insects (e.g., true flies, scorpionflies, and lacewings) and various extinct gymnosperm lineages, beginning in the Early Permian and increasing during the Middle Jurassic to Early Cretaceous. However, details on the morphology of lacewing proboscides and the relevant pollination habit are largely lacking. Here we report on three lacewing species that belong to two new genera and a described genus from mid-Cretaceous (Albian-Cenomanian) amber of Myanmar. All these species possess relatively long proboscides, which are considered to be modified from maxillary and labial elements, probably functioning as a temporary siphon for feeding on nectar. Remarkably, these proboscides range from 0.4–1.0 mm in length and are attributed to the most diminutive ones among the contemporary long-proboscid insect pollinators. Further, they clearly differ from other long-proboscid lacewings which have a much longer siphon. The phylogenetic analysis indicates that these Burmese long-proboscid lacewings belong to the superfamily Psychopsoidea but cannot be placed into any known family. The present findings represent the first description of the mouthparts of long-proboscid lacewings preserved in amber and highlight the evolutionary diversification of the ancient plant-pollinator interactions.

long-proboscid lacewing
ancient plant-pollinator interaction
insect
cretaceous
long proboscis


Eocene fossil is earliest evidence of flower-visiting by birds

Abstract: Birds are important pollinators, but the evolutionary history of ornithophily (bird pollination) is poorly known. Here, we report a skeleton of the avian taxon Pumiliornis from the middle Eocene of Messel in Germany with preserved stomach contents containing numerous pollen grains of an eudicotyledonous angiosperm. The skeletal morphology of Pumiliornis is in agreement with this bird having been a, presumably nectarivorous, flower-visitor. It represents the earliest and first direct fossil evidence of flower-visiting by birds and indicates a minimum age of 47 million years for the origin of bird–flower interactions. As Pumiliornis does not belong to any of the modern groups of flower-visiting birds, the origin of ornithophily in some angiosperm lineages may have predated that of their extant avian pollinators.

flower-visiting
stomach content
pollen grain


Diversity change during the rise of tetrapods and the impact of the ‘Carboniferous rainforest collapse’

Abstract: The Carboniferous and early Permian were critical intervals in the diversification of early four-limbed vertebrates (tetrapods), yet the major patterns of diversity and biogeography during this time remain unresolved. Previous estimates suggest that global tetrapod diversity rose continuously across this interval and that habitat fragmentation following the ‘Carboniferous rainforest collapse’ (CRC) drove increased endemism among communities. However, previous work failed to adequately account for spatial and temporal biases in sampling. Here, we reassess early tetrapod diversity and biogeography with a new global species-level dataset using sampling standardization and network biogeography methods. Our results support a tight relationship between observed richness and sampling, particularly during the Carboniferous. We found that subsampled species richness initially increased into the late Carboniferous, then decreased substantially across the Carboniferous/Permian boundary before slowly recovering in the early Permian. Our analysis of biogeography does not support the hypothesis that the CRC drove endemism; instead, we found evidence for increased cosmopolitanism in the early Permian. While a changing environment may have played a role in reducing diversity in the earliest Permian, our results suggest that the CRC was followed by increased global connectivity between communities, possibly reflecting both reduced barriers to dispersal and the diversification of amniotes.

tetrapod
carboniferous rainforest collapse
biogeography
endemism
sampling


Hagfish from the Cretaceous Tethys Sea and a reconciliation of the morphological–molecular conflict in early vertebrate phylogeny

Abstract: Significance Jawless, boneless, and virtually without fossil record, hagfish have long escaped systematists’ grip on their place among other fish. Yet their systematic resolution is critical to define vertebrates as a clade. Here we report an unequivocal fossil hagfish from the Cretaceous Mediterranean. Using this fossil to calibrate the evolutionary history of the group, our analysis supports hagfish and lampreys as sister groups, which likely diverged from one another in early Paleozoic times. As a result, vertebrates have a deep dichotomy, where some fossil jawless vertebrates sit closer to hagfish and lampreys than to jawed vertebrates. We showed that morphology-based analysis converged onto molecular inferences when characters are coded nonindependently, providing a case study for morphological–molecular conflicts in animal phylogeny. Hagfish depart so much from other fishes anatomically that they were sometimes considered not fully vertebrate. They may represent: (i) an anatomically primitive outgroup of vertebrates (the morphology-based craniate hypothesis); or (ii) an anatomically degenerate vertebrate lineage sister to lampreys (the molecular-based cyclostome hypothesis). This systematic conundrum has become a prominent case of conflict between morphology- and molecular-based phylogenies. To date, the fossil record has offered few insights to this long-branch problem or the evolutionary history of hagfish in general, because unequivocal fossil members of the group are unknown. Here, we report an unequivocal fossil hagfish from the early Late Cretaceous of Lebanon. The soft tissue anatomy includes key attributes of living hagfish: cartilages of barbels, postcranial position of branchial apparatus, and chemical traces of slime glands. This indicates that the suite of characters unique to living hagfish appeared well before Cretaceous times. This new hagfish prompted a reevaluation of morphological characters for interrelationships among jawless vertebrates. By addressing nonindependence of characters, our phylogenetic analyses recovered hagfish and lampreys in a clade of cyclostomes (congruent with the cyclostome hypothesis) using only morphological data. This new phylogeny places the fossil taxon within the hagfish crown group, and resolved other putative fossil cyclostomes to the stem of either hagfish or lamprey crown groups. These results potentially resolve the morphological–molecular conflict at the base of the Vertebrata. Thus, assessment of character nonindependence may help reconcile morphological and molecular inferences for other major discords in animal phylogeny.

morphological–molecular conflict
vertebrate
lamprey
cyclostomes
phylogenetic analysis


Modelling determinants of extinction across two Mesozoic hyperthermal events

Abstract: The Late Triassic and Early Toarcian extinction events are both associated with greenhouse warming events triggered by massive volcanism. These Mesozoic hyperthermals were responsible for the mass extinction of marine organisms and resulted in significant ecological upheaval. It has, however, been suggested that these events merely involved intensification of background extinction rates rather than significant shifts in the macroevolutionary regime and extinction selectivity. Here, we apply a multivariate modelling approach to a vast global database of marine organisms to test whether extinction selectivity varied through the Late Triassic and Early Jurassic. We show that these hyperthermals do represent shifts in the macroevolutionary regime and record different extinction selectivity compared to background intervals of the Late Triassic and Early Jurassic. The Late Triassic mass extinction represents a more profound change in selectivity than the Early Toarcian extinction but both events show a common pattern of selecting against pelagic predators and benthic photosymbiotic and suspension-feeding organisms, suggesting that these groups of organisms may be particularly vulnerable during episodes of global warming. In particular, the Late Triassic extinction represents a macroevolutionary regime change that is characterized by (i) the change in extinction selectivity between Triassic background intervals and the extinction event itself; and (ii) the differences in extinction selectivity between the Late Triassic and Early Jurassic as a whole.

mass extinction
extinction selectivity
global warming


New Age of Fishes initiated by the Cretaceous−Paleogene mass extinction

Abstract: Significance Ray-finned fishes are the most diverse and ecologically dominant group of vertebrates on the planet. Previous molecular phylogenies and paleontological studies have shown that modern ray-finned fishes (crown teleosts) radiated sometime in the Late Cretaceous or early Paleogene. Our data suggest that crown teleosts came into their current dominant ecological role in pelagic ecosystems immediately following the Cretaceous−Paleogene mass extinction 66 million years ago by filling newly vacated ecological niches and marking the beginning of an “age of ray-finned fishes.” Our study is, to our knowledge, the first geographically comprehensive, high-resolution study of marine vertebrate communities across the extinction. Ray-finned fishes (Actinopterygii) comprise nearly half of all modern vertebrate diversity, and are an ecologically and numerically dominant megafauna in most aquatic environments. Crown teleost fishes diversified relatively recently, during the Late Cretaceous and early Paleogene, although the exact timing and cause of their radiation and rise to ecological dominance is poorly constrained. Here we use microfossil teeth and shark dermal scales (ichthyoliths) preserved in deep-sea sediments to study the changes in the pelagic fish community in the latest Cretaceous and early Paleogene. We find that the Cretaceous−Paleogene (K/Pg) extinction event marked a profound change in the structure of ichthyolith communities around the globe: Whereas shark denticles outnumber ray-finned fish teeth in Cretaceous deep-sea sediments around the world, there is a dramatic increase in the proportion of ray-finned fish teeth to shark denticles in the Paleocene. There is also an increase in size and numerical abundance of ray-finned fish teeth at the boundary. These changes are sustained through at least the first 24 million years of the Cenozoic. This new fish community structure began at the K/Pg mass extinction, suggesting the extinction event played an important role in initiating the modern “age of fishes.”

cretaceous−paleogene mass extinction
ray-finned fish
shark
fish community
extinction event


Dietary and body-mass reconstruction of the Miocene neotropical bat Notonycteris magdalenensis (Phyllostomidae) from La Venta, Colombia

Abstract: Abstract. With 14 species recorded, the Miocene La Venta bat fauna is the most diverse bat paleocommunity in South America. It includes the oldest plant-visiting bat in the New World and some of the earliest representatives of the extant families Phyllostomidae, Thyropteridae, and Noctilionidae. La Venta's Notonycteris magdalenensis is an extinct member of the subfamily Phyllostominae, a group of modern Neotropical animalivorous bats, and is commonly included in studies of the evolution of Neotropical bats, but aspects of its biology remain unclear. In this study, we used multivariate dental topography analysis (DTA) to reconstruct the diet of N. magdalenensis by quantitatively comparing measures of molar complexity with those of 25 modern noctilionoid species representing all major dietary habits in bats. We found clear differences in molar complexity between dietary guilds, indicating that DTA is potentially an informative tool to study bat ecomorphology. Our results suggest N. magdalenensis was probably an omnivore or insectivore, rather than a carnivore like its modern relatives Chrotopterus auritus and Vampyrum spectrum. Also, we reconstructed the body mass of N. magdalenensis to be ∼95 g, larger than most insectivorous bats, but smaller than the largest carnivorous bat (V. spectrum). Our results confirm that N. magdalenensis was not a specialized carnivore. It remains to be demonstrated that the specialized carnivory ecological niche was occupied by the same lineage of phyllostomines from at least the middle Miocene. Combining our diet and body-mass reconstructions, we suggest that N. magdalenensis exhibits morphological pre-adaptations crucial for the evolution of specialized carnivory.


Metacryphaeus Tuberculatus and Metacryphaeus australis (Trilobita, Phacopida) from the Devonian of the Paraná Basin: Taxonomy and Paleobiogeography

Abstract:
Abstract.
 Calmoniid trilobites of the genus Metacryphaeus Reed include M. tuberculatus (Kozlowski), M. kegeli Carvalho, Edgecombe and Lieberman, M. meloi Carvalho, Edgecombe and Lieberman, M. rotundatus (Kozlowski), M. giganteus (Ulrich), M. convexus (Ulrich), M. curvigena Lieberman, M. branisai Lieberman, M. caffer (Salter), M. australis (Clarke), and M. allardyceae (Clarke). The geographic distribution of this genus comprises areas of Gondwana, including Bolivia, Peru, Brazil, the Falkland Islands, and South Africa. This study reports a new occurrence of M. tuberculatus in the Paraná Basin (Goiás and Paraná states, Brazil), in both the Alto Garças and the Apucarana sub-basins. Metacryphaeus tuberculatus was compared with all Metacryphaeus species and, especially, with M. australis, which also represents a new record for the Paraná Basin. These new records of M. tuberculatus imply a different dispersion interpretation from that which is known through the literature. Metacryphaeus tuberculatus seems to have originated during the Early Devonian and lived in areas such as Bolivia, Peru, and the Apucarana Sub-basin (Paraná Basin, Brazil). The dispersion between these areas is probably related to the Emsian transgression. During the Givetian, the species presumably migrated towards the Alto Garças Sub-basin (Paraná Basin, Brazil) and the Parnaíba Basin. These facts provide an alternative paleogeographic interpretation to that presented by Tropidoleptus Hall and Exaesiodiscus Moore and Jeffords in Gondwana and suggest a different pattern of migration during the Devonian of Brazil.

devonian
falkland islands
metacryphaeus tuberculatus