Jaw Adductor Research Articles

African wild dog (Lycaon pictus) masticatory apparatus and oral cavity morphology.

African wild dogs (Lycaon pictus) are unique among canids in their specialized hunting strategies and social organization. Unlike other, more omnivorous canids, L. pictus is a hypercarnivore that consumes almost exclusively meat, particularly prey larger than its body size, which it hunts through cooperative, exhaustive predation tactics. Its bite force is also among the highest reported for carnivorans. Here, we dissected an adult male L. pictus specimen and conducted diffusion iodine contrast-enhanced computed tomography (diceCT) scans to evaluate and describe its masticatory and oral cavity musculature. Muscles of mastication in L. pictus are separated by deep layers of thick intermuscular fascia and deep insertions. The superficial surface of m. masseter is entirely covered by an extremely thick masseteric fascia. Deep to m. masseter pars reflexa and superficialis are additional bellies, m. masseter pars profunda and zygomaticomandibularis. Musculus temporalis in L. pictus, divides into suprazygomatic, superficial, and deep bellies separated by a deep layer of thick intermuscular fascia, and it inserts along the entire rostral margin of the mandibular ramus. Musculus digastricus appears to comprise a single, large fusiform belly which appears to receive its innervation exclusively from CN V3 (nervus mandibularis, division of nervus trigeminus). Musculus pterygoideus medialis and lateralis are each composed of a single, deep belly. However, despite its great bite force, the jaw adductor muscle mass in L. pictus is not increased for its body size over other canid taxa. This finding suggests there are other architectural adaptations to hypercarnivory beyond increased muscle volume (e.g., pennation angle, greater strength, optimization of lever arms for mechanical advantage).

Bite hard: Linking cranial loading mechanics to ecological differences in gnawing behavior in caviomorph rodents.

The mammalian skull is very malleable and has notably radiated into highly diverse morphologies, fulfilling a broad range of functional needs. Although gnawing is relatively common in mammals, this behavior and its associated morphology are diagnostic features for rodents. These animals possess a very versatile and highly mechanically advantageous masticatory apparatus, which, for instance, allowed caviomorph rodents to colonize South America during the Mid-Eocene and successfully radiate in over 200 extant species throughout most continental niches. Previous work has shown that differences in bite force within caviomorphs could be better explained by changes in muscle development than in mechanical advantages (i.e., in cranial overall morphology). Considering the strong bites they apply, it is interesting to assess how the reaction forces upon the incisors (compression) and the powerful adductor musculature pulling (tension) mechanically affect the cranium, especially between species with different ecologies (e.g., chisel-tooth digging). Thus, we ran finite element analyses upon crania of the subterranean Talas' tuco-tuco Ctenomys talarum, the semi-fossorial common degu Octodon degus, and the saxicolous long-tailed chinchilla Chinchilla lanigera to simulate: (A) invivo biting in all species, and (B) rescaled muscle forces in non-ctenomyid rodents to match those of the tuco-tuco. Results show that the stress patterns correlate with the mechanical demands of distinctive ecologies, on invivo-based simulations, with the subterranean tuco-tuco being the most stressed species. In contrast, when standardizing all three species (rescaled models), non-ctenomyid models exhibited a several-fold increase in stress, in both magnitude and affected areas. Detailed observations evidenced that this increase in stress was higher in lateral sections of the snout and, mainly, the zygomatic arch; between approximately 2.5-3.5 times in the common degu and 4.0-5.0 times in the long-tailed chinchilla. Yet, neither species, module, nor simulation condition presented load factor levels that would imply structural failure by strong, incidental biting. Our results let us conclude that caviomorphs have a high baseline for mechanical strength of the cranium because of the inheritance of a very robust "rodent" model, while interspecific differences are associated with particular masticatory habits and the concomitant level of development of the adductor musculature. Especially, the masseteric and zygomaticomandibular muscles contribute to >80% of the bite force, and therefore, their contraction is responsible for the highest strains upon their origin sites, that is, the zygomatic arch and the snout. Thus, the robust crania of the subterranean and highly aggressive tuco-tucos allow them to withstand much stronger forces than degus or chinchillas, such as the ones produced by their hypertrophied jaw adductor muscles or imparted by the soil reaction.

Skull morphology of the enigmatic Genyornis newtoni Stirling and Zeitz, 1896 (Aves, Dromornithidae), with implications for functional morphology, ecology, and evolution in the context of Galloanserae

ABSTRACT The presence of Dromornithidae in the Australian Cenozoic fossil record was first reported in 1872, yet although eight species and hundreds of specimens are known, key information on their morphology remains elusive. This is especially so for their skulls, which contributes to a lack of resolution regarding their relationships within Galloanserae. The skull of the Pleistocene dromornithid, Genyornis newtoni, was initially described in 1913. Additional fossils of this species have since been discovered and understanding of avian skull osteology, arthrology, and myological correlates has greatly advanced. Here we present a complete redescription of the skull of Genyornis newtoni, updating knowledge on its morphology, soft-tissue correlates, and palaeobiology. We explore the diversity within Dromornithidae and make comprehensive comparisons to fossil and extant galloanserans. Furthermore, we expand on the homologies of skull muscles, especially regarding the jaw adductors and address the conflicting and unstable placement of dromornithids within Galloanserae. Findings support generic distinction of Genyornis newtoni, and do not support the close association of Dromornithidae and Gastornithidae. We thus recommend removal of the dromornithids from the Gastornithiformes. Considering character polarities, the results of our phylogenetic analyses, and palaeogeography, our findings instead support the alternative hypotheses, of dromornithids within, or close to, the Suborder Anhimae with Anseriformes.

Analysis of ontogenetic changes in head shape and diet in a catfish with moderately enlarged jaw adductors (Clariallabes melas)

Jaw adductor hypertrophy, or the presence of enlarged jaw-closing muscles, has arisen several times independently in African clariid catfish. Previous work has demonstrated that species characterized by enlarged jaw adductors may have unusual ways of foraging such as terrestrial foraging and prey capture, and often include a large proportion of hard and terrestrial prey in their diet. However, relatively little is known about species of the genus Clariallabes with an intermediate degree of jaw adductor hypertrophy. In the present study we present data on head shape and diet for a range of sizes of specimens of a poorly known species of Clariallabes, C. melas. Our data show that growth patterns in this species and the previously studied 
 C. longicauda are similar in some ways (e.g. positive allometry of the growth of the jaw muscles) but different in others (negative allometry in hyoid width). However, C. melas has a smaller head for its body size in all dimensions. Due to the large number of empty stomachs we encountered, dietary data remain preliminary, but suggest a varied diet including both hard and soft prey. Our data show that a large amount of variation in head shape may exist even among related species and that species with jaw adductor hypertrophy generally show positive allometry in the growth of the jaw adductors and associated structures. Whether jaw muscle hypertrophy is an adaptive trait in clariid catfish awaits further comparative analyses testing for the evolutionary association between jaw adductor hypertrophy and the inclusion of hard prey into the diet.

Cranial muscle architecture in wild boar: Does captivity drive ontogenetic trajectories?

The jaw system in mammals is complex and different muscle morphotypes have been documented. Pigs are an interesting group of animals as they are omnivorous and have a bunodont crushing dentition. Moreover, they have interacted with humans for over 10,000 years and grow nearly two orders of magnitude in size. Despite being a model system for studies on cranial form and function, data on the growth of the jaw adductor muscles are scant. Moreover, whether captivity impacts the growth and architecture of the jaw adductors remains unknown. Based on dissection data of the jaw adductors of 45 animals ranging from less than 1 kg to almost 100 kg, we show that muscle masses, muscle fiber lengths, and cross-sectional areas scale as predicted for geometrically similar systems or with slight negative allometry. Only the fiber length of the lateral pterygoid muscle grew with slight positive allometry. Animals raised in captivity in stalls or in an enclosure were overall very similar to wild animals. However, some muscles were larger in captive animals. Interestingly, variation in bite force in captive animals was well predicted by the variation in the size of the superficial masseter muscle relative to the overall jaw adductor mass.

Symphyseal morphology and jaw muscle recruitment levels during mastication in musteloid carnivorans.

In studies of mammalian mastication, a possible relationship has been proposed between bilateral recruitment of jaw adductor muscle force during unilateral chewing and the degree of fusion of the mandibular symphysis. Specifically, species that have unfused, mobile mandibular symphyses tend to utilize lower levels of jaw adductor force on the balancing (nonchewing) than the working (chewing) side of the head, when compared to related species with fused symphyses. Here, we compare jaw adductor recruitment levels in two species of musteloid carnivoran: the carnivorous ferret (unfused symphysis), and the frugivorous kinkajou (fused symphysis). During forceful chewing, we observe that ferrets recruit far more working-side muscle force than kinkajous, regardless of food toughness and that high working-to-balancing side ratios are the result of increased working-side force, often coupled with reduced balancing-side force. We propose that in carnivorans, high working-to-balancing side force ratios coupled with an unfused mandibular symphysis are necessary to rotate the hemimandible for precise unilateral occlusion of the carnassial teeth and to sustain laterally oriented force on the jaw to engage the carnassial teeth during shearing of tough foods. In contrast, the kinkajou's flattened cheekteeth permit less precise occlusion and require medially-oriented forces for grinding, thus, a fused symphysis is mechanically beneficial.

Ecological correlates of three-dimensional muscle architecture within the dietarily diverse Strepsirrhini.

Analysis of muscle architecture, traditionally conducted via gross dissection, has been used to evaluate adaptive relationships between anatomical form and behavioral function. However, gross dissection cannot preserve three-dimensional relationships between myological structures for analysis. To analyze such data, we employ diffusible, iodine-based contrast-enhanced computed tomography (DiceCT) to explore the relationships between feeding ecology and masticatory muscle microanatomy in eight dietarily diverse strepsirrhines: allowing, for the first time, preservation of three-dimensional fascicle orientation and tortuosity across a functional comparative sample. We find that fascicle properties derived from these digital analyses generally agree with those measured from gross-dissected conspecifics. Physiological cross-sectional area was greatest in species with mechanically challenging diets. Frugivorous taxa and the wood-gouging species all exhibit long jaw adductor fascicles, while more folivorous species show the shortest relative jaw adductor fascicle lengths. Fascicle orientation in the parasagittal plane also seems to have a clear dietary association: most folivorous taxa have masseter and temporalis muscle vectors that intersect acutely while these vectors intersect obliquely in more frugivorous species. Finally, we observed notably greater magnitudes of fascicle tortuosity, as well as greater interspecific variation in tortuosity, within the jaw adductor musculature than in the jaw abductors. While the use of a single specimen per species precludes analysis of intraspecific variation, our data highlight the diversity of microanatomical variation that exists within the strepsirrhine feeding system and suggest that muscle architectural configurations are evolutionarily labile in response to dietary ecology-an observation to be explored across larger samples in the future.

The dual function of prokinesis in the feeding and locomotor systems of parrots.

Prokinesis, a mode of avian cranial kinesis involving motion between the neurocranium and upper beak, has long been investigated in biomechanical analyses of avian feeding and drinking. However, the modern avian beak is also used for non-feeding functions. Here, we investigate the dual function of prokinesis in the feeding and locomotor systems of the rosy-faced lovebird (Agapornis roseicollis). Lovebirds and other parrots utilize their beak both during feeding and as a third limb during vertical climbing. Thus, we experimentally measured both force-generating potential and movement of the rosy-faced lovebird mandible and maxilla (via prokinetic flexion of the craniofacial hinge) during tripedal climbing and mandibular/maxillary adduction. We found that whereas the maxilla is primarily responsible for generating force during locomotion, the mandible is primarily responsible for generating force during forceful jaw adduction, hinting at a remarkable capacity to alter prokinetic function with differing neuromuscular control. The ability of the prokinetic apparatus to perform functions with competing optimality criteria via modulation of motor control illustrates the functional plasticity of the avian cranial kinesis and sheds new light on the adaptive significance of cranial mobility.

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'.

Energetic costs of feeding in 12 species of small-bodied primates.

There are no comparative, empirical studies of the energetic costs of feeding in mammals. As a result, we lack physiological data to better understand the selection pressures on the mammalian feeding apparatus and the influence of variables such as food geometric and material properties. This study investigates interspecific scaling of the net energetic costs of feeding in relation to body size, jaw-adductor muscle mass and food properties in a sample of 12 non-human primate species ranging in size from 0.08 to 4.2 kg. Net energetic costs during feeding were measured by indirect calorimetry for a variety of pre-cut and whole raw foods varying in geometric and material properties. Net feeding costs were determined in two ways: by subtracting either the initial metabolic rate prior to feeding or subtracting the postprandial metabolic rate. Interspecific scaling relationships were evaluated using pGLS and OLS regression. Net feeding costs scale negatively relative to both body mass and jaw-adductor mass. Large animals incur relatively lower feeding costs indicating that small and large animals experience and solve mechanical challenges in relation to energetics in different ways. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.

Shape analysis of the preorbital bar in caviomorph rodents.

The highly specialised masticatory apparatus of rodents raises interesting questions about how their skull withstands the intensive and sustained forces produced by biting on hard items. In these mammals, major systematics were explored for a long time based on the adductor muscles' architecture and the related bony structures. The infraorbital foramen stands out, where a hypertrophied head of the zygomaticomandibular muscle passes through-in hystricomorphous rodents-as a direct consequence of the lateral and posterior shift of the preorbital bar. Interestingly, this bar moved laterally and backwards-enlarging the foramen-but it never disappeared throughout evolution, even showing morphological convergence among rodents. Previous research proposed this bar as behaving mechanically similar to the postorbital bar in ungulates, i.e., a safety structure against torsion stress while chewing. We analysed its morphology by mathematically modelling it under bending and torsion scenarios (linearly and elliptically shaped, respectively), and as for biting load propagation (catenary curve). Although the preorbital bar primarily seems to be shaped for withstanding torsional stress (as the postorbital bar in ungulates) and as an escaping point for force propagation, these forces are not a consequence of chewing and grinding foods, but preventing the zygomatic arch from failing when the powerful laterally-displaced jaw adductor muscles are pulling the dentary upwards at biting.

Divergent mechanisms for crushing shelled prey in parallel lineages of durophagous stingrays (Myliobatoidei)

Durophagous vertebrates feed on shelled prey and share many morphological traits; in particular, high leverage jaws with robust jaw-closing muscles. Myliobatid stingrays are no exception, having reinforced jaw skeletons, large jaw adductor muscles, and with all species in the family consuming some combination of shelled prey. Myliobatid rays have a long evolutionary history (65–70 million years) and number over thirty species, yet despite similar performance demands given their diet ecology, exhibit considerable morphological diversity of the feeding apparatus. We examined how feeding performance changes in the bullnose ray, Myliobatis freminvillei over its ontogeny, and compared how these rays produce crushing bite forces relative to confamilial and co-occurring cownose rays (Rhinopterinae). Within the durophagous family Myliobatidae, rhinopterines are more generalist in their dietary preferences, feeding on small bivalves and infaunal crustaceans, whereas eagle rays (Aetobatinae) prey on gastropods and bivalves exclusively, while bullnose/bat rays (Myliobatinae) consume a size range of gastropods and hermit crabs. We found that bullnose rays show isometric changes in feeding performance over their ontogeny, relying on the high mechanical efficiency of their jaws to crush shelled prey throughout their ontogeny. This manner of generating high bite forces diverges from cownose rays and other durophagous elasmobranchs, which primarily rely on muscular hypertrophy to generate forceful bites. These two lineages evolved distinct means of dismantling hard prey, which further showcases how multiplicity of form can result in similar ecological function. These predators are frequently subsumed within a single ecological category, durophagy, which underestimates their diversity.

Updating the fossil record of the alligatoroid crocodylian Diplocynodon from the late Eocene of Transylvanian Basin

The basal alligatoroid crocodylian Diplocynodon kochi erected for incomplete three-dimensional skull from the late Eocene (Priabonian) Cluj Limestone Formation in Romania, represents one of the easternmost distributed European members of Diplocynodontidae. New isolated cranial and postcranial remains provide new insights into the diagnostic features, phylogenetic relationships and lifestyle of this taxon, extending its fossil record to four new localities situated on the north-western side of the Transylvanian Basin. Diagnostic traits of the holotype include an extended insertion surface of jaw adductors on the parietal and squamosal, whereas the newly referred mandibles possess an enlarged and procumbent first dentary tooth, and the posterior teeth and alveoli are mediolaterally compressed. These attributes might have been related to the prey capture approach of D. kochi, involved undoubtedly in the food chains of both continental (fluvial or marshy-lacustrine) and shallow marine environments, as indicated by the taphonomic settings of the surveyed fossil bearing deposits. The warm and humid climate is indicated for the late Eocene (Priabonian) of the Transylvanian Basin based on palynomorphs, however, the presence of “growth rings” and “lines of arrested growth” on the prezygapophyses of a dorsal vertebra referred to D. kochi, points to a seasonal climate, installed well before the Eocene–Oligocene terminal event. Positioned between western and eastern European faunas, the diplocynodontid populations from the Transylvanian Basin survived probably the Eocene–Oligocene transition, as suggested by the early Oligocene (Rupelian) fossil record from the area.

Differential growth of the adductor muscles, eyeball, and brain in the chick Gallus gallus with comments on the fossil record of stem-group birds.

The avian head is unique among living reptiles in its combination of relatively large brain and eyes, coupled with relatively small adductor jaw muscles. These derived proportions lend themselves to a trade-off hypothesis, wherein adductor size was reduced over evolutionary time as a means (or as a consequence) of neurosensory expansion. In this study, we examine this evolutionary hypothesis through the lens of development by describing the jaw-adductor anatomy of developing chickens, Gallus gallus, and comparing the volumetric expansion of these developing muscles with growth trajectories of the brain and eye. Under the trade-off hypothesis, we predicted that the jaw muscles would grow with negative allometry relative to brain and eyes, and that osteological signatures of a relatively large adductor system, as found in most nonavian dinosaurs, would be differentially expressed in younger chicks. Results did not meet these expectations, at least not generally, with muscle growth exhibiting positive allometry relative to that of brain and eye. We propose three, nonmutually exclusive explanations: (1) these systems do not compete for space, (2) these systems competed for space in the evolutionary past, and growth of the jaw muscles was truncated early in development (paedomorphosis), and (3) trade-offs in developmental investment in these systems are limited temporally to the perinatal period. These explanations are considered in light of the fossil record, and most notably the skull of the stem bird Ichthyornis, which exhibits an interesting combination of plesiomorphically large adductor chamber and apomorphically large brain.

Variation in the Jaw Musculature of Ratsnakes and Their Allies (Serpentes: Colubridae)

Snakes have a highly modified feeding apparatus. However, its associated musculature is often poorly known. In order to study variation in the cephalic musculature, we dissected specimens representing 28 snake species belonging to the New World clade Lampropeltini and their Old World relatives. The observed variation was analysed using a phylogenetic framework. We found that the pattern of their musculature is conservative. We observed no interspecific variation in the intermandibular muscles or in the posterior jaw adductors. Variation within the dorsal constrictors and lateral jaw adductors is relatively low. This could be explained by morphological (space) limitations and functional constraints.

Ontogeny of cranial musculoskeletal anatomy and its relationship to allometric increase in bite force in an insectivorous bat (Eptesicus fuscus).

Bite force is a performance metric commonly used to link cranial morphology with dietary ecology, as the strength of forces produced by the feeding apparatus largely constrains the foods an individual can consume. At a macroevolutionary scale, there is evidence that evolutionary changes in the anatomical elements involved in producing bite force have contributed to dietary diversification in mammals. Much less is known about how these elements change over postnatal ontogeny. Mammalian diets drastically shift over ontogeny-from drinking mother's milk to feeding on adult foods-presumably with equally drastic changes in the morphology of the feeding apparatus and bite performance. Here, we investigate ontogenetic morphological changes in the insectivorous big brown bat (Eptesicus fuscus), which has an extreme, positive allometric increase in bite force during development. Using contrast-enhanced micro-computed tomography scans of a developmental series from birth to adult morphology, we quantified skull shape and measured skeletal and muscular parameters directly related to bite force production. We found pronounced changes in the skull over ontogeny, including a large increase in the volume of the temporalis and masseter muscles, and an expansion of the skull dome and sagittal crest that would serve to increase the temporalis attachment area. These changes indicate that development of the jaw adductors play an important role in the development of biting performance of these bats. Notably, static bite force increases with positive allometry with respect to all anatomical measures examined, suggesting that modifications in biting dynamics and/or improved motor coordination also contribute to increases in biting performance.

Sustained force production by the jaw-adductor muscles of a megalophagous frog, Ceratophrys cranwelli.

Most frogs have weak jaws that play a relatively minor role in tongue-mediated prey capture. Horned frogs (Ceratophrys spp.), however, follow the projection of a large tongue with a vice-like grip of their jaws to hold and immobilize prey. Prey include relatively large vertebrates, which they may restrain for minutes to possibly hours. High endurance behaviors, such as prolonged biting, require that muscles be capable of sustained force production. The feeding behavior of Ceratophrys suggests that their jaw-adductor muscles may be capable of powering sustained bites for long periods. We examined the capacity for sustained bite force by conducting an in situ experiment during which we measured bite force while bilaterally and supramaximally stimulating the jaw-adductor muscles of euthanized Cranwell's horned frogs (C. cranwelli). Muscles were stimulated for at least 60 min with a series of tetanic trains, with one experiment lasting over 6 h. We found that a significant sustained force develops during the first few minutes of the experiment, and this force is present between tetanic trains when the muscles are not being stimulated. The sustained force persists long after tetanic forces are barely detectable. The observed sustained force phenomenon parallels that observed for the jaw-adductor muscles of alligator lizards (Elgaria), another animal capable of sustained biting. The ability to bite with sustained and significant force by C. cranwelli may be facilitated by a configuration of different muscle fiber types, such as slow tonic fibers, as well as specializations in the muscle fibers that mitigate the effects of fatigue.

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.

Multiple pathways to herbivory underpinned deep divergences in ornithischian evolution

The extent to which evolution is deterministic is a key question in biology,1,2,3,4,5,6,7,8,9 with intensive debate on how adaptation6,10,11,12,13 and constraints14,15,16 might canalize solutions to ecological challenges.4,5,6 Alternatively, unique adaptations1,9,17 and phylogenetic contingency1,3,18 may render evolution fundamentally unpredictable.3 Information from the fossil record is critical to this debate,1,2,11 but performance data for extinct taxa are limited.7 This knowledge gap is significant, as general morphology may be a poor predictor of biomechanical performance.17,19,20 High-fiber herbivory originated multiple times within ornithischian dinosaurs,21 making them an ideal clade for investigating evolutionary responses to similar ecological pressures.22 However, previous biomechanical modeling studies on ornithischian crania17,23,24,25 have not compared early-diverging taxa spanning independent acquisitions of herbivory. Here, we perform finite-element analysis on the skull of five early-diverging members of the major ornithischian clades to characterize morphofunctional pathways to herbivory. Results reveal limited functional convergence among ornithischian clades, with each instead achieving comparable performance, in terms of reconstructed patterns and magnitudes of functionally induced stress, through different adaptations of the feeding apparatus. Thyreophorans compensated for plesiomorphic low performance through increased absolute size, heterodontosaurids expanded jaw adductor muscle volume, ornithopods increased jaw system efficiency, and ceratopsians combined these approaches. These distinct solutions to the challenges of herbivory within Ornithischia underpinned the success of this diverse clade. Furthermore, the resolution of multiple solutions to equivalent problems within a single clade through macroevolutionary time demonstrates that phenotypic evolution is not necessarily predictable, instead arising from the interplay of adaptation, innovation, contingency, and constraints.1,2,3,7,8,9,18.

Lip service: Histological phenotypes correlate with diet and feeding ecology in herbivorous pacus.

Complex prey processing requires the repositioning of food between the teeth, as modulated by a soft tissue appendage like a tongue or lips. In this study, we trace the evolution of lips and ligaments, which are used during prey capture and prey processing in an herbivorous group of fishes. Pacus (Serrasalmidae) are Neotropical freshwater fishes that feed on leaves, fruits, and seeds. These prey are hard or tough, require high forces to fracture, contain abrasive or caustic elements, or deform considerably before failure. Pacus are gape-limited and do not have the pharyngeal jaws many bony fishes use to dismantle and/or transport prey. Despite their gape limitation, pacus feed on prey larger than their mouths, relying on robust teeth and a hypertrophied lower lip for manipulation and breakdown of food. We used histology to compare the lip morphology across 14 species of pacus and piranhas to better understand this soft tissue. We found that frugivorous pacus have larger, more complex lips which are innervated and folded at their surface, while grazing species have callused, mucus-covered lips. Unlike mammalian lips or tongues, pacu lips lack any intrinsic skeletal or smooth muscle. This implies that pacu lips lack dexterity; however, we found a novel connection to the primordial ligament which suggests that the lips are actuated by the jaw adductors. We propose that pacus combine hydraulic repositioning of prey inside the buccal cavity with direct oral manipulation, the latter using a combination of a morphologically heterodont dentition and compliant lips for reorienting food.