PeerJ:Paleontologyhttps://peerj.com/articles/index.atom?journal=peerj&subject=2200Paleontology articles published in PeerJTransformation of the pectoral girdle in pennaraptorans: critical steps in the formation of the modern avian shoulder jointhttps://peerj.com/articles/169602024-02-292024-02-29Qian WuJingmai K. O’ConnorShiying WangZhonghe Zhou
Important transformations of the pectoral girdle are related to the appearance of flight capabilities in the Dinosauria. Previous studies on this topic focused mainly on paravians yet recent data suggests flight evolved in dinosaurs several times, including at least once among non-avialan paravians. Thus, to fully explore the evolution of flight-related avian shoulder girdle characteristics, it is necessary to compare morphology more broadly. Here, we present information from pennaraptoran specimens preserving pectoral girdle elements, including all purportedly volant taxa, and extensively compare aspects of the shoulder joint. The results show that many pectoral girdle modifications appear during the evolution from basal pennaraptorans to paravians, including changes in the orientation of the coracoid body and the location of the articulation between the furcula and scapula. These modifications suggest a change in forelimb range of motion preceded the origin of flight in paravians. During the evolution of early avialans, additional flight adaptive transformations occur, such as the separation of the scapula and coracoid and reduction of the articular surface between these two bones, reduction in the angle between these two elements, and elongation of the coracoid. The diversity of coracoid morphologies and types of articulations joining the scapula-coracoid suggest that each early avialan lineage evolved these features in parallel as they independently evolved more refined flight capabilities. In early ornithothoracines, the orientation of the glenoid fossa and location of the acrocoracoid approaches the condition in extant birds, suggesting a greater range of motion in the flight stroke, which may represent the acquisition of improved powered flight capabilities, such as ground take-off. The formation of a new articulation between the coracoid and furcula in the Ornithuromorpha is the last step in the formation of an osseous triosseal canal, which may indicate the complete acquisition of the modern flight apparatus. These morphological transitions equipped birds with a greater range of motion, increased and more efficient muscular output and while at the same time transmitting the increased pressure being generated by ever more powerful flapping movements in such a way as to protect the organs. The driving factors and functional adaptations of many of these transitional morphologies are as yet unclear although ontogenetic transitions in forelimb function observed in extant birds provide an excellent framework through which we can explore the behavior of Mesozoic pennaraptorans.
Important transformations of the pectoral girdle are related to the appearance of flight capabilities in the Dinosauria. Previous studies on this topic focused mainly on paravians yet recent data suggests flight evolved in dinosaurs several times, including at least once among non-avialan paravians. Thus, to fully explore the evolution of flight-related avian shoulder girdle characteristics, it is necessary to compare morphology more broadly. Here, we present information from pennaraptoran specimens preserving pectoral girdle elements, including all purportedly volant taxa, and extensively compare aspects of the shoulder joint. The results show that many pectoral girdle modifications appear during the evolution from basal pennaraptorans to paravians, including changes in the orientation of the coracoid body and the location of the articulation between the furcula and scapula. These modifications suggest a change in forelimb range of motion preceded the origin of flight in paravians. During the evolution of early avialans, additional flight adaptive transformations occur, such as the separation of the scapula and coracoid and reduction of the articular surface between these two bones, reduction in the angle between these two elements, and elongation of the coracoid. The diversity of coracoid morphologies and types of articulations joining the scapula-coracoid suggest that each early avialan lineage evolved these features in parallel as they independently evolved more refined flight capabilities. In early ornithothoracines, the orientation of the glenoid fossa and location of the acrocoracoid approaches the condition in extant birds, suggesting a greater range of motion in the flight stroke, which may represent the acquisition of improved powered flight capabilities, such as ground take-off. The formation of a new articulation between the coracoid and furcula in the Ornithuromorpha is the last step in the formation of an osseous triosseal canal, which may indicate the complete acquisition of the modern flight apparatus. These morphological transitions equipped birds with a greater range of motion, increased and more efficient muscular output and while at the same time transmitting the increased pressure being generated by ever more powerful flapping movements in such a way as to protect the organs. The driving factors and functional adaptations of many of these transitional morphologies are as yet unclear although ontogenetic transitions in forelimb function observed in extant birds provide an excellent framework through which we can explore the behavior of Mesozoic pennaraptorans.Downsizing a heavyweight: factors and methods that revise weight estimates of the giant fossil whale Perucetus colossushttps://peerj.com/articles/169782024-02-292024-02-29Ryosuke MotaniNicholas D. Pyenson
Extremes in organismal size have broad interest in ecology and evolution because organismal size dictates many traits of an organism’s biology. There is particular fascination with identifying upper size extremes in the largest vertebrates, given the challenges and difficulties of measuring extant and extinct candidates for the largest animal of all time, such as whales, terrestrial non-avian dinosaurs, and extinct marine reptiles. The discovery of Perucetus colossus, a giant basilosaurid whale from the Eocene of Peru, challenged many assumptions about organismal extremes based on reconstructions of its body weight that exceeded reported values for blue whales (Balaenoptera musculus). Here we present an examination of a series of factors and methodological approaches to assess reconstructing body weight in Perucetus, including: data sources from large extant cetaceans; fitting published body mass estimates to body outlines; testing the assumption of isometry between skeletal and body masses, even with extrapolation; examining the role of pachyostosis in body mass reconstructions; addressing method-dependent error rates; and comparing Perucetus with known physiological and ecological limits for living whales, and Eocene oceanic productivity. We conclude that Perucetus did not exceed the body mass of today’s blue whales. Depending on assumptions and methods, we estimate that Perucetus weighed 60–70 tons assuming a length 17 m. We calculated larger estimates potentially as much as 98–114 tons at 20 m in length, which is far less than the direct records of blue whale weights, or the 270 ton estimates that we calculated for body weights of the largest blue whales measured by length.
Extremes in organismal size have broad interest in ecology and evolution because organismal size dictates many traits of an organism’s biology. There is particular fascination with identifying upper size extremes in the largest vertebrates, given the challenges and difficulties of measuring extant and extinct candidates for the largest animal of all time, such as whales, terrestrial non-avian dinosaurs, and extinct marine reptiles. The discovery of Perucetus colossus, a giant basilosaurid whale from the Eocene of Peru, challenged many assumptions about organismal extremes based on reconstructions of its body weight that exceeded reported values for blue whales (Balaenoptera musculus). Here we present an examination of a series of factors and methodological approaches to assess reconstructing body weight in Perucetus, including: data sources from large extant cetaceans; fitting published body mass estimates to body outlines; testing the assumption of isometry between skeletal and body masses, even with extrapolation; examining the role of pachyostosis in body mass reconstructions; addressing method-dependent error rates; and comparing Perucetus with known physiological and ecological limits for living whales, and Eocene oceanic productivity. We conclude that Perucetus did not exceed the body mass of today’s blue whales. Depending on assumptions and methods, we estimate that Perucetus weighed 60–70 tons assuming a length 17 m. We calculated larger estimates potentially as much as 98–114 tons at 20 m in length, which is far less than the direct records of blue whale weights, or the 270 ton estimates that we calculated for body weights of the largest blue whales measured by length.Plectronoceratids (Cephalopoda) from the latest Cambrian at Black Mountain, Queensland, reveal complex three-dimensional siphuncle morphology, with major taxonomic implicationshttps://peerj.com/articles/170032024-02-292024-02-29Alexander PohlePeter JellChristian Klug
The Plectronoceratida includes the earliest known cephalopod fossils and is thus fundamental to a better understanding of the origin and early evolution of this group of molluscs. The bulk of described material comes from the late Cambrian Fengshan Formation in North China with isolated occurrences in South China, Laurentia, Kazakhstan and Siberia. Knowledge of their morphology and taxonomy is limited in that most specimens were only studied as longitudinal sections, which are prone to misinterpretations due to variations in the plane of section. We describe more than 200 new specimens, which exceeds the entire hitherto published record of plectronoceratids. The material was collected by Mary Wade and colleagues during the 1970s and 1980s, from the lower Ninmaroo Formation at Black Mountain (Mount Unbunmaroo), Queensland, Australia. Despite the collecting effort, diverse notes and early incomplete drafts, Mary Wade never published this material before her death in 2005. The specimens provide novel insights into the three-dimensional morphology of the siphuncle based on abundant material, prompting a general revision of the order Plectronoceratida. We describe Sinoeremoceras marywadeae sp. nov. from numerous, well-preserved specimens, allowing investigation of ontogenetic trajectories and intraspecific variability, which in turn enables improved interpretations of the three-dimensional siphuncle morphology. The siphuncle of S. marywadeae sp. nov. and other plectronoceratids is characterised by highly oblique segments, an elongated middorsal portion of the septal neck (= septal flap) and laterally expanded segments that extend dorsally relative to the septal flap (= siphuncular bulbs). We show that this complex siphuncular structure has caused problems of interpretation because it was studied mainly from longitudinal sections, leading to the impression that there were large differences between specimens and supposed species. We revise the order Protactinoceratida and the families Protactinoceratidae and Balkoceratidae as junior synonyms of the Plectronoceratida and Plectronoceratidae, respectively. We reduce the number of valid genera from eighteen (including one genus formerly classified as an ellesmeroceratid) to three: Palaeoceras Flower, 1954, Plectronoceras Kobayashi, 1935 and Sinoeremoceras Kobayashi, 1933. We accept 10 valid species to which the 68 previously established species may be assigned. Sinoeremoceras contains 8 of the 10 plus the new species. Two species, previously referred to ellesmeroceratid genera, are transferred to Sinoeremoceras. This revised scheme groups plectronoceratids into distinct geographically and stratigraphically separated species, which better reflects biological realities and removes bias caused by preparation techniques. North China remains important containing the highest known diversity and was likely a centre of cephalopod diversification.
The Plectronoceratida includes the earliest known cephalopod fossils and is thus fundamental to a better understanding of the origin and early evolution of this group of molluscs. The bulk of described material comes from the late Cambrian Fengshan Formation in North China with isolated occurrences in South China, Laurentia, Kazakhstan and Siberia. Knowledge of their morphology and taxonomy is limited in that most specimens were only studied as longitudinal sections, which are prone to misinterpretations due to variations in the plane of section. We describe more than 200 new specimens, which exceeds the entire hitherto published record of plectronoceratids. The material was collected by Mary Wade and colleagues during the 1970s and 1980s, from the lower Ninmaroo Formation at Black Mountain (Mount Unbunmaroo), Queensland, Australia. Despite the collecting effort, diverse notes and early incomplete drafts, Mary Wade never published this material before her death in 2005. The specimens provide novel insights into the three-dimensional morphology of the siphuncle based on abundant material, prompting a general revision of the order Plectronoceratida. We describe Sinoeremoceras marywadeae sp. nov. from numerous, well-preserved specimens, allowing investigation of ontogenetic trajectories and intraspecific variability, which in turn enables improved interpretations of the three-dimensional siphuncle morphology. The siphuncle of S. marywadeae sp. nov. and other plectronoceratids is characterised by highly oblique segments, an elongated middorsal portion of the septal neck (= septal flap) and laterally expanded segments that extend dorsally relative to the septal flap (= siphuncular bulbs). We show that this complex siphuncular structure has caused problems of interpretation because it was studied mainly from longitudinal sections, leading to the impression that there were large differences between specimens and supposed species. We revise the order Protactinoceratida and the families Protactinoceratidae and Balkoceratidae as junior synonyms of the Plectronoceratida and Plectronoceratidae, respectively. We reduce the number of valid genera from eighteen (including one genus formerly classified as an ellesmeroceratid) to three: Palaeoceras Flower, 1954, Plectronoceras Kobayashi, 1935 and Sinoeremoceras Kobayashi, 1933. We accept 10 valid species to which the 68 previously established species may be assigned. Sinoeremoceras contains 8 of the 10 plus the new species. Two species, previously referred to ellesmeroceratid genera, are transferred to Sinoeremoceras. This revised scheme groups plectronoceratids into distinct geographically and stratigraphically separated species, which better reflects biological realities and removes bias caused by preparation techniques. North China remains important containing the highest known diversity and was likely a centre of cephalopod diversification.The decline and fall of the mammalian stemhttps://peerj.com/articles/170042024-02-272024-02-27Neil Brocklehurst
The mammalian crown originated during the Mesozoic and subsequently radiated into the substantial array of forms now extant. However, for about 100 million years before the crown’s origin, a diverse array of stem mammalian lineages dominated terrestrial ecosystems. Several of these stem lineages overlapped temporally and geographically with the crown mammals during the Mesozoic, but by the end of the Cretaceous crown mammals make up the overwhelming majority of the fossil record. The progress of this transition between ecosystems dominated by stem mammals and those dominated by crown mammals is not entirely clear, in part due to a distinct separation of analyses and datasets. Analyses of macroevolutionary patterns tend to focus on either the Mammaliaformes or the non-mammalian cynodonts, with little overlap in the datasets, preventing direct comparison of the diversification trends. Here I analyse species richness and biogeography of Synapsida as a whole during the Mesozoic, allowing comparison of the patterns in the mammalian crown and stem within a single framework. The analysis reveals the decline of the stem mammals occurred in two discrete phases. The first phase occurred between the Triassic and Middle Jurassic, during which the stem mammals were more restricted in their geographic range than the crown mammals, although within localities their species richness remained at levels seen previously. The second phase was a decline in species richness, which occurred during the Lower Cretaceous. The results show the decline of stem mammals, including tritylodontids and several mammaliaform groups, was not tied to a specific event, nor a gradual decline, but was instead a multiphase transition.
The mammalian crown originated during the Mesozoic and subsequently radiated into the substantial array of forms now extant. However, for about 100 million years before the crown’s origin, a diverse array of stem mammalian lineages dominated terrestrial ecosystems. Several of these stem lineages overlapped temporally and geographically with the crown mammals during the Mesozoic, but by the end of the Cretaceous crown mammals make up the overwhelming majority of the fossil record. The progress of this transition between ecosystems dominated by stem mammals and those dominated by crown mammals is not entirely clear, in part due to a distinct separation of analyses and datasets. Analyses of macroevolutionary patterns tend to focus on either the Mammaliaformes or the non-mammalian cynodonts, with little overlap in the datasets, preventing direct comparison of the diversification trends. Here I analyse species richness and biogeography of Synapsida as a whole during the Mesozoic, allowing comparison of the patterns in the mammalian crown and stem within a single framework. The analysis reveals the decline of the stem mammals occurred in two discrete phases. The first phase occurred between the Triassic and Middle Jurassic, during which the stem mammals were more restricted in their geographic range than the crown mammals, although within localities their species richness remained at levels seen previously. The second phase was a decline in species richness, which occurred during the Lower Cretaceous. The results show the decline of stem mammals, including tritylodontids and several mammaliaform groups, was not tied to a specific event, nor a gradual decline, but was instead a multiphase transition.The first gladius-bearing coleoid cephalopods from the lower Toarcian “Schistes Cartons” Formation of the Causses Basin (southeastern France)https://peerj.com/articles/168942024-02-262024-02-26Romain JattiotNathalie Coquel-PoussyIsabelle KrutaIsabelle RougetAlison J. RoweJean-David Moreau
The fossil record of gladius-bearing coleoids is scarce and based only on a few localities with geological horizons particularly favourable to their preservation (the so-called Konservat-Lagerstätten), which naturally leads to strongly limited data on geographical distributions. This emphasizes the importance of every new locality providing gladius-bearing coleoids. Here, we assess for the first time the gladius-bearing coleoid taxonomic diversity within the lower Toarcian “Schistes Cartons” of the Causses Basin (southeastern France). The material includes two fragmentary gladii, identified as Paraplesioteuthis sagittata and ?Loligosepia sp. indet. Just with these two specimens, two (Prototeuthina and Loligosepiina) of the three (Prototeuthina, Loligosepiina and Teudopseina) suborders of Mesozoic gladius-bearing coleoids are represented. Thus, our results hint at a rich early Toarcian gladius-bearing coleoid diversity in the Causses Basin and point out the need for further field investigations in the lower Toarcian “Schistes Cartons” in this area. This new record of Paraplesioteuthis sagittata is only the second one in Europe and the third in the world (western Canada, Germany and now France). Based on these occurrences, we tentatively suggest that P. sagittata originated in the Mediterranean domain and moved to the Arctic realm through the Viking Corridor to eventually move even farther to North America.
The fossil record of gladius-bearing coleoids is scarce and based only on a few localities with geological horizons particularly favourable to their preservation (the so-called Konservat-Lagerstätten), which naturally leads to strongly limited data on geographical distributions. This emphasizes the importance of every new locality providing gladius-bearing coleoids. Here, we assess for the first time the gladius-bearing coleoid taxonomic diversity within the lower Toarcian “Schistes Cartons” of the Causses Basin (southeastern France). The material includes two fragmentary gladii, identified as Paraplesioteuthis sagittata and ?Loligosepia sp. indet. Just with these two specimens, two (Prototeuthina and Loligosepiina) of the three (Prototeuthina, Loligosepiina and Teudopseina) suborders of Mesozoic gladius-bearing coleoids are represented. Thus, our results hint at a rich early Toarcian gladius-bearing coleoid diversity in the Causses Basin and point out the need for further field investigations in the lower Toarcian “Schistes Cartons” in this area. This new record of Paraplesioteuthis sagittata is only the second one in Europe and the third in the world (western Canada, Germany and now France). Based on these occurrences, we tentatively suggest that P. sagittata originated in the Mediterranean domain and moved to the Arctic realm through the Viking Corridor to eventually move even farther to North America.New records of immature aquatic Diptera from the Foulden Maar Fossil-Lagerstätte, New Zealand, and their biogeographic implicationshttps://peerj.com/articles/170142024-02-262024-02-26Viktor O. BaranovJoachim T. HaugUwe Kaulfuss
Background
The biogeographical and ecological history of true flies (Diptera) in New Zealand is little known due to a scarcity of fossil specimens. Here, we report a fauna of immature aquatic dipterans from freshwater diatomites of the early Miocene Foulden Maar Fossil-Lagerstätte in southern New Zealand.
Methods
We document 30 specimens of immature dipterans, mostly pupae, and compare their external morphology to extant aquatic Diptera. Based on the reconstructed paleoenvironment of Foulden Maar, we discuss taxonomic, ecological and taphonomic implications of this early Miocene fauna.
Results
Among Chironomidae, one pupal morphotype is attributed to Tanypodinae, one pupal morphotype and one larval morphotype are placed into Chironomus (Chironominae) and a further morphotype into Chironominae incertae sedis. Chaoboridae are represented by a pupal morphotype congeneric or very close to the extant Chaoborus, today globally distributed except for New Zealand. Additional immature specimens are likely larvae and puparia of brachyceran flies but cannot be identified to a narrower range. These finds document an aquatic dipteran fauna in New Zealand in the earliest Miocene and highlight Neogene extinction as a factor in shaping the extant Diptera fauna in New Zealand. Immature aquatic dipterans were a common and likely ecologically important component of the early Miocene Foulden Maar lake. Preservation of larvae and pupae may have been promoted by diatomaceous microbial mats and the light colour of the diatomite likely facilitated spotting of these minute fossils in the field.
Background
The biogeographical and ecological history of true flies (Diptera) in New Zealand is little known due to a scarcity of fossil specimens. Here, we report a fauna of immature aquatic dipterans from freshwater diatomites of the early Miocene Foulden Maar Fossil-Lagerstätte in southern New Zealand.
Methods
We document 30 specimens of immature dipterans, mostly pupae, and compare their external morphology to extant aquatic Diptera. Based on the reconstructed paleoenvironment of Foulden Maar, we discuss taxonomic, ecological and taphonomic implications of this early Miocene fauna.
Results
Among Chironomidae, one pupal morphotype is attributed to Tanypodinae, one pupal morphotype and one larval morphotype are placed into Chironomus (Chironominae) and a further morphotype into Chironominae incertae sedis. Chaoboridae are represented by a pupal morphotype congeneric or very close to the extant Chaoborus, today globally distributed except for New Zealand. Additional immature specimens are likely larvae and puparia of brachyceran flies but cannot be identified to a narrower range. These finds document an aquatic dipteran fauna in New Zealand in the earliest Miocene and highlight Neogene extinction as a factor in shaping the extant Diptera fauna in New Zealand. Immature aquatic dipterans were a common and likely ecologically important component of the early Miocene Foulden Maar lake. Preservation of larvae and pupae may have been promoted by diatomaceous microbial mats and the light colour of the diatomite likely facilitated spotting of these minute fossils in the field.Arthrological reconstructions of the pterosaur neck and their implications for the cervical position at resthttps://peerj.com/articles/168842024-02-212024-02-21Richard BuchmannTaissa Rodrigues
The lack of any pterosaur living descendants creates gaps in the knowledge of the biology of this group, including its cervical biomechanics, which makes it difficult to understand their posture and life habits. To mitigate part of this issue, we reconstructed the cervical osteology and arthrology of three pterosaurs, allowing us to make inferences about the position of the neck of these animals at rest. We used scans of three-dimensionally preserved cervical series of Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri for the reconstructions, thus representing different lineages. For the recognition of ligaments, joint cartilages, and levels of overlapping of the zygapophyses, we applied the Extant Phylogenetic Bracket method, based on various extant birds and on Caiman latirostris. We inferred that pterosaur intervertebral joints were probably covered by a thin layer of synovial cartilage whose thickness varied along the neck, being thicker in the posterior region. Ignoring this cartilage can affect reconstructions. According to the vertebral angulation, their neck was slightly sinuous when in rest position. Our analyses also indicate that pterosaurs had segmented and supra-segmented articular cervical ligaments, which could confer stabilization, execute passive forces on the neck and store elastic energy.
The lack of any pterosaur living descendants creates gaps in the knowledge of the biology of this group, including its cervical biomechanics, which makes it difficult to understand their posture and life habits. To mitigate part of this issue, we reconstructed the cervical osteology and arthrology of three pterosaurs, allowing us to make inferences about the position of the neck of these animals at rest. We used scans of three-dimensionally preserved cervical series of Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri for the reconstructions, thus representing different lineages. For the recognition of ligaments, joint cartilages, and levels of overlapping of the zygapophyses, we applied the Extant Phylogenetic Bracket method, based on various extant birds and on Caiman latirostris. We inferred that pterosaur intervertebral joints were probably covered by a thin layer of synovial cartilage whose thickness varied along the neck, being thicker in the posterior region. Ignoring this cartilage can affect reconstructions. According to the vertebral angulation, their neck was slightly sinuous when in rest position. Our analyses also indicate that pterosaurs had segmented and supra-segmented articular cervical ligaments, which could confer stabilization, execute passive forces on the neck and store elastic energy.Resolving taphonomic and preparation biases in silicified faunas through paired acid residues and X-ray microscopyhttps://peerj.com/articles/167672024-02-012024-02-01Gabriel S. JacobsSarah Monique JacquetTara SellyJames D. SchiffbauerJohn Warren Huntley
Paired petrography and acid maceration has shown that preferential silicification of shelly faunas can bias recovery based on taxon and body size. Here, silicified fossils from the Upper Ordovician Edinburg Formation, Strasburg Junction, Virginia, USA, were analyzed using X-ray tomographic microscopy (μCT) in conjunction with recovered residues from acid maceration of the same materials to further examine sources of potential bias. Results reveal that very small (<~1 mm) fossils are poorly resolved in μCT when scanning at lower resolutions (~30 µm), underestimating abundance of taxa including ostracods and bryozoans. Acid maceration, meanwhile, fails to recover poorly silicified fossils prone to disarticulation and/or fragmentation during digestion. Tests for patterns of breakage, however, indicate no significant size or taxonomic bias during extraction. Comparisons of individual fossils from 3-D fossil renders and maceration residues reveal patterns of fragmentation that are taxon-specific and allow the differentiation of biostratinomic and preparational breakage. Multivariate ordinations and cluster analyses of μCT and residue data in general produce concordant results but indicate that the variation in taxonomic composition of our samples is compromised by the resolvability of small size classes in μCT imaging, limiting the utility of this method for addressing paleoecological questions in these specific samples. We suggest that comparability of results will depend strongly on the sample size, taphonomic history, textural, and compositional characteristics of the samples in question, as well as μCT scan parameters. Additionally, applying these methods to different deposits will test the general applicability of the conclusions drawn on the relative strengths and weaknesses of the methods.
Paired petrography and acid maceration has shown that preferential silicification of shelly faunas can bias recovery based on taxon and body size. Here, silicified fossils from the Upper Ordovician Edinburg Formation, Strasburg Junction, Virginia, USA, were analyzed using X-ray tomographic microscopy (μCT) in conjunction with recovered residues from acid maceration of the same materials to further examine sources of potential bias. Results reveal that very small (<~1 mm) fossils are poorly resolved in μCT when scanning at lower resolutions (~30 µm), underestimating abundance of taxa including ostracods and bryozoans. Acid maceration, meanwhile, fails to recover poorly silicified fossils prone to disarticulation and/or fragmentation during digestion. Tests for patterns of breakage, however, indicate no significant size or taxonomic bias during extraction. Comparisons of individual fossils from 3-D fossil renders and maceration residues reveal patterns of fragmentation that are taxon-specific and allow the differentiation of biostratinomic and preparational breakage. Multivariate ordinations and cluster analyses of μCT and residue data in general produce concordant results but indicate that the variation in taxonomic composition of our samples is compromised by the resolvability of small size classes in μCT imaging, limiting the utility of this method for addressing paleoecological questions in these specific samples. We suggest that comparability of results will depend strongly on the sample size, taphonomic history, textural, and compositional characteristics of the samples in question, as well as μCT scan parameters. Additionally, applying these methods to different deposits will test the general applicability of the conclusions drawn on the relative strengths and weaknesses of the methods.Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremityhttps://peerj.com/articles/168212024-01-312024-01-31Ashleigh L.A. WisemanJames P. CharlesJohn R. Hutchinson
The force a muscle generates is dependent on muscle structure, in which fibre length, pennation angle and tendon slack length all influence force production. Muscles are not preserved in the fossil record and these parameters must be estimated when constructing a musculoskeletal model. Here, we test the capability of digitally reconstructed muscles of the Australopithecus afarensis model (specimen AL 288-1) to maintain an upright, single-support limb posture. Our aim was to ascertain the influence that different architectural estimation methods have on muscle specialisation and on the subsequent inferences that can be extrapolated about limb function. Parameters were estimated for 36 muscles in the pelvis and lower limb and seven different musculoskeletal models of AL 288-1 were produced. These parameters represented either a ‘static’ Hill-type muscle model (n = 4 variants) which only incorporated force, or instead a ‘dynamic’ Hill-type muscle model with an elastic tendon and fibres that could vary force-length-velocity properties (n = 3 variants). Each muscle’s fibre length, pennation angle, tendon slack length and maximal isometric force were calculated based upon different input variables. Static (inverse) simulations were computed in which the vertical and mediolateral ground reaction forces (GRF) were incrementally increased until limb collapse (simulation failure). All AL 288-1 variants produced somewhat similar simulated muscle activation patterns, but the maximum vertical GRF that could be exerted on a single limb was not consistent between models. Three of the four static-muscle models were unable to support >1.8 times body weight and produced models that under-performed. The dynamic-muscle models were stronger. Comparative results with a human model imply that similar muscle group activations between species are needed to sustain single-limb support at maximally applied GRFs in terms of the simplified static simulations (e.g., same walking pose) used here. This approach demonstrated the range of outputs that can be generated for a model of an extinct individual. Despite mostly comparable outputs, the models diverged mostly in terms of strength.
The force a muscle generates is dependent on muscle structure, in which fibre length, pennation angle and tendon slack length all influence force production. Muscles are not preserved in the fossil record and these parameters must be estimated when constructing a musculoskeletal model. Here, we test the capability of digitally reconstructed muscles of the Australopithecus afarensis model (specimen AL 288-1) to maintain an upright, single-support limb posture. Our aim was to ascertain the influence that different architectural estimation methods have on muscle specialisation and on the subsequent inferences that can be extrapolated about limb function. Parameters were estimated for 36 muscles in the pelvis and lower limb and seven different musculoskeletal models of AL 288-1 were produced. These parameters represented either a ‘static’ Hill-type muscle model (n = 4 variants) which only incorporated force, or instead a ‘dynamic’ Hill-type muscle model with an elastic tendon and fibres that could vary force-length-velocity properties (n = 3 variants). Each muscle’s fibre length, pennation angle, tendon slack length and maximal isometric force were calculated based upon different input variables. Static (inverse) simulations were computed in which the vertical and mediolateral ground reaction forces (GRF) were incrementally increased until limb collapse (simulation failure). All AL 288-1 variants produced somewhat similar simulated muscle activation patterns, but the maximum vertical GRF that could be exerted on a single limb was not consistent between models. Three of the four static-muscle models were unable to support >1.8 times body weight and produced models that under-performed. The dynamic-muscle models were stronger. Comparative results with a human model imply that similar muscle group activations between species are needed to sustain single-limb support at maximally applied GRFs in terms of the simplified static simulations (e.g., same walking pose) used here. This approach demonstrated the range of outputs that can be generated for a model of an extinct individual. Despite mostly comparable outputs, the models diverged mostly in terms of strength.New materials of plesiacerathere (Perissodactyla, Rhinocerotidae) from the late Early Miocene of Northern Chinahttps://peerj.com/articles/168222024-01-312024-01-31Danhui SunTao DengShiqi Wang
As a member of Aceratheriinae, the genus Plesiaceratherium in Europe is widely distributed and highly diverse. However, only one species of Plesiaceratherium (i.e., P. gracile) exists in China with a discontinuous distribution range. Recently, we have discovered new materials of Plesiaceratherium in the lower layers of the Zhang’enbao Formation exposed in Miaoerling in Tongxin County, China. The new materials are well-preserved and can be separated from other Plesiaceratherium species by the following combination of features: the long and generally flat skull, with closed frontoparietal crests; the deep nasal notch at the level of P4; the high supraorbital margin, with its anterior margin at the level of the M1/M2 boundary; the medium-sized upper I1, with an oval abraded surface; the semi-molarized upper premolars with the protocone and hypocone joined by a lingual bridge; the strong constrictions of protocone on the upper molars; the absent buccal cingulum on upper cheek teeth; the cheek teeth are covered by cement on the buccal walls; the convex base of mandibular corpus; the inclined backward ramus; and the mandibular foramen above the teeth neck. Based on the combination of characteristics and phylogenetic analysis, we herein establish the new species as Plesiaceratherium tongxinense sp. nov. living in the late Early Miocene. Phylogenetic analysis reveals that P. tongxinense is in the basal position of the genus Plesiaceratherium, providing more detailed morphological characteristics of the plesiaceratheres.
As a member of Aceratheriinae, the genus Plesiaceratherium in Europe is widely distributed and highly diverse. However, only one species of Plesiaceratherium (i.e., P. gracile) exists in China with a discontinuous distribution range. Recently, we have discovered new materials of Plesiaceratherium in the lower layers of the Zhang’enbao Formation exposed in Miaoerling in Tongxin County, China. The new materials are well-preserved and can be separated from other Plesiaceratherium species by the following combination of features: the long and generally flat skull, with closed frontoparietal crests; the deep nasal notch at the level of P4; the high supraorbital margin, with its anterior margin at the level of the M1/M2 boundary; the medium-sized upper I1, with an oval abraded surface; the semi-molarized upper premolars with the protocone and hypocone joined by a lingual bridge; the strong constrictions of protocone on the upper molars; the absent buccal cingulum on upper cheek teeth; the cheek teeth are covered by cement on the buccal walls; the convex base of mandibular corpus; the inclined backward ramus; and the mandibular foramen above the teeth neck. Based on the combination of characteristics and phylogenetic analysis, we herein establish the new species as Plesiaceratherium tongxinense sp. nov. living in the late Early Miocene. Phylogenetic analysis reveals that P. tongxinense is in the basal position of the genus Plesiaceratherium, providing more detailed morphological characteristics of the plesiaceratheres.