Feeding Biomechanics Research Articles

Tradeoffs between bite force and gape in Eulemur and Varecia.

In 1974, Sue Herring described the relationship between two important performance variables in the feeding system, bite force and gape. These variables are inversely related, such that, without specific muscular adaptations, most animals cannot produce high bite forces at large gapes for a given sized muscle. Despite the importance of these variables for feeding biomechanics and functional ecology, the paucity of in vivo bite force data in primates has led to bite forces largely being estimated through ex vivo methods. Here, we quantify and compare in vivo bite forces and gapes with output from simulated musculoskeletal models in two craniofacially distinct strepsirrhines: Eulemur, which has a shorter jaw and slower chewing cycle durations relative to jaw length and body mass compared to Varecia. Bite forces were collected across a range of linear gapes from 16 adult lemurs (suborder Strepsirrhini) at the Duke Lemur Center in Durham, North Carolina representing three species: Eulemur flavifrons (n = 6; 3F, 3M), Varecia variegata (n = 5; 3F, 2M), and Varecia rubra (n = 5; 5F). Maximum linear and angular gapes were significantly higher for Varecia compared to Eulemur (p = .01) but there were no significant differences in recorded maximum in vivo bite forces (p = .88). Simulated muscle models using architectural data for these taxa suggest this approach is an accurate method of estimating bite force-gape tradeoffs in addition to variables such as fiber length, fiber operating range, and gapes associated with maximum force. Our in vivo and modeling data suggest Varecia has reduced bite force capacities in favor of absolutely wider gapes compared to Eulemur in relation to their longer jaws. Importantly, our comparisons validate the simulated muscle approach for estimating bite force as a function of gape in extant and fossil primates.

A review of the role played by cilia in medusozoan feeding mechanics.

Cilia are widely present in metazoans and have various sensory and motor functions, including collection of particles through feeding currents in suspensivorous animals. Suspended particles occur at low densities and are too small to be captured individually, and therefore must be concentrated. Animals that feed on these particles have developed different mechanisms to encounter and capture their food. These mechanisms occur in three phases: (i) encounter; (ii) capture; and (iii) particle handling, which occurs by means of a cilia-generated current or the movement of capturing structures (e.g. tentacles) that transport the particle to the mouth. Cilia may be involved in any of these phases. Some cnidarians, as do other suspensivorous animals, utilise cilia in their feeding mechanisms. However, few studies have considered ciliary flow when examining the biomechanics of cnidarian feeding. Anthozoans (sessile cnidarians) are known to possess flow-promoting cilia, but these are absent in medusae. The traditional view is that jellyfish capture prey only by means of nematocysts (stinging structures) and mucus, and do not possess cilia that collect suspended particles. Herein, we first provide an overview of suspension feeding in invertebrates, and then critically analyse the presence, distribution, and function of cilia in the Cnidaria (mainly Medusozoa), with a focus on particle collection (suspension feeding). We analyse the different mechanisms of suspension feeding and sort them according to our proposed classification framework. We present a scheme for the phases of pelagic jellyfish suspension feeding based on this classification. There is evidence that cilia create currents but act only in phases 1 and 3 of suspension feeding in medusozoans. Research suggests that some scyphomedusae must exploit other nutritional sources besides prey captured by nematocysts and mucus, since the resources provided by this diet alone are insufficient to meet their energy requirements. Therefore, smaller particles and prey may be captured through other phase-2 mechanisms that could involve ciliary currents. We hypothesise that medusae, besides capturing prey by nematocysts (present in the tentacles and oral arms), also capture small particles with their cilia, therefore expanding their trophic niche and suggesting reinterpretation of the trophic role of medusoid cnidarians as exclusively plankton predators. We suggest further study of particle collection by ciliary action and its influence on the biomechanics of jellyfishes, to expand our understanding of the ecology of this group.

Evaluating modularity in the hominine skull related to feeding biomechanics.

Modular architecture of traits in complex organisms can be important for morphological evolution at micro- and sometimes macroevolutionary scales as it may influence the tempo and direction of changes to groups of traits that are essential for particular functions, including food acquisition and processing. We tested several distinct hypotheses about craniofacial modularity in the hominine skull in relation to feeding biomechanics. First, we formulated hypothesized functional modules for craniofacial traits reflecting specific demands of feeding biomechanics (e.g., masseter leverage/gape or tooth crown mechanics) in Homo sapiens, Pan troglodytes, and Gorilla gorilla. Then, the pattern and strength of modular signal was quantified by the covariance ratio coefficient and compared across groups using covariance ratio effect size. Hierarchical clustering analysis was then conducted to examine whether a priori-defined functional modules correspond to empirically recovered clusters. There was statistical support for most a priori-defined functional modules in the cranium and half of the functional modules in the mandible. Modularity signal was similar in the cranium and mandible, and across the three taxa. Despite a similar strength of modularity, the empirically recovered clusters do not map perfectly onto our priori functional modules, indicating that further work is needed to refine our hypothesized functional modules. The results suggest that modular structure of traits in association with feeding biomechanics were mostly shared with humans and the two African apes. Thus, conserved patterns of functional modularity may have facilitated evolutionary changes to the skull during human evolution.

Dynamic finite element modelling of the macaque mandible during a complete mastication gape cycle.

Three-dimensional finite element models (FEMs) are powerful tools for studying the mechanical behaviour of the feeding system. Using validated, static FEMs we have previously shown that in rhesus macaques the largest food-related differences in strain magnitudes during unilateral postcanine chewing extend from the lingual symphysis to the endocondylar ridge of the balancing-side ramus. However, static FEMs only model a single time point during the gape cycle and probably do not fully capture the mechanical behaviour of the jaw during mastication. Bone strain patterns and moments applied to the mandible are known to vary during the gape cycle owing to variation in the activation peaks of the jaw-elevator muscles, suggesting that dynamic models are superior to static ones in studying feeding biomechanics. To test this hypothesis, we built dynamic FEMs of a complete gape cycle using muscle force data from in vivo experiments to elucidate the impact of relative timing of muscle force on mandible biomechanics. Results show that loading and strain regimes vary across the chewing cycle in subtly different ways for different foods, something which was not apparent in static FEMs. These results indicate that dynamic three-dimensional FEMs are more informative than static three-dimensional FEMs in capturing the mechanical behaviour of the jaw during feeding by reflecting the asymmetry in jaw-adductor muscle activations during a gape cycle. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.

Functional morphological integration related to feeding biomechanics in the hominine skull

Quantifying and characterizing the pattern of trait covariances is crucial for understanding how population-level patterns of integration might constrain or facilitate craniofacial evolution related to the feeding system. This study addresses an important gap in our knowledge by investigating magnitudes and patterns of morphological integration of biomechanically informative traits in the skulls of Homo sapiens, Pan troglodytes, and Gorilla gorilla. We predicted a lower magnitude of integration among human biomechanical traits since humans eat a softer, less biomechanically challenging diet than apes. Indeed, compared to African apes, the magnitudes of integration were lower in H. sapiens skulls for form data (raw dimensions) but were similar or higher for shape data (raw dimensions scaled by geometric mean). Patterns of morphological integration were generally similar, but not identical, across the three species, particularly for the form data compared to the shape data. Traits that load heavily on the primary axis of variation in morphospace are generally associated with size and/or shape of the temporalis and masseter muscles and with dimensions related to the constrained lever model of jaw biomechanics. Given the conserved nature of morphological integration, skull adaptations for food processing in African apes and humans may have been constrained to occur along certain paths of high evolvability. The conserved pattern of functional integration also indicates that extant hominine species can operate as reasonable analogues for extinct hominins in studies that require population-level patterns of trait variance/covariance.

How do honeyeaters drink nectar?

We investigated the kinematics and biomechanics of nectar feeding in five species of honeyeater (Phylidonyris novaehollandiae, Acanthagenys rufogularis, Ptilotula penicillata, Certhionyx variegatus, Manorina flavigula). There is abundant information on honeyeater foraging behaviors and ecological relationships with plants, but there has never been an examination of their nectar-feeding from kinematic and biomechanical perspectives. We analyzed high-speed video of feeding in captive individuals to describe the kinematics of their nectar feeding, with specific focus on describing tongue movements and bill-tongue coordination, and to characterize the mechanism of nectar uptake in the tongue. We found clear interspecific variation in kinematics and tongue filling mechanics. Species varied in lick frequency, tongue velocity, and protrusion and retraction duration, which, in some cases, are relevant for differences in tongue filling mechanisms. We found support for the use of capillary filling in Certhionyx variegatus only. By contrast, Phylidonyris novaehollandiae, Acanthagenys rufogularis, Ptilotula penicillata, and Manorina flavigula employed a modified version of the expansive filling mechanism seen in hummingbirds, as there was dorsoventral expansion of the tongue body, even the portions that remain outside the nectar, once the tongue tip entered the nectar. All species use fluid trapping in the distal fimbriated portion of the tongue, which supports previous hypotheses describing the honeyeater tongue as a "paintbrush."

Nectar feeding beyond the tongue: hummingbirds drink using phase-shifted bill opening, flexible tongue flaps and wringing at the tips.

Hummingbirds are the most speciose group of vertebrate nectarivores and exhibit striking bill variation in association with their floral food sources. To explicitly link comparative feeding biomechanics to hummingbird ecology, deciphering how they move nectar from the tongue to the throat is as important as understanding how this liquid is collected. We employed synced, orthogonally positioned, high-speed cameras to describe the bill movements, and backlight filming to track tongue and nectar displacements intraorally. We reveal that the tongue base plays a central role in fluid handling, and that the bill is neither just a passive vehicle taking the tongue inside the flower nor a static tube for the nectar to flow into the throat. Instead, we show that the bill is actually a dynamic device with an unexpected pattern of opening and closing of its tip and base. We describe three complementary mechanisms: (1) distal wringing: the tongue is wrung out as soon as it is retracted and upon protrusion, near the bill tip where the intraoral capacity is decreased when the bill tips are closed; (2) tongue raking: the nectar filling the intraoral cavity is moved mouthwards by the tongue base, leveraging flexible flaps, upon retraction; (3) basal expansion: as more nectar is released into the oral cavity, the bill base is open (phase-shifted from the tip opening), increasing the intraoral capacity to facilitate nectar flow towards the throat.

How Triceratops got its face: An update on the functional evolution of the ceratopsian head.

Ceratopsian dinosaurs arguably show some of the most extravagant external cranial morphology across all Dinosauria. For over a century, ceratopsian dinosaurs have inspired a multitude of cranial functional studies as more discoveries continued to depict a larger picture of the enormous diversity of these animals. The iconic horns and bony frills in many ceratopsians portray a plethora of shapes, sizes, and arrangements across taxa, and their overall feeding apparatus show the development of unique specializations previously unseen in large herbivores. Here, I give a brief updated review of the many functional studies investigating different aspects of the ceratopsian head. The functional role of the horns and bony frills is explored, with an overview of studies investigating their potential for weaponization or defense in either intraspecific or anti-predatory combat, among other things. A review of studies pertaining to the ceratopsian feeding apparatus is also presented here, with analyses of studies exploring their beak and snout morphology, dentition and tooth wear, cranial musculature with associated skull anatomy, and feeding biomechanics.

Feeding biomechanics reveals niche differentiation related to insular gigantism.

Insular gigantism is an evolutionary phenomenon whereby small animals become bigger on islands compared to their mainland relatives. The abundance of insular giant taxa in the fossil record suggests the presence of a universal "giant niche" present on islands, with resource limitation as a potential driver for this process. However, insular habitats are ecologically diverse, suggesting that island taxa adopt different survival strategies, including adaptations for foraging behaviors. Here, we used finite element analysis to evaluate insular feeding niche adaptations in some of the most extreme examples of insular gigantism: Mediterranean giant dormice. We calculated stress, strain, and mechanical advantage during incisor and molar biting for 3 extinct insular giant species (Leithia melitensis, Hypnomys morpheus, and H. onicensis), an extant giant (Eliomys quercinus ophiusae), and their extant non-giant mainland relative, the generalist-feeder Eliomys quercinus. Our results show that dietary adaptations vary between giant taxa on different islands, and can occur relatively rapidly. Furthermore, the functional mandibular morphology in some insular taxa indicate adaptations moving away from a generalist feeding strategy toward greater trophic specialization. We show that the "insular giant niche" varies between islands and across time periods, arguing against a universal ecological driver for insular gigantism in small mammals.

Masticatory biomechanics of red and grey squirrels (Sciurus vulgarisandSciurus carolinensis) modelled with multibody dynamics analysis

The process of feeding in mammals is achieved by moving the mandible relative to the cranium to bring the teeth into and out of occlusion. This process is especially complex in rodents which have a highly specialized configuration of jaw adductor muscles. Here, we used the computational technique of multi-body dynamics analysis (MDA) to model feeding in the red (Sciurus vulgaris) and grey squirrel (Sciurus carolinensis) and determine the relative contribution of each jaw-closing muscle in the generation of bite forces. The MDA model simulated incisor biting at different gapes. A series of 'virtual ablation experiments' were performed at each gape, whereby the activation of each bilateral pair of muscles was set to zero. The maximum bite force was found to increase at wider gapes. As predicted, the superficial and anterior deep masseter were the largest contributors to bite force, but the temporalis had only a small contribution. Further analysis indicated that the temporalis may play a more important role in jaw stabilization than in the generation of bite force. This study demonstrated the ability of MDA to elucidate details of red and grey squirrel feeding biomechanics providing a complement to data gathered via in vivo experimentation.

Many-to-one function of cat-like mandibles highlights a continuum of sabre-tooth adaptations.

Cat-like carnivorans are a textbook example of convergent evolution, with distinct morphological differences between taxa with short or elongated upper canines, the latter often being interpreted as an adaptation to bite at large angles and subdue large prey. This interpretation of the sabre-tooth condition is reinforced by a reduced taxonomic sampling in some studies, often focusing on highly derived taxa or using simplified morphological models. Moreover, most biomechanical analyses focus on biting scenarios at small gapes, ideal for modern carnivora but ill-suited to test for subduction of large prey by sabre-toothed taxa. In this contribution, we present the largest three-dimensional collection-based muscle-induced biting simulations on cat-like carnivorans by running a total of 1074 analyses on 17 different taxa at three different biting angles (30°, 60° and 90°) including both morphologies. While our results show a clear adaptation of extreme sabre-toothed taxa to bite at larger angles in terms of stress distribution, other performance variables display surprising similarities between all forms at the different angles tested, highlighting a continuous rather than bipolar spectrum of hunting methods in cat-like carnivorans and demonstrating a wide functional disparity and nuances of the sabre-tooth condition that cannot simply be characterized by specialized feeding biomechanics.

Functional morphology of prey capture in stream-dwelling sailfin silversides (Telmatherinidae) based on high-speed video recordings

Understanding how ecology shapes the evolution of morphological traits is a major goal in organismal biology. By quantifying force of motion, hypotheses on the function of fundamental tasks of animals like feeding can be tested. Ray-finned fishes use various feeding strategies, classified into three main feeding modes: suction, ram and manipulation. While manipulation feeders are usually distinct in morphology and feeding behavior, differentiation between suction and ram feeders is often fine-scaled and transitional. Previous studies have identified different feeding modes and biomechanical adaptations on interspecific and intersexual levels in lake-dwelling sailfin silversides, species of a Sulawesi freshwater radiation. Functional feeding morphology of stream-dwelling species remained in contrast unstudied. We hypothesized that different requirements of riverine habitats favor the evolution of alternative functional adaptations in stream-dwelling sailfin silversides. To test this hypothesis, we investigated feeding of two phenotypically distinct riverine species, Telmatherina bonti and Marosatherina ladigesi, and their sexes, by high-speed videos and biomechanical models. The kinematic approaches identify T. bonti as ram feeder and M. ladigesi as suction feeder. Surprisingly, the biomechanical models of the jaw apparatus provide contradicting results: only one out of three studied parameters varies between both species. Contrarily to lake-dwelling Telmatherina, sexes of both species do not differ in feeding biomechanics. We conclude that T. bonti predominantly uses ram feeding while M. ladigesi primarily uses suction feeding as its main hunting strategy. Feeding biomechanics of stream-dwelling sailfin silversides are less distinct compared to lake-dwelling species, likely due to different trophic ecologies or less stable ecological conditions.

Biomechanics of nectar feeding explain flower orientation in plants pollinated by long-proboscid flies.

The function of flower orientation is much debated, with adaptation to pollinator mouthparts being a particularly compelling explanation, but also one that has lacked empirical support from broad-scale comparative studies. The two families of long-proboscid fly pollinators show similar hovering behaviour while feeding on nectar but differ in the biomechanics of their proboscides which can be up to 80mm in length: Tabanidae have a fixed forward-pointing proboscis while Nemestrinidae can swivel their proboscis downwards. We predicted that this difference has implications for the evolution of flower orientation. We established the flower angles of 156 South African plant species specialised for pollination by long-proboscid flies. Using a phylogenetically corrected analysis, we found that flowers pollinated by Tabanidae tend to be horizontally orientated, while those pollinated by Nemestrinidae tend to be more variable in orientation and more often vertically orientated. These results confirm the importance of pollinator biomechanics for the evolution of floral traits and highlight a potential mechanism of reproductive isolation between sympatric plant species pollinated by different long-proboscid fly families.

Morphometric and Genetic Description of Trophic Adaptations in Cichlid Fishes.

Simple SummarySkull and jaw shape are critical to how an animal eats. The goal of this work was to examine how facial variation evolves and the genetic basis of these changes. We used two species of Lake Malawi cichlids with different facial shapes, one which has evolved to eat prey by suction feeding, a second that bites algae from rocks, as well as hybrid individuals generated by artificial mating of the two species. We found a series of changes in craniofacial structure including the shape of the lower jaw and throat region that impact how animals perform at suction feeding and biting. We then identified genetic regions that regulate these facial shapes. These genetic regions suggested that evolution of the senses, among other traits, may play an important role in facial evolution. Also, evolution of different parts of the face are controlled by distinct genetic regions. Despite this, cichlids that eat similar ways have similar facial shapes, suggesting that the function of jaw movement places certain limits on facial evolution in cichlid fishes. Overall, this work provides insights into how the face evolves, how these changes relate to feeding, and the genes and molecules that regulate craniofacial variation.Since Darwin, biologists have sought to understand the evolution and origins of phenotypic adaptations. The skull is particularly diverse due to intense natural selection on feeding biomechanics. We investigated the genetic and molecular origins of trophic adaptation using Lake Malawi cichlids, which have undergone an exemplary evolutionary radiation. We analyzed morphological differences in the lateral and ventral head shape among an insectivore that eats by suction feeding, an obligate biting herbivore, and their F2 hybrids. We identified variation in a series of morphological traits—including mandible width, mandible length, and buccal length—that directly affect feeding kinematics and function. Using quantitative trait loci (QTL) mapping, we found that many genes of small effects influence these craniofacial adaptations. Intervals for some traits were enriched in genes related to potassium transport and sensory systems, the latter suggesting co-evolution of feeding structures and sensory adaptations for foraging. Despite these indications of co-evolution of structures, morphological traits did not show covariation. Furthermore, phenotypes largely mapped to distinct genetic intervals, suggesting that a common genetic basis does not generate coordinated changes in shape. Together, these suggest that craniofacial traits are mostly inherited as separate modules, which confers a high potential for the evolution of morphological diversity. Though these traits are not restricted by genetic pleiotropy, functional demands of feeding and sensory structures likely introduce constraints on variation. In all, we provide insights into the quantitative genetic basis of trophic adaptation, identify mechanisms that influence the direction of morphological evolution, and provide molecular inroads to craniofacial variation.

Inter‐relationships among body mass, body dimensions, jaw musculature and bite force in reptiles

AbstractBite force is important in understanding how feeding biomechanics impacts the life‐history of vertebrates. Bite force has been measured in many lizards and crocodilians but fewer turtles and snakes. Few comparative interspecific studies exist of the inter‐relationships between body size, jaw musculature and bite force. This study collated data reported in the literature for bite force and mass of the jaw musculature in reptiles and explored the relationships between these variables and body mass and size. Two hypotheses were explored: (1) bite force and mass of the jaw musculature scale with body size irrespective of order (as a broad taxonomic grouping) when controlled for phylogeny, and (2) bite force and jaw musculature mass would be directly related and be unrelated to order. Phylogenetically controlled Bayesian analyses showed that in relation to body mass there were different relationships with bite force for testudines compared to saurians and crocodilians. For snout–vent length, squamates and crocodilians exhibited a single relationship, but for head length, all reptile orders exhibited different allometric patterns. Although there was a strong phylogenetic signal in all cases, indicating that there are clusters of closely related species for a measure of size, there was no effect of order on the relationships between measures of body size and mass of the jaw musculature. The relationship between bite force and jaw muscle mass was isometric and did not differ among reptile orders. Several ecological factors may correlate with head morphology and bite force in reptiles. Further studies could explore the evolutionary drivers that shape the relationship between head size, jaw musculature and bite force. Similarities for the relationships between bite force and body size in different reptilian orders suggests that this could be a model system to demonstrate fundamental biomechanical principles of how jaw muscles generate a bite force.

Royal knifefish generate powerful suction feeding through large neurocranial elevation and high epaxial muscle power.

Suction feeding in ray-finned fishes involves powerful buccal cavity expansion to accelerate water and food into the mouth. Previous XROMM studies in largemouth bass (Micropterus salmoides), bluegill sunfish (Lepomis macrochirus) and channel catfish (Ictalurus punctatus) have shown that more than 90% of suction power in high performance strikes comes from the axial musculature. Thus, the shape of the axial muscles and skeleton may affect suction feeding mechanics. Royal knifefish (Chitala blanci) have an unusual postcranial morphology, with a ventrally flexed vertebral column and relatively large mass of epaxial muscle. Based on their body shape, we hypothesized that royal knifefish would generate high power strikes by utilizing large neurocranial elevation, vertebral column extension and epaxial shortening. As predicted, C. blanci generated high suction expansion power compared with the other three species studied to date (up to 160 W), which was achieved by increasing both the rate of volume change and the intraoral subambient pressure. The large epaxial muscle (25% of body mass) shortened at high velocities to produce large neurocranial elevation and vertebral extension (up to 41deg, combined), as well as high muscle mass-specific power (up to 800 W kg-1). For the highest power strikes, axial muscles generated 95% of the power, and 64% of the axial muscle mass consisted of the epaxial muscles. The epaxial-dominated suction expansion of royal knifefish supports our hypothesis that postcranial morphology may be a strong predictor of suction feeding biomechanics.

Bat Dentitions: A Model System for Studies at The Interface of Development, Biomechanics, and Evolution.

The evolution of complex dentitions was a major innovation in mammals that facilitated the expansion into new dietary niches that imposed selection for tight form-function relationships. Teeth allow mammals to ingest and process food items by applying forces produced by a third-class lever system composed by the jaw adductors, the cranium, and the mandible. Physical laws determine changes in jaw adductor (biting) forces at different bite point locations along the mandible (outlever), thus individual teeth are expected to experience different mechanical regimes during feeding. If the mammal dentition exhibits functional adaptations to mandible feeding biomechanics, then teeth are expected to have evolved to develop mechanically-advantageous sizes, shapes, and positions. Here, we present bats as a model system to test this hypothesis and, more generally, for integrative studies of mammal dental diversity. We combine a field-collected dataset of bite forces along the tooth row with data on dental and mandible morphology across 30 bat species. We (1) describe, for the first time, bite force trends along the tooth row of bats, (2) use phylogenetic comparative methods to investigate relationships among bite force patterns, tooth and mandible morphology, and (3) hypothesize how these biting mechanics patterns may relate to the developmental processes controlling tooth formation. We find that bite force variation along the tooth row is consistent with predictions from lever mechanics models, with most species having the greatest bite force at the first lower molar. The cross-sectional shape of the mandible body is strongly associated with the position of maximum bite force along the tooth row, likely reflecting mandibular adaptations to varying stress patterns among species. Further, dental dietary adaptations seem to be related to bite force variation along molariform teeth, with insectivorous species exhibiting greater bite force more anteriorly, narrower teeth and mandibles, and frugivores/omnivores showing greater bite force more posteriorly, wider teeth and mandibles. As these craniodental traits are linked through development, dietary specialization appears to have shaped intrinsic mechanisms controlling traits relevant to feeding performance.

Dental pathologies in lamniform and carcharhiniform sharks with comments on the classification and homology of double tooth pathologies in vertebrates.

Double tooth pathologies are important indicators of trauma, disease, diet, and feeding biomechanics, and are widely documented in mammals. However, diagnosis of double tooth pathologies in extinct non-mammalian vertebrates is complicated by several compounding factors including: a lack of shared terminology reflecting shared etiology, inconsistencies in definitions and key features within and outside of mammals (e.g., gemination, fusion, twinning, concrescence); differences in tooth morphology, heterodonty, regeneration, and implantation between mammals and non-mammalian vertebrates; and the unmet need for diagnostic criteria that can be applied to isolated teeth, which are common in the fossil record. Here we report on double tooth pathologies in the lamniform and carcharhiniform Cenozoic sharks Otodus megalodon (NCSM 33639) and Carcharhinus leucas (NCSM 33640, 33641). All three teeth bear a singular bifid crown with mirrored halves and abnormal internal microstructure—a single, bifurcating pulp cavity in C. leucas and a more than tripling of vessels in O. megalodon (from two to seven main ascending canals). We identify these abnormalities as likely examples of gemination due to their symmetry, which rules out fusion of tooth buds in one tooth file in different developmental stages in polyphyodont taxa; however, we note that incomplete forms of mesiodistal tooth fusion can be morphologically indistinguishable from gemination, and thus fusion cannot be rejected. We further compile and recategorize, when possible, the diversity of tooth pathologies in sharks. The identification of double tooth pathologies in O. megalodon and C. leucas has paleobiological implications. Such pathologies in sharks are largely hypothesized to stem from trauma to developing tooth buds. Carcharhinus leucas is known to feed on prey documented to cause feeding-related oral traumas (e.g., rays, sawfish, spiny fish, and sea urchins). However, O. megalodon, is considered to have largely fed on marine mammals, and perhaps turtles and/or fish, raising the possibility that the dietary diversity of this species is, as of yet, underappreciated. The genetic underpinnings of tooth morphogenesis and regeneration is highly conserved throughout vertebrate evolution, suggesting a homologous framework can be established. However, more research is needed to link developmental, paleobiological, and/or paleoenvironmental factors to gemination/fusion in polyphyodont taxa. We argue that the definitions and diagnostic criteria for dental pathologies in vertebrates require standardization in order to advance macroevolutionary studies of feeding trauma in deep time.

Inter‐relationships among body mass, jaw musculature and bite force in birds

AbstractBite force can provide an insight into the feeding biomechanics and ecology of vertebrates. There has been an increasing interest in bite force in birds with data being collected using force transducers although bite forces have also been calculated from the mass of the jaw musculature. Studies have sought to find ecological correlates between bite force and diet or feeding style as well as with morphology. This study collated data reported in the literature for bite force and mass of the jaw musculature in birds and explored the relationships between these variables and their relationships with body mass to test two hypotheses. First, bite force and mass of the jaw musculature would scale with body mass irrespective of the phylogeny. Second, bite force and mass of the jaw musculature would be directly related to each other and be unrelated to phylogeny. Phylogenetically controlled analyses showed that in relation to body mass there were different relationships with bite force, and with the mass of the jaw musculature, for non‐passerine (isometry and negative allometry, respectively) and passerine species (both positive allometry). By contrast, a single significantly positively allometric relationship described the relationship between jaw muscle mass and bite force. These relationships were driven in part by the diet of the species concerned but also may reflect morphological differences in jaw musculature. The few studies that compare measured and calculated bite force for the same species show that values derived from muscles were higher. Detaching muscles from their points of origin and insertion to calculate physiological cross‐sectional area may be biasing bite force calculations.

Mandible shape variation and feeding biomechanics in minks

European and American minks are very similar in ecology, behavior and morphology. Both species hunt terrestrial vertebrates and aquatic prey, but the American mink is a more generalist predator which, among other factors, allows it to outcompete the European mink in areas where it has been introduced. We used 3D geometric morphometrics and estimates of muscle mechanical advantage to assess the degree of variation in mandibular morphology, and to determine whether such variation reflects dietary differences between the two species. The three main axes of variation represented interspecific differences, a common allometric trajectory between species and sexes, and the interspecific effect of sexual size dimorphism, with males having overall stronger bites than females. Differences in mandible shape and biomechanical parameters suggest that American minks are better equipped for preying on terrestrial vertebrates, while the features seen in European mink could be related to tougher prey, fish capture, or both. Additionally, within each species, the larger specimens of each sex present indicators of a higher percentage of terrestrial prey in their diet. These results indicate a low potential dietary overlap between both species, suggesting that factors other than prey competition may have a role in the decline of the European mink.