Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa

We provide the first predictions of bite force (BS) in a wide sample of living and fossil mammalian predators. To compare between taxa, we calculated an estimated bite force quotient (BFQ) as the residual of BS regressed on body mass. Estimated BS adjusted for body mass was higher for marsupials than placentals and the Tasmanian devil (Sarcophilus harrisii) had the highest relative BS among extant taxa. The highest overall BS was in two extinct marsupial lions. BFQ in hyaenas were similar to those of related, non-osteophagous taxa challenging the common assumption that osteophagy necessitates extreme jaw muscle forces. High BFQ in living carnivores was associated with greater maximal prey size and hypercarnivory. For fossil taxa anatomically similar to living relatives, BFQ can be directly compared, and high values in the dire wolf (Canis dirus) and thylacine (Thylacinus cynocephalus) suggest that they took relatively large prey. Direct inference may not be appropriate where morphologies depart widely from biomechanical models evident in living predators and must be considered together with evidence from other morphological indicators. Relatively low BFQ values in two extinct carnivores with morphologies not represented among extant species, the sabrecat, Smilodon fatalis, and marsupial sabretooth, Thylacosmilus atrox, support arguments that their killing techniques also differed from extant species and are consistent with 'canine-shear bite' and 'stabbing' models, respectively. Extremely high BFQ in the marsupial lion, Thylacoleo carnifex, indicates that it filled a large-prey hunting niche.     

Tardigrades as ‘Stem-Group Arthropods’: The Evidence from the Cambrian Fauna

The Cambrian fauna can now reasonably be seen as containing many taxa that lie in the stem-groups of the extant phyla. As such, these fossils suggest how both the ‘body plans’ of extant phyla were assembled, and also how various ‘minor’ phyla relate to the larger groupings of today such as the arthropods and annelids.

The various arthropod and lobopod taxa of the Cambrian faunas have been controversial and have generally been considered either as lying in the crown or (occasionally) stem groups of the euarthropods, onychophorans and tardigrades. However, phylogenetic analysis strongly suggests that many of even the most euarthropod-like taxa do not lie within the euarthropod crown-group but are more basal. Further, the commonly expressed view that Cambrian lobopods are in effect stem- or crown-group onychophorans also seems not to be well supported. Lobopods in the Cambrian appear to be diverse and not particularly closely related to one another, and certainly cannot be combined in a monophyletic clade.

Both these advances offer hope that the tardigrades (placed as the sister group to the euarthropods in many analyses of extant taxa, here collectively named the Tactopoda) may be more closely related to some of these Cambrian taxa than others. The challenge for both neontologists and palaeontologists is to refine the systematic analysis of both living and fossil taxa in order to maximise the usefulness of the (admittedly few) characters that unite tardigrades to their Cambrian forbears.     

Using radii-of-curvature for the reconstruction of extinct South African carnivoran masticatory behavior

Paleoanthropologists have hypothesized that, during the evolution of increased carnivory in our lineage, hominins transitioned through a scavenging niche created by certain carnivoran taxa (especially sabertooths) that may have lacked the morphology necessary to utilize all parts of carcasses, thus leaving an open niche of high-quality scavengable remains. In this article, we examine the postcanine dentition of modern and fossil carnivorans using quantifications of occlusal radii-of-curvature (ROC) and correlate this morphology with feeding behavior to deduce the carcass-processing capabilities of the Plio-Pleistocene carnivores of South Africa. ROC data do a good job of separating taxa by dietary category, revealing possible differences in the carcass-processing abilities of fossil and modern members of some extant species, and confirming that Chasmaporthetes was probably a hypercarnivore and not a durophage like the modern hyenas. Contrary to previous hypotheses, sabertooth felids do not appear to have been more hypercarnivorous than modern felids based on these data.     

First description of milk teeth of fossil South American procyonid from the lower Chapadmalalan (Late Miocene–Early Pliocene) of “Farola Monte Hermoso,” Argentina: paleoecological considerations

The first record of milk teeth of South American fossil procyonids comes from the Late Miocene–Early Pliocene at “Farola Monte Hermoso,” Buenos Aires Province, Argentina. Five extant genera of Procyonidae inhabit South America (Bassaricyon Allen, Nasuella Hollister, Potos Geoffroy Saint-Hilaire and Cuvier, Procyon Storr, and Nasua Storr). Of these only Procyon and Nasua are present in the fossil record (Late Pleistocene–Holocene), in several localities in Brazil, Uruguay, and Bolivia. In addition, six other fossil genera were named, but only two are considered valid: Cyonasua and Chapadmalania. Thus, Cyonasua encompasses ten formally named species and Chapadmalania two. The new specimen, MLP 09-X-5-1, is assigned to cf. Cyonasua. In addition, anatomical evidence implies a much more carnivorous diet in Late Miocene–Early Pleistocene procyonids than that of extant South American taxa. Finally, I examine and discuss the “competitive displacement” hypothesis regarding the extinction of native marsupial carnivores after the arrival of immigrant placental carnivores in South America.     

A decade of progress in plant molecular phylogenetics

Over the past decade, botanists have produced several thousand phylogenetic analyses based on molecular data, with particular emphasis on sequencing rbcL, the plastid gene encoding the large subunit of Rubisco (ribulose bisphosphate carboxylase). Because phylogenetic trees retrieved from the three plant genomes (plastid, nuclear and mitochondrial) have been highly congruent, the ‘Angiosperm Phylogeny Group’ has used these DNA-based phylogenetic trees to reclassify all families of flowering plants. However, in addition to taxonomy, these major phylogenetic efforts have also helped to define strategies to reconstruct the ‘tree of life’, and have revealed the size of the ancestral plant genome, uncovered potential candidates for the ancestral flower, identified molecular living fossils, and linked the rate of neutral substitutions with species diversity. With an increased interest in DNA sequencing programmes in non-model organisms, the next decade will hopefully see these phylogenetic findings integrated into new genetic syntheses, from genomes to taxa.     

The Calcaneum—On the Heels of Marsupial Locomotion

The potential for making functional interpretations from a single postcranial element for marsupials was investigated through morphometric analysis of the calcanea of 61 extant species from Australia and New Guinea. Extant species were grouped into locomotor categories and a canonical variates analysis was carried out on measurements of their calcanea. A relationship between measurements of the calcanea and the locomotor behavior of species was found, allowing for prediction of locomotor behavior from calcaneum morphometrics. This was applied to fossil marsupial taxa, from early–late Miocene/?Pliocene deposits at Riversleigh, in an attempt to determine their locomotor behavior. Hopping (saltatorial) taxa are distinguished from quadruped terrestrial taxa and taxa capable of climbing (arboreal and scansorial) by their relatively longer tuber calcis and wider calcaneal head, by their dorso-ventrally thicker calcaneal head, and by their calcaneocuboid facet being less steeply angled antero-posteriorly. Taxa capable of climbing are distinguished from quadruped terrestrial taxa by their shorter tuber calcis relative to the calcaneal head and by their smaller calcaneo-astragalar facet. The locomotor categories distinguished in this study (arboreal/scansorial, quadruped terrestrial, and saltatorial) highlight differences between species in their use of available substrates and thus are informative with regards to the structural components of their habitat. The results of this analysis can be used, in combination with other data, to make inferences about the habitats of paleocommunities at Riversleigh through the Miocene. The calcaneum is a dense and very robust element and, therefore, has a good chance of being preserved. This method provides a quick and easy way of inferring locomotion and has a wide potential for application to many fossil deposits because it requires only a single element.     

Big cat,small cat: reconstructing body size evolution in living and extinct Felidae

The evolution of body mass is a fundamental topic in evolutionary biology, because it is closely linked to manifold life history and ecological traits and is readily estimable for many extinct taxa. In this study, we examine patterns of body mass evolution in Felidae (Placentalia, Carnivora) to assess the effects of phylogeny, mode of evolution, and the relationship between body mass and prey choice in this charismatic mammalian clade. Our data set includes 39 extant and 26 extinct taxa, with published body mass data supplemented by estimates based on condylobasal length. These data were run through ‘SURFACE’ and ‘bayou’ to test for patterns of body mass evolution and convergence between taxa. Body masses of felids are significantly different among prey choice groupings (small, mixed and large). We find that body mass evolution in cats is strongly influenced by phylogeny, but different patterns emerged depending on inclusion of extinct taxa and assumptions about branch lengths. A single Ornstein–Uhlenbeck optimum best explains the distribution of body masses when first‐occurrence data were used for the fossil taxa. However, when mean occurrence dates or last known occurrence dates were used, two selective optima for felid body mass were recovered in most analyses: a small optimum around 5 kg and a large one around 100 kg. Across living and extinct cats, we infer repeated evolutionary convergences towards both of these optima, but, likely due to biased extinction of large taxa, our results shift to supporting a Brownian motion model when only extant taxa are included in analyses.     

Examination of an Oligocene lacustrine ecosystem using C and N stable isotopes

The Late Oligocene (25.8 Ma) Enspel Fossillagerstätte in Westerwald, Germany, contains a comprehensive fossil ecosystem preserved with specimens retaining morphological detail and a concentration of organic material. Stable carbon and nitrogen isotope analyses were used to examine the lacustrine ecosystem preserved in one stratigraphic horizon. These data suggest the presence of several trophic levels, including primary producers (diatoms and higher plants), primary consumers (e.g., tadpoles and insects), and secondary consumers (e.g., the fish species Paleorutilus enspelensis). Terrigenous and aquatic plants were associated with the lowest δ¹³C and δ¹⁵N values (mean plant = − 26.28‰ ± 0.45, 3.18‰ ± 1.04), primary consumers such as flies are one trophic level higher, and carnivores such as fish are yet another level higher. The δ¹⁵N values for P. enspelensis also showed enrichment in ¹⁵N with increasing body length, implying a shift in diet or feeding strategy with size. P. enspelensis and tadpole (Pelobates decheni) samples showed intraorganism fractionation between ‘muscle’ and ‘bone’ tissues. Stable carbon and nitrogen isotope data from the measurement of components (shale, leaves and seeds) common to a number of different stratigraphic horizons showed significant variation between horizons. A number of the features of the stable isotopic data are similar to those relationships seen in modern ecosystems and therefore suggest that stable isotope analyses can contribute to understanding ancient ecosystems.     

Ancestral state reconstruction of body size in the Caniformia (Carnivora, Mammalia): the effects of incorporating data from the fossil record

A recent molecular phylogeny of the mammalian order Carnivora implied large body size as the ancestral condition for the caniform subclade Arctoidea using the distribution of species mean body sizes among living taxa. "Extant taxa-only" approaches such as these discount character state observations for fossil members of living clades and completely ignore data from extinct lineages. To more rigorously reconstruct body sizes of ancestral forms within the Caniformia, body size and first appearance data were collected for 149 extant and 367 extinct taxa. Body sizes were reconstructed for four ancestral nodes using weighted squared-change parsimony on log-transformed body mass data. Reconstructions based on extant taxa alone favored large body sizes (on the order of 10 to 50 kg) for the last common ancestors of both the Caniformia and Arctoidea. In contrast, reconstructions incorporating fossil data support small body sizes (< 5 kg) for the ancestors of those clades. When the temporal information associated with fossil data was discarded, body size reconstructions became ambiguous, demonstrating that incorporating both character state and temporal information from fossil taxa unambiguously supports a small ancestral body size, thereby falsifying hypotheses derived from extant taxa alone. Body size reconstructions for Caniformia, Arctoidea, and Musteloidea were not sensitive to potential errors introduced by uncertainty in the position of extinct lineages relative to the molecular topology, or to missing body size data for extinct members of an entire major clade (the aquatic Pinnipedia). Incorporating character state observations and temporal information from the fossil record into hypothesis testing has a significant impact on the ability to reconstruct ancestral characters and constrains the range of potential hypotheses of character evolution. Fossil data here provide the evidence to reliably document trends of both increasing and decreasing body size in several caniform clades. More generally, including fossils in such analyses incorporates evidence of directional trends, thereby yielding more reliable ancestral character state reconstructions.     

Use of gas chromatograms of essential leaf oils to compare eight taxa of genus Angophora (Myrtaceae): possible relationships to the genus Eucalyptus

Essential oils were extracted from leaves of eight taxa of the genus Angophora, and then analysed. As expected the individual components of these oils were essentially the same as those found in the Eucalyptus species of our earlier studies (Bignell et al., 1997b. Flavour Fragrance J. 12, 423-432). In addition, as is also the case with the bloodwood eucalypts, only relatively low yields of oil were obtained. In all cases the Cineole component was extremely small, but the oils of six of the eight contained very large concentrations of the sesquiterpene Bicyclogermacrene. A table of the 52 major oil components is included. Principal components analysis (PCA) was performed on the gas chromatograms (GC) of the essential oils and the resulting scores plots compared with the cladistic classification of Thiele and Ladiges (1988). Cladistics 4, 23–42). Because of the close relationship between genus Angophora and Eucalyptus ‘subgenus’ Corymbia, the GC data for the eight Angophora taxa were combined with corresponding data for eleven randomly chosen taxa from ‘subgenus’ Corymbia (Bignell et al., 1996b, Flavour Fragrance J. 11, 339–347; 1997a, Flavour Fragnance J. 12, 277–284) and a PCA performed on the total system.     

Parallel evolution in the hominoid trunk and forelimb

The evolutionary history of the living hominoids has remained elusive despite years of exploration and the discovery of numerous Miocene fossil ape species. Part of the difficulty can be attributed to the changing nature of our views about the course of hominoid evolution. In the 1950s and 1960s, individual Miocene taxa were commonly viewed as the direct ancestors of specific living ape species, suggesting an early divergence of the modern lineages.^1–5 However, in most cases, the Miocene forms were essentially “dental apes,” resembling extant species in dental and a few cranial features, but possessing more primitive postcranial features that suggested arboreal quadrupedalism rather than suspensory habits. With the introduction of molecular methods of phylogenetic reconstruction and the increasing use of cladistic analysis, it has become apparent that the radiation leading to the modern hominoids was somewhat more recent than had been believed, and that most of the Miocene hominoid species had little to do with the evolutionary history of the living apes. © 1998 Wiley-Liss, Inc.     

Fossil evidence of evolutionary convergence in juvenile dental morphology and upper canine replacement in sabertooth carnivores

The convergent suite of morphological traits characterizing the mammalian sabertooth ecomorphology is well documented, including modifications of the dental and osteological portions of the masticatory apparatus from a less‐specialized carnivore condition. Equally important is how those specialized adult morphologies developed through ontogeny because previous studies have shown that growing such specialized craniodental traits may require evolutionary modification of growth patterns and tooth replacement mechanisms. Despite the understanding of convergent morphological specialization in adult sabertooth carnivores, the possibility of a convergent ontogenetic trajectory toward those adult morphologies has not been rigorously examined. The present study examines numerous previously undescribed juvenile nimravid specimens. The results provide insights about nimravid ontogeny and show, for the first time, that the nimravid sabertooth lineage included species in which the permanent upper canine erupted within a lingual concavity of the deciduous upper canine until it reached comparable crown height beyond the alveolar border. Furthermore, this investigation assesses the juvenile morphology and upper canine replacement of felid and barbourofelid sabertooth taxa. The results provide evidence of convergence in deciduous upper canine morphology of three sabertooth carnivore lineages (i.e., nimravid, felid, and barbourofelid), as well as preliminary evidence of convergence in the upper canine replacement process. It might be beneficial for studies of extreme morphological specialization to simultaneously consider convergence in adult morphologies and how morphologies change through ontogeny.     

Extinct large colobines from the Plio-Pleistocene of Africa

Three genera and six species of extinct African Plio-Pleistocene Colobinae are discussed. One new genus, Rhinocolobus and three new species, R. turkanaensis, Cercopithecoides kimeui and Paracolobus mutiwa are named. These colobines show a diversity in postcranial and dental morphology not seen among extant species. Rhinocolobus was most similar to extant Colobus in postcranial morphology and had similar high-cusped shearing teeth. Cercopithecoides shows a number of postcranial skeletal features typical of terrestrial cercopithecid species and has lower cusped teeth. Paracolobus, while generally more similar to Rhinocolobus than to Cercopithecoides, is intermediate in some features of its postcranial morphology. The distribution of the various taxa among East and South African sites with different palaeoenvironments is generally consistent with the morphological interpretation. With the exception of Cercopithecoides kimeui, which persisted a little longer, these large colobines disappear from the fossil record about 1.8 million years ago. Their disappearance coincides with an interval of increasing aridity documented at Olduvai Gorge, the Omo Valley, and East Turkana.     

Cranial mechanics compared in extinct marsupial and extant African lions using a finite-element approach

Few species have generated more or longer running controversy than Australia's extinct marsupial lion Thylacoleo carnifex ( Owen, 1859 ). Over the last century and a half, feeding behaviours as disparate as osteophagy and specialized herbivory have been suggested and T. carnifex has been placed in both phalangeriform and vombatiform clades. Phylogenetic placement remains uncertain, but broad consensus has been achieved regarding diet, with all recent authors agreeing that T. carnifex was a carnivore. However, the marsupial lion's extraordinary cranial and dental morphologies remain without clear analogy, leaving many questions unanswered regarding how this most atypical mammalian predator killed its prey. Here I apply a rapidly emerging new approach in comparative biology, finite-element analysis, to the examination of cranial mechanics in T. carnifex . Comparisons are made with an extant lion Panthera leo (Linnaeus, 1758) under simulations designed to model stress distributions imposed by biting (intrinsic loads) and interaction with struggling prey (extrinsic loads). Modelling that approximates the 3-D architecture of jaw adductors suggests that both the placental and marsupial lions could generate considerably greater bite forces than has been predicted using 2-D approaches, but with relatively greater forces in the marsupial. The distribution of cranial stress is in many respects similar in both species, but results from simulations of extrinsic forces suggest that the marsupial was particularly well adapted to resist the high stresses that would be expected in dealings with relatively large prey. On the other hand, relatively high stress recorded in the rostrum of T. carnifex under intrinsic loadings suggests that it may have deployed a very different modus operandi , wherein the carnassial teeth played an active role in effecting a kill.     

The femur of extinct bunodont otters in Africa (Carnivora,Mustelidae, Lutrinae)

This study compares fossil femora attributed to extinct African bunodont lutrines with extant mustelids and ursids to reconstruct locomotor behavior. Due to the immense size differences among taxa, shape data were used to compare morphology. Based on morphological differences, the fossil femora are suggested to belong to different taxa with different locomotor abilities and habitat preferences. The Langebaanweg femur is the oldest and has a typical mustelid morphology suggesting that it was a locomotor generalist like most mustelids. The West Turkana form is more like extant nonbunodont otters, but much larger, and may have belonged to a semiaquatic taxon. The enormous Omo femur shares some features with truly aquatic taxa (e.g., Enhydra) and is the most likely to have been fully aquatic. The same may hold true for the Hadar species as it is most similar to that from the Omo. If these femora truly belong to bunodont lutrines, then they are more diverse in postcranial morphology than in dental morphology.     

Double infection experiments with echinostomatids (Trematoda) in Lymnaea stagnalis by implantation of rediae and exposure to miracidia

1972. Double infection experiments with echinostomatids (Trematoda) in Lymnaea stagnalis by implantation of rediae and exposure to miracidia. International Journal for Parasitology, 2: 409–423. Echinostomatid species parasitizing Lymnaea stagnalis as first intermediate hosts in a South German Lake have been found present in natural double infections, but at frequencies lower than expected. Simultaneous double infection and superinfection experiments in Lymnaea stagnalis with Isthmiophora melis, Echinoparyphium aconiatum and Echinostoma revolutum were performed by redial implantation and by exposure to miracidia. All three combinations possible of these echinostomatids proved to be unstable, one species being eliminated by another ‘stronger’ one after an invariable suppression order. The degree of vigour of Isthmiophora melis in this suppression order is greater if mother rediae (macropharyngeate) are present, i.e. after miracidial invasion instead of daughter redial implantation. Snails parasitized by rediae of a ‘weak’ type could be superinfected by implantation of rediae of a ‘strong’ type, but not if the first (‘weak’) infection had reached the stage of shedding cercariae. Superinfection by implantation of Echinoparyphium aconiatum rediae (‘strong’ type) was not successful when the first infection consisted of sporocysts of plagiorchiids, or of Apatemon sp. (Strigeidae) that had reached the stage of shedding cercariae.     

New partial skeletons of Palaeocene Nyctitheriidae and evaluation of proposed euarchontan affinities

Small-bodied, insectivorous Nyctitheriidae are known in the Palaeogene fossil record almost exclusively from teeth and fragmentary jaws and have been referred to Eulipotyphla (shrews, moles and hedgehogs) based on dental similarities. By contrast, isolated postcrania attributed to the group suggest arboreality and a relationship to Euarchonta (primates, treeshrews and colugos). Cretaceous–Palaeocene adapisoriculid insectivores have also been proposed as early euarchontans based on postcranial similarities. We describe the first known dentally associated nyctitheriid auditory regions and postcrania, and use them to test the proposed relationship to Euarchonta with cladistic analyses of 415 dental, cranial and postcranial characteristics scored for 92 fossil and extant mammalian taxa. Although nyctitheriid postcrania share similarities with euarchontans likely related to arboreality, results of cladistic analyses suggest that nyctitheriids are closely related to Eulipotyphla. Adapisoriculidae is found to be outside of crown Placentalia. These results suggest that similarities in postcranial morphology among nyctitheriids, adapisoriculids and euarchontans represent separate instances of convergence or primitive retention of climbing capabilities.     

江西萍乡上栗县晚更新世哺乳动物化石发现

2014~2015年在江西萍乡上栗县长平乡发现的杨家湾哺乳动物化石点,是江西省为数不多的更新世化石点中出土化石最为丰富多样的:目前已在杨家湾1号洞发现数千件哺乳动物牙齿化石,初步鉴定出40个属种;在化石数量上,野猪占绝对优势,其它常见种类还有鹿类、豪猪、黑熊、小型食肉类、鬣羚、水牛、犀牛及猴类等,而长鼻类及貘相对较少;在本地区同时代化石点中,猴类化石最丰富,食肉类属种也更多样。灭绝种类有巴氏大熊猫、最后斑鬣狗、德氏狸、剑齿象及巨貘等。从动物群组合来看,杨家湾1号洞与湖南道县福岩洞古人类化石点最为相似,其时代也应当属于晚更新世。