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bioRxiv2021

Orientations of Mistaken Point fronds indicate morphology impacted ability to survive turbulence

The Ediacaran organisms of the Mistaken Point E surface have provided crucial insight into early animal communities, including how they reproduced, the importance of Ediacaran height and what the most important factors were to their community dynamics. Here, we use this iconic community to investigate how morphological variation between eight taxa affected their ability to withstand different flow conditions. For each of Beothukis, Bradgatia, Charniodiscus procerus, Charniodiscus spinosus, Plumeropriscum, Primocandelabrum and Fractofusus we measured the orientation and length of their stems (if present) and their fronds. We statistically tested each taxons stem and frond orientation distributions to see whether they displayed a uniform or multimodal distribution. Where multimodal distributions were identified, the stem/frond length of each cohort was tested to identify if there were differences in size between different orientation groups. We find that Bradgatia and Thectardis show a bimodal felling direction, and infer that they were felled by the turbulent head of the felling flow. In contrast, the frondose rangeomorphs including Beothukis, Plumeropriscum, Primocandelabrum, and the arboreomorphs were felled in a single direction, indicating that they were upright in the water column, and were likely felled by the laminar tail of the felling flow. These differences in directionality suggests that an elongate habit, and particularly possession of a stem, lent greater resilience to frondose taxa against turbulent flows, suggesting that such taxa would have had improved survivability in conditions with higher background turbulence than taxa like Bradgatia and Thectardis, which lacked a stem and which had a higher centre of mass, which may have fared better in quieter water conditions.

paleontology↗

bioRxiv2021

Morphometric Analysis of Lungfish Endocasts Elucidates Early Dipnoan Palaeoneurological Evolution

Lungfish (Dipnoi) are lobe-finned fish (Sarcopterygii) that have persisted for over 400 million years from the Devonian Period to present day. They are the extant sister group to tetrapods and thus have the ability to provide unique insight into the condition of the earliest tetrapods as well as their own evolutionary history. The evolution of their dermal skull and dentition is relatively well understood, but this is not the case for the central nervous system. While the brain itself has very poor preservation potential and is not currently known in any fossil lungfish, substantial indirect information about it and associated structures such as the inner ears can be obtained from the cranial endocast. However, before the recent development of X-ray tomography as a palaeontological tool, these endocasts could not be studied non-destructively, and few detailed studies were undertaken. Here we describe and illustrate the endocasts of six Palaeozoic lungfishes (Iowadipterus halli, Gogodipterus paddyensis, Pillararhynchus longi, Griphognathus whitei, Orlovichthys limnatis, and Rhinodipterus ulrichi) from tomographic scans. We combine these with six previously described lungfish endocasts (4 fossil and 2 recent taxa), also based on tomographic studies, into a 12-taxon data set for multivariate morphometric analysis using 17 variables. We find that the olfactory region appears to be more highly plastic than the hindbrain, and undergoes significant elongation in several taxa. Further, while the semicircular canals covary as an integrated module, the utriculus and sacculus of the inner ear instead vary independently of each other. The functional and phylogenetic implications of our findings are discussed.

paleontology↗

bioRxiv2021

Dinosaur bonebed amber from an original swamp forest soil

Dinosaur bonebeds with amber content, yet scarce, offer a superior wealth and quality of data on ancient terrestrial ecosystems. However, the preserved palaeodiversity and/or taphonomic characteristics of these exceptional localities had hitherto limited their palaeobiological potential. Here we describe the amber from the Lower Cretaceous dinosaur bonebed of Arino (Teruel, Spain) using a multidisciplinary approach. Amber is found in both a root layer with amber strictly in situ and a litter layer namely composed of aerial pieces unusually rich in bioinclusions, encompassing 11 insect orders, arachnids, and a few plant and vertebrate remains, including a feather. Additional palaeontological data - charophytes, palynomorphs, ostracods- are provided. Arino arguably represents the most prolific and palaeobiologically diverse locality in which fossiliferous amber and a dinosaur bonebed have been found in association, and the only one known where the vast majority of the palaeontological assemblage suffered no or low-grade pre-burial transport. That has enabled unlocking unprecedentedly complete and reliable palaeoecological data out of two complementary windows of preservation from the same site.

paleontology↗

bioRxiv2021

The scratch-digging lifestyle of the Permian "microsaur" Batropetes as a model for the exaptative origin of jumping locomotion in frogs

Recent studies have shown that the Triassic stem-frog Triadobatrachus lacked the ability to jump off, but nonetheless had the forelimb strength to withstand the impact of landing from a jump. We propose a hypothesis to resolve this pseudoparadox: the strengthened forelimbs are former adaptations to forelimb-based digging that later made jumping possible by exaptation. Micro-CT data from a skeleton of Batropetes palatinus reveal thin cortical bone, confirming Batropetes as terrestrial. Combining adaptations to walking and digging, confirmed by statistical analyses, Batropetes is thought to have searched for food in leaf litter or topsoil. We interpret Batropetes as having used one forelimb at a time to shove leaf litter aside. Batropetes may thus represent an analog or possibly a homolog of the digging stage that preceded the origin of Salientia. We discuss the possibility of homology with the digging lifestyles of other "microsaurs" and other amphibians.

paleontology↗

bioRxiv2021

Belemnite phylogeny and decline during the mid-Cretaceous

Belemnites are common fossil coleoid cephalopods of the Mesozoic. They began to diversify in the Triassic-Early Jurassic and maintained this diversity until the early Early Cretaceous. During the mid-Cretaceous, they declined in diversity and distribution, being restricted to only the Boreal and Austral Realm since the Turonian. Here, I present the first cladistic analysis of belemnite phylogeny, spanning taxa representative of the whole diversity and stratigraphic range of the group. This analysis shows that the usually applied subdivision of all belemnites into "Belemnitina" and "Belemnopseina" is not supported. A newly identified clade, the Pseudoalveolata, is suggested here. Pseudoalveolate belemnites represent the last remaining belemnites after the Aptian. Oceanic anoxia and warming are likely the main cause of the mid-Cretaceous belemnite decline, resulting in the Aptian-Albian dominance of the warm-adapted pseudoalveolate genus Neohibolites. The rise of teleost fish diversity during the mid-Cretaceous is discussed and its relevance for belemnite evolution. Some teleosts (e.g., Enchodus) might have taken over the mesopredator niches left by belemnites during the mid-Cretaceous, being better adapted to warming seas. Belemnites were not able to recover their earlier widespread distribution and diversity and the last remaining, disjunctly distributed families, the northern Belemnitellidae and southern Dimitobelidae, became extinct at the K/Pg-boundary.

paleontology↗

bioRxiv2021

Early evolution of beetles regulated by the end-Permian deforestation

The end-Permian mass extinction (EPME) led to a severe terrestrial ecosystem collapse. However, the ecological response of insects--the most diverse group of organisms on Earth--to the EPME remains poorly understood. Here, we analyse beetle evolutionary history based on taxonomic diversity, morphological disparity, phylogeny, and ecological shifts from the Early Permian to Middle Triassic, using a comprehensive new data set. Permian beetles were dominated by xylophagous stem groups with a high diversity and disparity, which probably played an underappreciated role in the Permian carbon cycle. Our suite of analyses shows that Permian xylophagous beetles suffered a severe extinction during the EPME largely due to the collapse of forest ecosystems, resulting in an Early Triassic gap of xylophagous beetles. New xylophagous beetles appeared widely in the early Middle Triassic, which is consistent with the restoration of forest ecosystems. Our results highlight the ecological significance of insects in deep-time terrestrial ecosystems.

paleontology↗

bioRxiv2021

Relationships between the hard and soft dimensions of the nose in Pan troglodytes and Homo sapiens reveal the nasal protrusions of Plio-Pleistocene hominids

By identifying similarity in bone and soft tissue covariation patterns in hominids, it is possible to produce facial approximation methods that are compatible with more than one species of primate. In this study, we conducted an interspecific comparison of the nasomaxillary region in chimpanzees and modern humans with the aim of producing a method for predicting the nasal protrusions of ancient Plio-Pleistocene hominids. We addressed this aim by first collecting and performing regression analyses of linear and angular measurements of nasal cavity length and inclination in modern humans (Homo sapiens; n = 72) and chimpanzees (Pan troglodytes; n = 19), and then by performing a set of out-of-group tests. The first test was performed on two subjects that belonged to the same genus as the training sample, i.e., Homo (n = 1) and Pan (n = 1), and the second test, which functioned as an interspecies compatibility test, was performed on Pan paniscus (n = 1), Gorilla gorilla (n = 3), Pongo pygmaeus (n = 1), Pongo abelli (n = 1), Symphalangus syndactylus (n = 3), and Papio hamadryas (n = 3). We identified statistically significant correlations in both humans and chimpanzees with slopes that displayed homogeneity of covariation. Joint prediction formulae were found to be compatible with humans and chimpanzees as well as all other African great apes, i.e., bonobos and gorillas. The main conclusion that can be drawn from this study is that regression models for approximating nasal projection are homogenous among humans and African apes and can thus be reasonably extended to ancestors leading to these clades.

paleontology↗

bioRxiv2021

On the dynamics of spatial updating

Most of our knowledge on the human neural bases of spatial updating comes from fMRI studies in which recumbent participants moved in virtual environments. As a result, little is known about the dynamic of spatial updating during real body motion. Here, we exploited the high temporal resolution of electroencephalography (EEG) to investigate the dynamics of cortical activation in a spatial updating task where participants had to remember their initial orientation while they were passively rotated about their vertical axis in the dark. After the rotations, the participants pointed towards their initial orientation. We contrasted the EEG signals with those recorded in a control condition in which participants had no cognitive task to perform during body rotations. We found that the amplitude of the P1N1 complex of the rotation-evoked potential (RotEPs) (recorded over the vertex) was significantly greater in the Updating task. The analyses of the cortical current in the source space revealed that the main significant task-related cortical activities started during the N1P2 interval (136-303 ms after rotation onset). They were essentially localised in the temporal and frontal (supplementary motor complex, dorsolateral prefrontal cortex, anterior prefrontal cortex) regions. During this time-window, the right superior posterior parietal cortex (PPC) also showed significant task-related activities. The increased activation of the PPC became bilateral over the P2N2 component (303-470 ms after rotation onset). In this late interval, the cuneus and precuneus started to show significant task-related activities. Together, the present results are consistent with the general scheme that the first task-related cortical activities during spatial updating are related to the encoding of spatial goals and to the storing of spatial information in working memory. These activities would precede those involved in higher order processes also relevant for updating body orientation during rotations linked to the egocentric and visual representations of the environment.

paleontology↗

bioRxiv2021

Linking host plants to damage types in the fossil record of insect herbivory

Studies of insect herbivory on fossilized leaves tend to focus on a few, relatively simple metrics that are agnostic to the distribution of insect damage types among host plants. More complex metrics that link particular damage types to particular host plants have the potential to address additional ecological questions, but such metrics can be biased by sampling incompleteness due to the difficulty of distinguishing the true absence of a particular interaction from the failure to detect it--a challenge that has been raised in the ecological literature. We evaluate a range of methods for characterizing the relationships between damage types and host plants by performing resampling and subsampling exercises on a variety of datasets. We found that the components of beta diversity provide a more valid, reliable, and interpretable method for comparing component communities than do bipartite network metrics. We found the rarefaction of interactions to be a valid, reliable, and interpretable method for comparing compound communities. Both beta diversity and rarefaction of interactions avoid the potential pitfalls of multiple comparisons. Lastly, we found that the host specificity of individual damage types is challenging to assess. Whereas some previously used methods are sufficiently biased by sampling incompleteness to be inappropriate for fossil herbivory data, alternatives exist that are perfectly suitable for fossil datasets with sufficient sample coverage.

paleontology↗

bioRxiv2021

High diversity of pimpline parasitoid wasps (Hymenoptera, Ichneumonidae, Pimplinae) from the lowermost Eocene Fur Formation

With an estimated 100,000 extant species, Darwin wasps (Ichneumonidae) are more specious than all vertebrates together. However, only 288 fossil species have been described to date, with hundreds more awaiting formal description in palaeontological collections. One of the largest gaps in our knowledge concerns the ~12 million years between the K-PG mass extinction and the late Ypresian, from which only two species have been formally described, including Pimpla stigmatica Henriksen from the Danish Fur Formation (~55 Ma). Ie here redescribe and reclassify this species in the genus Epitheronia Gupta, n. comb., and describe nine new species from this fossil locality that are consistent with a placement in Pimplinae: Crusopimpla collina n. sp., C. elongata n. sp., C. minuta n. sp., C. rettigi n. sp., C. violina n. sp., Theronia? furensis n. sp., T. nigriscutum n. sp., Xanthopimpla ciboisae n. sp., and X. crescendae n. sp. The diagnosis of the genus Crusopimpla Kopylov, Spasojevic & Klopfstein is amend in the light of the new species. By comparing the preserved colouration between and within specimens of different species, we draw conclusions about the taxonomic usefulness of colour patterns observed in Fur Formation ichneumonids. The number of described species of pimpline parasitoid wasps from Fur is very high when compared to any other fossil deposit, but low with respect to numbers of extant species. Further study and excavation of Fur ichneumonids will certainly reveal an even higher diversity.

paleontology↗

bioRxiv2021

New fossil wasp species from the earliest Eocene Fur Formation has its closest relatives in late Eocene ambers (Hymenoptera, Ichneumonidae, Pherhombinae)

Darwin wasps (Ichneumonidae) are one of the most species-rich insect families, but also one of the most understudied ones, both in terms of their extant and extinct diversity. We here use morphometrics of wing veins and an integrative Bayesian analysis to place a new rock fossil species from the Danish Fur Formation ([~]54 Ma) in the tree of Darwin wasps. The new species, Pherhombus parvulus n. sp., is placed firmly in Pherhombinae, an extinct subfamily so far only known from Baltic and Rovno-Ukranian ambers, which are estimated to be 34-48 Ma and 34-38 Ma, respectively. Our phylogenetic analysis recovers a subfamily clade within the higher Ophioniformes formed by Pherhombinae, Townesitinae and Hybrizontinae, in accordance with previous suggestions. Due to the placement of the new species as sister to the remaining members of Pherhombinae, we argue that our finding is not at odds with a much younger, late Eocene age ([~]34-41 Ma) of Baltic amber and instead demonstrates that Pherhombus existed over a much longer period than previously thought. Our results also exemplify the power of wing vein morphometrics and integrative phylogenetic analyses in resolving the placement even of poorly preserved fossil specimens.

paleontology↗

bioRxiv2021

A community-developed extension to Darwin Core for reporting the chronometric age of specimens

Darwin Core, the data standard used for sharing modern biodiversity and paleodiversity occurrence records, has previously lacked proper mechanisms for reporting what is known about the estimated age range of specimens from deep time. This has led to data providers putting these data in fields where they cannot easily be found by users, which impedes the reuse and improvement of these data by other researchers. Here we describe the development of the Chronometric Age Extension to Darwin Core, a ratified, community-developed extension that enables the reporting of ages of specimens from deeper time and the evidence supporting these estimates. The extension standardizes reporting about the methods or assays used to determine an age and other critical information like uncertainty. It gives data providers flexibility about the level of detail reported, focusing on the minimum information needed for reuse while still allowing for significant detail if providers have it. Providing a standardized format for reporting these data will make them easier to find and search and enable researchers to pinpoint specimens of interest for data improvement or accumulate more data for broad temporal studies. The Chronometric Age Extension was also the first community-managed vocabulary to undergo the new Biodiversity Informatics Standards (TDWG) review and ratification process, thus providing a blueprint for future Darwin Core extension development.

paleontology↗

bioRxiv2021

A new early-branching armoured dinosaur from the Lower Jurassic of southwestern China

The early evolutionary history of the armoured dinosaurs (Thyreophora) is obscured by its patchily distributed fossil record and by conflicting views on the relationships of its Early Jurassic representatives. Here, we describe an early-diverging thyreophoran from the Lower Jurassic Fengjiahe Formation of Yunnan Province, China, on the basis of an associated partial skeleton that includes skull, axial, limb and armour elements. It can be diagnosed as a new taxon based on numerous cranial and postcranial autapomorphies and is further distinguished from all other thyreophorans by a unique combination of character states. Although the robust postcranium is similar to that of more deeply nested ankylosaurs and stegosaurs, phylogenetic analysis recovers it as either the sister taxon of Emausaurus or of the clade Scelidosaurus+Eurypoda. This new taxon, XXXX XXXX, represents the first valid thyreophoran dinosaur to be described from the Early Jurassic of Asia and confirms the rapid geographic spread and diversification of the clade after its first appearance in the Hettangian. Its heavy build and distinctive armour also hint at previously unrealised morphological diversity early in the clades history.

paleontology↗

bioRxiv2021

An early Cambrian polyp reveals an anemone-like ancestor for medusozoan cnidarians

Extant cnidarians are a disparate phylum of non-bilaterians and their diploblastic body plan represents a key step in animal evolution. Anthozoans (anemones, corals) are benthic polyps, while adult medusozoans (jellyfishes) are dominantly pelagic medusae. A sessile polyp is present in both groups and is widely conceived as the ancestral form of their last common ancestor. However, the nature and anatomy of this ancestral polyp, particularly of medusozoans, are controversial, owing to the divergent body plans of both groups in the extant lineages and the rarity of medusozoan soft tissues in the fossil record. Here we redescribe the enigmatic Conicula striata Luo et Hu from the early Cambrian Chengjiang biota, south China, which has previously been interpreted as a polyp, lophophorate or deuterostome. We show that C. striata possessed features of both anthozoans and medusozoans. Its stalked polyp and fully encasing conical, annulated organic skeleton (periderm) are features of medusozoans. However, the gut is partitioned by [~]28 mesenteries, and has a tubular pharynx, resembling anthozoans. Our phylogenetic analysis recovers C. striata as a stem medusozoan, indicating that the enormously diverse medusozoans were derived from an anemone-like ancestor, with the pharynx lost and number of mesenteries reduced prior to the origin of crown group Medusozoa.

paleontology↗

bioRxiv2022

Global ecomorphological restructuring of dominant marine reptiles prior to the K/Pg mass extinction

Mosasaurid squamates were the dominant amniote predators in marine ecosystems during most of the Late Cretaceous. Evidence from multiple sites worldwide of a global mosasaurid community restructuring across the Campanian-Maastrichtian transition may have wide-ranging implications for the evolution of diversity of these top oceanic predators. In this study, we use a suite of biomechanical traits and functionally descriptive ratios to investigate how the morphofunctional disparity of mosasaurids evolved through time and space prior to the Cretaceous-Palaeogene (K/Pg) mass extinction. Our results suggest that the worldwide taxonomic turnover in mosasaurid community composition from Campanian to Maastrichtian is reflected by a notable increase in morphofunctional disparity on a global scale, but especially driven the North American record. Ecomorphospace occupation becomes more polarised during the late Maastrichtian, as the morphofunctional disparity of mosasaurids plateaus in the Southern Hemisphere and decreases in the Northern Hemisphere. We show that these changes are not associated with strong modifications in mosasaurid size, but rather with the functional capacities of their skulls, and that mosasaurid morphofunctional disparity was in decline in several provincial communities before the K-Pg mass extinction. Our study highlights region-specific patterns of disparity evolution, and the importance of assessing vertebrate extinctions both globally and regionally. Ecomorphological differentiation in mosasaurid communities, coupled with declines in other formerly abundant marine reptile groups, indicates widespread restructuring of higher trophic levels in marine food webs was well underway when the K-Pg mass extinction took place.

paleontology↗

bioRxiv2022

40 new specimens of Ichthyornis provide unprecedented insight into the postcranial morphology of crownward stem group birds.

Ichthyornis has long been recognized as a pivotally important fossil taxon for understanding the latest stages of the dinosaur-bird transition, but little significant new postcranial material has been brought to light since initial descriptions of partial skeletons in the 19th Century. Here, we present new information on the postcranial morphology of Ichthyornis from 40 previously undescribed specimens, providing the most detailed morphological assessment of Ichthyornis to date. The new material includes four partially complete skeletons and numerous well-preserved isolated elements, enabling new anatomical observations such as muscle attachments previously undescribed for Mesozoic euornitheans. Among the elements that were previously unknown or poorly represented for Ichthyornis, the new specimens include an almost-complete axial series, a hypocleideum-bearing furcula, radial carpal bones, fibulae, a complete tarsometatarsus bearing a rudimentary hypotarsus, and one of the first-known nearly complete three-dimensional sterna from a Mesozoic avialan. Several pedal phalanges are preserved, revealing a remarkably enlarged pes presumably related to foot-propelled swimming. Although diagnosable as Ichthyornis, the new specimens exhibit a substantial degree of morphological variation, some of which may relate to ontogenetic changes. Phylogenetic analyses incorporating our new data and employing alternative morphological datasets recover Ichthyornis stemward of Hesperornithes and Iaceornis, in line with some recent hypotheses regarding the topology of the crownward-most portion of the avian stem group, and we establish phylogenetically-defined clade names for relevant avialan subclades to help facilitate consistent discourse in future work. The new information provided by these specimens improves our understanding of morphological evolution among the crownward-most non-neornithine avialans immediately preceding the origin of crown group birds.

paleontology↗

bioRxiv2022

Computed tomographic analysis of the dental system of three Jurassic ceratopsians: implications for the evolution of the tooth replacement pattern and diet in early-diverging ceratopsians

The dental system of ceratopsids is among the most specialized structure in Dinosauria, and includes high angled wear surfaces, split tooth roots, and multiple teeth in each tooth family. However, the early evolution of this unique dental system is generally poorly understood due to a lack of knowledge of the dental morphology and development in early-diverging ceratopsians. Here we study the dental system of three of the earliest-diverging Chinese ceratopsians: Yinlong and Hualianceratops from the early Late Jurassic of Xinjiang, and Chaoyangsaurus from the Late Jurassic of Liaoning. By using micro-computed tomographic analyses, our study has revealed significant new information regarding the dental system of these early ceratopsians, including no more than five replacement teeth in each jaw quadrant; at most one generation of replacement teeth in each alveolus; nearly full resorption of the functional tooth root during tooth replacement; and occlusion with low-angled, concave wear facets that differs significantly from the shearing occlusal system seen in ceratopsids. Yinlong displays an increase in the number of maxillary tooth alveoli and a decrease in the number of replacement teeth during ontogeny as well as the retention of remnants of functional teeth in the largest individual. Early-diverging ceratopsians thus display a relatively slow tooth replacement rate compared to late-diverging ceratopsians. Combined with paleobotany and palaeoenvironment data, Yinlong likely uses gastroliths to triturate foodstuffs, and the difference in diet strategy might have influenced the pattern of tooth replacement in later-diverging ceratopsians.

paleontology↗

bioRxiv2022

AI-based graptolite identification improves shale gas exploration

Graptolites are fossils from the mid-Cambrian to lower Carboniferous periods that inform both our understanding of evolution and the exploration of shale gas [1-4]. The identification of graptolite species remains a visual task carried out by experienced taxonomists because their fine-grained morphologies and incomplete preservation in multi-fossil samples have hindered automation. Artificial intelligence (AI) holds great promise for transforming such meticulous tasks, and has already proven useful in applications ranging from animal classification to medical diagnostics [5-15]. Here we demonstrate that graptolites can be identified with taxonomist accuracy using a deep learning model [16-18]. We develop a convolutional neural network to classify macrofossils, and construct a comprehensive dataset of >34,000 images of 113 graptolite species annotated at pixel-level resolution to train the model. We validate the models performance by comparing its ability to identify 100 images of graptolite species that are significant for rock dating and shale gas exploration with 21 experienced taxonomists from research institutes and the shale gas industry. Our model achieves 86% and 81% accuracy when identifying the genus and species of graptolites, respectively; outperforming taxonomists in terms of accuracy, time, and generalization. By investigating the decisions made by the neural network, we further show that it can recognise fine-grained morphological details better than taxonomists. Our AI approach, providing taxonomist-level graptolite identification, can be deployed on web and mobile apps to extend graptolite identification beyond research institutes and improve the efficiency of shale gas exploration.

paleontology↗